Research Interests

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Research Interests and Approach

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Parrotlets 

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Demographich Models & PVA

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Marbled Murrelets & Song Sparrows

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Snail Kite

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Conservation Biology

 

I.  Research Interests and Approach

 My research incorporates conservation biology, and behavioral and population ecology.  Most of my work has been with birds, although my students have studied aquatic invertebrates, plants, herps, and mammals.  This work has been conducted in terrestrial, wetland, and near shore marine ecosystems in the U.S. (California, Connecticut, Florida, Illinois, Ohio, and Michigan), the Caribbean (Cuba and Puerto Rico) and South America (Guyana and Venezuela). 

The interaction of animal ecology and natural resource management has been a common denominator of much of my work.  My conservation biology publications have emphasized the management and recovery of endangered species, the use of models for analyzing risks of extinction for threatened populations, and the potential for conservation gains and losses from sustainable harvesting and international trade.  In ecology, my work has centered on understanding why particular parental care strategies occur, patterns of demography and population dynamics, and how diet specialization is maintained. 

I approach research by combining intensive field studies based on natural history and quantitative sampling with field experiments and modeling, and expect the same of my graduate students.  A dominant theme in my research has been to determine the influence of environmental variation on behavior, to link behavior to population ecology, and to use this knowledge in conservation.  I have made contributions to both applied conservation biology and basic ecology because I chose systems to study that were amenable to both kinds of research, and I conducted studies that both tested theories and yielded data that was relevant to management or policy options.  This approach formed the basis for a review I published on how behavioral ecologists can make their research programs more useful for conservation (Beissinger 1997) and is illustrated below by my recent work on parrotlets. 

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II.  Parrotlets as Models for Understanding Hatching Asynchrony, Demography of Tropical Birds and Sustainable Harvesting  

(Forpus passerinus Chicks)

      My current studies of Green-rumped Parrotlets in Venezuela have important implications for both conservation biology and behavioral ecology, and have been supported by two grants from the National Science Foundation.  One award was to investigate the functional significance of the early onset of incubation behavior leading to hatching asynchrony in this small parrot, while the other was collaborative research to evaluate genetic and demographic approaches to estimate population viability.  The Green-rumped Parrotlet is an ideal species for studies of hatching asynchrony and demography because it lays unusually large clutches (x = 7 eggs), begins incubation on the first egg so chicks in a brood hatch extremely asynchronously (up to two weeks apart), nests in artificial boxes facilitating clutch and brood manipulations, and is easily captured for banding (Beissinger & Waltman 1991).

  Hatching asynchrony can be considered one of the long-standing puzzles in avian behavioral ecology.  Birds are unusual, compared to other vertebrates, in having some control over birthing patterns but often select a pattern that leads to mortality of last hatched young.  As we have pointed out in an extensive review paper, no less than 17 hypotheses have been proposed to explain why birds initiate incubation before a clutch has been completed (Stoleson & Beissinger 1995).  Most previous work concentrated on demonstrating the adaptive significance of the resulting hatching patterns (and the brood reduction it causes due to age differences among nestmates), while ignoring the behaviors and events that occur before incubation is initiated.  When birds hesitate for several days before initiating incubation, they leave their eggs unattended and susceptible to fluctuating temperatures and predators.  We measured an exhaustive set of fitness correlates at experimental synchronously and asynchronously hatching nests, and could find no evidence that asynchrony benefited hatching patterns in one of the most comprehensive analyses of the adaptive significance of brood reduction (Stoleson & Beissinger 1997a).  Using doubly labeled water, we also found no differences in the energy expenditures of parents that reared synchronous and asynchronous broods (Siegel et al. 1999). 

Finding little evidence that asynchrony directly benefited young or adults after hatching, we then conducted experiments to examine the consequences of not caring for eggs during laying.  Field experiments tested the ability of newly laid eggs to survive when exposed to ambient air temperatures (Stoleson & Beissinger 1999).  Parrotlet eggs were exposed for only several days to warm air temperatures before their hatchability started to decline rapidly.  A second experiment examined the consequences of not guarding nests (Beissinger et al. 1998).  Unguarded parrotlet nests had a high rate of egg destruction by nonbreeding male-female and male-male parrotlet pairs due in part to limited breeding opportunities (Beissinger 1996). 

