RAPTOR RESEARCH

Volume 6. Supplement Part A. 1972

SPECIAL CONFERENCE ON CAPTIVITY

BREEDING OF RAPTORS-A REPORT
Sponsored by Raptor Research Foundation, Inc.
Held at Sioux Falls, South Dakota, U.S.A., November 22-23, 1971

Part A. Introduction.

by
Byron E. Harrell
Biology Department
University of South Dakota
Vermillion, South Dakota 57069

This conference was planned to assess the situation, to explore in depth the
problems encountered, and to discuss possible future directions in raptor breed-
ing. Formal talks or papers were limited; topics were organized into ten panel
discussions on Biology and Management followed by a session on Evaluation
and Planning. The program was intended to aid participants in breeding pro-
jects. Announcements were sent to all Raptor Research Foundation members
and to all zoos in the United States and Canada.
The conference was held at the Holiday Inn in Sioux Falls, South Dakota.
There were three sessions on Monday and two on Tuesday, November 22 and
23, 1971. Byron E. Harrell was the coordinator. Arrangements and aid at the
meeting were provided by Donald V. Hunter, Jr., Dan O’Brien, Lee Eberly, and
Joyce Harrell. Program suggestions were made by Tom Cade, James Enderson,
Robert Berry, and Richard Fyfe. We are indebted to the ten panel chairmen
who made arrangements for their own topics and to all of the panel members
for their presentations. They are all listed below. The conference was tape re-
corded so a transcript could be prepared. Another conference in 1972 is antici-
pated.
The program was organized into the following panels.
1. Breeding Stock Inventory (Monday moming)-Donald V. Hunter, Jr.,

Chairman, Bob Hinckley, Roger Thacker, Bruce Wolhuter.

2. Housing and General Management (Monday morning) -Richard Fyfe,
Chairman, James Enderson, Heinz Meng, James Weaver.
3. Health and Nutrition of Breeding Pairs (Monday morning)— William H.
Halliwell, Chairman, David E. Allen, C. Wendell Carlson (his presentation on
Monday afternoon), James McIntyre, Richard D. Porter.

A1 Raptor Research 6(Supp.):Al-A6, 1972

 .

A2 RAPTOR RESEARCH Vol. 6, Supp. A

4. Photoperiod Problems and Management (Tuesday afternoon)— Stanley
Temple, Chairman (substituting for Tom J. Cade who could not attend).
5. Reproductive Endocrinology (Tuesday afternoon)— Erich Awender, Chair-
man, Timothy Lawson, James C. McIntyre, Stanley Temple.
6. Behavioral Consideration to Egg Laying (Tuesday afternoon)— R. Wayne
Nelson, Chairman, Richard Fyfe, Frances Hamerstrom, Richard R. Olendorff,
Dr. and Mrs. Noel Snyder (not present; contribution read by Nelson) [at the
conclusion of this panel J. Gregory Thomas spoke about closed circuit televi-
sion recorders, their use in observation, and the kinds of equipment available]
7. Artificial Insemination (Tuesday afternoon)— Robert Berry, Chairman,
Frances Hamerstrom, Walter Morgan, Stanley Temple.
8. Incubation, Natural and Artificial (met with Panel 9 Tuesday morning)—
Richard Olendorff, Chairman, Robert Berry, James Enderson, John Snelling,
Stanley Temple.
9. Rearing, Parental, Foster, and Hand (met with Panel 8 Tuesday morning)—
James Enderson, Chairman, Robert Berry, Richard Olendorff, John Snelling,
Stanley Temple.
10. Health and Nutrition of Young (Tuesday afternoon)— Richard D. Porter,
Chairman, William H. Halliwell.
11. Evaluation and Planning (Tuesday afternoon)— Donald V. Hunter, Jr.,
Chairman, Panel Chairmen and members.

A packet of 58 pages of material was given to all who had preregistered for
the conference. Packet materials included:

1. Program (1 page).
2. Proceedings of a discussion Meeting on Captive Hawk Breeding (8 pages).
3. [6 pages; this was a copy of the MS of a paper in press in RRN; it was in-
cluded without specific permission, but in response to the author’s desire that
his views get full consideration; he resented this inclusion, withdrew the paper
from publication, and submitted it elsewhere; it is therefore omitted from this
list.]

4. Galicz, G. and others,“Autumn Egg Production by Captive Peregrine Fal-

cons” (3 pages, see RRN 5(6): 150-151, 1972).
5. Breeding Project Questionnaire (1 page; this and the replies (6-12) are
published in Raptor Research 6(1): 16-18, 21-31 as B.P.I.E. Nos. 28-34).
6. Campbell, J. A. Report on Peregrine, Merlin, Red-tail (3 pages).
7. Allen, David. Report on White-tailed Sea Eagle (3 pages).
8. Meng, Heinz. Report on Peregrine (3 pages).
9. Hunter, Donald V. Report on Goshawk (1 page).
10. Hunter, Donald V. Report on Peregrine (1 page).
1 1. Hunter, Donald V. Report on Gyrfalcon (1 page).

12. Hunter, Donald V. Report on Red-tailed Hawk (1 page).

13. Thacker, Roger. “Estimations Relative to Birds of Prey in Captivity in
the United States of America” (27 pages, RRN 5(5): 108-122, 1972).
14. List of birds of prey currently held at Cornell— Tom J. Cade (1 page).
1972 Breeding Conference— Introduction A3

15. Snelling, John C. Abstract— “Artificial incubation, and temperature as-

pects of hand rearing downy
raptors” (2 pages).
16. Snelling, John C. Abstract— “Artificial incubation of American Kestrel
eggs” (1 page).
17. Enderson, James. Abstract— “Experiences with artificial incubation and
hand-rearing of falcons” (1 page).
18. Olendorff, Richard. Abstract— “Hand-rearing of Buteos” (1 page).
19. Olendorff, Richard. Abstract— “Certificial incubation of Buteo eggs” (1
page).

Additional material (30 pages) handed out at the meeting included:

1-6. Additional questionnaire reports (published in Raptor Research 6(1):

31-41, 1972, as B.P.I.E. Nos. 35-40).
1 Shultz, Philip L. Report on Prairie Falcon (6 pages).
.

2. Canadian Wildlife Service. Report on Prairie Falcon (3 pages).

3. Canadian Wildlife Service. Report on Prairie Falcon (2 pages).
4. Canadian Wildlife Service. Report on Peregrine Falcon (3 pages).
5. Canadian Wildlife Service. Report on Peregrine Falcon (3 pages).
6. Canadian Wildlife Service. Report on Peregrine Falcon (3 pages).
7. RRF Pathology Committee, names and addresses of members (1 page, part
of annual report).
8. Zu/Preem flier (1 page).
9. “The Nutrient Requirements of Chicks” (2 pages).
10. Morgan, Walter. “Artificial Insemination” (outline of procedure; 1 page).
1 1. Artificial Insemination Panel; Practical considerations in artificial insemi-
nation; Berry, Robert B. Abstract— “Artificial insemination of captive gos-
hawks” (4 pages).
12. Berry, Robert B. Abstract-“Foster Rearing of Raptors in Captivity” (1
page).

Those recorded in attendance are listed below.

Allen, David E.,11332 Sunnyslope, Kansas City, MO 64134 (Panel 3).

Andersen, William C., Otero Jr. College, La Junta, CO 81050.
Awender, Erich, 1317 La Cresta Dr., Freeport, IL 61032 (Panel 4 Chairman).

Berry, Robert B., RD 1, Chester Springs, PA 19425 (Panel 7 Chairman, Panel 8,

Panel 9).
Beske, Alan E., 83 North 300 East, Springville, UT 84663.
Blankespoor, Gilbert W., Dept, of Biology, Augustana College, Sioux Falls, SD
57102.
Bond, Frank M., 540 Camino Rancheros, Santa Fe, NM 87501.

Cade, Tom J., Sec. of Ecol. and Syst., Bldg. 6, Langmuir Lab., Cornell Univer-
sity, Ithaca, NY 14850 (not present; originally Chairman of Panel 4).
A4 RAPTOR RESEARCH Vol. 6, Supp. A
Campbell, John, Box 130, Black Diamond, Alberta, Canada.
Carlson, C. Wendell, Poultry Science Dept., South Dakota State University,
Brookings, SD 57006 (Panel 3).
Carnie, S. Kent, 206 Sheridan Rd., El Paso, TX 79906.
Cranson, Babette, RR 2, La Junta, CO 81050.
Crawford, J. Jeff, RR 2, Box 53, Durand, IL 61024.
Crouse, Rodney N., 4311 S. 71st St., Apt. 3, Milwaukee, WI 53220.
Crowe, Sam, 152 Monte Key, Los Alamos, NM 87544.

Dague, Phyllis R., Lab. of Ornithology, 159 Sapsucker Woods Rd., Ithaca, NY
14850.
Donald, Tom, c/o 1930 Cowan Cres., Regina, Saskatchewan, Canada.

Eberly, Lee, Rt. 2, Box 94, Vermillion, SD 57069.

Enderson, Jim, 10070 Otero Ave., Colorado Springs, CO 80908 (Panel 2, Panel
8, Panel 9 Chairman).

Freitag, Gary, 230 Daphne Way, Broomfield, 80020. CO

Fuller, Mark, 1231 Raymond Ave., St. Paul, MN 55108.
Fyfe, Richard, RR 1, Fort Saskatchewan, Alberta, Canada (Panel 2 Chairman,
Panel 6).

Galicz, George, 13281 60 Ave., Surrey, BC Canada.

Goben, Bill, S., Renton, WA 98055.
3016 Benson Rd.
Gorrell, Wallace, 1709 S. Duluth Ave., Sioux Falls, SD 57105.
Graham, Dick, Rocking R Rd., Colorado Springs, CO 80915.
Greichus, Yvonne, Station Biochemistry, South Dakota State University, Brook-
ings, SD 57006.
Grier, Jim. Lab. of Ornithology, Cornell University, Ithaca, NY 14850.

Halliwell, William H., Dept, of Pathology, School of Vet. Med., Columbia, MO

65201 (Panel 3 Chairman, Panel 10).
Halverson, Keith J., Great Plains Zoo, Sioux Falls, SD 57104.
Hamerstrom, Fran, Plainfield, WI 54966 (Panel 6, Panel 7).
Harrell, Byron, Biology Dept., University of South Dakota, Vermillion, SD
57069 (Coordinator).
Hinckley, Bob, RR 2, Monee, IL 60449 (Panel 1).
Hoback, Frank, 942 Canyon Rd., Santa Fe, NM 87501.
Hughes, David, Bat worthy, Chagford, Devon, UK
Hughes, Francis, Apt. 1417, 100 De Gaspe, Nun’s Island, Montreal, Quebec,
Canada.
Hunt, Grainger, Dept, of Zoology, University of Texas, Austin, TX 78712.
Hunt, Deidre, Dept, of Zoology, University of Texas, Austin, TX 78712.
Hunter, Donald V., RR 3, Centerville, SD 57014 (Panel 1 Chairman, Panel 11
Chairman).

Ketchum, Sherm, 1 1610 W. Lincoln, West Allis, WI 53227.

 .

1972 Breeding Conference— Introduction A5

Lawson, Tim, 7013 Hawks Dr., Olympia, WA 98501 (Panel 5).

Layman, Steve, 813 East 4th, Ellenburg, WA 98926.

MacLeod, Mark, 52 Lake Park, Champaign, IL 61820.

McIntyre, James C., 2937 Country Club Drive, Colorado Springs, CO 80909
(Panel 3, Panel 5).
Marcus, Stanley A., Rt. Coleman, MI 48618.
2,
Mattingly, Edwin, University of New Mexico, 5 OS /! B. Prosperity Ave. SE, Al-
1

buquerque, NM 87102.
Meng, Heinz, State University, New Paltz, NY 12361 (Panel 2).
Mesch, Ken, Dept, of Biology, University of Colorado, Boulder, CO 80302.
Morgan, Walter, Poultry Science Dept., South Dakota State University, Brook-
ings, SD 57006 (Panel 7).

Nelson, Alora L., Dept, of Biology, University of Calgary, Calgary, Alberta,

Canada.
Nelson, R. Wayne, Dept, of Biology, University of Calgary, Calgary, Alberta,
Canada (Panel 6 Chairman).

Oar, Jack, RR 5, Box 226, Rockford, IL 61108.

Oberg, John, Route 1, Owen Center Rd., Rockton, IL 61072.
O’Brien, Dan, Rt. 3, Meckling, SD 57044.
Olendorff, Richard R., 3317 Olympus Dr., Bremerton, WA
98310 (Panel 6,
Panel 8 Chairman, Panel 9).

Platt, Joseph B., Dept, of Zoology, Brigham Young Univ., Provo, UT 84601.
Porter, Richard D., Patuxent Wildlife Research Center, Laurel, MD 20810 (Pan-
el 3, Panel 10 Chairman).

Rafuse, Bob, 135 Mayfair Cres., Regina, Saskatchewan, Canada.

Richards, Gerald L., 510 Idaho Ave., Provo, UT 84601.
Ricken, Christine, 1200 College Ave. No. 107, Boulder, CO 80302.
Russell, W. C., Omaha Zoological Society, 10th & Deerpark Blvd., Omaha, NE
68107.

Schubert, Teddy, 20 Spring Lane, West Caldwell, NJ 07006.

Schwartz, Charles IL, Box 385, Rawson, OH
45881
Sherrod, Steve, Box 282, Springville, UT 84663.
Shultz, Philip L., 107 Cienega St., Santa Fe, NM
87501.
Simons, Bob, Dept, of Zoology, University of Illinois, Urbana, IL 61801.
Simonyi, Joseph S., 456 Sparton Rd., Victoria, B.C., Canada.
Smylie, Tom, Box 174, Timeras, 87059.NM
Snelling, John C., Cornell Lab. of Ornithology, 159 Sapsucker Woods Rd.,
Ithaca, NY 14850 (Panel 8, Panel 9).
Snyder, Mrs. Noel, 5113 127th Ave., Tampa, FL 33617 (not present, contrib-
uted material to Panel 6).
A6 RAPTOR RESEARCH Vol. 6, Supp. A
Snyder, Noel, 5113 127th Ave., Tampa, FL 33617 (not present, contributed
material to Panel 6).
Stoddart, John, 430 S. Franklin, Denver, CO 80209.
Stoudenmire, T. Malory, 4233 E. 189th, Cleveland, OH 44122.
Swartz, L. G., Dept, of Biology, Bunnel Bldg., University of Alaska, College,
AK 99701.
Taggart, Joe, 143 Browning Lane, Rosemont, PA 19010.
Temple, Stanley A,, Lab. of Ornithology, Cornell University, Ithaca, NY 14850
(Panel 4 Chairman, Panel 5, Panel 7, Panel 8, Panel 9).
Thacker, Roger, Dept, of Animal Labs., Ohio State University, 400 W. 12 Ave.,
Columbus, OH 43210 (Panel 1).
Thomas, J. Gregory, 15 Rolling Springs Court, Carmel, IN 46032 (extra presen-
tation following Panel 6).
Torgersen, Tom, 7844 Torgersen Rd., Eureka, CA 95501.
Trefry, Philip A., c/o R. Fyfe, RR 1, Fort Saskatchewan, Alberta, Canada.

Walker, Skip, Dept, of Biology, University of Alaska, College, AK 99701.

Weaver, Jim, 159 Sapsucker Woods Rd., Ithaca, NY 14850 (Panel 2).
White, Clayton M., Dept, of Zoology, Brigham Young Univ., Provo, UT 84601.
Wilbowr, Craig R., Box 878, Chelan, WA
98816.
Wolhuter, Bruce R., 1405 Wiltshire, Colorado Springs, CO 80906 (Panel 1).

This report will be issued in parts corresponding to the first ten panels. Per-
tinent material from the last panel will be included in the appropriate part. The
editors of the individual parts utilized the tapes and the transcription mentioned
above or their own to prepare the material. In some cases the transcript has
been modified for clarity, reduction of repetition, and to provide for more logi-
cal sequence. In other cases the individual panel members have rewritten their
presentations. The individual reports will be published in the sequence in which
they are completed. Pagination will be continuous in this supplement to Vol-
ume 6 of Raptor Research but page numbers will be preceded by the letter des-
ignating the part to distinguish the Supplement from the regular issues.
RAPTOR RESEARCH
Volume 6. Supplement Part B. 1972

SPECIAL CONFERENCE ON CAPTIVITY

BREEDING OF RAPTORS-A REPORT
Sponsored by Raptor Research Foundation, Inc.
Held at Sioux Falls, South Dakota, USA, November 22-23, 1971

Part B. Incubation, Natural and Artificial (Panel 8)

edited by
Richard R. Olendorff
3317 Olympus Drive
Bremerton, Washington 98310

Panel Members: Richard R. Olendorff, Chairman; Robert Berry; James Ender-

son; John Snelling; Stanley Temple.

INTRODUCTION

OLENDORFF. One aspect of the wholly artificial method of rearing

critical
birds of prey is artificial incubation. Really, it should not be a problem at all
considering the state of the art in the poultry business. I would venture a guess
that if we gave some fertile eggs to an avian scientist specializing in incubation,
he would chuckle at our floundering and hatch out a high percentage of the
birds. Berry and Temple will talk about using conditions suitable for chickens
or hawks. For this discussion I have considered what literature was available on
the various aspects of incubation by falconiforms only, and will review this
briefly as we go. I will invite discussion by the panel and the audience of each
point separately. Some of the things we will be talking about include tempera-
ture, humidity, egg-laying, attentiveness to the first egg of the clutch, candling,
and turning. Some other considerations we might want to talk about are egg
weight loss, length of the incubation period, the roles of the sexes in incubation,
the mechanics of hatching, the analysis of failure to hatch and incubation of
hawk eggs by chickens.

TEMPERATURE CONSIDERATIONS
OLENDORFF. Beginning with temperature, all we have to go on is what has

B7 Raptor Research 6(Supp) B7-B3 5 :

, 1 972
B8 RAPTOR RESEARCH Vol. 6, Supp. B

been tried and what small successes have resulted. First we should dispense with
the notion that there is a single incubation temperature for a species. We should
probably speak of ranges of incubation temperatures and optimum incubation
temperatures. Huggins (1941) studied incubation temperatures of wild Marsh
Hawks and found that during attentive periods the eggs were incubated at an
average temperature of 90.1 F. The temperatures varied from 82.9 to 95.7 F
over an extended period of observation. In the laboratory Stanley and Witschi
(1940) used a temperature of 96.0 F to incubate Red-tailed and Cooper’s Hawk
eggs when older developmental stages were required for their embryological
studies. I incubated two Swainson’s Hawk eggs full term at 96.0 F while con-
ducting my dissertation research, but the embryos died in the very late stages
of incubation several days beyond the expected hatching date. I understand
that this is one result of too low an incubation temperature, but it is difficult
to attribute failure to any one cause. Moving up the temperature scale, Hunter
(1970) attempted to get incubation temperature readings from captive Gos-
hawks. The average of ten readings was 95.4 F, with a maximum of 97.2 F.
Hunter thought that this was quite low. Enderson (1971) used a temperature
of 97 to 98 F to artificially incubate Prairie Falcon eggs with some success. He
also placed a dummy egg with a thermister in it under some Prairie Falcons. At
no time did the temperature exceed 95 F in the center of the egg. I will let him
expand on this when I finish. Lawson and Kittle (1970) incubated American
Kestrel eggs at 99.0 F and got large embryos, but none hatched. Kish and Clark
(1971) used 99.5 to 99.8 F to incubate 5 Golden Eagle eggs, but all were appar-
ently infertile. Dr. H. Mendelsohn of Tel Aviv, Israel, hatched a clutch of Long-
legged Buzzard eggs at 100.4 F. Enderson (1971), again with Prairie Falcons,
used 101 to 102 F in a still air incubator and 99.2 F in a forced air incubator
to achieve success. Again, I will leave the details to him. Hancock (1971) used
temperatures of 102 to 103 F top of the egg and 99 F bottom of the egg to in-
cubate Peale’s Falcon eggs, but the eggs were infertile. Finally, for comparison,
the optimum for chicken eggs according to Bellairs (1960) is 101.3 F which
corresponds to the body temperature of the hen. Along these lines, I have seen
the body temperature of Marsh Hawks, Peregrines, Red-tails and Swainson’s
Hawks published as roughly 105 F. Kestrels were somewhat higher at 107 F.
Some unpublished data I have indicate that body temperatures of Red-tails and
Swainson’s Hawks are actually lower than that, about 99 to 101 F depending on
the time of day. Perhaps someone can recall other representative body tempera-
tures. Those I took were over a twenty-four hour period Vi to % inch deep in the
cloaca. The figures of 105 F may have been deep core temperatures, but the
point should be made that incubation temperatures probably do not exceed
peripheral body temperature. Another temperature consideration is the matter
of allowing eggs to cool once or twice a day. Certainly the eggs of wild birds are
cooled periodically during inattentive periods. Eric Stauber reported that Fes-
ner (see Stauber (1971)) cooled eggs to room temperature twice a day and I
have seen other references to it. It is difficult for me to see any advantage in
doing so. Perhaps there is a greater exchange of oxygen and CO2 during the
cooling and reheating period. If anyone can shed any light on this, please do so.
1972 Breeding Conference— Panel 8, Incubation B9

So much for my discourse on temperature. I’d like to turn it over to Jim Ender-
son now.

ENDERSON. In the packet that you have received I have spelled out some of
the temperatures I have used. These are as follows: In 1970 four Prairie Falcon
eggs incubated at 99.7 F, forced air temperature, 75 percent humidity and tilt-
ed through 1 50 degrees at three-hour intervals, while resting on the small end
of the egg, died at ages varying from one to three weeks. In 1971 another cap-
tive-produced egg hatched after full-term incubation at 99.3 F still air tempera-
ture at the upper surface of the shell, 70 percent humidity and about five turn-
ings per day using freshly washed hands. About 80 percent humidity was reach-
ed in the last three days of incubation with an air-pump driven power atomizer.
Two other embryos died at about 7 days of incubation when the incubator rose
to about 101 F because of a faulty thermometer. Hatching in Prairie Falcons
requires 50-60 hours; eggs were not rotated at this time. Hatching was aided
successfully in one case when it was clear that the chorionic vessels were dry.
The eggs were candled daily; embryos appear most active around 21 days, the
air space enlarges rapidly after 30 days and hatching of four captive Prairie Fal-
con eggs required about 35 days. Chicks hatched from the pipped condition in
less than 30 minutes. There are a couple of other things I would like to men-
tion. Thermisterized whole eggs appear to give readings 1 to 2 F below the air
temperature immediately surrounding the egg. I believe that John Snelling ex-
perienced that as well with thermisterized American Sparrow Hawk eggs. An-
other thing you have to watch is the distinction between still air and forced air
temperatures. Forced air temperatures are usually below those used in still air
incubators. Another problem is where do you measure the temperature sur-
rounding the egg? In a still air incubator my technique has been to measure the
temperature at the upper surface of the egg. In a still air incubator the tempera-
ture there is much different than at the lower surface of the egg, the egg being
progressively cooler as one goes down through the thermal stratification in a
still air incubator. You get into all sorts of complexities because the embryo, at

least in early stages, is within a quarter inch, perhaps, in an egg the size of a
Peregrine (maybe a little more), of the upper surface of the egg. The forced air
incubator is quite a different matter. I strongly suspect that temperatures ex-
ceeding 99.5 F are too high for incubation of the eggs of most large raptorial
birds.

HINCKLEY. Is that temperature a still air or a forced air temperature?

ENDERSON. I would not raise the temperature over 99.5 F in any case, even
at theupper surface of the egg in a still air incubator. The standard temperature
used by Cornell in a forced air incubator for chicken eggs is 99.7 F.

HUNTER. That is equivalent to 101.6 still air, right?

ENDERSON. I don’t know. Where do you measure the still air temperature?
BIO RAPTOR RESEARCH Vol. 6, Supp. B

Actually, think if the temperatures are of the order of 99.2 to 99.4 in a forced
I

airincubator, the eggs can stand some latitude temperature wise. Probably even
temperatures like 98.8 will work. The only difference there would be the time
for development.

HUNTER. I think it is important to define exactly where we are measuring

the still air temperature. We don’t need to worry about forced air. Don’t you
think some kind of a standardization is necessary? Where do you measure still
air temperatures? I measure it 3/8 of an inch below the top of the egg. And then
it depends on whether your heating element is above or below, because in a still

air incubator the hottest air rises because of the thermal qualities. There are a
lot of variables here which I really think need to be defined.

OLENDORFF. The incubator I used had two vents in the top which I left
open assuming that that was for circulation of air so that there should be a cer-
tain amount of circulation even in a still air incubator.

LAWSON. Ithink the poultry people measure their temperature one third of
the way from the top of the egg.

SWARTZ. Jim, do you find the disparity between air temperature and therm-
ister temperature to be the same in forced air and still air?

ENDERSON. I don’t have experience with still air thermisterized eggs.

WALKER. How does this disparity come to be? Do you calibrate these to air
temperature? If you did so, you would get a direct read out of air temperature.

ENDERSON. A possibility is that the lower temperature of the thermister-

ized egg due to evaporative cooling, although this is a little hard to explain.
is

It would seem to me to take considerable loss of moisture to cool an egg 1-2

degrees at that temperature.

OLENDORFF. Let’s get a little more input on temperature.

BERRY. The incubator

I used to incubate Northern Goshawk eggs was a
forced air, Roebuck type incubator. The temperature ranged from
Sears and
99.0 to 99.5 F throughout the hatching period. Relative humidity was kept be-
tween 86 and 90 percent. I would say the average was between 86 and 88 per-
cent. Since the relative
humidity outside was so high, it was difficult to get the
relativehumidity down below 90 percent in the incubator. I also hatched some
Peregrine Falcon eggs which were taken from the Canadian arctic. I hatched
them at the same temperature, 99.0 to 99.5 F. I took some temperature read-
ings of both the Northern Goshawk and the Bantam hen. This involved
taking
the temperature about 2 dozen times throughout the nest to get a median tern-
 1

1972 Breeding Conference— Panel 8, Incubation B1

perature within the nest area. The Goshawk temperature ranged from 96 to 99
F with median of 98.5 F. The chicken ranged from 97 to 101 F with a median
a
of 99.5. The chicken was a full degree warmer than the Goshawk which, accord-
ing to A. van Tienhoven from Cornell and John Skinner from the University of
Wisconsin, may have contributed to the death of several Goshawk embryos dur-
ing the 1970 year— that one degree temperature difference. The maximum skin
temperatures of the brood spots were also measured for the Goshawk and the
chicken. The Goshawk had a temperature of 101 F and the chicken 103 F.
These are not necessarily very accurate temperatures, but it does indicate that
the chicken is somewhat higher in incubation temperature than the Goshawk,

and apparently some of these other hawks do have a lower incubation tempera-
ture than we might expect in domestic fowl.

UNKNOWN. Did you cool the eggs which were successful?

BERRY. Yes. All eggs were cooled twice per day every day including the
days immediately preceding hatching for 15 to 20 minutes in a room tempera-
ture of approximately 70 F. Werner Fesner suggests that this cooling is very
important to allow the reabsorption of oxygen.

HUNTER. The Goshawk under which I took the temperatures would not
tolerate a remote sensing device in the nest (Hunter, 1970). I put a temperature
thermometer in the nest and she would throw that out except when I went in
at night and worked it up through the nest to the desired height. In this way I
got the unexpectedly low temperatures. I put a chicken egg under her and it
took her 28 days to hatch it. This indicates that the Goshawk is probably incu-
bating at a lower temperature than the chicken.

SWARTZ. The poultry people abandoned this cooling business some years
ago, finding that it is not only dispensible, but that the implications are that it
may very slightly lower hatchability. The last is in doubt.

BERRY. I think just the reverse of that. John Skinner at the University of
Wisconsin also corroborated Werner Fesner and said he thought that cooling
would definitely increase hatchability of the eggs.

FYFE. Relative to some of our observations on the prairies, the impression

we have is that the birds very seldom remained off of the eggs for any more
than a short time. There was a definite attempt for the male to get immediately
back to the nest as soon as the female came for food.

TURNING

OLENDORFF. It is generally agreed that eggs should be turned several times

daily to prevent adhesion of membranes and, according to the Lanyons, to pro-
B12 RAPTOR RESEARCH Vol. 6, Supp. B

vide exercise for the developing embryo.

NELSON. Bob Berry, were you turning the Goshawk and Peregrine eggs a
certain number of times per day?

BERRY. All eggs were turned every three to four hours every day, with a re-
duction in frequency 3 to 4 days prior to hatching. They were turned through-
out the day and night, 1 80 degrees at a turn.

FYFE. We didn’t actually observe any turning as such with our Prairie Fal-
cons, but the birds did shuffle on the nest to readjust their position. This would
probably roll the eggs. This often happened several times in an hour. Other
times the bird would sleep for an hour or so at a time.

NELSON. I have written to a few people here about the studies of gulls
which were done I believe in Europe by Dr. Rudy Drent of the University of
British Columbia at Vancouver, and there have been a couple of other studies as
well of wild birds. They marked the eggs with lines at varying degrees around
the egg so that they could determine what part of the egg is uppermost. All of
the shuffling we are seeing in the nest, the various shifting of the feet which
appears to be threatening the egg, seem to be to keep the egg in exactly the
same position so that the embryo is uppermost. If you take a reasonably well-
incubated egg and mark a line on it and you put it in a bowl of water, the egg
will take up a particular position because a certain part of the egg is heavier. It
happens that the embryo part of the egg is lighter and it always floats upper-
most. This is the whole purpose apparently of why birds of a variety of species
shuffle the eggs. It is shuffling to keep the embryo uppermost and closest to the
brood patch of the adult. Artificial incubation in incubators has been done with
uncountable numbers of chicken eggs. It is very likely that in chickens we have
selected for those birds that lay eggs for which it doesn’t matter whether they
are upside down or not. Jim Enderson said awhile ago that there is a very great
difference between the temperature at the top of the egg and the temperature
at the bottom. With the gulls and with the godwits in Europe it has been proven
quite conclusively that all of the movement of the eggs in the nest are to keep
the embryo uppermost against the brood patch.

TEMPLE. Did that fellow ever candle an egg? You can follow an embryo all

the way around an egg as you turn it. The embryo floats in the egg.

NELSON. They have opened a variety of eggs at various stages and it is quite
of the supplement for Behaviour.
so. It is in the latest issue

HAMERSTROM. This all begins to make some sense to me. I marked my

Golden Eagle eggs on the bottom as they were laying so I could tell which egg
was which. As I went to the nest day after day I never could find those mark-
ings. I thought that they had worn off. When I took the eggs away, the marks
1972 Breeding Conference-Panel 8, Incubation B13

were still there. This suggests, although my eyes are not very good without
glasses, that those markings were usually at the bottom.

WOLHUTER. Do you have any theories as to why the markings were down?
Could it have been the bird’s reaction in wanting to put the markings down, or
do you think that this was just the position the eggs happened to be in.

HAMERSTROM. I don’t think it was the bird’s reaction.

HUNTER. This brings up a good point that somebody really ought to check
out. Does the position of the shell have anything to do with where the embryo
is? I think that no matter where you turn the shell the embryo will stay on top,
because it is floating in fluid inside. It is not attached to the eggshell in any
way. I have often questioned in my own mind why an egg needs to be turned,
but everybody does.

TEMPLE. That is why you

turn it. That embryo remains free. If you
exactly
don’t turn membranes will adhere to the shell. Then, later on, if the
it, the
membranes adhere to the shell and you turn it, you get rupturing of blood ves-
sels.

ENDERSON. The embryo won’t float if you don’t turn it. I think in this
case one picture or 3600 pictures is worth a thousand words. In the summer of
1970 Gerald Swartz, Stan Temple and I took some time-lapse motion pictures
of Peregrine Falcons incubating eggs along the Yukon River. I want to run a
section of that film for you now. This is just the part where the bird is incubat-
ing eggs. This is, of course, greatly speeded up. The frames were taken about
2V2 minutes apart. You get a feel for the activity over the eggs. The change in
position of the bird on the nest averages out to about once every 30 minutes.
I presume that changes such as these involve some movement of the eggs. We

did see active turning as well. The manuscript dealing with these films is being
submitted to the Living Bird. Only less than one percent of the time are the
eggs exposed.

TEMPERATURE AND TURNING

TEMPLE. This is a summary of our data from seven Peregrine eyries at which
we had time-lapse cameras. A couple of interesting things showed up including
the one that Jim Enderson just mentioned. If we assume that Peregrine Falcons
in the wild do it the right way, we can probably get some pretty good insight
into some of these questions. One very important point is this thing on periodic
cooling of the eggs. I think the thing that was surprising to us is that inattentive
periods— times when the eggs were uncovered and not actively incubated— were
less than one percent of the time. The average time that these eggs were ex-
posed was less than three minutes. In other words, Peregrine Falcons do not
B14 RAPTOR RESEARCH Vol. 6, Supp. B

periodically cool their eggs. Less than three minutes of exposure does not allow
the eggs to cool down hardly at all. The other thing you will notice -here is that
the inattentive periods drop off in the last five days before hatch, during which
the embryo breaks into the air sac. You will then hear the embryo peeping and
scratching around in the egg. With regard to the role of the sexes in incubation
for those of you who are going to allow your birds to incubate, quite surprising
again to us, the male incubated approximately one-third of the time. The times
they were on the eggs per shift were about two hours for the male and. 3%
hours for the female. The male definitely (in wild pairs) contributes substan-
tially to the incubation of the eggs. This is quite contrary to impressions of
quite a few people who have published on this for Peregrine Falcons. The im-
portant thing to note is that the inattentive periods are very insignificant.

HAMERSTROM. How many eyries were involved in your study?

TEMPLE. Seven. Not all of them were involved in each of the periods into
which we broke the incubation period.

SMYLIE. Did the inattentive periods correspond to the times when they
were changing?

TEMPLE. The information would suggest that it is usually when the male
brings food in and the falcon leaves the eggs to gomeet him. The eggs are left
alone while the food exchange takes place away from the eyrie.

SMYLIE. When he comes in to incubate, how long does it take to make the
change from male to female? Even if it is 30 seconds, if it happens enough times
during the day, you are going to get up to a 20 to 30 minute total that the eggs
might be exposed.

TEMPLE. Here we have the percentage of the total time those eggs were ex-
posed and it is less than one percent.

FYFE. The exchange is a short thing.

TEMPLE. It is very short. The eggs are not allowed to cool. Anyone who has
worked with eggs knows that in three minutes that egg is not going to cool very
much.

UNKNOWN. Have you every seen the male incubate through the night?

TEMPLE. Yes. The male incubated during the night in approximately the
same proportion as he incubated— about a third of the nights.

WOLHUTER. Perhaps Dr. Porter can agree or disagree with me, but it seems
that male kestrels do more of their incubating at night.
1972 Breeding Conference— Panel 8, Incubation B15

PORTER. We didn’t record data on what happened at night, although the

literature has one or two references to it. I believe that Tom Cade published on
this.

TEMPLE. Ours seemed to be completely random. In other words, about one-

third of the nights— about what you would expect— the male incubated.

UNKNOWN. What was air temperature at these nests?

TEMPLE. This was on the Yukon River so the air temperature was down to
60 to 70 F.

PORTER. Is there any indication of abnormality which might be expected

from the pesticide situation?

TEMPLE. We didn’t observe anything we could directly relate to pesticides.

We did find that at the two eyries out of the seven that failed to produce
young, eggs were broken. Thin eggshells did accompany failure. In one nest
where eggs were broken and young did not hatch, the tiercel did not fulfill his
responsibility during incubation. We only had a camera at this eyrie for a short
period because we had a malfunction, but during that time when the female
would get off apparently to feed, the male would not incubate his whole atten-
tive period.

ENDERSON. Dr. Porter, that is true. At the two nests that failed, we found
the thinnest shelled eggs. Egg breakage occurred at other eyries as well. Small
sample size. I’m sorry.

NELSON. When you are talking about one percent or so of the time, this
amounts to about 1 5 minutes per day.

TEMPLE. Right. The inattentive periods averaged about three per day. There
were three intervals when the eggs were left uncovered each day. The eggs were
left uncovered for an average of less than three minutes each time.

NELSON. How often did you expose a frame?

TEMPLE. About every minutes. This gives us a less than three minute in-
terval. The bird is incubating in one frame, he is gone the next and he is back
the next. This gives you a maximum time of five to six minutes off per ex-
change, but we are averaging.

GRIER. You have thousands of pictures there and they are using a sampling
procedure.

NELSON. In any case, someone made the point that cooling of the eggs was
B16 RAPTOR RESEARCH Vol. 6, Supp. B

required for oxygen reabsorption. Fifteen minutes per day may be very impor-
tant.

TEMPLE. But it is not 1 5 continuous minutes.

NELSON. But for oxygen reabsorption it doesn’t matter.

TEMPLE. Who brought up this oxygen reabsorption business?

BERRY. Werner Fesner is the source of that little bit.

TEMPLE. Fesner is not an expert in avian biology, and I would question that
he had done anything to indicate this.

OLENDORFF. From the laws of physics, most of the cooling will occur dur-
ing the three to five minutes that the eggs are uncovered. In other words,
first
when the difference between ambient temperature and egg temperature is the
greatest, the rate of cooling will be the greatest, and the rate will decrease as
cooling continues. I cannot tell you if it is significant or not, but the cooling is

faster at first.

TEMPLE. The studies they have done on the cooling of eggs indicate that the
albumen is a tremendous insulator. Uncovered eggs retain their temperature for
quite a while.

HUNTER. It looks like we are just conjecturing on this. No one really knows.
Maybe someone ought to get some data on this.

GOBEN. I watched with my Ferruginous Hawks just to see

this fairly closely
this cooling period. On
the exchange they were very careful not even to let cool
air get to the egg. If the female was on, the male would stand right beside her,
slip a wing half underneath her and she would lift off. He would then slip a
wing over the egg and move over on to the eggs. The opposite exchange would
occur the same way. You never get to see the eggs.

FYFE. Our field observations closely parallel what you have here
would very
for the Peregrines. Most of the movement of eggs was just with the foot move-
ments. The time off the eggs was usually just the time it took for the female to
go and get food and the male to come back and go on to the eggs. It would cer-
tainly be less than three minutes. We spent a lot of hours observing, but we
could have missed periods. I don’t think, however, that we would have missed a
long inattentive period. With our captive birds we found the same thing. They
made very sure that the eggs were incubated virtually all of the time.

SWARTZ. On the general point of cooling, I think we have a relatively fool-

ish situation. The point doesn’t deserve too much discussion. As for movement,
1972 Breeding Conference— Panel 8, Incubation B17

I wonder if we are not, in the incubator, substituting with infrequent large

movements for very frequent small movements. It is probably not necessary

for a bird to roll an egg clear over.

TEMPLE. I might add that most poultry people now have automatic turners
in their incubators. This is the way I handled the red-tail egg we got from artifi-

cial insemination. It was moved through a 90 degree arc every hour. The egg
was never completely rolled over. It was rocked back and forth through a 90
degree arc. It was lying on its side in a normal incubating position.

ENDERSON. Someone might experiment with a clock drive turner using

rollers.

TEMPLE. That would save getting up in the middle of the night to turn eggs.

SNELLING. In 1970 I addressed myself to the problem of artificial incuba-

tion in the American Kestrel, specifically to all of the questions we are asking
today. For logistical reasons I only decided to look at temperature relations and
cooling. I had at my disposal a sample of wild laid American Kestrel eggs, 36
fertile ones to be exact. I used a still air incubator, a very simple unit which can
be constructed by hand for less than $15.00. 1 divided my eggs into three tem-
perature treatments. All eggs were in the same incubator. The middle level was
at 101 F (38.5 C). The low level was at 97 F (36.0 C). The upper level was at
104 F (40.0 C). These eggs were divided another way. It was a three by two ex-
perimental design actually. Half of the eggs were cooled twice a day to about
21 C (morning and evening). The results are shown in Tables 1 and 2. These
are the hatching successes for the various temperature treatments. At 38.5 C
(101 F), 11 out of 1 1 eggs hatched normally and they were all raised to fledg-
ing. At 36.0 C, five eggs hatched, while eight did not hatch. At 40.0 C, three
eggs hatched; nine did not hatch. If you perform a Chi Square test on these
data, the conclusion is that 38.5 C (101 F) was the optimum incubation tem-
perature. Bear in mind that there is a distinct difference between still air and
forced air temperatures. This was a still air unit in which normally you would
have a higher incubation temperature. This would correspond to about 99.1 ¥
which is the proper forced air incubation temperature. Apparently American
Kestrel eggs, with respect to temperature, react like chicken eggs. The sensor
was simply a dry bulb thermometer at the top of the eggs. The heating element
was throughout the entire incubator on the sides.

OLENDORFF. You got better hatching at the lower temperature than you
did at the higher. Was that statistically significant?

SNELLING. No. Table 3 shows you what we have with respect to cooling.
The eggs were cooled for 30 minutes twice a day down to room temperature
around 21 C. Of the eggs which were cooled, 1 1 hatched and seven didn’t hatch.
Eggs not cooled, eight hatched; ten did not hatch. The Chi Square test perform-
B18 RAPTOR RESEARCH Vol. 6, Supp. B

Table 1 . Effects of Incubation Temperature and Periodic Cooling on

Artificially Incubated American Kestrel Eggs (Snelling)

Cooling Treatment
Temperature 30 Minutes
Treatment Twice Daily Not Cooled
36 C
no. fertile eggs 6 7
no. eggs hatch 3 2
mean age 1 when taken 10.3 11.6
mean hatch time 2 49.3 78.8
condition unhatched eggs
no. pipped, fully formed 1 3
no. fully formed, yolk sac external 1 -

no. % developed -
1

no. x
h developed 1 1

38.5 C
no. fertile eggs 6 5
no. eggs hatch 6 5
mean age when taken 13.3 12.2
mean hatch time 50.7 44.2
40 C
no. fertile eggs 6 6
no. eggs hatch 2 1

mean age when taken 9.8 13.0

mean hatch time 47.0 62.2
condition unhatched eggs
no. pipped, yolk sac external 4 -

no. pipped, fully formed -

1

no. fully formed -

1

no. % developed -
2
no. Vi developed -
1

Mays
^hours from pip to hatch

Table 2. Hatching Success of Artificially Incubated American Kestrel Eggs

as a Function of Incubation Temperature.

Incubation Temperature
36 C 38.5 C 40 C
no. hatch 5 11 3
no. unhatched 8 0 9

Chi 2 (2 d.f.) = 14.63, p <.001

1972 Breeding Conference- Panel 8, Incubation B19

Table 3. Hatching Success of Artificially Incubated American Kestrel Eggs

as a Function of Periodic Cooling.

Eggs Cooled for 30 Minutes Eggs Not Cooled

Twice Daily
no. hatch 11 8
no. unhatched 7 10

Chi 2 ( I d.f.) = 1 .02, p» .05 n.s.

ed on these data shows no significant difference. My conclusion is that cooling

American Kestrel eggs has no effect on the hatch whatsoever. Cooling is not
detrimental or helpful.

HUNTER. Is there a difference in the time of incubation at the different

temperatures?

SNELLING I cannot say for sure, because I was not sure how old the eggs

were when got them. Table 4 summarizes data I got from thermisterized eggs.
I

Once again, the general pattern of a two to three degree (C) drop with therm-
isterized eggs is seen. First of all, concern yourself only with the right-hand
column. An experimental pair of birds which was fed on pesticides incubated at
34 to 36 C. The control pair which was not fed pesticides incubated their eggs
at 34 C. This is a two to three C drop from what we would expect from incuba-
tor temperature. In the next three figures down— temperatures with respect to
still air incubator— eggs thermisterized in the incubator at the 36 C, 33 C was

Table 4. Temperatures of Eggs Incubated by Captive American Kestrels

and in a Still-air Incubator.

Thermister Coil Thermisterized Egg

under Eggs
Captive Kestrels
experimental! 31 C 34-36 C
control — 34 C
Still-air Incubator
36 C level — 33 C
38.5 C level — 36 C
40 C level — 38.5 C

1 Fed on day-old cockerels injected with 5 ppm Arochlor 1254 and 15 ppm
DDE.
B20 RAPTOR RESEARCH Vol. 6, Supp. B

obtained from the middle of the egg by thermister. At the 38.5 C level, 36 C
was obtained by thermister. At the 40 C level, 38.5 C was obtained. Consistent-
ly lower temperatures at the center of the egg than in the air surrounding the
egg were obtained. I would like to reiterate the point that people should be
very cautious attempting to “naturally” incubate eggs from what they read
from the center of the egg by thermister. If you will recall, the temperature
data showed that a low incubation temperature can be just as bad as a high in-
cubation temperature. There are some data from poultry studies that this is not
the case with fowl, but I think that at this point in time we had better conclude
that a low incubation temperature could be detrimental.

ENDERSON. In 1970 placed a thermisterized egg under a captive Prairie

I

Falcon which was incubating eggs. The highest temperature I could read was
35 C, which agrees fairly well with Snelling’s data. That would mean that the
other eggs in the clutch may have been reaching on the order of 36.5 to 37.0 C.

OLENDORFF. Has anyone ever placed thermisters on the surface of the egg?

ENDERSON. No. This is the kind of detail we are going to have to do on it.

SNELLING. Figure 1 just shows the incubator I used. This unit was designed
by under $ 1 5.00 and
a poultry scientist at Cornell University. It can be built for
has many could also be used as a brooder for young raptors. You can
uses. It
see that the resistor wires are strung out along the entire length of the sides. It
is made out of a styrofoam ice chest. It is a simple unit to use. It can also be

adapted for battery powered use. While I still have the floor, let me complete
the results of this particular study. The mean time of hatching from the time
which the eggs pipped until they actually emerged was 55.4 hours. This agrees
well with what the Snyders found with their Cooper’s Hawk eggs. Everybody
here who has watched an egg hatch will bear me out in this— one is strongly
tempted to hatch them artificially, to start picking away at eggshells. This is the
worst thing you possibly could do. The natural situation is perhaps two days. I
think you should just be patient and wait. As Jim Enderson has aptly pointed
out, assistance to the young birds at hatching can be disastrous. The embryonic
circulatory system may not be reabsorbed, resulting in massive bleeding, and
unbilical infections.

LAWSON. What is an estimate of the age of the youngest egg you started
with?

SNELLING. The eggs ranged from about two to about 22 days of age when
they were taken from the wild. The grand mean would run around 1 2 days, so
they were about one third incubated when I got them. Statistical tests on the
age of the egg with respect to the temperature treatments showed no signifi-
cance. They were randomly placed in the different treatments.
1972 Breeding Conference— Panel 8, Incubation B21

GRIER. You might point out that Jeff Lincer has incubated a lot of eggs in
captivity using the same technique.

SNELLING. Jeff Lincer is a graduate student at Cornell who has done his re-
search on Kestrel-pesticide relationships. He had a very bad hatch this year. Less
than 50 percent of his eggs hatched. He was using 97 F for his incubation tem-
perature.

ENDERSON. Eggs can be easily turned without actually handling them by

placing them on
a sheet of paper toweling and sliding the paper toweling, the
eggs being held in the rack by wires on either side.

GRAHAM. Talking with poultry people, they indicate to me that the largest
mortality is at about nine to 1 1 days. Eggs obtained later on might be expected
to have a higher hatch rate than natural.

SNELLING. That is very true. However, 1 1 of 1 7 eggs which did not hatch
reached either completely developed embryos in the egg or pipping. I think
that the most critical time was at hatching.

GRAHAM. The poultry people indicate two times of high mortality at nine
to 1 1 and then at 1 8 days in a chicken egg.

HUNTER. With Prairie Falcons, two clutches with which

I cooperated were

hatched at 101 F. One hatched within two days were removed from
after they
the nest in the wild. The other hatched seven days after removal from the nest.
This was still air incubation.

OLENDORFF. From my own experience, I found in two cases, young which

had difficulty hatching were also difficult to raise during the first five days.
They simply died.

SMYLIE. Snelling, did you have a fan in your unit?

SNELLING. No, it was completely a still air unit.

SMYLIE. Could you put one in? Have you had any plans for that?

SNELLING. No, it could be done I suppose.

NELSON. How humid was the environment in your incubator?

SNELLING. I didn’t mention humidity or turning. I did turn the eggs four
times per day at least 1 80 degrees. The humidity I figured I couldn’t control
very well. In a situation where I had cloth wicks projecting out of the pans of
water in the bottom of the incubator I have a rough humidity measurement of
B22 RAPTOR RESEARCH Vol. 6, Supp. B

Figure 1 The still-air incubator employed to incubate American Kestrel eggs.

.

The size of the unit depends on the type of styrofoam ice chest from which the
body is constructed (Snelling, from Schano, 1969).

Abstract of “How to make a still-air incubator” by E. A. Schano.

(4-H Poultry Science Incubator Project, pamphlet 1-8-1 a, 6 pp., rev. 1969.
N.Y. State Coll, of Agric., Cornell Univ., Ithaca, N.Y.)

1. Use styrofoam plastic ice chest (12-16”x20-24”xl2-15”).

2. Cut welded wire 6” wider and 6” longer than inside bottom; cut 3”
squares from comers, fold the sides down, test for loose fit, tape edges to pre-
vent punctures.
3. Place three rings of masking tape inside— 2Vi” from top, 1” above screen,
and half way in between.
1972 Breeding Conference— Panel 8, Incubation B23

4. Install microswitch assembly (see illustration): a place assembly so center

.

hole (1) is ca. 514” down from the top on one end with outlet box (2) on top;
b. with pencil make three holes for two bolts (3) and control bolt (4), insert
bolts from inside, add washers (5), tighten nuts (6) until firm; c. insert control
bolt (4) with locking wing nut (7), screw into either wafer (8); d. insert heating
cable plug (9) into outlet (10) and pilot light (11) into other outlet; e. to check,
plug cord (13) to 1 10V source, pilot light glows when wafer (8) in contact with
micro-switch (14), cable warms, turn control bolt clockwise until click heard
and pilot out, turn control bolt counterclockwise until click heard and pilot
goes on and cable warms;/ disconnect. (Heating cable, thermostat, and therm-
ometer available from Lyon Rural Electric Company Box 30, San Diego, CA
92012; ask for New York Kit and No. 66 Incubator Thermometer; as of July
1971 price including postage and handling was $1 1.55.)

MICRO-SWITCH ASSEMBLY

1. Centerhole
2. Convenience Outlet Box
3. Bolts

4. Temperature Control Bolt

5. Washers
6. Nuts
7. Self-locking Nut
8. Ether Wafer
9. Plug
10. Convenience Outlet
11. Pilot Light

12. Convenience Outlet

13. Lead Cord Plug
14. Micro-Switch

15. Heating Cable

6. Make observation window on top of chest: cut around glass on cover 14”
into styrofoam, cut hole in cover %” inside this cut, and cut out strip 14” thick
and %” wide, insert glass, fasten with tape.
7. Make ventilation holes: on long sides make four holes with a pencil 2”
from top and 4” apart and four holes 3” from bottom and 4” apart.
8. Test: put cake tin (9x14x1 14”) under screen with 14” of warm water, put
thermometer 1 ” above screen, plug in, turn control bolt until pilot on, adjust
to desired temperature. (Abstract— B.E.H.)
B24 RAPTOR RESEARCH Vol. 6, Supp. B

70 percent.

WOLHUTER. You mentioned that the birds which had the most difficulty
hatching were the hardest to raise after hatching. Could this be a natural selec-
tion against weak birds?

OLENDORFF. I think that those which had a difficult time hatching were
simply too exhausted to even accept food and swallow it.

MENG. I noticed with my Peregrine when she laid, in fact she had three eggs

at the time, that the eggs were not in the center. They were around her body.
She was sitting on the sand and her wings were just covering the eggs to keep
the eggs from freezing. It was cold outside. After she had a full clutch, I found
the eggs under her. I was wondering if in the wild the bird could be setting on
the eggs and the eggs could be at the bird’s side under her wings and they could
be cooled that way.

ENDERSON. We have time-lapse photos of Prairie Falcons at the time of egg-

laying in Colorado in 1971 and it is clear that at this' one eyrie the degree of

incubation, i.e. the time spent incubating per day, increases from about ten per-
cent or so after the laying of egg 1, until about 90 percent after the laying of
egg 5. We saw that at dawn and at dusk the frequency of which the bird was on
the nest increased rapidly. Apparently, the birds do sit on the eggs at night even
after the laying of the first egg.

MENG. Could the eggs be under the wings so the temperature would not be
up?

