Captive Breeding and Reintroduction: an Overview Bill Konstant

of Contents

rticles Captive Breeding and Reintroduction: An Overview Bill Konstant

Propagation and Reintroduction of Imperiled Plants, and the Role of Botanical Gardens and Arboreta Linda R. McMahan

Overview of the Goals and Activities of the IUCN Captive Breeding Specialist Group and International Species Information System Suzanne R. Jones, Editor

Translocations of Captive-Reared Terrestrial Vertebrates, 1973-1986 Brad Griffith, J. Michael Scott, James W. Carpenter, and Christine Reed

L 5 14 The Role of Genetics in Captive Breeding and Reintroduction for Species Conservation Alan R. Templeton

18 Decision-Making About A Reintroduction: Do Appropriate Conditions Exist? Devra G. Kleiman

20 Is Captive Breeding an Appropriate Strategy for Endangered Species Conservation? Tony Povilitis

24 Measures of the Value and Success of a Reintroduction Project: Red Wolf Reintroduction in Alligator River National Wildlife Refuge Michael K.Phillips

27 Ethical Perspectives on Captive Breeding: Is It For The Birds? Gary E.Vamer and Martha C. Monroe

30 Captive Propagation and the Conservation of Species: A U.S. Fish and Wildlife Service Perspective James Tate, Jr.

Case Histories Birds 32 The California Condor: Current Efforts for its Recovery Michael Wallace

36 The Decline and Restoration of the Guam Rai1,Rallus owstoni Gregory J. Witkman, Robert E. Beck, Jr., Stuart L. Pimm, and Scott R. Demckson

40 Peregrine Falcon Recovery Tom J. Cade

46 Captive Propagation in the Recovery of Whooping Cranes James C. Lewis Reptiles and Amphibians Endangered Species 48 Conservation of Crocodilians: The Release of Captive- UPDATE A forum for information exchange on Reared Specimens ehngered species hues F. Wayne King November 1990 52 Captive Breeding and Reintroduction: Recovery Efforts Vol. 8 No. 1 Suzanne Jones ...... Editor of the Virgin Islands Boa, Epicrates monensis grand Dr. Terry Root ...... Faculty Advisor Peter J. Tolson Jon Jensen ...... Staff Advisor

54 Release and Translocation Strategies for the Puerto Instructions for Authors: Rican Crested Toad, Peltophryne lemur The Endangered Species UPDATE Robert R. Johnson welcomes articles related to species protection in a wide range of areas including but not limited to: research and Plants management activities for endangered Central Florida Scrub: Trying to Save the Pieces species, theoretical approaches to species conservation, and habitat pro- Susan R. Wallace tection and preserve design. Book reviews, editorial comments, and an- Down But Not Out: Reintroduction of the Extirpated nouncements of current events and pub- lications are also welcome. Malheur Wirelettuce, Stephanomeria malheurensis Robert L. Parenti and Edward 0. Guerrant, Jr. Readers include a broad range of professionals in both scientific and policy Reintroduction of the Texas Snowbell, Styrax texana fields. Alticles should be writ& in an easily understandable style for a knowl- Paul Cox edgeable audience. Manuscripts should be 10-12 double Translocation of an Otherwise Doomed Population of spaced typed pages. For further information please contact Smne Barrett's Penstemon, Penstemon barrettiae Jones at the number listed below. Edward 0. Guerrant, Jr. Subscription Information: The Endangered Species UPDATE is Fish published approximately ten times per 68 Colorado Squawfish Reintroduction Efforts in the Lower year by the School of Natural Resources at The University of Colorado River Basin Michigan. Annual rates are $23 for Buddy L. Jensen regular subscriptions, and $18 for students and senior citizens (add $5 for postage outside the U.S.). Students A Fish Faunal Conservation Program: The Lake please enclose advisor's signature on Victoria Cichlids university leuehead; senior citizens Les Kaufman endose proof of age. Send check or money order (made payable to The University of Michigan) to: Apache Trout Culture: An Aid to Restoration Robert E. David Endangered Species UPDATE School of Natural Resources The University of Michigan Mammals Ann Arbor, MI 48109-1 1 15 79 Red Wolf: Recovery of an Endangered Species (3 13)763-3243 Michael K. Phillips Front Cover: Red wolf (Canis ruficr) Photo by U.S. Fish and Wildlife Service The Conservation Program for the Golden Lion Tamarin, Leontopithecus rosalia Back Cover: Devra G.Kleiman, Benjamin B. Beck, Andrew J. Baker, Peregrine falcon (Falco peregrinus) Jonathan D. Ballou, Lou Ann Dietz, and James M. Dietz Photo by Mike Riley The views expressed in the Endangered Black-Footed Ferrets On The Road To Recovery Species UPDATE are those of the author Tim Clark and may not necessarily reflect those of the U.S. Fish and Wildlife Service or The University of Michigan.

Extra copies of this and other past Special Issues are available for sale. Please Printed on Re~ycledPaper contact the Endangered Species UPDATE office for more information. @ Note From The Editor

My vision for this Special Issue began quite simply. The idea started with the observation that as the public's realization of the seriousness of today's environmental problems and the plight of a growing number of species has increased, so has the use of conservation management techniques, such as captive propagation and reintroduction, requiring resource-intensive manipulation of individuals, populations, and communities. On the one hand, I am excited about this rapidly growing body of management knowledge -- such as that concerning genetics, population viability analysis, and captive breeding protocols -- and the options that such new skills may offer to snatch back species teetering on the brink of extinction. Yet, on the other hand, I am wary of this human tendency of ours to try to solve environmental problems like habitat degradation and loss of biodiversity -- which are ultimately caused by human technology and disruption of ecological systems -- with increased human technology and even more "invasive" (from the viewpoint of the species) management methods. Thus, even while I applaud the management progress being made with endangered species, I wonder to what avail is the perfection of captive breeding techniques such as double-clutching, artificial insemination, and cross-fostering,if we still are unable to find workable resource use solutions to the problems which caused most species to become endangered in the first place. Therefore, with both interest and reservations about the increased use of captive propagation and . reintroduction, I began collecting articles from authors (many more than I originally intended!) which explore the whole spectrum of the topic: what captive breeding and reintroduction programs are currently underway, what are their goals and strategies, are they appropriate -- both in terms of effectiveness and ethics -- and finally, how well they actually are working in the field. The Special Issue begins with two articles providing general overviews of large-scale captive breeding and reintroduction initiatives for both plants and animals, followed by several pieces evaluating the effectiveness of past reintroductions, the genetics of captively breeding populations of rare species, and guidelines for determining when reintroductions are appropriate. Next comes a handful of articles addressing the appropriateness of captive breeding and reintroduction as a means of preserving endangered species, including discussions of effectiveness,measures of value and success of such programs, and ethical considerations. The issue then concludes with 18 case histories of representative bird, reptile and amphibian, plant, fish, and mammal species or species groups currently involved in captive breeding and reintroduction programs. Each synopsis briefly outlines the history of such captive breeding and reintroduction efforts and discusses the appropriateness of this management strategy in each instance. It is my hope that this Special Issue will contribute to the international discussion concerning the appropriateness of various management strategies for preserving endangered species, and provide insights to those working around the globe to preserve biodiversity.

Suzanne R. Jones UPDATE Editor Special Thanks to UPDATE Supporters: Wildlife Preservation Trust International Wildlife Preservation Trust International is a nonprofit organization dedicated to the preservation of species threatened with extinction. This is done through a multifaceted approach of captive breeding of endangered species, research in the wild and in captivity, reintroduction of species to the wild, professional training of conservation biologists and community conselvation education. National Fish and Wildlife Foundation The National Fish and Wildhfe Foundation is an independent not-for-profit foundation established by Congress in 1984 to encourage private sector involvement in conservation by matching private contributions with federal funds. The Foundation has developed part- nerships between public and private conservation organizations, corporations, individuals, and government agencies, leading to \ investment in over 290 projects in all 50 states, Mexico, Latin America, the USSR, and Tanzania. Major initiatives include: wetland conservation through the North American Waterfowl Management Plan, leadership training for wildlife professionals, and an annual assessment of federal natural resource agency budgets for Congress. Chevron Corporation Chevron Chevron salutes the ongoing efforts of the School of Natural Resources to share the latest ideas and opinions in the urgently compelling field of conservation biology. Our continued support for publication of the Endangered Species UPDATE is grounded in the belief that exchanging credible information is essential for developing sound resource management strategies and making effective public policy decisions.