These exciting findings have suggested a new avenue for understanding the significance of hatching asynchrony — the idea that unincubated bird eggs have a shelf life that may vary with environmental conditions.  It was an important theme at a symposium I co-organized at the International Ornithological Congress in South Africa in 1998.  I am now pursing this idea on two fronts.  First, I have begun to use models to examine how the onset of incubation affects the potential for interactions among egg viability, predation and brood reduction (Beissinger 1999).  Second, I designed a set of critical experiments to determine if and how temperature constrains viability of unincubated eggs using a climate gradient along the slope of a mountain in Puerto Rico with another bird, the Pearly-eyed Thrasher.  This project was funded by NSF, and work was begun in February 2000 in collaboration with a postdoctoral student, Dr. Mark Cook. The thrasher project incorporates studies of the temperature and humidity effects with studies of microbiology to determine which factors critically affect the shelf life of eggs.

Our parrotlet work in Venezuela is providing the first detailed study of a New World parrot, having banded over 5,000 birds and tracked over 1500 nesting attempts during the past 14 years (Waltman & Beissinger 1992; Curlee & Beissinger 1995; Sandercock et al. 2000).  It is also one of the longest studies of a South American bird, now in its 15th year, and provides important comparisons with well-studied temperate counterparts.  Dr. Brett Sandercock, a former postdoc of mine, and I are in the midst of analyses of demography and population dynamics of parrotlets.  We used mark-recapture statistical models to estimate survival for 1334 adults that were color-banded and released during the first decade of study (Sandercock et al. 2000).  Both sexes had low annual local survival rates, similar to temperate birds.  However, this conclusion obscured substantial heterogeneity in survival caused by a large number of nonbreeding males.  Using dynamic multi-state models, we found that survival was higher for breeders than nonbreeders, and higher than similar temperate species.  Estimates were not biased by dispersal because adults showed strong site-fidelity.  This work has important implications for understanding latitudinal changes in survival and life history of birds.    

The next step in my research program is to integrate behavior into a holistic understanding of parrotlet mating and social systems, and population dynamics.  I am pursing this through: (1) collaborative studies of the genetic mating system including microsatellite DNA analyses of parentage with Dr. Colin Hughes at the Universit6y of Miami and his former doctoral student, Dr. Rebecca Melland (Hughes et al. 1998); (2) detailed behavioral studies of the social system of nonbreeding parrotlets being pursued by one of my current Ph.D. students, Eric Punkay; (3) continued long-term analyses of behavioral ecology conducted by my current postdoctoral student, Dr. Amber Budden,;and (4) analyses of the temporal and spatial dynamics of parrotlet populations using matrix population models that incorporate the mating and social structure of the population. 

Several applications for conservation have grown directly from our parrotlet studies.  First, parrots are one of most threatened groups of birds, due to harvesting for the pet trade and habitat destruction.  Biological and conservation information was so lacking for parrots that I co-organized a symposium on this topic for the American Ornithologists' Union.  The resulting book (Beissinger & Snyder 1992) is not only an important reference for parrots, but serves as a benchmark evaluation for a variety of techniques used in conservation biology: reintroduction, captive breeding, ecotourism, sustainable harvesting, and education.  Second, nestbox schemes patterned after our work in Venezuela may offer a way to both conserve habitats and manage parrots in a sustainable manner.  My project in Venezuela developed a conservative model for sustainably harvesting parrots (Beissinger & Bucher 1992, Stoleson & Beissinger 1997b).  A portion of young could be harvested without impacting population size by providing nest boxes, since many parrots are limited by nest site availability.  In addition, young can be sustainably harvested by taking last-hatched chicks that typically die due to sibling competition caused by hatching asynchrony.  We have also evaluated the accuracy and precision of methods for monitoring parrot populations in one of the few bird studies that have compared survey techniques with a reference population (Casagrande & Beissinger 1997).  Our studies have provided the basis for sustainable harvesting provisions in the Wild Bird Conservation Act of 1992, for which I testified twice before the U.S. Congress (Beissinger 1992).  The development of biological principles to guide the international trade, as part of the work of a subcommittee of the American Ornithologists’ Union that I chaired (Beissinger et al. 1991), helped to strengthen the act.   I summarize this work by addressing potentials, principles and practices of sustainable use of birds at a symposium of the London Zoological Society (Beissinger 2001).