ENDERSON. I don’t know.

PORTER. In our kestrels we found that although the females were sitting on
the eggs, they were not warming the eggs until late in egg-laying. They would
probably keep them from freezing, however.

WHITE. It is important to remember that the genus Falco has brood patches
on the sides, rather than in the middle.

MENG. The eggs were quite far apart, and later when she was really actively
incubating they were together.

HALLIWELL. It would seem to me, just thinking of it from the practical

standpoint that the eggs would be less vulnerable to gross temperature changes
immediately after the laying of the egg, at the moment you have a blastoderm
with one or two cell layers. Just a chilling of an embryo in the early stages will
not cause death. Later on you probably will.
1972 Breeding Conference— Panel 8, Incubation B25

TEMPLE. This is well known to poultry people. In fact, right after laying,
they put them in a refrigerator and hold them until they have a large batch that
they can start incubating at the same time. They store these eggs for up to six
days without a decrease in hatchability.

ENDERSON. I think 40 F is the preferred temperature.

HALLIWELL. The other thing probably is that the falcon gets more mater-
nal the more eggs she has.

RICHARDS. At the end of March, one of my kestrels laid two eggs. About
this time the temperature went down to -12 C. I was worried about the eggs
freezing so I stayed up a whole night and a day to watch to see what happened.
The female would go into the nest box and stay there for a while and then
leave. Every time she left I put a thermometer underneath the eggs. She would
take the egg up to about 16 C, but it wouldn’t get higher than that. I never re-
corded a temperature lower than about 0 C. As soon as several more eggs were
laid the temperature went up to 25 C and higher.

HUMIDITY

OLENDORFF. Humidity another problem. Most people who have tried

is

artificial incubation thus far have not bothered measuring it. I simply kept a
shallow pan the size of the bottom of my incubator filled with water at all
times. Enderson and Mendelsohn in separate experiments used humidity as high
as possible. Kish and Clark (1970) measured it at 86 percent. Stauber (1971)
stated that Fesner actually sprayed a fine mist of water on the eggs he incubated
and had nearly total success when he did so. Although there are some data that
submerging eggs quickly in water decreases hatchability, a fine spray may in-
deed increase hatchability. If we talk about the microclimate of the egg during
incubation, it is certain, according to Wesley Lanyon who knows a great deal
about hand-rearing passerines, that considerable moisture is supplied to the eggs
from the skin of the brood patch. Furthermore, incubating birds have been ob-
served to bathe and almost immediately return to the eggs. During my recent
stay at the American Museum of Natural History I spoke with Jean Delacour
who has artificially raised countless birds of scores of species in France, and to
Wesley Lanyon. Both Delacour and Lanyon spoke of using the highest humidity
possible and spraying or painting the eggs periodically with water. In the Lan-
yons’ article on hand rearing young passerines, they state that “When the eggs
begin to pip, the moisture should be increased. The glass on the chamber door
should show some moisture condensation during the hatch. If necessary, one
can place sponges in the water pans at this time to increase evaporation. . . .

Too little moisture restricts movement within the shell and may make the mem-
branes too dry and tough for the chicks to penetrate.” I note along this line
that incubation attentiveness in Enderson, Temple and Swartz’s study increased
B26 RAPTOR RESEARCH Vol. 6, Supp. B

in the five days preceding hatching. In the still air incubator I used for the

Swainson’s Hawk eggs there were vents for circulation of air and the eggs were
simply laid on 14 inch mesh hardware cloth. The eggs were, for all practical
purposes suspended in air. I would suggest that this may have had a consider-
able drying effect on the eggs in spite of my attempts to keep the humidity
high. Never did water vapor condense on the glass portions of the incubator.
It is possible that the embryos met with the fate that the Lanyons suggest. This

is pure speculation on my part.

TEMPLE. With the red-tail eggs we incubated absolutely as though they were
chicken eggs, we, of course, had a very close measure of the humidity in the in-
cubator. During the major portion of the incubation period, from laying up to
five days before hatch, the humidity was maintained between 75 and 80 per-
cent. During the five days just prior to hatch, after the young had broken into
the air sac, the humidity was raised and held as high as possible, which usually
meant slightly over 90 percent relative humidity. This is standard procedure
with the poultry industry. They have a separate chamber that they call a hatch-
er where it is possible by the use of cloths suspended and soaked in water to
increase the humidity. As Olendorff said, this is very important to prevent dry-
ing of the membranes.

OLENDORFF. We have had a lot of failure of incubation in the very late

stages. Remember Frank Beebe and his work with Peale’s Falcons. Everyone
was questioning humidity at that time several years ago. I feel that that was a
fair analysis of the situation.

HUNTER. Is there any danger of too high a humidity at any stage?

TEMPLE. Actually, I think the poultry people would say no. You cannot
overdo relative humidity in the air. There have been a number of experiments
performed on this dipping of eggs and spraying them.

HUNTER. With goose eggs this is apparently necessary.

TEMPLE. That goose egg experiment was done again, and what they found
was that they increased infection of the embryo by moistening the surface of
the egg. Getting the surface of the egg moist allowed bacteria to enter the pores
of the egg and cause infection in the developing embryo. So, probably it is not
a good idea to get the surface of the egg moist. There is a different situation
with waterfowl where, as with a goose, every time the bird gets on the egg she
brings water.

HUNTER. Ducks hatch so easily that if you keep them moist and put them
under the register behind the stove they’ll hatch.

NELSON. There is some good information for Ospreys as well. They actually
1972 Breeding Conference-Panel 8, Incubation B27

bring water. Welty cites some literature example concerning this. Has anyone
actually seen with raptor incubations that wetting eggs will cause infection?

TEMPLE. This was a general sort of thing with many poultry eggs. I might
add that with the Ospreys there was some artificial incubation done this last
summer. I think the sample size was somewhere around one dozen eggs. The
hatchability was as good as you would expect from the wild. The incubation
was done once again precisely as with chickens. I think seven out of the 12
hatched.

RICHARDS. I would like to comment on something that impressed me in

reading about Lanner Falcons in the Sahara. It said that for 100,000 square
miles or something like that in any direction there was hardly any vegetation let
alone a water hole. Still they are nesting out there. It would be impossible for
them to get a drink, let alone bring back water on their feathers.

TEMPLE. Just an observation on those nesting Peregrines, there weren’t very

many times during incubation when the falcon came back to the eggs appearing
to be wet like she had been in the water. This was not true during the brooding
period. During brooding the falcon would often come in obviously just return-
ing from a bath.

ENDERSON. I wouldn’t say it was very often even during brooding.

WHITE. One point that is fairly clear with nearly all types of birds, for any-
one who has done anything with passerines, that under the brood patches birds
have a very jelly-like area. The brood patch is a highly vascular area with a lot
of moisture and lipid there. This does provide moisture through the skin to the
egg.

PORTER. I noticed in the American Kestrel just after the eggs were laid,
they had a rather rough surface. As incubation started, the eggs developed a
glossy, almost waxy-like, texture on the surface. This was probably involved
with this brood patch.

WHITE. The brood patches are a very important physiological phenomenon

where the blood vessels engorge just below the skin.

OLENDORFF. I wonder if it wouldn’t be a good index for high humidity to

use the technique of Lanyon of observing water condensing on a piece of glass
on the door of the incubator.

TEMPLE. That’s of course, just a function of the gradient between the incu-
bation temperature and the outside temperature.

OLENDORFF. Yes, but if the incubators are in rooms near room tempera-
B28 RAPTOR RESEARCH Vol. 6, Supp. B

ture and the incubation temperatures are roughly 99 F, condensation would

occur at nearly the same relative humidity each time.

GOBEN. In 1965 I raised two prairies. For what it is worth, twice a day for
an hour, I would take a wet cloth and wrap the two eggs. I always had a pan of
water in the incubator. I also had problems of the chick sticking to the mem-
brane at hatching and I took a dropper with water and moistened them.

GRAHAM. In Germany a very successful hatcher has a heating element in his

water. Five days prior to hatching he turns it on. By heating the water you get
more evaporation and a higher relative humidity.

ENDERSON. I used a pump driven atomizer to obtain 80 percent relative

humidity in an incubator in Colorado Springs. In our climate 70 percent humid-
ity is no small chore, since the humidity of the outside air is about 1 5 percent.
Another possibility which occurred to me was to use an aquarium pump and air
stone submerged in the water pan at the bottom. You will have trouble getting
too much humidity.

SWARTZ. There are available very convenient electronic hygrometers.

ENDERSON. We had one hooked into the circuit for the atomizer so when
the humidity fell below 80 percent, the pump came on.

INDETERMINA TE EGG-LA YING

OLENDORFF. Egg-laying has not presented major difficulties except where

females have become egg bound. We have spoken to this point earlier. Just to
point out a few facts about egg-laying— it is a fairly widespread characteristic
that falconiforms lay their eggs every 48 to 60 hours on the average. Variation
does exist and it is not uncommon for 72 hours to pass between eggs. In the
wild, birds of prey desert most easily at egg-laying, a point to remember when
disturbing your birds. One interesting aspect of egg-laying is whether falcons
and hawks are determinate or indeterminate layers. Determinate layers produce
a fairly constant number of eggs (say four in the case of most shore birds) and
then stop laying— period. Indeterminate layers will continue to lay eggs as their
clutches are removed one by one at the time of laying. Ratcliffe (1963) states
that Peregrines do not usually continue to lay if an incomplete set of eggs is
taken. Davenport ( 1 900) was able to collect as many as 1 6 European Sparrow-
hawk eggs from a single nest by removing each one soon after it was laid every
other day for a month. Dr. Porter has some evidence that this is also the case
with American Kestrels and I would like to give him the opportunity to expand
the comments he made yesterday about indeterminate laying.

PORTER. I think that Green, who studied the Peregrines in the Aleutians
1972 Breeding Conference— Panel 8, Incubation B29

early in the century removed eggs and got renesting in very short periods of
time. would suspect that the Peregrine follows the same pattern as the kestrel.
I

I ran an experiment using four single female kestrels from which I removed each

egg as it was laid. I used four other females, but I left one egg in the nest at all
times. Three other females I let incubate normally. The latter females laid the
usual five-egg clutch and incubated them consistently even beyond the normal
incubation period. Those from which I removed eggs, but left one egg in the
nest, one female laid eight eggs at the normal two-day interval and then had a
gap of 1 2 days before laying five more eggs. After a gap of 1 1 days, five more
eggs followed; then, after 14 more days without laying she laid four more eggs.
She laid 22 eggs in all. Another female which was always left one egg laid 23
eggs with the greatest interval of only four days between eggs. A third one laid
1 8 eggs with the greatest interval being six days. A fourth female laid eight eggs

with the greatest interval of four days. This suggests that if they are indetermin-
ate, they are just now evolving the capability. There is some variation within the
species. Those from which I removed all eggs as they were laid, one female laid
only three eggs and laid them very late. Another laid 13 eggs with the largest
interval between eggs being five days. Another laid 26 eggs with the largest in-
terval being four days. The fourth female laid 1 7 eggs with the largest interval
being five days. A five- or six-day interval is probably not a recycling interval.
The latter ranges from nine to 17 days in the kestrel. The following year I
placed four eggs in a kestrel nest after the female had laid her first egg. She con-
tinued laying until she had laid a total of four eggs. This was a second nesting
for the female for the season. Very frequently, on the second nesting, one fewer
(four) eggs are laid. I feel that the four-egg clutch was complete. Thus, she laid
a full clutch despite the fact that additional eggs were added to her nest. This
supports the hypothesis that the American Kestrel is an indeterminate layer.

OLENDORFF. It makes us wonder if there really is much difference between

the concept of indeterminate laying and recycling. In the case of your kestrels
there is represented some middle ground between the two concepts.

THOMAS. What happened with regard to fertility in these cases?

OLENDORFF. Davenport (1900) did not mention fertility. The point is that
with artificial insemination one could fertilize the excess eggs.

PORTER. All of the birds I used were single females.

GRIER. The ones at Cornell last summer were almost all fertile. This involv-
ed 23 and 26 eggs from different pairs of kestrels.

HUNTER. Did you have any problems with calcium, Dr. Porter? Was there
any dietary supplementation? Was there any thinning of the eggshells?

PORTER. We didn’t get any appreciable differences in texture on eggshells

B30 RAPTOR RESEARCH Vol. 6, Supp. B

until we reached about the 19th egg in 1967. The thickness of the shells re-
mained fairly constant,while the texture began showing signs of deterioration.
In 1 970, even the 26th egg appeared to be in relatively good condition.

OLENDORFF. We don’t have to be that greedy. If we just got two or three

more Peregrine eggs—.

HUNTER. Did you do anything to the diet?

PORTER. No. We just gave them our normal kestrel diet.

OLENDORFF. It seems that if you did want to get more eggs by these meth-
ods, you would supplement the diet with calcium.

HALLIWELL. The calcium which goes into the eggshell indirectly comes
from the diet, but the bird must take it in in the correct ratio. It must be ab-
sorbed and then stored in the bone. It is withdrawn from the bone back into
the serum and then goes to the shell-forming portions of. the body. Feeding
more calcium during the laying period is not sufficient. You must have a build-
up of calcium previous to egg-laying.

ENDERSON. In 1971 my
nine year old Peregrine laid a clutch of four eggs.
I removed them and she then started on a clutch which eventually yielded 13
eggs. I left a single egg in the nest at all times. This went on until the 11th egg
at which time I took the bird out of the room and blocked her down. Two
more eggs were laid while she was on the block. Both of these had ultra-thin,
soft shells; the last one was unpigmented.

OLENDORFF. The Peregrines, then, seem to be indeterminate layers.

GOBEN. With my Ferruginous Hawks this year, the seventh egg was obvious-
ly lacking. The color was completely abnormal. It was odd-shaped and broke
the second day.

BERRY. would appear that at least one female Goshawk is not an indeter-
It
minate layer. For five separate years she was allowed to retain only one egg in
every clutch which she incubated. In several of these years all eggs were taken
and she failed to recycle or lay additional eggs.

OLENDORFF. We are all sorry to hear that.

BERRY. There was also appreciable shell thinning in the three eggs she laid
in 1971 from the first to the last shell. The last shell seems to be only about
half as thick as the first shell. She was vitamin supplemented for at least two
months prior to egg-laying with calcium glutonate about three times a week.
1972 Breeding Conference— Panel 8, Incubation B31

RECYCLING

OLENDORFF. It is well known from egg collecting days that many raptors
have the capability of completing a successful nesting even if their first clutch
of eggs is taken. Bond (1946) mentions two observers taking 12 fertile Peregrine
eggs from the same eyrie in the s^me year (three clutches of four) and a fourth
clutch of undetermined number was laid. The potential for recycling seems to
be related to the time invested in any given set of eggs. If young are destroyed,
birds will rarely, if ever, renest. An interesting question for study would be to
figure out how long you can let the parents incubate before recycling is cur-
tailed. The reasoning here is that we have had trouble with full-termed incuba-
tion, but relatively less with short-termed incubation. The Lanyons and avicul-
turists in general have little or no luck incubating passerine eggs full term. The
possibility remains, however, that we could allow captives to incubate for two
weeks, then recycle and increase our chance of successful artificial incubation
by taking advantage of natural incubation of the parents at first. Perhaps we
will be able to leave them under the parents until candling is feasible and then
to recycle the parents.

ANDERSEN. We have been working with Swainson’s Hawks for about three
years now. In any case, if the nest is destroyed during the first week of incuba-
tion, there will invariably be a renesting. We have yet to find any case where a
successful renest has been made after two weeks of incubation. We see nests re-
lined,but never relaying. If we use this approach, probably no more than one
week of incubation by the parents will be possible with the buteos.

NELSON. In the wild Peregrines this year we had two which lost young
nests
and one of the week after it hatched.
pairs lost their youngster within the first
They slipped right back into courtship behaviors we would expect prior to egg-
laying. They did not lay eggs, however. I mention the point because we may
think that if we get the early courtship activities, we will get eggs. If this hap-
pens during a recycling procedure after one clutch of eggs hatches, there is not
much hope for a second clutch.

HUNTER. I did not take the eggs away from my Peregrine until the 13th
day. They went through all of the courtship again, but no eggs were laid.

GOBEN. Boyd had three clutches of Prairie Falcon

Lester eggs from a single
pair and he removed eggs on the 14th day each time.

PORTER. We have some instances with the kestrels in which the female laid
the egg of her second clutch before the young of the first clutch had fledg-
first

ed or left the nest box. Apparently, the recycling was initiated at some earlier
time even when the nestlings were younger, probably 14 days prior to fledging.

WOLHUTER. I question the fact that birds that tend to nest early in the sea-
B32 RAPTOR RESEARCH Vol. 6, Supp. B

son or late in the season have smaller clutches. I wonder if the time of year we
are trying to get the birds to recycle is important. If it gets to be too late in the
summer when you try to recycle, are you missing your whole chance? Should
you start the whole process early so you have more time later in the year for a
second clutch?

PORTER. Our data on removing eggs suggest that the later the birds start
laying, the fewer eggs will be laid in a sequence.

GRIER. Part of this might be tied in to photoperiod considerations. Later

on, day length is starting to drop off. It might be possible to keep things going
by maintaining a long photoperiod.

GALICZ. I have one pair of Peregrines that laid in late March and incubated

for 39 days. Fourteen days later she began laying a clutch of three eggs. Again,
after a second full incubation period on August 18 she began a clutch of three
eggs. I did not use artificial light.

PORTER. As you found with Peregrines, we also found with kestrels that
about 12 to 13 days was the average period for recycling, from the time the
first clutch was removed until the first egg of the second clutch was laid. We
found also that the period from the time that they were paired and placed into
pens with a nest hole to the time it took to stimulate reproduction was about
1 2 to 13 days. This is in press in The Condor at this time.

HUNTER. How many of the kestrels which successfully raised young recycl-
ed after they raised young?

PORTER. I don’t have those data. The percentage was rather low.

HUNTER. Does anyone know if that is common in the wild?

ANDERSEN. We some work on clutch size of Swainson’s Hawks two

did
years ago and looked at replacement clutches. The replacement clutches were
the same size as the original clutch in every case. As far as raising them, we get
a much lower success ratio of second clutches.

WOLHUTER. I your southern birds that would have a higher inci-

think it is

dence of recycling with kestrels. I think it would be interesting to take

at least
eyass birds from very far south and put them in a breeding project and see if
there is a higher tendency for them to recycle.

WHITE. There is one literature citation in the publication of the London

Academy of Sciences where it said that Peregrines in Africa near the equator
do lay twice a year. We need to substantiate this statement.
1972 Breeding Conference— Panel 8, Incubation B33

TEMPLE. In arctic Peregrines, this year on the Colville Riverwe discovered

for the first time an arctic Peregrine with a replacement clutch. The bird broke
her first set of eggs, and laid a second clutch.

PORTER. How far along were those birds in incubation?

TEMPLE. We could not tell because the eggs were broken when we got there.

UNKNOWN. What happened to the second clutch?

TEMPLE. The second clutch hatched and the young were dead when we
made our second float.

BERRY. In the Ungava region of northern QuebecI took four Peregrine eggs

this year and artificiallyincubated them. They were taken at the mid-point of
incubation. Two of the four eggs hatched. I went back a month later to see if
those birds had recycled, and while both adult birds were on the actual eyrie
ledge there was no indication of a recycling. At least a recycling attempt did
not result in eggs being laid.

CANDLING

OLENDORFF. I realize that candling is a very simple procedure. I’ve never

done it and I’m sure many of you haven’t. Perhaps Jim Enderson would tell us
a little about it.

ENDERSON. My experience with candling is limited to the eggs of Prairie

Falcons and Peregrines. Only the Prairie Falcon eggs contained embryos. The
presence of an embryo can be determined using a standard commercial candler,
or a two-lb coffee can with a 40-watt bulb inside and with a small hole about
the size of a quarter in the lid. By rotating the egg after four days of incubation,
that is four days after the completion of the clutch, and waiting for the yolk to
come up, you can see behind the pigment layer a distinct shadow cast on the
inside surface of the shell. You must be careful not to overheat the egg, because
with the coffee can device a great deal of heat is generated. I turn the light off
between candlings. Bob Berry has mentioned that the problems of candling
Goshawk eggs are not the same as with falcons. As the embryo gets bigger, after
about the 24th or 25th day, it is nearly opaque and very difficult to see motion.
The easiest thing in that case is to turn the egg so the air sac end is down. As
the light comes up along the side of the air sac, you may see motion where the
embryo pushes against the air sac and casts a shadow on the side of the egg. By
about 32 days the air sac enlarges rapidly and you must avoid the impression
that the embryo is dead and that the egg is suddenly drying out. On about the
34th day the embryo will break the air sac and motion is again easy to see. Pip-
ping in Prairie Falcons usually takes place around the 34th or 35th day. At
B34 RAPTOR RESEARCH Vol. 6, Supp. B

least two more days are required before hatching occurs.

EBERLY. How large is the embryo at 1 0 to 15 days?

ENDERSON. By about the 10th day it is still under one cm long.

BERRY. At six to seven days in Goshawk eggs you can see a pea-sized dark
spot. You can see the blood vessels around that spot. The circumference of the
area of blood vessels might be the size of a quarter.

ENDERSON. Especially with the pale Prairie Falcon eggs, the vessel system
is obvious by two weeks.

TEMPLE. People have been worried about the possible deleterious effects of
candling. The poultry literature suggests that as long as candling is done quickly
and you don’t expose the egg to the heat of the candler, candling has no adverse
effect. I candled the red-tail egg I hatched sometimes twice a day every day.

PORTER. At times when would open the nest boxes of the kestrels the
I

bright sunlight would had the same thing occur with

hit the eggs directly. I’ve
Prairie Falcons in the wild. Does anyone know if this has deleterious effects on
them?

FYFE. One time we set a blind up and the eggs were exposed to direct sun
for quite a while. The outside temperature was between 50 and 60 F. The bird
was off the nest for an hour and 20 minutes. All eggs hatched.

HUNTER. I use a slide projector for candling. You just use your hand. In
seven days you see blood vessels all over the inside of the egg. In my limited
experience, if you can see anything inside the egg at seven days, it is probably
fertile.

NELSON. In the wild Peregrine on the coast where the

air temperature al-
most never gets over 68 to 70 F, very obvious, possibly without exception,
it is

that the female incubates during the heat of the day. She just will not let the
male come on until the eyrie gets into the shadow. Are there changeovers in the
taiga Peregrines that will allow the sun to hit the eggs?

TEMPLE. At one which had a very sunny exposure, there frequently

eyrie
occurred over-heating of the young and an adult was often present to shade the
young. I do not recall about changeovers during incubation, but they changed
about every three hours, so there must have been changeovers during the sunny
part of the day.

NELSON. One thing that alarmed me one day when I went to an eyrie the
day that one of the eggs hatched (the adult female was off the ledge for less
1972 Breeding Conference— Panel 8, Incubation B35

than a minute by the time I got to the scrape) the young bird was on its side
thrashing and squirming. As soon as I shaded was perfectly all right. You
it, it

would think it would lie motionless. I think some of the reaction was to sun
light.

TEMPLE. We will show exactly that in some of our films for the next panel.

OLENDORFF. Let’s use that as a lead in for the next panel.

REFERENCES
Bellairs, R. 1960. Development of birds. In: Biology and Comparative Physiol-
ogy of Birds, A. J. Marshall (ed.). New York: Academic Press, pp. 127-
188.
Bond, R. M. 1946. Peregrine population of western North America. Condor
48:101-116, 1946.
Davenport, H. S. 1900. Nesting habits of the Sparrow-Hawk. Zoologist 1900:
426-428.
Enderson, J. H. 1971. Account of Prairie Falcons bred in captivity— 1970. Rap-
tor Res, News 5:44-47.
Hancock, D. 1971. B.P.I.E. No. 23. Raptor Res. News 5:24-26.
Huggins, R. A. 1941. Egg temperature of wild birds under natural conditions.
Ecology 22:148-157.
Hunter, D. V. 1970. B.P.I.E. No. 16. Raptor Res News 4: 1 1 5-1 19. .

Kish, F., and G. K. Clarke. 1971. B.P.I.E. No. 24. Raptor Res. News 5:26-27.
Lawson, P. T., and E. L. Kittle. 1970. Artificial incubation [of American Kes-
trel eggs] Raptor Res. News 4: 136-137.
.

Ratcliffe, D. A. 1963. Status of the peregrine in Great Britain. Bird Study 10:
56-90.
Schano, E. A. 1969. How to make a still-air incubator. N. Y. State Coll, of
Agric., Cornell Univ., Ithaca, NY, 4-H Poultry Science Incubation Project,
Pamphlet 1-8- la, reb. 6 pp.
Stanley, A. and E. Witschi. 1940. Germ cell migration in relation to asym-
J.,

metry in the sex glands of hawks. Anat. Rec. 76:329-342.

Stauber, E. 1971. Reproduction of birds of prey in captivity. Hawk Chalk 10
(l):35-37.
RAPTOR RESEARCH
Volume 6 Supplement Part C 1 972

SPECIAL CONFERENCE ON CAPTIVITY

BREEDING OF RAPTORS-A REPORT

Sponsored by Raptor Research Foundation, Inc.

Held at Sioux Falls, South Dakota, U.S.A., November 22-23, 1971.

Part C. Behavioral Considerations to Egg-laying (Panel 6)

edited by
R. Wayne Nelson
Department of Biology
University of Calgary
Calgary 44, Alberta, Canada

Panel Members: R. Wayne Nelson, Chairman, Richard W. Fyfe; Frances Hamer-

strom, and Richard R. Olendorff. A paper submitted by Noel and Helen Snyder
was read; unfortunately they were unable to attend the conference.

INTRODUCTION by R. Wayne Nelson

By considering the behavior of captive birds of prey prior to and during the
time when eggs might be laid, it should be possible to‘ learn much which could
be applied toward the goal of achieving reliable production of captive-bred off-
spring. One of the major problems confronting captivity breeding is the acquisi-
tion of fertile eggs. In the many instances in which no eggs, or only infertile
eggs have been laid, it could be presumed that at some point in the courtship
behavior or sexual development one or both members of the captive pair had
been blocked. Observations of the behavior of the captive birds should indicate
where some of the problems lie.
This panel attempted to outline the courtship behavior of some wild and cap-
tive raptors, and discussed various methods whereby the captive birds might
better be induced to take on the behavior and condition necessary for reproduc-
tion. The emphasis on the larger falcons reflects the urgency of the problem
with those species; the information from other species provides very useful sug-
gestions for captivity breeding of the larger falcons, due to similarities which
appear within many of the species, and due to techniques which have been
worked out with the other species.

C37 Raptor Research 6(Supp.) C37-C68, 1972

C38 RAPTOR RESEARCH Vol. 6, Supp. C

Unfortunately, we must (literally or figuratively) leave the owls in the dark.

There is a general lack of knowledge concerning much of the behavior of the
owls prior to egg-laying.
The panel also attempted to indicate what some of the behavioral problems
are, and how they might be avoided. The aim of the panel was to stimulate and
encourage a better appreciation of what it takes to get birds of prey to behave
appropriately in captivity, so that their behavior will bring them to reproduce
successfully.

THE NEED FOR A CHANGE IN DIRECTION- TOWARD THE SAME GOAL

by Richard Olendorff

Introduction. Before getting started on the topic of behavior I would like to

say that the other people on this panel, and certainly manyof you in the audi-
ence, have spent more time contemplating the behavioral aspects of the captiv-
ity breeding of raptors than I. I suppose I am here because I wrote a lengthy
review article on falconiform reproduction with emphasis on behavior, and car-
ried out a very small captivity breeding project with one pair of American Kes-
trels during which rather detailed behavioral observations were made. Never-
theless, I consider my knowledge of falconiform behavior rather superficial and
propose only to speak in generalities for the next several minutes to set the
stage for the other panel members. In doing so, I would like to combine some
aspects of the review article with some of the results of the kestrel breeding
experiment.

Importance of Behavior. First of all let me point out that animal reproduc-
tion has often been researched with too little emphasis on ethology or behav-
ior. Falconiform behavior must be important to us, but there is danger, too, in
holding to stringent paradigms in the name of esthetics or ethology. So we must
strike a balance which will result in production, a balance which may involve
considerable artificialization (if I may coin a word) of the breeding sequence
and conditions in general.
I used to think that habitat simulation and forced fulfillment of complete

repertoires of reproductive behavior were the answers to all of our problems. I

used habitat simulation with my kestrels with some success. As it turns out,
though, this technique works very effectively for kestrels, but it is neither an
effective nor an efficient method of rearing large falcons. This follows directly
from the fact that copulation-producing behavior is an immensely complex se-
quence of events. Thus, we should glean only what is necessary from wild fal-
con behavior, and environment for that matter, emphasizing as we go those
aspects which will increase the production of captive birds.

Introducing the Birds to Their Mates. Introduction of males and females has
been discussed at great length. Although introduction is probably a problem in
1972 Breeding Conference— Behavior C39

itself, it is We have two types of pairing

closely related to pairing in general.
situations to consider: (1) the male with the female, and (2) the male and fe-
male with man in the artificial case. In raptors, as with any species of bird, pair-
ing requires a great deal of adjustment, probably more than usual with the pug-
nacious raptorial species.
The worst failure is the situation in which the female will not tolerate a mate
in the same room, the problem being introduction.

Pair-bonding. Assuming that the birds are successfully introduced, the next
behavioral requirement is and strengthening of the pair-bond. Of al-
initiation
most equal importance hormonal considerations discussed earlier at this
to the
conference are all forms of courtship such as mutual roosting, cooperative hunt-
ing, courtship flights, courtship feeding, and nest scraping (or building) to men-
tion but a few. I think we can skip the matter of induction of physiological
which has been substantially
readiness, (albeit inconclusively) discussed already,
and move on to eliciting copulation.

Toward Copulation. Pairing and copulation-producing behavior have one very

important thing in common; they involve sequentially presented stimuli which
result in a sequence of responses. If the sequence is interrupted, reproduction
may be inhibited through the neuroendocrine system. If the sequence develops
adequately, reproductive processes will continue.
Let’s consider the components of the sequence leading to copulation.

Mutual Roosting. Cade observed it in wild arctic Peregrines; Nelson has not
observed it in wild Peale’s Falcons. My kestrels engaged in it by the hours. In
any of the behavioral sequence does not pose much of a problem.
case, this part
The opportunity for mutual roosting exists with the simulated habitat approach;
it is probably unnecessary if artificial insemination is to be used.

Courtship Feeding should not be a problem provided that (1) the food is not
tied to a stump or board, so that the male can take it to the female, and (2) the
behavior is allowed to develop by the female. By the latter I mean that the fe-
male be kept adequately fed and/or be previously handled such as to prevent
her from immediately stealing the food from the male. One such oversight could
stop the sequence. Courtship feeding seemed very important with my kestrels,
and I invite you to read about it in the article in Raptor Research News. Note
particularly its relationship to the pair bond and its copulation-producing con-
tributions.
Many other familiarities seen in the wild are also permitted and have been
seen in captive situations; again, in the case of wholly artificial means they
appear to be unnecessary. Billing, vocalizations, mutual preening, bowing, nib-
bling of the feet, and wing fluttering do not present major blocks to the chain
of behavior patterns leading to copulation, since they are readily allowed in
confinement.
With cooperative hunting excursions and courtship display flights we run into
C40 RAPTOR RESEARCH Vol. 6, Supp. C

problems. They are virtually out of the question in the captive situations. Dis-
play flights are important. This can be seen from the facts that (1) courtship
flights in many birds are so species- or genus-specific that they are being used in
systematic studies of closely related bird groups, (2) birds of prey fall into this
category, having quite distinctive courtship flights, and (3) in many cases, as
with my kestrels, courtship flight behavior patterns are very stereotyped, a pro-
perty which is shared by most instinctive behavior patterns. Courtship is instinc-
tive in its basic pattern; and one instinctive behavior often acts as a stimulus for
another instinctive behavior.
Assuming that your falcons paired reasonably well, it is possible that inhibi-
tion of courtship flights was the pitfall of your project. The external environ-
ment was the problem. Suffice it to say that the elicitation of instinctive, se-
quential behavior patterns will always be a problem, even at the outset of the
wholly artificial paradigm.
Jn spite of the pitfalls and the failures, there have been some successes. In
cases where birds have produced fertile eggs, they were able to abbrevi-
at least
ate their courtship displays. They most certainly performed displays, neverthe-
less, but adequate courtship is not happening often enough with large falcons.

The Alternativeto ‘Natural’ Captivity Breeding. The bright spot among the
failures the increased interest in artificial means. The disappointment of hav-
is

ing a six year old falcon that will attack an old tiercel on sight certainly has led
many to ponder the thought of bypassing pairing, and bypassing the territorial
aspects of falconiform behavior. This leads directly to the question, “What be-
havior patterns, natural or aberrant, should be stressed if we forget about pair-
ing bird to bird, and pair birds to man?”
The falconers among us realize the great extent to which falconiform behav-
ior is plastic. We have seen the whole gamut from screamer to seasoned game
hawk. As our discussion continues, we should reflect on how individual birds
turned out, and how their behavior patterns might be used in captivity breed-
ing. Realize, though, that behavior patterns which are undesirable in a bird used
for falconry may be very desirable in a captivity breeding program.
As a young falconer I was taught that the only behavior patterns required of
a game hawk which were not part of her natural repertoire were (1) for the bird
to come down to you out of a tree or the sky, and (2) for her to let you ap-
proach her on a kill. Anything more was gravy, or simply unnecessary.
If we reduce captivity breeding to its simplest terms, the only unnatural be-
havior patterns required are (1) for the male to accept a man as a mate or at
least as a sexual stimulus, and (2) for the female to lay eggs in a captive situa-
tion. As we have seen today, it’s not that easy, but there is reason for optimism
in both of these situations. Couple (1) and (2) with artificial insemination, and
possibly also with artificial incubation and hand- or foster-rearing, and we have
a plan of some importance developing.

Summarizing. In summing up, and to emphasize the importance of artificial

means, I would like to make the following points.

1972 Breeding Conference— Behavior C41

Simulation of natural nesting conditions in breeding lofts has until now occu-
pied the efforts of many raptor aviculturists. This method has already proved
too inefficient to keep pace with declines in wild populations and, as a result of
the latter, in the numbers of birds available for captivity breeding and falconry.
Unless some breakthrough occurs, we need an alternative to habitat simulation
and forced breeding behavior.
Man has been augmenting the productivity of domestic birds for decades by
bypassing the necessity of developing complex, copulation-producing behavior
patterns. In light of recent successes with artificial insemination with raptors,
and assuming that there are far more similarities between falcons and domestic
birds than differences, we should place emphasis on behavior patterns which
will increase the production of progeny by artificial means. It is irrelevant whe-
ther or not these behavior patterns are “typical” of falconiform birds. We
should take advantage wherever practical of known behavior patterns of wild
birds,but we should not be bound to them in the name of ethology or esthetics.
Contrary to some purists’ beliefs, two captive falcons will not a chicken
make, either morphologically, physiologically, or psychologically. Such prob-
lems of the progeny will be solved when the time comes. Right now we need to
produce— artificially, if we can, at least for the time being.

TRIGGERS FOR EGG-LA YING

by Frances Hamerstrom

Sean Morris, the brilliant young British biologist, recently told me of his
work filming Blue Tits. As he spoke, one thing after another that I had noticed
with raptors fell into place: for example, triggers that lead to egg-laying.
The Blue Tit must time its egg-laying so that caterpillars will be available to
feed the young. There are early springs and late springs so she cannot use day-
length as a guide. She prepares her nest early in the season and then loiters.
When the male brings caterpillars to her in large enough quantities she starts
laying— having started, she completes the clutch.
Balfour has stressed the close connection between the sex act and food in
harriers. In my experience, harriers only breed well when mice are abundant,
and furthermore it is my hunch that food is more important than aerial displays
in harriers. Recently harriers have been breeding successfully without sky danc-
ing in good vole years. There was one exception: many voles and almost no
breeding. I suspect it was the year that they were carrying the highest pesticide
residues.
For captive breeding, it appears we do not need all the triggers, but we do
want to time them. Presentation of food is the easiest of all to time.
Falconers are accustomed to offering food once a day. I can think of few
things less likely to trigger egg-laying. Notice I am not talking about availability
of food, but of presentation of food. A number of species of raptors lay small
clutches or fail to breed when food is scarce. When food is scarce it is less often
C42 RAPTOR RESEARCH Vol. 6, Supp. C

presented.
Essentially all falconers understand the advantages of tid-bitting in training.
Holding half a Plymouth Rock hen on the glove gives at best one flight a day. I
am coming back to that Blue Tit female— repulsive as it may be to some to learn
from dickey-birders— the Blue Tit lays when caterpillars are presented often
enough. Let’s give our breeding birds the idea that hunting is good when we
want eggs. Tid-bit.
Furthermore, a trigger may lose its effectiveness if it is used constantly. I do
not believe that my Golden Eagle, Chrys, would have laid six clutches in five
years if she had not had frequent nest visits from me just before laying. She did
not get this kind of attention until I figured egg-laying was near. Like the Blue
Tit male, I spent several weeks in lack-a-daisical attention and then pulled out
all the stops at once: frequent visits, nest building, neck stroking and using my

voice. The most powerful response that she gave was to hay for nest lining. If
hay was withheld and then dumped on her nest, she went straight into a copula-
tory reflex— possibly a releaser for ovulation.
Birds that have stick nests are fascinated by sticks. If you work with them
with sticks and are getting along well with them with sticks, tease them— don’t
always give them the sticks, hang onto the sticks, make it a little difficult for
the birds.
As falcons and owls do not build their own nests, they are not apt to be
“turned on” by hay or sticks. Old pigeon wings and such may stimulate sex play
and ovulation in these groups.
Heinz Meng, when he mentioned how he crouched and ran along outside his
falcons’ breeding room, immediately struck a bell as far as I was concerned—
think we need to vary things. Which of our breeding birds have a chance to de-
fend their nest? We spare them this as hard as we can. We shy away the public,
frighten away the dogs, and even hassle the children. Now, if you let the birds
get scared sometimes, maybe it does them good as long as they don’t get too
scared. Consider this possibility, because then they are defending a nest, which
is the natural thing for them to do from time to time.

Many of us are inclined to keep trying everything that we can think of to

produce eggs. Perhaps we need to learn to deny some of the stimuli for a time.
They may work a week later.
It has become plain at this conference that raptors are not strongly deter-
minate layers. To get more eggs— once you have some— two courses are open:
keep taking away eggs (I’d leave one in the nest), or take the whole clutch and
hope the bird will lay another.

References

Balfour, E. 1962-1963. The hen harrier in Orkney. Bird Notes 30(5): 145-1 53.
Hamerstrom, Frances. 1970. An Eagle to the Sky. Iowa State Univ. Press, Ames,
Iowa. 143 p.

(Fran’s book devotes almost half of its pages to her efforts toward captivity
1972 Breeding Conference— Behavior C43

breeding of her Golden Eagles. During the last several years, she and her eagles
have come extremely close to producing captive-bred eaglets by the artificial
insemination method. For a fascinating account of the ideas, methods, and tech-
niques surrounding the courtship and artificial insemination of these birds of
prey, I would strongly recommend that everyone interested in captivity breed-
ing carefully read this book. And, by the way, it is in its third printing. R.W.N.)

BREEDING BEHA VIOR OF CAPTIVE AND WILD PRAIRIE AND

PEREGRINE FALCONS
by Richard Fyfe

Introduction. One aspect of the raptor studies being carried out by the Toxic
Chemicals Section of the Canadian Wildlife Service has been field studies of fal-
con behavior in relation to our pesticide investigations. These studies have been
oriented toward documenting normal behavior patterns so that we would be in
a position both to identify and interpret behavioral changes observed in the
field. Detailed observations have been made on several pairs of Prairie Falcons
(Falco mexicanus ) which were known to be carrying very low residues of toxic
chemicals and on one pair of anatum Peregrines {F. peregrinus anatum ) in 1969
and 1970.
In addition to our field work I should indicate that for several years I have
tried unsuccessfully to breed both tundra {F. p. tundrius ) and Peale’s Pere-
grines {F. p. pealei) in captivity. The pair of pealei laid two clutches of infertile
eggs in 1970 and again in 1971. Obviously, since the eggs were not fertilized,
something was wrong, but what? Since we had observed that our presence al-
tered the behavior of the birds we felt that it was not possible to observe nor-
mal behavior so long as the birds were aware of our presence. Therefore, in
1971 we built two observation blinds and were able to photograph and carry
out detailed observations on the pair of pealei and on one pair of Prairie Fal-
cons. Fortunately for us the latter pair were successful in producing four fertile
eggs, hatching these and raising all four young to fledging.
For this report I have gone through our field notes and have tried to list the
main behavior patterns for the Prairie Falcon in the wild, and have compared
these with what we observed in the captive Prairies up to the time of egg-laying.
I have done the same for the Peregrines, but since we did not have as much in-

formation on this species I have also relied on descriptions in “Der Wander-

falk” by Fischer (1968).
The behavior patterns described below and in Table 1 are listed in the order
in which they were observed or described; however, each pattern does not
necessarily follow in this exact sequence. In some instances several aspects may
be ongoing at any one time with variation in intensity depending on the stage
of the nesting cycle; for example, courtship feeding, nest display, mating, and
male and female nest scraping may all be observed during the same day or series
of days.
C44 RAPTOR RESEARCH Vol. 6, Supp. C

The observations described were made by Bob Gibbon, Keith Hodson, Tom
Donald and myself, the majority of the observations of the captive birds being
carried out by Tom Donald and of the wild Prairie Falcons by myself.

Initial Indications of Breeding. Initially, with both the captive Prairie Falcons
and Peregrines, our indication that something was going to happen was
first
“caching”, or territorial behavior, by the male in the case of the Peregrines, by
the female in the case of the Prairie Falcons. This occurred from a month and a
half totwo months before anything else and was the only early indication that
something was happening. In the case of the Prairies, about one month later we
heard our “chupping” (February 26). (We do not describe it as “eechip-
first
ping”; not the “eechip” of the Peregrine, but rather a “chup”, an audibly
it is

different call.) By the middle of the next month everything had gone complete-
ly quiet with the Prairies, and as far as we could tell the birds were not paying
any attention to one another. Then, very suddenly, we heard the chupping again
(April 3), and we then saw what I describe as preliminary pair bond behavior.

Table 1 . Observed Courtship Behavior Patterns in Captive and Wild Prairie and
Peregrine Falcons.

F. mexicanus F. peregrinus
captive wild captive wild

A. Attraction of mates
m. initially present on territory
-prominent perching na X? na X
-visiting potential nest sites na X na X
f. initially present on territory

-prominent perching na X? na X
-visiting potential nest sites na — na X
B. Establishing pair bond
pair remains on territory X X X X(F)
aerial courtship
-mutual soaring na X na X
-courtship flight na - na X(F)
-territorial delineation na X na X
cooperative hunting na — na X(F)
mutual roosting
-in territory X X X X
-at nest ledge — — — X(F)
m. visits potential nest sites X X X X
m. wailing X X
m. & f. display at nest together (1) X X —
m. tries to attract f. to nest by
carrying food to nest ledge X X X X
1972 Breeding Conference— Behavior C45

F. mexicanus F. peregrinus
captive wild captive wild

m. ledge display X X X X
f. remains on territory (2) X X X X
m. brings food to f. (courtship feeding) X X X X
m. & f. feed on food together X — — —
familiarities (mutual preening, etc.) X — — X(F)

Nesting preliminaries
f. begging food with much wailing ? X - X
m. begins making nest scrape X X X X
m. & f. display at nest site (3) - — X X
f. works on nest scrape
— X X X
mating
-f. wails X(faint) X — X
-f. solicits by posturing X X - X
-following courtship feeding X X — X
-comes directly from ledge display X X — —
-following defense of territory - X - X
-no observed preliminaries X X — X
. Egg-laying
f. dozing at length, appears sickly X X X X
first egg laid, f. shapes scrape X X X X
noticeably more aggressive X X X X
m. = male f. = female
;

X = observed
- = not observed
na = not applicable
? = not clearly defined
(1) Preliminary pair bond behavior observed in Prairie Falcons.
(2) Female perching near nest site.
(3) Male and female ledge display in Peregrine Falcons.
(F) From Fischer’s “Der Wanderfalk.”

our hope that this table will serve as a rough guide to the behavior pat-
It is
terns which might be expected in Prairie and Peregrine Falcons. Without ques-
tion, others will be able to add to the listed behavior patterns, and hopefully
those with data on other species will also be able to compare their observations.
It would seem useful for similar tables to be drawn up for other species, perhaps

using the above as a base and merely adding or deleting behavior patterns where
applicable.
C46 RAPTOR RESEARCH Vol. 6, Supp. C

PRAIRIE FALCONS

Preliminary Breeding Behavior. Preliminary behavior as described for the

PrairieFalcon in the literature is listed as aerial courtship however, other than
;

limited mutual soaring, we have yet to see any aerial courtship in Prairie Fal-
cons in the field. We have seen what we believe was described in the literature;
however it was not aerial courtship at all, but rather the interaction between
two pairs fighting over a cliff site.
The preliminary breeding behavior that we did observe occurred early in the
season as follows. A male and a female visited a series of nest ledges— the male
going to the ledge first, then followed shortly by the female. At the ledge they
would turn in a small circle, head down and side-by-side, both making the chup-
ping sounds. It appeared that each was trying to force the other one off its feet
by what appeared to be one trying to get its head underneath the other’s body
to lift it up. This they repeated over and over again. In the wild I have seen this
behavior only twice (Wayne Nelson has also observed something similar), and
with our captive pair we observed this behavior on April 3rd and 4th and then
it was all over. Its occurrence was early in the season, was very brief, and was

something that could have been easily missed. I have not observed this behavior
later in the breeding seasonand therefore describe it as preliminary pair bond
behavior.

Male at the Ledge. The preliminary pair bond behavior seemed to initiate
things and was followed by increased activity by the male at nest ledges. Initial-
ly males have been observed visiting two or more ledges, looking at them, mak-
ing scrapes, apparently paying little or no attention to the female and seemingly
satisfying himself with the ledges. We have observed this to some extent in our
bird in captivity as well as in the wild.

Attracting the Female to the Ledge. Next, in general, the male tries to attract
the female to the ledge of his choice by going to the ledge, displaying and call-
ing to the female. This occurs very often in the early phases as over and over
again the male goes to the ledge to display and he then comes out to look at the
female, or simply stands, chups and then looks up Very often he calls
at her.
with his head actually face down.
Frequently, in the case of the Prairie Falcons, the males also apparently tried
to attract the females to the nest by carrying food directly to the nest ledge,
and this in turn signaled the onset of “courtship feeding.” The male would take
the food to the ledge, get the female to come to him for food, and then take
off, leaving her there. These two activities were observed in both captive and
wild birds.

Food-transferring. In the wild the male Prairie brings food to his territory,
following which the female runs or flies to him, wailing loudly and mantling as
she comes. He then relinquishes the food to her when she grabs it, letting go
with barely a struggle. However, in our captive pair the male at first would not
1972 Breeding Conference— Behavior C47

relinquish the food and for the most part hung onto it. Sometimes he would
just stand there and female would approach, grab the food as well, and
eat, the
there would be a bit of a struggle. Usually they would end up eating it together,
side by side.

Female at the Ledge. At this stage in the wild, occasionally the female would
go to the nest ledge of her own accord. Although the male still tried to attract
her there whenever she was elsewhere, as soon as she arrived he would take off
and leave her, following which she would usually work on the scrape or perch
beside the ledge.
In captivity we did not observe the female nest scraping in the early stages.

Wailing. mentioned how the female in the wild can be heard wailing.
I’ve
This is incessant, and if you were to visit an area where Prairies are courting you
would be able to hear the wailing up to half a mile away whenever the female
calls to the male for food or is wailing prior to copulation (copulation most
often follows immediately after courtship feeding). In the captive situation we
could hardly hear the female as her wailing, though continuous, was very, very
faint. This may have been a result of the birds always being so close together in
confinement.

Copulation. Mating begins several days to a month or more before egg-laying,

and increases in frequency towards the laying of the eggs. (Captive Prairies first
mating April 9, first egg April 15.) Usually mating took place in one of the fol-
lowing three sequences observed in both captive and wild situations.
1. Immediately after the male had fed the female and she had finished the

meal, she would feak, put her head down and start to call (wailing) to him— he
would then come directly to her.
2. The male would be at the nest ledge, the female sitting elsewhere. She
would posture and call, and the male would come directly from the nest ledge.
3. Sometimes, with no preliminary that we could see other than the female
bowing, the male would come directly to her and mate with her.
The most obvious behavior here was that the female literally postured (or so
it appeared) any time the male flew towards or looked at her. I believe she ini-

tiated mating by the male in this way.

Often we saw the male simply look towards the female, and her head would
go down, she would then posture, head down, wings out— and appeared to soli-
cit the male to come. In the beginning he did not always come to mate, but as
egg-laying approached, the mating increased in frequency. The most matings
that we have observed in captivity was five times in a single day. In the wild the
most that I have observed was three times in one hour. In Peregrines mating has
been observed up to five times in one hour.

Lethargy near Egg-laying. Just before egg-laying there is a period when the
females appear very sickly for several consecutive days at which time they just
sit quietly with feathers puffed out. At this time the captive female was observ-
C48 RAPTOR RESEARCH Vol. 6, Supp. C

ed dozing on her perch, and whenever the male tried to attract her attention
she appeared to ignore him as if past much of the sexual stimulus.
This was also very obvious in the wild and although I have not noticed the
females ignoring the males, I have seen the females perch on cliffs for great
periods of time, all the while looking very sickly. This is very obvious in both
Prairie Falcons and Peregrines just prior to and during egg-laying.

Egg-laying and Aggression It was apparent that after the first egg was laid,
.

the captive birds became much more aggressive (Heinz Meng mentioned the in-
creased aggression in his Peregrines before and during egg-laying, and it intensi-
fied after the eggs were laid). Our captive male Prairie previously was not aggres-
sive, had always been shy, and was never very tame, therefore, in this case, the
increased aggression was quite obvious. Usually aggression in the wild appears
to intensify with egg-laying but is not as obvious or intense as observed in the
captive birds.
Serious incubation did not start until the last egg, on April 2 1 with incuba-
,

tion following for 39 days (i.e. until the last egg hatched). We will discuss incu-

bation periods in another panel, but I would suggest that you don’t pay too
much attention to some of the incubation periods that are listed in the litera-
ture for falcons or you could be in trouble.

Differences between Wild and Captive Birds. There were several things which
we noticed were very different between the captive and wild birds in the last
part of courtship, and in egg-laying. One activity that appeared to be normal to
both captive and wild pairs was the male bringing food to the female, or bring-
ing food to the ledge and the female coming for it. In the wild there seemed to
be an obvious regularity to much of this behavior. The male would bring food
within a set period of about an hour in the early morning, later in the morning,
again in the afternoon, and so on. In the captive situation there seemed to be no
regularity. Also in relation to incubation in the wild, male and female incuba-
tion periods appeared fairly well-regulated, but not in captivity, as our female
Prairie simply didn’t want to let the male have his turn at the nest for either
incubation or brooding. Still another area of abnormal behavior appeared in
courtship feeding, as mentioned, when the male was reluctant to give prey to
the female. Later we observed the male feeding the female on the nest ledge,
also the female feeding the male on the nest. I have seen nothing like this in the
wild to date.

PEREGRINES

Early Activities. As mentioned, the first sign we had of behavioral change was
“cacking” in our captive Peale’s Peregrines— this was also the first indication
that they were being territorial and that breeding behavior was beginning. This
was followed by an extended period when the male would stand by the wire
and wail (the “waaaik” call, which we have also heard very early in the season
in the wild). For Peregrines, Fischer lists several preliminary courtship activities:
1972 Breeding Conference— Behavior C49

the attraction of a mate, mutual roosting, courtship flights, cooperative hunt-

ing, wailing, and the male moving from ledge to ledge. Some of these aspects
we did see in preliminary courtship in our captive birds. We did not, however,
observe mutual roosting at the ledge, and I personally have not yet observed
this nor cooperative hunting in the wild.

Attracting the Female to the Ledge. Next in sequence is the activity of the
male at the ledge, examining ledges and displaying, apparently trying to attract
the female. Once again this seems to be very closely tied with courtship feeding.
I have observed both captive and wild males carrying food to the nest ledge and

the females coming to retrieve it from them. The male actually brings food to
the ledge, apparently to attract the female there. Often, as with the Prairies, the
male would go to the nest ledge and while bowed, call (“eechip”) to the female,
very obviously looking towards her, or alternately he would call, turn around,
and then go out and apparently look to see if she was attracted. This seemed to
be a very definite attempt to attract the female to the ledge and, again, was ob-
served in both wild and captive situations.