2 Endangered Species UPDATE Vol. 8 No. 1 Captive Breeding and Reintroduction: An Overview

The idea that captive breeding can contribute to the survival of wild plant and animal species is not new. In fact, several large and charismatic animals driven to extinction in the wild within historic times survive today only in captivity and only because of successful efforts to breed them there. While they have been saved, however, thousands - perhaps hundreds of thousands - of other wild species have vanished without a trace, leaving no genetic reservoir. But can we measure the effectiveness of captive breeding as a tool to preserve biodiversity based solely upon the survival of species in zoological and botanical gardens, arboreta, and private collections? Through reintroduction efforts, individuals born and raised in captivity can help re- establish extirpated wild populations or prevent rapidly diminishing populations from dying out. This strategy for recovery is proposed for a growing number of species each year. The increased importance of these conservation techniques is apparent in the addition of the Captive Breeding Specialist Group and Reintroduction Specialist Group within the KJCN's Species Survival Commission. Similarly, the growing emphasis on captive propagation is illustrated by the establishment of the National Collection of Endangered Plants, maintained by a consortium of botanical gardens under the auspices of the Center for Plant Conservation. Those who would minimize the role of captive breeding and reintroduction in the preservation of biodiversity point to the larger picture, the need to save large areas of habitat and intact ecosystems. The contribution of saving single species, they argue, is insignificant by comparison and much less cost-effective. It is difficult, however, to derive a cost-to- benefit ratio for programs which focus public attention on "flagship species," programs that ultimately heighten awareness of larger environmental issues. Although many zoo professionals have a largely favorable attitude toward the one-two punch of captive breeding and reintroduction, it has never been proposed as a panacea to prevent extinction. Neither has it been suggested as an alternative to other, more promising strategies. However, some notable successes - the golden lion tamarin, peregrine falcon, and Arabian oryx among them - demonstrate that we can use captive breeding and reintroduction programs as stopgap measures when time is short and other strategies cannot immediately be brought into play. The task for today's conservation biologists is to determine how, when, and where captive breeding and reintroduction make the most sense in a holistic approach to conservation management. This Special Issue of the Endangered Species UPDATE represents an important step in that direction by putting a fair number of projects under the microscope. By considering the factors affecting the implementation of captive breeding and reintroduction efforts, as well as analyzing the progress which different agencies and organizations have achieved to date to save a variety of endangered species, we can certainly hope to develop a better yardstick for the future.

Bill Konstant Executive Director of Wildlife Preservation Trust International

Vol. 8 No. 1 Endangered Species UPDATE 3 Propagation and Reintroduction of Imperiled Plants, and the Role of Botanical Gardens and Arboreta by Linda R. McMahan --

Threats to Imperiled Plant~ natural rarity due to limitations of cli- based. The extraordinary efforts of The mate, geography, and the plant's own Nature Conservancy and other similar I find the lack of readily available natural history characteristics. The other land conservation organizations to ac- information about endangered plants type of rarity is the increased endanger- quire land that harbors endangered spe- curious, since there are so many of them. ment of ma, which were at one time cies is particularly relevant to the plants Farmore attention is given to endangered more wide-spread and common, due to within these habitats. Although we as the animals, even those less "glamorous" habitat loss. Classic examples of the conservation community do not know like desert pupfish and bats, than to the enough about plants and their conserva- approximately 5,000 species, subspe- tion to always be effective, we do know cies, and varieties of plant taxa in the that an essential part of what we must do United States that one might qualify as "Far more attention is is to preserve land upon which endan- rare, endangered,or threatened (these are given to endangered ani- gered plants can grow, preferably their taxa ranked as GI, G2, or G3 - or the mals, even those less own habitat "selected" through the evo- equivalent - by The Nature Conser- "glamorous" like desert lutionary process over many generations. vancy). This 5,000-plus figure is particu- pupfish and bats, than to larly staggering because it represents Strategies Used to Conserve about 20% of the native flora of the con- the approximately 5,000 Plants tinental United States, Hawaii, and species, subspecies, Puerto Rico. and varieties of plant Despite the herculean effort to acquire But how many people can list ten taxa in the United States land for rare and endangered plants, it has endangered plants? hobably only those that one might qualify as become clear over the last few decades botanical specialists in private organiza- that land acquisition alone is often not tions and government entities who ac- rare, endangered, or sufficient to secure a rare plant species tively work with the species themselves. threatened.. ." from extinction. Other factors affecting This lack of information by itself is wildlands, even protected ones, send one threat to the future survival of plant plant conservationists scrambling to species, as a lack of information also latter are plants from the central Florida learp more about species and how to breeds a lack of vocal advocates for plant scrub habitat which is being developed manage them. Critical needs include conservation. But the other, more tan- for housing and agriculture (see related information about germination, growth gible threats which occur are basically case study), along with plants from the requirements, and the basic life history the same as for animal ma. Such threats Los Angeles basin, southern tip of Flor- information so often lacking on rare and include human conversion of wildlands ida, and Oregon's Willarnette Valley. understudied species. Information on re- to urbanization, farming, water projects, Species from prairie remnants of the establishmentof rare taxa (even common and mining, as well as the less direct en- Midwest also fit this category. ones for that matter), seems singularly croachment into wildlands by exotic The concentration of rare species in lacking. Yet we must continue trying to flora and fauna and introduced diseases, the United States varies somewhat in lo- find the answers while we are working and interference with natural succession. cation, being lowest in the Midwest and towards conserving these same species. In addition, outright collecting affects Northeast and highest in the tropical or The roles of botanic gardens comple- many of the showier species, and is par- sub-tropical areas of Hawaii, Puerto ment those of other organizations that ticularly harmful for slow-growing Rico, Florida, and Texas. Other "hot specialize in inventory, land protection, plants such as cacti. At least one rare spots" of rarity are those areas of gener- and legal protection. Their contribution, species, Astragalus agnicidus, was ally high plant diversity such as the however, greatly exceeds the raising of nearly extirpated as a pest, presumed to Appalachians of the East and the coastal public awareness about rare plants, as be toxic to sheep. states of the West. important as that is. What is most impor- Tangible threats are further compli- Plant conservation is closely tied to tant is the basic study of rare plants, cated by two general types of rarity. (I am the land on which the plants grow. Plant conserving their germplasm, and learn- oversimplifying this categorization due species often have rather specific re- ing about establishment of plants in cul- to space restrictions, but it will serve to quirements, being adapted to the soil and tivation and in the wild. These tasks are emphasize the complex factors that con- climate of their habitat, and thus their not new to botanic gardens, as they are servationists face.) One overall type is ultimate conservation must be land- the same skills required to maintain the

4 Endangered Species UPDATE Vol. 8 No. 1 gardens maintain sons I will not explain here, seed banks germplasm collections are less applicable for seeds which are of over 300 of the large, oil-rich, or from tropical species. nation'srarest taxa, with When prepared properly, appropriate the number growing by seeds stored at sub-zero temperatures 50 to 100 taxa each year. remain viable many times longer than Extensive cooperative they would in the wild. The loss of efforts have evolved be- germplasm is slow, and if the initial tween the CPC and its sampling is large enough there is member gardens, and little need for breeding -- which would in- federal agencies such as evitably result in the loss of genetic diver- the Bureau of Land sity -- to replenish supplies. Seed banks Management (BLM), are perhaps the only efficient method of U.S. Fish and Wildlife offsite storage for annual and short-lived Vials of seeds in the Seed Bank for Rare and Endangered S cies at Service, U. S. Depart- perennial species, because maintaining The Berry Botanic Garden photo by L.R. z~nhrn ment of Agriculture static germplasm of species which would (USDA -- including the have to be bred constantly is virtually other plant collections in cultivation. Forest Service and National Germplasm impossible. Yet, these skills, combined with a healthy Repository), state agencies, heritage Many of the CPC member gardens, measure of scientific rigor and methods, programs, and The Nature Conservancy. including The Berry Botanic Garden, become useful conservation tools. maintain sub-zero seed banks. In addi- The advent of the Center for Plant Botanic Garden Conservation tion, seeds are stored at the USDA Na- Conservation (CPC) in 1985 was an Met hods tional Seed Storage Laboratory in Ft. important step in creating scientifically Collins,Colorado, under aMemorandum credible plant conservation at U.S. bo- Botanic garden methods vary depend- of Understanding between the CPC and tanic gardens and arboreta. The CPC, a ing upon the region of the country and the the USDA. This allows storage of seed private conservation group supported types of plants they conserve. This from rare and endangered plants in a primarily by individuals and founda- summary will only skim the surface of variety of optimal conditions, including tions, was founded as a consortium of these methods, but should provide some storage at the temperature of liquid nitro- botanic gardens (beginning with 18 and idea of their scope and applicability. gen. now standing at 20) to maintain offsite 1. Seed:Seed banks are good 2. Living collection^: Living collec- collections (called the National Collec- methods of conserving germplasm where tions are appropriate for some taxa, such tion of Endangered Plants) to use for they are applicable. They work best with as long-lived herbaceous perennials, conservation purposes. Gardens for the small seeds of low oil content from tem- shrubs, and trees. If space is no object, network were selected to represent as perate regions, where plants are adapted sufficient germplasm of living plants can many different geographic regions of the to periods of winter dormancy. For rea- be maintained long-term, up to several country as possible, in order to hundred in many cases, with fulfill the objective of being able careful maintenance of to grow any species outdoors if germplasm through backup cut- necessary. An important criterion tings or other propagation meth- for selection of institutions was ods. The living collections ap- that each participator have a proach can also be augmented by strong commitment to plant con- seed bank strategies where this is servation. The member gardens appropriate. At The Berry Bo- vary from the botanical giants like tanic Garden, for example, we are the Arnold Arboretum, New York currently contracting with the Botanical Garden, and the Mis- state of Idaho to conserve souri Botanical Garden, to small germplasm of disjunct popula- ones like The Berry Botanic Gar- tions of western dogwood (Cor- den in Portland, Oregon, the Bok nus nuttallii). Although common Tower Gardens in Lake Wales, elsewhere, this dogwood is in Florida, and the Mercer Arbore- serious decline in Idaho, possibly tum in Houston, Texas. due to fungal infection. While The founders of the Center botanists in Idaho work out pos- probably did not realize how sible conservation measures extensive these propagation ef- Seedlings of the endangered Sidulcea nebonia~in the green- there, the Garden will store wild- fOa house of The Berry Botanic Garden, being grown as pan of the become in a mere five conservation program for the Center for Plant Conservation collected seed and raise plants for years. In 1990, the 20 member photo by L.R. McMahan distribution to public facilities in