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III.  Demographic Models of Population Viability and Management of Endangered Species

Population viability analysis (PVA) uses models to estimate risks of extinction and has become an important tool in conservation decisions.  Yet, PVA is relatively untested and is still far from reaching its potential.  In a widely read paper (Beissinger and Westphal 1998), I summarized various types of PVA models and examined the strengths, weaknesses and optimal uses of PVA.  The paper catalyzed a highly successful international conference I co-organized with Dr. Dale McCullough, Leopold Chair in Wildlife Ecology at U.C. Berkeley, in San Diego, California in March 1999 that was attended by 350 participants (Mann and Plummer 1999).  We edited a book from the conference that was published by University of Chicago Press (Beissinger and McCullough 2002).

I have tried creatively to use demographic models and PVA to make management decisions for the recovery of endangered species.  I formalized an approach to modeling extinction in periodic environments and applied it to water management problems affecting the Snail Kite in the Everglades (Beissinger 1995a).  To model the likelihood of extinction, I needed a stochastic population viability model that did not project demographic rates randomly, because Everglades water levels have long-term periodicity of droughts and floods (Beissinger 1986).  I developed a method using the cyclic occurrence of environmental states that may also be applied to other systems.  This is one of the few PVAs that calibrates population projections against census data.

I have also developed demographic models to evaluate management strategies for endangered birds, orchids and sea turtles.  I conducted demographic analyses and developed a simple model while serving on a National Research Council Committee to develop a plan to recover the highly endangered Mariana Crow on the island of Guam (NRC 1996).  I also evaluated the demographic conditions when relocating loggerhead sea turtle nests from beaches to protected sites is a useful strategy (Grand & Beissinger 1997).  Recent work (Meretsky et al. 2000) examined levels of mortality that are permissible for the California Condor reintroduction effort.  One of my former graduate students, Rich Shefferson, adapted mark-recapture models to estimate dormancy in a threatened orchid (Shefferson et al. 2001).  While serving on the Marbled Murrelet Recovery Team, I developed a model using the ratio of juveniles to after-hatch-year birds and survivorship estimated from allometric relationships (Beissinger 1995b, USFWS 1995) to estimate rates of population decline for Marbled Murrelets in the Pacific Northwest.  As a result of my experiences, I teach the small population biology portion of a training course in scientific principles of endangered species management to USFWS biologists.

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IV.  Field Studies of the Marbled Murrelets and Song Sparrows in California  

(Marbled Murrelet nest)

  My graduate students and I have been conducting field studies of an endangered seabird, the Marbled Murrelet, in central California.  This bird is extremely difficult to study because its nests are hard to find in old growth forests 35-75 m above the ground and individuals are difficult to catch and band while spending most of their life at sea.  Until recently, very little was known about this bird except that it was threatened by the cutting of old growth forests and oil spills and had declined in numbers throughout the Pacific Northwest

The first goal of our research was to develop robust methods to estimate murrelet population trends.  We adapted line-transect sampling to avian surveys at sea (Becker et al. 1997).  This resulted in a new sampling method for murrelets, spawned a workshop that I ran for the U.S. Fish and Wildlife Service to train at-sea researchers in the techniques, and yielded a research contract to test various sampling designs for a regional monitoring program in the Pacific Northwest (Beissinger et al. 1999).  Ben Becker, a recent Ph.D. graduate from my lab and now a research scientist with the National Park Service,  developed a detailed analysis of the habitat use and factors affecting the distribution of murrelets at sea as part of his dissertation.  He also analyzed the stable isotope contents of murrelet feathers in order to determine the diet composition of these birds.