Nest-scraping. In our captive pealei the male was the first of the pair observed
to be making nest scrapes. In so doing he would go to the nest ledge, start bob-
bing, forming with his body, and foot scraping. (The same actions were used by
The female also made scrapes although generally
the Prairies.) a little later in
the period when visiting the nest ledge.

Copulation. In the wild, we have observed copulation both following “court-

ship feeding” and following the male and female nest display on the nest ledge.
In our captive birds we did not observe copulation, and this is apparently where
the courtship broke down. Although lack of mating was perhaps partly the fault
of the male, our observations suggest that it was primarily the fault of the fe-
male. In both captive and wild situations the male and female Peregrines shared
in incubatingand had regular changeovers. Also, both birds became aggressive
at the onset of egg-laying.

Pairing. Our observations suggest that the preliminary courtship ritual with
Prairies and Peregrines may occasionally be extremely short. At some sites we
have had Prairie Falcons perching in pairs at cliffs throughout the winter and
into the spring, at others lone males at the cliffs and at still others lone females.
In the latter cases as soon as the mate arrives everything apparently is “go”
suggesting that we have a situation in which we have long established pairs.
(To some extent this has been confirmed by trapping and identifying the pairs
several years in succession. Contrary to some of Jim Enderson’s early findings
in Colorado, we have found that our Prairies are staying with the same mates at
the nest sites.)
To be more specific we have examples in which we have been watching an in-
dividual Prairie Falcon at a cliff for a week or more when suddenly a mate
C50 RAPTOR RESEARCH Vol. 6, Supp. C

arrives and the same day they have been observed copulating. There appears to
be little or no preliminary activity in such cases.
Similarly with Peregrines, we watched a cliff for two weeks, on which the fe-
male was back first. Her mate from the previous year did not return, but a new
male did, and, again, on the first day he was there they were observed mating.
It is therefore not clear just how necessary some behavior patterns are, particu-
larly the preliminary ones such as courtship flying.

Failure at Copulation. We feel there are several reasons why we did not get
copulation in our captive Peregrines. One obvious observation was simply that
our female Peregrine at no time solicited the male. On one occasion we did ob-
serve her bowing, but when the male came toward her there was no posturing.
The male was observed on at least four occasions to attempt copulation, i.e. he
tried to mate with the female; on only one occasion did we see her head go
down, but that was all, and the male was not allowed to mate with her.
To further complicate matters the male apparently was afraid of the female
(something that we found rather difficult to tell with certainty). Several times,
for example, we observed that when the two were sitting close together, if the
female turned toward the male he would immediately back off or fly. Also
several times we observed him backing away from her when she approached. As
a result of these and other observations we believe that the male was definitely
afraid of the female. At no time did we observe familiarities (billing, preening,
etc.) describe^ by others for Peregrines and observed with our Prairies. All of
which suggests our pair were not truly compatible.
These are some of the factors which we feel prevented copulation and pre-
vented us from getting fertile eggs with the Peregrines, in contrast to the captive
Prairies, which were compatible, mated, and from their first clutch raised four
healthy young.

Blinds for Observing Behavior in Captivity. Initially it should be noted that

with regard to our captive Peale’s, because the female was laying eggs we be-
lieved that she was ready but the male was not. If we had not had the blind
there is no way we would have realized that the lack of fertilization may not
have been the male’s fault since we could not have assessed this without seeing
what the birds were doing. The preliminary activities, vocalizing, and egg-laying
apparently do not necessarily tell the whole story.
A blind allows you to see everything that is happening. With the Prairies we
photographed and observed mating at about 1 2’, and Tom actually watched the
female laying the individual eggs. We therefore knew what was in the nest and
what was happening at all times. Some of the activities occur so briefly that un-
less you can spend a considerable amount of time watching you may very well
miss some of the important things. Nevertheless, with some record of sequence
of behavior patterns it is possible to watch for and see the more important as-
pects.
If you establish a blind, don’t be too alarmed if at first you notice that your
birds are a bit upset every time they hear you, because they probably will be,
and may take a few days to adjust. However, after about a week they just seem
1972 Breeding Conference— Behavior C51

to forget about you. We found that we could talk and do almost whatever we
wanted in the blind.However, watch cigarettes, lighted matches, cigarette light-
ers or flashlights in the blind because these bright lights can be seen through the
mirrors. It is absolutely necessary to keep the inside of the blind at least twice
as dark as the outside in order for the mirrors to function.

Distractions. In apparent contrast to the adjustments the birds make to

sounds, we have noticed that the captive birds are easily distracted visually.
With our Prairie Falcons, when they could see out of the building, any sudden
movement would make them cease whatever they were doing. For this reason
we covered one end of the building with fiberglass, and immediately activities
proceeded without disturbance. Noises do not seem to bother the birds to the
same extent, whereas we felt anything new visually was a very definite distrac-
tion to them.

Disturbance, Desertion, and Damage to Eggs and Young through Visits and
Gunshots. My data from the field suggest that visits to eyries or the vicinity of
eyries immediately prior to egg-laying or at about the time of the first or second
egg may be very critical and the birds will readily desert. In contrast, once incu-
bating, the falcons at least will take a considerable amount o/ human disturb-
ance. (This appears to vary with individuals, and tolerance may be influenced
by toxic chemical residues.)
Also in relation to disturbance, one aspect that we observed will be of impor-
tance to anyone working in the field. We noticed that if a person comes to an
eyrie unobserved by the incubating bird, even while talking and making noise,
the bird will remain tight on the scrape. This holds true most often with Pere-
grines, Prairies, and Golden Eagles, so that the incubating bird usually remains
on the scrape until it actually sees the intruder. Even if one bird is flying around
making a great racket, the other will usually sit tight, with the result that on
sighting the intruder the incubating bird springs from the nest. Since in incubat-
ing eggs or brooding the small young, the adults work their feet under and be-
tween the eggs and/or chicks, when startled off, these may be thrown up to
four feet from the scrape. We have observed and photographed the results, and
actually have pictures of eggs and chicks three to four feet out in front of the
scrape. We have also come to a cliff and have seen a chick go down to its death
just because the brooding bird was startled, even though there was talking and
noise from above to warn the adult. I suggest that anything which startles the
bird on the nest with eggs or small downy young is actually endangering those
eggs or chicks. As mentioned, the eggs and small chicks apparently are resting
on top of the adult’s toes and tarsus, and may be flung out as the adult flushes.
If the ledge is wide enough, and not of jagged material, this may not be too
serious since the falcons can roll the egg back, or they will go as far as three and
four feet and pick up even a day old chick and carry it back. Nevertheless the
danger is a very real one, and I have recorded the results over and over again.
On the brighter side, we have also been able to observe the results when one
approaches directly so that the birds can see you coming. First they sit up to
C52 RAPTOR RESEARCH Vol. 6, Supp. C

look at you, then raise, and finally stand. They, therefore, are able to leave the
nest when they want to, not when they have been startled.
For people doing surveys I suggest this is a very real concern. If you are firing
guns at cliffs, or if you are going to the tops of cliffs and looking over, when
there are eggs or small young, you are risking the possibility of cutting down
production by one or more, depending on how often it happens. We have re-
corded these results, and I mention them because it was a real shock when we
realized just how frequently this occurred.

Literature Cited

Fischer, W. 1968. Der Wanderfalk. A. Ziemsen. (Die Neue Brehm-Bucherei.)

Wittenberg Lutherstadt.

IMPRINTING, AND OTHER BEHA VIORAL CONSIDERATIONS

by Wayne Nelson

Introduction. The discussions of imprinting at this conference were, to this

observer, among the most important. of the egg-infertility problems may
Many
be solvable by carefully considering that aspect of behavior. I will attempt to
throw some more light on imprinting and its problems at the end of this section.
This report is not a written account of my presentation to the conference;
rather, it is an attempt to synthesize and evaluate some of the discussions and
some of the questions which arose and which were concerned with the behavior
prior to and during egg-laying.
(An outline of courtship behavior in Peregrines (pealei) in the wild was pre-
sented to the conference, with 35 mm
slides and 16 mm
film. From this it is
hoped that the audience received a “feel” for the type of behavior patterns
which normally lead up to egg-laying in the coastal Peregrines.)
Based upon the courtship behavior seen in wild Peregrines, a discussion and
a number of suggestions for captivity breeding have been presented {Raptor Re-
search News, March- April 1971). That article considered a variety of factors
which appeared to be of importance to captivity breeding— e.g. spacious ledges
of a variety of types; one-way glass for viewing Undisturbed behavior; avoidance
of disturbance; necessity of water at laying and hatching. Several further points
must be added to that article’s discussion.

Some Behavioral Considerations. 1 Aggression. Heinz Meng described at this

.

conference how he “encouraged” his tiercel Peregrine to “defend” the breeding

chamber. Dr. Meng would run, crouching, past the chamber, and the tiercel
would chase along the ledge inside the windows of the chamber. (This pair of
Peregrines was given food through the windows of their building.) This situation
is very similar to what sometimes occurs in the wild coastal Peregrines; prior to

egg-laying one or both adult falcons sometimes, for no apparent reason, sudden-
1972 Breeding Conference— Behavior C53

ly will fly up to a quarter mile from their cliff to harass viciously a perched Bald
Eagle. may well
It be, as Dr. Meng suggested, that stimulating aggression (NOT
fear) may increase the pair-bonding of these birds— in captivity and in the wild.
2. Disturbance. There is a very definite difference between aggression and
fear, between the response of a bird to a distant intruder and to an intruder
which is at or in the eyrie. At the conference a number of interesting observa-
tions were reported.
(a) R. Fyfe described how captive Prairie Falcons (being watched from a
blind) ceased their courtship activities when human beings or other animals
were in view in the yard outside the breeding quarters; a translucent divider,
obstructing the view of outside activities, allowed the birds sufficient privacy
that their activities were not subsequently interrupted. These birds and others
became very accustomed to sounds in the nearby area and from the blinds from
which the birds were being watched.
(b) Maj. R. Graham told how a captive Peregrine laid a clutch (which was re-
moved), then recycled. At the time when the second clutch was expected, the
falcon’s abdomen distended somewhat, indicating that the first egg of the sec-
ond clutch was soon to be forthcoming. He felt that his visits to the nest ledge,
several times per day, prevented the falcon from laying that egg (and the second
clutch), and that it had been absorbed.
(c) G. Galicz noted one instance in which his Peregrines’ breeding quarters
was visited on one day— on the same or the following day an egg with no shell
was laid. He suggested that the disturbance of the visit might have caused the
production of the abnormal egg.
In view of these and other observations, it would seem wise to avoid entering
the breeding quarters occupied by a defensive (territorial) pair, especially just
prior to and during the laying of a clutch. Other means are available for check-
ing on egg-laying.

3. Food-transferring and Diet. A food appears necessary

plentiful supply of
for egg production. In a variety of raptor species has been observed that the
it

males usually eat the heads of prey items before giving the food to the female
or nestlings (Osprey— Ames 1966; Cooper’s Hawk-N. and H. Snyder, this con-
ference; European Sparrowhawk— Herren 1970; Peregrines— Nelson 1971; and
others). While the reason for this is not yet obvious, it appears to be of some
importance— and it suggests that we might be wise to avoid giving our captive
birds such unnatural diets as strictly or largely heads and necks. It is possible
that extra-thick eggshells are resulting from diets heavily laden with calcium
(bone), and that a few instances of nestlings dying upon hatching may be due
to difficulties in breaking out of very strong eggshells.

4. Vocal Stimulation from Nearby Pairs. From the behavior and laying dates
of wild Peregrines I have found no suggestion that the sounds (or other activi-
ties) of one pair either stimulates or inhibits the breeding of nearby pairs.
C54 RAPTOR RESEARCH Vol. 6, Supp. C

5. is some suggestion that clutches laid in the late spring or

Lighting. There
early summer may be due to relatively poor lighting within the breeding quar-
ters. Some pairs which lay infertile first clutches (followed by fertile second
clutches), and some pairs which refuse to come into any breeding condition,
may be doing so because of various degrees of inadequate light stimulation (see
Koehler 1969). Observations of the behavior of such late-laying or non-laying
or infertile clutch pairs could indicate what their difficulties are-and observa-
tions made on them when brighter lighting is provided should give additional
suggestions as to why these birds are behaving abnormally.

6. Photoperiod. Numerous writers in Raptor Research News and elsewhere

have indicated the importance of long daylength and changing daylength upon
captive raptors. The general lack of courtship behavior reported for tundra Pere-
grines (and Gyrfalcons) is obvious (see BPIE reports). One means of stimulating
courtship (and egg-laying) in these birds, that apparently has not yet been at-
tempted seriously, is that of giving the captive northern birds a photoperiod re-
gime which follows as closely as possible the changes which the birds in the
wild would receive— not simply giving them a long day-length only in the spring
and summer, but giving them a changing photoperiod which judiciously dupli-
cates that which the birds are seeing in the wild. This method must be tested,
since all other methods appear to be failing with these northern birds. I hope
to be able to expand on these thoughts at a later date.

7. The Variety of Repertoires. It seems fairly safe to say that no two pairs of
birds will necessarily behave in an identical manner. Variations in behavior, lead-
ing to the same results, need not be considered as signs of gross abnormality.
Fyfe (this conference) noted a number of similarities and differences between
the behavior of wild and captive Prairie Falcons; in general the results were the
same-e.g. in captivity the male may hang onto the food when he brings it to
the female, but she gets food from it whether he gives it to her or whether they
feed from it together; both male and female may incubate, side by side, but the
eggs get incubated.
Also, should be remembered that the birds may have to “learn” how to
it

carry out some of the more intricate behavior patterns— e.g. they may have to
“practice” a number of times before being able to complete copulation; the
first feeding of the nestlings may be very clumsily carried out by the female.
Don’t panic if the birds do not do something correctly or perfectly the first
time you see it.

IMPRINTING AND CAPTIVITY BREEDING

by Wayne Nelson

Introduction. To this writer, one of the most potentially useful concepts to

fall into place at this conference concerned the imprinting of captive breeding
stock. For the most part in the following discussion I will be considering the
1972 Breeding Conference— Behavior C55

larger falcons, but many of these observations and suggestions undoubtedly are
applicable to other species. (I must thank Butch Olendorff for providing me
with the initial stimulus which has brought the following thoughts and observa-
tions to the surface in my mind. While little of the following discussion is origi-
nal information, the way it all fits together does fall into the category of—“How
foolish of me not to have thought of that long ago.” Because many captive
breeders have not apparently seen the problem in this way, I am going to treat
it in some depth.)

Two Types of Captive Birds. By observing the courtship behavior of the cap-
tive birds we should be able to recognize two types of birds:
1
. those birds which are imprinted to their own species, and
2. those birds which are imprinted to human beings.
The former type, if given reasonable facilities, minimum disturbance, adequate
lighting and photoperiod, etc., should be capable of breeding on their own. The
latter type (imprinted to human beings) will possibly never “revert”, never com-
plete a courtship phase with a natural mate, and never produce a fertile clutch;
they are probably good for only one thing— artificial insemination reproduc-
tion— and they should be very good for that. There may be some gradation be-
tween these two types, and there may be some reversibility of the imprinting
in either direction. I would guess that the majority of the captive birds taken as
eyasses fall into the latter category— imprinted to human beings.
From a hurried review of some animal behavior literature, it appears that
little is known about imprinting in birds of prey. Konrad Lorenz (1937:263)
noted:
“Heinroth failed to breed hand-reared Great Horned Owls, Ravens and
other birds, for no other reason than that these tame individuals re-
sponded sexually to their keepers instead of to each other. In a very few
cases known, the birdrwhose sexual reactions were thus directed toward
man, finally accepted a fellow-member of the species which, however,
was always regarded ajs a rather poor substitute for the beloved human
and was instantly abandoned whenever the latter appeared.”

Mendelssohn and Marder (1970), in discussing captivity breeding problems,

pointed out:
“One of the reasons for lack of success may be that one or both part-
ners may be imprinted on humans. Many birds of prey kept in captivity
have been taken from the nest as young nestlings and been hand-reared.
If they are reared without the company of conspeciflcs, they will, if
young enough, develop imprinting on humans; the human companion
will be accepted at first as a parent companion, later as a social compan-
ion and eventually as a sex companion (Lorenz, 1935). A bird may re-
main imprinted for many years or even for life. Young owls may be-
come imprinted on humans until a relatively late age, since with these
species identification of the companion seems to be visual, and visual
C56 RAPTOR RESEARCH Vol. 6, Supp. C

acuity develops in owls only after their intermediate plumage has grown.
. .Sometimes even rearing owls together with conspecifics does not
.

prevent their becoming imprinted on the human foster parent (Heinroth

and Heinroth, 1926).”

Age of Imprinting. Most readers are probably aware that at a very tender age
(hours) ducklings, geese, chickens, etc., will fixate onto nearby moving objects
(sometimes the appropriate sounds must be given, also) and exhibit the “follow-
ing response” (a form of imprinting) in which the object— person, wagon, moth-
er bird, etc.— is followed and treated as the mother bird. In birds which are not
mobile soon after hatching, and which may even be blind for some days, the
imprinting process does not occur until a later date in the nestling’s life.
In the birds of prey it apparently is not yet clear exactly when the critical
period occurs, when the young birds acquire the “image” of their species, their
parent, and the features of their future sexual partner. There are some good
hints as to when this occurs, however.
McElroy (1971, and various Hawk Chalk articles) has described behavioral
differences between the “imprint” and the “brancher” in Cooper’s Hawks train-
ed for falconry. The “imprints”, taken from the nest at about 18 days or less
remained relatively tame, whereas branchers (fairly well feathered, quite mobile
but not quite flying) never became very tame and were almost as difficult to
train as wild-caught passage (migrant young) or adult Cooper’s Hawks. In gener-
al terms, with some adjusting due to differences in the rates of development,

this situation probably applies throughout the raptors.

In the Blond Ring Dove ( Streptopelia risoria ), Klinghammer and Hess (1964)
conducted experiments to try to discover when the critical period for imprint-
ing occurred. These doves first fly at about 14 days of age, and are fed by the
parents until at least 21 days of age. They were tested when about 10 months
old to see whether they would choose another dove or a person as a mate, after
having been reared under a variety of conditions (naturally by the parents,
hand-reared and isolated, etc.). The optimum imprinting (to people) was found
to occur when the young doves were taken from the nest and reared by people
at 7-9 days of age. If taken when somewhat younger, the doves were slightly
less predictable in their choosing between a human and a dove for a mate. If
taken when older than nine days, the doves usually chose another dove instead
of a person. In isolated doves, which saw no other doves and saw only a man at
feeding time and when pens were cleaned, the imprinting to people was more
likely to occur. Some doves, even having been imprinted during infancy to their
natural parents, did show some imprinting to people due to association (admit-
tedly brieD through their life in isolation. This study of imprinting in doves may
be the closest thing with which comparisons can be made with raptors. If the
general situation of imprinting in doves sounds at all similar to some captive
breeding attempts with raptors, I do not think it is coincidental. Indeed, it
seems probable that many or most of the captive breeding stock, which we are
trying to get to breed with a mate of their own species, are actually imprinted
to human beings.
1972 Breeding Conference— Behavior C57

Some Successful Breeding Attempts. Following from the suggestions seen in

the above information, does it seem entirely coincidental that:
1. Renz Waller’s twice successful Peregrines (1942 and 1943) involved a fe-

male of unstated age and origin and a wild-caught adult male? (Waller,
1968.)
2. Philip Shultz’s haggard pair of Prairies laid eggs in 1971 (one clutch of
four, infertile however)? (Shultz,BPIE 35, Raptor Research 6:31-35.)
3. Richard Fyfe’s successfully breeding Prairies (first clutch in 1971 hatched
all four eggs) were taken from the wild as “five week old eyas” (male) and

“four or five week old eyas” (female))? (Fyfe, BPIE 3 6 Raptor Research
,

6:35-36.)
4. Heinz Meng’s successfully breeding Peregrines (pealei ) (second clutch in
“
1971 hatched all four eggs) were taken from the wild when they were al-
most branchers ”? (Meng, BPIE 30, Raptor Research 6:25-28.) (Emphasis
is mine.)

The ages of capture of Beebe’s (1967) Peregrines (fertile eggs, 1967), Schramm’s
successful Peregrines (Peterson 1968), and Kendall’s (1968) and Enderson’s
(1971) successful Prairies were not specifically stated in their reports.

Some Apparent Contradictions. In American Kestrels it has been shown in

several instances that some hand-reared eyasses (even reared from eggs) eventu-
ally pairwith their own species (Koehler 1969, Porter and Wiemeyer 1970).
Porter and Wiemeyer (1970) mentionssome deaths of male kestrels which were
caused by their female partners, but the histories (backgrounds) of those birds
were not specifically stated.

The Captive Imprint. There are two situations in which “imprints” are found
in captivity.The first situation involves imprinted birds which we do not realize
are imprints because they are in a paired captive breeding effort. The second in-
volves birds which in general are held singly, and which show signs (sometimes)
of broodiness and probably courtship toward man, especially in spring. (Through-
out these discussions I am using the terms “imprint” and “imprinting” rather
loosely, mainly because I have been unable to find an adequate definition for
these terms as they apply to altricial birds.)

1 . The Imprint in Probably most eyass-taken falcons (of the large spe-
a Pair.
cies, at least) in captive breeding projects fall into this category. The two sexes
probably show little interest in each other; the female probably dominates at
feeding time. They both may become breedy, noisy, and very defensive in the
spring, and infertile clutches (even after recycling once or twice) are the only
result. They make good foster-parents as a rule.
Their apparent compatibility, their sharing of defense, incubation, and foster
parent activities, and their disrespect toward human intruders in their room— all
this does not tell you that they are paired, and it does not tell you that they are
not imprinted to human beings. In fact, if the female does lay eggs, it tells you
C 58 RAPTOR RESEARCH Vol. 6, Supp. C

that (a) your conditions are ext mely close to producing fertile eggs, or, more
likely, (b) your conditions are reasonably conducive to reproduction, but those
birds are indeed imprinted to people and not to their own species.
a. An apparent example of an imprint in a pair. By watching the birds (see

below for methods) during the courtship phase you should be able to determine
if the pair has formed a pair-bond and is in fact cooperating in their activities.
Fyfe’s observations of the point of pair break-down in his Peregrines are very
significant (Fyfe, BPIE 38, Raptor Research 6:38-39, and this panel report).
This pair in 1971 showed most of the expected courtship activities. But, at the
point several weeks prior to egg-laying when copulations should have begun,
this pair failed; the female failed to respond to the approaching male. Had she
responded to the stimulus “Approaching Male Peregrine,” she would have bent
forward into the copulatory posture. The male would then have been able to
alight on her back, and copulation could have been completed. This female laid
eggs— obviously she was sufficiently stimulated by the other features of the cap-
tive environment, and she even recycled when the first clutch was removed. She
was sufficiently stimulated for egg-laying to occur— but the male apparently was
the wrong stimulus for the final and most essential behavior pattern— copula-
tion.
b. Possible solutions to the problem of imprints in pairs. Assuming that the
captive breeder desires progeny from his birds, and assuming that he has spent
some time watching his birds and has seen no sign of copulation nor any sign
that it might occur, what should he do?
( 1 ) The arrival of the first egg is probably the first positive sign that we can

react to. At that point, entering into the breeding room (perhaps at night), cap-
turing the birds, anduiilizing (rough) artificial insemination (A.I.) methods may
be the only means of acquiring some fertile eggs in the first clutch. (In other
words, one should have practiced A.I. beforehand to be prepared.) Handling of
the female (with a partially-formed second egg within) would have to be care-
fully done. Candling of eggs at about 10 days of incubation should show wheth-
er or not fertility was achieved.
(2) The breeder could observe the behavior of the pair through the laying of
the first clutch, and positively discover whether copulations were occurring,
whether the male was just a few weeks slow (e.g. due to insufficient light stimu-
lation), whether the female was not responding to his approaches, or whether
he was not responding to her solicitations. Candling of the first clutch after
about 10 days of incubation should confirm the infertility of the eggs, and re-
moval of eggs should cause the birds to recycle.
This may be a very appropriate way of synchronizing the pair— causing them
both to incubate for a spell, and develop the appropriate hormonal and behav-
ioral responses, then stealing the eggs so that they both start from that point
into the processes toward egg-laying and sperm production. Recycling could be
expected to take about two weeks. (When recycling, one should be certain the
female has an adequate diet and a supply of clean fresh water at all times.)
At about the 12th day after the first clutch was taken (this length of time
appears to be fairly variable), if she is going to recycle, the female should again
1972 Breeding Conference— Behavior C59

enter into “egg-laying lethargy.” Again, by observing over a number of hours

(read a book behind your one-way glass if things arenot too active in the bird-
room), the person should attempt to confirm that the pair is indeed incompat-
ible, that copulation is not occurring. At this point, or immediately after the
first egg is laid, A.I. methods could be imposed upon the birds, to attempt to
obtain fertile eggs in the second clutch.
(3) A possible (but considered unlikely) long-term solution might involve at-
tempting to leave the two imprinted birds entirely isolated from people for a
long period of time, feeding and watering them via trap doors, or in absolute
darkness. There is a possibility that, lacking any reinforcement of the imprint-
ing to people (that is, no person presenting food, etc.), the birds might “revert”
back, might successfully mate with the other bird in the enclosure. Aside from
a small number of instances with kestrels, there does not seem to be much sug-
gestion that such reversion will occur in other species.

2. The Imprint as a Companion = the Human Being as a Bird’s Mate. This

method appears to offer the best means whereby the imprints can be brought
into productivity. It takes some time, effort, and devotion to the cause— but
these are some of the usual characteristics of the people involved with captivity
breeding.
Apparently this method has not yet been seriously attempted with the fal-
cons. Olendorff s comments on the Behavior Panel of this conference certainly
apply at this point.
This method has been very seriously attempted for several years, with some
success, by Fran Hamerstrom (1970) with her Golden Eagles. In 1970 Bob
Berry got fertile eggs from his Goshawk by this method. In 1971 Stan Temple
got fertile Red-tail eggs, and Bob Berry was completely successful with this
method, getting three full-grown youngsters from his Goshawk “pair.”
By pairing the imprint with a human being over a period of time (manning,
handling, etc.), and by manipulating photoperiod, nest or ledge materials, food
availability— the human mate should be quite capable of synchronizing the po-
tential breeders, which, of course, should be very definitely separated, lest they
kill each other fighting over possession of the shared human mate.

The A. I. involved actually has two possible techniques: (a) massage = rough
A.I. (also required for the imprints which are in pairs and are fairly wild), and
(b) cooperative technique— in which the bird attempts copulation with the hu-
man mate, and semen is cooperatively received from the male and shortly after-
wards transferred via syringe to the copulating female. Various degrees of grada-
tion between these two methods are possible and sometimes necessary (e.g. with
eagles)! We certainly look forward to detailed reports on the recent progress
with this most promising method.
Prolonging fertility for A.I. One behavioral aspect mentioned above merits
repeating in this slightly different context— this concerns the means of prolong-
ing semen production in the imprint male. To get the female to again produce
eggs we simply have to remove her clutch relatively early in incubation (if taken
late in incubation, the recycling period may be longer and less likely to occur).
C60 RAPTOR RESEARCH Vol. 6, Supp. C

The length of time the male bird of prey is actually producing semen is not
known to this writer. It may
be only a matter of several weeks or less. If the
first (presumably A. I. clutch is to be removed to an artificial incubator
fertile)
in order to obtain a second clutch, by that time the male may have “dried up”
as far as semen production is concerned. He may still be copulating with the
human mate, but his semen production may have ceased (we definitely need
some more information on this aspect of A.I. with raptors). And, in fact, the
male may have been somewhat out of phase with the female (either ahead of,
or behind her) for successful A.I. of the first clutch with his semen.
Syndhronization of the separated pair might be brought about through caus-
ing the female to incubate her first clutch for 10-14 days— and getting the male
also to incubate at this same time. The male might require some preliminary
courtship and nestbuilding play activities with the human mate, but he should
be motivated and prepared to incubate if he is provided with a nest and nest
materials just prior to when the female begins laying. When she has begun laying
the male could be provided with eggs for his nest (perhaps chicken eggs, tinted
to roughly match his species’ egg color). Once the female is laying, the need for
his semen diminishes. With eggs in his nesh, and perhaps with some help from
the human mate, he should begin incubating those eggs, just as the female incu-
bates the ones she has been laying. With. both sexes incubating at the same time,
presumably they will be rather closely synchronized, hormonally and behavior-
ally, even if they were out of phase prior to egg-laying. After both have been
incubating 10-14 days (this would allow candling of the female’s clutch, and
would allow both to become firm incubators), by taking away the eggs from
the male and the female (perhaps from the male a day or two earlier), it might
then be possible to have the pair perfectly synchronized in egg and semen pro-
duction, so that in the second time around the human intermediary could en-
sure successful A.I. of the second clutch.
Another possibility exists for those birds which are held under artificial light.
This method involves simply finding and keeping the birds at the photoperiod
at which they produce eggs and semen. These methods are used to some extent
with domestic poultry (I have not checked into the details with poultry). In
Slate-colored Juncos ( Junco hyemalis ) trapped in Illinois on migration, Wolfson
(1959) found that a day of 12 hours light— 12 hours dark prolonged testis activ-
ity for about nine months. Schwab (1970), with European Starlings ( Sturnus
vulgaris ) from California, prolonged sperm production in these birds for over
15 months by keeping them at either 10.5 hours light— 13-5 hours dark or 1
light— 13 dark. He found that 12 light — 1 2 dark did not cause those starlings to
continue sperm production for a long period- of time. In natural conditions in
California the starlings produce sperm for perhaps three months.
With raptors from various latitudes, of course the day-night ratio would have
to be modified slightly to keep the birds producing semen and eggs. This meth-
od holds considerable promise for producing multiple successful A.I. broods
per season. Until we know more about the effects on the birds of such prolong-
ed stimulation, it might be very wise to give the birds some months each year
in which they are not subjected to the demands of the breeding season.
1972 Breeding Conference— Behavior C61

The Captive “Non-imprint.” When a pair of birds has been taken from their
parents late in nestling life, beyond the major imprinting period, their chances
for successful captivity breeding should appear to be very great— provided that
(1) the pair is given appropriate and sufficient stimuli in their quarters, and (2)
they are not imprinted to human beings through frequent or constant associa-
tion such as at feeding times (remember that imprinting may not be confined
life, but may also occur by “associa-
solely to the “critical period” of nestling
tion” through life). The same general situation may hold true for passage and
haggard birds, in which cases the minimizing of disturbances may be extremely
important.
Knowing the histories of the pair should give the person a good idea of whe-
ther the birds are imprints or not.
Candling of the first clutches of such birds (any breeding birds) for fertility
is essential, unless successful copulations of the pair have been positively ob-
served. If first clutches are infertile, the birds should be forced to recycle if at
all possible, so that by the arrival of the second clutch (or third) the male might

be sufficiently stimulated by the external environment that he might do his

part. This allows the observer more time in one season in which to determine
whether the behavior of his pair is appropriate, whether they are indeed “non-
imprints.”
Caution. Some care must be taken so that misinterpretations of the efforts
of the breeding pair do not occur in the human mind. There is a definite likeli-
hood that if the lighting is insufficient, the female will be farther advanced than
the male, that copulations will not be seen until near the second or third (?)
clutch— until the male becomes sufficiently stimulated or becomes synchronized
with the female (see Koehler 1969). Some caution must be exerted here, so that
the “case” is not incorrectly “diagnosed”— so that a case in which the male is
insufficiently stimulated (e.g. by poor lighting) is not incorrectly diagnosed as
being a case in which the male is imprinted to people. To avoid making such a
mistake, and blundering into the breeding quarters during the laying of the first
clutch to attempt A. I. at that point, it would seem wisest to watch closely the
birds’ behavior through the laying of the first clutch, and through the first 10
days or so of incubation, then steal the clutch and watch their behavior as the
second clutch nears its time of laying— to learn positively whether the pair is in
fact cooperating in their activities, to learn whether copulations are occurring
or are being attempted.

Foster-parent Problems. Too little experimentation appears to have been

done to date to be certain of the effects of foster-rearing on potential captive
breeding birds. What good is a Prairie Falcon for captive breeding if the bird was
reared by, and is imprinted to a Goshawk? It might be wise to use rough A.I. in
such situations, but the problem would be less difficult if the Prairie had never
imprinted to another bird species but had imprinted to man instead— or better
yet, to Prairie Falcons. And think of the poor bird if it should ever be lost into
the wild— trying to court a wild Goshawk! Foster-rearing by a different bird spe-
cies may well create more problems by imprinting than it will solve. The scien-
tific literature concerning imprinting in waterfowl illustrates some interesting
C62 RAPTOR RESEARCH Vol. 6, Supp. C

examples of similarly very confused birds.

Watching the Captive Birds. It is obvious from the foregoing that a number
of problems in captivity breeding are solvable if the behavior of the birds is ob-
served in the courtship phase. In the larger species at least, it is unlikely that the
birds will be seen to do much of anything if the human observer is at all obvious
(even through small peepholes) and within any reasonable distance. The court-
ing birds seem to be rather shy.
How to watch. One-way glass (two-way mirror) is the obvious answer to this
problem. While there was some criticism of one-way glass at the conference, it
definitely has been shown by a number of people to be very useful (with wild
and captive birds), as long as (1) the glass is aimed in such a way that the birds
cannot see their reflections in it, and (2) the person is on the darker side (it
should be almost twice as bright on the birds’ side— otherwise they may see you
through the glass). Richard Fyfe has mentioned seeing a lighted match and a
glowing cigarette through a “mirror” in a store. Presumably birds could be dis-
tracted temporarily by such activities behind the one-way glass.
At first the observations from behind the viewing port should not be very
regular if there is any chance that the birds can hear the person there, or when
arriving or leaving; the first visits should be irregular and of varying lengths so
that the birds are unable to establish any regularity of sounds outside the enclo-
sure. Ideally, they should be accustomed to sounds from the nearby area so that
they pay no attention whatsoever to them. Tire quiet sounds of a radio, on and
off during the day at various times during the year may condition the birds to
expect any type of sound at any time from outside their breeding quarters. In
this way, the sounds made by the watcher would in no way inhibit the birds
from carrying out their normal daily activities and courtship activities.
What to look for in the pair. The various behavior patterns— or the lack of
them— are the keys to solving the imprint— non-imprint problem. Of special in-
terest should be those behavior patterns associated with the nest. Which bird
does the building or nest-scraping? Which spends time perched at the nest? Do
they perch close by one another? Which dominates at feeding? Does the male
take food to the female? Do they attempt copulation?
In some species we now have at least an outline of what behavior patterns
can be expected and in roughly what order they might be expected to occur.
Among the outlines of courtship behavior which we have at present are those of
the kestrel (e.g. Olendorff 1968), Prairie Falcon (Fyfe, this conference), Pere-
grine (e.g. Nelson 1971), Golden Eagle imprints (Hamerstrom 1970), Goshawk
imprints (Berry 1968, 1970, this conference), and others. And certainly there
are a great many clues and observations scattered through many of the BPIE
reports and Raptor Research News articles for those who will reread them.
As our knowledge of raptor behavior increases, we will inevitably be able to
shed light on more of the problems in captivity breeding. The assistance of all
of the people involved with captivity breeding projects, in making a few notes a
day on the undisturbed behavior of their birds, would undoubtedly be very
helpful in the future in solving problems which we cannot yet see.
1972 Breeding Conference— Behavior C63

Summary and Conclusions— Imprinting. 1 Probably most eyass-taken raptors

.

human beings; possibly some passage- and haggard-taken birds

are imprinted to
are also imprinted to people due to frequent association with man.
2. Itseems to be very unlikely that an imprinted bird in a pair will ever breed
with its feathered companion; the imprints consider human beings as “their spe-
cies”, and only human beings are likely to be considered as potential mates.
3. The age at which imprinting occurs in a nestling raptor is apparently not
yet known; some evidence suggests that the critical period extends to % of the
fledging (flying) age; pealei fly at 41-43 days of age— imprinting might be ex-
pected to occur in this subspecies up to 30-34 days of age, the point when the
nestlings are really beginning to feather out all over.
4. By observing the undisturbed behavior of a captive pair during the period
of time prior to egg-laying, the behavior patterns should tell the observer whe-
ther the birds are imprinted to their own species (i.e. they are attempting to
cooperate in their breeding efforts), or whether they are imprinted to man (i.e.
they are ignoring each other, or refusing to complete the courtship and mating
behavior with each other, and generally appear to be attempting to breed sep-
arately).
5. The behavior patterns which will provide the best clues for solving the
imprint-or-non-imprint problem will be those concerned with the nest, food
(sharing), and especially copulation.
6. Several possible solutions to the problem of imprints in pairs are discussed:

(1) rough A.I. in the first laying; (2) (best short-term solution) rough A.I. just
prior to, or during, laying of second clutch; (3) (least likely to succeed) leaving
the pair together in total isolation for a long time (years), hoping that they
might revert, or lose the imprinting to man, and mate with their feathered com-
panion.
7. A.I. appears to be the only means whereby imprints will be brought to
breed successfully. The cooperative A.I. technique with tame and manned birds
appears to offer the best opportunities.
8. Some caution must be employed so that some non-imprints (which are in-
sufficiently stimulated by a poor captive environment) are not incorrectly classi-
fied as imprints; behavioral clues are suggested.
9. Rearing birds with foster parents of another species may cause many diffi-
culties due to the nestlings probably imprinting to the species of the foster
parents, rather than to man or their own species.
10. It willbe by watching the undisturbed behavior of the captive pairs that
we will really begin to understand what their problems are, and why some of
them are not breeding in captivity.

References

Beebe, F. L. 1967. Experiments in the husbandry of the Peregrine. Raptor Re-

search News l(4):61-86.
Berry, R. B. 1968. Captive breeding behavior: American Goshawk-Part 1. Rap-
tor Research News 2(3): 58-72.
 .

C64 RAPTOR RESEARCH Vol. 6, Supp. C

Berry, R. B. 1970. Captive breeding behavior: American Goshawk— Part 2. Rap-

tor Research News 4(1): 1 8-3 1
Enderson, J. H. 1971. An account of Prairie Falcons bred in captivity— 1970.
Raptor Research News 5(l):44-47.
Hamerstrom, F. 1970. An Eagle to the Sky. Iowa State University Press, Ames,
Iowa. 143 p.
Heinroth, O., and M. Heinroth. 1926. Die Vogel Mitteleuropas. Berlin. (Cited in
Mendelssohn and Marder 1970.)
Kendall, H. 1968. Breeding Prairie Falcons in captivity. J. North Amer. Falcon-
ers’ Assoc. 7:57-64.
Klinghammer, E., and E. H. Hess. 1964. Imprinting in an altricial bird: the
Blond Ring Dove Streptopelia risoria. Science 146:265-266.
Koehler, A. 1969. Captive breeding of some raptors. Raptor Research News
3( 1 ):3-l 8 (translation by Frances Hamerstrom of Koehler 1968. Uber die
Fortpflangzung einiger Griefvogelarten in Gefangenschaft. Der Falkner 18:
28-33, 1968).
Lorenz, K. Z. 1935. Der Kumpan in der Umwelt des Vogels. Jour, f Ornithol.
83:137-213, 289-413. (Cited in Mendelssohn and Marder 1970.)
Lorenz, K. Z. 1937. The companion in the bird’s world. Auk 54:245-273.
McElroy, H. 1971. Desert Hawking. Peregrine Press, College Station, Texas.
216 p.
Mendelssohn, H., and U. Marder. 1970. Problems in reproduction in birds of
prey in captivity. Internat. Zoo Yearbook 10:6-11.
Nelson, R. W. 1971. Captive breeding of Peregrines: suggestions from their be-
haviour in the wild. Raptor Research News 5(2): 54-82.
Olendorff, R. R. 1968. A contribution to the breeding behavior of the Ameri-
can Kestrel in captivity. Raptor Research News 2(4): 77-92.
Peterson, R. 1968. The domestic raising of the Peale’s Peregrine Falcon. J.
North Amer. Falconers’ Assoc. 7:64-67.
Porter, R. D., and S. N. Wiemeyer. 1970. Propagation of captive American Kes-
trels. /. Wildl. Mgmt. 34(3):594-604.
Waller, R. 1968. A report of the first successful breeding of Peregrine Falcons
in captivity. North Amer. Falconers’ Assoc. 7:45-57 (translation by E.
J.

H. Spuhler of Waller 1962. Der Wilde Falk ist Mein Gesell. Melsunger:
Verl. Neumann-Neudamm. p. 285-304).
Wolfson, A. 1959. The role of light and darkness in the regulation of spring
migration and reproduction in birds, p. 679-716 In R. B. Withrow (ed.),
Photoperiodism. Amer. Assoc. Adv. Sci., Washington, D. C.
1972 Breeding Conference— Behavior C65

APPENDIX: USING CLOSED CIRCUIT TELEVISION FOR THE

OBSER VATION OF RAPTORS IN CAPTIVITY BREEDING PROJECTS
by J. Gregory Thomas

The use of closed circuit television for observation purposes in a captivity

breeding project can yield several substantial benefits to the researcher. First,
the quality of the observation can frequently be enhanced. The silent video
camera will not annoy or otherwise disturb the subjects of the observation.
Hence, observed behavior should be more truly reflective of the raptors’ reac-
tions to each other and to their physical surroundings. Possibly, the researcher
would then have the information needed to alter pairing of the raptors or to
make a change in some physical parameter of the experiment that would lead
to both breeding success and a better understanding of why or why not success
was achieved. Second, with the addition of video tape recording equipment, the
amount of observation the researcher could reasonably be expected to do can
be greatly increased. In the terms of the statisticians, the “sample size” has
been enlarged. This should help to improve the accuracy of conclusions drawn
from the observations. Third, another important benefit comes with the use of
video tape recorders. Observations can be saved and stored for later reflection
and study. Also, the observations may be shared with other researchers. This
would give a heretofor unknown depth to communication between researchers.
It might also aid in preventing costly and time consuming “reinventing of the

wheel.” A more detailed discussion of the costs, techniques involved, and bene-
fits to be derived follows this introduction. Systems and accessories both sim-

ple and complex will be included.

Preliminary Remarks. Most video equipment within a reasonable price range

is of Japanese manufactore. Experience in selling and servicing such equipment
has led the author to believe that it provides good to excellent results for a
relatively small outlay of money. It is used extensively throughout educational
institutions and in the business community. Therefore, it is sometimes possible
to pick up good used equipment at only a fraction of the original cost. The key
to getting a good buy in new or used equipment is to have access to a quality
video dealer who can both advise on hardware selection and service it. Between
five and ten percent of the initial investment on any video gear should be allow-
ed for maintenance each year. Because of cost considerations, only black and
white systems will be considered in this paper.
Briefly, other considerations that should not be overlooked include the fol-
lowing. Cable runs between cameras and display and/or recording gear shouldn't
exceed one thousand feet. Longer distances often require the use of expensive
line amplifiers. The general physical environment of the camera should be dry
and have temperatures not below 32 degrees F or above 100 degrees F. Devia-
tions from this will usually require the use of special environmental enclosures.
Because of the high light levels used in most breeding chambers, light is rarely
a problem. However, subjects should be frontlighted, and the camera can not
look directly at very bright light sources without damage to itself. Theft preven-
C 66 RAPTOR RESEARCH Vol. 6, Supp. C

tion should also be part of over-all system plans.

Single Camera Systems. The simplest system consists of one camera, wall or
ceiling mounted, connected to one monitor. The least expensive setup of this
kind involves using an existing TV receiver as the monitor. Most cameras have
what is called an RF output signal as well as a pure video signal. The RF signal
is adjustable for a range of channels usually running from Channel 2 through

Channel 6. The coaxial cable from the camera is terminated with an impedence
matching transformer which in turn is connected to the VHF antenna terminals
of the TV set. The picture being transmitted by the camera will appear on the
TV set when it is turned to the proper channel. TTiis system can be had for $300
or less, if one already has a TV to use as the RF monitor. If it is not desirable
or practical to use a home set in this manner, a separate video video monitor
should be purchased. The video monitor makes use of the video output of the
camera, and it generally will have a sharper picture than the RF receiver. Com-
plete systems that include a camera, monitor, cable, and in some cases even an
intercom cost $400 and up. Caution should be exercised when considering the
lowest priced systems. Picture quality and system reliability sometimes leave a
lot to be desired. All in all such simple systems should greatly increase the ease
and amount of observation the raptors receive.

Multiple Camera Systems. For more complete coverage of a breeding enclo-

sure, or the observation of more than one enclosure, more than one camera
may be needed. Each camera may have its own monitor, or a multiple-input-
single-output switcher may be used to display the desired camera output on a
single monitor. More details on switchers will be found in the section on camera
accessories. The use of more than one camera per enclosure would likely in-
volve using one camera with a wide angle lens for observation of general activ-
ity, and a second camera with a telephoto lens for detailed observation of the
nesting site.

Camera Accessories. There are many camera accessories available that will
greatly extend the usefulness of any video camera.
Lenses. Lenses come in a variety of forms. Wide angle lenses give a large field
of view with little detail. Telephoto lenses give a close, detailed view of a small
area. A zoom lens allows the user to adjust over the whole range, from wide
angle to telephoto. Fixed focal length lenses usually cost less than $100. Man-
ual zoom lenses start around $200, and remote controlled zoom lenses start
at about $900. Special lenses for use in very low light levels run between $200
and $350. Lens choice is important since it plays a large part in what the obser-
ver will see.
Scanners. Scanners are devices that allow the camera to pan back and forth
over the scene. They may be either automatic or remotely controlled to allow
for stopping the camera on a particular part of the scene. Thus, a lens that is
more telephoto than wide angle in nature may be used without losing the abil-
ity to see the whole scene. Automatic scanners run $125, and remote controlled
1972 Breeding Conference— Behavior C67

ones around $250.

Pan and tilt devices. Pan and tilt devices permit both horizontal and vertical
positioning of the camera from a remote location. They cost between $700 and
$1000. Coupled with a remote controlled zoom lens, this is the ultimate in con-
trol the researcher may have in choosing the scene to be observed.
Environmental enclosures. Temperature extremes or exposure to the ele-
ments requires placing the camera in an environmental enclosure, which costs
around $200 with a heater and blower.
Switchers. Switchers for routing video signals to monitors or video tape re-
corders may be simple, inexpensive ($50 or less) push button devices, or sophis-
ticated sequential electronic switchers costing $300 or more. The simple switch-
er will suffice in many cases, but the sequential switcher offers several special
advantages. With the simple switcher, the camera output desired may be punch-
ed up for display on the monitor at will. With the sequential switcher, the moni-
tor will sequentially display however many cameras are connected to it. The
time between switching is adjustable, and if any one scene is desired for exten-
sive observation, the sequencing can be stopped at that input. See the section
on video tape recorders to see how this switcher may have further uses.
Video tape recorders. For cost reasons, compatibility, and availability, only
video tape recorders (VTR) using half inch video tape will be considered. They
are capable of making high quality recordings and are usually quite reliable.
SONY, Panasonic, Javelin, and Hitachi are quality brands of both VTRs and
cameras. The simplest VTRs record and play back a video signal and usually
have a stop action feature. The next step up adds electronic editing and slow
motion playback capability. The simple VTR costs between $695 and $800,
while the more sophisticated models run $1000 to $1200. The maximum re-
cord time for these units is one hour with a conventional 2400 foot, seven inch
roll of video tape. Special thin tapes may extend this time by 25%, but give de-
creased service life. A standard one hour roll lists for $40, but a dealer with any
kind of volume usually makes them available for about $25. The tape may be
erased, recorded on, or played 500 times or more before becoming unusable.
The one hour time limit may be avoided by using a special time-lapse VTR.
These record at a slower speed than a standard VTR, and therefore a one hour
tape may be stretched to a seven, 12 or more hour tape. The author recom-
mends that no greater than a 7:1 or 12:1 reduction be used if any degree of
clarity is to be maintained for the picture. Besides being able to record extended
periods of time, the time-lapse VTR can greatly aid analysis of the material re-
corded. The recorder is played back at standard speed until some unusual activ-
ity flashes by. Then it may be slowed down for detailed observation. Thus, a
whole day’s activity may be recorded and reviewed in less than two hours time!
Nothing is missed and researchers’ time is saved for other duties. If the output
of a sequential switcher is connected to the time-lapse VTR, a sampling of a
whole day’s activity for many cameras can be obtained. Time-lapse recorders
cost between $1700 and $3000. The author personally feels the Javelin X-400
at $2000 offers the most in features and performance, especially slow playback.
Its only drawback is that when service is required, only a Javelin dealer is likely
C68 RAPTOR RESEARCH Vol. 6, Supp. C

to have the tools to do the job. On all the VTRs discussed an audio input is

available. With the use of microphones in the breeding quarters as well as cam-
eras, nothing is missed.

Summary. From this discussion of closed circuit television, it can be seen

that anything from a shoestring budget to a federal grant is needed for financ-
ing of a system, depending on the degree of sophistication required by the re-
searcher. Closed circuit television can give the researcher the tools needed to do
an in-depth analysis of the subject’s behavior. From this should come better
understanding of the complex processes involved in getting raptors to breed in
captivity. Following this should be more successes and easier duplication of
them.

[The author is in a position to offer new equipment at a 10-15% discount

(plus shipping) to serious researchers. Also, the author will offer limited (time-
wise, not ability) consulting services free to those seriously interested in the
design and purchase of closed circuit television systems.]
RAPTOR RESEARCH
Volume 6 Supplement Part D 1972

SPECIAL CONFERENCE ON CAPTIVITY

BREEDING OF RAPTORS-A REPORT

Sponsored by Raptor Research Foundation, Inc.

Held at Sioux Falls, South Dakota, U.S.A., November 22-23, 1971

Part D. Rearing, Parental, Foster, and Hand (Panel 9)

editedby
James Enderson
Department of Biology
Colorado College
Colorado Springs, Colorado 80903

Panel Members: James Enderson, Chairman, Robert Berry, Richard Olendorff,

and John Snelling.

Although the pre-hatching period in the artificial rearing of raptoral birds is

usually attended by a good deal of apprehension on the part of the person in
charge, the rearing of hatched young to adulthood can be accomplished quite
easily. For perspective, this panel outlined some of the characteristics of the
rearing of young by wild adult Peregrines and then explored the various tech-
niques and possible pitfalls involved in captive rearing.
Stan Temple presented time-lapse motion picture data obtained from wild
nesting Peregrines on the Yukon River in the summer of 1970. A more com-
plete report on this work is in press ( Living Bird— Enderson, Temple, and
Swartz). Throughout the first week after hatching young Peregrines were brood-
ed by the adult over 95% of the time where ambient temperatures averaged
57 F.
After the first week, there was a rapid drop in the attentiveness of the brood-
ing adults. Brooding occurred 10% of the time after the age of three weeks.
Correlations in the amount of brooding of young by the adults appeared to
occur with changes in weather conditions. As one would expect, brooding was
more frequent during rainy periods, and when temperatures were below the
average for the period. The implications of these data are that young Peregrines
will require the artificial application of heat almost constantly until they are
five to seven days old, and thereafter can tolerate room temperatures and lower.