Vol. 8 No. 1 Endangered Species UPDATE 5 the Pacific Northwest to be maintained as 4. we do not have a base of published or germplasm repositories and "backup" to &: An area more difficult to character- practical data on which to base these the wild populations. ize in a single phrase is the role of botanic efforts; we are, in fact, learning as we go Another plant maintained as a living gardens in refining conservation meth- along. collection at The Berry Botanic Garden is ods -here referred to as development of After the initial agreement to under- Barrett's penstemon (Penstemon barret- conservation policy, for lack of a better take the effort, the appropriate seed or tiae). We maintain cuttings of plants that label. One example is the 1989 sympo- cuttings from nearby populations (from were destroyed in lock construction at sium hosted by the nearby Bonneville Dam. The exact CPC at the Mis- germplasm is being used to re-establish souri Botanical plants near the site (see related case Garden to learn study). state-of-the-art 3. Research: Research goes hand in means of obtaining hand with maintaining germplasm col- and conserving ge- lections. Routine studies of germination netically represen- and growth are undertaken by the CPC tative germplasm gardens to better understand the species from the experts in in their care. Seed viability is tracked. the field. A book Increasingly, however, the CPC gar- based on the sym- dens are entering other areas of research posium is in devel- as well. Botanists at the Rancho Santa opment, but the Ana Botanic Garden, for example, have CPC did not wait to used chloroplast DNA analysis and apply information isozyme studies to look at the taxonomic gleaned from the status of several rare species. This year, participants. and Endaneered S~eciesof the Pacific-Northwest The Berry Botanic Garden undertook an From this sym- Photo by LR.~c~aha: isozyme study of the rare Gentner's posium, the CPC mission bells (Fritillaria gentneri) under revised recommendations regarding the the exact population if material is avail- contract with the Rare Plant Conserva- amount of germplasm to collect from able) are sown and grown to transplant tion Program of Oregon's Department of each population of a rare taxon, and the size at the botanic garden. Sowing wild Agriculture. number of populations that should be seed might work, but may take as much Gardens are also looking into the role sampled to "capture" a large part of the as 100 times the seeds used to establish of tissue culture in conservation. At the genetic diversity represented by that plants, which is wasteful of germplasm North Carolina Botanical Garden, taxon in the wild. In response to these and less effectiveoverall. tissue culture of rare pitcher plants helps recommendations, the CPC and its These cultivated plants are then put establish living collections and raise member gardens have already begun into nature and receive a great deal of plants for the specialty trade. The recollection of some of the taxa already care, at least for their first year, includ- Arnold Arboretum is using tissue culture in collections to ensure adequate genetic ing watering, weeding, and perhaps to try to establish disease-free plants of representation. protection in wire cages to guard against Florida torreya (Torreya tarifolia), a 5. predation. This care is necessary species in serious decline in Florida's &: Finally, botanic gardens are in- because the plants did, after all, begin panhandle. The Berry Botanic Garden creasingly becoming involved with re- their lives in cultivation, and are not uses tissue culture to establish disease- establishment of wild populations of rare particularly predisposed to survive free living collections of endangered lil- and endangered species. This is a natural under the whims of nature. Seeds pro- ies. At some future time, it may even be relationship between botanic gardens duced by these transplants, however, possible to maintain tissue culture lines and land managing agencies, as botanic are typically left to fare on their own of trees and tropical species otherwise gardens are set up to propagate and grow and re-establish naturally. Often, several difficult to maintain as living collections. large numbers of plants from docu- "pulses" of horticulturally grown plants Such cultures would require much less mented sources. are needed to get the population off to a space, but much research remains to be Typically, a land management agency good start. done before this kind of germplasm col- or organization such as the BLM or The When the establishment is complete, lection becomes a reality. Nature Conservancy will approach a monitoring is essential to follow the Other areas of botanic garden re- botanic garden for assistance in a re- population and determine the need for search include population demographic establishment program, whether it be additional material. Similarly, managing studies, classic taxonomic research, seed reintroduction of a species to a former the factors that threatened the original germination studies (such as the role of site or introduction to a new site. Cur- population is critical. Plants left on their fire in germination), and life history rently, I would have to say that all such own may easily succumb to the same studies. establishments are experimental, since threats as their predecessors.

6 Endangered Species UPDA TE Vol. 8 No. 1 Effectiveness of Plant the species fiom central Florida is nearly be acquired to conserve habitat for a rare Reintroduction impossible due to their poor suitability plant than for an animal of comparable for seed storage and a typically short life population size. Even research, an ex- The question of whether plant reintro- span. Establishing germplasm reposito- pensive component, generally requires duction really works is difficult to answer ries in nature, i.e., establishing intro- less money for plant than animal species. because of the lack of experience to date. duced populations, may be the only ef- The public constituency for rare plants, We can be cautiously optimistic based on fective way of conserving them. even though still small, is growing. Quite a few good experiments now underway Another consideration, first articu- frankly, I do not think it is yet time with critically rare plants. The answer lated to me by Robert Jenkins of The to apply the principles of triage to rare will probably be that it works under cer- Nature Conservancy, is that of global plant conservation in the United States. tain conditions, although we have yet to warming. If global warming does in fact determine exactly what those conditions occur, most rare plants will be unable to Additional Sources of Information are. We can only be certain of success for survive in nature without our help. They those re-established populations that are simply will not be able to evolve or make Two goal sources of infomation, frequently containing references and discussions on plant reproductive and seem to produce age long geographic "treks" between habitats reintrodudion and restoration include: the Center structures like their counterparts else- quickly enough to survive so rapid a for Plant Conservation's quarterly newsletter where. Long-term monitoring for at least climatic change. In this instance, learn- Plant Cornervation (CPC,125 Arborway, Jamaica several decades will be necessary. ing about reestablishment of rare plants Plain, MA 02130-3520); and the Natural Areas Association's quarterly publication Natural Areas It is insufficient, for example, for will be essential if they are to survive. Journal (320 South Third Street, Rockford, IL seeds or plants to be simply placed out in 61 104). the wild and forgotten. Such a project is Conclusion Specific anicles are contained in: likely to fail if the original causes of Elias, T.E., ed. 1987. Conservation and Manage- ment of Rare and Endangered Plants. California extirpation of the population or species One last word - in my opening para- Native Plant Society, Sacramento, CA. 630 pp. are not controlled. Additionally, such graph I talked about the sheer volume of efforts will not succeed if the plant mate- rare plants. The enormity of the problem rial reintroduced is not sufficient to es- causes some people to wonder if we can Linda R. McMahan is Executive Director of 'he Berry Botanic Garden, 11505 SW Surnrne~lle tablish a reproductively stable popula- save them all. However, the great efforts Avenue, Portland, OR 97219. tion. made in the 1980s should give us all Successful reintroductions appear to be those set up as outlined in the previous section, i.e., those in which living plants are used and cared for in the wild, threat- ening factors are controlled, and ongoing monitoring is conducted. Examples include an experimental introduction - now in its fourth year- of Knowlton's cactus (Pediocactus knowltonii) in New Mexico, introduction of Texas snowbells (Styrax texana) at two sites in Texas' Edward's Plateau (see case study), and re-establishment of Penstemon barrettiae at Bonneville Dam. Many factors determine success, and many considerations are important in re- establishment efforts. Such elements include maintenance of genetic diversity needed to ensure establishment, the use of local vs. non-local germplasm and the effects of germplasm origin on the establishment effort, the success of various vegetation forms (e.g., annuals vs. perennials), and similar factors. Ultimately, of course, learning how to establish wild populations of rare plants may be essential to their survival in the sive by al- Pediocacfm krowltonii (Knowlton's cactus), over-collected for wild. Susan Wallace of Bok Tower the canmercial market, is known from only one population in lhoughthe cost of land northwest New Mexico. Reintroduction into an ecologicnlly Gardens argues persuasively that mainte- quisition is high. But even similar location has yielded survival rates of more than 75%. nance of germplasm offsite for many of then, usually less land must Photo: Peggy OIweU. Center for Plant Conservation