The second objective of our work is to determine the causes of the murrelet population decline.  Zach Peery, a doctoral student in my lab, and I began a mark-recapture study and a radio-telemetry study in 1999 that should yield demographic and behavioral clues to determine whether murrelets are declining because of high rates of nest failure or because many individuals fail to nest.  We radio-tagged a total of 48 murrelets in April and May 2000 and 2001, and  followed their movements and survival closely. Telemetry has allowed us to find nests and to estimate the proportion of birds nesting and their nesting success.

The Song Sparrows of San Francisco Bay offers a unique glimpse of evolutionary biology in a threatened ecosystem close to Berkeley.  Three subspecies of Song Sparrow are found in restricted, nonoverlapping portions of the bay wetlands at densities an order of magnitude greater than neighboring upland subspecies.  Letitia Grenier is a doctoral student in my lab who is studying the behavior and ecology of these birds in order to understand the factors that promote the causes and consequences of such high densities.  Her work includes detailed studies of the territorial and social system, DNA analyses of parentage, and analyses of food habits using stable isotopes.  Dr. Cully Nordby, a postdoctoral student in my lab who is a Smith Postdoctoral Conservation Fellow sponsored by the Nature Conservancy, is working on the effects of an introduced plant on the nesting biology and competitive interactions of these sparrows and marsh wrens.

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V.  Mate Desertion, Diet Specialization and Conservation of the Endangered Snail Kite  

(Snail Kite - Rostrhamus sociabilis)

      My dissertation and postdoctoral field studies of the Snail Kite in Florida and South America from 1977-1989 yielded data crucial to the conservation of this endangered hawk, including the discovery of key biological processes relevant to its persistence in the Everglades ecosystem.  Four papers (Beissinger 1986, Snyder al. 1989, Takekawa & Beissinger 1989, Beissinger 1995a) offer valuable data on kite demography, and show the effects of Everglades water management through detailed field studies and analysis of long-term data on water level and demography.  I also examined the problems of protecting small, temporarily-used wetlands under the Endangered Species Act, which are critical to the kite's survival during drought and have been rapidly developed.

While conducting the above work, I also studied how differences in reproductive investments and demography between the sexes affected the mating system and parental investments of these hawks.  I discovered with Dr. Noel Snyder that the Snail Kite has an extremely unusual mating system: if snails are abundant, either parent may desert its mate about half-way through the period of parental care and may find a new mate to begin another nesting attempt.  My early work in Florida elucidated the proximate characteristics of this mating system, and is often cited as one of the first detailed studies of mate desertion by either sex in birds (Beissinger & Snyder 1987).  It also provided strong tests of parental investment theory by relating expenditure of reproductive investments to the sex of the deserting parent at kite nests (Beissinger 1987a,b).  Later experiments in Venezuela tested a heuristic model that related both brood size and food demand to a monoparental threshold for desertion in one of the few papers to manipulate the occurrence of mate desertion (Beissinger 1990a).  In an important paper (Beissinger 1986), I modeled environmental predictability (Beissinger & Gibbs 1993), showed how ecosystem management affected both short- and long-term environmental cycles, and illustrated how environmental unpredictability in the form of fluctuating water levels could select for mate desertion.  To understand mate desertion from a comparative perspective, I conducted the only study in the past 40 years on the biology of the Slender-billed Kite (Beissinger et al. 1988), a little known South American congener of the Snail Kite that also feeds primarily on snails.  Interestingly, unlike the colonial, marsh-nesting Snail Kite, the territorial Slender-billed Kite nests solitarily in flooded forests and did not appear to practice mate desertion.

Vertebrates that eat few kinds of foods are extremely rare.  My earliest studies of the Snail Kite in Guyana were the first to quantify hunting behavior, snail choice and energetics of this highly specialized forager (Beissinger 1983).  Later I documented the ecological circumstances and consequences for kites of taking nonsnail prey - mainly small turtles in Florida and crabs in Venezuela (Beissinger 1990b).  Behavioral experiments conducted with free-flying birds in Venezuela showed that kites preferred snails to crabs, even when snails were much less profitable (Beissinger et al. 1994).  This work suggested a role for behavioral conservatism, in the form of risk-averse foraging and neophobia, in maintaining diet specialization. 