D69 Raptor Research 6( Supp.) : D69-D 72, 1972

D70 RAPTOR RESEARCH Vol. 6, Supp. D
The time-lapse films showed that young Peregrines were fed within four hours
after hatching, and that the average rate of feeding by adults approximated two
feedings per young per day in the first five days of life, and that this rate in-
creased to nearly three feedings per day per young at least until the young were
25 days old. The interval between feedings was as high as seven hours within the
first few days of life, and then decreased to about four to five hours thereafter.
On the average, each feeding session by the adult lasted only about ten minutes.
There were no instances where young Peregrines up to the age of four weeks
were observed feeding themselves. Unfinished prey items were never left on the
nest ledge by the adults. The time-lapse films also showed that there was a very
marked decrease in the participation of the male on the nest ledge after hatch-
ing. The female did virtually all of the feeding and brooding. Two day old young
Peregrines were very sensitive to over-heating in direct sunlight and actively
sought shade at that age.
In regard to foster rearing in captivity, Bob Berry commented on numerous
recorded instances of the adoption of the young raptors by foster parents of
their own or other species. It also appears that lone adults will adopt young of
various ages. Generally, these adoptions have followed an unsuccessful attempt
by a broody pair or lone adult to produce young of their own in captivity.
There have been unsuccessful attempts to cause adoption after the foster par-
ents have stopped “brooding.” There have also been the cases of a captive Gyr-
falcon and a captive Peregrine Falcon, that have adopted and successfully raised
young without entering into a breeding cycle at all. Successful foster rearing
has occurred in accipiters, buteos and falcons.
Richard Graham described an instance in 1971 in which a pair of Prairie Fal-
cons, after the incubation of infertile eggs for four weeks, readily accepted a
pair of very young Prairie Falcons and reared them. On the other hand, a pair of
Peregrine Falcons, that had not incubated eggs, killed a Prairie Falcon chick as
soon as it was placed with them. Fran Hamerstrom related an account of a Gol-
den Eagle that accepted a one-day-old chicken, and one of a Red-tail, which
after accepting an adopted Red-tail chick, also adopted a white mouse.
According to Bob Berry, there are three major pitfalls in allowing captive
adult raptors to rear young. The first is that after a diet of day-old chickens,
cannibalism of the young raptor by the adults can occur. This circumstance was
seen in Henry Kendall’s Prairie Falcons, where the adults attempted to eat the
young. Adult Sparrow Hawks cannibalized their young in a project operated by
Tom Cade. Fran Hamerstrom also cautioned against feeding captive adults food
such as day-old chickens that are similar to birds they will later be expected to
adopt. A second problem, experienced by Berry, was the failure of a Goshawk
to brood her nine-day-old young. A third problem, experienced by Heinz Meng
in 1971, was the failure of the adults to feed the young, even though they
appeared to be doing so. Don Hunter cited his own experience in which an
adult Goshawk adopted a very small chicken even though it had been fed day-
old chickens prior to that time. It was pointed out that the peeping of young
raptors, or for that matter, young chickens, may inhibit the adult from killing
them. Hunter also cited an instance where a pair of Red-tails incubated and
1972 Breeding Conference— Rearing D71

raised a young goose until it was large. Phil Shultz said that his pair of Prairie
Falcons immediately brooded orphan Prairie Falcon chicks, but did not feed
them for two or three days, and they had to be fed by hand until that time.
On the subjects of artificial brooding and hand-feeding of young raptoral
birds, John Snelling spoke of hand-rearing 19 artificially hatched Kestrels on
day-old chickens, pigeons, mice, and sparrows. The food items were plucked or
skinned, and then the bones finely chopped with scissors. The temperature of
the still-air brooder was 95 F for the first week after hatching, 90 F up to three
weeks, and 72 F thereafter. He also said that he had had a good deal of trouble
rearing eagle chicks in Africa, where he had no means of accurately controlling
brooding temperature.
Richard Olendorff reported on a project involving the hand-rearing of Red-
tailed Hawks, Ferruginous Hawks and Swainson’s Hawks. Immediately after
hatching the young buteos were left in the incubators at 99 F, and showed no
distress. After 1 2 hours, heat was applied by means of incandescent lamps and
the proper temperature was obtained by trial and error. If the birds were too
cold, they shivered, and if too hot, they panted. The appropriate brooding tem-
perature from the first to the third day appeared to be about 88 F and between
the fifth and the sixth day, it appeared to be about 81 F and about the eighth
day, 77 F. Thereafter, the application of artificial heat was unnecessary. The
birds were brooded separately and it was pointed out that if three or four had
been brooded together, perhaps less heat would have been necessary. The young
buteos were fed every four hours, beginning within a few hours after hatching.
James Enderson mentioned that the down on artificially brooded young fal-
cons does not appear to fluff readily after hatching and that stroking with a
feather can raise the down and reduce the heat loss. Brooding temperatures on
several freshly hatched falcon chicks he reared were held between 90 and 95 F.
Chicks that are too hot tend to pant and stretch those that are too cool tend to
;

chirp, and pull their head under their abdomen. After the fourth day, the
brooding temperature was lowered to 87 or 88 F for the next four days. Bob
Berry also reported that young Goshawks and Peregrines show stress, at temper-
atures below 88 or over 92 F in the first week of life. In regard to temperature,
Heinz Meng mentioned that he placed a newly hatched Peregrine chick in a long
box with a heat source, a light bulb, at one end, so that the bird could regulate
its own temperature by moving toward or away from the bulb. After about the
first week, the chicks can tolerate room temperatures. A still-air styrofoam

brooder was used by Enderson, and a thermometer kept at the level of the up-
per surface of the chick at all times.
Young falcon chicks kept on a relatively smooth surface such as paper towel
seem unable to keep their legs beneath them or to raise their heads readily,
while those kept on coarse gravel do not experience that trouble. Bob Berry
mentioned that he had experienced the same problem in Peregrines and Gos-
hawks, and solved it by placing the birds on a soft towel in a bowl. Fran Ham-
erstrom mentioned that she has provided emergency heating for the young by
placing it inside of her shirt! She noted that chicks covered with a piece of
D72 RAPTOR RESEARCH Vol. 6, Supp. D
cloth are usually resting and when the cloth is removed they quickly raise their
head in anticipation of being fed.She has used an unorthodox brooding tech-
nique in the cases of very young harriers, horned owls and other species. The
young are carried under clothing just above the belt, supplying warmth, contact
and motion to the chick. When older, the chick is placed in a strawberry box
on a cloth pushed into the box. The chick is then covered with a cloth to pro-
vide weight because “when a brooding raptor rises to feed its chick, the young-
ster is exposed to light and lack of tactile pressure. Light and the release from
pressure are the dinner bell. When I take a chick out from inside my shirt, or re-
move the sock from the basket, I am giving the same signal, and the heavy-
headed chick has not wasted its precious time and energy in vain waving its
head about nor struggling for footing before this time.” According to Berry,
artificially brooded chicks do not readily submit to being brooded by adult
birds, although they eventually learn the source of the heat.
Richard Fyfe noted that wild Prairie Falcon chicks are negatively photo-
trophic in that they turn away from the light and huddle towards the rear of
the ledge.
RAPTOR RESEARCH N

Volume 6 Supplement Part E 1972

SPECIAL CONFERENCE ON CAPTIVITY

BREEDING OF RAPTORS-A REPORT

Sponsored by Raptor Research Foundation, Inc.

Held at Sioux Falls, South Dakota, U.S.A., November 22-23, 1971

Part E. Housing and General Management (Panel 2)

edited by
Richard Fyfe
Canadian Wildlife Service
10015 1 03rd Avenue
Edmonton, Alberta, Canada

Panel Members: Richard Fyfe, Chairman, James Enderson, Heinz Meng, James
Weaver.

HUNTER. I’ll introduce the next panel which is Housing and General Man-
agement and Richard Fyfe is chairman of this panel.

FYFE. Thank you, Don. Since several of our panel members won’t be able
to make it, I’ve asked Dr. James Enderson and Mr. Jim Weaver to fill in and I’m
very pleased that these people have accepted. Of the original panel Dr. Meng
and myself are here and we thought that we would begin by first generally out-
lining our facilities and what we feel are the strengths and weaknesses of these.
Then go through the various aspects of raptor facilities point by point and hope-
fully open this to some discussion from the floor. The panel will be Dr. Meng,
Jim Weaver, Jim Enderson and myself, and I think if you’re ready, Heinz, we’ll
go right on into the description of your facilities.

MENG. My work with raptor breeding and the development of my breeding

facilities started many
years ago. In 1946 I caught a passage female Red-tailed
Hawk that I have kept for the past 26 years. She has been kept on an outside
perch throughout the year next to an enclosure into which she can fly when it
is stormy. She has laid two or three eggs each spring since she was four years

old. Usually I substitute these for glass eggs and allow her to incubate them one
year. She even incubated the eggs in snow following a late storm in April. She

E73 Raptor Research 6(Supp): E73-E88, 1972

E74 RAPTOR RESEARCH Vol. 6, Supp. E

istethered throughout the year and builds her nest on the ground next to the
perch. Several times we have provided her with foster young and she has fed and
cared for these birds conscientiously. One year I removed her first clutch of
three eggs and three weeks later she had recycled and laid two more eggs. Many
hawks when they become older become sort of like brood hens and will raise
almost anything that is put under them. She has even raised a chicken which
she hatched and raised to about three weeks of age. Whenever the chick got
cold it would go under the red-tail just as if it were the mother hen. Unfortu-
nately one night there was a thunder storm and the chick walked away and died
of exposure.
My breeding chamber initially began with a small white-painted hawk house
built in 1953. In 1963 I built an addition to the hawk house specifically design-
ed as a breeding chamber for a pair of passage tundra Peregrines. The breeding
chamber is located in our back yard and is very secluded. It is surrounded by
many shade trees so that it is quite cool in the summer, also there is a swampy
woods in back. As illustrated the breeding chamber was attached to the original
hawk house.

At first the 1964 pair of passage Peregrines ( Falco peregrinus tundrius ) had
the entire chamber to themselves, but in 1 969 this chamber was partitioned to
make two separate chambers. The partition was made out of 2x4 lumber and
covered with burlap on each side. Entrance is through the hawk house into
chamber A; in order to get to chamber B one must go through chamber A.
Therefore, by going through the hawk house there is always a closed door so
Window areas, without glass, are provided
that the birds cannot possibly escape.
in the northand south walls of the chambers. The windows are 6’xl0’ and are
provided with 3/8” vertical hollow aluminum rods placed 2 Yi” apart center to
center. The window areas facing north are 4’xlO’. On the outside the aluminum
bars are covered with 2”x4” mesh welded wire coated with vinyl to prevent
escape should one of the falcons be able to push between the bars. At the bot-
tom on the window areas on the south there is a 3’ wide shelf with a lip 2” high.
This whole area is filled with sand. At the end of each shelf there is a large bath
pan. At 3’ above the bottom shelf there is another shelf 3’ wide covered with
synthetic rug and astroturf material. The north window areas are smaller but
 .

1972 Breeding Conference— Housing E75

have no shelves. The upper half of each area is covered with burlap to provide
some protection from the north wind but the open lower half allows some wind
to circulate through. The floor is of cement. Perches and shelves are arranged so
that the birds cannot defecate on them from above. There are only three rafters
in each chamber and these are covered with rug or astroturf. The birds usually
use these to roost on since they are the highest perches available. The interior
of the breeding chamber is unpainted, and the walls are of homosote board, a
pressed fiber material. Although there are lights in each chamber they were not
used during the 1971 season. In chamber A, which houses the Peale’s Peregrines,
the nest ledge runs north and south along the eaves of the original hawk house.
The ledge is two feet wide and has a lip of rug covered 2x3 inch wood, sand
about 2 inches deep fills the entire ledge (dimensions 2x15 feet). The part of
the hawk house roof that is covered by the breeding chamber forms a darker
area above the nest ledge. Half way up this sloping roof is a horizontal shelf
about one foot square; often both birds sit on this ledge together. A one-way
mirror has been installed in the peak to facilitate observations of the pair. The
perches are all padded. In addition to the ledges there are a couple of stone
ledges (cement blocks) where the birds can sit if they so desire

Successful Peregrine Breeding in Captivity. The Peale’s Peregrines used in the

successful breeding attempt were taken from the Queen Charlotte Islands in
British Columbia in 1967. They were taken almost as branchers and each bird
was from a different eyrie. The birds were initially handled for about a month
and were then put in a small holding chamber which was about 1 2 feet x 4 feet
x 7 feet high (the south end of the original hawk house), and here they were
kept for the next two years. At two years of age, the pair was put into the
breeding chamber which was 10 feetx 15 feet x 10 feet high at the eaves (cham-
ber A).
The was fed from the outside of the breeding chamber and as early as
pair
the of 1970, the male became quite aggressive. Their food consisted mostly
fall

of fat, healthy, freshly killed homing pigeons. As a precaution against tricho-

moniasis the heads and crops are removed, also the feet. The pigeons are then
torn into three pieces by pulling the wings apart, which tears off breast from
the breast bone, and then removing the back and legs from the breast piece
which is still attached. Viscera are not removed. The food is then fed on the
upper window shelf from the outside. Both birds come to the ledge and take
the food from my hand. Also day old chicks and an occasional rabbit have been
fed to this pair of birds. No drugs or vitamins were used, although wheat germ
oil was mixed with yolk and put on the meat for about a week during the end
of February, but they don’t like the taste. At about the time of egg laying, the
male became even more aggressive towards me and would actually try to get me
through the bars. Several times I would run along the side of the hawk house in
sort of a crouching position and he would be very vocal, run along the window
shelf, and try to attack me. I feel that this stimulated him sexually and perhaps
had something to do with his being fertile. Often I could look in from the out-
side and see the falcons glaring down at me with their wings poised for a Hying
E76 RAPTOR RESEARCH Vol. 6, Supp. E

attack.
The birds were both very vocal and four eggs were laid at two day intervals
starting March 4, 1971. After eight days of incubation the eggs were candled
and found to be infertile. These eggs were then removed and 13 days later the
falcon had recycled and laid four more eggs. For each clutch incubation started
with the third egg and the female did most of the incubation at the beginning,
but as time went on the male did more and more. Copulation was not observed
and it is felt that it probably occurred on or above the nest ledge. Incubation of
the second clutch was not interrupted and three eggs hatched on May 8th and
one young on May 10th. During the incubation period the male took food to
the nest ledge and gave it to the female. It was noticed that the female went to
the nest site and started plucking and eating at the time the young were peep-
ing. On May 10th, two days after I heard the first peeping I finally climbed to
the nest ledge. I hadn’t disturbed the falcons since the laying of the third egg of
the second clutch. When I climbed to the ledge (1:00 PM) I found two dead
young, one almost dead, and another just hatching. I brought the young into
the house, but the one which was almost dead died shortly after, and the one
that was hatching was out of the egg at 2:00 PM and immediately ate three
small pieces of fresh pigeon breast meat. For the next ten days he was fed most-
ly skinned eviscerated day old chicks. These were cut up and fed with round
tipped forceps; all of the bone and cartilage was fed along with the meat. A 40
watt bulb was used for heat and was so regulated that the temperature stayed at
about 35 C (95 F). The bird was able to move away from the light if it felt too
warm.
It appears as though the female fed herself at the nest ledge but did not feed

the eyasses although she brooded them well. I did not use any supplementary
vitamins and on occasion I did give him little slivers of fresh pigeon breast. How-
ever the main diet was day old chicks. During his 14th day of life he consumed
eleven chicks. The skin and feathers were removed so that there would be no
pellet forming material. Every two hours I simply fed him a full crop and I did
not have to wait for pellet regurgitation.
In summary, the results of the 1971 breeding attempt are as follows:
First clutch— 4 infertile eggs laid, removed on eighth day.
Second clutch-4 eggs laid, all fertile, all hatched-three of the four young
died apparently due to inexperience of female, the fourth young was
successfully hand-reared with the result that one Peale’s tiercel was pro-
duced. He is in perfect health and plumage, with no hunger streaks and
weighs 24 ounces.

THOMAS. At what age was the eyas removed from the parents?

MENG. He was removed before he came out of the egg, just at hatching. Ac-
tually the three eggsbegan hatching on May 8th. I heard a lot of peeping and I
provided food on the window ledge. The male came and brought the food up to
the female and I could see her plucking it. I could see the back of her tail and
the young ones were peeping so I thought everything was going fine and I pro-
1972 Breeding Conference— Housing E77

vided them with a pigeon twice a day tom into three sections. On the following
morning I went out again and as soon as she got off the young ones started
peeping and the same thing happened, that is the male brought the food up and
she began eating. On the third morning I didn’t hear any peeping so I decided
that I had better look because I hadn’t been up there since the third egg of the
second clutch. I didn’t want to disturb them and when I went up there again I
saw that two young were dead and another was practically dead. The fourth was
just coming out of the egg. Apparently the female had gone up there and fed
herself but hadn’t fed theyoung ones although she was brooding them. As soon
as Isaw that I grabbed everything and hand-raised the last young in the kitchen,
I put him under a lamp and fed him about every two hours. It was unfortunate

that that happened. Several others have had young prairies and the females fed
their eyasses all right, butfemale apparently fed herself but didn’t feed the
this
young. Next spring what I plan to do is take them when she hatches them and
put additional eggs under her for about one week, raising the young inside.
Then I will re-substitute so she’ll be getting a group of young that are a week
old already. Maybe she will be mature enough then so that in the future perhaps
I won’t have to continue in this substituting.

VOICE. What is the floor covering?

MENG. Just cement, but they rarely go down onto the floor unless it is to
catch a live chick that may have fallen down from the feeding shelf.

ENDERSON. Do you keep the pair together year round?

MENG. Yes. They have been together for as long as I have had them. When I
firstgot them they were put in the original hawk house in a four foot wide sec-
tion that I had partitioned off. They were there for three years while the tundra
birds had the complete run of the breeding chamber. Since then they have been
in chamber A.

VOICE. How much manning have these birds had?

MENG. The first had them, for a period of about a month and a
year that I

half, perhaps two months I would take them out and put them on block perches
and take them in at night, putting them on my elevated shelf perches.

VOICE. What about the diet of the adults prior to egg laying.

MENG. The diet consisted mostly of freshly killed homing pigeons, also day
old chicks.

THOMAS. Did you have problems with wind, etc., and disturbance?

MENG. Not too much. If you think back

' I mentioned that on the northern
E78 RAPTOR RESEARCH Vol. 6, Supp. E

exposure, the north windows are 4x10 feet with the top two feet covered by
burlap woven between the bars. So there is just a 10x2 foot wide strip that is
open on the north exposure, with the result that there is no difficulty and they
are out of the wind.

THOMAS. Did you ever, like Jim Enderson has done, try using barrels to try
and stimulate more of an eyrie cave type situation? I have seen a prairie project
where there were ledges and nothing was happening, but the day they put one
barrel in the prairies were in there clucking and just going crazy. They seem to
like the protective nature of such a situation.

MENG. I haven’t used barrels, but as I mentioned the nest site was back in
the comer in the most hidden area of the breeding chamber.

WEAVER. The new facility at Cornell University, Ithaca, New York is a steel
sheeted building 227 feet long and 47 feet wide. It is divided inside to house 38
pairs of birds in 38 separate chambers. Thirty-six of the chambers are approxi-
mately. 10x20x17 feet at the apex of the roof Hoping to 14 feet at the eaves.
The entire area is surrounded by a six fpot Cyclone fence with a barbed-wire
top. The outside openings of the pens are screened first by one-half inch steel
conduit bars. Six inches beyond the bars the outside open area is covered with
half inch hardware cloth and the bottom two feet are covered with fiberglass
light panel. The fiberglass is primarily to take up the extra space left from the
conduit as the conduit lengths are only 10 feet. However it turns out that it is a
good idea to keep blowing snow off the bottom of the cages. Inside, the walls
of the pens are paneled in plywood. The roof has at least one sheet of light
panels that are white. Each room has two or three wall perches, most of which
are covered with cocoa matting. We were using hay bales, but with the expense
combined with the problem of molding, we have decided to replace these with
cocoa mats, which seem to closely resemble the texture of perches used in the
wild (that is tundra heather, dwarf birches and that sort of thing). Each room
has at least two ledges that can be used for nesting; these are filled with gravel.
The edges of all the perches are padded with Tartan Turf, similar to Astro Turf.
Each room is vented at the top with half inch hardware cloth behind the wood-
en bars, situated above the lights. Because the temperature immediately under
the steel roof can exceed 1 00 degrees in the summer we have installed roof ven-
tilators. We feel that with three big ventilator fans the length of the building,
we should be able to draw enough air from the front to cool it considerably.
Each room has the facility for eight 150w flood lights for artificial lighting, and
a pane of one-way glass for observation. There is also a double decked observa-
tion hall that runs the length of the center of the building. Of course it is also
heated to an extent in the winter so that it will not freeze, and it affords good
opportunity for observation. The floors are covered with gravel and we furnish
a little bit of straw underneath the perches to facilitate cleaning. Our plans are
simply to go in once or twice a year to take the straw out and work the gravel
over with a weak solution of formalin as is done in the poultry industry. We
1972 Breeding Conference— Housing E79

have been feeding a number of birds on the fist in an effort to try to keep them
as tame as possible. We don’t know whether it is going to be really valuable or
not, but in most cases it would seem that the smaller the disturbance caused by
entering the pen, the better your chances are going to be. I think in some of
them we will completely isolate the bird during the breeding season. That is,
we will just put food in through the observation port or something of that sort.

ENDERSON. I am describing the new facility that I built in the past summer
at Colorado Springs, which may
be of interest to many of you, because the cost
involved is considerably less than the larger facilities such as at Cornell. This
building consists of five parallel rooms, each six feet wide. Each room is 18 feet
long except that the nest ledge on one end adds three more feet to that dimen-
sion. There is an enclosed alley- way adjoining one end of all five rooms, with
doors opening into the breeding chambers. The outside dimensions are 30x24
feet; the cost is around $1,300 for materials.
Each room is equipped in the same manner. The interior walls are painted
white, the floor is dirt. The perches are straw bales, except for the lip of the
nest ledge which is covered with nylon carpet. The nest ledge is covered with
two inches of coarse sand and small pebbles. In Colorado bales of straw tend
not to mold, so they are suitable for use as perches. A lower perch projecting
from the wall at eye level, enabling the birds to fly from the floor to the higher
perches in two steps is provided.
The opening in the roof is 3x12 feet and is covered with 2x4 inch welded
wire.
Behavior of Captive Pairs. The four pairs of Prairie Falcons vary in their ad-
justment to each other. Members of compatible pairs seem to notice each other;
in fact, at times they step on each other’s wings and tails without paying much
attention to it. I have never noticed any kind of antagonism among two pairs.
Another pair, including birds which are about eight years old, appear to me to
be poorly adjusted to each other. They are very tense in captivity and easily
frightened, even though the female has laid eggs and the male copulated with
another female in captivity in 1967. They are tried and proven in that sense,
but they seem not to be compatible. In another pair of Peregrines, the female
perhaps is dominant over the tiercel and forces him to move. Also with the birds
which seem compatible and adjusted in the nesting situation, a play -type of be-
havior has been observed where the birds appear to play with the wings of three
week old chicks which had been a steady in the diet of these birds.

FYFE. In contrast to the previous discussions of breeding projects and breed-

ing facilities,we have been working with a very different situation in that we
have been forced to use a large number of different breeding chambers, each
with different dimensions, exposures and natural lighting. Due to such factors
as availability of buildings, incompatibility of birds or apparent non-adjustment
of birds to a given situation, we have had to improvise a great deal and in so do-
ing have experimented with a variety of different situations for perches, ledges
and lighting. We have moved our birds around to different rooms and have de-
E80 RAPTOR RESEARCH Vol. 6, Supp. E

amount of natural lighting to which they would be expos-

liberately altered the
ed. Wehave experimented with a variety of perches and perch locations and
have even painted the insides of the chambers white in order to increase light
intensity.

Room Size Our breeding chambers have ranged

. in size from the smallest
rooms approximately 12x12x6 feet high to rooms as large as 30x20x18 feet
high. In general our observations of the larger falcons suggest that the smaller
rooms contribute to a high degree to the anxiety of the birds. We have had Pere-
grines, Prairies and Gyrs in small rooms and it appeared that their inability to
avoid the nearness of any intrusion created a great deal of anxiety. In such a
situation these birds were observed to fly directly into walls, the bars of win-
dows or into the weld wire. These large birds also appeared to have a tendency
to fly just below the ceilings of these rooms with the result that most were con-
stantly tipping their wings as they flew from one perch to another. In contrast,
the smaller birds, kestrels and Merlins, did not seem to have the same degree of
difficulties in these small rooms, although they also exhibited a degree of anxi-
ety and would fly into the walls or the wires covering the windows, when we
entered the pen, otherwise they seemed well adjusted and did not have the same
tendency to tip their wings on the ceiling as they flew. One pair of Prairie Fal-
cons kept in a small room of this type did however lay five infertile eggs in a
barrel-type nest ledge in 1 97 1 Initially this pair exhibited the same sort of anxi-
.

ety which we attempted to minimize by staying away from the pen for the most
part; however, once egg laying began, aggressive behavior became more domi-
nant, and it was extremely dangerous to enter this small room with this particu-
lar pair of birds. In contrast, we had our pairs of adult Peregrine and Prairie Fal-
cons in rooms which were much larger by dimension, roughly 27x17 feet and
about seven feet high. Although these birds were not as easily disturbed by our
presence there still was a tendency to fly from one end of the room to the other
and into the wire screen covering the window. There was not quite the same
tendency to fly immediately next to the ceiling and a minimum of feather dam-
age was recorded. In 1970 and 1971 we did not have any birds attempting to
lay in these pens although we did provide them with both barrel and open
ledges and we believe the primary reasons for the lack of breeding was the fact
that we had immature birds included in each pair in these rooms. (This has been
borne out since in 1972 we had two pairs lay in these pens.) The next largest
chamber that we used was the chamber in which we had successful breeding.
This was a room 27 feet long by 12 feet at the apex sloping to about seven feet.
The most obvious behavior of the pair in this room was simply that they had a
tendency to fly from one end to the other; however, the mere provision of high-
er perches seemed to result in a major adjustment. It was particularly noticeable
that within a couple of days of the placement of the Prairie Falcons into the
pen, they would sit above us with relative indifference, in sharp contrast to the
frantic flying which had been so prevalent with this pair in the previous pen
which had a low ceiling. This chamber was situated on the north side of a bam
and consequently received only a small amount of sunlight. Initially, the east
1972 Breeding Conference— Housing E81

end of the room was covered with one inch weld wire but was later covered
with fylon when we observed the birds altering their behavior with the slightest
movement or activity observed outside of the building. In order to increase
light intensity, the roof of this building had also been opened into long sections
which were covered with fylon and the inside of the room had been painted
white with the exception of the small area immediately behind the nest ledge.
The west end of the building had a large window, 4x6 feet. This window afford-
ed some opportunity for these birds to sit in the sun in the late afternoon or
evening. Two small windows on the north side were only about five feet in
height and were seldom used by the birds.
The largest pen was the loft of a barn which had been renovated and was
felt to be the best of our breeding chambers for the larger falcons. The rough
dimensions were 27x28x18 feet high; i.e., it was large enough to provide an
opportunity for these larger birds to fly around inside. The birds in this pen did
fly a great deal, receiving a considerable amount of exercise and were in excel-
lent condition, judging by their case in making the floor or food ledge to the
higher perches. This room was open for the full length on the south side and
openings were covered, also part of the east end of the building and a small por-
tion on the roof at the southeast peak of the roof. All open areas were covered
with one inch weld wire. Perches were installed specifically so that the birds
could sit in these open areas to receive a maximum amount of sunlight. This
room was also judged to be the most suitable because of the obvious adjustment
of. the birds to the breeding chamber. We believe that the height provided the
necessary security for the birds and the size of the room, perches and available
nest sites seemed to meet the basic requirements. Unfortunately, the pair which
were in this room for two years laid infertile eggs both years, as discussed later
in the section on behavior.
Lighting and Enclosure Coverings. All of the breeding chambers in the pro-
ject have been lighted through natural lighting conditions, primarily because the
natural lighting at our latitude provides the natural photoperiod for the Prairie
Falcon, Peale’s and anatum Peregrines. A serious attempt was made in all of our
buildings to create as large an opening or alternately flight pens on the south
and west sides of the buildings.These were generally in the form of open win-
dows covered with either fylon or weld wire. We also used fylon extensively to
cover open areas of the roof or to screen the larger windows so that the birds
would not see activities which might disturb them. We have found that the clear
fylon is a very suitable material which allows a maximum amount of light, al-
though it does screen ultraviolet we are told. Because of the problem of birds
flying into the wire of windows or other openings, we have tried to minimize
this by placing perches adjacent to the wire screening. We found that the birds
used these extensively and very often this has solved the problem of an individ-
ual bird flying into the wire. With Merlins and with Prairie Falcons that were
prone to fly into wire, we have used snow fencing on the outside of the wire
which creates a visible barrier. The birds tend either to perch in front of the
snow fencing, or if they do strike the wire, they strike feet first. This has greatly
decreased the number of times the birds have flown directly into the fence and
E82 RAPTOR RESEARCH Vol. 6, Supp. E

has virtually eliminated the problem of cut ceres and broken or frayed feather
ends.
Ledges. We have used three different types of nest sites. For the larger falcons
we have used open ledges with a narrow lip which were filled with turf, gravel
or sand or a combination of each. Most recently we have been using a fine gra-
vel and have found this eminently suitable. We have also barrel ledges with a
similar substrate; however, in 1971 where we had both barrel ledges and an
open ledge in the pens, both the Peregrines and Prairie Falcons chose the open
nest ledges. (In 1972 all three pairs of adult birds chose barrel ledges.) The
Peale’s Peregrines were provided two open nest ledges and one barrel nest ledge
the same height in 1971. The second nest ledge is actually lower than the barrel
ledge. In the two nesting attempts this pair chose the open ledges. We have also
provided barrel ledges in rooms that were without other provision and in the
case of one pair of Prairie Falcons they laid in the barrel ledge provided. The
third nest site provided was simply an old Magpie nest for our year old Merlins
and although we did observe some activity in association with the nests, the
birds did not lay. Since this is the normal nest selection in the wild, we felt that
this was the logical choice for these birds in captivity; however, because of the
difficulty in moving these nests, in the future we will construct our own using
twigs and wire frames.
Perches. As indicated, we have tried several types of perches and locations.
In general, these have ranged from normal flat wooden perches of 2x4 or 2x6
to round perches usually in the form of natural tree branches. We. have tried
swinging perches which are simply tree branches tied at each end and suspended
from the ceiling and also straw bales, both suspended on walls and left on the
ground. Our flat perches have been covered using such materials as cocoa mat-
ting and sand or gravel.
By preference, we have now settled pretty well on straw bales, and tree
branches for perches. Straw bales are very excellent in our area because they re-
main dry, can be changed readily, and we believe, they provide some relief to
the birds’ feet. Tree branches are also suitable, easily obtained and because of
the round shape, provide a perch where the bird is not sitting on the ball of the
foot, which again we believe is of some assistance in the elimination or preven-
tion of foot problems. We have also used swinging tree branch perches particu-
larly for larger birds such as the Gyrfalcons, and although these are suitable,
they have their limitations and are difficult to suspend properly.
Any discussion of perches inevitably comes back to the question of foot
problems and generally I can only say we have had very few foot problems.
Curiously the majority of these problems with us have been with Peregrines and
not with Gyrfalcons or Merlins which have so traditionally been prone to foot
problems. For the most part, we have been able to solve foot swellings by pro-
vision of alternate perches although in cases where a corn was involved, we have
had to go to minor surgery.
It is our observation that the choice of location of perches is very important

and that three basic considerations should be met; i.e., (a) they should provide
security for the birds (height with large falcons) (b) there should be provision
1972 Breeding Conference— Housing E83

for the female to perch near the nest ledge prior to egg laying and (c) perches
should be situated so that any attempt at mating would not be thwarted by the
location (e.g. closeness of the perch to the ceiling). We have attempted to pro-
vide two or three perches near each nest ledge, one adjacent to the ledge and
two to one side or the other and also a perch across the mouth of the nesting
ledge itself. We have found that the birds appear to prefer to eat on straw bales
or a flat surface of this nature, and will carry their food to straw bales or a flat
ledge for plucking and eating. High perches at or near windows are designed to
provide an element of security and at the same time allow the birds to spend a
good deal of time in the sun.
Blind. If you wish to know what is happening and unless you have very tame
birds, the provision of a blind or a suitable viewing area is absolutely essential
in establishing new breeding pairs of birds. Although the benefits are described
in some detail in the section on behavior, it is worth indication that a blind
should have the following features. First it should provide you with the oppor-
tunity to approach and observe the birds without being seen by them so that
they will not be disturbed nor aware of your presence. We have also found that
blinds provide suitable opportunity for recording and photography; however, if
this is going to be done, the holes and photoports should be provided well in
advance so that the birds will not be disturbed when they are used. Blinds
should be large and comfortable enough to provide an opportunity to sit and
observe for a considerable period of time without discomfort. We have found
one-way mirrors allow optimum viewing provided they are not situated so that
the bird is not looking at itself in the mirror. The mirrors should be removable
so that they can be kept clean and should be large enough to provide adequate
viewing and possibly photography if you so desire. The inside of the blind
should be dark as you need a two to one differential in light between the inside
and the outside of the blind so that the mirrors function properly. And finally,
you should not smoke or, that is, have a lighted cigarette or any bright object
such as a match, lighter, flashlight or anything that would reflect light immedi-
ately behind the mirrors as this is readily observable and will upset the birds. If
you are in a cool climate, the blind should also be provided with some ventila-
tion and possibly a fan so that in the early spring the mirrors do not become
covered with condensation. There has been some question as to disturbance due
to the blind and I suggest that the birds will adjust readily to sound in most in-
stances; therefore, the blind should be set up in such a way that it can be ap-
proached. However, once in use, a little bit of preliminary work, perhaps visiting
the blind and moving around for a few minutes a day until the birds gradually
get adjusted to it, the use of a radio in the blind turned on very quietly, or

something of this type will help to adjust the birds to sound from the blind and
they will soon become accustomed to this. One additional suggestion is simply
that the blind be situated in such a way that you can look directly into the nest
ledges and can therefore be in a position to determine whether the birds are lay-
ing or not. Your blinds should also allow you to keep the birds under observa-
tion in all areas of the pen so that it is possible to observe the various behavior
patterns which might occur in one area or the other.
E84 RAPTOR RESEARCH Vol. 6, Supp. E

Floor Covering. The floor covering in all of our pens has been wheat straw.
Although straw and hay are apparently considered taboo in some areas, we very
much prefer wheat straw as it is clean, relatively dust free and since we are living
in a very dry area, it is very easy to keep clean. We do not have problems with
moldy material and straw is very suitable since it absorbs the droppings from
the birds, is readily cleaned and is very inexpensive.
Perhaps we can now go into the questions. You will have noticed that we
have described different types of housing and in checking the literature, it ap-
pears that most of the facilities were actually enclosed buildings or large out-
door cage type aviaries or a combination of the two.

SWARTZ. Imight comment a little bit just to add to the data on some of
these things. have noticed the same sort of thing with Gyrfalcons which you
I

mentioned with height and found that immediately when the bird had an op-
portunity of getting away and looking down he was much happier. I have sub-
sequently built some additional quarters with this provision. Your comment on
blinds also leads me to remark that for some years now. I’ve had a closed circuit
television but since I just use a domestic set, it has to compete with Porky Pig
and other such things. So I propagate with R.F. (that qualified as closed cir-
cuit). This has been an elegant system and I have boxes in all the quarters that
permit rotation of the camera and it’s really much cheaper than you probably
have thought. Merlins somewhat to my surprise have preferred the kestrel box
sort of thing by and large. For somewhat over three years now I have been
working with both the progeny of a ground nesting pair and a pair of birds
atypical to the Alaskan situation which use an old Magpie nest. I have provided
ground situations, ledge situations, ledge situations with overhanging roof and
ultimately a giant gigantic chickadee box sort of arrangement made out of old
blazo boxes (a wooden gasoline box with a hole cut in it) very much like a kes-
trel box with a slightly larger opening. As mentioned, it appears that they have
preferred the kestrel box sort of thing.

VOICE. What do you mean by swinging perches?

FYFE. These are perches that are tethered from the roof. They are a branch
or a trunk of a small tree tethered at either end by a rope so that when the bird
lands on the perch it has a tendency to give. We have used these particularly
with Gyrfalcons.

VOICE. What’s the hypothesis behind the choice of this perch?

FYFE. This perch has been used to permit the birds to land without striking
too hard. We have noticed that with Gyrfalcons in particular that they have a
tendency to hit very hard when they land on perches and the idea behind the
swinging perch is to lessen the shock and thereby prevent injury to the ball of
the foot.
1972 Breeding Conference— Housing E85

GOBEN. have observations on height that is contrary to what you have in

I

that my Hawks have behaved very differently from the fal-

pair of Ferruginous
cons. I had them in an outdoor aviary where they could go indoors and out-
doors and they had a considerable height where they could get up and look
down on the surrounding yard. That year I did not get anything and I removed
the birds and the next spring I put them into a 4x6 cubicle and got fertile eggs.

EBERLY. Did you find any practices or were there any practices that could
have influenced the preference of the ledges over the barrels, or did you pretty
well arrange things so that there was enough variety?

FYFE. We have tried to arrange the ledges and barrels in such a way that
they were backed up to the wall at relatively the same height. In the largest
all

pen where the Peale’s were put up we had two ledges and one barrel. Of the two
open ledges one was at the same height as the barrel and one was much lower.
In the two nesting attempts these Peregrines used the two open ledges in prefer-
ence to the barrel. The barrel was used just for stashing food.

STODDART. Are you trying to look for a formula which would be applic-
able to all raptors or individual species rather than for an individual bird? You
seem to be paying a lot of attention to nest boxes or ledges to height and so
forth.

FYFE. The thinking behind what we have been doing has been hopefully to
find a formula for a given species, in this case for the Peregrine or for Prairie
Falcons. The basic idea behind it is to provide security and a situation in which
the birds were at ease. I fully realize that birds have different temperaments,
but what we have tried to do is to provide them with as many situations as pos-
sible and to let them select the situation which they might prefer. We have tried
to give them more than we felt was needed in the way of perches and ledges as
mentioned, to let the birds select so that we might learn where the preferences
may be.

STODDART. Are the birds you are working with from the same source? I
would think that if you are trying to find a formula for a species, it would be
necessary that all of the young you are working with and the parents would
have to be from the same source with the same temperament, and so on. As
for example, in the case of Jim Enderson’s Prairie Falcons, I think that they
would nest in almost anything although this might be a situation which would
be very uncomfortable, birds whose history was different from Jim’s birds. I
personally think everybody seems to be paying too much attention to finding
the secret whether it is a nest box, a ledge or height. I don’t think that there is a
secret and I think the solution lies directly in the birds’ securities or insecurities
and it will depend on the individual bird species.

FYFE. Basically, I think that I agree with you in that we are trying to make
E86 RAPTOR RESEARCH Vol. 6 Supp.
,
E

provisions which will provide for the necessary security of the birds. We cannot
tell by looking at a bird what its particular requirements may be; therefore, we
have been trying to make as many provisions as possible and are letting the bird
more or less select its own preferences.

HUNTER. think that this poses a very interesting question and one which I
I

had hoped someone would take on. It poses the ecological problem of whether

or not an individual will choose the situation in which he grew up, or whether
the choice of a site is innate. I think that someone perhaps in the position of
Dr. Porter with his experience with kestrels might be able to add something
along these lines.

HAMERSTROM. Perhaps in partial answer to the last question, I have found

that two horned owls which I have came from different nest sites. One came

from a stick nest and the other came from a hole nest. The female selected the
stick nest and the male selected the hole nest. I have no owlets.

VOICE. Relative to the use of wire on the pens, have you had problems with
the birds flying into the wire?

FYFE. Yes, to some extent we have, particularly with tiercels of all species
and definitely with Merlins. When I was in Patuxent last year, I noticed that
they had their Merlins in wire pens, I believe, weld wire in construction. How-
ever, they had put snow fencing on the outside of the pen and when I inquired
they advised me that this kept the birds from flying directly into it. We have
tried this with some of our Merlins and with Prairie Falcons and it definitely
does help. They still fly into the wire, but they land feet first on it and they do
not try to go head first into the wire and damage themselves accordingly. In the
same regard, we have not observed any problems with the fylon covered areas.

HUNTER. I have one more question that

I would like to ask and I don’t

know who to direct Has anyone in the group had experience using snow
it to.
fence on top and what happens when you have a heavy snowfall in such a situa-
tion?

SHULTZ. It gets quite heavy. The snow fencing is on wire with 2x4 supports
and we haven’t had any break-through to date. We have, however, gone out
early in the morning and knocked snow off because we felt it does present a
potential hazard.

CAMPBELL. I would like to ask Jim Weaver whether or why they use spot-

lights as opposed to floodlights at Cornell?

WEAVER. They are floodlights.

SHERROD. Also, a question related to the floodlights. Can you tell me what
1972 Breeding Conference— Housing E87

the intensity would be?

WEAVER. do not know what the exact intensity would be. We are primar-
I

ily giving them a good source of light and are not worried about intensity.

LAWSON. Is there any reason why everybody wants to use anything besides
artificial light? Why is there thiscommon bond to give them some sort of natur-
al lighting; what is the reason behind it? Someone suggested vitamin D. Do you
mean from a supplement?

VOICE. Yes.

VOICE. question how much value we are going to get from sunlight through
I

this opaque material since it cuts out all the ultraviolet light.

SHULTZ. Several of us have built pens with opaque or plywood walls with
screened or snow fence tops. My facility is staggered down a hillside with a 4Vi
foot difference between the uphill and bottom hillside. It is built with plywood
walls so that the bird has no visibility of dogs, children and so forth from the
outside and the top is screened with snow fence. We have never had our birds
fly into the wire at the top. They have all the sunlight in the world and they can
see the blue sky and trees. They seem to be extremely happy in this situation
and are comfortable.

SWARTZ. A comment on your point. In contrast, I have not made any ef-
fort to use natural light and this is dictated in part by the fact that I’ve got 50
to 60 degree below winter to get through in Alaska. The more windows I have,
the more troubles I have. To date I have not got eggs, but in other respects I

feel that I am getting complete response out of incandescent light as the sole
source of illumination. To this point I have not considered that there is any
super unique magic in any particular wave lengths of light. Although talking
with Joe Simony i shakes my confidence a little. I was clearly within three or
four days of laying with my Merlins just a few weeks ago (judging from follicle
size) when disaster struck. And that’s total incandescent light in a converted
coal bin.

STODDART. With regard to artificial light I still think that it might work
well with birds taken out of the wild specifically for breeding. That is, if they
were kept under artificial light for three or four years. I have tried it on old fal-
conry birds because I thought it would really enhance their environment. In this
regard I have done it with two Peregrines and a Prairie all of these birds were ;

old, the youngest being four years when I put him under artificial light. The
birds’ condition dropped. The weight didn’t drop because I adjusted this, but
the birds lost feather bloom so when I put the birds up to mate I didn’t consider
total artificial light, but this year we are going to have artificial light as supple-
ment. I think for a bird who is three or four or seven years old that has been
weathered using falconry methods and is stuck under artificial lights as I have
E88 RAPTOR RESEARCH Vol. 6, Supp. E

done three times, that it may be an emotional strain on them. They don’t feel
right and they don’t preen enough and so forth. You know if your bird is losing
bloom, when it’s baby bloom in the first place, you know something is the mat-
ter. Maybe an imprint Peregrine or something that was used to the lights all of
the time would compensate for it. But it has adverse effects on a bird that is
used to being weathered.

FYFE. I would just like to make one comment in relation to light. We have
stayed with natural lighting because we are at the same latitude as the majority
of the birds we are dealing with and we felt that in this manner we w^re in a
position to provide them with their natural photoperiod.

HUNT. We should keep in mind that day length is only one cue that birds re-
ceivefrom the environment. There is also the position of the sun, the stars and
the moon and I would think that when in doubt, we would be best advised to
use natural light.

FYFE. A final note which may be of interest is that as mentioned we are at

the same latitude as the wild birds we are dealing with and in our case, both the
Peregrines and the Prairies were laying at the same time as the wild birds.
RAPTOR RESEARCH
Volume 6 Supplement Part F 1 972

SPECIAL CONFERENCE ON CAPTIVITY

BREEDING OF RAPTORS-A REPORT
Sponsored by Raptor Research Foundation Inc. ,

Held at Sioux Falls South Dakota U.S.A., November 21-23 1971.

, , ,

Part F. Artificial Insemination (Panel 7)

edited by
Robert B. Berry
Yellow Springs Road
Chester Springs, Pennsylvania 19425

Panel Members: Robert B. Berry, Chairman, Frances Hamerstrom, Walter Mor-

gan, Stanley A. Temple.

HUNTER. If there are no more questions I think perhaps we can move on to

the next thing on our program which is artificial insemination. One of our guests
who is here to give us a demonstration has a limited amount of time so we will
proceed to that. Now, Bob Berry.

BERRY. I’ll be the chairman for this artificial insemination panel discussion.
I want to pass out some papers that I have prepared. For those of you who
don’t get them, you might get them from someone else or write to me and I will
send you a copy. Now, if Bill Halliwell is ready with the slide projector.

HALLIWELL. Yes, I’ll turn it on for you.

BERRY. Well, I’m not quite ready. I want to say that what we hope to cover
here basically is the two techniques of artificial insemination. Both Stan Tem-
ple and Dr. Walter Morgan from State University of South Dakota, will demon-
strate the massage technique on chickens. I will talk about cooperative insemi-
nation in the American Goshawk, a technique that does not require massage or
manipulation of the gonads. Fran Hamerstrom is also on the committee and she
will discuss thetechniques that she has applied to the Golden Eagle. Perhaps the
easiest way for me to show you or explain the technique of cooperative insemi-
nation is to show you some very few slides. If we can have the lights now we

F89 Raptor Research 6(Supp.):F89-Fl 10, 1972

F90 RAPTOR RESEARCH Vol. 6, Supp. F

can begin.
Before there can be any cooperative insemination there must be a pair bond
established between the researcher and the experimental subject, in this case a
female Goshawk which was raised in captivity as a household pet. She is four
years old at this time, and she is accepting a stick from me in probably the same
fashion as she would accept sticks or food items from a male Goshawk. She
builds her own nest. She fulfills all the requirements for a natural mating ex-
cept, of course, she does not have a natural mate. All during this nest building
stage and during the period of copulation which precedes the eggs, incidentally,
by about three weeks, she is extremely defensive about other people, other
human beings than myself, and against conspecifics. In this case the male Gos-
hawk, which happens to be housed in an adjoining chamber and is also what we
can refer to as a conditioned or imprinted eyass bird. The male also builds a
nest and carries out at least the prenesting cycle. I have never given him young-
sters to raise. But I presume he would probably incubate eggs in the same fash-
ion as the female. The male was conditioned to ejaculate semen voluntarily. I
started in February to condition the male. He was four years old and had never
displayed any indication of sexual response. There was no display in previous
years, no chirping, and no territorial aggression. I merely started feeding him
from the fist. He was a typical eyass Goshawk. He mantled violently and he
attempted to escape with the food. Within two weeks he stopped mantling and
within another week he started chirping towards me. Within a month when I
gave him the food he would hop off two or three feet, and he would hop back
to me and give the food back. He would step off, look at it and chirp. He would
stick his tail up in the air and chirp some more. So I knew that he was coming
into breeding vigor. When I approached the cage, his tail would be straight up in
the air as I think you will see in another picture and he was chirping violently—
all the time being completely and totally hostile towards the female. The female

was doing the same thing at the same time in the adjoining pen.

VOICE. They could see one another, is that right?

BERRY. They could see one another during the 1970 year. This was dupli-
cated during the second year with no visual contact between the two birds, and
they behaved precisely the same at the precise same time of year. They dis-
played vigorously. He was obviously very excited. One day I was in there feed-
ing him in the early evening which was my general practice and I put a paper bag
full of food on the ground next to me and I went to feed the female. He came
down and attempted to copulate with the paper bag. That evening I was devis-
ing all sorts of artificial female Goshawks in my mind, trying to figure out how
the devil to get semen from him. I didn’t want to handle him physically because
one of the very important aspects of maturation of the gonads is the absence of
stress. I feel that if you grab a bird you are going to have one chance and only
one chance to take semen. So I was trying to devise a technique whereby semen
could be taken without shaking him up. These are actual photographs of his
first clumsy attempt to copulate with me as the mate. This is the bird landing
1972 Breeding Conference— Artificial Insemination F91

on my leg. Notice how his feet are balled up so that they do not damage the
mate. He is calling violently. He kind of works his way down to my shoe. This
is a trifle bizzare, but it works [laughter] . Now he is down on the shoe. He is

just about to copulate with the shoe as you can see. This is actual copulation.
The wings are down. He is on his tarsi, he is straddling the shoe. And his abdo-
men is revolving in a circular fashion and pressed down against the shoe. This
forces the papilla to penetrate the panel feathers, the papilla being the rudimen-
tary copulating organ in a bird of prey. They have no structure comparable to
the mammalian penis. So this papilla is forced down against the shoe which of
course would be pressed against the female’s cloaca or the everted oviduct in
natural mating. This is immediately after— I guess this is a withdrawal process,
a sort of post-copulating depressive mood [laughter] It wasn’t very many days .

until I devised a workable technique and sort of mutually agreed that my hand
would be the desired copulating object. I would go in to his chamber, place my
hand and fingers on this block which was his feeding block at about a 45 degree
angle and he would fly to the block, placing one foot on one side of my hand
and the other foot on the other side and copulate with the back of my hand.

VOICE. Facing your arm?

BERRY. Facing my arm. Exactly the way it occurred right here. He literally
works his way right up on the hand, the papilla penetrating the panel feathers.
I guess it’s about a quarter of an inch long. It’s merely a lymph fold. I imagine
Stan could probably fill you in on this.

VOICE. Is it just blood that distends it?

TEMPLE. It develops during the breeding season. Not blood.

BERRY. But has some erectile tissue in it. It is not very much. It is quite
it

short. But anyway this lodges between the fingers, and he ejaculates. The semen
is collected from the fingers in a tuberculine syringe and taken to the female.

Maybe you can see it better in this picture, a better idea of the way the feet are
balled up and he gets down on his tarsi and so forth. During the 1970-71 season
we went through this routine 117 times. We only got semen 15 times during
thiswhole process. There are many, many, many dry runs preceding a one-week
period during which you get most of your semen. And the week period corres-
ponds precisely to the egg laying period of the female. Collectable amounts of
semen were only secured three times during the 1970-71 season. Three times
for each season.

VOICE. How much is collectable, cc-wise?

BERRY. semen amounted to about 0.02 cc which is equivalent

Collectable
to about a good amounts were ejaculated.
sized drop of water. Very, very small
However, examination with a microscope disclosed the semen was very concen-
 1

F92 RAPTOR RESEARCH Vol. 6, Supp. F

trated and very motile. When I approached the female Goshawk, she assumed a
precopulatory posture. This is taken with a telephoto lens. When I was 40 or 50
yards away the female would become quite excited and fly to the door of the
pen. She assumed this position with her head down and her wings out a little
bit. The only physical part of this whole routine by me is the stroking of the
female on the back which stimulates her to spread her panel feathers and expose
the cloaca. Actually right there I am not really working with the cloaca, I am
palpitating the bird to find out just where the eggs really are in the bird. You
can feel an egg as the abdomen extends. It becomes enlarged about two to three
days before an egg is laid. You know when that egg is coming. For maximum
fertilization, insemination should be 36 to 48 hours preceding that first egg.
This photo illustrates the minimal stimulation on her back. You notice the tail
is going the other direction and unfortunately the wing in this particular picture

is drooped down to support the conspecific male, obscuring the division in the

panel feathers. She is actually exposing her cloaca. She just wouldn’t tolerate
anybody else around when this whole thing was going on; I can understand that.
Generally I would do this with the left hand, the minimal amount of stroking
of the back, then I would grasp the tail in the left hand and lift it up, exposing
the cloaca. In this picture there is no eversion of the oviduct. In the year 1970
there was no eversion of the oviduct. The syringe was placed in the upper left
hand portion of the cloaca and inserted about % inch. The left side corresponds
to the functional left oviduct in a raptor. In the 1 97 1 year the cloaca turned in-
side out on the bottom exposing the small orifice of the oviduct. In the 1 97
year the semen was placed about xh inch into the actual oviduct. It didn’t seem
to make any difference because there were four fertile eggs in 1970 and three
fertile eggs in 1971, 1 00% fertility in both years.

THACKER. Bob, when you took the semen specimen, did you immediately
run into the next pen and implant it?

BERRY. Immediately. The semen was immediately transferred and was not
diluted. There is always residue. If you collect perhaps .03 cc, there is always
some residue in the syringe which was mixed with physiological saline and given
to the bird about 1 5 to 20 minutes after the original insemination.

VOICE. Did you give it three times in a row then for three days?

BERRY. No, if you got one of those papers, it shows that the maximum, the
optimum insemination occurs a day or two before each of the eggs. With prob-
ably any one of those inseminations the whole clutch would have been fertil-
ized.

THACKER. Did you do any motility work on the semen, any work like that?