Vol. 8 No. 1 Endangered Species UPDATE 7 Overview of the Goals and Activities of the IUCN Captive Breeding Specialist Group and International Species lnformation System by Suzanne R. Jones, UPDATE Editor [Regrettably, due to time conflicts, Ulys- volunteers from all over the world, sew- ties and personnel for international col- ses S. Seal, CBSG Chairman, and Tom ing three-year volunteer appointments. laborative captive propagation pro- Foose, CBSG Executive Oficer, were Among other things, these volunteers are grams. unable to meet our press deadline with responsible for procuring their own fi- 2) Establish a global network of pro- their intended article on "The Role of nancial support to fund their conserva- fessionals in captive and wildlife man- Zoos and the Captive Breeding Specialist tion endeavors. agement, population biology, and other Group in Captive Breeding and Reintro- disciplines to advise on the development duction of Endangered Species." (Look The Captive Breeding Specialist and conduct of recommended captive for an article in a later issue of the Group propagation programs. Collaborative UPDATE.) Although we cannot begin to programs are already underway for such replace their insights and broad under- One specialist group, the Captive species as the Sumatran rhino, kouprey, standing of the field, this short article is Breeding Specialist Group (CBSG), is and lemurs. an attempt to provide a brief overview of unique in that it cuts across taxonomic 3) Prepare Captive Breeding Action the goals and activities of the CBSG and categories. The IUCN maintains that Plans for all of the vertebrates in collabo- ISISfor those who are unfamiliar with the habitat protection alone is likely to be ration with the appropriate SSC and In- important role which they undertake. insufficient to maintain biological diver- ternational Council for Bird Preservation The following material was compiled by sity, especially for species numbering Specialist Groups. These plans are to pro- the UPDATE Editor directly from public 1,000 or less in the wild. Consequently, vide analyses of the status of the species relations material put out by the CBSG self-sustaining captive populations need in captivity and the wild, and make rec- and ISIS, and from converiations with Nate Flesness, ISIS Executive Director.] habitat protection alone is likely to be insufficient to Background Organizational ". .. lnformation maintain biological diversity, especially for species numbering 1,000 or less in the wild. Consequently, self- The International Union for the Con- sustaining captive populations need to be established servation of Nature and Natural Re- sources (IUCN) is the largest interna- before species are reduced to critical numbers . . ." tional professional conservation organization - active in over 120 countries - in which states, government agencies, to be established before species are re- ommendations of priority species for and non-governmental organizations duced to critical numbers, and thereafter captive breeding programs. Action Plans participate equally in conservation ef- need to be "coordinated internationally generally focus on broader taxons than forts. Within the IUCN are six commis- according to sound biological principles, the species level. As of yet, only a hand- sions, one of which is the Species Sur- and a view to the maintaining or re- ful of taxonomic orders have Captive vival Commission (SSC). The SSC is establishment of viable populations in Breeding Action Plans in some stage of comprised of a network of 100 specialist the wild." The CBSG is thus charged development; participants are currently groups, each of which generally focus on with advising the SSC and the IUCN on writing drafts for primates, parrots, a particular group of species, e.g., the the uses of captive propagation for con- carnivores, and antelopes. Primate Specialist Group. servation purposes, and the organizing, Similar recovery plans are being The primary responsibility of the facilitating, and monitoring of interna- developed by international, national and IUCN Specialist Groups is to develop tional captive propagation programs. regional professional associations under Action Plans for taxa under theirjurisdic- The CBSG primarily addresses animal the auspices of the American Association tion. Action Plans are specific proposals species, particularly vertebrate species, of Zoological Parks and Aquariums. for stabilizing and recovering imperiled within the SSC. These Species Survival Plans (SSP), and species, with an emphasis on recovery in Specific CBSG objectives include the other equivalent international plans, re- the wild. Specialist groups receive, at following: quire close coordination between partici- best, minimal resources from the IUCN, 1) Establish a global network of zoo pating institutions, e.g., for moving ani- and are staffed and run by 3,000-plus professionals and zoos to provide facili- mals between zoos, coordinating quaran-

8 Endangered Species UPDA TE Vol. 8 No. 1 tine zones, etc. Currently, there are a total of about 153 SSPs (53 in North America and an additional 100 from around the world) in some stage of development, with a goal to expand this number to 1,000. The CBSG tries to coordinate these various types of plans and efforts within the captive breeding community. 4) Assist in the organization and implementation of captive breeding programs recommended in Action Plans, including preparing studbooks, assisting in collaboration with Specialist Groups to obtain animals from the wild, conducting workshops on conservation biology and population viability analysis of individual species, and making specificman- agement recommendations. 5) Assist in the adoption and use of effective information systems and networks, such as ISIS and ARKS, by all of the world's zoos. These information ~eveilyendan~ereddue to poaching and habitat destruction, fewer than 800 Sumatran (or hairy) activities are essential to the work of the rhinos, bicrrorhinur swnaliensir, remain on earth. In an emergency effolt to preserve this species, CBSG, and so are discussed below in four U.S. zoos (including the Bronx Zoo) and the Indonesian Government established the Sumatran further detail. Rhino Trust in 1987, to aid in rhino conservation in southeast Asia and capture doomed rhinos for captive breeding programs in Indonesia and the U.S. Photo: New York Zoological Society Captive Breeding Information Systems graphic management of captive popula- software tool for creating and maintain- tions. Additionally, data on total captive ing studbooks, performing demographic The International Species Informa- holdings for each species allows partici- and genetic analyses, and supporting tion System (ISIS) is a global zoo animal pating institutions to better allocate population management. information system which works very available zoo space among imperiled One final informational activity of the closely with the CBSG, and is likewise species, and thus better fulfill their in- captive breeding community is the housed at the Minnesota Zoological creasing conservation responsibilities. preparation and distribution of the CBSG Garden. This computer-based database Every six months, ISIS distributes to News. This newsletter, recently estab- contains information on over 117,000 dues-paying members registries of lished to promote communication of living vertebrates, plus a greater number "who-has-what," and short summaries of news and information among people of their ancestors, from more than 363 the captive status of the over 4,200 regis- with "special interest in the uses, re- zoological member institutions in 38 tered taxa. Such information can aid sources, and problems of captive breed- countries (in particular North America, captive managers in searching for a ing as a tool for the conservation of en- Europe, Australasia, and expanding to needed specimen or locating people with dangered species," is sent to to all mem- Latin America, Asia, and Africa). Basic expertise on a particular species. Also bers of the CBSG, chairpeople of Spe- biological information such as age, sex, maintained by ISIS are partial zoological cialist Groups, wildlife agencies and parentage, place of birth, and circum- studbooks for all of the 4,200 taxa regis- departments with captive breeding pro- stance of death is recorded to use in tered with ISIS. (For 95% of all captive grams, and to all of the world's zoos. various captive population analyses. species,noformal studbookexists,hence The long-term success of these nu- According to the CBSG, properly ISIS is the only available source for this merous initiatives, and the effectiveness managed captive populations are impor- information.) of these ambitious and multifaceted or- tant in providing biodiversity insurance Related systems also available from ganizations remain to be seen. There is in case species go extinct in the wild, and ISIS include: no doubt, however, that the CBSG and to prevent captive populations from 1)ARKS (Animal Record Keeping ISIS are and will continue to be pivotal being a burden on wild populations (ISIS System), a computer system for main- contributors to captive breeding efforts data indicate that 92% of new zoo mam- taining standardized zoological inven- to preserve endangered species. mals, 7 1% of birds, and a "majority" of tory records - currently used by over Additional information may be ob- herps are now captive-bred.) The line- 250 zoos and related institutions; and tained by writing to the CBSG or ISIS at age information available on ISIS is par- 2) SPARKS (Single Population 12101 Johnny Cake Ridge Road, Apple ticularly useful for genetic and demo- Analysis & Records Keeping System), a Valley, MN 55124, USA.

