Egg Size, Fertility, Hatchability, and Chick Survivability in Captive California Condors (Gymnogyps Californianus)
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Zoo Biology 23:489–500 (2004) Egg Size, Fertility, Hatchability, and Chick Survivability in Captive California Condors (Gymnogyps californianus) Nancy C. Harvey,1n Jenine D. Dankovchik,1 Cynthia M. Kuehler,2 Tom Levites,3 Susie Kasielke,4 Lloyd Kiff,5 Michael P. Wallace,1 and Michael E. Mace3 1Center for Reproduction of Endangered Species, Zoological Society of San Diego, San Diego, California 2Keauhou Bird Conservation Center, Volcano, Hawaii 3San Diego Wild Animal Park, Escondido, California 4Los Angeles Zoo, Los Angeles, California 5The Peregrine Fund, Boise, Idaho California condors are one of the most endangered species native to the mainland United States and are subject of intense effort regarding captive breeding and reintroduction. We analyzed 20 years of California condor egg records from the wild and from three captive propagation facilities for fertility, hatchability, and chick survivability, along with changes in egg size due to multiple clutching. Overall annual mean percent of fertile eggs was 80.2%, hatchability was 87.3%, and chick survivability to Z60 days was 95.5%. One egg-laying site had a significantly lower fertility rate (P r 0.0001) than the other sites, which was probably due to pair incompatibility rather than any physiological factors. Egg volume of the first egg was significantly greater than both the second (t ¼ 6.73, P ¼ 0.0001) and third egg (t ¼ 6.62, P r 0.0001) of the season, while the second egg had a significantly greater volume (t ¼ 3.20, P ¼ 0.0084) than the third egg. Chicks from the second egg (t ¼ 3.24, P ¼ 0.029) and third egg (t ¼ 7.94, P ¼ 0.0014) of the season were significantly smaller than chicks from the first egg of the season. The decrease in egg measures and chick hatch weight due to multiple clutching did not affect hatchability or chick survivability. There were significant positive relationships (Po0.0001) between fresh egg weight and chick hatch weight and between egg volume and chick hatch weight, as well as between fresh egg weight and egg volume. In spite of the decrease in fresh egg weight, egg volume and chick hatch weights, due to egg removal to stimulate double and sometimes triple clutching, the captive propagation program has been nCorrespondence to: Nancy C. Harvey, Center for Reproduction of Endangered Species, Zoological Society of San Diego, P.O. Box 120551, San Diego, CA 92112. E-mail: [email protected] Received for publication July 1, 2003; Accepted October 26, 2003. DOI 10.1002/zoo.20015 Published online in Wiley InterScience (www.interscience.wiley.com). c 2004 Wiley-Liss, Inc. 490 Harvey et al. successful in producing birds for the restoration of this species. Zoo Biol 23: 489–500, 2004. c 2004 Wiley-Liss, Inc. Key words: endangered; Gymnogyps californianus; multiple clutching; egg volume INTRODUCTION Successful captive breeding programs for the restoration of critically endangered avian species, such as the Peregrine falcon (Falco peregrinus) [Cade et al., 1988], California condor (Gymnogyps californianus) [Kuehler and Witman, 1988], San Clemente loggerhead shrike (Lanius ludovicianus mearnsi) [Farabaugh et al., 2002], and a variety of Hawaiian forest birds [Kuehler et al., 2000, 2001] require high rates of viable fertile eggs, hatchability, and chick survivability for such programs to be successful. Although detailed records on egg weight, length, breadth, fertility, and/or viability, along with hatchability and chick survivability, are routinely maintained in captive propagation programs, few of the data from such records have been published. For seven endangered Hawaiian forest bird species, hatchability for viable eggs was 82.0%, and survivability to 30 days of age was 86.0% and (Kuehler, personal communication). Egg records from another captive passerine species, the San Clemente loggerhead shrike, revealed 82.7% fertility, 96.9% viability, 72.2% hatchability, and 54.4% chick survivability for eggs that were removed for artificial incubation and hand-rearing (Farabaugh, personal commu- nication). When shrike eggs were left with the parents for incubation and chick rearing, 31.3% of the eggs disappeared or were broken, so their fertility and viability could not be determined. Of the total eggs from captive parent-reared nests, known fertility was only 66.9%, hatchability was 81.4%, and chick survivability was 92.9% (Farabaugh, personal communication). Captive propagation programs for the restoration of endangered species often resort to double and sometimes triple clutching to rapidly increase the population. For captive Peregrine falcons [Burnham et al., 1984], this strategy resulted in a significant decline in mean egg weight and mean egg breadth from the first to the second, and again from second to the third clutches laid in the same year for captive Peregrine falcons [Burnham et al., 