BERRY. Yes, Motility with undiluted semen did not seem drastically altered
after three hours at room temperatures. After six hours motility obviously de-
1972 Breeding Conference— Artificial Insemination F93

creased and perhaps half the sperm no longer moved. At nine hours, 90% of the
sperm were immobile and the rest were not very active.

TEMPLE. I have got a lot of data on that.

BERRY. OK. Let Stan worry about the motility. This is the first chick hatch-
ing. The chicks were hatched in an incubator and given back to the adult female
to raise. At just one day of age here is the huge Goshawk feeding the tiny little
chick. She was allowed to incubate one of the eggs for 3 1 days. I gave her this
chick and took the egg away from her. The next chick was hatching three days
later and she was given the second chick, and six days after that the third chick
hatched. I allowed the third egg to remain in my study for several days hoping
to get her to recycle. So perhaps a rather bizarre technique. It’s something that
you certainly can’t practice with the ordinary raptor. But I do believe it can be
practiced with most any imprinted raptor that is properly conditioned to accept
you as mate. And I think probably the only requirement for this is total and
complete isolation from other raptors and of course a lot of communication be-
tween you and the bird during the normal process of maturation of the gonads.
Stan Temple, would you present the massage technique of artificial insemina-
tion?

TEMPLE. I think I’ll comment a little on my work with cooperative insemi-

nation This was done with Red-tailed Hawks. It’s the same basic type of
first.

procedure- there’s not much you can vary in the procedure. I would comment
on this imprinting technique. Imprinting in birds is accomplished very early in
their life. And when we use “imprinting,” we hear that the bird has become
fixed on some object during its early development in the nest; and it will come
to regard that object as a conspecific, something it should associate with. We do
this— falconers do this all the time— when we take a young eyass from the nest
and handraise it. The bird sees you and thinks of you as its parent. It becomes
imprinted to you, it thinks that you are the type of object that it should regard
as a parent image. The next thing, when a bird has regarded you as a parent
image-as long as they are kept away from other members of their same spe-
cies— the next step in this progression is for them to assume that you are a social
partner, and falconers do this by hunting with their birds, by manning them and
by hunting them. The bird cooperates in a social interaction, hunting, which is
a partnership between you and the bird. The next logical step from a social in-
teraction, when the bird becomes sexually mature, is a sexual interaction. And
this imprint is nothing unique to birds of prey; all birds will do this. Most of
you, I think, have probably read some of the popular accounts that have come
out on territoriality, like Konrad Lorenz’s On Aggression and so on. Ducks, for
instance, have a following response when they’re imprinted. You raise a duck
from hatching and it sees you and thinks you’re its mother, and it’ll follow you
around. These ducks, when they’re imprinted, will also do exactly the same
thing that we have happening with birds of prey— they’ll become socially and
F94 RAPTOR RESEARCH Vol. 6, Supp. F

sexually fixed on human beings.

I repeated essentially the same type of technique that Bob had. I had a male
that was 14 years old. He started copulating on my gloved hand, putting semen
on the glove when he was four years old. It was way back when I was in high
school. However, I was never able to get a female that would lay eggs— and of
course, this is one thing we should stress right off the bat in artificial insemina-
tion. Artificial insemination isn’t worth a darn unless you have a bird that has
fully functional gonads. So the male would copulate fine with the glove. We
could collect semen. I might say that the male Red-tail produced semen volumes
and frequencies that far exceeded Bob Berry’s Goshawk. I collected semen daily
and I had volumes that ranged from .1 up to almost .4 cc. This is a big drop, not
a little drop. I injected it daily into the female. The female responded the same
way to stroking her back. This female was a little different, she’d let anyone
stroke her. This may have been because Bob was the only person who handled
his Goshawk, whereas this bird was handled by many people. Two fertile eggs
were produced. One hatched by artificial incubation. The other was put under
a broody hen. I’ll have more about that later. So basically what I would like to
emphasize is that falconers as a group probably have many birds that, given the
right amount of handling, are likely candidates for this type of work with arti-
ficial insemination. Most of these birds have been taken as eyasses trained for
falconry and then retired. You saw Bob Berry handing a stick to the female. It
is very important that you go in there daily and help them build their nest,

stroke them. Be around so you strengthen the pair bond. The other thing that I
tried last spring was manual ejaculation of the male. I knew that my male Red-
tail was in full reproductive condition. Therefore, I thought it would be valu-

able to try a forced ejaculation technique. Now this massage technique is a little
deceptive using it on a bird such as this Red-tail that will ejaculate voluntarily,
because it doesn’t take much to get him to ejaculate. But I decided I would go
all the way and try doing just like you would have to do to a bird that wasn’t

going to ejaculate. I trained the bird so I could lift him up holding him by the
legs. Unlike Bob Berry’s bird I did get my bird right in the middle of produc-
tion. It didn’t seem to bother him at all. He had to be trained very much like
you train a Cooper’s Hawk to sit on the hand. The training technique involved
first holding his wings closed; forcing him down on my lap holding him there
;

for a few minutes and offering him a bit of food as I let him up. Much like
breaking a bird to a hood. After about a week and a half of this there was no
stress at all involved with picking him up right off the perch and holding him.
He just didn’t seem to mind it at all, once he had been broken in to it. The bird
was held by the legs; the cloaca was exposed, and the technique that we will
show you after we are finished here with chickens, because we don’t have any
raptors in breeding condition, worked very adequately on him. The technique
involved physically pulling the papilla out and what you do is strip or pull the
semen out of the very end of the sperm duct— the seminal glomulus. This is
where the sperm is stored before ejaculating. Using this technique, this forced
technique, I wasn’t able to get as large an ejaculate as I was with the voluntary
 1

1972 Breeding Conference— Artificial Insemination F95

technique. The ejaculates were much smaller. They ranged down to less than 0.
cc, more in the range of what Bob was able to get using the voluntary technique,
but was able to get perfectly viable semen that looked in every way healthy,
I

just likeit would in a normal ejaculate from a hawk. There is one other tech-

nique that can be used for collecting semen, and this involves the technique
called electro-ejaculation. This technique is used frequently with farm animals,
bulls and rams and hogs. It involves giving the bird an electric shock. What you
are trying to do is cause the muscles that squeeze the seminal glomulus to cause
an ejaculation. You are causing those muscles to contract violently and force an
ejaculation. You all know that you get a muscle spasm when you get shock-
ed. The machine that was used is right over here. Basically, it is a pair of elec-
trodes. You control the voltage and the amperage of the shock that you give
the bird. They found doing work of this type with chickens that approx-
imately 30 volts at a range of about 1/10 or lower amperage, given to the bird
at three second intervals, three to five times, three seconds off, three seconds
on, etc., usually stimulated an ejaculation. The electrodes are attached at two
points. One electrode is attached to the rump area just above where the testes
would be or just above the leg joint. The other electrode is a probe and the
probe is inserted right into the cloaca. Give him the shock. This is one draw-
back perhaps with the birds of prey. It does cause muscle spasms in the hip
region, but the bird seems to recover right away. My work has only been with
chickens using this technique so far. It seems to be a very good technique for
getting semen from a male that is in full reproductive condition but that would
not cooperate like our imprinted birds would. I don’t think it is going to be
necessary to apply this with birds of prey. But I present it here. I have circuit
diagrams for anyone who would like to build one and try it on some bird. You
can have a crack at it, and see what happens. I haven’t yet tried it.

SWARTZ. You found that placement of the cloacal electrode was not criti-

cal. I failed to succeed with an adapted ranger electrode stimulator. And I think
maybe Skip Walker and Jim unsuccessfully tried it with pigeons or discovered
that placement was critical.

TEMPLE. Placement is critical, yes. What I did was bypass the papilla. If I

had a blackboard, could make a diagram. The papilla is on the ventral side, the
I

lower side of the cloaca. You can think of the cloaca as being like a rubber
glove and the papilla rolling out like you sticking your finger through the rub-
ber glove. Sticking it out through the cuff, sort of like that, the papilla comes
out; you insert the electrode back almost into the rectum of the bird. It’s be-
hind the papilla so that the placement of the electrode isn’t hindering the pap-
illa from being exposed.

VOICE. Are you trying in essence to get it as close to the testes as you can?

TEMPLE. No, no. The idea is not to get it close to the testes. That’s not it at
all. You are trying for the muscles that control the seminal glomulus which is
the area of the sperm duct just before it gets to the cloaca.
F96 RAPTOR RESEARCH Vol. 6, Supp. F

SMYLIE. Do you use any kind of anesthetic while you are doing this?

TEMPLE. You can. If you put it on your own hand you can feel what it’s
like.When you get it in contact with the muscles as you are by clamping it,
you get muscle spasms. It’s not the type of thing that is . . .

SMYLIE. What about trapping a wild raptor. Anesthesia . . .

TEMPLE. I wouldn’t recommend it, no.

SMYLIE. I was just wondering if any experimenting has been done with birds
that are not tame.

TEMPLE. Yes. This technique is one way. They have used it with uncoopera-
tive and untrained birds. But the thing I’m saying is that even for an untrained
bird, you can get the bird to ejaculate using the massage technique that we will
demonstrate later on. In fact something that Fran Hamerstrom alluded to in a
littlenote in Raptor Research News it can be used effectively. The one other
,

thing that I’ve got some information on I think is rather unique information,
since it’s probably the only case where we’ve gotten enough semen from a bird
of prey to try to use a certain diluent for preserving the semen. Let me put this
slide on. I was able to get enough semen from a male Red-tail that it was worth-
while to try some very simple experiments on how to extend the life and to di-
lute the semen so it might be able to go around to between seven or eight more
birds. There are two— think of them as things you can dilute the semen with or
perhaps preservative— that are very easy for you to prepare. Most Extenders or
diluents that are used by the poultry industry are not the type of things that it
would be very easy for the average person to put together. To make up I tried
two that are very easy. I tried Ringer’s Solution which is a physiological saline
type solution, the mixture of which you can get out of any standard physiology
book. And one that has worked with chicken semen and turkey semen is steril-
ized whole milk. I diluted the semen in 1 1 dilutions— one part of semen to one
:

part of my diluent in each of these trials, and I compared it at two different

temperatures: ice box temperature about six degrees C and body temperature
at about 40 degrees C.
What I looked at here was how well the semen survived under these condi-
tions as compared to undiluted semen. Can these techniques be used perhaps to
extend the life of the semen sample so that we could ship semen around the
country to different people. What I did: The semen in a good ejaculate is very
dense, and the individual sperm cells are very active. It’s a very simple-minded
type of experiment. In a microscope field (under 430 power) I placed samples
of each of the different treatments described above. In each field of view I
counted 100 sperm cells. Out of the 100 I counted how many of these sperm
cells were dead, the ones that were not at all motile, the ones that were not
moving at all. I then did that, initially. The first column here is undiluted
 1972 Breeding Conference— Artificial Insemination F97

Table 1 . Semen Extension Experiments (after Temple).

Percentage of Dead Cells

0 hrs. 3 hrs. 6 hrs.

Undiluted at 6 C 0 21 30
Undiluted at 40 C 0 39 54

semen (Table 1). This is fresh semen that has just been collected. Zero hours of
collection I found no dead cells of the 100 cells counted three hours later un-
;

diluted semen at six degrees 21 at six hours 30. So that by six hours after col-
;

lection at six degrees about a third of the sperm cells are no longer functional.
At 40 degrees body temperature, this is much worse. The three samples were 0
at collection time, 39% at three hours, over 50% at the end of six hours 54%
were dead. Obviously, keeping it cool but not frozen is the best way to do it. I
also looked at sperm motility. What I did here is a very simple thing. I picked 10
random sperm cells in the microscope field and scored their motility-how fast
they were moving, how vigorously they were moving. I scored it 0, 1 or 2. Two
if it was very active; 1 if it was moderately active; 0 if it was practically inac-
tive. I then got a percent of what full activity would be for those ten. It’s 100%
right away after collection. It drops off to 40% activity, which is pretty low
really. If you get anything less than half of that motility, you might as well for-
get it. The activity is probably so low that the sperm is not active-even worse
with warm semen, 100% to start out with. It goes down to six hours with only
10% activity of the sperm cells. That’s what happens when you are using undi-
luted semen just as you collect it. If you dilute it with Ringer’s Solution, the
same thing occurs, generally. I think I won’t go through all these data. You see
that in both cases it is worse. Ringer’s Solution didn’t do the trick. It didn’t ex-
tend the life of anything. The sperm motility drops right off, much sharper than
it does with undiluted semen. Actually, approximately the same for warm se-

men, but this is a small sample size. There is only one sample, so you can’t really
say. Sterilized milk is merely milk that has been heated to destroy the lactic
acid in it, which is harmful to semen. And here the results are roughly compar-
able to the Ringer’s Solution. Still nowhere near as good as undiluted semen.
The only thing you can say is that at higher temperatures these diluents did
seem to preserve motility of the sperm a little bit better. OK, you can have the
light on now. I want to emphasize that it is probably not going to be very prac-
tical tosend or distribute avian semen, raptor semen, through the mail. In other
words, someone has a female in California that is laying eggs and someone has
if
a male in New York that they can collect semen from, it is probably not practi-
cal to send it to the fellow. Probably if you are going to get hawks together the
way to do it is to bring the two birds together somehow.

HUNTER. You didn’t try freezing it immediately in liquid nitrogen?

F98 RAPTOR RESEARCH Vol. 6, Supp. F

TEMPLE. I was advised by the poultry people that it would have been much
worse 'Freezing avian semen is not like mammalian semen where you can quick
freeze it and preserve it. Quick freezing avian semen for some reason is very,
very hard; it kills something like 90% of the sperm cells right off the bat.

HUNTER. One wonders why.

TEMPLE. Who knows. might say the literature on artificial insemination in

I

poultry is voluminous. People are just working on this. The publications are in
the hundreds every year. They are actively working on it with some of the wild-
est diluents you could ever imagine. Stuff that is carbonated and aerated. It’s
unbelievable what they have tried. So I might also say if any of you want to try
this massage technique, we are going to show it in a few minutes, on any of
your birds that you think might be in reproductive condition, there are a couple
of things that you can look at in the microscope to tell whether it’s a good se-
men sample. We’re going to show you a couple of these things.

BERRY. We are going to look at some of the semen under the microscope in
back after Dr. Morgan shows us the technique. If you’re done, Stan, thank you
very much. I think we have to hold questions, we just don’t have the time. Dr.
Morgan has a tight schedule. Fran, could you take the floor and tell us your ex-
periences with the eagles.

HAMERSTROM. Bob Berry asked me to talk on the massage technique. I

had never heard of We
always called it rough artificial insemination. I have
it.

successfully gotten semen from four species: Horned Owls, Red-tails, Golden
Eagles and Broad-winged Hawks. Some of these birds were prepared in advance.
Others were fresh caught wild birds. We just grabbed them and got semen. I’ll
tell you how one does this assuming you all know how to catch them in the first
place. You grab their legs. If it’s a Red-tail or smaller you put the legs between
your knees with the bird’s backup. Then you start stroking its back and putting
pressure on its belly until the tail comes up. Then you slip your fingers on eith-
er side of the vent. If it’s a male, out comes semen, if that male was just about
ready to copulate. With the Golden Eagle one prefers to have several assistants
[laughter] I tried it alone for quite a long time. And my eagle kept mounting
.

me and fortunately not my little hand. I would rip off my jacket and look at it
carefully to see if there was semen on it. I had some disadvantages that you
didn’t all have.
I would like to talk about some of the things that work up to semen produc-
tion. Youcan use a combination of these things. You can do all this coaxing at
the beginning and then if you need semen because you know your female is
about to lay and you can’t get it by the cooperative technique, why you may
wish to resort quickly to the massage technique. What leads up to this? In the
birds that build stick nests, it’s playing with sticks. Tease them. Don’t let them
have sticks all the time or you will lose your advantage. This works with owls,
and I don’t think it works with falcons. They don’t build stick nests. They are
1972 Breeding Conference— Artificial Insemination F99

not fascinated by sticks. I think they are fascinated by things like wings of
pigeons. I use my voice working with the eagle and neck stroking. My Red-tail

copulates first on my ankle and after that he much prefers my head. My Horned
Owl copulated ohly once and that was on my head at night [laughter] My diur-
.

nal raptors tend to copulate first thing in the morning. Then they’re keen on it
and then again late in the afternoon but their enthusiasm wanes near noon. With
the Golden Eagle you have to wait about 20 minutes between attempts to get
semen. If you fail the first time, go get coffee— spend 20 minutes doing some-
thing else and then go again. With rough artificial insemination I am using the
equivalent of that little box but it is cheaper. You just take the bird and you
keep stroking and you keep working and you get exhausted both emotionally
and physically, because you think you are being rough on the bird. And just
about the time you want to give up, the bird bates. That’s when the semen
comes, right at that bate. That’s when you keep going. This is the quick shot
treatment. When that bird has had enough of this monkey business and wants
to get away is when he is going to ejaculate semen. I’d like to clear up one thing.
I have a paper in Raptor Research News (5(3) 9 1 : 1971) on semen extenders—
,

a recipe for it. My purpose for semen extender was totally different. It was, if I
didn’t get enough semen, to extend the quantity rather than the time. Now if
we could show my slides, that would be very fine. I might mention that I have
a paper coming out in Die Vogelwarte on the male Red-tail’s capabilities. And
the Golden Eagle material is largely in my book. (I would like to give one bit of
advice to breeders. When you go in for any of these monkey-shines, take the
bird away from the breeding pen if you are going in for rough artificial insemi-
nation and then walk the bird back to the pen. The bird will forgive you in
about three paces while you are carrying it. There is nothing more soothing to a
bird than to be carried.) Could I show the slides, please. Here is my Golden
Eagle, Grendel. He is a little uncertain but it is early in the season. He is prepar-
ing to mount. You cannot imagine the conversation in the Hamerstrom house-
hold. This happened every day after breakfast. I would go out and get mounted.
I’m the very absent-minded type. Sometimes I would go running out in a little
thin sweater or something or other and my husband says, “Fran, did you re-
member your copulation jacket?” [laughter] The bird is really cued in on that
.

jacket. At any time of year I believe he will mount me or show considerable in-
terest if I have that on. Next. Here he comes flying. And eventually he gets up
onto my head. I think that’s next or there aren’t any more. I don’t know which.
Oh, yes, preparation for artificial insemination. I go and catch the bird. This
poor chap comes to our place, and he had never seen an eagle near to before and
he said, “Oh, I wouldn’t want to get near that;” and in about five minutes he
was helping because we were short-handed. Next. He is picking up the bird. This
is the female, and I always run away because I don’t want to spoil my beautiful

relationship with that female eagle, Chris. So I watch from the window. I may
be foolish but there is no point in taking a chance. Next. I’ve gotten five eggs
out of her in one clutch. And it brings me to a point I would like to make. Con-
sider the possibility of not just recycling. Consider egg stealing. My hunch is
birds of prey are far less determinate than we have supposed. And you may get
F100 RAPTOR RESEARCH Vol. 6, Supp. F

more eggs. Put the first eggs in an incubator or under something else. But con-
sider this possibility, too. Some experimental work needs to be done on it. Arti-
ficial insemination. Two professors from the poultry division of the University
of Wisconsin. always admire their courage. They handle eagles as though they
I

are turkeys. Until finally they saw my man get nailed. And then they had a very
different viewpoint. Next. This is about two minutes after a really rough go.
Two attempts at getting semen. The second was successful. Everybody waS ex-
hausted. And if you notice the eagle is perfectly placid. They get over this right
away. And so do my other birds. Next; good, that’s the end.

BERRY. Thank you, Fran. Can we now have Dr. Walter Morgan come up
here for a minute. He has hopefully a few comments or recommendations and
suggestions.

MORGAN. I’m happy to be with you today. Brookings is about 60 miles

north of here, and my schedule isn’t really so tight. There are a few comments
I’d like to make before the demonstration. Actually I don’t know if I am going
to stand on these chairs or what. I was sitting in the back row, and I am a little
sympathetic. It’s kind of difficult to see the demonstration; I am afraid it will
be quickly done, and we will have the result from it, and we won’t have gotten
enough information from it. First, we might ask when is artificial insemination
used. And we use it in poultry for special single sire mating when we want a par-
ticular male, a lot of progeny from that and known identity. We use it if we
have cages. If we have birds in cages which are breeding birds. Normally we had
floor matings for many years. Now as we use cages more, we do not mix the
sexes often. And we can use artificial insemination as a fine tool here for when
the sexes are separated. We can keep a pen of males. When you put a group of
males together after they are sexually mature normally they are going to start
fighting. But if you rear them together, the males will be harmonious and you
can take one male out, collect semen, and put him back in. Nothing will hap-
pen. There are more copies of this if anyone needs them. Another kind of arti-
ficial insemination that is used and commercially it is used very much now in
turkey production. Practically all of our turkey producers use this. The reason
for this is that the geneticists have done such a good job in breeding for big full
breasts and for short legs and for meaty legs that the male Tom turkeys are not
able to mount the hens any more. This is kind of ridiculous to breed for one
specific desire in our economy to the extent that natural matings are not likely.
The more successful breeders I think have the natural breeding plus artificial
insemination. Some of them use only artificial insemination. Considering when
it’s used, we consider when there is maturity. What Stan said was true about

having to have mature birds before you can collect any semen or before it is
worthwhile to inseminate them; at least they should be inseminated a short
period before they come into production. Now with turkeys for instance, the
pens of the males are lighted for about a month— they have artificial illumina-
tion. The day is lengthened artificially about a month before the semen is col-
lected. This way spermatogenesis is complete and mature sperms are produced.
1972 Breeding Conference— Artificial Insemination FI 01

With the females, about the same time is needed.

Why insemination used? One reason for using it is precision so

is artificial

that you know exactly what male has been mated with what female. I think
that is important to all of us here. The raptor breeders. For establishing geno-
types, for establishing what particular lines for selection you might want to
have. To overcome natural mating obstacles. And it seems to me that that is
what the whole bag of tricks is about here. And to conserve space by using arti-
ficial insemination if you have them, for instance, in cages.
As far as storageof the semen is concerned, the duration of potency for
chicken semen is normally about three or four hours, if it is in vitro. After you
have collected the semen you have two or three hours, three hours perhaps, to
inseminate. You can carry it around and move it if you want to. But as Stan in-
dicated, you cannot ship it long distances. We do not yet have means of preserv-
ing semen the way they can, for instance, bull semen, where they can use it a
year and a half, perhaps 20 years after. I don’t know how much longer, after
they have collected the semen. But we can only use it a very short period with
poultry semen. I think that the longest that it has been demonstrated is some
semen was sent from Maryland, College Park, over to England. They inseminat-
ed some hens there. But it was a very rapid trip in less than two days.
One of the cautions that I would mention to you, and this stems from some
work which I did in Belgium a couple years ago. I had an opportunity to work
quite a bit with semen motility at that time. First I mention that, in order to be
potent, good semen has to have good motility. But the opposite is not true.
If it has good motility, this does not mean that it is potent semen. I was inter-
ested in what dead semen was in your slides. A lot of us think of semen that
has lost its motility as dead semen. We had some irradiation studies over there.
When semen is irradiated with over 4,000 roentgens, semen that had been ejac-
ulated, it lost its potency but it had good motility. We had semen that was con-
trol semen that had not been treated that continued its motility for up to 10-12
days but it was not potent the first day after it had been collected. So you can
see motility, and it’s not necessarily a criterion for saying that it’s good semen.
Some of the cautions that We take with artificial insemination are that we have
to watch for the quick ejaculator. I think you have been exposed to that in the
discussion that we have had before. Sometimes after you have been training the
male, training the roosters for a while, you have them in the cage. You reach in
for them, if you happen to hit them the wrong way— it’s gone like that. You
haven’t even put your receptacles, your collector up there to get the semen.
Another caution would be cleanliness of your operation. Now I have distrib-
uted a page telling what the procedures are going to be. I also have here the
equipment that is necessary for artificial insemination. Normally we use a pair
of scissors, any pair of scissors, to cut the feathers off the rooster around the
vent on the abdomen. The reason for this and the only reason for this is visibil-
ity, so you can see what you are doing and can have a clear field to work with.
For our chickens this is a good type of collector to use. It’s a plastic container,
for turkeys and for chickens also. The normal amount of semen that we collect
from a well trained male will be up to one cc. We need approximately, or nor-
F102 RAPTOR RESEARCH Vol. 6, Supp. F

mally use, .2 cc for each insemination. So that if we have a good producer of

semen, we can take care of five hens without using an extender or extending
the semen. The semen in hens will be effective for a week. Now I don’t think
there are any cases with good semen where you need to inseminate more fre-
quently than a week. A week apart for inseminations is good. They can last up
to two weeks, possibly to four weeks. In the turkey the semen insemination
will last for practically the whole breeding period. So we have differences in
species. And this is something you are concerned with, but you have such a
small clutch with your raptors. I would suggest that a good insemination a day
or two before they start laying would last for the whole clutch. There is no rea-
son to be concerned about multiple inseminations. I’m saying this off the top
of my head without a lot of knowledge, but from the experience we have had
with chickens. The amount of semen that is inseminated is important because
you probably won’t have too little semen; you may have too little semen in
chickens. If you have 0.2 cc, mainly you have enough. It is protected and main-
tained within pockets which are in the oviduct. And the actual insemination
takes place as the egg is ovulated from the ovary as most of you know, as it
starts down the oviduct. And these little individual bits of semen are available
to fertilize if another egg is coming. So it is a matter of livability, if you want to
use that term— viability. I would rather use viability of the semen after it is in
the oviduct. In turkeys we find it lasts a long time. In chickens it doesn’t last
for such a long time. Now the tools that we use for collecting the semen. We
collect it in a small tube. After it has been collected (this is ABC on your hand-
out) we transfer the semen from the tube that we collect it in, into a syringe.
And normally what I do in practice is take out .2 ml for each injection insemi-
nation. because if you have .8 out and you try to inseminate .2 down to .6, it is
quite precarious because you have to try to look at it, it’s pretty difficult. If
you just have .2 in there and make a complete plunge you are all set. Those are
really the only instruments that you use. In addition to those today, if we can
we will set up a demonstration of motility and for that we use cover slips and
depression slides, and I have the slide here. It will probably be five or ten min-
utes after I finish the demonstration that we will have the sperm available to
look at the motility.

BERRY. Dr. Morgan, would you comment on temperature shock?

MORGAN. Temperature shock on semen?

BERRY. The outside temperature when you collect semen. Should you take
precautions before transferring it to the female?

MORGAN. No. I would reiterate that in storage the coldest semen retains its
motility much we used we had it refrigerated tem-
longer. In the samples that
perature. But retaining motility doesn’t mean retaining viability. They are more
heat susceptible than they are cold susceptible. I would make that comment
coming from body temperature they go to the cooler room temperature and
1972 Breeding Conference— Artificial Insemination F 03
1

this is fine. So I don’t think any precautions need to be taken there, if you do
it within a couple of hours.

THACKER. Dr. Morgan, would you recommend sterilized equipment?

MORGAN. I would, yes. I would be sympathetic toward the posture of the

veterinarians and I would say that sterilized equipment is highly desirable. I do

recommend it. I don’t follow it at all times. I’ve been fortunate not
In practice
to have problems. could use those chickens if somebody wants to bring those
I

boxes up and that lab coat that’s there.

BERRY. I was looking at semen under a microscope. Extended over a period

of hours, noticed that some sort of growth appeared in the semen. It was al-
I

most like they became too crowded to swim after a while. Something was grow-
ing in there. Was that bacteria?

MORGAN. There is a clumping of the semen which I’ve noticed, they clump
around the semen.

BERRY. They clump around that, what is that?

MORGAN. It’s some kind of starch artifact, I don’t know what it is.

HUNTER. Greg Thomas is going to take this on video tape. We are going to
kill two birds with one stone if you will excuse the expression, demonstrating
closed circuit TV.

MORGAN. What has been done previously to these birds— you recognize that
they do not have all their feathers. The first two here are trained males. I hope
they produce. They have been clipped. The third one I will clip and it hasn’t
been trained. I have less hope for good results with the others. Now you will
notice on your sheet that this can be done with two operators or one operator.
Frequently they have one person hold the bird and the other person stroke
it and collect the semen. I have come to do it as a single person operation, and

it normally works pretty well. The testes as you know are in this area of the

bird up near the back. Stroking of the back like so produces the ejaculate. Now
there is some fecal material there. And there is a sort of stripping of the duct
that I think Stan mentioned in some of his birds, too. Usually after you have
tried it three times and if you haven’t gotten anything, then you might as well
give up. He produced a pretty good supply of semen and there was some fecal
material in there as well. So this sample you would have to let the fecal material
sediment down to the bottom. And I will remove it with a syringe that I have
here. As I said, it is anticlimactic because it is all over. I’ll remove the semen.
Now that one produced about 40 hundredths cc of semen.
GRAHAM. How much pressure did you use on that?
FI 04 RAPTOR RESEARCH Vol. 6, Supp. F

MORGAN. I put a lot of pressure.

TEMPLE. I have an untrained rooster that any of you can practice with.

HUNTER. I have 300 of them.

MORGAN. OK, there are some things, the first was without telling many of
the particulars about the technique. I think with your raptors you’ll have dif-
ferent types of reactions than we would have here. Reactions in terms of meth-
ods of holding them. For this I put my little finger between the legs, hold the
legs and have the container near the cloaca. Now, actually, I stroke him two or
three times, and then I push down; and I’m kind of pushing up on the abdomen,
too. If you can watch and see that perhaps, stroke him like that. Now that is
fecal matter, that’s not semen. I hope you didn’t get a picture of that.

VOICE. Is this a trained bird again?

MORGAN. Yes, this is one of the two trained ones. There is quite a bit of
fecal material but there is a good supply of semen, too. We’ll set this semen up
under the microscope.
I will demonstrate now the insemination into a hen. And for that purpose I

will return some of the semen into the container. It is a very simple process. The
semen is not as clear as I would like it to be. So whether it’s our personal rela-
tionship I’m not sure. Now this rooster has all of its feathers, you see. He hasn’t
been clipped or anything. It’s possible to collect semen without clipping them,
but I like to clip the feathers off. Whoops [laughter] It is a little bloody. There
.

is a little bit of semen, there is a very little bit. But there is some. This is norm-

ally what we would expect. You can see where it drained down the sides here.
We would expect to get a little bit the first time and an increase with daily
trials. I’ll set up some of this semen.

TEMPLE. While you have that chicken, there is a technique that I use on
other birds such as pigeons. This seems to work a little better for me, a little
better to have someone hold it and hold the tail up and massage the back. Then
for actually stripping the bird that is going to be very uncooperative, like for
instance a hawk that’s not imprinted. Using the other hand and approaching
ventrally instead of dorsally is to reach in until you can feel the end of the pa-
pilla and bringing it out; he has just ejaculated and just sort of letting your fin-
gers slide along. I don’t know, for me that seems to have gotten more from an
untrained bird than just the stroking of it. You might want to comment on
that technique. Is that common to pull the papilla out?

MORGAN. The chicken has about as little papilla as any of your raptors
would. They are practically without it, too. If you have a duck or goose, they
have a nice long one. You can really strip that. There is not much to strip a rap-
tor. And I think there isn’t much to stripping chickens.
1972 Breeding Conference— Artificial Insemination F105

TEMPLE. It’s small, pulling it out.

MORGAN. I put it in my
notes here-manipulation is stripping of the duct,
so we have that in common.
would be optimistic about these techniques with
I

your birds particularly after hearing what we’ve heard with previous two speak-
ers. It’s a really simple technique. It’s something that you work on— well, if you
know when the female is going to start laying, you would start on your male a
week to 10 days or maybe two weeks before that. Then if you inseminate, if
you’re able to inseminate her, just before she starts laying, you are through in
two weeks. So it’s not a long tedious process. And if you can collect semen, the
insemination of the female is extremely easy.

GRAHAM. Would you explain in more detail just the pressure you used and
exactly where when you stripped.

MORGAN. With the hens there is quite frequently, there usually is, an ex-

pulsion of fecal material. Now, for holding the hen normally, I rest her on my
thigh like this, too, and her feet will go right on my thigh. And I will press down
on her back when I get ready because
I all my hands to manipulate
will need
here.The hen has mentioned previously, an oviduct. With the chickens the
as
oviduct everts. There is quite a large opening. It is just a matter of putting the
tube into the everted oviduct. That’s mentioned on your sheet; it’s really men-
tioned there better than I can say now because you relax after you have insert-
ed the syringe. And when the hen has relaxed, I push the plunger. And in all
this we also realize the sensitivity of the birds we are working with and we are
trying to avoid stress. I think I use the word effort in the first line there; several

places I said without too much effort, or without much effort. Without stress, I
am trying to say. I encase her head under my arm and go in like this. There is
some fecal material there which is interfering with the oviduct. There are two
openings. You see there is an oviduct and there is also an anal opening. You
have to insert it into the oviduct. We have had some chickens in Brookings
which had two oviducts. This is a genetic difference where they have a right
and left oviduct. Very unusual. Normally, they just have the left. If they have
two oviducts and we have inseminated into the right oviduct, we have never
had any fertility. So it should be into the left functional oviduct. And I noticed
you said you went on the left direction when you were into the raptors so this
I would expect, so this is as it should be. If anyone else would like to practice

with these, why they are welcome to, I consider it kind of a challenge for Don
saying that he wanted one that was untrained and I didn’t know they were go-
ing to bring one; so I have one that is untrained, or I’ll take his.
In the training period these have been trained for about a week. I have a stu-
dent who works with me on this. And
the first time that he tried it, there were
1 5 roosters, and four of them ejaculated semen. So it is not unusual to ejaculate
the first time, to getsemen the first time.

EBERLY. How do you train them?

FI 06 RAPTOR RESEARCH Vol. 6, Supp. F

MORGAN. Just by trying to collect semen from them. This is what the train-
ing would expect this would be true in raptors. You would have to train
is. I

them. have heard several people speak of the very precise and close association
I

people have with raptors. Perhaps this is a necessary part of it. But at least with
chickens any of us who try it can usually get results if we use the proper tech-
nique.
OK, I strip this male, Eve got the tube in my hand so make it all realistic.
as
I have him kind of sitting down like that just like the hen was sitting
like to
down. Then I can bring pressure to bear up here and with this hand I am push-
ing down here. I am pushing on the abdomen and with these two fingers I am
stripping in here There is quite a bit of pressure that is given in the excitement.
Here I think one of the dangers I might be suggesting, that we might be reluct-
ant to squeeze them too much. You do squeeze them quite a bit. I’ve found
with students that they think maybe I am hurting the bird.

VOICE. As hard as a hand shake? [laughter]

MORGAN. Yes, at least.

VOICE. When you get a small amount of semen coming down the side, do
you use a dilution material to get that out?

MORGAN. This I let settle for quite a while. This will eventually settle, may-
be after two minutes. Then I try to pull what I can up into the syringe. I haven’t
diluted it. We don’t practice dilution. I’m for dilution. I think it’s OK. But we
haven’t needed to use dilution.

VOICE. You get just that little bit that you got out of that first ejaculation.
Will that be enough to draw up?

MORGAN. No, on this one I didn’t have enough.

VOICE. But that’s just about as much as you are going to get out of a small
raptor.

TEMPLE. Look at a Red-tailed Hawk, a Red-tail gives as much. Bob, you

have never stripped your bird to see whether you could get more out.

BERRY. No.

VOICE. Is it too viscous to come up in capillary tubes?

TEMPLE. doing this on Starlings which are roughly comparable to

I tried
I could bring the papilla out and get an ejaculation. And that
kestrels as to size.
is what I used in capillary tubes. I just touched it and it was sucked up into the

capillary tube. I’m almost certain that if you have a raptor in full reproductive
 .

1972 Breeding Conference— Artificial Insemination F107

condition using the stripping technique, you should get plenty. I don’t think
you have to worry about not getting enough.

VOICE. What about [inaudible]

TEMPLE. I’ve used a pipette for the insemination; I didn’t use a syringe, I

used a pipette.

HUNTER. For whatever it is worth, about four years ago he told me that
Pheasants were very difficult. I didn’t happen to have any chickens at home,
but I did have some Pheasants. It was breeding season. A cock pheasant was ob-
viously in breeding condition. I went over and picked him up, got semen from
him immediately, it surprised me, quite easily. I never really tried it on chickens.

MORGAN. Some people use a

tube to get a greater distance in inser-
little

tion; this is particularly true in turkeys, where there

is a longer oviduct distance

and the sperm really has to go down to the area of the ovary, but usually the
sperm is potent enough to do it without this assistance. I am finished then, un-
less there are any questions

GRAHAM. What about injection of fecal material with the sperm?

MORGAN. We like to be cautious and avoid that but if you do, that will not
kill the sperm.

VOICE, [inaudible] bird that is laying eggs already and then you inseminate?

MORGAN. OK. is about when there is a delay in when you in-

The question
seminate, and don’t give the answer to this, come back to it. With laying
if I
hens particularly if there is an egg in the oviduct, this egg will not be fertilized,
but the egg that she lays day after tomorrow will be fertilized. The most desir-
able time to do this in chickens is right after they have laid an egg, because their
oviduct is free then. It takes about 24 hours for a new egg to come through. If
there is an egg in the oviduct, this does not prevent the sperm from going up
though. So it can be done when there is an egg in there. If you do make an in-
semination today and your raptor lays an egg tomorrow, I don’t think the egg
is going to be fertile.

VOICE. You mentioned that if you can feel the egg in the bird, it is already
in the oviduct, right?

MORGAN. Somebody mentioned that.

BERRY. I think I mentioned that. If you can feel an egg in the oviduct, you
could probably get that egg. That’s before the shell. You can feel it enlarge long
before the shell is laid down around the soft part of the egg. So you can get that
F108 RAPTOR RESEARCH Vol. 6, Supp. F

egg after you feel it.

MORGAN. is ovulated, it’s the yolk which is ovulat-

In chickens after the egg
ed, the actual size going to be when it’s laid. After the yolk is ovulated and
it’s

has started down the oviduct, it does not become fertile at any time. In chick-
ens we can palpate the egg. You can feel in the cloaca. If there is a hard shelled
egg, you know it right now. And that egg will never be fertilized by an insemi-
nation today.

MENG. Have you experimented with insemination after an egg has been laid?
Drilling a small hole in the shell and putting the sperm in.

MORGAN. I haven’t and I think it’s impossible.

TEMPLE. Very difficult. People have tried to with chickens and it’s very,
very difficult to do. What happens is the membrane that’s laid down around the
yolk as it goes down through the oviduct is very difficult.

BERRY. I think it has been done though.

VOICE. A fellow out in California is doing a lot of work on it right now.

TEMPLE. It has been done. Your chances are pretty slim. I might add, when
isthe right time to start insemination if you have a hen that is going to lay eggs?
On the female Red-tail you can feel the pubic symphysis which is where the
two halves of the pelvic girdle come together. It’s between the legs almost to
the vent. That separates and widens as the female comes to the time when she
is going to lay an egg. This is a real easy thing if you have handled your bird
when she is not in reproductive condition— you can feel those two bones very
close together. When she comes into reproductive condition and approaches the
time she is going to ovulate (in the female Red-tail it is about a week before she
ovulated) those bones come really wide apart. In the female Red-tail you could
put three fingers in between the halves of the pubis. Really opens right up, but
you feel it coming apart. Once you can feel it coming apart that is probably the
time you want to start your insemination. That is a sure sign she is going to be
laying real soon.

APPENDIX: Summary Issued at Conference

PRACTICAL CONSIDERATIONS IN ARTIFICIAL INSEMINATION

by Robert B. Berry

Anatomy. Female: In most hawks, both ovaries are functional but only the
left oviduct is functional. Anatomically, the oviduct is comprised of five parts:
1972 Breeding Conference— Artificial Insemination FI 09

the infundibulum (fertilization occurs here), the magnum, the isthmus, the shell
gland and the vagina.
Male: The genital tract consists of paired testes, epididymides, deferent ducts
and in hawks, a rudimentary copulatory organ called a papillae. Sperm is stored
at the base of the deferent ducts (Petrak, 1 969).

Artificial Insemination.
Advantages Disadvantages
1 . Improved fertility. 1. Stress.
2. Cross breeding. 2. Urates reduce viability of sperm.
3. Reduction in males. 3. Requires a skilled operator.
4. Determine potency in individuals

Semen Production.
1. Vitamin A and E deficiency affect spermatogenesis adversely.
2. Coccidiostat at 0.0125% in feed does not affect semen production.
3. While semen can generally be taken from a male fowl at any time of year,
one cannot expect similar results from a raptor since maturation of the
gonads is a cyclic occurrence (they may expand 200 to 500 times their
size during the mating season).
4. Greater semen yields occur in the late afternoon— domestic fowl.
5. The greater the frequency, the lower the yield.

Collection of Semen.
1. Optimum response is secured from a fowl if he is picked up quickly and
immediately manipulated— ostensibly before surprise turns to fear.
2. Do not allow semen to cool quickly, even though fowl semen is more resis-
tant to temperature shock than the semen of other domestic animals.
3. The best fertility is achieved within one hour after collection.
4. Maximum fertility in a chicken requires insemination every seven days.
5. Duration of fertility in a chicken is 10-13 days.

Semen Quality.
1. with motility, percentage of dead sperm,
Fertility appears to be correlated
methylene blue reduction time and numbers of live sperm per unit volume
of semen.
2. Fertility does not appear to be correlated with pH, density of sperm and
volume of ejaculation.

Semen Quantity.
1 . Chicken (large) 1 to 4 cc.
2. Turkey 0.3 to 1 cc.
3. Goshawk 0.01 to 0.05 cc.
4. Red-tailed Hawk .10 to .40 cc.
 .

FI 10 RAPTOR RESEARCH Vol. 6, Supp. F

Dilution and Storage.

1 Many attempts have been made to arrive at a diluent medium and ideal tem-
perature and other physical conditions for the prolonged storage of semen,
all without success. Storage beyond 48 hours produces such reduced fer-

tility as to be impractical.

Insemination.
1 Optimum time to inseminate is just following oviposition (egg laying).
.

2. Fertilization will generally occur the second day following insemination.

3. Maximum fertility in chickens requires insemination every seven days.

4. Duration of fertility in chickens— 10 to 13 days.
5. A hard-shelled egg in the shell gland reduces the chances of good fertility for
other eggs (easily discovered by palpating the abdomen just behind the
breastbone).

Is artificial insemination the panacea we are looking for in captive raptor re-

production? Certainly not! Artificial insemination should be attempted when

all else has failed, and then probably only when intraspecific hostility endangers

the life of one of a pair. Maturation of the gonads must take place before sperm
or egg production is possible— a complex and lengthy process brought about by
a combination of external stimuli, including the weather, photoperiod, terri-
tory, absence of stress, stimulation by a mate. Therefore, if a pair of birds is
physiologically capable of mating successfully, a prerequisite for artificial in-
semination, then they should be allowed to do so. Exceptions might involve a
laying female with a displaying male that appears on the threshold of copula-
tion, but for some unknown reason, cannot take that last step. Extreme care,
however, must be taken in the introduction of stress which might destroy the
reproductive cycle.

Reference Material

Burrows, W. H. 1939. “Artificial Insemination of Chickens and Turkeys.” U. S.

Dept, of Agriculture Circular No. 525.
Perry, E. J. 1968. The Artificial Insemination of Farm Animals. New Bruns-
wick, N. J.: Rutgers University Press, pp. 258-300.
Petrak, M. L. 1969. Diseases of Cage and Aviary Birds. Philadelphia: Lea and
Febiger, pp. 95 and 322.
Marshall, A. J. Biology and Comparative Physiology of Birds. 1961, Volume II.
New York and London: Academic Press, pp. 120-122.
Maule, J. P. 1962. Die Semen of Animals and Artificial Insemination. Technical
Communication No. 5 of the Commonwealth Bureau of Animal Breed-
1

ing and Genetics, Edinborough. Published by Commonwealth Agricultural

Bureau, Aberdeen.
RAPTOR RESEARCH
Volume 6 Supplement Part G 1972

SPECIAL CONFERENCE ON CAPTIVITY

BREEDING OF RAPTORS-A REPORT

Sponsored by Raptor Research Foundation Inc. ,

Held at Sioux Falls, South Dakota U.S.A., November 21-23, 1971.

,

Part G. Breeding Stock Inventory (Panel 1)

edited by
Donald V. Hunter, Jr.
Rural Route
Centerville, South Dakota 57014

Panel Members: Donald V. Hunter, Jr., Chairman, Robert Hinckley, Roger

Thacker, and Bruce R. Wolhuter.

HUNTER. The subject of Breeding Stock Inventory is a general topic that we

chose as a starting subject. First of all, I think everyone is interested at least to
some degree, in what everyone else has done. But to have each person come up
here and describe his breeding project would take our full two days, so we have
asked that questionnaires be filled out and sent in. As many as came in on time
have been duplicated and are in the packet. The first panelist will be Roger
Thacker.

THACKER. The paper that you have got in your folder there, I finished
approximately three months ago, entitled Estimation of Birds of Prey in Cap-
tivity in the United States of America. It is, to put it quite bluntly, now com-
pletely out of date because inventories changed so quickly. It is a piece of work
that was completed over approximately 18 months of sending out surveys to
various organizations and I think in the few minutes that I’ve got to go through
it, it would be best to start from the beginning and just work straight through

the paper. [Thacker then went through the paper, which is printed in Raptor
Research News 5(4): 108-1 22, 1971.]

HUNTER. Thank you very much, Roger. First of all I’d like to stress the
importance of finding out how many birds of prey are in captivity other than

Gill Raptor Research 6(Supp.):Gl 1 1-G1 15, 1972

G1 1 RAPTOR RESEARCH VoE 6, Supp. G
the Peregrines that most of us are interested in right now. I think it is quite ob-
vious that through correllary experiments, knowledge that we gain from other
birds of prey may help us in those that are in danger at the moment. From the
Lincoln Park Zoo we have Mr. Robert Hinckley who will say a few words to us
about the availability of birds in zoos and what the general attitude of people in
zoos is as far as doing research in captivity breeding.

HINCKLEY. I’ll speak about our zoo to begin with. We don’t keep birds as

singles at our zoo where it is physically possible to get mated pairs. We don’t
accept contributions of single birds at our zoo, we accept only mated pairs and
we don’t display single birds at our zoo. We display only mated pairs wherever
it is physically possible to have them. That’s our position.

We have outdoor flight cages for large birds. We have a pair of Golden Eagles
that have laid eggs for two years and the male has copulated with the female
for two years. These birds are on display and we have good hopes for this
spring. We have a new pair of Bald Eagles and White-headed Eagles, certainly of
breeding age. We have high hopes for them. We have a pair of Andean Condors
both in adult plumage and they have nesting facilities available. We have a pair
of Griffon Vultures with nesting facilities available. We have a pair of Verreaux’s
Eagle Owls with nesting facilities available.
The problems of displaying birds at zoos go into several categories. The first
is, after all, we do have birds on display, and that is one of the primary func-

tions of the zoo. One of the next problems with display is security. We can dis-
play Golden Eagles and we don’t worry about people sneaking in overnight and
stealing them. We don’t have any doubt, however, that if we put a pair of Gyr-
falcons on display in a breeding project that within a week they’d be stolen. So
one of the very legitimate problems that any zoo has is that you cannot keep
attractive, exotic birds of prey and expect to keep them, so we don’t attempt to
keep those kinds of birds. The next problemthe zoo has is the expense of pro-
viding outdoor Right cages. There no such thing as taking a few posts and
is

some chicken wire and for 60 bucks putting up a display at a zoo. You have to
get bids through the park district and then you have to have union people put
the displays up and then you’re talking about thousands of dollars just for cy-
clone fence. So it’s not an easy project from that view. One of the major stum-
bling blocks without any question is knowledge and the competence about birds
and this is a problem to which this group could readily address itself. Another
one of the problems on setting up breeding pairs is the Fish and Wildlife people.
We tried for three years unsuccessfully to get an immature male Bald Eagle to
go with our immature female Bald Eagle and we still don’t have one. There is
no question that there is a good deal that Raptor Research Foundation could
do to assist zoological parks. One would be some kind of information bulletin,
just on the basics of getting your birds to breed. Some people don’t realize it’s
wrong to keep several species in the same flight cage. A consultation committee
might legitimately be set up through Raptor Research to assist zoos with breed-
ing projects. It might be possible to set up some kind of breeding award for
1972 Breeding Conference— Breeding Stock G113

zoos, since the public in general does not always recognize how much time and
effort and endeavor on many people’s part went into successful breeding in
front of the public. It might be well to offer some kind of recognition to those
people who have succeeded, such as the Topeka Zoo with Golden Eagles nest-
ing this year.

HUNTER. Thank you very much, Bob. We have one more panelist who would
like to say a few words, Bruce Wolhuter from the Cheyenne Mountain Zoo in
Colorado Springs.

WOLHUTER. I think the other zoo people here really expect a rebuttal, but
I have to agree with everything that’s been said. In the zoos we are woefully
ignorant, for one, on how to take care of birds, and two, as Thacker pointed
out, some of them don’t even know what the difference is between the raptors.
Zoos basically have several problems; one is that for the most part, zoos are dis-
play oriented and they’re more interested in displaying the birds, the largest
number of birds, without thought to conditions or what they’re putting in with
what. And is disastrous if you’re trying to do any kind of breed-
of course this
ing at would say, well over 90% of zoos I’m familiar with have at least one
all. I

or two exhibits where they have to have at least six or seven species put to-
gether. And I think most of you realize this pretty well precludes any chances
of breeding. If you want to have birds that are breeding it’s necessary, first of
all to start out with a pair and some zoos don’t know whether they have a pair

or not. When I went to the Cheyenne Zoo (I started about a month ago), they
had about six female Kestrels in a cage. Obviously it’s going to be kind of a
rough breeding project unless they can change their ways on that. And those are
the birds that are easier to sex. So, you’ve got to get qualified people that can
sex ,a bird to the point to know whether they have a pair or not. And then you

have to isolate the birds, meaning not mixing them with other species. You have
to set up conditions that will at least encourage breeding and a lot of zoos will
fight this, depending on whether the zoo is run by the City Park Commission or
by its own zoological society, but there is a need to get these breeding condi-
tions set up. There’s a question in my mind as to the value of removing these
birds from public view during the breeding, whether this public contact is detri-
mental or not. It remains to be seen; some zoos have bred birds where the pub-
lic has complete access. I know Topeka Zoo with their Golden Eagles, this par-

ticular year was the year that they didn’t really do that much work with the
eagles and the first thing they knew, they had eggs. As soon as they got the eggs
they isolated the cage, whereas in the past they isolated the cage entirely all the
way through as soon as the courtship activity was started. And something Bob
mentioned, the problem in zoos of singles. So many zoos say well, if you’ve
seen one you’ve seen them all; we’ll just put one on display; it’s too expensive
to get more. I think this is one thing where the zoos can work together, bv
cooperation. Bob mentioned that they have a pair of Andean Condors and
G114 RAPTOR RESEARCH Vol. 6, Supp. G

there are several zoos who, just for example, have just one Andean Condor and
I think it would be great if they could get together, you know, loaning birds,
strictly for the purpose of breeding.
Let me just mention that the majority of zoos are not feeding the birds food
that, in my mind, is even good for maintaining them in healthy condition, and
it certainly isn’t doing much to encourage breeding.

HUNTER. Thank you very much, Bruce. We will entertain a few questions
from the floor now.encourage We
participation as much as you would like. I
think Joe has his hand up first back there.

PLATT. Yes, Mr. Thacker, the idea of tighter restrictions, has this gone
through? As far as impounding birds?

THACKER. Yes, it went through, yes.

WOLHUTER. I want to make a comment— you're asking about the importa-

tion of birds. They’re really tighteningup controls; right now it’s impossible.
They’re particularly watching the falconiforms and also the strigiforms and any
more if you import a bird, first you have got to have papers from the country
which it was imported from, saying it was taken legally. Otherwise, no soap.
Also this has to be cleared with our government before you can bring them in.
Eve also seen a rough draft of legislation that’s being prepared something . . .

like the endangered bird, a group that’s interested in this. And they’re trying
to tighten it up even further in such a way that there will be complete control.
So I think the era of bringing birds in for pets is about over. What effect this
will have on bringing birds in for breeding purposes or anything else remains to
be seen. But it’s going to get tougher and think it’s probably a good thing as
I

far as getting rid of the commercialism.