Vol. 8 No. 1 Endangered Species UPDATE 9 Translocations of Captive-Reared Terrestrial Vertebrates, 1973-1986 by Brad Griffith, J. Michael Scott, James W. Carpenter, and Christine Reed

A translocation is the intentional re- to maintain species diversity. (Griffith et al. 1989) analyzed factors lease of animals into the wild in an at- Several species (e.g., black-footed associated with success or failure of tempt to establish (introduce), re-estab- ferret, California condor) were relegated contemporary translocations of native lish (reintroduce),or augment (restock) a to captive populations as a result of sys- terrestrial vertebrates. Our analysis free-ranging population (IUCN 1987), tematic pressures on their populations combined translocations of both exclu- and may consist of single or multiple and habitats. The ultimate goal of main- sively captive-reared and exclusively releases at one or several sites. An in- taining these captive populations is even- wild-caught animals. In this article we creasing appreciation of biological diver- tual re-establishment of viable free-rang- focus on analysis of all translocations sity has focused attention on transloca- ing populations. Successful re-establish- that includedat least some captive-reared tions of rare native species to restore or ment of wild populations requires an animals and compare the results to preserve diversity in communities of understanding and effective manage- translocations of exclusively wild- &-ranging animals. If current patterns ment of all aspects of reintroductions. caught animals. of habitat loss continue, natural commu- Several studies analyzed factors that nities may become restricted to disjunct influenced success or failure of introduc- Translocated Species and habitat fragments. Rates of species tions and colonizations of non-native Release Sites extinction may be expected to increase in species (Crawley 1986, Moulton and these small fragmented habitats, and Pirnm 1986, Newsome and Noble 1986, In our 1987 survey (Griffith et al. periodic translocations may be required O'Connor 1986). In an earlier paper, we 1989), we obtained detailed reports on 405 translocations of native terresmal Table 1. Tranel~onsaf native species fmm or within Aurtralia, Canada,New Zealand, vertebrates from Australia, Canada, New Zealand, and the United States that were and the United States, t973-1986, that included at least sane captive-reared animals. conducted during 1973-1986. Eighty Species F"9mQJ Species F"1m~r (20%) of these translocations consisted exclusively of captive-reared animals (1 8 apies) 78 $dam,&(10 species) 19 and 23 (6%) consisted of both captive- reared and wild-caught animals. The Andean wndor 2a Arabian oryx 1 a remaining 302 (74%) translocations were of exclusively wild-caughtanimals. Antipodes Island parakeet la Bighorn sheep zb The purpose of translocations was simi- Bald eagle 15 Brush-railed bemg 5 lar for captive-reared and wild-caught White-tailed Barn owl. 1 deer lb animals: 47% to reestablish extirpated BlaJi stilt 4a Elk lb populations; 32% to augment existing Brown teal 6' Golden lion tamarin 1 populations; and 21% to establish new Bumwing owl 2 Numbat 1 populations. Canada goose 13~ Red wolf la The 103 translocations of captive- Masked bobwhite la Swift fox 4a reared animals included 32 species Northern black duck 1a Woods bison 2 (Table 1). Most (56%) species were Newall's manx shearwater 1 birds, and these comprised 76% of Nene la 6 translocations that included captive- Petegrine fdm 1 (4 speciesf reared animals. Raptors comprised 22% Puerto Rican parrot 1 a of the species and 46% of the transloca- Redmedparakeet 3% Houston t6ad la tions among captive-reared birds. Among buds, 11%of speciesand 19%of SandhiU crane 5 Johnston's crocodile 1a translocations were native game birds; Trumpeter swan 1 Kanp's ridey sea turtle 3% the remaining birds were threatened, Wild tudcey zb Puem Rican crested toad 1 endangered, or sensitive species, or their surrogates. a No tranrlocations of exciusively wild-caught animals were conducted for these species. Mammals comprised 31% of the b Native game animals; remainder are threatened, endangered, sensitive, or non-game sutroga~esfor endangered species. species and 18% of the 103 translocations that included captive-reared ani-

10 Endangered Species UPDA TE Vol. 8 No. 1 mals (Table 1). Ungulates, (19%), or by mixed methods marsupials, and canids were (22%). The remainder (9%) about equally represented. received no rearing assis- Among these mammals, 30% tance. of the species and 21% of the translocations were native Comparisons Between game species and the remain- Translocations of Cap- ing mammals were threat- t ive-Reared and Wild- ened, endangered or sensitive Caught Animals species (Table 1). Reptiles and amphibians accounted for Many characteristics of 13%of the species and 6% of translocations of captive- the translocations. reared animals were empha- Most (73%) of the 103 sized by comparison to translocations including cap translocations of wild-caught tive-reared animals occurred animals. Differences between in the United States and Can- translocations that included at ada (Table 2). At least 9 Cana- least some captive-reared ani- dian provinces or territories mals and translocations of ex- and 28 states released captive- clusively captive-reared ani- reared native species during mals were insignificant. 1973-86. The remaining However, significant differ- translocations were distrib ences between translocations uted among Australia (6%), that included any captive- New Zealand (14%), and reared animals and transloca- other countries (8%). Cap tions of exclusively wild- tive-reared animals were re- caught animals were numer- leased at6 locations in Austra- ous (Table 3). lia, l l locations in New Zeal- There were differences be- and, and in 6 other countries. tween the kinds of animals translocated, but not between Unique Properties the characteristics of areas of Translocations of where they were released Captive-Reared Animals (Table 3). Translocations of exclusively wild-caught ani- Several characteristics of mals were in equal propor- translocations were unique to tions for game species com- captive-reared animals. pared to threatened, endan- These included source and gered, or sensitive species and supplementation of brood for birds compared to mam- stock, source and incubation mals. Translocations of cap- methods for bird eggs, and tive-reared species were rearing methods. dominated by threatened, en- The brood stock for translocated cap- Incubation of source eggs in captivity dangered, or sensitive birds. However, tive-reared vertebrates was exclusively was by mechanical means (27%), by there was no difference between translo- captive-propagatedfor 42% of transloca- source species (34%),by a foster species cations of captive-reared and wild- tion~,exclusively wild-caught for 27%, (5%), or by mixed methods (34%). In caught animals regarding location of the and composed of both wild and captive captivity, captive-propagated young release area in relation to the historic stock for 31% of translocations. In 60% were reared by their biological parents species range or release area habitat of translocations of captive-reared ani- (49%), by same-species foster parents quality. mals, the captive brood stock had been (4%), by different-species foster parents The increased care of captive-reared supplemented with animals from other (3%), by puppets (3%), directly by hu- animals was evident in several compari- populations of either wild- or captive- mans (12%), or by mixed methods sons (Table 3). At release, captive- propagated vertebrates. Most (86%) (29%). Once released, captive-propa- reared animals were in better physical eggs yielding captive-reared birds for gated young were reared by their biologi- condition than were wild-caught ani- translocation were from captive popula- cal parents (22%),by same-speciesfoster mals. Habitat improvement prior to re- tions, an additional 11% were from the parents (16%), by different-species fos- lease was more likely for captive-reared wild, and 3% were from mixed sources. ter parents (12%), directly by humans animals, and captive-reared animals