1984]. Similar data are unavailable for the San Clemente loggerhead shrike and the Hawaiian forest birds captive propagation programs. Field biologists recognized California condors as critically endangered by the early 1980s [Snyder and Snyder, 1989, 2000]. Starting in 1982, the Condor Recovery Team (CRT) agreed to transfer wild eggs and immature birds to the Zoological Society of San Diego’s (ZSSD) facility at the San Diego Zoo (SDZ) and Los Angeles Zoo (LAZ) to minimize extinction risks. Following additional condor losses in the wild, the U.S. Fish and Wildlife Service agreed with CRT recommendations and began removing the remaining nine birds from the wild and placing them in captive facilities [Snyder and Snyder, 1989]. The entire population of condors numbered 27 at the start of the captive propagation program in 1987, with 15 birds at the ZSSD’s facility at the Wild Animal Park (WAP) and 12 birds at the (LAZ). In 1995, The Peregrine Fund (TPF) became the third organization to participate in the captive- breeding program. California condors become sexually mature at around 6 years of age [Koford, 1953; Snyder and Snyder, 1989] and typically lay a single egg every 2 years. California Condor Egg Size and Fertility 491 California condors have the capacity to lay a replacement egg if the previous egg/ chick is lost or removed early in the season [Snyder and Hamber, 1985]. From 1987 to 1994, the condor staff routinely removed eggs from the captive pairs on the day of oviposition for artificial incubation and hand rearing, allowing the birds the possibility of laying a second clutch, and in some cases a third clutch, within a single season. Protocols for artificial incubation, egg weight loss, adult and chick diets, feeding schedules, and record keeping were established early in the program [Kuehler and Witman, 1988; Toone and Risser, 1988]. In 1995, the CRT recommended that birds hatch and rear their second egg of the season, therefore eliminating the possibility of them laying a third egg within one breeding season. Almost two decades of egg records from three captive propagation facilities dedicated to the recovery of the California condor provide a unique opportunity for analyzing information on fertility, hatchability, and chick survivability. MATERIALS AND METHODS Housing and Diet Each captive pair is housed in a large outdoor flight pen, with an adjacent roost area leading to a secluded nest box. At all three facilities, the birds are fed rabbits, rats, and fish. At the WAP and TPF, the diet also includes chicken. In addition, the WAP provides canine diet and beef spleen, while the LAZ provides feline diet and horsemeat. Water is provided ad libitum. In an effort to control weight gain and to mimic food availability in the wild, the birds at the WAP are fasted 2 to 3 nonconsecutive days per week, and at the LAZ they are fasted during the 2 weekend days plus 1 nonconsecutive day during the week. Source of Eggs and Sample Size A total of 46 females and 42 males in 53 different pairing combinations contributed 406 eggs to the database from 1983 to the end of the 2003 breeding season. Of the 406 eggs, 16 eggs came from the wild, 141 from the WAP facility, 131 from the LAZ facility, and 118 from TPF facility. Incubation From 1983 to 1985, modified Lyon incubators were used for incubation [Kuehler and Witman, 1988]. After 1985, Peterson Model 1 incubators were used at both the WAP and the LAZ. The Lyon incubators were used as hatchers at the WAP, while at the LAZ the Peterson Model 1 incubator tray was maintained at a level position for hatching. Finally, TPF used a Humidaire Model 21 for both incubation and hatching. The incubation parameters used for the first 15 eggs brought in from the wild [Kuehler and Witman, 1988] included a theoretical mean egg weight loss of 12.0% (fresh egg to pip). Out of 298 fertile eggs, 253 eggs hatched, and 238 chicks survived to Z60 days of age. Chicks were hand-reared (n ¼ 192), parent-reared (n ¼ 45), or cross-fostered (n ¼ 16) to either a different pair of California Condors or to a pair of Andean Condors (Vultur gryphus). 492 Harvey et al. Egg Measurements Upon removal from the nest, the aviculturalists weighed the eggs to the nearest 0.10 gram on Satorius (WAP), A&D Co. Ltd. (LAZ), and Denver (TPF) digital scales. Fresh egg weights were acquired on 239 eggs. There were records for 338 eggs for which both length and breadth measures were available to calculate egg volume. Aviculturalists measured the length and breadth of each egg to the nearest 0.10 mm. Length and breadth measures for the wild and captive eggs at the WAP were acquired with a Mitutoya digital caliper, while the LAZ and TPF facilities used SPI and Craftsman dial calipers, respectively. Since we had more length and breadth measures than fresh egg weights, we also calculated egg volume (V) using Hoyt’s 2 [1979] equation (V ¼ Kv  LB ) where the estimated volume coefficient (Kv ¼ 0.507) is applicable to all eggs which are not very pointed.