THACKER. The other tiling you have got to remember on importation of

birds is as I understand it, when they’re brought in and they’re presented at the
dock or wherever they’re coming, in, the person who inspects that bird is not

somebody who oriented towards them, since he’s a customs official, and if he
is

doesn’t have an interest in birds of prey he could be looking at a bantam chick-

en. Now this is serious. He has no idea what’s coming in. If the guy tells him,
well it’s a red-footed fighting cock, he’s going to put down on the paper a red-
footed fighting cock in many cases. I’ve seen lots of these papers come through.
But I do believe the bureau will be sending people down to instruct these cus-
toms people in the falconiforms and the strigiforms.

HINCKLEY. There is one more point is today perhaps as much as

here, there
20% chance that some time in the next twelve months there will be a total ban
on importing all avian species into the United States because of Newcastle’s
disease.
1972 Breeding Conference— Breeding Stock G115

THOMAS. Roger, about what you just said. I can remember last year at the
NAFA convention someone brought up the point that the federal people had
five or six different stations around the country that all the birds of prey or
anything like that that could remotely resemble it had to go through those sta-
tions and these people . . .

THACKER. They are on call ... the customs people, as I understand it, if
they get something they do not understand, they can get on the telephone and
say, come down and take a look at it.

SCHUBERT. I have a point about what Mr. Hinckley was saying. 1 think
Raptor Research should be more in touch with zoos. I used to work at a zoo.
We oftentimes got raptors in that nobody wanted. We had too many Red-tails;
we tried to trade them off with other zoos, but nobody wants Red-tails . . .

just put into a cage until they die, or else overcrowded, 10 or 15 Red-tails in a
cage. I think Raptor Research could be in touch with these zoos and with other
zoos to provide more information service where they can send those they don’t
want Right now we have a Crane Hawk we don’t want; if anyone here wants a
Crane Hawk they can have it; they don’t know what to do with it. They have a
Krider’s Hawk that isn’t doing well in captivity, they want to send it out west.
I think there definitely are lots of raptors in zoos.

HUNTER. Thank you very much. And

as far as Raptor Research is concerned
we’d be very happy to do of thing and we would accept volunteers.
this sort
Any more questions? I think maybe we’ve gone over this. We’ve got lots of very
interesting ground to cover and unless there are any more questions we will
close this and go on to the next panel.
RAPTOR RESEARCH
Volume 6 Supplement Purl H 1972

SPECIAL CONFERENCE ON CAPTIVITY

BREEDING OF RAPTORS-A REPORT

Sponsored by Raptor Research Foundation Inc. ,

Held at Sioux balls South Dakota, U.S.A., November 21 -2d,

, 0 71
Part H. Health and Nutrition of Young (Panel 10)

edited by
Richard D. Porter
Patuxent Wildlife Research Center
U. S. Bureau of Sport Fisheries and Wildlife
Laurel, Maryland 20810

Panel Members: Richard D. Porter, Chairman, William H. Halliwell, and David

E. Allen; contribution of John Serafin was presented by R. D. Porter.

PORTER. Health and nutrition of young raptors or nestling raptors is a poor-

ly known field of investigation. Although there is a considerable amount of in-
formation oh the nutrient requirements of young chickens, there is not much
known on this subject as regards young raptors.
Although Dr. John Serafin, the nutritionalist for the Rare and Endangered
Program at Patuxent, was unable to attend, he authored the following discus-
sion on nutrition of young poultry which has a degree of applicability to all
young birds in process of rapid growth and development.

PRINCIPLES OF NUTRITION, by John Serafin*

Nutrition is the process of assimilating food. The chemical reactions that

cause the metabilie functions of the animal to occur are supplied with raw ma-

Nutrition is the process of assimilating food. The chemical reactions that

cause the metabolic functions of the animal to occur are supplied with raw ma-
terial from the environment, through processes lumped under the term “nutri-
tion." This definition includes the ingestion, digestion and absorption of the
chemical elements which make up food, and the distribution of these nutrients

*Patuxent Wildlife Research Center, U. S. Bureau of Sport Fisheries and Wild-

Rare and Endangered Program.
life,

HI 17 Raptor Research 6(Supp.):H 1 1 7-H 28, 1972

1
HI 18 RAPTOR RESEARCH Vol. 6, Supp. H

within the animal organism in the proper form for use by the cells.
The complexity of the chemical substances that an animal must obtain from
his environment varies widely throughout the animal kingdom. Some unicellu-
lar organisms have very simple nutritional needs, mainly some inorganic ele-
ments, a source of carbon and energy and a few nitrogen compounds. The bird
has very complex nutritional needs with over forty specific compounds, classes
of compounds, or specific elements required from the environment for the in-
ternal chemistry of this animal to function properly. Even this is a relatively
simple list of raw materials from which to build the myriad of compounds pre-
sent in the animal body.
Because of the economic importance of the chicken for food through the
production of meat and eggs, much effort has been expended to determine ade-
quately the nutritional needs of this animal. Because the young chick is an ex-
cellent experimental animal for fundamental nutritional studies, more informa-
tion is available about the nutrition of the chicken than any other animal today.
This reservoir of nutritional knowledge has placed the formulation of poultry
feeds on a scientific basis and made possible scientific formulation of diets for
other species of birds as well. Today feeds for many species of birds in zoos,
birds captured- from the wild and ornamental and show birds can be successfully
reared using scientifically formulated diets.

Qualitative Nutritional Needs

These needs of birds can be divided into categories. These are:

1. Sources of energy. Many types of compounds can supply energy to birds

but the primary energy sources are classes of compounds called fats and carbo-
hydrates. Not all carbohydrates and fats can be digested by birds, so that only
carbohydrates and fats that can be digested and absorbed by the digestive tract
can be considered nutrients.
2. Amino acids. Proteins of the animal body are really polymers of some 22
amino acids that are arranged in an almost infinite variety of combinations to
produce the specific proteins of the structural and enzymatic systems of the
body. Some of these amino acids can be synthesized by the bird from simpler
compounds, but others must be included in the diet of the bird.
3. Vitamins. This term encompasses a group of compounds widely differing
in chemical structure. They are all required links in the chain of metabolic reac-
tions that occurs in the animal body. The chief properties that set vitamins
apart are the minute amounts in which they are required, and the types of meta-
bolic functions they perform.
4. Inorganic chemical elements. Many individual chemical elements are re-
quired for proper metabolic functioning of the bird. Amounts required and
functions of these elements vary tremendously from the large amounts of cal-
cium required for bone structure and egg shells, to infinitesimal amounts of
selenium required for presently unknown functions.
5. Water and oxygen. No description of the chemical relationship of an ani-

mal with its environment would be complete without mention of these two
1972 Breeding Conference— Health and Nutrition of Young HI 19

very important components. Life is not possible without water and oxygen.

Water forms the medium in which body chemistry functions and oxygen is re-
quired for the fire which releases energy from the foodstuff.

Quantitative Nutritional Needs

Although it is possible to define the qualitative nutritional needs of an ani-

mal in terms of individual chemical compounds, elements or classes of com-
pounds, the quantitative nature of nutrition is not so simple. The amounts of a
nutrient required in the diet depend on many factors. For example, the require-
ments for amino acids vary with the total amino acid content of the diet and
also some amino acids specifically influence requirements of others. The satu-
rated fatty acids content of the diet may influence the linoleic acid needs of the
chick. Pantothenic acid and vitamin B 1 2 requirements may vary with levels of
each. The phosphorus content of diets, particularly for young, rapidly growing
birds is dependent upon the calcium level in the feed and excesses or deficient
amounts of one or the other can bring about severe skeletal deformities. As
a percentage of the diet, most nutrient requirements vary with energy level.
Chickens tend to regulate feed intake to a certain caloric consumption. From
available evidence this seems to hold for other species as well. In high energy
diets less feed is required to meet the energy needs than in diets with low ener-
gy concentration. Therefore a specific daily intake of a protein requires less
protein as a percentage of the diet with a low energy concentration than with a
higher energy concentration.
Ideally there is undoubtedly a specific quantitative combination of amino
acids to give the most efficient conversion of dietary protein to body protein or
egg protein. There is a specific relationship between amino acid levels, fat levels
and carbohydrate levels to give maximum utilization of dietary energy for pro-
ductive purposes with a minimum loss of heat.
Palatability and nutritional adequacy also are interrelated. Palatability is a
rather ill-defined term that usually means that consumption of a given diet or
substance by a bird is affected by taste, physical factors, and any other term
usually felt to be unrelated to the nutritional addquacy of the diet. Through the
years, increased research has shown that nutritional adequacy of a diet influ-
ences voluntary feed consumption far more than certain vaguely defined pala-
tability factors. Diets in which pure amino acids supplied the protein needs of
the chick used to be considered “unpalatable” for chicks because chicks would
only consume small amounts of these diets. However, as research discovered
better combinations of these amino acids, feed consumption improved. A diet
with good quantitative balance between nutrients present is in most cases also
a highly “palatable” one, that will be consumed readily in large amounts.

Supplying Nutrients

Nutrients can be supplied to birds in several ways. It is possible to produce

pure forms of nearly all the nutrients required and to make diets using these
H120 RAPTOR RESEARCH Vol. 6, Supp. H

Table 1 . The Nutrient Requirements of Chicks (0-8 weeks of age).

Nutrient Amount required in diet

Metabolizable energy, Cal/kg diet 3000-3650

Protein (N x 6.25), 0-4 weeks 22%-28%
Protein (N x 6.25), 4-8 weeks 20%-24%
ME/P (ME Cal/gm protein), 0-4 weeks 13.2-13.5
ME/P, 4-8 weeks 15.0-15.3

Amino acids: Percent of the protein

Arginine 6.0
Lysine 5.0
Methionine 3.5
or Methionine 2.0
+ Cystine 1.5
Tryptophan 1.0
Glycine 5.0
Histidine 2.0
Leucine 7.0
Isoleucine 4.0
Phenylalanine 7.0
or Phenylalanine 3.5
+ Tyrosine 3.5
Threonine 3.5
Valine 5.0
Fat +
Linoleic acid 2.0
Fiber
Minerals: * Percent
Calcium 1.0
Phosphorus 0.6
Available phosphorus 0.45
Sodium 0.25
Potassium 0.3
Chlorine 0.15
Magnesium 0.05 (500 mg/kg)
Manganese 0.005 (50 mg/kg)
Zinc 0.005 (50 mg/kg)
Iron 0.0025 (25 mg/kg)
Copper 0.0003 (3 mg/kg)
Molybdenum 0.0002 (2 mg/kg)
Selenium 0.00001 (0.1 mg/kg)
Iodine 0.000035 (0.35 mg/kg)
Cobalt (required only as part of the vitamin B 1 2 molecule)
Sulfur (requirement supplied in form of methionine and cystine; also
present in thiamine and biotin)
 1 1

1972 Breeding Conference -Health and Nutrition of Young HI 2

itrient Amount required in diet

tamins: *
Vitamin A (stabilized) 1320 lU/kg
Vitamin D 3 550 ICU/kg
Vitamin E (acetate is more stable
than alcohol) 15 lU/kg
Vitamin K] (menadione sodium bisul-
fite-Vitamin K 3 -approximately
same requirement) 0.8 mg/kg
Thiamine (Vitamin B ) 1
1 mg/kg
Riboflavin (Vitamin B 2 ) 4 mg/kg
Pantothenic acid 1 3 mg/kg
Niacin 33 mg/ kg
Pyridoxine (Vitamin Bfc) 4 mg/kg
Biotin 0. 2 mg/kg
Folic acid ( Folacin) 0.60 mg/kg
Vitamin B] 2 (Cobalamin) 0.01 mg/kg
Choline 1 200 mg/kg
Ascorbic acid (Vitamin C) Not required
p-Amino benzoic acid Not required
i Inositol Not required
1 hioctic acid Not required
Mevalonic acid Not required

^Mineral and vitamin requirements as given apply to diet containing 3000 Cal
ME/kg. Requirements increase in proportion as energy content of diet is in-
creased. (For example, at 3650 Cal/kg of diet, available phosphorus require-
ment is 3650/3000 x 0.45 = 0.55',*.

Possible unidentified required nutrients:

1. Water-soluble-organic
(a) Spares zinc requirement
2. Water-soluble
(a) contains selenium
3. Fat-soluble
(a) Present in vegetable fats, egg yolk fat and lard
4. Water-soluble
(a) Present in intact proteins

iThe “requirements” presented here are based upon interpretation of research

evidence to date. Undoubtedly, they do not represent the optimum “balance”
of nutrients. Further research on interrelationships between the various nutri-
ents will lead to changes in the “required” levels of many of these nutrients.
H122 RAPTOR RESEARCH Vol. 6, Supp. H

pure nutrients. This is the method commonly used to study qualitative nutri-
tional requirements. Vitamins and essential elements were discovered as diets of
increasing purity were found to be deficient in factors present in impure ingre-
dients.
Ordinarily, however, nutrient requirements of birds are met by making use
of plant materials or animal by-products, mineral elements, along with some
chemically synthesized vitamins, and amino acids. Many sources of various nu-
trients are available for use in bird feeds. Where it is possible to supply a vitamin
such as riboflavin more economically through chemical synthesis rather than by
milk products, for example, this then is the method ordinarily used.
Feed ingredients vary in their ability to supply nutrients required by birds.
The amino acid composition of fish meal, for example, is closer to the needs of
the chicken than the amino acid composition of peanut meal. Corn is a far bet-
ter energy source than wheat bran. Niacin is poorly available in grains but is
supplied readily in pure form or by yeasts. Feed ingredients differ markedly in
their ability to meet nutritional needs of the bird because of their composition,
digestibility, presence of toxic materials and many other factors. Part of the
problem q/ applying scientific nutritional information is the evaluation of avail-
able feed ingredients, and their ability to supnly the nutrients required. Table 1
lists the nutrients required by birds and gives the approximate amounts needed^

in the dietsof chicks to support satisfactory growth. While the table happens to
show the quantitative requirements for the chicks, it is possible to formulate

for many species using these values as guidelines. Requirements between species
vary only slightly in basic nutrient needs and while the form these nutrients are
ingested in may vary greatly in some instances, the basic nutrients and amounts
remain much the same. Thus diets can be formulated for almost all species of
birds using ingredients which will supply ample amounts of these nutrients to
support growth, well-being, and reproduction. By considering the nutrient com-
position of ingredients and blending those ingredients which are acceptable and
will be consumed by a bird, one can provide wholesome, complete diets for
nearly any species, be it a quail, chicken, goose or eagle.

A DIGESTIVE TRA CT BL OCKA GE B Y FECA L ITUS

.

PORTER. We fed our captive American Kestrels chicken parts and whole
ground rodents. Even though the diet seemed to be very adequate for the
growth and development of young Kestrels, we ran into a problem in 1967 in-
volving four out of about 130 young. The fur that was present in the food
accumulated in their stomachs and they were unable to form castings so as to
regurgitate it. As a result, fur blocked their digestive tract and they apparently
starved to death. This affected only the very young nestlings. This was a very
low mortality rate, involving only a small percentage of the young. The follow-
ing year this kind of mortality was not evident.

HALLIWELL. A Golden Eagle at the Topeka Zoo at six or eight weeks of age
1972 Breeding Conference-Health and Nutrition of Young HI 23

died due to a fecalith made up of undigestible materials of fur, feathers and

bone. These materials reformed into a hard calcareous mass and obstructed the
intestinal tract. This same problem has been encountered in other young birds
and seems to be reasonably prevalent. At the time the eaglets were hatched at
the Topeka Zoo they were removed from ZuPreem and placed on rabbits and
rats. In eagle nests visited in Colorado, Wyoming and Montana, there were con-
siderable numbers of rabbits and some young foxes and coyotes. The percent-
ages of fur, bone, teeth and other indigestible parts in the eagles’ natural prey is
about 30 percent of the total. Is there more parenchymous organs and less bone
and fur in these prey species which would make them more digestible to the
eaglets and thus eliminate the fecalith problem?

SCHUBERT. A zoo [name not given that received an immature Bald Eagle
1

from an animal dealer subsequently seemed to get the cramps and died. An au-
topsy revealed a ball of “Stay-dry” (a commercial preparation to put on the
bottom of the cage), all balled up in its digestive tract.

OLENDORFF. Golden Eagles do the same thing; a bird that had an amputat-
ed leg and constantly was eating off the floor had a ball of Stay-dry two-thirds
of the way down the intestine. Stay-dry is dried sugar cane; pieces of sugar cane
passed all the way through the intestine; I guess one stuck and they kept cling-
ing, even though the eagles were casting up a great deal in proportion.

WOLHUTER. Birds have a tendency to find their own castings, if they are
not fed food containing casting material; if the bird is fed on the fist and given
plenty of castings, there is no reason for it to pick them up from the floor.

B.NESTL/NGSAND CASTINGS

1 . Captive Birds

HUNTER. A captive jerkin picked off the pieces of chicken having feathers
and ate them himself and fed the young only parts having no casting or bone.

OLENDORFF. Young captive buteos (hand fed) started casting when they
were days old; they were given bone when they were three to five
six to seven
days old. Their diet was ground very finely and consisted of three pounds of
mice, three pounds of pigeon, three pounds of jack rabbit and three pounds of
cottontail. Vertebrae, bones and skull bones of sparrows were fed to them very
early after hatching. The bones were taken very readily by the young hawks.
Cracking of the bones seemed to stimulate the feeding response.

FYFE. The captive Peregrines were fed Coturnix quail. The male would pluck
the bird surprisingly clean and usually in plucking he would eat the head. He
would then take the prey to the nest or to the female; there wouldn’t be a great
HI 24 RAPTOR RESEARCH Vol. 6, Supp. H
deal of castings at that time. With the captive birds there was very little casting
going into the nest from the very beginning other than perhaps bones which
would be in the food itself.

HAMERSTROM. Young Red-tails seem to get their castings as soon as they

start to care for themselves.

2. Wild Birds

TEMPLE. Yukon River Peregrines plucked their prey species up to a week

before fledging, but they obviously had some castings before this.

GAL1CZ. Peregrines, observed from a blind in the Queen Charlottes some 10

years ago, did not appear to be feeding any casting material, only small pieces
of lean meat.

C. TYPES OF FOOD GIVEN NESTLINGS IN THE WILD

HALLIWELL. During the initial period after hatching, hawks seem to feed
their young quite a bit on the innards, particularly liver of the prey species; the
liver seems to have quite a laxative effect.

HUNTER. When my jerkin came to the liver he made sure that the young got
it;even when he was feeding away from the other birds and they were full, he
would take the liver and feed it to them. The liver seems to have some special
significance. Color had something to do with choice of food for the young:
month old chicks have very light colored meat. The jerkin seemed to be avid
when pigeons were fed and fed on all parts except the skin, whereas with the
chickens he ate the bony structures.

GALICZ. My observations from a blind some 10 years ago with Peregrines on

Queen Charlottes seem to support those of Hunter’s. Peregrines were feeding
the inner parts of Merlins to their very small young.

TEMPLE. Yukon Peregrines definitely fed their young a combination of mus-

cle meat and viscera; on some larger shorebirds and ducks they definitely were
feeding on breast muscle.

ENDERSON. A distinct impression from a frame (on a feeding sequence with

Peregrines— time lapse photography in the Arctic) suggested that when the
young were given small unidentifiable passerines the (adult) bird worked right
down through the lungs, everything.

FYFE. Young, not quite a week old (Prairie Falcons), were getting mostly
muscle meat from ground squirrels and birds at about four days of age. Captive
 5

1 972 Breeding Conference— Health and Nutrition of Young HI 25

Peregrines fed young breast meat, possibly bits of liver, no entrails.

NELSON. On coast Peregrines— when the young were a very young age, their
parents fed them breast muscle, and during the first couple of weeks the parents
were extremely meticulous about not letting the young have any of the guts;
they would offer the entrails to the young, then take them back, eat maybe 1
inches of the intestine, then resume feeding. Innards apparently were not fed
the young at the beginning; it was almost entirely breast muscle— normally skin-
ned, fairly well intact.

[Editorial Note: Figure 1 is a photograph I took at a Utah Peregrine eyrie in

1947 showing a youngster being fed entrails. R.D.P.]

HAMERSTROM. The
subject of raptor baby food has been approached from
several standpoints. Again I try to imitate what the parents would do. The rap-
tors that I have watched closely by sharing the nest as a mate— Red-tailed Hawks
and Golden Eagle— tend to feed liver, heart, lungs, and kidneys— but not intes-
tines— of birds and mammals to small chicks. They fed some muscle even in the
first few days after the hatch— especially that of young birds or mammals. Dr.
Heinz Meng mentioned a preference for feeding dark meat to young chicks and,

Figure 1. Peregrines at 1947 Utah eyrie feeding entrails to young. Photograph

by Richard D. Porter.
HI 26 RAPTOR RESEARCH Vol. 6, Supp. H
as I think back, I agree. It seems to me that pigeon breast was fed before rabbit
meat by both species.
Myfemale Golden Eagle partially swallows and then “regurgitates” morsels
to small chicks. My male Red-tailed Hawk and Dr. Meng’s female Red-tailed
Hawk do not regurgitate, but feed tidbits well moistened with saliva.
I feed any very young raptor that I hand rear the above diet and both mois-

ten and warm each morsel in my mouth before giving it to the chick.

[More information is available in F. Hamerstrom, 1970, “An Eagle to the

Sky,” Ames, Iowa: Iowa State University Press, and F. and F. Hamerstrom,
1971, Potential eines mannlichen Greifvogels ( Buteo jamaicensis ) in Bezug auf
Nestbau, Briiten und Jungenaufzucht, Die Vogelwarte 26:192-197; English
translation in Raptor Research 6(4): 144-149, 1972.]

SIMONYI. In the wild the adult birds (Peregrines) never give feathers to the
young ones, but after about four weeks the young begin playing with the left-
overs at which time some feathers are eaten.

FYFE. Regarding early feeding, two things that were very obvious both in
wild and in captive Peregrines was the extent to which the adults cleaned the
prey before they take it to the nest. The general pattern was that the male
would return to the young with the head removed from the prey.

TEMPLE. Peregrines on the Yukon plucked their prey so thoroughly that it

was difficult to identify the prey items brought into the nest, indicating only a
remote chance for the young to be given extraneous casting materials in large
quantities.

OLENDORFF. Red-tails, Swainson’s and Ferruginous Hawks get pieces of

fur and pieces of bone from the very beginning.

STODDART. About two years ago a Swainson’s Hawk and a Prairie Falcon
were given pine shavings for castings without apparent problems. The shavings
formed good castings and appeared to cast up normally.

FYFE. Young, not quite a week old [Prairie Falcons] were getting mostly
muscle meat from ground squirrels and birds at about four days. Captive Pere-
young breast meat, possibly bits of liver, no entrails.
grines fed

D. DISEASE, PATHOLOGY AND OTHER HEALTH FACTORS

1 . Rickets

HALLIWELL. In rickets, legs are bowed and feet turn in excessively; caused
by feeding nothing but muscle meat.
1972 Breeding Conference— Health and Nutrition of Young HI 27

2. Miscellaneous

OLENDORFF. Referred to a paper by Andre Brosset, which he translated

from French (/ N. Amer. Falconers' Assn. 6:38-42, 1967). Brosset raised his
young raptors on the floor because he experienced a bird falling off the table.

3. Frounce (Trichomoniasis)

OLENDORFF. Two young Red-tailed Hawks, age about 20 days, contracted

frounce. They were treated with “Erntril” and recovered without any problems.

HALLIWELL. Description of Frounce: it’s caused by a very small flagellated

protozoan. Must be placed under a microscope to be seen. It causes a lesion in
the mouth which is a result of the lining of the mouth being irritated. It forms a
cheesy-like growth there. Following treatment it would probably take a day or
two for the lesions to drop off.
To diagnose trichomoniasis: with a well-moistened Q-tip, swab the bird’s
mouth, particularly the back throat region. Put it in a drop of water on a micro-
scope slide. An examination under the microscope will reveal millions of rapid-
ly moving organisms. After treatment you should not see trichomonads after
24 hours, although you may still have scales.

HALLIWELL. I have never seen a case of Frounce that has been refractory
to Erntril. One treatment of Erntril is literally a “sure cure” for trichomoniasis
and it works super fast.

McINTYRE. agree with Dr. Halliwell with but one exception: a Prairie Fal-
I

con inDenver that was treated twice, skipped a 24-hour period with three tab-
lets of Erntril, one of each dosage and still had evidence of Frounce. He was
then placed on Enheptin and this got rid of it. Why did the Erntril not work?
One can not give it to the bird in the first bite of food because it goes past the
crop, where it is needed and on to the stomach. To administer, take the bird’s
bill, close its throat, and poke the food into the esophagus and down into the

crop. I do not worry about the size of dose.

HALLIWELL. Erntril comes in two forms— one is powdered and’ the other is
a tablet. The tablet contains 125 mg. I use 125 mg/2 lbs of bird. Best to use
powdered form, which you have to place in a gelatin capsule; however, may
have problem with gelatin capsule dissolving in quickly enough to release the
drug while the medicant is still in the crop where it will do some good.
Cattle which were given one or two tablets every day, on post mortem, con-
tained 3-5 tablets; hence some tablets were in the digestive tract 24-48 hours.
There is need to develop a better method of administration of medicant since
either form is a problem.

LAYMAN. I use a catheter tube (orally) to administer drugs and liquid foods
HI 28 RAPTOR RESEARCH Vol. 6, Supp. H

to a Prairie Falcon.

SHERROD. When my Prairie Falcon got Frounce, I gave her one-third of a

tablet of Emtril, but after three days the falcon still had Frounce. I then gave
her an additional third of a tablet. This seemed to have adverse effects on the
falcon causing her to go crazy.

HALLIWELL. Dosage (35-40 mg) given the Prairie Falcon was too low; treat
with the entire 25 mg of the drug. However, indiscriminate use of the drug as
1

a cure-all for everything is not wise.

 :

RAPTOR RESEARCH
Volume 6 Supplement Part I 1972

SPECIAL CONFERENCEON CAPTIVITY

BREEDING OF RAPTORS-A REPORT

Sponsored by Raptor Research Foundation, Inc.

Held at Sioux Falls, South Dakota, U.S.A., November 21-23, 1971

Part I. Photoperiod Problems and Management (Panel 4)

edited by
Byron E. Harrell
Biology Department
University of South Dakota
Vermillion, South Dakota 57069

Panel Members Stanley Temple, Chairman

TEMPLE. Photoperiod one type of external stimulus that we are definitely

is

able to manipulate, something that we have under our control when we put the
bird in a captive situation. This subject is going to be particularly critical with
the falcons, since different populations nest at different latitudes. I might start
out by telling you how birds evolved a photoperiodic or light induced control
or timing mechanism for their reproductive cycle. There are several require-
ments that they would probably want in terms of what function they would
use. On£, it must be something that is easily perceived by the animal. Another
thing, it must be something fairly constant from year to year, it shouldn’t be
changing drastically. Probably another thing is that it should be something that
would vary geographically, so that the birds can adapt themselves to a specific
geographic region; I think it doesn’t take much to see that the varying daylight
length throughout the year fits all these requirements admirably. It doesn’t vary
from year to year. March first is the same period of daylight from one year to
the next. Light is something that is very easily perceived by a bird— birds are
primarily visually oriented animals. Also it is something that varies quite a bit
geographically, particularly with latitude. You all know that if you go north,
you are going to have longer days proportionately during the summer breeding
season of most birds. So, as Peregrine Falcons— we might as well address our-
selves to Peregrine Falcons right off— as Peregrine Falcons evolved in the tun-
dra ecosystem they had to adapt their endocrine system specifically so they

1129 Raptor Research 6(Supp.):I129-I140, 1972

1130 RAPTOR RESEARCH Vol. 6, Supp. I

were using the light regime, the photoperiod, that is present in the tundra. They
are specifically adapting themselves for that condition.
I think, very simply, you don’t have to get anything very complicated, on
what kind of regime to put a bird on in terms of light and dark. You can
light
pretty much point blank say that. You should try to duplicate as closely as
possible the light conditions of the latitude of the region from which the bird
came. For tundra birds this means that during the breeding season you should
have them on long periods of light, probably 20 hours of light a day is going to
be necessary. For birds such as Peale’s Falcon or an anatum type of Peregrine,
if you keep them at the latitude where they were normally taken or fairly close
to it, there is no need to supplement with artificial light. I think that it is abun-
dantly clear from the success with Peale’s Falcons and Jim Enderson’s anatums.
Natural light is perfectly adequate unless you have a closed breeding chamber.
For tundra birds, and I suppose this is where most of our potential breeders are
probably going to come from since that is the only viable continental race of
Peregrines left, you are going to have to go to supplemental light in your breed-
ing chamber.
What do we know about the type of light that should be given artificially?
There are a couple of things that are definitely known about a bird’s receptive-
ness to different types of light. This has been shown conclusively in studies of
photoperiodicity on many species of passerine birds; not much has been done
with birds of prey. But all of the other kinds of birds that have been looked at
once again show this uniformity— they are all pretty much the same in their
adaptation to photoperiod. One thing that comes up right away is the intensity
of the light. Every experiment that has been done to measure the intensity of
light that is necessary to trigger a photoperiodic response in a bird shows that
there is a threshold level above which more intensity does nothing to increase
the photoperiodic response. In other words, if you get the light to a certain
level, more light is not going to strengthen the photoperiodic response. The
threshold level that has been discovered in such birds as House Sparrows,
finches, Starlings, is quite low, below what I’ve seen at most breeding facilities.
Just take a regular camera light meter; you can approximate the light that you
would get from daylight with your photofloods. Take a light meter reading off
the floor; if you can duplicate the intensity with photofloods, you are in fine
shape as far as intensity of light goes. There is no evidence to suggest that
brighter intensity is going to help you out. The other thing that is probably cri-
tical is the color or the colorimetric spectrum or character of the light. The
spectral quality of artificial light will vary from light source to light source.
They have done quite a few experiments once again on different species of
birds to find out which wave lengths are most stimulating to the gonads. And
I don’t know whether you want to call it a night club effect or something— red

light is the most stimulatory. If you submit. a bird to light that is rich or has a
high red wave length to it you are more likely to get a stimulatory response
from the bird. Blue light, light that tends toward the ultraviolet, is the worst
that you can use. Infrared light is not effective, it’s invisible and the things that
are stimulatory are in the visible spectrum.
1972 Breeding Conference— Photoperiod 1137

birds, should be putting their birds onto two summers a year. And they may
well get in one of these summers production, as it is. Now they’re not even get-
ting eggs with just one exception over here.

TEMPLE. assume a transequatorial migrant, then you

Well, if you’re going to
are going to put the bird two summers. And if you suspect that,
on essentially
then what I mentioned earlier about this photorefractory period is protecting
the bird from the second cycle. I would be very surprised if you could recycle
that tundra Peregrine. Not until a year later. I’m almost certain.

GRIER. Dr. Porter, have your American Kestrel populations from different
shown the same thing? You’ve got different times under the same regime
areas
with the breeding.

PORTER. Right, we, in our colony, have Kestrels that were obtained as nest-
lings from Massachusetts, New York, Ohio, Pennsylvania, and Maryland. We
also had a group of Kestrels, female Kestrels from Florida, that were obtained
in the winter. The weights of the Florida Kestrels from Florida were not signifi-
cantly different from those from the northeast which suggests that they/mprob*
ably not of the small Florida race, the resident race. In any event the Florida
Kestrels nested a month later than did the birds from the northeast. They main-
tained this despite the fact that they were right in the middle— even the Florida
Kestrels that were placed right in the middle of the colony with the birds from
the northeast, still maintained this month’s difference in nesting data, a month
later. However, as we looked over our records, there was a tendency for this
gap to decrease slowly each year. We really need some experiments with the
Kestrels to determine what they’re actually behind in.

TEMPLE. I think it’s a little bit too much to hope that all tundra birds will
quickly adapt, you know, in a matter of several years.

HUNTER. Would you think then that maybe just a total increase in light
suddenly would be as good as a gradient?

WHITE. That probably may not be the answer.

HUNTER. What I’m getting to is, is a sudden increase in the total light equal-
ly as good as a gradient increase?

TEMPLE. Once again if we’re safe in extrapolating from other species there
doesn’t seem to be any advantage to giving the birds graded increase in light.
For most species the sudden jump up to a stimulatory photoperiod is sufficient,
unless raptors are very differentfrom the other birds who have been examined;
there’s no reason to think going to bring this on your tundra
that. If you’re
birds at the appropriate time, sometimes perhaps during May, up into 20 hours,
I would think that would be perfectly adequate to do it in one increase.
1138 RAPTOR RESEARCH Vol. 6, Supp. I

HUNTER. Some chicken people do this though. They bring the light up
slowly, graded, to increase productivity.

WOLHUTER. I’m wondering if there’s a difference between just all of a sud-

den throwing a long photoperiod, versus gradual, when you talk about the thres-
hold, the bird reaching a threshold, is there any difference in that? Also would
that differ with some birds, like our transequatorial migrants?

TEMPLE. I can’t say that we know for sure with birds of prey, but with oth-
er birds that are easier to breed in captivity, the sudden increase in duration is

sufficient.

WOLHUTER. There’s no need to build up— you can start them long?

TEMPLE. No, so it is with other birds. I’m not saying it’s the same as raptors,
but seemingly we can apply information from other birds to raptors, just as
well.

GRIER. We can’t say that poultry takes a 14-hour day and all you have to do
is put them at that and hold them at that and they’ll keep producing.

MOLT PROBLEMS
HUNTER. I have another comment I would like to make. Something on the
order of seven or eight years ago, I put Dakota Belle, a tundra Peregrine, on 24-
hour light on January 1; 43 days later she began to molt. The next year I did
exactly the same thing but I waited one month later until February 1 44 days ;

later she dropped her first feathers with the full light, 24 hours. Then the next
year I gave her no light: she dropped her feathers as she always had before, on
the 12th of July. Now we’re only assuming that there is some tie-up between
molt and reproductive activity, but . . .

TEMPLE. Yes, the correlation between reproduction and molting, photo-

period molt, probably just a sort of a spurious correlation. It’s pretty clearly
is

shown that the thyroid is probably the organ that is most directly related to
inducing molt. And it is being stimulated by photoperiod, you’re right, but the
stimulation going to the hypothalamus and it’s releasing another hormone
is

that has effect on the gonads; this is the thyroid stimulating hormone
little

which stimulates the thyroid in the same way that the gonadotropins stimulate
the gonads.

HUNTER. On the questionnaires, one of the things I left out I am afraid was

a question of when the first feathers dropped in molt; now I do have some of
these data from various people, many of them having observed the first feathers
to be dropped in the molt after the second egg was laid in the first clutch.
1972 Breeding Conference-Photoperiod 1133

MARCUS. I’m a little conversation about total expo-

confused about all this
sure and total light, integrated light exposure versus light periods. I think we’re

getting mixed up about how many Langleys, for example, does a bird get ex-
posed to? All this seems to be affected by weather and it seems like an unusual
condition in which it would be clear short days that would stimulate, perhaps
help stimulate activity. Then you’d have all kinds of false starts possibly.

TEMPLE. Remember what I said, there is a threshold level above which more
intensity doesn’t stimulate the bird any more strongly, you know.

MARCUS. Energy, you have to measure the light energy. The temporal light
has to do something, it has to cause, let’s say, a photochemical reaction and so
it has to be a total energy quantum, or something that . . .

TEMPLE. No, that’s exactly what these experiments disproved, that there is
not a direct correlation between intensity of light and response. There is none.
In other words, you don’t get it below a threshold; as soon as you reach that
threshold, you get that response.

MARCUS. I’m just disturbed about this business of length of period versus
total exposure. I just plain cannot get the question out of my mind.

TEMPLE. why transequatorial migrants ever leave South

In order to explain
America, there an hypothesis that it is the total exposure to light on the win-
is

tering ground that triggers them to leave. I don’t see how this would work on
stimulating the gonad in light of what we know with other birds.

GRIER. I think what you’re saying is whatever it is, the total amount of rad-
iation isn’t as constant aswhat you were saying initially was necessary from an
evolutionary sense. Let’s say the day length is going to be the samfc thing from
year to year on the 1 5th of May. The amount of light can depend on fog or
cloud cover and all that.

MARCUS. But you have to measure the exposure in some way. It’s not a
timed period and how are you going to measure total exposure time? You have
to integrate it in some way.

TEMPLE. In other words you’re saying that two days out of two days and a
foggy day is worth one day in a sunny day.

MARCUS. I’m not saying that’s so. I’m saying it doesn’t make sense that
way.

TEMPLE. No, it doesn’t. I agree.

HUNTER. What about the problem of light intensity for threshold?

1134 .
RAPTOR RESEARCH Vol. 6, Supp. I

TEMPLE. I don’t think that is a problem, Don. I think all the breeding cham-
bers I have seen or heard of are sufficiently bright.

HUNTER. One thing that bothers me is that it was said by someone and I got
the impression that we need the approximate light outdoors inside the building;
a little bit difficult.

TEMPLE. The evidence from sparrows and other species indicates no.

HUNTER. It might be helpful— I would like to suggest this— it might be help-

ful if we take candlepower which we can do with a regular photographic light
meter. If anyone could do this and take light meter readings, it may be of some
value.

TEMPLE. Certainly as bright as possible.

CAMPBELL. Don, I went into this quite extensively, not being much of a

photographer. I went into this with the poultry peopld; the intensity of artifi-
cial light comparable to daylight is four watts per square foot of floor area. I
also went into this in the schools and the same principle applies in the schools,
your overhead lights or movable, I think, is four watts per square foot, this is

what I am told.

HUNTER. At what distance from the light would this be? It seems there
would have to be a distance factor.

CAMPBELL. No, I don’t think so; if you put your light meter on the ground.
In my own case each breeding chamber has roughly 145 square feet, 12 by 12,
so I can put 600 watts, equally distributed in that pen.

SIMONYI. May I say something, since for the past four years I have had Red-
tailsand the past two years, Peregrines. They need from 95 to 1 10 candle foot
power, per square foot, and that’s it, nothing else.

HALLIWELL. And you run yours 13V6 hours a day?

SIMONYI. Well, I start off at nine and from there I work it up; you can go as
high as you want, up to maybe 20 hours.
*

THACKER. You were talking about colored lights, reds and blues. Four years
ago in Europe there was some work done by Manholds, mostly on lab mice and
rats. If they were exposed to different colored lights, you could almost tell
what sex the offspring would be. The various colored lights would produce vari-
our sexes, like blues would produce one sex, reds would produce another.

TEMPLE. I know that experiment; a lot of people were very skeptical about
the results. It’s the type of thing where your sample sizes weren’t large enough,
1972 Breeding Conference— Photoperiod 1135

there’s a possibility those were spurious.

THACKER. Well, I think he repeated them after he did them the first time,
because there were so many people that were doubtful.

STODDART. I understand that for mammals to reach puberty they have to

have so many hours of infra-red.

TEMPLE. I’m not really up on the mammalian literature, but as far as I

know, that pretty much a theory.

is still

GEOGRAPHIC PROBLEMS

GALICZ. Just for a point of information on our birds in British Columbia

that did lay late in the fall, there was no artificial light used, both last year and
this year.

TEMPLE. This is a phenomenon that is sometimes seen in many species and

it’s very difficult to explain, especially in male birds. You often do see some
type of sexual activity in the fall and the reason for this isn’t clear— some spe-
cies even increase their gonadal size.

GALICZ. These birds are very sexy, they produced fertile eggs.

TEMPLE. That’s very interesting.

MENG. have a feeling that if we have tundra

Stan, in this whole regard I

probably react like four year old

birds, eyasses, that are four years old, they’ll
Peale’s. I don’t think there have been any that have been kept for four years,
are there any?

FYFE. have an eyass female and the eyass male

I will be three next year. But
the female is eight, and she was raised as an eyass.

TEMPLE. We have six year olds, Alaskan birds at Cornell. This is one thing
I’d want to A bird will probably not adapt to a differ-
clarify right off the bat.
ent photoperiod from that of the population it is taken from. The experiments
here were taken from White-crowned Sparrows which breed, as you know, from
California right on up to Alaska and they were not able to stimulate White-
crowned Sparrows from Alaska to the summer photoperiod in California.

WHITE. I’d like to make a couple of comments here that I would hope may
be germaine at this time, because I think a lot of us think photoperiod is really
not understood and we’d hope that those of you working especially with birds
of tundra origin would take very good notes on what happens to your birds and
the way in which they respond. As Stan mentioned, there are often double
cycles. Ptarmigans are a good example of birds that in September set up terri-
1136 RAPTOR RESEARCH Vol. 6, Supp. I

tory for about a week. This may be what happened in British Columbia with
these resident birds. Secondly, as Stan also mentioned, when you have trans-
equatorial migrants, we don’t really know what it is that triggers off the return
of migration. For example, there are voluminous data to demonstrate that birds
from tundra origin moving north in the spring as in April and May across breed-
ing grounds of birds that are already breeding. Many of these birds do not have
developed gonads, and when they arrive in the tundra, many of them still do
not have developed gonads. And so one would suggest that rather than the hour
photoperiod, say a 16-hour photoperiod, or whatever, but rather it is the total
accumulation of light over a given period that causes gonadal cycling. So with
these long-distance migrants I would hope that those who are working with
them would take very, very good notes on what they do in terms of this be-
cause the same phenomenon has been noted in birds of the Soviet Union in
birds here.

NELSON. On these arctic birds would you care to speculate as to the time
ranging on them. I’m thinking particularly of Bob Berry’s notes in one of the
BPIE reports in which he said the birds started getting hepped up in the fall.
Now as I see it, if the tundra birds are down in South America, something is
getting them to move back north; this may be why the tundra Peregrine stimu-
lated with hormones over here was much. Maybe she was trying to
exercising so
migrate back north as Richard Fyfe’s captives apparently tried doing a few
years back. The transition between South America and the Canadian Arctic,
let’s say, is going from an autumn to a spring and summer very rapidly. Now
would you care to speculate on what type of a light regime people should be
using in their captive building and how they might stimulate the birds as they
would be stimulated in the wild.

TEMPLE. As you can do probably no better than to approximate the

I say
natural condition. would probably put the birds onto a stimulatory photo-
I

period. For a tundra Peregrine, probably sometime around the first of May, the
end of April. I would suggest something on the order of 20 hours of light, four
hours of darkness. I would probably hold the birds on that light until probably
late July or August and then drop them down to a winter photoperiod, perhaps
drop them down to the normal photoperiod for the area where they were.

NELSON. That’s the problem. I think that’s what everybody has been trying
to do. They give them one summer and then put them back into a winter which
the birds never ever have seen.

TEMPLE. That’s not true, because tundra birds, some of them, are transequa-
torialwhere they do go down there, but they certainly do winter on the Texas
coast and Mexico.

NELSON. Yes, but we don’t know where those birds are coming from neces-
sarily. My point is that these people, at least some of them with high arctic
1972 Breeding Conference— Photoperiod 1131

Now how do you stand when you start using incandescent bulbs or fluores-
cent light? You’re actually in pretty good shape on both of these. They both
tend to produce light of a wave length that will give you a good red component.
And actuallyI can’t think that you can improve very much on normal indandes-

cent and fluorescent light. They’re both very adequate. There are some that
may be better. You may be familiar with light called Vita-light. It is a fluores-
cent bulb that has specially tinted glass and very closely approximates the natu-
ral spectral composition of daylight. If you want to do it naturally, one of these
Vita-lights isprobably one of the best things you can get for it. We’ve got a
couple of other considerations . . .

CRAWFORD. Is that the same as Gro-light?

TEMPLE. Gro-light, yes, the same thing. It’s a different product name. Any
of these, there are several brand names, but these are ones that are filtered so
they approximate a natural spectrum. You’re basically pretty safe with just a-
bout any of the artificial light sources.
One other thing that you have to worry about is the phenomenon that’s been
called the refractory period. Let’s just follow a Peregrine, she’s migrating up to
the tundra, increasing her photoperiod, the daylight per day stimulating her
gonads, she reaches a point at the end of the breeding season where her gonads
start to regress. As her gonads start to regress, she enters a photorefractory
period. This photorefractory period varies from species to species in how long it
is, how long it lasts. During this period further increases in the photoperiod will

not stimulate gonadal growth. This is very adaptive; consider what happens to
that tundra Peregrine when she leaves the arctic in late summer. There is a de-
crease in photoperiod; by heading south, she crosses the equator, going down to
Argentina or wherever she might want to winter. Once she crosses the equator,
she’s in the Southern Hemisphere and she starts increasing her photoperiod a-
gain. It’s obviously nonadaptive to have gonads enlarging on the winter grounds.
And this photorefractory period, you can almost assuredly say, in a species like
the Peregrine Falcon where they’re known to be transequatorial migrants, is
probably quite long, It probably takes them on through to early winter, prob-
ably mid-winter, on the order of four or five months.
Now what does this mean in terms of recycling your bird? We haven’t really
discussed this at all, but it’s something that really bears consideration. Using the
photoperiod and artificially manipulating the photoperiod, it is possible to put
your bird on more than one stimulatory light cycle a year. There have been a
number of people who’ve tried this; I think Jim Enderson has been the most
successful in getting some reproductive behavior from his Prairie Falcon by giv-
ing it a stimulatory light regime in the late fall. This is fine for a Prairie Falcon.
The Prairie Falcons, since they aren’t highly migratory, probably have a very
short photorefractory period. I would definitely say that it’s not going to be
possible with tundra falcons to get them to recycle, to go through more than
one breeding cycle in a given calendar year because of this photorefractory
period. It may be very different with Peale’s Falcons and I understand the group
1132 RAPTOR RESEARCH Vol. 6, Supp. I

in British Columbia has some evidence that they are able to recycle their Peale’s
Falcons so they can get an extra reproductive cycle in the fall. Now the way
you would accomplish this if you’re going to do it by artificially manipulating
the light, would be to allow your bird perhaps in the spring to follow a normal
stimulatory photoperiod, once she had finished a reproductive cycle, drop the
lights down to a nonstimulatory or winter photoperiod, keep the bird on that.
I would probably say extend that period; without any basis for how long, you

should extend it, but for most species I would imagine you should probably ex-
tend it for a good three or four months, I would say 90 days at least, before
you started increasing the birds’ photoperiod again, to restimulate. This way I
think by careful manipulation you should be able to get more than one clutch
out of a bird a year. I think it hasn’t been said so far, but I think you’re prob-
ably all aware that in falcons in particular you can remove a first clutch of eggs
and they will replace with a second and sometimes a third clutch. Once again if
you’ve got a Peregrine that will produce fertile eggs, you can probably get 25
Peregrine eggs out of two Peregrines by constantly causing them to recycle, as
Jim Enderson has shown very clearly, even though these were infertile. I think
that’s probably about it for photoperiod. It’s really fairly simple. There’s no
mystique about it.

LIGHT-QUANTITY AND QUALITY

LAWSON. I think there have been some experiments done in poultry where
they actually put discs in the eyes of the birds and they claim that the light
stimulus actually goes through the skull.

TEMPLE. This is serious controversial work. I might summarize that because

a lot of that was done at Cornell. What
they’ve done is to remove the eyeball
and stimulate the optic nerve directly, using a quartz rod transmitting the light.
You probably know that quartz will transmit the light in a'straight line, and
they transmit it right on to the surface of the optic nerve and they’re able to
stimulate gonadal development doing this. Another common theory, if it is still
in a theory stage, is that the organ called the pineal body in the brain which is
the vestige of an actual photosensitive structure in lower vertebrates, such as
amphibians and reptiles, is still present in birds and mammals and is located in
such a position in the skull that it’s possible that light is transmitted through
the skull. This pineal body may be a very important internal clock mechanism
to the animal; they don’t think that it has much to do with reproduction. All
vertebrates have daily cycles of events that are very precisely controlled. We get
longer cycles that have some type of internal control and' it’s been theorized
that the pineal body may be responsible for some of these controls.

HUNTER. Iread some work with Mallard ducks, in which they remove the
and subjected the light directly on the skull, and they
eyeball, cover the eye up,
got gonadal development without even going through the optic nerve.
1972 Breeding Conference— Photoperiod 1139

TEMPLE. some interesting observations that Jim Enderson

Well, we’ve got
and Jerry Swartz and made up in Alaska using a time-lapse camera of nesting
I

Peregrine Falcons throughout their nesting season. The photographs were gpodd
enough that we could monitor the molt of these Peregrines throughout the re-
productive period. Basically what we found was that the molt was not sort of a
steady molt throughout the breeding period. The female at the hatching of the
eggs stopped h£r molt; her molt arrested right there and she did not continue
her molt until much later on after the young were fairly close to fledging. This
may be very adaptive for the birds because when the young are growing of
course the female is at a handicap if she’d molt; she’s going to have to go out
and help provide food, you know. They’re going along very steadily; undoubt-
edly it is tied in to a certain extent with the reproductive cycle. Dr. Awender,
haven’t you done some work on inducing molts with hormones?

AWENDER. Yes, but that was female sex hormones; it was very closely re-
lated to the progesterone, Noroleucate. There are a couple of new ones which I
haven’t tried yet. This is a strong pregestational and antiestrogenic effect that
does it, nothing to do with thyroid.

TEMPLE. Of course, you molt in most birds seems to be asso-

realize that the
ciated with a decrease in the gonads or a decrease in the circulation titre of
steroid hormones. This may be a spurious correlation.

GRAHAM. I have a quick question maybe someone can throw some light on.
Has anyone thought of why a Gyrfalcon from the arctic brought down will
molt in Alaska, whereas the tundra Peregrines brought down from an arctic reg-
ime having this different molting season . . .

HUNTER. That’s not just the same.

WHITE. No, the tundra birds I brought down molted the same time the rest
of the wild population do. I’m talking about eyasses. Eyass tundra birds brought
down in this example kept in mid-latitude for three or four years molt at the
same time that the wild tundra populations do. Now you’re talking about trapp-
ed eyasses.

GRAHAM. No, I’m comparing trapped passage Gyrfalcons and trapped pas-
sage tundra Peregrines.

WHITE. Eyasses molt at the same place regardless. You take an eyass tundra
bird— it’ll molt the same time the tundra populations does— they’ll start in April
and end, you know, in August.

STODDART. I was just wondering with all this discussion, what were you
talking about when you talk about tundra birds? Are you classifying the passage
1140 RAPTOR RESEARCH Vol. 6, Supp. I

Peregrines trapped on the beach?

WHITE. Of course, when you get a migrating bird,you can say is that it is
all

from some place further north. I don’t want to comment because many tundra
birds do appear on the Texas coast, and many, in fact probably some of the
western birds that are raised right in Colorado occur on the coast; nobody really
knows until we get banding data. What Dick asked was whether birds taken
from the Colville River, for example, molt at the same time that the wild popu-
lation molts along the Colville River. If you were to have a passage bird from
the Colville I would wager you that it would not molt at the same time as the
eyasses.

STODDART. Wouldn’t you think if we are going to birds especially the pas-
sages, that you should match molts.

TEMPLE. The molt of the male and the female are drastically difficult.

STODDART. No, compared to the wild.

HUNT. In regard to bird trapped on the Texas coast, I’ve never seen one
that looks like an anatum or molts like an anatum.
RAPTOR RESEARCH
Volume 6 Supplement Part J 1972

SPECIAL CONFERENCE ON CAPTIVITY

BREEDING OF RAPTORS-A REPORT

Sponsored by Raptor Research Foundation, Inc.

Held at Sioux Falls, South Dakota, U S. A., November 22-23, 1971

Part J. Reproductive Endocrinology (Panel 5)

edited by
Richard R. Olendorff
3317 Olympus Drive
Bremerton, Washington 98310

Panel Members: Erich Awender, Chairman, Timothy Lawson, James C. McIn-

tyre, and Stanley Temple.

AWENDER. We here because this is going to

will try to get right into things
be a more nebulous problem and may be more I can already see things
divisive.
shaping up from two sides, the practical and theoretical. They do not rule each
other out, but I think we will have quite a bit of discussion and not quite so
much formal presentation. Mr. Temple will present the basics so that we will
know what we are talking about and see in general outline what endocrinology
is all about. I am not too qualified to say anything here, because I am mostly

versed background-wise in human medicine and I have done very little work
with birds. I will tell you what little I know later, but let’s start with Mr. Tem-
ple, please.