Vol. 8 No. 1 Endangered Species UPDATE 1 I likely to be present in the release area for TaUe 3. Camprison of chnracteristics between native bird an& mammal mslocariaas with translocations of captive-reared animals. at least some captive-reared animals and translocations Q£ exclusively wild-caught Mim& Captive-reared species also were more from or within Ausuatia, Canada, New Zealand, Md the United States, 1973-1986. Data obtained 041 GdTith a aL (1989); 10 reported tran$Wonsof ~ptifesot amghbians were likely to be late breeders with small aoluded fmanalysir. Animals that first gave birth at age 2 or younger with averagedutch clutches or litters, were less likely to be size of three arman are amsidered enrly bRedwa with large clutches; all others are late herbivores, and were more likely to be breeden with mall &tches, carnivores than wildcaught species. .,, Operational differences between ceof Ca-give-d wwught --Test Statistics translocations of captive-reared and wild-caught animals were not always Vaaable n Percent n Percent G P statistically different. An equivalent proportion (14%) of released animals in al Dedignatim 9ative game spedes 15 15.5 148 49.7 38.8 ~0.001 both groups were radio-tagged. There 'lha~,endangered, was a non-significant tendency for the or semitiwe species 82 84.5 150 9.3 average translocation of a captive-reared Taxa species to consist of more releases (8.2 Birds 78' 80.4 154 513 26.7 4.001 Mammals 19 19.6 144 48.3 vs. 5.5, P = 0.237) and to release more Location of =lease area individuals (143 vs. 87, P = 0.223). The Core of hisuric nmge 7 1 76.3 199 69.8 1.5 0.221 average length of the release program for Peripheryoroutside 22 23.7 86 30.2 captive-reared animals (3.7 yrs) was Release area habitat greater than for translocations of wild- Excellent 35 39.3 88 30.6 2.6 0.278 Good 41 46.1 158 54.9 caught animals (2.9 yrs., P= 0.043). The Fair or Poor 13 14.6 42 14.6 sex ratio of released animals was slightly Habitat impvanent prior higher for captive-reared (1.13) than for No 48 55.8 181 ~,3 3.1 0.080 wild-caught animals (0.85, P = 0.04). Yes 38 44.2 92 33.7 Released same day delivered to ana Factors Associated With Success Yes 37 41.6 229 80.4 46.3 d.001 of Translocations of Captive- No 52 58.4 56 19.6 Reared Animals Hard or soR releasc Hard 27 29.7 m n.7 69.1 d.001 Soft 64 70.3 65 22.3 Translocations that included captive- reared animals were only about one-half Avem c physical condition m rAse as likely to be successful as were translo-. Excellent 58 66.7 116 41.6 17.7 0.001 cations of exclusively wild-caught ani- Good 26 29.9 151 54.1 Fair or poor 3 3.4 11 3.9 mals (Table 3). However, translocations Potential armpetiton of captive-reared animals shared five 32 35.2 55 19.2 28.9 4.001 characteristics that were associated with a? 36 39.6 68 23.8 Neither 23 25.3 163 57.0 reduced probability of success among all translocations (Griffith et al. 1989). Adult food habits Camivon 49 50.5 72 24.2 40.0 4.001 Translocated captive-reared animals Htibivare 36 37.1 216 72.5 were more likely (P 2 0.001): a threat- Omdivar~ 12 12.4 10 3.4 ened, endangered, or sensitive species; a Early breeder, la e clutch 14 14.4 127 42.6 28.1 d.001 bird instead of a mammal; a late breeder Latebreeder, rm$ clutch 83 85.6 171 57.4 with a small clutch; a carnivore or omni- Result of translocation vore rather than an herbivore; and to have Failure 24 61.5 41 25.2 17.9 d.001 Success 15 38.5 122 74.8 morphologically similar or congeneric competitors present in the release area than were translocated wild-caught were more likely to have been held on the than wild-caught translocated animals to animals (Table 3). Countering these ef- release area prior to liberation and to have die due to capture and handling prior to fects, translocations of captive-reared been "soft released" (food and/or shelter release (1% vs. 6% respectively, animals shared two characteristics that provided on site) than were wild-caught P = 0.164). were associated with increased animals. If held on the release area, there Release communities and ecologi- probability of success among all translo- was a non-significant tendency to hold cal characteristics of translocated species cations (Griffith et al. 1989). Transloca- captive-reared animals longer (14.8 vs. differed between translocations of cap- tions of captive-reared animals lasted 2.6 days, P = 0.18) than wildcaught tive-reared and wild-caught animals longer and tended to release more ani- animals. There was also a non-signifi- (Table 3). Congeneric or morphologi- mals than translocations of wild-caught cant tendency for fewer captive-reared cally similar competitors were more animals.

12 Endangered Species UPDA TE Vol. 8 No. I Limited data precluded rigorous Appropriateness of Translocation have not been reduced) did not exceed analysis of factors associated with suc- of CaptlveReared Animals 10% for releases as large as 400 birds cess or failure of translocations of cap- (Griffith et al. 1989). Persistence of the tive-reared animals. Only 39 of the 103 Translocation of captive-reared ani- species is enhanced more by keeping the translocations could be classified as a mals may be preferable to translocation animals in captivity and by establishing success or failure; the survey respon- of wild-caught animals when a captive disjunct captive populations than by re- dents could not ascertain success or fail- population exists and removal from a leasing them in unsuitable habitat. ure of the remainder of translocations at single wild population significantly re- Because release areas outside or on the time of data collection. duces theviability of the wild population. the periphery of historic ranges of species Among the 39 translocations of cap- Translocation of captive-reared animals are associated with a marked reduction in tive-reared animals that could be ana- is also warranted when IUCN (1987) translocation success, movement of cap- lyzed, only legal designation and food guidelines are followed, and when a tive-reared threatened, endangered, or habits showed significant differences in comprehensive analysis indicates a rea- sensitive species to such areas should be success rates. Only 29% of the 31 sonable chance of: re-establishinga free- limited to situations without alternatives. translocations of threatened, endan- ranging population that enhances diver- Regardless of the number released, re- gered, or sensitive species were success- sity of native species at an appropriate leasing animals in "good" habitat quality ful whereas 75% of the 8 translocations scale; establishing a satellite population in the core of the species range is a better of native game species were successful that reduces susceptibility of a species to alternative than releasing them in "excel- (P .= 0.018). The success rate was 21% extinction from catastrophic loss of a lent" habitat quality on the periphery or for 14 translocations of carnivores, 61% single wild population; speeding the re- outside of the species historic range for 18 translocations of herbivores, and covery of a species after limiting factors (Griffith et al. 1989). 14% for 7 translocations of omnivores have been ameliorated; or effectively We implore people engaged in (P = 0.022). There was anon-significant augmenting genetic heterogeneity of a translocations of captive-reared animals tendency for translocations of birds to be small population with depleted genetic to keep detailed and accurate records. less successful (32% of 31 transloca- diversity. In the latter case, the admoni- Currently, we have insufficient data for a tion~)than mammals (62% of 8 translo- tions of Greig (1979) and IUCN (1987) to rigorous analysis of specific factors such cations; P = 0.121). However, because avoid genetic "pollution" of disjunct as genetic heterogeneity, sex and age data were limited, expected cell populations shouldbe heededif possible. composition, or specific rearing and frequencies were low enough to cast Reasonable chance of success is handling procedures that may affect the doubt on the statistical validity of all situational. Ideally, greater than a 50% success of translocations of captive- these univariate tests. chance of successis aminimum criterion. reared animals. Only by increasing the Limited data on translocations of However, a lower probability of success quantity and quality of the database can captive-reared animals precluded multi- may be acceptable if the only alternative we hope to fine-tune the process and variate analyses of all potentially impor- is to maintain the species exclusively in increase the effectiveness of the tech- tant variables. However, the absence of captivity. Acceptance of a less than 50% nique. differences in success rates between chance of success should not be due to translocations of exclusively captive- releasing an insufficient number of ani- Acknowledgements reared and exclusively wild-caught animals, Exclusively captive population(s) mals in multivariate analyses (Griffith et should be increased prior to a transloca- The original work was conducted when B. al. 1989), permits evaluation of alternate tion to provide enough animals to maxi- Griffith was in the Department of Fisheries and Wildlife Resources, Univenity of Idaho, Moscow translocation strategies for captive- mize the chance of success given the ID 83843, and J. C. Carpenter was a Research reared animals with Griffith et al.'s characteristics of the available release Veterinarian with the U. S. Fish and Wildlife Serv- (1989) model. Success of translocations area and the translocated species. ice at Patuxent Wildlife Research Center, Laurel for threatened, endangered, or sensitive Conversely, for species relegated to MD 20708. Conducted under the auspices of the Idaho Cooperative Fish and Wildlife Research species was enhanced by excellent habi- captivity, removing animals for translo- Unit, which is funded and supported by the Idaho tat quality in the release area, releasing cation should not appreciably reduce Department of Fish and Game, U. S. Fish and animals in the core of the historic range viability of the captive population. Ide- Wildlife Service, and the Wildlife Management rather than on the periphery or outside, ally, multiple viable captive populations Institute. This is contribution number 562from the University of Idaho Forest, Wildlife, and Range increasing the number of released ani- should be established prior to transloca- Experiment Station. mals, and increasing the number of years tion to minimize the chance of species in which animals were released. Scott extinction. Literature Cited and Carpenter (1987) and Kleiman Except for experimental determina- (1989) emphasized that protected and tion of limiting factors, captive-reared Crawley, M.J. 1986. The population biology of maintained habitat, identification and animals should not be released in poor or invaders. Phil. Trans. R. Soc. Lond. B314:711- control of limiting factors, and proper fair habitat. Estimated success for 731. care and training of captive-reared ani- Greig, J.C. 1979. P~ciplesof genetic conserva- aanslocations of threatened, endan- tion in relation to wildlife management in mals are prerequisites for successful gered, or sensitive birds to poor or fair Southern Africa. S. Afr. J. Wildl. Res. 957-78. translocations. habitat (i.e., areas where limiting factors Griffith, B., J.M. Scott, J.W. Carpenter, and C. (continued on bottom of next page)