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VI.  Contributions to the Development of Conservation Biology

A final dimension of my work has been to help the newly emerging science of conservation biology develop as a discipline that can offer “tools” to assist in the protection, restoration and sustainable use of biological diversity.  In a widely read paper that has had a large influence on endangered species recovery strategies, a diverse group of colleagues and I analyzed the use of captive breeding as a conservation tool (Snyder et al. 1996, 1997a,b).  I also developed an approach to set conservation priorities among ecosystems that uses differences in the distributions of threatened and nonthreatened species as indicators of ecosystem integrity, and the distribution of parks and reserves and nonprotected lands as indicators of ecosystem protection (Beissinger et al. 1996).  We applied it to an analysis of the biomes and avifauna of South America, showing how the most diverse biomes (tropical humid forests and montane) may not be as threatened as less diverse but poorly protected biomes (tropical dry forests and subtropical rainforests).  Finally, I have worked with a group of economists to examine why economics matters for endangered species protection (Shogren et al. 1999), and with a committee of ornithologists to evaluate a widely-used protocol for prioritizing species for conservation (Beissinger et al. 2000).

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Literature Cited

Becker, B. H., S. R. Beissinger, and H. Carter. 1997.  At-sea density monitoring of Marbled Murrelets in central California: methodological considerations.  Condor 99:743-755.

Beissinger, S. R.  1983.  Hunting behavior, prey selection and energetics of Snail Kites in Guyana: consumer choice by a specialist.  Auk 100:84-92.

Beissinger, S. R.  1986.  Demography, environmental uncertainty, and the evolution of mate desertion in the Snail Kite.  Ecology 68:1445-1459.

Beissinger, S. R.  1987a.  Mate desertion and reproductive effort in the Snail Kite.  Animal Behaviour 35:1504-1519.

Beissinger, S. R.  1987b.  Anisogamy overcome: female strategies in Snail Kites.  American Naturalist 129:486-500.

Beissinger, S. R.  1990a. Experimental brood manipulations and the monoparental threshold in Snail Kites.  American Naturalist 136:20-38.

Beissinger, S. R.  1990b.  Alternative foods of a diet specialist, the Snail Kite.  Auk 107:327-333.

Beissinger, S. R.  1992.  Statement before the joint hearing of the U.S. House of Representatives Subcommittee on fisheries and Wildlife Conservation and the Environment of the Committee on Merchant Marine and Fisheries, and the Subcommittee on Trade of the Committee on Ways and Means.  Congressional Record Serial 102-84, Pp. 16-18 and 110-119.

Beissinger, S. R.  1995a.  Modeling extinction in periodic environments: Everglades water levels and Snail Kite population viability.  Ecological Applications 5:618-631.

Beissinger, S. R.  1995b.  Population trends of the Marbled Murrelet projected from demographic analyses.  Pages 385-393 in Ecology and Conservation of the Marbled Murrelet (C. J. Ralph, G. L. Hunt, Jr., M. G. Raphael, and J. F. Piatt, Eds.).  General Technical Report PSW-GTR-152, Pacific Southwest Research Station, U.S.D.A. Forest Service, Albany, CA.

Beissinger, S. R.  1996.  On the limited breeding opportunities hypothesis for avian clutch size.  American Naturalist 147:655-658.

Beissinger, S. R.  1997.  Integrating behavior into conservation biology: potentials and limitations.  Pages 23-47 in Behavioral Approaches to Conservation in the Wild (J. R. Clemmons and R. Buchholtz, Eds.).  Cambridge University Press, Cambridge, U.K. 

Beissinger, S. R.  1999.  Interaction of egg viability, brood reduction and nest failure on the onset of incubation.  Pages 638-646 in Proceedings of the 22nd International Ornithological Congress (N.J. Adams, & R.H.Slotow, Eds.).  BirdLife South Africa, Durban: Johannesburg, South Africa.  