TEMPLE. As Dr. Awender has said, when we are talking about endocrinology
of breeding birds, we are talking about the control mechanism. Endocrinology
is the study of hormones. Hormones are internal secretions of the body and are

the mechanisms by which birds, mammals, and all vertebrates control their re-
productive processes. We are dealing with the very intimate control of all of the
acts that are associated with breeding. It is a very complicated picture and, un-
fortunately, the information for birds as a group is very scanty. Our under-

standing of even mammalian endocrinology, especially of the cycles of various

hormones that control reproduction, is scanty; with birds we are even more at
a loss. We are fortunate that the evidence collected so far indicates that most

J141 Raptor Research 6(Supp):J 141-1 61, 1972

J142 RAPTOR RESEARCH Vol. 6, Supp. J

vertebrates have the same (certainly all birds have the same) basic endocrine

control of their reproductive cycles. Information that we might gain from

studying the endocrinology of a pigeon or a chicken or some other bird is un-
doubtedly perfectly applicable to birds of prey. Basically, because they are so
important to the birds, there has not been a lot of modification of these sys-
tems. It is one basic system that has worked. So what I am going to do is go
through the endocrine control of a bird’s reproductive system, and point out,
when I finish explaining it, possible places in this system where problems im-
posed by captivity may be disrupting the normal function. After I have done
this, I might comment a little on ways of helping the birds with reproductive
endocrinology problems. I think Dr. Awender and Tim Lawson will follow up
with some of the approaches that they have tried.
First of all let’s start out with what it is that triggers the reproductive cycle
in birds. What starts the cycle is an external stimulus. The responses to external
stimuli are probably where birds have evolved all the variation that we see be-
tween species. Different species have evolved different sets of external stimuli
which trigger reproduction. For many birds, photoperiod— increasing day length
—is one of the most critical external stimuli triggering the reproductive cycle.
For other birds, climatic factors, such as rainfall, temperature, and other things
also come into play, as do important factors such as the interaction between
pairs, nest-building and courtship. All of these external stimuli come from out-
side the individual bird. The external stimuli are received through the bird’s
senses. Be they visual, auditory, tactile, or whatever, the stimuli are received by
the central nervous system— i.e. by the brain. They are then transferred to the
region of the brain called the hypothalamus. The hypothalamus you can think
of as being an integrating system. This is where all the information is synthe-
sized, and the appropriate messages, if you will call them that, are relayed to
the different portions of the brain that will ultimately trigger the production of
the hormones which in turn will control the reproductive cycle. The hypothala-
mus secretes very little known substances called neurohormones. These hor-
mones are secreted by nerves, a very unusual condition, and they are very, very
hard to identify. These neurohormones are secreted by the hypothalamus after
it has integrated all the information from the external environment. They are

then relayed to the pituitary gland.

If you remember your high school biology, the common designation for the
pituitary gland is the master gland of the body. Many of the important hor-
mones that control body processes are produced in the pituitary. This is exactly
the case with reproduction. The whole thing gets started in the pituitary. This
is the gland that will secrete, after it has been triggered, gonadotrophic hor-

mones. Gonadotrophic hormones, very simply, are hormones that will stimulate
the gonads. With birds we are very much at a loss in identifying the gonado-
trophic hormones. For mammals it is fairly well established. There are two main
ones— Follicle Stimulating Hormone, commonly called FSH, and Luteinizing
Hormone, commonly known as LH. These two hormones have been identified
in mammals as the hormones that control the maturation of the gonads. We
1972 Breeding Conference— Endocrinology J143

have not yet been able to isolate two separate hormones from birds. In birds
there may be one hormone that has both of these functions. But, suffice it to
say, the anterior pituitary secretes gonadotrophic hormones which are released
into the blood stream and circulated through the body to the gonads.
FSH and LH (or whatever they might be in birds) have two separate func-
tions. FSH in the female causes development of the ovaries. It causes the ovaries
to enlarge, to become mature, preparing them for ovulation. In the male, FSH
stimulates the development of testes. Birds are cyclic in their gonadal develop-
ment. During the winter the gonads regress; during the breeding season they en-
large. The enlargement of avian gonads is in direct response to these gonado-
trophic hormones. LH, the other gonadetrophic hormone that we know exists
in vertebrates, has the function in the female of causing ovulation. Once the
gonad has matured, a spike or sudden spurt of LH from the anterior pituitary
causes the egg to be ovulated, to leave the ovary, and pass into the oviduct
where it will be developed into an egg and laid. In the male, LH does not peak
quite the same way it does in the female, or we do not think it does. LH prob-
ably increases along with FSH and this is also responsible for the development
of secretory products in the gonads.
The gonads not only produce gametes, sperm and eggs, but they are also im-
portant in that they secrete sex steroids, the sex hormones, as well. These play
a large role in the development of secondary sexual characters, such as the comb
in a male chicken. They have a tremendous role in controlling male and female
behavior. The way the gonads elicit this is by secreting the sex steroids into the
blood stream. We are talking about estrogens. The estrogen that is most active
in birds is estradiol. In males we are talking about androgens produced by the
testes. The main androgen produced by the bird’s testes is testosterone. When
released from the gonads into the blood stream, the sex steroids circulate and
cause appropriate changes that will prime what you might think of as the acces-
sory breeding organs. It will cause, for instance, the development of the sperm
duct that will eventually carry the semen from the testes to the outside. In the
female it causes the development of the oviduct, the structure which is going to
be secreting the albumen and the eggshell around the egg. These structures are
also cyclic, decreasing in size during the winter and increasing in size during the
reproductive season. The increase is in response to steroid hormones secreted by
either the testes or the ovary. Also, estradiol and testosterone control reproduc-
tive behavior. They do this once again by going back through the bloodstream
to the brain, to the hypothalamus. At this point, the information on the circu-
lating levels of these hormones causes appropriate behavior. As the testosterone
levels in the blood increase, a male bird is stimulated, for instance, to defend
his territory or to go through courtship display; a female responds by building
nests and all the other courtship displays that are associated with the breeding
cycle.
That is basically the cycle of hormones that we are talking about. It is very
important that we realize that this is a cycle of hormones. All of these are inter-
related; think of it as a circular thing. We are dealing with a situation known as
J144 RAPTOR RESEARCH Vol. 6, Supp. J

a negative feedback mechanism— something like a thermostatic control. Let’s

go back over the system and show how this negative feedback system works.
Early on in the breeding season, when the external stimuli become sufficient to
cause secretion of the gonadotrophic hormones, the hypothalamus is also moni-
toring the level of the steroidhormones from the gonads. I am going to be a
little anthropomorphic here, but just for convenience suppose the hypothala-

mus is not detecting very much estradiol or testosterone. The gonads are small;
they need the gonadotrophic hormones. The hypothalamus secretes a lot of
gonadotrophic hormones to cause development of the gonads. As the gonads
increase in size in response to this, they start secreting steroid hormones, either
testosterone or estradiol. The blood levels increase. The hypothalamus reads
this and says, “There is a high level of steroid hormones; therefore it is time to
shut off or decrease the gonadotrophic level.” So you can see how this is a
thermostatically controlled mechanism. They balance each other out, very cri-
tically. This has important implications which I will talk about in a few minutes.
The other thing that is important about hormones and their control of repro-
duction in birds is that the timing of these hormones is very, very critical. The
hormones have to come into play at exactly the right time to cause a synchro-
nous series of events to occur. As I mentioned before, you have to have a grad-
ual build-up of gonadotrophic hormones to gradually build up the gonads.
Once the gonads are mature, you need a big spark, or peak, of the hormones
that will cause ovulation, the release of the egg. You also have to have approp-
riate changes in the gonadal hormones, the steroids. These have to build up to a
certain level. The intensity of the behavior that they are going to elicit is, of
course, a direct function of how much of this hormone is circulated. Behavior,
as you know, starts out slowly early in spring and builds up to a peak. It is prob-
ably at this peak time when reproductive behavior is peaking. We are talking
about copulation and fertilization and the actual events that mean success or
failure of breeding.
Now, where might this system break down in a captive bird? There are several
places. The first one involves the external stimuli. When you put a bird into cap-
tivity, you are obviously depriving it of many of the external stimuli he would
normally be getting in the wild. As of yet, no one has cataloged the complete
array of external stimuli necessary for a Peregrine Falcon to be successfully
stimulated to reproduce. I think we can safely say that photoperiod, the increas-
ing of the day length, is very important. Also, probably, courtship, the inter-
play between the mates, is also a very important stimulus. Probably, having an
adequate nest site is very important. Outside of these very obvious ones, we just
do not know. This is where the items discussed earlier on the basic require-
ments for a successful breeding chamber come into play. It very well could be
that some things associated with the deprivation of stimuli in captivity are
blocking the whole system right at the start. Another possible place where you
could get into problems is in the hypothalamus. It could be that the birds are
picking up other stimuli from the captive situation that are blocking the hypo-
thalamus from triggering production of gonadotrophic hormones. Certainly you
1972 Breeding Conference— Endocrinology J 145

know that there are many things that we do with birds in captivity that are un-
usual in terms of stimuli. At the level of the gonads things can also go wrong. It

is very possible that the gonads may not be producing enough of the steroid
hormones, the sex hormones, to elicit proper behavior. When we get to the
level of the gonads, we can go to an endocrinologist who is very versed in these
matters and ask, “We have a bird in captivity; we have done everything we can
to make conditions ideal for it, but it will not breed; what is wrong?” First
thing he is going to ask is, “What were the conditions of the gonads after you
gave this bird all of these sufficient stimuli— or what you thought was suffici-
ent?” This is one place where we are sadly lacking. We have had many failures
breeding Peregrines, and I would venture to say that we probably do not even
have one or two instances where we knew what conditions of the gonads accom-
panied the failures. Were these gonads enlarged? Were they anywhere near func-
tional? This is a critical question. If we find, for instance, that they are not en-
larged— they are remaining very small and undeveloped— then we can say the
problem is with the hypothalamus. If they are enlarged and we are still not get-
ting appropriate reproduction, then we might think of looking at the sex hor-
mones. Is there production of those to elicit the behavior that must accompany
reproduction?
These are two points where we might consider helping birds out. We have two
groups of hormones at our disposal, the gonadotrophic hormones and the sex
hormones. I would first of all, as a physiologist, caution anyone from using
these hormones in a bird for which you have not examined the gonads. Remem-
ber, this is a negative feedback system. You must know what the condition of
the gonads is in your bird that is not breeding. You may well screw up the bird
even more by giving it an inappropriate hormone. For instance, let’s say that
your bird is not doing anything. You say, “Well, I’ll give the male a shot of tes-
tosterone; that ought to make him reproductively inclined.” In fact, it will. If
you castrate a bird and give it supplemental injections of testosterone, it will go
through all of the appropriate reproductive behaviors, including mounting and
copulation, but, of course, if the testes are removed, it has no sperm to contrib-
ute. So, let’s say you go ahead and try injecting testosterone; let’s say that was
not the problem; let’s say that the bird actually did not have gonads that were
enlarged. What you have done then is add testosterone. It goes back to the
hypothalamus, and the hypothalamus says, “Oh, here is all this testosterone,”
and cuts off all the gonadotrophic hormones. What you have done, if anything,
is to decrease the size of your bird’s gonads, because you have knocked out the

gonad stimulating hormone. Let’s say you do it the other way and make a mis-
take; you inject gonadotrophic hormones, when, in fact, the gonads are fully
developed. What you are going to do, then, is harm the cycle of the testes or
the ovaries; the cycle has to build up gradually. What I would caution is that
before you attempt any kind of wholesale use of these hormones on the bird,
you are going to have to know what the condition of the gonads is in the bird
that did not breed for you.
Now, the other thing that makes manipulation with these hormones very
J146 RAPTOR RESEARCH Vol. 6, Supp. J

difficult is that for no species of birds, including chickens, do we have any in-
formation on what the normal circulating levels of these hormones are. We have
ballpark estimates, but we have no information at all on what the level is, for
instance, during the peak in the breeding season. We have no idea what the
concentrations of sex hormones are in the blood of even a breeding chicken.
As I said, these hormones are very precisely involved in this cycle. If you are
going to attempt to induce breeding in a bird by injecting exogenous hormones,
you are going to have to be very sure that you are going to be able to inject
these hormones in the proper sequence, at the proper concentrations, at the
proper time. Otherwise, you should not expect to get sure-fire results. There
have been reports in some of the literature of people who have stimulated cer-
tain birds. In fact, the only successes that I know of that have been published,
have been achieved with small finches. They have injected sometimes artificial
gonadotrophic hormones, sometimes naturally occurring mammalian hormones.
These successes are not really adequate to say that it was the injection of hor-
mones that ultimately triggered the birds to lay eggs. The birds grew up under
conditions that were definitely stimulatory to gonadal development, and it was
not a very well controlled experiment. What I am trying to say is that the re-
searchers did not show that, in fact, these hormones were the responsible agent.
One very interesting report that I should mention was carried out on Pintails.
If you happen to be a waterfowl breeder, Pintails are your Peregrine Falcons.
They are very difficult to breed in captivity; why, they do not know. A very
well designed experiment was carried out at the Delta Waterfowl Research Sta-
tion in the late 50’s on this problem. The fellow who carried out this study had
huge sample sizes of Pintails to work with. He could experiment; he had room
to play around, which we do not have with Peregrine Falcons, obviously. He
tried every trick in the book; he tried giving them increased photoperiod; he
tried varying all the external stimuli he could possibly vary. He still could not
get full gonadal development. Gonads would develop to a sub-optimal level,
taper off and decrease. He then decided to try the hormones. He looked at the
gonads. He knew that the gonads did not reach full condition, so he thought,
“Well we’ll try gonadotrophic hormones to see if I can beef up the hormone tit-
ers and stimulate the gonads to become fully mature.” He tried a wide variety
of doses on many birds. He tried FSH alone, LH alone and combinations of the
two. He tried another hormone which is very difficult to get hold of. He also
tried a hormone called pregnant mare’s serum. The serum from a pregnant horse
has a hormone in it that, as of yet, is unidentified. This hormone possesses the
dual qualities of FSH and LH. It is sort of a hybrid form. You can get this very
easily from the blood of a pregnant horse. He tried giving this because it’s read-
ily available. He gave it in varying doses— up to massive doses— in a complete
range and, even with this, he was not able to get the gonads to come into full
condition. This is probably because we have no knowledge as to how this hor-
mone should be injected— what the cycle of it should be. He went point blank
injecting, and did not get results.
Now, I am not saying that this approach is not going to work; what I am say-
1972 Breeding Conference— Endocrinology J147

ing is, if you apply the artificial exogenous you are going
hormones to a bird,
to have to be very careful what you you may ruin the bird.
are doing because
I do not want to sound pessimistic, because I know Dr. Awender and Tim Law-

son have both done some experiments with exogenous hormones that indicate
that perhaps there is as yet some response in birds of prey to these hormones.
However, just to caution you, realize that in these experiments they were not
controlled to show that it was actually the injection of the hormones that was
causing the development of the gonads. I hope somebody in this room can pick
up the obvious project and look at it with a common species like the Kestrel.
If you get a bird that will not breed in captivity, laparotomize it, look at the
gonads, and then work from there. That is the first experiment that has to be
done.

AWENDER. I will be brief with my own project. I have a very small sample
towork with and I am not willing to take any chances to harm the birds, kill
them or operate on them; that is the main reason I shied away from injections
to begin with. I have ruled out injections. Injectable stuff is very hard to come
by, it is expensive, and we do not know for sure whether it would have any sig-
nificance for birds. Most of the laboratory work that has been done so far in-
volves laboratory animals, usually a rat, mouse or guinea pig, very occasionally
a chicken and, once, maybe a pigeon. But the bulk of everything has been done
on mammals and has been applied mostly in human medicine with variable re-
sults. I do not think many, if any, groups, as such, have applied something ther-
apeutic to birds. It was always on the scientific level. We do not care too much
if a chicken does not lay eggs; we kill it and eat it. But with falcons it is a little
different. So I have ruled out shots and went to tablet .

My experimental pair, just to tell you what I was working with, are passage-
tundra falcons. The tiercel is going on four; the falcon is going on nine, years of
age. They were ordinary passage falcons. They were trapped at the usual time
at the usual place. The female was flown in falconry for about five years, the
tiercel for one and a half years. From then on I put them in this project and
they have been together since, with one interruption. The tiercel was gone last
year for a short time. There is not too much
of non-injectable things
in the line
to stimulate the pituitary. As I said before, I am drawing only from what I can
deduce from human medicine, and I have had very little, if any, contact with
veterinarians. I do not have the foggiest idea where the gonads of my falcons
are. Are they big, little, or indifferent; are they even existent? The falcons were
acting in a very neutral way. If the size would not have betrayed the tiercel, I
would not have even known which sex they were.
So, at any rate, I thought I would start at the top, from the pituitary. I went
ahead last year, in 1970, and put the two tundra falcons in their room where
they were going to stay, and exposed them to light, beginning abojit the first of
March. They had light for about a month— about 20 hours a day. It amounted
to roughly 500 watts plus daylight. After the first month I put them on 24-hour
daylight, figuring if a little is good, a lot might be better. This is a common
J148 RAPTOR RESEARCH Vol. 6, Supp. J

thing applied by lay people. This did not do much; there was no change in be-
havior at all; they were very passive to me and to each other, and stayed out of
each other’s way. They never perched together. Then I heard about a drug
which was producing quintuplets, sextuplets and even eight babies in humans
and I thought, great. The more eggs it produced in a falcon, the better. It may
kill a woman, but not a falcon: they lay one at a time. This drug is called clomi-
phene-citrate or Clomid for short. The trade name is Clomid. It is a synthetic
thing, comes in tablet form, 50 mg per tablet, and women take it for only five
days. Then they are supposed to ovulate, and some of them really do. They go
overboard and eventually they abort; they cannot carry them to term— not
enough room. So I thought this was ideal for falcons. I figured that if a woman
takes 50 mg a day for five days, a falcon needs 0.75 mg per day on a commen-
surate per weight basis. I could not make it; it was so little. I had to go to the
pharmacy and they had trouble weighing out that little for me, but we thought
we came close to 0.75 mg. I put it in a little capsule, put it in a piece of meat
and gave it to the falcon for five days, figuring to do it just like for the ladies.
Nothing happened; absolutely nothing happened. The falcon started her molt
a little bit late. That was a pretty complete washout. In order to be good to the
tiercel, too— I did not want to inject him— there is testosterone which is fairly
effective by mouth if a methyl group is attached to it; it is then methyl-testos-
terone. In human medicine we use it in doses of 10 mg per tablet and usually
one, two or three tablets will do. I gave the tiercel a human dose. It did not
kill him; it did not do anything. Nothing! It may have inhibited, as we may
conclude from Temple’s presentation. At any rate, he had the testosterone for
the same time the falcon had her drug. I stopped the experiment and that was
that for 1970.
This year I thought, well, let’s go one better and prime the falcon with estro-
genic hormone. It did not work from the top, so I started from the bottom and
gave her something by mouth. The ordinary estrogenic substance is not effec-
tive by mouth so I went to stylbestrol, diethyl stylbestrol to be precise, and
that comes in my mg tablets. I primed this falcon with diethyl
office in half
stylbestrol, half mg
per day, given by mouth. I started on May 9 and gave it to
her until May 28, roughly three weeks. That did not do much at first, but to-
wards the end of that period the falcon became just a little restless. I could not
explain on what basis and I did not know if the drug had anything to do with
it. She was peaceful usually, but then she sat on the highest perch and beat her

wings like crazy, exercising many hours a day. I am not aware of what this
meant, but otherwise she left the tiercel alone. It was the same falcon on the
same photoperiod. Then, on May 29, I stopped abruptly the diethyl stylbes-
trol and switched over to the Clomid that I gave her the year before. Again, I
gave daily doses, but I really went up high, probably too high because I figured
I did not have enough time. I gave her 1 2Vi mg; this is a quarter of a tablet. That

is an enormous dose on a weight basis. She took it from May 29 to June 9 and

that made a significant behavioral change. She became extremely broody, which
she had never done before. She went into her brooding comer and made a
1972 Breeding Conference— Endocrinology J149

scrape there. She became extremely passive. I could touch her on the back and
she did not get up. She acted like Mr. Nelson mentions several times— she acted
sick— but she stayed in good physical health. She ate and I could practically
feed her like a little eyass falcon; she was that tame and that quiet. I palpated
her abdomen to see if she would enlarge things. This drug is supposed to induce
ovulation or, at least, get the follicles to mature more. I could not palpate any
increase in her abdomen. Laparotomy was out of the question.
I stopped this experiment at that time. After about a month or so she chang-

ed her behavior again and this time with a vengeance. She became extremely
aggressive to me and to the tiercel. This persists to date. The tiercel has to re-
treat many times a day. She has crabbed with him several times. My room is 10
feet wide, 1 5 feet long and the highest point is about 1 2 feet. If she was nimble
enough she could really hurt him; nothing has happened so far, but she has re-
mained, for about a month after I stopped those drugs, extremely aggressive
and downright mean to the tiercel. The tiercel received no medication this year.
I have just left him alone. He behaved passively, as usual, and now he is con-

stantly a fugitive. The female again postponed her molt even more. She started
extremely late. She started about the first of September. Now, of course, she is
almost finished; she molted extremely rapidly.
There is not much else I can add to this particular pair. The Clomid is some-
thing that has been used in humans successfully, as I said. In the ordinary lab-
oratory animals, and as far as I know definitely on mice and laboratory rats, it
did inhibit. It inhibited follicle maturation and ovulation. I thought, perhaps, if
it works in people, it may not work in lower mammals. As you can see, with

falcons nothing happened. I do not know why, but I think if somebody had
more time and wanted to titrate the dosages a little more accurately, perhaps
more could come of it. It has one advantage; it can be given by mouth. Most of
the other hormones we are talking about here can only be injected and again
we just never know where we stand with the dosage. Too much is not good;
too little is not good; and whether we start on the top or on the bottom we may
antagonize the other part.
I have tried other pairs, but they have not been receiving any hormone treat-

ment. One is a pair of Peale’s. I think the passage tundras needed it most. That
is why I picked this pair for experimentation. Passage tundras are known to be

hard to get into any kind of breeding condition, much less to lay fertile eggs.
The others will probably do it naturally. I have a pair of Peale’s and a pair of
Prairies. The Prairie Falcons have been stimulated with light only and I think
they did not do it on a natural basis because the tiercel was too young. The tier-
cel is a home-bred bird that Henry Kendall bred three years ago. He is one of
the surviving tiercels. He has been mated with a falcon that was then three years
older. Now he is three; she is six. She has laid eggs on photophasing. I turned
the light on one year on December first and left it on day and night. Six weeks
later, January 16 she started to lay eggs at the usual two-day interval. She laid
a clutch of four. I took the eggs away and put them in an incubator; they were
not fertile. I gave her two banty chicken eggs. She alone incubated them well
J150 RAPTOR RESEARCH Vol. 6, Supp. J

enough to start them growing. could see the embryo two weeks later. The
I

tiercel was totally passive. This year the same thing happened, except I started
the light later— on March first. The eggs were laid while we were gone skiing at
the end of March. When we came back, she had a clutch of four again and the
tiercel helped incubate. But, again, they were not fertile. They behaved very
well. The tiercel was very aggressive; the falcon was not. She was calm and
broody, so I did not disturb the eggs for 32 days. Then I realized they were no
good and it was too late to recycle. The moral of this story is: no matter how
good it looks, candle on the ninth or tenth day.
The third pair are the Peale’s and there is not much to say about those be-
cause that pair I will leave alone for natural breeding attempts, not even any
light stimulation, because I think they have the potential to do it on their own.
The tiercel is three; the female is six. They are eyasses. The female has raised
foster babies of her own species in British Columbia. This year, just for the
heck of it, I gave her a two week old Prairie Falcon. She immediately adopted
it and fed it, but the tiercel did not participate in the feeding. She is a good fos-

ter mother. She is of the right age. The tiercel has been broody for the last two
years even though he is young, so I hope that next year, just by leaving them
alone and giving them good food, they might do it the natural way. It seems
like the natural way may prevail over the injectable way or tablet drug culture
way, but time will tell. We are just at the beginning.

LAWSON. I think first of all the key in the chart from Stan Temple’s presen-

tation the very first block, the external stimuli. Unfortunately it is not always
is

practical to create rock cliffs and Colville River in your breeding room and I
think that most people will agree now, as we have talked about earlier, that the
birds seem to realize where they have come from, where they were born, and
they seem to always do better in a familiar environment. After they get there,
the external stimuli that both initiate this process and keep it going after it is
initiated are vital. For example, if you take Kestrels and put them in a breeding
room and do not supply them with a nest box, you are wasting your time. They
just will not do anything. The nest hole seems to be essential to success. When
you take these birds out of their normal habitat, you are confusing them, com-
pletely. What we try to do with the gonadotrophic hormones, at least my ra-
tionale is, that we are trying to bypass the external stimulus needed to initiate
breeding. I have been criticized, and very rightly so, by Stan Temple and John
Snelling in that my experiments are not controlled so that I can say 1 00% that
my hormones are doing the job. I am giving them hormones, increased light,
warmer temperatures, and total isolation so that the limited success I claim
could be any of these or all of them together. I go along with Dr. Halliwell; I
do not care if I have to hang an Indian medicine bag in there to get them to lay
eggs; I will do anything and this is the basis for my using these hormones. My
other big misunderstanding with all breeding projects is that everybody seems
to feel that these birds are not affected by human presence. Dr. Meng is living
proof that I am completely wrong on this, but I am going to say it anyway.
1972 Breeding Conference— Endocrinology J151

Adrenalin, epinephrin, is proven to be directly antagonistic to follicle stimulat-

ing hormone. When you walk into the room to feed the birds and they jump
around being really ill-at-ease, you are destroying everything that you have tried
to build up in these birds. It has been shown that adrenalin is directly antagon-
istic to follicle stimulating hormones. Something else that Stan said was that if
you are going to try these hormones, you must inject at the proper times and in
the proper amount. I will go along with proper time, but I do not think the
amounts are that critical. I do not think you are going to kill them with preg-
nant mare’s serum. When I did this with Kestrels I gave these four-ounce birds
one cc of pregnant mare’s serum each day, injected intermuscularly, for five
days running. That is about half of the dose you would give a 50-pound dog. So
I am really whopping it to them. Dr. Awender seems to correlate that you have

to increase the dosage far above what you would calculate for the weight of the
bird. You have to do it at the proper time or you are defeating the bird’s natur-
al mechanism. If the gonads are beginning to develop and you give them follicle

stimulating hormone or testosterone, the chances of success are nil. But I just
do not believe that the amount that we are giving is that critical. I started off
with Kestrels and we did perform a laparotomy on these birds to check the gon-
ads before we started. The birds were captured about 35 miles apart on purpose
to preclude their being a naturally mated pair, and we did this in mid- December.
We gave the birds five injections and then left them alone, trying to initiate a
response with hormones, and then let nature take its course. We let increased
photoperiod and lack of disturbance take over. We have done this with Kestrels
about five times, with Red-tails twice. With the Kestrels we got copulation in
I I days from capture and eggs laid in 33 days. With the Red-tails we never did

see copulation, but we got fertile eggs 47 days from capture.

STODDART. When did you give them more light?

LAWSON. Immediately, 16 hours of daylight.

STODDART. You gave them injections of hormones?

LAWSON. We gave them one injection a day for five days. Before we even
put them in the breeding room, we gave them the first injection.

VOICE. The tiercel as well as the falcon?

LAWSON. Both.

MARCUS. Tim, when you gave five shots did you keep them jessed up for
five days and catch them?

LAWSON. No sir. These were wild birds; they were not manned. They were
kept behind a one-way glass all the time with no external disturbance whatso-
J152 RAPTOR RESEARCH Vol. 6, Supp. J

ever as far as sight is concerned, just the noise that normally occurred.

MARCUS. How did you give them shots for five days?

LAWSON. Catch them at night. They never saw us.

STODDART. Do you think that causes adrenalin production, i.e. handling

them at night?

LAWSON. Not nearly like going into the room would cause. It takes two
minutes to do this.

HUNTER. In conversation with you before, we discussed the site of injec-

tion with Peregrines.

LAWSON. We were giving these birds 6 cc a day which is a whopping dose

we went to the subcutaneous tissue be-
for the breast muscle of a Peregrine, so
hind the neck where you are going to get equal absorption. I am sure that you
are going to get the same results.

SCHUBERT. You put it all in one area and it absorbed?

LAWSON. We did not have any trouble with soreness or any sign of discom-
fort whatsoever. In correspondence with Dr. Graham, I am sorry he is not here,
he mentions that he had trouble with what he thought was a bad reaction to
this foreign protein. Pregnant mare’s serum is serum from a pregnant mare, just
like it says, and it is a foreign protein. I think everybody is familiar with serum
sickness or has heard about it. It has been shown that you can take a guinea pig
and give it a shot of pregnant mare’s serum, wait ten days and give it another
shot and it will die in 30 seconds right before your eyes. But we have had abso-
lutely no problems with this in our birds.

STODDART. Where were the Peregrines from that were used?

LAWSON. Don Hunter’s Peregrines.

STODDART. Were they tundras or Peale’s?

LAWSON. They were tundras.

HUNTER. Maybe you ought to explain what the Peregrines did.

LAWSON. I’m sorry you asked that. The first pair of birds Don sent me, the
female had been in captivity I don’t know how long, but she was pinioned on
one side.
1972 Breeding Conference— Endocrinology J153

HUNTER. She had a bad wing.

LAWSON. The tiercel was fine. I wish I had my notes here with me, but I do
not.I forget how many days it was exactly. It was around the normal time that

we would expect to see some action. The female began to cluck and show amor-
ous tendencies toward the tiercel and beg to be fed. She started to scrape and
the tiercel would not run away from her. He would sit and eat within a few
inches of her feet and a couple days later she died of what was diagnosed by
the pathology department at Ohio State as gout. But, on autopsy, the follicles
on her ovaries were about an inch in diameter, which is a lot bigger than they
should have been if she was regressed.

MARCUS. Was the biggest one an inch in diameter?

LAWSON. That was the biggest one. And they were graded all the way down.
Somebody asked me when that was. It was the 29th of February.

MARCUS. Have you ever done any repeats on this?

LAWSON. Just with the Kestrels. Just the five trials with Kestrels and twice
with Red-tails. The second time with Red-tails was with the same pair. I gave
the hormone without doing another laparotomy to see the stage of the gonads.

MARCUS. What do you mean by a laparotomy?

LAWSON. A laparotomy is just to make an incision and look at the gonads,

physically, to see what size they are.

MARCUS. Is there any work similar to yours without exogenous introduc-

tion of hormones?

LAWSON. Cade and Willoughby of Cornell did this with Kestrels. We pat-
terned our experiment after them almost totally with the exception of the hor-
mones. They got an average of 54 days with variants of 52 and 61 days before
eggs were laid by their Kestrels. That is the sole basis on which I base my deci-
sion that maybe these hormones had something to do with getting the first egg
in 33 days in our experiments. I have no controlled experiments using eight
hours of daylight and hormones to prove that it was my hormones that did it
exclusively.

TEMPLE. Tim, were your birds maintained in a heated building?

LAWSON. The temperature in the building was about 40 degrees.

TEMPLE. Temperature definitely is shown to affect the rate of development

J154 RAPTOR RESEARCH Vol. 6, Supp. J

of gonads. The pilot experiments with American Kestrels that Cade and Wil-
loughby did, were done in January outside in Syracuse, New York, which was
very cold. This might explain the time difference.

LAWSON. Absolutely.

SNELLING. Tim, could you tell us what happened after the Red-tail’s eggs
were known to be fertile?

LAWSON. They were put in an incubator and the incubator went wild at
night and killed three of them. What we should have done, hindsight is al-
all

ways better, was just leave them alone, and let them recycle on their own, but
we panicked and initiated new injections. I am sure we fouled them up.

HALLIWELL. was unfamiliar, or did not catch exactly what you did with
I

these Peregrines. injected 6 cc in one massive dose, subcutaneously, behind

You
the neck-one dose and then sat back to see what would happen.

LAWSON. Not one dose, five doses.

HALLIWELL. Six cc for five days?

LAWSON. Right.

HALLIWELL. Whereas in your Kestrels and in your Red-tails you gave one
cc per day for five days. All of the injections were subcutaneous, not intermus-
cular?

LAWSON. No, the Kestrels’ doses were intermuscular, the Red-tails’ were in-
termuscular, and the Peregrines started out to be intermuscular for the first two
days, and then we got to being afraid that maybe we were going to cause some
kind of myasitis by giving this in the breast muscle, so we went to the skin be-
hind the neck, which is voluminous. You could inject probably 1 0 or 1 5 cc back
there and still have a little room. It is a much better place.

SHULTZ. If you had success with one cc in the Red-tails, why would you
use a six times larger dose with Peregrines?

LAWSON. The Red-tails were given three cc for five days. We took that fig-
ure right out of the air; no basis behind it whatsoever, other than it was a mas-
sive dose for a bird of that weight. We had previous luck with a massive dose in
Kestrels, so we just took three cc and then we went to six in the Peregrines
after we failed to get results the second time in the Red-tails.

SHERROD. Did you use five doses of six cc each, or five doses of three cc on
1972- Breeding Conference— Endocrinology J155

the Peregrines?

LAWSON. We gave five daily doses of six cc per bird.

PLATT. In your opinion, what do you think happened to the female Pere-
grine? It was not gout.

LAWSON. I do not know. I found her in trouble in the morning at 10:00 and
at noon she was dead.

STODDART. Would follicle size have any effect on it? Were they not partic-
ularly large follicles?

LAWSON. Oh no, that would not make her die, if that is what you mean.
That was the response I was trying to get— increased follicle size.

STODDART. Was that a normal-sized mature follicle?

LAWSON. Not quite; it was growing; it had to grow half again that big before
it would be ovulated.

STODDART. I thought you meant that it was overly large.

LAWSON. No.

STODDART. Now what?

LAWSON. Now what? I came in the Army in July and I have been on TDY
since then and have not had enough time to get back to it, but I am going to.

STODDART. This was done on a Ph.D. at the University?

LAWSON. No, this was done on my own time while I was attending veterin-
ary school, but not in association with any university. I used some university
facilities because I begged for them, but that is the only connection.

STODDART. Is anyone continuing the work there?

LAWSON. Nobody is there now to do it, no.

MENG. Just one comment. You mentioned before the adrenalin that my
Peregrines secreted when they were frightened. Actually they were not fright-
ened. I did not subject them to fright actually; I did not go in there and scare
them because I feel that is very bad; that just fouls everything up. I approached
the cage from the outside, put food there in a very cautious manner and then
J156 RAPTOR RESEARCH Vol. 6, Supp. J

he started defending his territory. I cowered away and just took off, but it was
not to the point of his getting frightened. He did not secrete that much adrena-
lin. If I went in the enclosure he would get frightened and that would probably

have defeated the whole purpose. It was just like in the wild where they defend
their territory a little bit, but they were not frightened to the point where he
went the other way.

TEMPLE. I might add— Tim has seen the paper— the paper that proves that
adrenalin will block reproduction only if given in unphysiological or massive
doses. The doses were given attempting to use it as a birth control technique
in mammals. So, probably, the adrenalin secretion that you would cause by get-
ting a bird to defend is probably insignificant. I want to propose something to
you to see what the general opinion I am doing my Ph.D. work on just this
is.

problem. I ammeasuring all four of these hormones, FSH, LH, estrogen and
testosterone in Starlings because I can get hold of lots of Starlings around here
and I have the facilities at hand to measure the amount of these hormones in
the blood. How many of you—just to see whether it would at all be workable-
how many of you who next spring fail in a breeding effort, would be willing to
allow me to take two cc of blood from your bird and analyze it for these sex
steroids? All I need is about two cc of blood and I can do an analysis. For Pere-
grines that is nothing; they can take that loss of blood very easily. It does not
even have to be Peregrines; it can be other species. If I could measure the sex
steroid levels in the blood, I could probably say whether the bird does in fact
have gonads that are functioning normally.

STODDART. What about the females that are laying eggs?

TEMPLE. I would not want to touch it; that is why I said an unsuccessful
attempt.

STODDART. What if the eggs are infertile?

TEMPLE. Okay, then.

STODDART. If you candle those eggs on the ninth day, let’s say, and you
analyze the blood that day, what are the chances of establishing good baseline
levels for that point in the breeding cycle?

TEMPLE. What it would do is give us a standard and I could then compare. I

could say, “Okay, this is what her sex steroid level is; let’s take a sample from a
bird that does not lay eggs; what is her sex steroid level?” If it is lower, then
you would definitely say that her ovaries are not developing; she is not produc-
ing sufficient estrogen to supply the vesicle.

HUNTER. If you can work this technique out first, you can do it with one
1972 Breeding Conference— Endocrinology J157

drop if you wantto. Could you check the phospholipid levels? There is a corre-
lation, too, between the phospholipid, apparently, and the state of gametogene-
sis.

TEMPLE. is especially true of the male. Phospholipids are natur-

Right. This
ally occurring fatty types of chemicals. They are precursors for estrogen and
testosterone. They are involved in the production of these hormones.
*
s

SMYLIE. What would be the requirements for getting the blood to you?

TEMPLE. Physically what you have to do is collect the blood and immediate-
ly centrifuge it to separate the plasma portion of the blood. You have to have a
doctor do this who has a centrifuge. It is very easy to do. You freeze the plasma
immediately.

AWENDER. I would like to give a chance to our other panel member, Dr.
McIntyre, to present his case and then we will have a little more time for ques-
tions and answers. I know this topic has a lot more questions than answers, but
let’s get to one more panelist, please.

McINTYRE. I really do not have very much, because we have not fooled
much with hormones. I am going to throw out something here that perhaps
some of you know and some do not. When I first got into this business, Dr.
Berthrong, a pathologist at Penrose in Colorado Springs, had done a lot of post-
mortems on falcons. He.asked us to look at the adrenals every time we posted a
bird. He thought the adrenals were much smaller in captive birds than they were
in the wild. We tend, also, to believe this. What effect do small adrenals have
on the stimulation of sex hormones, ACTH, corticosteroids, and so forth, I real-
ly do not know .but some of you experts may, perhaps.

TEMPLE. On this point, the one naturally occurring stress hormone, a hor-
mone that is produced when an animal is under stress, is corticosterone pro-
duced by the cortex of the adrenal. This is known to inhibit production of gon-
adotropin. As its name implies, corticosterone is a steroid. Structurally it is
very similar to the sex steroids. This circulates in the blood. When a bird is
stressed, the titers are high, it goes to the hypothalamus. The hypothalamus is
sometimes (pardon me for being anthropomorphic) is sometimes sloppy in read-
ing the message. Many different types of steroids will be identified by the hypo-
thalamus. It is shown that if you inject corticosterone in physiological doses
into a breeding bird, you knock out the negative feedback system because the
hypothalamus interprets this steroid coming from the adrenal as a sex steroid.
This is another thing we are going to do at Cornell. We have Red-tails that breed
perfectly normally— that are at ease, and we have others that do not. We are
going to take blood samples from them to measure the corticosteroid level and
see if there is a contrasting difference. This will be done next year.
J158 RAPTOR RESEARCH Vol. 6, Supp. J

EBERLY. Is there any variation at all in the various brand names of pregnant
mare’s serum?

LAWSON. Let’s see, the one I was using is called Gonadin. I can not remem-
ber who makes it.

VOICE. Upjohn.

LAWSON. I think Donomone is another one.

AWENDER. I think Entruitrin-S might be another one.

HUNTER. If you have any more questions, you can direct them to some of
the other people here for just a minute. We are going to proceed next with the
artificial insemination part of the program during which there will be some
demonstrations. Immediately afterward we will take a break and there will be
some slides outside which you can look at, hopefully some live sperm under
the microscope.

MATTINGLY. In your presentation, Dr. Awender, at a certain stage you said

that laparotomy was out of the question. Might I ask you why this was?

AWENDER. Yes, I simply did not want to hurt them. I could do it easily. I
cut and sew in the morning, every day in the morning; I could do it at home in
the afternoon, too. But I do not like to do it without anesthesia. I know birds

can stand it. have operated on them when it was necessary. I would much
I

rather do an open reduction and internal fixation on a broken bone of a hawk

without anesthesia or very little anesthesia. They can take pain better than anes-
thesia, I think. I thought this was going to throw them too far off to grab them;
to make a little incision there, then look, and then sew them up again. I thought
it would not be the right thing for me to do with one sample, with one pair. If

you have a bigger sample to work with, then it becomes statistically important.
Here I know my explanation would have been she did not lay eggs because I cut
her.

MATTINGLY. The question is being mumbled around back here. I think we

saw the same thing when Stan asked about taking blood. I think we are a little
too over-protective about our birds. Actually, do you hurt a bird when you lap-
arotomize it?

AWENDER. I think they have some pain and, probably, it involves enough
handling that if you do it with or without an anesthetic it may have far reaching
effects the next day or two days later. We simply do not know. I favor experi-
mentation with dispensable birds. Take buteos; take Kestrels if you must. I was
not going to use my Peregrines. That’s all.
1 97 2 Breeding Conference- Endocrinology J 1 5

MATTINGLY. But, we are not getting any information.

AWENDER. I admit this is quite a shortcoming. I was not about to do it. I

am not that scientific.

MATTINGLY. By the way I have been working out dosages for a new very
effective anesthetic called Ketelar or Ketamine.

AWENDER. Ketamine. Yes, we use it.

MATTINGLY. Very, very good. I have no worries about it. You can double
the doses.

AWENDER. Yes, it splits the mind. I have not tried it, but I know people
who do and it’s OK. On people it works fine. You hurt them and they do not
know it.

HALLIWELL. I think concerning anesthesia in these birds also used we have

Ketamine and have had very good success with it and sizes of birds
in all ranges
from Golden Eagles all the way down to several Kestrels. We feel we have had
better success using the inhalation anesthetics of which you are familiar. There
are probably two on the market today: methoxy fluorane which we have not
had as good a success with as we have had with the also halogenated ether called
Halathane. And here, again, with Halathane we have done an extensive amount
of work, although I will admit this has been done in a veterinary clinic where
we are able to monitor blood pressure, heart rate, EKG, and so on. Because we
have had excellent success with it, I feel reasonably competent to use it in a
field situation. We have done over two dozen Kestrels, without a death. I pinn-
ed a wing on a baby owl approximately six months ago; we have also used it on
the larger birds. I think very definitely that it would be worth using and I would
be willing to talk to anyone who would like to undertake this sort of project. I
think with either of these drugs the risk is not great at all used under judicious
care by somebody who has some reasonable amount of experience.

SNELLING. With respect to laparotomy I must admit that I feel the same
sort of twinges of pain when I think about laparotomizing a Peregrine. How-
ever, there has been some extensive laparotomy done on haggard Red-tails by a
man who could not come today. He is a graduate student at Cornell now. And
he did a similar study with haggard Red-tails injecting levels of PMS into them
several years ago and he laparotomized these birds, I think, two or three times
a week, completely putting them under with anesthesia every time he did it. His
sample size was approximately 25 to 30 haggard Red-tails. He never experienced
any difficulty. So, judging from the experience of poultry people who will tell
you that a laparotomy is nothing, I really think that the risk is pretty minimal.
Perhaps some of us should begin to think about this technique.
J160 RAPTOR RESEARCH Vol. 6, Supp. J

TEMPLE. might add that when we do laparotomies on poultry or pigeons,

I

for instance, we do not even use anesthesia. The incision that you have to make
is in the very thin wall of the rib cage. It is not an area where the animal is go-

ing to experience a large amount of bleeding as long as it is done correctly. It is

a very simple thing, but it takes experience.

AWENDER. I think we need a of help; more work needs to be done.

lot
What the Cornell people are doing wonderful, and I would suggest that they
is

go into it with all their might and do a lot of experimenting. I expressed here
before that I was unwilling or emotionally unable (or name it what you want)
to experiment with the birds in my possession, and I think a lot of small breed-
ers may feel this way. We are too close to the birds. I think an impersonal ap-
proach is needed. One thing that was not mentioned here in endocrinology is
this— we have been working on material that has been derived from veterinary
sources or from human medicine. It is possible that there is a species specificity
of some of these things. FSH as we know it may not work in birds and the only
analogy I can draw is this: about 15 years ago or so I was dreaming; I thought
in those days I would never have a Gyrfalcon in my life so I was going to make
a Prairie Falcon grow bigger, and I was going to give it growth hormone. I could
not get any, so the closest thing to growth hormone was a human pituitary ex-
tract closely resembling the pregnant mare’s serum, a thing called Entuitrin-S.
This was supposed to be contaminated with growth hormone and I thought I
would utilize the contamination and I shot this Prairie. I got this eyass as small
as I could; it still was about a ten day old chick, but I thought the skeletal
growth was still backward enough. I gave it daily injections of Entuitrin-S and
it developed into a beautiful, normal, lovely Prairie, no bigger, no smaller. And

then I contacted Dr. Riddle who had done a lot of research in the early 1930’s
with prolactin; this hormone was not mentioned here. It is a very definite avian
thing. It does not have much practical value and so we thought prolactin might
be a growth stimulating factor or the growth hormone for birds. It just goes to
show that what may be growth hormone for mammals may not be the same
thing at all for birds, and that goes for all other hormones. Some of them are
definitely showing effects; some of the others are not; I think this making ex-
tract from pituitaries of birds will become a major project. It has been suggested
to me to hire some high school kids and let them take the things out of chicken
hatcheries and so forth, but the little fellows cannot do it. I would obviously
appreciate some help from Cornell, and I would gladly inject some stuff and
watch one or two birds. But, I cannot make the stuff and I do not think most
amateurs can.

HUNTER. I looked into this avian derivative of Follicle Stimulating Hor-

mone several years ago and I ran into a brick wall.

AWENDER. Right.
1972 Breeding Conference— Endocrinology J161

TEMPLE. We do at Cornell. I might say that to get usable amounts you go

through tons of chicken heads.

HUNTER. Yes, that is I had a serum company tell me they would make
right.
it for me, but I hate to tell you what they said it would cost.

SHULTZ. Stan Temple has mentioned that if we would send him two cc of
blood from our birds that he could run some analyses, and I think this would
be extremely useful and extremely helpful. From this group, I am sure that we
could get enough samples to make it significant. Would you repeat this for the
benefit of the people who were not listening?

TEMPLE. I think what I will do is publish a note in the next issue of Raptor
Research News that is going to get to everyone, rather than right now.

VOICE. Do you only want falcons or do you want buteos and accipiters or
what?

TEMPLE. I can assume the cost of some of these analyses, but every sample
isgoing to cost me about $10 and about 16 hours of work to analyze it, so I
would prefer to limit it to Peregrines for the time being because of the time and
expense involved.

HUNTER. It certainly was my feeling during this reproductive endocrinology

session and has been for quite a long time that this particular field does have
some real possibilities, although I do not think that it is anything for, as Dr.
Awender says, the little fellow. It needs to be done under controlled conditions
and I am hoping that Dr. Lawson will find time to pursue his work a little fur-
ther.
RAPTOR RESEARCH
Volume 6 Supplement Part K 1972

SPECIAL CONFERENCE ON CAPTIVITY

BREEDING OF RAPTORS-A REPORT
Sponsored by Raptor Research Foundation Inc, ,

Held at Sioux Falls South Dakota, U.S.A., November 22-23, 1971

,

Part K. Health and Nutrition of Breeding Pairs (Panel 3)

edited by
Byron E. Harrell
Biology Department
University of South Dakota
Vermillion, South Dakota 57069

Panel Members: William H. Halliwell, Chairman, David E. Allen, C. Wendell

Carlson, James McIntyre, Richard D. Porter.

INTRODUCTION-NUTRITION

HALLIWELL. I think nutrition is well exemplified by a little joke I heard a

while back. It is a story about a farmer. One of his banker friends said, “Farmer
Jones, why don’t you go back to school and learn more about farming and then
you could come back and do a better job of farming and have better crops and
increased productivity.” And Farmer Jones said, “Shucks, Banker, that won’t
do any good, I don’t farm as well as I know how to now!” I think the same
thing applies to those of us trying to breed birds of prey, and I speak in particu-
lar reference to the nutrition of birds of prey. What I have tried to do today is
to go through some of the scientific basis of fact for the nutrition of birds of
prey.
Pure and simple, nutrition is the transformation of food into living tissue,
but this takes a lot of things from the time you lay it out to the bird until the
time the bird' transforms it into living tissue. In order for a diet to be adequate,
it must first provide food for sufficient energy. Second, it must contain all the

essential nutrients in correct quantities. Thirdly, it must be digestible and have

a high biological value. By biological value we mean how many nutrients it has—
vitamins, minerals, carbohydrates, proteins, fats. All essential nutrients must be

K163 Raptor Research 6(Supp):K163-K187, 1972

1972 Breeding Conference- Health and Nutrition of Adults K1 67

CARLSON. No. What you are doing by the high protein diet is increasing the
number of R-complex vitamin requirements. But if they are satisfied it should
not really hurt the animals. Protein is just as good a source of energy as starch.

HUNTER. I mean during the reproductive stage . . .

CARLSON. I suspect the first thing you might notice is smaller egg size if

energy is really markedly deficient. I think if they had plenty of protein they
would still be able to concentrate it.

GRIER. I don’t remember the trace experiment. There was a man who studi-

ed and there were two groups one of which was fed chopped up feath-
rickets,
ers from chickens that had been irradiated and got rid of their rickets. The other
group had not been irradiated and did not lose their rickets. So I think there is
some possible Vitamin D synthesis going on here.

TEMPLE. One thing you have to be aware of, you are talking about a bird
where this ultra-violet light is not reaching the skin. A bird has very little expos-
ed skin-the synthesis is taking place apparently in birds on the feathers them-
selves not reaching the skin. There is no way that ultra-violet light could reach
the bulk of the skin on birds. It reaches of course on the face and beak.

WHITE. In birds it is hypothesized that the reason for preening is to take oil
out from the uropygeal gland, put it on the feather, and then as they repreen,
the oil on the feather is irradiated and as they repreen they then take this back
into their mouth.

GRIER. I think that whole thing has been pretty well discounted. For exam-

ple, the experiments were repeated but without success, and there have been
recent literature and reviews pretty well discounting the whole thing; it seems
they do get enough radiation through the feet and lores and ceres.

TEMPLE. What he did when he discounted was to say that probably the
irradiation of oilon the feathers into Vitamin D is minimal compared to the
dietary sources. The primary sources of Vitamin D are dietary, coming from
whatever the bird is feeding on.

HALLIWELL. The and Vitamin D 3 or usable

relationship of ultraviolet light
metabolically active Vitamin D and the absorption of calcium from the gastro-
intestinal tract was brought up and I wonder if you would speak in the general
area of Vitamin D production in birds as well as the utilization of Vitamin D.

CARLSON, If birds are exposed to about 1 5 minutes of sunlight a day the

kind of sunlight we have most of the time in South Dakota you shoudn’t have
to worry about Vitamin D. Of course this isn’t always the case and when you
1972 Breeding Conference— Health and Nutrition of Adults K165

that egg, the eggshell must sustain of the embryo through hatching. Maybe
life
that’s the reason we are getting some of the
early embryonic death; that is what
I am really trying to say. I went through the literature for 1 969, 1 970, and 1971

over the past months and I have tried to pull out facts from the literature per-
taining to early embryonic death as well as to the adult male and the adult fe-
male.
I would just like to list these specific entities that were directly related to

health and nutrition of adult birds that would be interesting to you, such as
Vitamin A. In studies with poultry when you decrease the amount of Vitamin
A, you decrease fertility of males, decrease fertility of females, get embryonic
death at two days after laying the egg. The young embryo metabolizes almost
all of the Vitamin A in the period of time from the laying of the egg until that

embryo pops out of the egg. And this is one of my reasons for being somewhat
reluctant to suggest or even condone the feeding of day-old chicks. I think Dr.
McIntyre has corrected this situation himself very excellently and I really think
I agree with him as far as ease to the individual falconer. You can buy the baby

chick locally or even buy them from Sears and Roebuck for a very reasonable
fee, put them in a brooder, grow them to young adulthood about four to six
weeks of age. Feed them on a good quality starter feed and follow the recom-
mendations. Feed their best feed, take good care of these chicks until they are
four to six weeks of age; at that age take them out of their brooder, wring their
necks, put them in a baggie and put them in your freezer. Don’t gut them, don’t
pull the feathers off them, cut the feet off them, leave the head on and put
them in your freezer. As you need, pull them out and feed your birds. What are
you doing here? First thing, trichomoniasis, what we call frounce and the pigeon
fancier calls canker, will not exist after 18 hours of freezing so you can take the
most trichomonad infected pigeon, freeze it for 1 8 hours and you can feed it to
my birds.
The next thing I want to discuss is Vitamin C. You are all familiar with Vita-
min C or ascorbic acid— it’s been known since the day of the limeys. Low qual-
ity, low amounts of Vitamin C in the diet are reflected in poor albumen quality
meaning low protein for the albumin of the egg. Now I don’t know whether
this has any effect at all on the livability of the young chick that’s developing
in the egg but it seems to us that if the protein quality is down, the chick has
less chance of survival. So I’m just mentioning it for what it’s worth.
Vitamin D can be fed as an inactive form which can be activated by the ac-
tion of ultra-violet light with the skin. But you can also feed the active form.
So I think you ought to make amends one way or the other to have vitamin D
in this bird. Vitamin D is absolutely necessary for calcium absorption by the
intestinal tract. You can feed a bird pure calcium phosphate rock and if he
doesn’t have enough Vitamin D, he won’t absorb any of it, it comes out the
other end the same kind of rock. The lack of Vitamin D inhibits egg production
which again is tied up with the calcium.