VoI. 8 No. 1 Endangered Species UPDATE 13 The Role of Genetics in Captive Breeding and Reintroduction for Species Conservation

by Alan R. Templeton

Biodiversity is the variety of life, in- pace (e.g., global warming). Long-term ing program, as well as deal with practi- cluding variation at the genetic, species, survival, therefore, depends upon the cal constraints. and community levels. Today there is a released population having sufficient crisis in biodiversity at all of these levels. genetic variability to provide adaptive PRESERVING GENETIC The natural habitats of many species flexibility in an uncertain future. Hence, VARIABILITY have already been completely destroyed, captive populations must be managed to and those of many others have been so preserve genetic diversity. Founder Populations reduced and fragmented that the species A second goal of genetic management are in danger of extinction. Even when is to alter the genetic composition of the In order to preserve a species' genetic habitat preservation is possible, it fre- species as little as possible from its initial diversity in captivity, it is obviously quently requires the reintroduction of state. Unfortunately, these two goals are necessary to carry over much of that diversity from the natural population into the initial captive population. The founding size of the captive population is a ". . .satisfying the goal of preserving genetic diversity critical parameter in determining the makes it difficult to satisfy the goal of not altering the extentofthiscarry-over,~iththeamount species' genetic composition. consequently, the ge- of genetic diversity increasing with increasing initial size. Unfortunately, netic manager must often balance contradictory goals many captive breeding programs are es- in the breeding program. . ." tablished only after the natural population has already been reduced to a very few individuals (e.g., the California con- propagules. Thus, captive breeding is sometimes contradictory. Captivity may dor), so much of the species' genetic needed topreserve a species that has gone induce selective forces, and populations diversity has already been lost. More- extinct in nature, to provide a backup for that respond to these novel selective re- over, with small founding numbers, the habitat preservation efforts, or to serve as gimes will be altered from their wild genetic manager will also frequently a source of propagules for reintroduction. genetic constitution. The capacity to have to deal with additional genetic prob- The goal of most captive breeding respond to altered selective regimes lems such as inbreeding depression programs is to maintain the species in depends upon genetic diversity in the (Templeton and Read 1984). Hence, captivity until release back into nature is population. Hence, satisfying the goal of rather than establish captive populations possible. However, the released popula- preserving genetic diversity makes it as acts of desperation, it is better to estab- tion will often experience an environ- difficult to satisfy the goal of not altering lish them as "insurance policies" when ment that is different from its original the species' genetic composition. Conse- the natural populations are still suffi- one, and all environments are subject to quently, the genetic manager must often ciently large to contain much genetic change, often at an increasingly rapid balance contradictory goals in the breed- diversity.

(continuedfrom preceding page)

Reed. 1985). Translocation as a species conser- Case, eds. Community Ecology. Harper and Brad Griffith is Assistant Leader of the Maine vation tool: Status and strategy. Science Row Publishing, Inc., New York. Cooperative Fish and Wildlife Research Unit, Uni- 245:477-480. Newsome, A. E.,and I.R. Noble. 1986. Ecological versity of Maine, U.S. Fish and Wildlife Service, IUCN. 1987. The NCN Position Statement on and physiological characters of invading spe- Orono, ME 04469; J. Michael Scott is Leader of Translocation of Living Organisms. 22nd cies. Pages 1-20 in R. H. Groves and J. J. the Idaho Cooperative Fish and Wildlife Research IUCN Council Meeting, Gland, Switzerland, 4 Burdon, eds. Ecology of Biological Invasions. Unit, U.S. Fish and Wildlife Senice, University of September 1987. 20 pp. Cambridge Univ. Press, New York. 166 pp. Idaho, Moscow. ID 83843; James W. Carpenter is Kleiman, D.G. 1989. Reintroduction of captive O'Connor, R. J. 1986. Biological characteristics Associate Professor at the Department of Clinical mammals for conservation: Guidelines for rein- of invaders among bud species in Britain. Phil. Sciences, College of Veterinary Medicine, Kansas troducing endangered species into the wild. Trans. R. Sw. Lond. B 314:583-598. State University, Manhattan, KS 60506, and BioScience 39:152-161. Scott, J. M., and J. W. Carpenter. 1987. Release of Christine Reed is Conservation Officer (Agricul- Moulton, M.P., and S.L Pimm. 1986. The extent captive-reared or translocated endangered ture) at the Department of Conservation, Twizel, of competition in shaping an introduced birds: What do we need to know? Auk 104544- New Zealand. avifauna. Pages 80-97 in J. Diamond and T.J. 545.

14 Endangered Species UPDA TE Vol. 8 No. I "Simply using a large number of founders does not ensure that genetic diversity is being preserved. Without a knowledge of how the species' genetic diversity is divided within and between local populations, it is impossible to design a sampling program that is ensured of preserving the species' genetic diversity."

When the numbers in the founding this species' genetic diversity is to ultimate carrying capacity as rapidly as population are low, the situation is more sample a few animals from many differ- possible. For example, a captive popula- difficult genetically, but not hopeless. ent glades, rather than large numbers of tion of Speke's gazelle (Gazella spekei) Neiet al. (1975) showed that populations animals from a few glades. was founded in 1969 from one male and established from a single mated pair can This example illustrates the impor- three females (Templeton and Read carry over a sizable portion of the genetic tance of genetic surveys on endangered 1984). From an analysis of inbreeding diversity found in the ancestral popula- species. Simply using a large number of depression in this species, Templeton tion when the ancestral population is founders does not ensure that genetic (1987) inferred that the natural popula- panmictic (not genetically subdivided) diversity is being preserved. Without a tion was large and panmictic. Therefore, and randomly mating. These conditions knowledge of how the species' genetic much of the species' genetic variability ensure that much of the species' overall diversity is divided within and between would be in the form of individual genetic diversity is present in the form of local populations, it is impossible to heterozygosity. The population was also individual heterozygosity. Thus, a few design a sampling program that is en- expanded at a rapid rate after its estab- individuals can carry over much genetic sured of preserving the species' genetic lishment (Templeton and Read 1984). diversity. Unfortunately, one or both of diversity. Because of these two factors, the theory these conditions are frequently violated. of Nei et al. (1975) predicts this captive For example, my laboratory and the Preventing Loss of herd should have much genetic variation. Missouri Conservation Commission Genetic Diversity Direct genetic surveys (Templeton et al. have been involved with sampling natu- 1987) revealed that the current captive ral populations of the collared lizard, Once the founder population has been herd is polymorphic for 14%of its loci, a Crotaphytus collaris, for reintroduction established, it is also necessary to prevent figure that is typical for large grazing on restored habitats in the Missouri the loss of the genetic diversity that has mammals (1 1.4%) or large mammals in Ozarks (Templeton et al. 1990). These been carried over from the natural popu- general (13.2%) (Baccus et al. 1983). lizards live on Ozark glades: rocky, tree- lation. Once again, population size has a Hence, despite an extreme founder event, less, outcrops that provide xeric habitats great effect. The larger the ultimate the herd is not depauperate in genetic separated from one another by the pre- captive population size, the more irn- variability relative to other mammals. dominant oak-hickory forest found in the mune it is from the loss of genetic vari- Genetic diversity is best maintained Ozarks. Hence, the natural habitat of ation through the action of genetic drift. when each founder contributes more or these lizards is highly fragmented. Ge- It is thereforeoptimal to make the captive less equally to the population. It is criti- netic surveys have revealed no or very population as large as is practically pos- cal that this equalization be done from the little genetic diversity within a glade but sible. However, there are also other para- onset of the captive breeding program. If extensive differentiation among glades meters affecting the preservation of ge- not, much of the genetic variability pres- - even glades just a few hundred yards netic variation. ent in some of the founders will be lost by apart (Templeton et al. 1990). Hence, Nei et al. (1975) showed that the loss chance alone. Equalizing the founder even a large sample of lizards taken from of genetic variation decreases as the rate contributions in later generations can a single glade subpopulation would miss of population growth after the founder never undo this damage; once the vari- almost all the genetic diversity present in event increases. Consequently, it is best ation of a particular founder is lost, it is this species. The best way to preserve to increase the captive population to its gone forever. The analysis of the Speke's