Beissinger, S. R.  In press.  Trade in Live Wild Birds: Potentials, Principles and Practices of Sustainable Use.  In Conservation of Exploited Species (J. D. Reynolds, G. M. Mace, K. H. Redford, and J. G. Robinson, Eds.).  Cambridge University Press. 

Beissinger, S. R. and E. H. Bucher.  1992.  Can parrots be conserved through sustainable harvesting?  BioScience 42:164-173. 

Beissinger, S. R. and J. P. Gibbs.  1993.  Are variable environments stochastic? A review of methods to quantify environmental predictability.  Pages 132-146 in Adaptation in Stochastic Environments (J. Yoshimura and C. W. Clark, Eds.).  Lecture Notes on Biomathematics Springer-Verlag, Berlin.

Beissinger, S. R. and D. R. McCullough (Editors).  2002.  Population Viability Analysis.  University of Chicago Press, Chicago, IL. 

Beissinger, S. R. and N. F. R. Snyder.  1987.  Mate desertion in the Snail Kite.  Animal Behaviour 35:477-487. 

Beissinger, S. R. and N. F. R. Snyder (Editors).  1992.  New World Parrots in Crisis: Solutions from Conservation Biology.  Smithsonian Institution Press, Washington, D.C. 

Beissinger, S. R. and J. R. Waltman.  1991.  Extraordinary clutch size and hatching asynchrony of a neotropical parrot.  Auk 108:863-871.

Beissinger, S. R., and M. I. Westphal.  1998.  On the use of demographic models of population viability analysis in endangered species management.  Journal of Wildlife Management 62:821-841.

Beissinger, S. R., B. T. Thomas, and S. D. Strahl.  1988.  Vocalizations, food habits, and nesting biology of the Slender-billed Kite with comparisons to the Snail Kite.  Wilson Bulletin 100:604-616.

Beissinger, S. R., N. F. R. Snyder, S. R. Derrickson, F. C. James, and S. M. Lanyon.  1991.  International trade in live exotic birds creates a vast movement that must be halted.  Auk 108:982-984. 

Beissinger, S. R., T. J. Donnay, and R. Walton.  1994.  Experimental analysis of diet specialization in the Snail Kite: the role of behavioral conservatism.  Oecologia 100:54-65.

Beissinger, S. R., E. C. Steadman, T. Wohlgenant, G. Blate, and S. Zack.  1996.  Null models for assessing ecosystem conservation priorities: threatened birds as titers of threatened ecosystems.  Conservation Biology 10:1343-1352.

Beissinger, S. R., S. Tygielski, and B. Elderd.  1998.  Social constraints on the onset of incubation in a neotropical parrot: a nest box addition experiment.  Animal Behaviour 55:21-32.

Beissinger, S. R., B. H. Becker, L. Rachowicz and A. Hubbard.  1999.  Testing and designing methods for developing an at-sea monitoring strategy for the Marbled Murrelet.  Unpublished Report to the U.S. Fish and Wildlife Service.  99 pages.

Beissinger, S. R., J. M. Reed, J. M. Wunderle, Jr., S. K. Robinson, and D. M. Finch.  2000.  Report of the AOU Conservation Committee on the Partners in Flight species prioritization plan.  Auk 117: 549-561.

Casagrande, D. G., and S. R. Beissinger.  1997.  Evaluation of four methods for estimating parrot population size.  Condor 99:445-457.

Curlee, A. P., and S. R. Beissinger.  1995.  Experimental analysis of mass change in female green-rumped parrotlets (Forpus passerinus): the role of male cooperation.  Behavioral Ecology 6:192-198.

Grand, J. and S. R. Beissinger.  1997.  When relocation of loggerhead sea turtle (Caretta caretta) nests becomes a useful strategy.  Journal of Herpetology 31:428-434.

Hughes, C. R., R. R. Melland, and S. R. Beissinger.  1998.  Polymorphic trinucleotide microsattelite loci for a neotropical parrot, the green-rumped parrotlet, Forpus passerinus.  Molecular Ecology 7:1247-1248.

