Dr. Kushlan’s research and conservation programs on waterbirds, carried out over the past fifty years, have substantially expanded the knowledge base and conservation approaches to waterbirds, especially colonial waterbirds.
Studies were carried out both in the Everglades and at study sites in the tropics.
His waterbird research programs have engaged diverse questions of distribution, behavior, and conservation. He has particularly focused on the foraging and reproductive biology and conservation of storks, ibises, and herons. Beyond specific research and conservation results derived from the studies, he has engaged more widely in conservation issues especially playing a leading role in institutionalizing waterbird conservation nationally and internationally.
In the Everglades, the primary waterbirds studied were four species of herons, the white ibis and the wood stork. A common question of these studies involved the responses of these species to seasonal hydrological fluctuations.
Studies also included determining waterbird historical and current status, monitoring protocols, indicator roles, foraging behavior and prey selectivity, functional morphology, individual and population energetics, nestling growth and survival, population biology, and seasonal landscape use.
In addition to research papers, several syntheses brought together available information on the biology, behavior and conservation of waterbirds including four monographic books on herons, ibises and storks.
In other studies, Dr. Kushlan addressed information gaps for lesser known species, including the Lava Heron in the Galapagos, Whistling Heron in Venezuela and Argentina, Capped Heron in Venezuela, Boat-billed Heron in Mexico, Bare-throated Tiger Heron in Panama, Agami Heron in Costa Rica and French Guiana, Little Egret in the West Indies, and Green Heron in Bermuda. His surveys of seabirds defined their status in the Bahamas and Panama. These studies and application of waterbird conservation protocols have had global implications, especially through Dr. Kushlan’s leadership within waterbird research and conservation organizations, and authorship of regional and global conservation plans
For access to the articles cited below, please go to the Articles page. For more information about cited books, please visit the Books page.
Kushlan, J.A. 1972. Aerial feeding in the Snowy Egret. Wilson Bulletin 84:199-200.
Kushlan, J.A. 1973. Promiscuous mating behavior in the White Ibis. Wilson Bulletin 85:331-332.
Kushlan, J.A. 1973. Bill-vibrating: a prey attracting behavior of the Snowy Egret, Leucophoyx thula. American Midland Naturalist 89:509-512.
Kushlan, J.A. 1973. Least Bittern nests colonially. Auk 90:685-686.
Kushlan, J.A. 1973. Black-crowned Night Heron diving for prey. Florida Field Naturalist 1:27-28.
Kushlan, J.A. 1973. White Ibis nesting in the Florida Everglades. Wilson Bulletin 85:230-231.
Kushlan, J.A. and J.L. Schortemeyer. 1974. Glossy Ibis nesting in southern Florida. Florida Field Naturalist 2:13-14.
Robertson, W.B. Jr. and J.A. Kushlan. 1974. The southern Florida avifauna. Pp 414-452. In: Environments of South Florida: Present and Past. (P.J. Gleason, ed.). Miami Geological Society, Coral Gables FL.
Kushlan, J.A. and M.S. Kushlan. 1975. Food of the White Ibis in southern Florida. Florida Field Naturalist 3:31-38.
Kushlan, J.A., J.C. Ogden, and A.L. Higer. 1975. Relation of water level and fish availability to Wood Stork reproduction in southern Everglades, Florida. U.S. Geological Survey Report 75-434, Tallahassee FL, USA.
Kushlan, J.A. 1976. Site selection for nesting colonies by the American white ibis Eudocimus albus in Florida. Ibis 118:590-593.
Kushlan, J.A. 1976. Wading bird predation in a seasonally fluctuating pond. Auk 93:464-476.
Kushlan, J.A. 1976. Feeding behavior of North American herons. Auk 93:86-94.
Kushlan, J.A. 1976. Feeding rhythm in nestling White Ibis. Wilson Bulletin 88:656-658.
Ogden, J.C., J.A. Kushlan, and J.T. Tilmant. 1976. Prey selectivity by the Wood Stork. Condor 78:324-330.
Kushlan, J.A. 1977. Growth energetics of the White Ibis. Condor 79:31-36.
Kushlan, J.A. 1977. Population energetics of the White Ibis. Auk 94:114-122.
Kushlan, J.A. and D.A. White. 1977. Nesting wading bird populations in southern Florida. Florida Scientist 40:65-72.
Kushlan, J.A. 1977. Significance of plumage colour in the formation of feeding aggregations of ciconiiforms. Ibis 119:361-364.