Choline methionine, and Vitamin B ]2> high levels of these nutrients, increase
,

egg production and increase egg size in birds. Vitamin E is involved in feather
K166 RAPTOR RESEARCH Vol. 6, Supp. K

coat; it’s involved in ovarian and testicular function. It seems to have a very
good ameliorative effect on stress, meaning if these birds go into a captive situa-
tion after being trapped, perhaps they are utilizing Vitamin E at a faster rate
and an adequate supply of it may help us acclimate these birds to captivity.
Calcium I have already alluded to, but to stress it further, normal laying hens
when put on a deficient diet, reduced their laying ability by more than 50%
within a week or ten days and that’s a pretty significant thing to consider.
Diethyl stilbestrol and methyl testosterone seem to have a relationship to egg
production and fertility.
Phosphorus and calcium go together. They have to be present in sufficient
quantities, they have to be present in the correct ratio, and they have to have
Vitamin D for this. Sodium is also required in the diet. Selenium seems to be
important in raising young chickens; on a selenium deficient diet they got poor
growth in young chicks. Manganese is another mineral; it again is tied in with
the calcium and phosphorus ratio. Magnesium is the same type of thing.
Protein. Poultry grow best on a ratio of approximately 20% protein. This can
be lowered with commercial poultry. However, they find that for unadulterated
protein, you need about 18 to 20% for the chickens to maintain good growth
rate or good egg laying rate.

SWARTZ. Arginine as well as methionine are I believe essential amino acids

in birds. They are also important structural components in feathers and hair. At
least it occurred to me as a possibility that maybe here when the ingestion of
feathers might have some meaning in terms of a specific nutrient.

HALLIWELL. To further elucidate that, methionine is the first limiting ami-

no acid meaning when you have decreased protein synthe-
in protein synthesis,
sis it is probably due to a deficiency of methionine. Now I am not willing to

express myself on whether all parts of the body contain methionine and other
amino acids, but you are correct in that feathers do have quite a bit of methio-
nine.

VITAMINS

CAMPBELL. Why do you add Vitamin A and carotene to the diet?

HALLIWELL. I frankly don’t know why they added both. Carotene is a pre-
cursor of Vitamin A, meaning that carotene is converted into Vitamin A. It may
also go into some other enzymatic reactions and just about every reaction in the
body is controlled by enzymes— it may fit in there or some other reason un-
known to me.

HUNTER. Do you think there would be any chance of getting into difficulty
when there is an imbalance between protein and energy, in other words you
have very high protein and low net energy?
 iil v^flrLKU J- pi
LJU- uy
;

1972 Breeding Conference-Health and Nutrition of Adults K1 67

CARLSON. No. What you are doing by the high protein diet is increasing the
number of B-complex vitamin requirements. But if they are satisfied it should
not really hurt the animals. Protein is just as good a source of energy as starch.

HUNTER. I mean during the reproductive stage . . .

CARLSON. I suspect the first thing you might notice is smaller egg size if
energy is really markedly deficient. I think if they had plenty of protein they
would still be able to concentrate it.

GRIER. I don’t remember the trace experiment. There was a man who studi-
ed rickets, and there were two groups one of which was fed chopped up feath-
ers from chickens that had been irradiated and got rid of their rickets. The other
group had not been irradiated and did not lose their rickets. So I think there is
some possible Vitamin D synthesis going on here.

TEMPLE. One thing you have to be aware of, you are talking about a bird
where this ultra-violet light is not reaching the skin. A bird has very little expos-
ed skin-the synthesis is taking place apparently in birds on the feathers them-
selves not reaching the skin. There is no way that ultra-violet light could reach
the bulk of the skin on birds. It reaches of course on the face and beak.

WHITE. In birds it is hypothesized that the reason for preening is to take oil
out from the uropygeal gland, put it on the feather, and then as they repreen,
the oil on the feather is irradiated and as they repreen they then take this back
into their mouth.

GRIER. think that whole thing has been pretty well discounted. For exam-
I

ple, the experiments were repeated but without success, and there have been
recent literature and reviews pretty well discounting the whole thing; it seems
they do get enough radiation through the feet and lores and ceres.

TEMPLE. What he did when he discounted was to say that probably the
irradiation of oil on the feathers into Vitamin D is minimal compared to the
dietary sources. The primary sources of Vitamin D are dietary, coming from
whatever the bird is feeding on.

HALLIWELL. The relationship of ultraviolet light and Vitamin D 3 or usable

metabolically active Vitamin D and the absorption of calcium from the gastro-
intestinal tract was brought up and I wonder if you would speak in the general
area of Vitamin D production in birds as well as the utilization of Vitamin D.

CARLSON. If birds are exposed to about 1 5 minutes of sunlight a day the

kind of sunlight we have most of the time in South Dakota you shoudn’t have
to worry about Vitamin D. Of course this isn’t always the case and when you
K168 RAPTOR RESEARCH Vol. 6, Supp. K
are in captivity you do have to supply it in the diet. As you mentioned there
are Vitamin D’s. Any of the Vitamin D’s that are used for most of our large ani-
mals are not potent for birds. Birds must have Vitamin D3 or calciferol and we
find just recently that this is modified by the animal. And even a more potent
form could be used, but calciferol or Vitamin D3 certainly at the present time
is the most acceptable type.

TEMPLE. I talked with the nutrition people at Cornell and asked them about
the calcium-phosphorus-Vitamin D complex. They came up with a very simple
dietary supplement that answers a lot of the problems. That is powdered non-
fat dry milk. That milk product has a very high and a very well-balanced cal-
cium-phosphorus ratio. Perfect for birds of prey, in fact. It also has calciferol in
it.

CARLSON. Well, I wouldn’t expect it to have very much. I would certainly

not rely on milk as a source of Vitamin D. But as far as calcium-phosphorus,
you can’t beat milk.

THOMAS. The person that wants to use supplements would like to know
what is the best supplement to get your D3 from and what do you have to have
to make it available to the bird? I wonder what
I need to look for as far as caro-

tene. If I get Vitamin A form, is it available to the bird biologically

in liquid
once it gets in the system? You can get pellets or tablets, whatever, it’s got A
and D, but if it doesn’t say is it D2? Is it D3? The only thing that I’ve been able
to find is a stuff called Linotone, it’s a coat conditioner for dogs, and it’s very
high in Vitamin A and D3, it’s got a lot of it in it, and I give the birds a half a
teaspoon split between them every day but I don’t know whether the D3 in this
is biologically available to the bird. I suppose if we could know more about

supplements, as to availability this might be one more practical solution that we

could use.

GALICZ. What may be a source of Vitamin D, artificially or supplementary?

CARLSON. Well, we buy it in ton lots as irradiated calciferol. We have any

number of vitamin supplements that are used in poultry feeds that you can buy
at your veterinary supply house that have potent Vitamin D.

VOICE. What about what you can pick up at a drug store that has the cod
liver extract type of thing?

CARLSON. That would be fine.

VOICE. Does it have calciferol? ,

CARLSON. It has calciferol. You want to stay away from irradiated yeast
1972 Breeding Conference— Health and Nutrition of Adults K169

and irradiated vegetable products. That is Vitamin D2.

HALLIWELL. This brings another question to mind again. People are supple-
menting I have my own opinion but I would
their diets with additive vitamins.
like yours on the danger of hyper-vitaminosis or hyper-nutritionalism due to if
one drop is good five drops is five times as good, a type of syndrome which we
all fall prey to.

CARLSON. Well, of course, of most of the commercial feeds we see we don’t

see a stated potency as far as vitamin activities are concerned. For example, if
we were thinking about Vitamin D in terms of the laying hen about 100 inter-
national units per day should be plenty adequate. We could go as high as, how-
ever, let’s see it’s about a factor of 20, in other words up to about 2000 inter-
national units and still not be toxic to the bird. You can’t do that with some of
the others, however. Vitamin A will cause you difficulty rather quickly if you
get above about 25-50,000 international units per day. So again T would like to
temper that by saying it depends on the type of multiple excesses that you
might give the bird. Chances are that you could go to three or four times the
recommendation and still not hurt the bird or its reproductive potential because
there is quite a wide range between requirements and toxicity as far as most of
the organic vitamins. The same thing cannot be said for the minerals. You can
certainly put in an excess of minerals very rapidly. This should be of greater
concern as far as I’m concerned.

HUNTER. In ruminants it is very important for the utilization of Vitamin A

to have Vitamin E. Now
you suggesting
are that in these frozen chick diets, we
might supplement with Vitamin E? And D?

HALLIWELL. I Frankly don’t think so. Now this is my own personal opinion.
Vitamins A, D, E, and K are all fat soluble vitamins, that is they exist in the sub-
cutaneous, the abdominal, the heart fat and unless some oxidant has gotten into
this chicken, such as long freezer storage, there should be sufficient amount of
these vitamins in the fat on, again over four week chicks.

CARLSON. As Vitamin E and Vitamin A interactions are concerned,

far as
basically it is Vitamin A
that the bird is really needing and Vitamin E is sort of
an antioxidant that protects Vitamin A; at least that’s the latest. We have had a
lot of go arounds as far as Vitamin E as to whether it is really essential or not.
About 15 years ago some work came out showing that it was not essential. Then
five years later some people reported that it was essential and so on. But at the
present time it is thought that selenium, one of the minerals, apparently pro-
duces or carries on all of the activity normally we thought Vitamin E was doing.
If we have an adequate amount of selenium available for the animal we can very
definitely show that the Vitamin E requirement is greatly reduced. So that bas-
ically the large amounts of Vitamin E that we usually put into animal feeds is
K170 RAPTOR RESEARCH Vol. 6, Supp. K

there to prQtect the Vitamin A to a large extent. Well that means of course we
could use other things besides Vitamin E. We could use any one of a number of
commercial anti-oxidants that are available today.

HALLIWELL. Many of the birds that we have in captivity today are over five
years of age and they or may not have been at some time during the first
may
years on a deficient diet. Could you discuss the effect on the reproductive or-
gans of previous nutritional deficiencies. In other words, the bird that is meta-
bolically nutritionally deficient two years ago and on adequate diet today,
would you expect damage to the reproductive organs of a bird today? Or do
you assume that they will rejuvenate?

CARLSON. It depends only on the deficiency that you encounter. As far as

Vitamin E deficiency is concerned, and again it depends on the extent of the
deficiency, if it were an extreme deficiency, of course, it is irreversible. It
wouldn’t do any good to try to correct a bird that had been so handled. But I
don’t suppose that they would have survived either. Generally speaking I would
say unless it is one of these more critical nutrients, if it’s just a marginal de-
ficiency, I believe when you get a bird back on a normal diet that their repro-
ductive capacity would recover as well.

HUNTER. Itseems to me that there is an area here that we need to hit quick-
ly and this is supplementation of vitamins to whatever diet we are using. Can
you go wrong in having too much vitamins?

HALLIWELL. Yes, you can have hypervitaminosis.

MINERALS AND WATER

HALLIWELL. For minerals to be utilized by the body they have to be there

in sufficient amounts and there are interrelationships between them, they have
to be in some particular form so they can be absorbed. For instance, fats will
combine with calcium and form an insoluble soap that is unable to be absorbed
by the intestinal tract, as a matter of fact. If you put calcium in high quantity,
it seems to be absorbed by the intestinal tract by diffusion, in other words there

is no metabolic expenditure of energy, cellular energy to absorb this calcium,

but under situations of low calcium intake there is an active expenditure of

energy by these cells to absorb the calcium that is present within the gastro-
intestinal tract and take it into the body. So this whole thing is wrapped up
again and you can’t talk about one individual mineral as a point.

TEMPLE. An interesting case with calcium would be the female Red-tail that
I used in the insemination work. She was obtained from a very naive farm boy
in Iowa who had had the bird for nine years as a pet. He was not a falconer, just
kept her as a pet; for nine years the bird never received casting. She received
1972 Breeding Conference— Health and Nutrition of Adults K171

nothing but beef heart and beef liver as the staple diet. That as we know is
about the worst food you could put into a bird. When we got her at Cornell we
only had her for about one month before she entered a reproductive cycle for
us and she laid one egg normally. The next egg was laid without a shell. She
went into a hypocalcemic condition similar to what poultry people call egg lay-
er’s cramp from chickens laying too many eggs and getting rid of too much cal-
cium in it. What this bird did was wipe out her calcium reserve with one egg and
the second egg was laid without a shell. In this case we administered calcium
gluconate in the diet in large quantities and the third egg came normally and
was fertile. As I say there is a good chance that early abuse in terms of a bad
diet something like this is easily remedied.

CARLSON. Especially in terms of calcium, if you give them too much cal-
cium during their growing period this can be very serious.

PLATT. You have mentioned the problem of calcium with young birds ex-
posed to overdosage. What’s that?

CARLSON. The bird gets used to excess amounts of calcium and basically
the kidneys are just overworking and you end up with just an overloaded kid-
ney. not going to be able to metabolize calcium later. There is a lot of ne-
It is

crosis as the veterinarians refer to it and gout and things of that nature that the

bird would eventually succumb to. What we would like to do of course as soon
as the bird is about ready to come into production, precede that for about a
week or so with a diet that does have in the neighborhood of, for your type
bird I would suppose about 2% calcium rather than the usual .8% that you are
routinely feeding during the growing period.
Someone asked the question and I will repeat it for him. In terms of what
could be deficient as far as a diet of meat or basically animal food is concerned?
First as a poultry nutritionalist my experience is largely in the area of that type
of an animal where they eat mostly grain or other byproducts of the vegetable
kingdom and very little animal food. In fact today we probably have no food of
animal origin in most of our poultry diet. But as far as meat protein is concern-
ed there is one mineral that could perhaps be deficient and that is zinc. Normal-
ly when we feed high meat diets to poultry one of the things we look out about

or worry about is the possibility of deficiency of zinc and this could be a factor

of course, particularly in reproduction, in that it is required for sexual maturity,

precocity and so on. So this is perhaps something that some of you might want
to give some thought to. The natural diet where they get mostly meat just isn’t
going to carry a whole lot of this mineral for the animal. You may have to sup-
plement with a supplement commercial type feed.

CRAWFORD. Is a deficiency of manganese important?

CARLSON. This is true if manganese or some of these trace minerals were in

K172 RAPTOR RESEARCH Vol. 6, Supp. K

the diet at rather marginallevels. You can increase the requirements for almost
any of them by going excessive in terms of calcium-phosphorus ratio. If you
have a good supply of manganese, zinc, and other trace minerals then you can
vary your calcium levels quite widely without any great detriment. That is how
we first discovered the importance of manganese. We put in extra calcium and
phosphorus and sure enough the birds came down with sclerosis. When we put
in 30 ppm of manganese, we don’t have to worry about it.

CRAWFORD. What supplemental dose of manganese is used?

CARLSON. In the poultry case you put in a half ounce of manganese sulfate
per ton of feed, 30 parts per million. Use a trace mineral salt that contains well
about 1 % manganese.

MARCUS. Did I hear you say you found that zinc was important for the re-
productive process?

CARLSON. Yes. It’s one of the more recent findings as far as mineral nutri-
tion is concerned.

MARCUS. Where exactly is that concentrated?

CARLSON. In the ovary. The size of the yolk is greatly reduced in the case
of a zinc deficiency. In fact if the animal is very markedly deficient you don’t
get any reproductive activity at all, even the sexual characteristics.

MARCUS. What I’m trying to find out is, what is a natural source of zinc for
a raptor, a logical natural source for a natural supplement.

CARLSON. That’s a real good question. Meat is very poor. Bone meal might
have some. But zinc oxide, zinc sulfate, zinc salts of various sorts could be in-
corporated into a trace mineral mix to provide you with enough.

OLENDORFF. Does anybody have any ideas about egg binding, egg bound-
ness in females. I understand this has been a problem.

HALLIWELL. Dr. McIntyre, has anybody ever called you about egg bound
birds?

mcintyre. no.

HALLIWELL. Has anybody else experienced this? Frankly, I don’t feel the
expert either; as far as I know it is inability to lay the egg. The ovum is released
from the ovary, the white is secreted, the shell is formed, but the egg is not laid
and that is as much as I know about it. I’ve never been consulted about the
 :

1972 Breeding Conference— Health and Nutrition of Adults K173

malady.

WOLHUTER. Some of the literature of European zoos that are raising birds
mentioned that birds that breed early in the season during cold temperature are
more frequently egg bound, but whether it is indeed a function of temperature,
I don’t think has been proven, but this is one possibility.

LAWSON. Some work done in poultry indicates that it might also be due to
water deprivation. You have to give adequate quantities of water.

HALLIWELL. Which would tie in with frozen water pans.

TEMPLE. I wouldmake another point more closely related to breed-

like to
ing and water intake. think anyone that has had any egg laying females has
I

noted, and it is well-documented in poultry, that water intake goes up tremen-

dously during egg production, because the bird is mobilizing a lot of water to
go into the egg. It is very important that the bird has easy access to a lot of
water when breeding.

COMMERCIAL BIRD-OF-PREY DIET

HALLIWELL. To my knowledge time there are two commer-
at the present
cially produced bird of prey diets and I am quite familiar with one of them. It
is a product called Zu-Preem put out by Hill’s Packing Company (Table 1). I

don’t get any commission, I am not peddling it. I’m just trying to use it as a

Table 1 . Manufacturer’s ingredient list and guaranteed analysis of Zu/Preem

Bird of Prey Diet (Hill’s Division Riviana Foods Inc., P.O. Box 148, Tope-
ka, Kansas 66101).

Ingredients Horse meat, horse meat byproducts, meat byproducts, chicken,

fresh liver, ground corn, ground wheat, fish meal, dried whole egg, dicalcium
phosphate, brewers dried yeast, iodized salt, choline chloride, Vitamin A palmi-
tate, carotene, D-activated animal sterol, a-tocopherol, menadione, niacin, cal-
cium pantothenate, thiamine, riboflavin, pyridoxine hydrochloride, folic acid,
biotin, Vitamin B-12 supplement, ferrous carbonate, manganous oxide, zinc ox-
ide, copper oxide, cobalt carbonate, magnesium oxide.

Guaranteed Analysis
Crude protein min. 18.0% Ash max. 4.0%
Crude fat min. 5.0% Calcium min. 0.4%
Crude fiber max. 0.5% max. 0.8%
Dry matter min. 40.0% Phosphorus min. 0.3%
Moisture max. 60.0%
K174 RAPTOR RESEARCH Vol. 6, Supp. K
good example where we are falling down. The bird of prey diet put out by Hill’s
Packing Company has about 60% moisture which comes up again to the general
area of 70% of body weight being water; protein, it’s up about where the Na-
tional Research Council recommends, 20%; fat, it has an adequate quantity; ash,
remember in muscle meat ash is way down around 1%— here it is 4%. Vitamins—
they do exist in specific quantities in this bird of prey diet and this is calculated
with each batch that they mix up. And so we do have some idea what we are
feeding the birds with this diet. Dave, would you like to talk about your pro-
grams at the Kansas City Zoo and any recommendations you have?

ALLEN. We have a pair of White-tailed Sea Eagles that have been on this diet
for 23months and they have reared progeny in 1970 and 1971. I would like to
show you some pictures and add a few comments to what Dr. Halliwell said.
This the exhibit of the Kansas City Zoo where the pair of Sea Eagles is exhib-
is

ited. This was an old mammal enclosure that was built by WPA in 1940 and
these birds came to the zoo in 1964. We covered the bars with wire and the top
with chain link. That is the pair on the nest. Now there are two nest frames and
I turned in a breeding project questionnaire and there is a diagram in that BPIE

No. 69 (.Raptor Research 4(2): 23-25, 1970). The birds chose the front nest
which I believe is 1 5 or 1 8 feet from the public. And that is where they have

nested and raised their two offspring. Here’s a picture of the nest frame from
the access door. The bottom is about six feet tall and the dimensions are in the
BPIE. Both the 1970 female and 1971 male offspring were much larger than
either of the parents and I think this is probably due to fat, lack of exercise,
and I hope, superior diet; it worked very well. Now the back nest frame against
the wall is the one they used mostly for perching. They can get back on the
back edge under that ledge for shelter from the rain, sun, snow, it’s the only
protection they have.
The diet comes frozen, a five pound package, eight packages to the case. We
feed two portions. A Kestrel portion is about the size of a ball we form and a
larger portion for Golden Eagle. I did not know about the Nebraska product
until today. 1 had heard they were working on it, but didn’t know they had it
out. The price is $12.85 per case and eight five-pound frozen chunks. Now a
few falconers in the Kansas City area had a little difficulty in getting this pro-
duct, but what we do is buy it in carload lots, store it in commercial cold stor-
age and draw it off. Others can draw off it too, and they are charged for it.
But this would have to be worked out with whoever uses Zu-Preem. You have
to have a place to store it and freight on it will be quite expensive. I think you
have to buy 40,000 pounds before they pay the freight. Birds have been on it
for 23 months and they have produced two very large and healthy offspring. I
would like to add that the manufacturer states that the pesticides DDT, DDD,
and DDE are controlled during processing to .088 parts per million. Now I don’t
have any comparison figures today on what food of chickens, fish, horsemeat
contain, but I know this will be considerably lower.
1972 Breeding Conference-Health and Nutrition of Adults K175

HALLIWELL. DDT in regular dog food and other meat byproducts varies,
this ranges somewhere around 20 and 30 parts per million, so you can see that
this is one-tenth.

ALLEN. Now the way it was explained to me is that they use a fish meal,

human grade fish meal, and they have developed a process to remove these resi-
dues to this level, which I thought was significant considering we are talking
about White-tailed Sea Eagles; their previous diet was fish, prepared fish.
To give you an example of what Dr. Halliwell mentioned, in mackerel, which
was one of their favorites, the calcium to phosphorus ratio was 1 to 34. They
were also fed smelt. We had a big problem with smelt to do with sea lions; now
this is not other fish eating raptors, but I think it is significant in that smelt con-
tains thiaminase and we had four deaths of sea lions: we had to give them sup-
plementary injections of Vitamin B even to get them to eat. And further mack-
erel and smelt were used at the St. Louis Zoo and the high unsaturated fats
caused death by steatitis in crocodilians. So our whole approach at Kansas City
was to get not only the Sea Eagles but the rest of the birds off the fish. We have
other carnivorous birds on the bird of prey diet and flamingos as well; we feel
it’s pretty important. You might be interested to know how we got the birds on

this diet. It looks like hamburger, but it’s not ground as finely and if the bird
foots it or carries it, it sometimes falls apart, but Hill’s is working on a chemical
process to bind it together better.

HALLIWELL. Today brought some samples of the product we’re talking

I

about, Zu Preem. It’s ground up and it comes to you frozen. When you serve it
and it thaws, it becomes almost mush and they have trouble carrying it. Hill’s
has now added a binder to this and I have brought samples of that packed in
dry ice. I’ll put out their present product and their new improved product on
one of the tables in the back so you can examine them.

ALLEN. In introducing the birds to the product, we didn’t really have any
difficultiesexcept with one Bateleur Eagle; she still hasn’t accepted it as the sole
ration. We accomplished it in various ways. Birds that had been kept on horse
meat took it right away, the Sea Eagles accepted it in a week. We chopped fish
with chunks of Zu-Preem we even took the viscera out of rodents and stuffed
;

it with Zu-Preem; it works. You’d be surprised how fast you can get them on it.

We had one Kestrel on the diet for 16 months; it was killed by a predator. The
other Kestrels in the zoo now have been on it about four months, so we feel
we’ve greatly improved the diet because prior to this these birds were fed
chunks of horse meat, typical zoo approach to raptors which we’ve already
talked about. Growth is a factor of Zu-Preem that we haven’t gotten into. Some
young raptors were raised on freshly killed rats; as soon as they left the nest, we
started transferring them over to Zu-Preem. We did have three downy Horned
Owls brought in this year that we raised on Zu-Preem, and they were beautiful
birds, had good color, appeared normal, and had no bone abnormalities. And
K176 RAPTOR RESEARCH Vol. 6, Supp. K
we also feed roughage once a week, rats and mice as supplement. Frank Kish,
former Curator of Birds at Topeka, said the Golden Eagles that bred and raised
offspring this year have also been maintained on this diet. And he says that the
castings are normal without any supplementary feeding, but he didn’t raise this
eagle on Zu-Preem, he raised it on rats. We don’t have anything much to say
about the growth.

THACKER. Did I understand you correctly? You said you transferred some
of your fish eating eagles from a fish diet onto the Zu-Preem diet. Have you
found that the birds prefer this diet to a fish diet?

ALLEN. No, that’s interesting. If you offer them any other food in connec-
tion with the Zu-Preem, they will eat the other food first. They will eat fish

first, rats, not horse meat; they completely ignore horse meat.

McINTYRE. I can make one comment on this. I talked the salesman out of
a case for a trial and the birds I’ve got won’t eat it. I’m sure that I could get
them on it in time, but it’s real difficult to feed a bird in your fist with this Zu-
Preem, I can tell you that.

HALLIWELL. I’m not about to suggest to any of you that this hamburger
stuff works very well tied on the end of a lure swung around your head at the
end of a ten foot cord. All your nearby friends will look like they have been
through a hamburger melee. But I think all of us do put our birds up for the
molt. The feathers and the other portions of the body will benefit from this
diet. I think you cannot find a better diet to molt your birds out on.

ENDERSON. Is any casting formed with Zu-Preem?

ALLEN. Frank Kish of Topeka in the International Zoo Yearbook, Volume

Golden Eagle were normal with a strict diet of Zu-
10, states that castings in
Preem; now I’m quoting him. I can see that in Kansas City we feed once or
twice a week whatever the schedule is, rats or mice.

VOICE. A
normal casting would have feathers and things like this— what
would be castings in Zu-Preem?

ALLEN. In Zu-Preem, there are whole chunks of bones and skin, but I have
to confess I haven’t tried analyzing the castings at all.

HUNTER. Now one of the things that you talked about was the dietary dry
matter. I really don’t understand why that is important if all you are going to
do is increase the intake.

HALLIWELL. Dry matter does several things. Dry matter is a portion of this
1972 Breeding Conference-Health and Nutrition of Adults K177

casting. It is also methyl being utilized as a binder to hold this

cellulose that is

mush which is the only

together. Binder can also be and in this particular case
one I am familiar with, Zu-Preem is a fiber base that just provides extention to
the stomach. Really, many of us could get along quite well on an astronaut diet
of pills and a little bit of juice.

MENG. I am
going to say something about casting. Some birds would be
killed by giving them too much casting. A Sparrow Hawk, for example: if you
feed them mice, they'll sometimes swallow all that fur and what does the fur
do in the stomach? It just fills them up and they don’t desire to eat any more.
If you skin the mouse out, then they eat too much and they’ll have that much
more food in there. One thing that hasn’t been brought out, if you feed this
ground up material, I just don’t feel right feeding my birds something like that;
I want freshly killed pigeons, they’re warm. I cut the head and crop off to avoid

this trichomonad problem and the feet off, too. The birds just enjoy plucking
the feathers off and eating this fresh material and I wouldn’t subject my birds
to these meat patties. I just can’t see it. They have to have something like in the
wild.

HALLIWELL. I certainly can sympathize with you and agree with you very

well, but the point of this meeting as far as I’m concerned, is to improve cap-
tive breeding of raptors, and if this means feeding them horse manure that we
can preserve this species, I’m going to be out there with a pitchfork!

STODDART. I agree. Whatever is possible to keep the birds alive so we can

breed them. I’m not trying to cut any comers. What I want to know: has any-
one fed this stuff to a Peregrine for three or four years? I have a Peregrine that
is seven years old that was just given to me for breeding purposes and the bird

was given Zu-Preem for two years, the last two years. That Peregrine does not
have the feather quality, foot quality, or anything like it should be, and I’ve
been pumping whole birds into it and the change is phenomenal.

MENG. Just one quick comment. When these birds feed on, say, the back-
bone of pigeons, they are constantly wearing their beak down, and there are
many birds like my Peales’ that I couldn’t catch and cope their beaks off, it
would upset them too much. By feeding fresh material like this where all the
bones are there they have to pull their beaks are down. My Red-tail is 25 years
old, I’ve never touched her beak, she’s been living on pigeon heads for 25 years,
all the pigeon heads I pull off.

GOBEN. Hiave something on the birds and their beaks growing and eating
this pattie routine. Iwas feeding a lot of beef heart for a while, at one time I
had a bird grow a long beak, and I threw in a rib of a deer; it was a Peregrine,
and that thing just messed around with that thing until its beak wore down, it
looked beautiful.
K178 RAPTOR RESEARCH Vol. 6, Supp. K
HALLIWELL. Thank you. Dave Allen, do you have anything to add to that?

ALLEN. Well, prior to 1 968 in the zoo these birds, a lot of eagles, they had a
lot of Golden and Bald Eagles then that were shot up, Fish and Wildlife Service
brought in, were fed straight muscle horse meat. Now they survived for five to
ten years without any casting material. So it occurred to me from a zoo stand-
point, exactly what is the relationship to health of the bird and casting materi-
al? Eve never understood this and we’ve gotten a lot of arguments about it. I
am sure there is a relationship.

VOICE. Can we get an answer on casting? We have no definitive answer, we

are all around the subject.

HALLIWELL. think there is no definitive answer. I have no personal an-

I

swer myself and think from the wise rash of discussion, unless you’ve got
I

scientific facts all you are talking about is hearsay and personal experience, and
hearsay and personal experience hasn’t been doing it in the past and it probably
isn’t going to do it in the future.

PATUXENT BIRD-OF-PREY DIET

PORTER. I’m here basically as an ecologist who has been involved in the
breeding of American Kestrels, primarily. We do have a group of Peregrines that
we are attempting to breed; these are tundra birds. I am from Patuxent Wildlife
Research Center. We have done some work with Kestrels. We started a colony
of Kestrels there in 1 964 for the purpose of carrying out experiments with pesti-
cides and we are still maintaining this colony. So far as their nutrition is con-
cerned, we started out there, this was before my time, giving the birds ground
beef which was primarily muscle, mixed with some liver and tongues and parts
of beef supplemented with vitamins and minerals, and then we incorporated
turkey breeder crumbled for the purpose of soaking up the excess fluids; these
were ground up. Then the next diet that was used starting in 1966 was a horse
meat diet supplemented with vitamins primarily with Vionate. The first year of
reproduction in 1965 was fair. There were quite a number of the eggs hatch,
and young were produced, but the adult birds ate a number of the young. They
even captured some of the young after they had fledged, so there was something
awry there somewhere. Then in 1 966 with the horse meat diet there was little
if any reproduction. There was also a foul-up, so to speak, in the behavior of the
birds because in some the males were tethered after the females laid their third
egg. The males were tethered because it was thought that in 1965 the males
were responsible for the killing and eating of young birds and the destruction
of eggs. This was before I arrived on the scene and the reproduction was nil. We
had only a few birds hatch eggs.
In 1967 just prior to the nesting season, I placed the birds on a diet contain-
ing whole white rats, one third part, whole white hamsters or mice one third, and
1972 Breeding Conference— Health and Nutrition of Adults K179

one third part chicken necks. We removed the skins from the chicken necks to
get rid of the excessiveamounts of fat and I am not sure after learning what Dr.
Halliwell has to say that this was a good idea. We included chicken heads, and
we put one-sixth part. And we put them in a large cutter-mixer, Hobart cutter-
mixer and cut them up into a very fine texture. Then we added to a 22 kilogram
batch 75 grams of Vionate, well known vitamin supplement, and 126 grams of
bone meal to give just in case the calcium-phosphorus content of the diet was
insufficient. By utilizing this diet we obtained a reproductive success in the first
year it was used equal to that which you would expect in wild Kestrels. And it
turned out to be very successful. However, in the second year with this diet we
ran into embryonic mortality in the colony, and we determined that at least
part of the embryonic mortality was due to bacteria entering the egg and ap-
parently killing the embryo. When the eggs are laid very early in the season they
develop a film of condensed moisture around them and the bacteria enter from
the nest box and apparently multiply in the egg and kill the embryo; however,
there were a number of eggs that have not shown this. We have been unable to
culture bacteria from eggs and we don’t know just what to think about those.
This situation has worsened in recent years. We have sterilized thoroughly the
nest boxes prior to nesting seasons, put clean nest material in and still we get
some embryonic mortality. This last year, 1971 breeding season, the embry-
onic mortality was quite bad. So after what Dr. Halliwell said I think that our
diet, our ground diet may be suspect, I think it should be given a very thorough
chemical analysis to determine whether there were deficiencies in the diet or
not.
We have given our birds water, we have
placed water trays on heaters through-
out the winter so the water is available. Sometimes the birds had no water. At
least they had water in the winter but we had no heater under the water pan
and we viewed no detectable difference in their reproduction or in the birds.
We have been feeding our Peregrines this ground rodent and bird diet. Some
birds have taken to it rather readily. One particular female, a large female that
was captured off the beach at Assateague, very voraciously accepted it; within
the first day it was accepted. And another female trapped the same day on the

beach took literally weeks to accept this diet. We now feed our Peregrines white
rats during the non-breeding season supplemented with an occasional Coturnix
Quail and an occasional Mallard duck. We do not feed them the Kestrel diet. 1
don’t know just what to say about the ground Kestrel diet. It has been very
successful in the past but as I have indicated I believe that it is very much in
need of a good chemical analysis to determine just exactly what it has in it,
what might be missing to cause our unexplained embryonic mortality. It may
be due to this factor. We have tried Vionate. I think Vionate is an excellent diet
supplement. We got good hatchability with Kestrels using Vionate. The shells
of the egg appeared to be in somewhat better condition than with our regular
Kestrel diet; this is just from cursory observation. However, after hatching we
have had problems. I shall leave this to the final discussion on Health and Nu-
trition of the Young.
K180 RAPTOR RESEARCH Vol. 6, Supp. K
OTHER FOOD FOR CAPTIVE RAPTORS
McINTYRE. I suppose you all know that I’m from the Air Force Academy.
When I took over this job they were feeding these birds nothing but frozen
beef hearts. Well, me being a neophyte in the falconry business, I couldn’t see
feeding these birds beef heart as a complete diet. Every once in a while, maybe
once a year, they’d go out and get some pigeons and feed these birds pigeons.
Well, a month after I took over, I threw out 400 pounds of frozen beef heart.
You know, I can do this, being a veterinarian; I declared them unfit for human
consumption and threw them out. At first I started using quail, Coturnix Quail.
Then the guy I was buying the Coturnix Quail from got outrageous in his price
and we started using our own chickens. We’d get day-old chickens and raise
them to three weeks. The feed we were putting into them, the only non-medi-
cated feed that you can put into a chicken, I think, is game bird feed. Actually
the medicated feed has about 3% Auromyacin in it and I cannot see that this is
going to hurt any bird of prey. I also used a water soluble multi-vitamin, it’s
called Head Start, and put this in the water of the chicks. This way I don’t have
to worry about giving my hawks multivitamins. During the winter time, since I
have access to the hospital at the Academy, I get a bottle of multivitamins and
shove a tablet down the birds every other day. Of course, I noted a great im-
provement in the condition of the birds and the sheen of the feathers and so
forth, so I thought that we were on the right gimmick, and we’ve had some birds
now that we’ve had for six, seven, eight years that have been on this chicken
diet almost exclusively. Occasionally we feed them some venison that we get off
a road kill at the Academy. Of the breeders that we have, a pair of anatums, the
tiercel we had was in lousy shape when we got him. He was supposed to have
been five years of age. His feet and cere were completely white; he couldn’t
jump off the ground to a block perch. Two weeks after we had him on a strictly
chicken diet, and we fed the whole chicken, just took nothing off. We have
learned now to cut the feet off because the Peregrines particularly won’t eat the
shank of the legs and this saves the cadets from going around picking them up.
Now we got eight eggs out of the Peregrine last year in two different clutches.
The tiercel did not have much to do with it, but I think it was because he was
in such lousy shape when we got him. I am looking for better signs this year,
hopefully, than we had last year, because he is in real good shape. Now we fly
him on a string, and we jump him straight up, and he’s in pretty good shape.
And he is real tame, you can put him in your coat pocket, carry him any place
you want to. Sometimes we take him to football games and we never tie him
up in a hotel room. Of course this shakes a lot of people up— they walk in there
and there is a Peregrine sitting on the back of a chair. I think, I hope this is an
adequate diet. After looking at these figures Bill gave, I am not too sure. In the
breeding pen of course we keep a bath of water which was a wading pool; we
try to keep it full of water all the time. Fortunately we’re lucky we have water
available; the big problem is that it freezes in the wintertime. So this is about all
I have on the nutrition of these birds.
 1

1 972 Breeding Conference-Health and Nutrition of Adults K 1 8

HUNTER. What would be the cost? Would the cost be prohibitive to do a

comprehensive assay on four week old chick? And wouldn’t it be fruitful to do
so?

HALLIWELL. I think it probably would be fruitful to do so. I don’t know

whether I am capable of doing it or not, but I will certainly check into it and

see.

TEMPLE. We have the nutrition people at Cornell doing an analysis of day

old chicks, horse meat, and the various things that we are feeding the birds.
They are doing one at a time. They started on the day old chicks and the four
week old chicks. This information should be available soon.

THACKER. Iwant to make a comment on thiamine deficiency in Pere-

just
grine Falcons. Bill was saying that Prescott Ward diagnosed it in a Peregrine Fal-
con last year which was in A VMA Journal. Near Ohio State there is a bird sanc-
tuary. It’s run by one of the Audubon groups and based on Prescott Ward’s
article all of the birds fed there are fed on chicks; this is cockerels up to 12 days
old. They get them from a local dealer, kill them and feed direct. And these
birds have been on these chicks now for at least a year and a quarter to a year
and a half. And in the last four months we have had two Great Horned Owls,
a Red-tail, a Red-shoulder, and a Prairie Falcon all exhibiting signs of thiamine
deficiency. That is one point I wanted to bring up about chicks.

SWARTZ. I have my own conviction about snowshoe hares— they’re poor

food. Would you comment on beef heart?

McINTYRE. Beef I think we’re all agreed, but maybe not,

heart of course,
basically is low and phosphorus. Of course, with beef heart you get
in calcium
no castings at all. I think you can supplement an adequate diet with this occa-
sionally, an adequate diet of pigeons, chickens, maybe rats. I’m sure that people
who fee4 pigeons regularly feel that this is the best diet. And I can’t argue with
them. The only argument I’ve got is that you always have to worry about tricho-
monas or frounce. And since people don’t want to disturb breeding pairs par-
ticularly you are taking a chance to lose a bird. Of course I am a great believer
in casting material; this is why we have gone strictly to the chicken. And it’s a
whole chicken, feathers and all, and they get good castings every day. I would
stay away from wild birds because I don’t know how much pesticide they may
have in them: Starlings and so forth. Dick Graham, I think, had some chickens
analyzed that he is raising. And three of us in the Colorado Springs area have
chickens from the same hatchery. I forget now, but I think he said the highest
part pesticides was in the liver, about .03 parts per million, that right?

GRAHAM. Yes, that was the highest of the whole business.

K182 RAPTOR RESEARCH Vol. 6, Supp. K
GALICZ. For have been experimenting by feeding strict-
the last few years I

ly rabbit heads. These are strictly fresh rabbit heads acquired fresh at the butch-
er’s three times a week and each pair of Peregrine receives three heads per day.
They appear healthy, they have extremely good egg production both last year
and particularly this year. One pair had as many as three clutches and incubated
all three, but there is a problem of fertility. Now what is the answer, this is my

question.

HALLIWELL. won’t try to pick out specific factors, but I certainly think
I

you whole animal diet. One thing you are missing and you
are not feeding a
haven’t seen it in these birds yet, maybe they are picking it up someplace else.
Dr. Prescott Ward at Edgewood Arsenal in Maryland a year ago diagnosed a case
of thiamine deficiency in a Peregrine Falcon, most of you are familiar with the
work. So far as we can tell, the only place that thiamine is found in the natural
diet of birds of prey is in the intestinal tract, in the ingesta of the intestinal
tract, so I would say you are missing this. 1 frankly think you are missing quite
a bit.

GALICZ. They are producing eggs, and they are beautiful eggs; the birds
appear healthy.

HALLIWELL. These eggs, have you checked them? Have any of them been
fertilized?

GALICZ. Last year there was a possibility that one hatched but it isnot con-
clusive. The chick disappeared, that was the only case but it is not conclusive.
Itappeared to be there, but four days later when I actually checked back the
other three eggs were there and they were rotten, they were not fertile.

HALLIWELL. My comment to you would be to switch diets and I don’t

know where you’re going to switch to.

GALICZ. My reason for feeding the rabbit heads was because I was assured
they were reasonably free from various pesticides; these people were feeding
very carefully. The only other thing I did use was Vitamin E at one period after
they finished setting and 14 days after nesting they immediately laid again.
Now whether it is the Vitamin E that prompted the laying so quickly after in-
cubating, I don’t know.

MENG. I feed my birds pigeons; I take the two breasts off and then give them
the backs and the intestines and all the gonads of mature pigeons. They seem to
get the hormones from the gonads they feed on. So if you could do that in addi-
tion to the rabbits perhaps that could be the answer.

GALICZ. I’ll give it a try next season.

1972 Breeding Conference— Health and Nutrition of Adults K183

HALLIWELL. Why not just combine, feed two or three varied diets.

NELSON. have a possible answer to George Galicz’s question, and Dr. Meng
I

might have some suggestions or ideas here. In birds in the wild the female for
several weeks prior to egg laying doesn’t get any heads. The male eats the heads,
brings the rest to the female in courtship feeding activities. The female, while
she is laying and right through apparently until the chicks are a couple weeks
old, doesn’t get any heads or virtually none, although there may be some. Now
Dr. Meng might have some ideas here and others who have fed whole prey items.
Do the birds select certain parts from their kill? I think they do and certainly if
you could arrange it to feed them whole pigeons the birds are going to medicate
themselves. They’ve done this for millions of years.

SWARTZ. I would like to comment on the point of varied diet and also to
parasites; I’ve been playing a trick lately. Skip Walker is our stock feeder in
maintaining our mouse colony, when they get to a certain size I take over the
feed lot operation and feed them up on laboratory chow, then take them off
of the laboratory diet and put them on pure carrot. The skin then turns yellow
in just three days. This may serve to flush some of the bad things out of them
in this way. The Merlins for example get one of these mice every day; this does
produce a very lovely yellow fat, interesting yellow ceres. The feet are also
yellow; it’s been, so far, very satisfying and solves this intestinal problem, too.

HALLIWELL. I would like to have Mr. Kent Carnie talk about his birds that
have gone on a more adequate diet, and you can even see the difference in the
tail feathers.

CARNIE. I was intermewed passage European Goshawk. She had

flying an
been fed on a diet in essense of what she caught, jackrabbits, all last winter. She
took straight muscle plus heart, that was it. She seemed healthy, she flew ex-
tremely well, the diet was fine, her appetite was fine, but midsummer she came
down with what would have said was frounce ten years ago. However, there
I

were no trichomonads. She might have had spots in the lungs, it might have
been aspergillosis, but it seems very possibly that from this straight meat muscle
diet it might well have been the vitamin deficiency, something that Dr. Halli-
well published about in Hawk Chalk; it sounded like Vitamin A. Anyway we
tested for aspergillosis, we treated for frounce, we treated for vitamin deficien-
cy. At the same time with the vitamin deficiency, gave her one-a-day vitamin
I

then every other day, then twice a week. I changed her chow
tablets, first daily,
from straight jackrabbit muscle to an entire pigeon cut up in pieces which un-
fortunately at that time had to be forced down her throat. This bird went en-
tirely off her feed; she had to be force fed for nine days straight. She went
from 50 to 44 ounces and things looked bleak. Nine days straight, she finally
bottomed out at 44 and began to perk up. Up to that time incidentally her
feet were pale gray and very, very scaly and very dry in contrast to the yellow
K184 RAPTOR RESEARCH Vol. 6, Supp. K
waxy A1 Nye’s Gos for example. He had her on a diet I’m not familiar
feet of
with, I think young chicks. Incidentally I tried the chick route with this bird,
she just won’t have it. Prior to this disease she had shown a perfectly normal
molt except that she had only dropped her decks and she dropped nothing else
in the train. With the onset of the disease and the change in diet, she began to
snap out of it, she dropped her train. You can now look at this bird’s train and
see her decks in comparison with the brand new feathers. They are as different
as night and day. The feathers are drastically different in color, in quality, in
brittleness and this thing speaks to a natural diet to me. It is my fault she got
into this condition. It is because of Raptor Research Pathology Committee that
she is alive today. It surely shows by looking at this one bird’s train the visible
evidence of the different environment.

HALLIWELL. Most of the people raising birds of prey or keeping them in

captivity have been feeding a somewhat jury-rigged diet containing either chick-
en parts or chicken wings or something like that and trying to supplement the
diet with products, usually veterinary products such as Vionate and diet supple-
ments. And I wondered if you would discuss for a moment the rationale of try-
ing to supplement a basically inadequate diet by adding to it these supplements.
In other words what is your opinion in this direction? My personal approach is
to try to put into the diet to begin with everything you need, in other words to
try to provide in the diet by what you are feeding rather than supplement, let’s
say, horse meat with large amounts of Vitamin E, Vitamin A, minerals, and so
on down the line. Do you understand?

CARLSON. A
number of years ago we used to do what we call a free choice
feeding along with our poultry feeds and we put several things out there for the
birds to pick at. We found that most of them might do a fair job, but there were
always those few that were just too gluttonous in eating all grain or all mash or
all and wouldn’t really balance their diet and so we came to the con-
this or that
clusion that was necessary to put everything in one meal and I think that is
it

what you were getting at. And so I think where you have considerable invest-
ment that this is the way I would go. I would try to formulate a complete ration
and expect the animal to get that. Of course all of one thing— they may not like
it, it’s not very much variety, but we have found in commercial poultry produc-

tion that that is what gives us the return.

CAMPBELL. Would you care to comment on

egg yolk as a source of vitamin,
trace minerals, and supplementary food for raptors?

CARLSON. As I would have to say that egg yolk is

a poultry nutritionalist
the perfect food. does contain all the essentials at least for the growth of that
It

embryo throughout that period that it is in the shell and even for a few days
afterward. Generally there are pretty ‘fair quantities, so I would say that egg
yolk ought to be a good supplement.
1972 Breeding Conference-Health and Nutrition of Adults K185

HALLIWELL. Quite a few people have been feeding day old cockerels to
their adult raptorial birds. Would you comment on what you think the nutri-
tional value is of day old cockerels?

CARLSON. They have a good amount of egg yolk in them. Then again
still

you are getting about everything that was in the egg to begin with except the
loss of a little oxygen and maybe some other things that volatilize. It should be
a pretty fair diet.

HALLIWELL. Do you think that this would be a nutritionally staple diet

again considering carbohydrates, fats, proteins, minerals, vitamins, to maintain
the healthy bird?

CARLSON. It should be.

VOICE. Isn’t a four week old chick more nutritional source of food than a
day old chick?

CARLSON. I would say that a day old chick would be better than
a four
week one. The only reason I would say not quite as good
that the other bird is

is that we have diluted it perhaps with nutrients that your raptor is not going

to need. Zinc, for example. You don’t build up much zinc in that bird and yet
the day old chick probably contains a lot more in parts per million.

HALLIWELL. What would you think of the mineral contents of day old
chicks? What I’m really trying to say is that this chick had to utilize metabolic-
ally many of the nutrients that were in that egg the day it was laid in order to
create a baby chick at the time of hatching. What is the relationship of this to a
chicken that has grown to four or so odd weeks on an adequate diet?

CARLSON. Well, there is one thing that the day old chick would not have
and that is calcium and phosphorus. Even though the shell is almost 100% cal-
cium carbonate, not much of it gets into the chicken itself. The bones are prac-
tically all protein; there is very little calcified tissue there. Certainly the older
bird would be much better for the massive minerals, in other words calcium,
phosphorus and so on. Other than that I think you would be better off with
the day old chick.

SHERROD. It seems to me that since they naturally prey on older birds, it

would seem natural to use other than day old chicks.

CARLSON. This is true. Certainly the older bird is probably going to be a

pretty good diet. Maybe as good as the bird needs.

EBERLY. I read one report by a gentleman, I believe in Europe, who had

K186 RAPTOR RESEARCH Vol. 6, Supp. K

success breeding owls and he switched his diet over to cockerels, probably
around five weeks old, and he did not have success that year. He thought maybe
itwas because he had used medicated pre-starter for the chicks. What do you
think of that?

McINTYRE. I know, but I don’t see the logic behind this. The
really don’t
medication in medicated feed is so small, it’s 3% aureomyacin and you try
this
to correlate this and say well there probably wouldn’t be so many humans on
earth if this had something to do with fertility, because there are a lot of people
that get shot with antibiotics today and a lot of people take it orally without a
prescription or with a prescription and this is the only basis I know, I really
don’t know.

EBERLY. Is that all that’s in it, aureomyacin?

McINTYRE. That’s all that’s in it.

HALLIWELL. If you have something else to add on medicated feed we’ll

listen to you.

FYFE. I was under the understanding that some of the things that are going
into chicken starter and turkey starter include arsenic in very small quantities,
and I wonder if this would have any effect or should be of a concern as far as
the next step up in birds of prey which would concentrate something like this.

HALLIWELL, Yes, everything you have said is true. My

only answer is they
are dumping just as much in the rivers and on the and the prey species
fields
contain some also. Beyond this I am unwilling or unable to comment.

SMYLIE. What I would like to know is, in this starter feed are coccidiostats
Are there any hormones or anything else that could be carried through
in there?
to show changes in the raptors? Also what is the amount of DDT found in com-
mercial chicken, etc., that is available at the grocery store now?

CARLSON. Well, as far as the chick starter being carried over to the raptor,
I don’t believe there is anything there that you need to worry about as far as
hormones are concerned. We don’t add hormones at the present time. We do
have antibiotics and coccidiostat to cut down on coccidiosis. But at the level it
is used and by the time it goes through the bird I’m sure that this would not

interfere with the bird that you are concerned with. As far as the amount of
DDT and other chlorinated hydrocarbons in poultry, undoubtedly there is some
but Food and Drug is doing a pretty good job in terms of monitoring this and I
don’t think it’s going to hurt you as much to eat that bird as it hurt the bird in
the beginning. In other words we find that avian species are much more suscep-
tible to chlorinated hydrocarbons than people are. Of course that brings another
1972 Breeding Conference— Health and Nutrition of Adults K187

question— do we get enough in them to hurt your bird? That’s entirely another
subject. At the present time under conditions where we do have control over
our animals I don’t believe this is a problem. If, for example, you have your
aviary close to where they are burning some transformers, I would be a little
bit concerned about PCB’s and things.

GALICZ. I understand that stilbesterol isused in some cases to promote rapid

growth in poultry. Now if such poultry was fed to the hawks, would this cause
sterility?

CARLSON. If you fed nothing but livers I suspect after a few years this
might be a problem, but normally I don’t think so. We used to be able to inject
a pellet into the bird but this was prohibited about 1 5 years ago. Now the only
way we can get into a bird is through the diet or through the drinking water and
then in very minute quantities. I don’t think it would be a problem. Then you
might be concerned with feeding a lot of cow liver or something where you’re
getting a real potent supply.

TEMPLE. In this regard, too, of the steroid hormones including diethyl

all

stilbesterol in the chickens that you would be feeding to your hawk is fairly
rapidly broken down. The residues that would be in birds will probably be so
small, unless you were giving a massive dose I wouldn’t worry about it.

GALICZ. The reason I was asking that is, at one stage I happened to be en-

gaged commercial mink farming and some years back in Canada we did have
in
some serious problems with lack of production in mink directly due to feeding
chicken.

CARLSON. But you fed them the whole head and that head had a pellet in
the back of the neck so they got a real slug of it.