Mann, C. C. and M. L. Plummer.  1999.  A species’ fate by the numbers.  Science 284:36-37.

Meretsky, V. J., N. F. R. Snyder, S. R. Beissinger, D. A. Clendenen, and J. W. Wiley.  In press.  Demography of the California Condor: implications for restablishment.  Conservation Biology.

National Research Council (NRC).  1996.  The Scientific Basis for the Preservation of the Mariana Crow.  National Academy of Sciences, Washington D.C.  91 pp. 

Sandercock, B. K., S. R. Beissinger, S. H. Stoleson, R. R. Melland, and C. R. Hughes.  2000.  Survival rates of a Neotropical parrot: implications for latitudinal comparisons of avian demography.  Ecology 81:1351-1370. 

Shefferson, R. P., B. K. Sandercock, J. Proper, and S. R. Beissinger.  2001.  Estimating dormancy and survival of a rare herbaceous perennial using mark-recapture models.  Ecology 82:145-156. 

Shogren, J. F., J. Tschirart, T. Anderson, A. W. Ando, S. R. Beissinger, D. Brookshire, G. M. Brown, Jr., D. Coursey, R. Innes, S. M. Meyer, and S. Polasky.  1999.  Why economics matters for endangered species protection and the ESA.  Conservation Biology.13:1257-1261.

Siegel, R. B., W. W. Weathers, and S. R. Beissinger.  1999.  Hatching asynchrony reduces the duration, not the magnitude, of peak load in breeding green-rumped parrotlets (Forpus passerinus).  Behavioral Ecology and Sociobiology 45:444-450.

Snyder, N. F. R., S. R. Beissinger, and R. Chandler.  1989.  Reproduction and demography of the Florida Everglade (Snail) Kite.  Condor 91:300-316.

Snyder, N. F. R., S. R. Derrickson, S. R. Beissinger, J. W. Wiley, T. B. Smith, W. D. Toone and B. Miller.  1996.  Limitations of captive breeding in endangered species recovery.  Conservation Biology 10:338-348.

Snyder, N. F. R., S. R. Derrickson, S. R. Beissinger, J. W. Wiley, T. B. Smith, W. D. Toone and B. Miller.  1997a.  Captive breeding and conservation (reply to Hutchins, Wiese and Willis).  Conservation Biology 11:3-5.

Snyder, N. F. R., S. R. Derrickson, S. R. Beissinger, J. W. Wiley, T. B. Smith, W. D. Toone and B. Miller.  1997b.  Limitations of captive breeding: reply to Gippoliti and Carpaneto.  Conservation Biology 11:808-810.

Stoleson, S. H. and S. R. Beissinger.  1995.  Hatching asynchrony and the onset of incubation in birds revisited: when is the critical period?  Current Ornithology 12:191-270.

Stoleson, S. H., and S. R. Beissinger.  1997a.  Hatching asynchrony, brood reduction, and food limitation in a neotropical parrot.  Ecological Monographs 67:131-154.

Stoleson, S. H., and S. R. Beissinger.  1997b.  Hatching asynchrony in parrots: boon or bane for conservation.  Pages 157-180 in Behavioral Approaches to Conservation in the Wild (J. R. Clemmons and R. Buchholtz, Eds.).  Cambridge University Press, Cambridge, U.K. 

Stoleson, S. H., and S. R. Beissinger.  1999.  Egg viability as a constraint on hatching synchrony at high ambient temperatures.  Journal of Animal Ecology 68:951-962.

Takekawa, J. E., and S. R. Beissinger.  1989.  Dispersal, cyclic drought, and the conservation of the Snail Kite in Florida: lessons in critical habitat.  Conservation Biology 3:302-311. 

U. S. Fish and Wildlife Service (USFWS).  1995.  Draft Recovery Plan Marbled Murrelet (Brachyramphus marmoratus) (Washington, Oregon and California population).  U.S. Fish and Wildlife Service, Portland, Oregon.  171 pages.

Waltman, J. R. and S. R. Beissinger.  1992.  The breeding biology of the Green-rumped Parrotlet.  Wilson Bulletin 104:65-84.

 

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