Kushlan, J.A. 1977. Foraging behavior of the White Ibis. Wilson Bulletin 89:342-345.
Kushlan, J.A. 1977. Sexual dimorphism in the White Ibis. Wilson Bulletin 89:92-98.
Kushlan, J.A. 1977. Differential growth of body parts in the White Ibis. Auk 94:164-167.
Kushlan, J.A. and W.B. Robertson, Jr. 1977. White Ibis nesting in the lower Florida Keys. Florida Field Naturalist 5:41-42.
Kushlan, J.A. 1978. Feeding ecology of wading birds. Pp. 249-296. In: Wading Birds (A. Sprunt IV, J.C. Ogden, and S. Winckler, eds.). National Audubon Society, New York NY, USA.
Ogden, J.C., J.A. Kushlan, and J.T. Tilmant. 1978. The food habits and nesting success of wood storks in Everglades National Park in 1974. U.S. National Park Service, Natural Resources Report 16, Washington, DC, USA.
Kushlan, J.A. and L.C. McEwan. 1978. Colonial birds in South Florida national parks, 1976-1977. Report T-545, US National Park Service, South Florida Research Center, Homestead, FL, USA.
Kushlan, J.A. 1978. Wading bird use of the east Everglades. Florida Field Naturalist 6:46-47.
Kushlan, J.A. 1978. Commensalism in the Little Blue Heron. Auk 95:677-681.
Kushlan, J.A. 1978. Nonrigorous foraging by robbing egrets. Ecology 59:649-653.
Kushlan, J.A. 1979. Arboreal foraging by Cattle Egrets. Wilson Bulletin 91:471-472.
Kushlan, J.A. 1979. Prey choice by tactile-foraging wading birds. Proceedings of the Colonial Waterbird Group 3:133-142.
Kushlan, J.A. 1979. White Ibis. Pp. 83-84. In: Rare and Endangered Biota of Florida. Vol. 2, Birds (H.W. Kale, ed). University Presses of Florida, Gainesville, FL, USA.
Kushlan, J.A. 1979. Effects of helicopter censuses on wading bird colonies. Journal of Wildlife Management 43:756-760.
Kushlan, J.A. 1979. Wading bird preservation in Everglades National Park. Pp. 483-487. In: Proceedings of the First Conference on Scientific Research in National Parks, 1976. U.S. Department of Interior, Washington, DC, USA.
Kushlan, J.A., M.C. Baumann, and L.C. McEwan. 1979. A bibliography of south Florida wading birds. South Florida Research Center Report T-514, National Park Service, Homestead, FL, USA.
Kushlan, J.A. 1979. Feeding ecology and prey selection in the White Ibis. Condor 81:376-389.
Kushlan, J.A. 1980. Prey choice by tactile-foraging wading birds. Proceedings of the Colonial Waterbird Group 3:133-142.
Ogden, J.C., J.A. Rodgers, R. Andrews, S.A. Nesbitt, M.C. Landin, and J.A. Kushlan. 1980. Current management of colonial waterbirds in the United States. Pp. 32-38. In: The Management of Colonial Waterbirds (J. Parnell and R.F. Soots, eds.). North Carolina Sea Grant Publication UNC-SG-80-06. North Carolina Sea Grant, Wilmington NC, USA.
Kushlan, J.A. 1981. Resource use strategies in wading birds. Wilson Bulletin 93:145-163.
Kushlan, J.A., J.A. Hancock, J. Pinowski, and B. Pinowska. 1982. Behavior of Whistling and Capped Herons in the seasonal savannas of Venezuela and Argentina. Condor 84:255-260.
Kushlan, J.A. 1983. Pair formation behavior in the Galapagos Lava Heron. Wilson Bulletin 95:118-121.
Kushlan, J.A. 1983. Special species and ecosystem preserves: colonial waterbirds in U.S. national parks. Environmental Management 7:201-207.
Hancock, J. and J.A. Kushlan. 1984. The Herons Handbook. Harper & Row, New York, NY, USA; Croom-Helm, London UK.
Kushlan, J.A., L.G. Morales, and P.C. Frohring. 1985. Foraging niche relations of wading birds in tropical wet savannas. Pp. 663-682. In: Neotropical Ornithology (P. Buckley et al., eds). AOU Ornithological Monograph 36.
Kushlan, J.A. and P.C. Frohring. 1986. History of the southern Florida Wood Stork population. Wilson Bulletin 92:368-386.
Kushlan, J.A. 1986. Colonies, surveys, and sites: the terminology of colonial waterbird research. Colonial Waterbirds 9:119-120.
Kushlan, J.A. 1986. Responses of wading birds to seasonally fluctuating water levels: strategies and their limits. Colonial Waterbirds 9:155-162.
Kushlan, J.A. 1986. The management of wetlands for aquatic birds. Colonial Waterbirds 9:246-248.
Kushlan, J.A. 1987. External threats and internal management: the hydrologic regulation of the Everglades. Environmental Management 11:109-119.
Parnell, J.A., D.G. Ainley, H. Blokpoel, B. Cain, T.W. Custer, J.L. Dusi, S. Kress, J.A. Kushlan, W.E. Southern, L.E. Stenzel, and B.C. Thompson. 1988. Colonial waterbird management in North America. Colonial Waterbirds 11:129-169.
Frohring, P.C., D.P. Voorhees, and J.A. Kushlan. 1988. History of wading bird populations in the Florida Everglades: a lesson in the use of historical information. Colonial Waterbirds 11:328-335.
Kushlan, J.A. 1988. Impact of water management on wildlife in the Florida Everglades. Pp. 231-243. In: Ecologia y Conservacion del Delta de los rios Usumacinta Y Grijalva (M. Chevez, ed.). Instituto Nacional de Investigacion sobre Recursos Bioticos (INIREB) – Divison Regional Tabasco, Tabasco, Mexico.
Kushlan, J.A. 1989. Wetlands and wildlife, the Everglades perspective. Pp.773-796. In: Freshwater Wetlands and Wildlife (R.R. Sharitz and J.W. Gibbons, eds.) Dept. of Energy Symposium Series No. 61. Oak Ridge, TN, USA.
Kushlan, J.A. 1989. Avian use of fluctuating wetlands. Pp.593-604. In: Freshwater Wetlands and Wildlife (R.R. Sharitz and J.W. Gibbons, eds.). Dept. of Energy Symposium Series No.61. Oak Ridge, TN, USA.
Hancock, J. and J.A. Kushlan. 1989. Guide des hérons du monde. Delachaux & Niestlé. S.A., Neuchâtel, Switzerland.
Coulter, M.C., K. Brouwer, A.L. Bryan, Jr., M.P. Kahl, C.E. King, J.A. Kushlan, C.S. Luthin, M.v. Wieringen, and D.C. Young. 1991. A Bibliography of Storks, Ibises, and Spoonbills. Savannah River Ecology Laboratory, Aiken, SC, USA.
Hancock, J.A., J.A. Kushlan, and M.P. Kahl. 1992. Storks, Ibises, and Spoonbills of the World. Academic Press, London, UK.
Kushlan, J.A. and K.L. Bildstein. 1992. White Ibis. Birds of North America No. 9 (A. Poole, P. Stettenheim, and F. Gill, eds.). American Ornithologists’ Union, Philadelphia, Pennsylvania, USA.
Kushlan, J.A. 1992. Population biology and conservation of colonial wading birds. Colonial Waterbirds 15:1-7.
Kushlan, J.A. 1993. Waterbirds as bioindicators of wetland change: Are they a valuable tool? Pp. 48-55. In: Waterfowl and Wetland Conservation in the 1990s – a global perspective. (M. Moser, R.C. Prentice, and J. Van Vessem, eds.), IWRB Special Publication No. 26. Slimbridge, UK.
Kushlan, J.A. and H. Hafner. 1993. Status and conservation of the herons of the world. Species 19:68.
Kushlan, J.A. 1993. Colonial waterbirds as bioindicators of environmental change. Colonial Waterbirds 16:223-251.
Davis, W.E., Jr., and J.A. Kushlan. 1994. Green Heron. Birds of North America No. 129, (A. Poole, and F. Gill, eds.). American Ornithologists’ Union, Philadelphia, Pennsylvania, USA.
Kushlan, J.A. 1997. The conservation of wading birds. Colonial Waterbirds 20:129-137.
Kushlan, J.A. and D.A. McCrimmon. 1997. Wading birds: advances in knowledge since 1976. Colonial Waterbirds 20:105.
Mikuska, T., J.A. Kushlan, and S. Hartley. 1998. Key areas for wintering North American herons. Colonial Waterbirds. 21:123-134.
Kushlan, J.A. and H. Hafner (eds.). 2000. Heron Conservation. Academic Press, London, UK.
Butler, R.W., J.A. Kushlan, and I. J. Davidson. 2000. Herons in North America, Central America and the West Indies. Pp. 151-175. In: Heron Conservation (J.A. Kushlan and H. Hafner, eds.). Academic Press, London, UK.
Kushlan, J.A. 2000. Heron feeding habitat conservation. Pp. 219-235. In: Heron Conservation (J.A. Kushlan and H. Hafner, eds.). Academic Press, London, UK.
Kushlan, J.A. 2000. Research and information needs for heron conservation. Pp. 331-342. In: Heron Conservation (J.A. Kushlan and H. Hafner, eds.). Academic Press, London, UK.
Hafner, H., R.V. Landsdown, J.A. Kushlan, R.W. Butler, T.W. Custer, I.J Davidson, R.M. Erwin, J.A. Hancock, A.M. Lyles, M. Maddock, L. Marion, F. Morales, T. Mundkur, C. Perennou, O. Pineau, D. Turner, P. Ulenaers, J.v. Vessem, and L.Young. 2000. Conservation of herons. Pp. 343-375. In: Heron Conservation (J.A. Kushlan and H. Hafner, eds.). Academic Press, London, UK.
Kushlan, J.A. and H. Hafner. 2000. Reflections on heron conservation. Pp. 377-379. In: Heron Conservation (J.A. Kushlan and H. Hafner, eds). Academic Press, London, UK.
Kushlan, J.A., et al. 2002. Waterbird Conservation for the Americas. The North America Waterbird Conservation Plan. Waterbird Conservation for the Americas, Washington DC, USA.
Kushlan, J.A. 2004. Ibises and spoonbills. Pp 112-117. In: New Encyclopedia of Birds (C. Perrins, ed.). Oxford University Press, Oxford, UK.
Kushlan, J.A. and J. Hancock. 2004. Herons and bitterns. Pp. 98-105. In: New Encyclopedia of Birds (C. Perrins, ed.). Oxford University Press, Oxford, UK.
Kushlan, J.A. 2004. Storks. Pp. 106-109. In: New Encyclopedia of Birds. (C. Perrins, ed.). Oxford University Press, Oxford, UK.
Reynaud, P.A. and J.A. Kushlan. 2004. Nesting of the Agami Heron. Waterbirds 27:308-311.
Kushlan, J.A. and J.A. Hancock. 2005. The Herons. Oxford University Press, Oxford UK.
Kushlan, J.A., et al. 2006. Conservación de las Aves Acuáticas para las Américas. Waterbird Conservation for the America and US Fish and Wildlife Service. Washington, DC, USA.
Kushlan, J.A. 2006. Integrating waterbird conservation: populations, habitats and landscapes. P. 700. In: Waterbirds around the World. (G.C. Boere, C.A. Galbraith, and D. A. Stroud, eds.). The Stationary Office Ltd., Edinburgh, UK.
Kushlan, J.A. 2007. Conserving Herons: A Conservation Action Plan for Herons of the World. Station Biologique de la Tour du Valat, Arles, France.
Kushlan, J.A. 2007. Sympatric foraging of Little Egrets and Snowy Egrets in Barbados, West Indies. Waterbirds 30:614-617.
Kushlan, J.A. and M.J. Steinkamp. 2007. Seabird nesting and conservation in the northern Bahamas. Waterbirds 30:613-623.
Angehr, G. and J.A. Kushlan, 2007. Seabirds and colonial wading bird nesting in the Gulf of Panama. Waterbirds 30:335-357.
Kushlan, J.A. and G.R. Angehr. 2007. Seaside tiger herons. Waterbirds 30:278-283.
Kushlan, J.A., W. Burke, M. Frost, and E. Massiah. 2007. Little Egret conservation in Barbados. Birds Caribbean 05-06:1-2.
Kushlan, J.A. 2009. Foraging and plumage coloration of the Galapagos Lava Heron (Butorides striatus sundevalli). Waterbirds 32:415-422.
Kushlan, J.A. and J.W. Prosper. 2009. Little Egret (Egretta garzetta) nesting on Antigua: a second nesting site in the Western Hemisphere. Journal of Caribbean Ornithology 22:108-111.
Wingate, D.B., J.L. Madeiros, and J.A. Kushlan. 2009. Green Heron colonizes Bermuda. Waterbirds 32:162-168.
Heath, J.A., P.C. Frederick, J.A. Kushlan and K.L. Bildstein. 2009. White Ibis (Eudocimus
albus), Ver. 2.0. The Birds of North America On Line (A. Poole, Ed.). Cornell Lab of Ornithology, Ithaca, New York. https://doi.org/10.2173/bna9
Kushlan, J.A. 2009. Status and conservation of the Magnificent Frigatebird (Fregata magnificens) on Barbuda, West Indies. Journal of Caribbean Ornithology 22:41-43.
Kushlan, J.A. 2009. Feeding repertoire of the Boat-billed Heron. Waterbirds 32:408-411.
Kushlan, J.A. 2010. White Ibis (Eudocimus albus) nesting in the Bahamas. Florida Field Naturalist 38:68-70.
Kushlan, J. A. 2010. Seabirds of Great Isaac Cay, The Bahamas. Florida Field Naturalist 38:110-112.
Kushlan, J.A. 2011. Longevity of the Tricolored Heron (Egretta tricolor). Journal of Heron Biology and Conservation. 1:2. www.HeronConservation.org/vol1/art2.
Kushlan, J.A., D.B. Wingate, and J.L. Madeiros. 2011. Conservation of the Green Heron (Butorides virescens) on Bermuda. Journal of Heron Biology and Conservation. 1:4. www.HeronConservation.org/vol1/art4.
Kushlan, J.A. 2011. Heron count protocols: Inventory, census and monitoring of herons. HeronConservation, www.HeronConservation.org.
Kushlan, J.A. 2011. Bibliography of the herons of the world. HeronConservation, www.HeronConservation.org.
Kushlan, J.A., C. Hickey, and A.W. White. 2011. The American Oystercatcher (Haematopus pallliatus) in the northern Bahamas. Journal of Caribbean Ornithology 24:48-55.
Kushlan, J.A. 2011. The terminology for courtship, nesting, feeding, and maintenance in herons. HeronConservation, www.HeronConservation.org.
Kushlan, J.A. 2012. A history of conserving colonial waterbirds in the United States. Waterbirds 35:608-625.
Angehr, G R., J.A. Kushlan, and K.N. Hines. 2014. Nesting sites and population estimates of seabirds and other waterbirds of the Gulf of Chiriquí, Panamá. Waterbirds 34:426-431.
Stier, A. and J.A. Kushlan (eds.). 2015. Agami Heron (Agamia agami) Conservation Plan. GEPOG Association, Cayenne, French Guiana.
Kushlan, J.A. and K. N. Hines. 2016. Behavior of the Agami Heron. Waterbirds 39:187-192.
Kushlan, J.A., G. Angehr, and K. Hines. 2017. Seabirds and other colonial waterbirds of the Caribbean coast of Panama. Journal of Caribbean Ornithology 30:145-153.
Stier, A., A. Ricardou, S. Uriot, N. de Pracontal, and J.A. Kushlan. 2017. Breeding season, home range and migration of the Agami heron (Agamia agami). Waterbirds 40:289-296.
Kushlan, J.A. 2018. Heron conservation – a history. Waterbirds 41:345-354.
Barlow, C.R., and J.A. Kushlan. 2020. First observations and film of the courtship display of White-backed Night Heron (Nycticorax leuconotus) in The Gambia, West Africa, Journal of Heron Biology and Conservation 5: Art 1. www.heronconservation.org/JHBC/vol05/
Heath, J.A., P.C. Frederick, J.A. Kushlan, and K.L. Bildstein (2020). White Ibis (Eudocimus albus), version 1.0. In Birds of the World (A.F. Poole, Editor). Cornell Lab of Ornithology, Ithaca, NY. USA.
Davis Jr., W.E. and J.A. Kushlan (2020). Green Heron (Butorides virescens), version 1.0. In Birds of the World (A. F. Poole and F. B. Gill, eds.). Cornell Lab of Ornithology, Ithaca, NY. USA.
Browning, M.R., and J.A. Kushlan. 2021. The Great White Heron is a species. Journal of Heron Biology and Conservation. www.heronconservation.org/JHBC/vol07/
Sangster, G., J. A. Kushlan, S. M. S. Gregory, and E. C. Dickinson 2023. Agamiinae, a new family-group name for the Agami Heron Agamia agami (Ardeidae). Avian Systematics 1 (4): 45–49.
Kushlan, J. A. 2023. Herons. In A. B. Hodgson, editor. Florida Breeding Bird Atlas II. Special Publication Number 9. Florida Ornithological Society, Tampa, USA.