Vol. 8 No. 1 Endangered Species UPDATE 15 gazelle herd supports this prediction duce the population's response to selec- When a population is brought into captiv- (Templeton et al. 1987). Before the ini- tion. For example, large founding sizes ity, all three of these factors are com- tiation of genetic management, one promote genetic diversity and reduce the monly altered, resulting in adaptation to a founding female (F5)was bred very little chances of genetic transilience - rapid new genetic environment. The best way during the early 1970s. Starting in 1979, adaptive shifts induced by the founder to avoid adaptation to an altered genetic the founder representation was deliber- event (Templeton 1980). Equalizing re- environment is to minimize changes in ately equalized (Templeton and Read productive success in captivity to maxi- the genetic environment under captivity. 1984), but by that time over 50% of the mize the maintenance of genetic diver- To do this we need to know what the genetic diversity carried by F5 would sity also reduces the opportunity for se- genetic environment is. This knowledge have already been lost (MacCleur et al. lection in the captive population. Finally, can be acquired from genetic surveys on 1986). By using the @gee data, population subdivision, the best strategy the natural populations. If genetic sur- Templeton et al. (1987) found that none veys on natural populations are not fea- of the allelic diversity present in the herd sible, inbreeding and outbreeding de- in 1982-1985 traced back to F5 despite a pressions can be distinguished through fairly equal founder representation by pedigree analysis (Templeton and Read that time. This example shows that it is 1984, Templeton et al. 1986), and this critical to equalize founder representation as soon as possible after the estab lishment of captivity. Mistakes made contrary to the usual early in the breeding program cannot be ". . . corrected unless additional sampling advice of no artificial se- from the natural population is possible. lection, it is sometimes Once the herd has grown to its carry- necessary to deliberately ing capacity, genetic drift can still erode manipulate the genetic the genetic variation that has survived to fate of the population to that point. The best way of minimizing the impact of genetic drift in a closed ensure its survival under population is to subdivide the population captivity." into breeding units that have minimal gene flow (Maruyama 1970). We have therefore established several subherds of information can be used to infer the natu- Speke's gazelle at various zoos. Animals ral genetic environment (Templeton are exchanged between subherds only 1987). when they are absolutely needed for If the natural population is subdivided management purposes. In this way, we into inbred demes, one can maintain minimize gene flow between herds and genetic diversity by sampling individuals maximize the global herd's ability to from several local populations. One can retain variation. then minimize alterations in the genetic environment by establishing subdivided PREVENTING EVOLUTIONARY breeding units, with the founders of each CHANGES breeding unit ideally coming from the for long-term maintenance of genetic same local natural population. If the Adaptions to the captive environment variation, tends to result in subpopula- natural population is not subdivided or may imperil the chances for success after tions with little genetic variation within inbred, the management situation is more release. As long as the captive breeding them. Hence, there will be little response difficult. The best way to avoid changing population is genetically variable, it can to selection under this breeding design. the genetic environment is to avoid in- adapt to the captive environment. Guard- Unfortunately, population subdivi- breeding within the captive population. ing against any inadvertent selection for sion frequently changes the genetic envi- This requires using the entire captive domestication can help, but we simply ronment of the species. When a captive population as a single breeding unit. cannot anticipate or monitor all the ways population adapts to its new environ- However, this strategy also maximizes in which a population can adapt to a ment, it is not just the external environ- the long-term loss of genetic diversity in captive environment. Hence, the goals of ment that is important. Because genes the population and maximizes the poten- maintaining genetic diversity and pre- interact with one another, the selective tial of the captive population to adapt to venting adaptation to the captive envi- fate of a gene depends critically upon its the external captive environment. Con- ronment are sometimes in tension and genetic environment (Templeton 1979). sequently, some compromise is neces- complicate each other in practice. The genetic environment in turn is deter- sary. The fundamental decision deter- Nevertheless, some of the strategies mined by the system of mating, popula- mining which strategy to implement for preserving genetic variation also re- tion size, and population subdivision. depends on the expected length of time

16 Endangered Species UPDA TE Vol. 8 No. 1 the population will remain in captivity. depression (Templeton and Read 1984). captive population to evolve away from Basic population genetic theory predicts Since inbreeding could not be avoided, inbreeding or outbreeding depressions that neutral genetic diversity is lost at a we decided to evolutionarily reduce the both in captivity and after release. With- rate of 1/(2N) per generation (Crow and inbreeding depression by a breeding pro- out genetic variability, the options for Kimura 1970), where N is the variance gram, and were successful in achieving both the species and the genetic manager effective size. Using this neutral rate loss this goal (Templeton and Read 1984). are closed. Therefore, genetic manage- as an index, the breeding program should This success illustrates that inbreeding ment must keep as its primary goal the last no more than 0.6(N) generations in depression is not an inherent attribute of maintenance of genetic diversity; all order to preserve at least 75% of the inbreeding, but rather it is a genetically other management policies must be initial genetic diversity carried over into determined response to an altered genetic evaluated in terms of this central policy. captivity. Moreover, by keeping the environment. Hence, as long as the cap- breeding program of short duration, the tive population retains genetic variation, ACKNOWLEDGEMENTS adaptation of the population to its captive it should be capable of evolving a reduc- conditions is also minimized. These tion in the inbreeding depression. This work was supported by NIH grant R01 considerations imply that avoidance of As this example shows, inbreeding GM31571. inbreeding is the better strategy when the depression can be managed. Conse- breeding size is large and when the ex- quently, when necessary, it is possible to LITERATURE CITED pected duration in captivity is short. The implement the inbred, subdivided breed- Baccus, R., N. Ryman, N.M. Smith, C. Reutenvall, subdivided strategy is better when one ing strategy on an outbred species. By and D. Cameron. 1983. Geneticvariabilityand can only maintain a small number of efficiently preserving genetic diversity, differentiation of large grazing mammals. J. breeding individuals or when the antici- it is hoped that any future released popu- Mammal. 64: 109-120. pated breeding program exceeds 0.60 lation will have theevolutionary flexibil- Cmw, J.F., and M. Kimura. 1970. An Introduction toPopulation Genetics Theory. Harper & Row, generations. ity to readapt to its old genetic environ- New York. Unfortunately, practical constraints ment as well as to its restored habitat. To MacCleur, J.W., J.L. Vandeberg, B. Read, and often dictate the breeding strategy much re-establish genetic variation in any fu- O.A. Ryder. 1986. Pedigree analysis by com- more than theoretical considerations. puter simulation. Zoo. Biol. 5:147-160. ture reintroduction, it would be neces- Maruyama, T. 1970. Rate of decrease of genetic For example, in our Speke's gazelle sary to mix animals from several sub- variability in a suMivided population. Biomet- breeding program, our founding popula- herds. It is possible that the different rika 57:299-312. tion consisted of one male and three subherds might have evolved incompat- Nei, M., T. Maruyama, and R. Chakraborty. 1975. females. Once the original founders were The bottleneck effect and genetic variability in ible coadapted complexes during captiv- populations. Evol. 29:l-10. no longer reproducing, all possible ity, which in turn could lead to an out- Templeton, A.R. 1979. The unit of selection in matings in this herd had to be between breeding depression when the animals Drarophila mercatorwn. 11. Genetic revolu- close relatives. Hence, there was no are mixed. To minimize the outbreeding tions and the origin of coadapted genomes in parthenogenetic strains. Genetics 921265- possibility of avoiding strong inbreeding depression, it is best to perform the initial 1282. in this captive population, so we had no mixing in captivity and release the Templeton, A.R. 1980. The theory of speciation choice but to opt for the inbred, subdi- healthy F, or later generation individuals via the founder principle. Genetics 94:1011- vided population strategy. since the brunt of the outbreeding depres- 1038. Templeton, A.R. 1987. Inferences on natural If practical constraints or long-term sion is borne by the F,(Templeton et al. population structure from genetic studies on considerations lead one to implement an 1986). In this manner, the release popu- captive mammalian populations. Pages 257- inbred, subdivided population captive lation does not have to suffer from the 272 in B.D. Chepko-Sade and Z.T. Halpin, ed. breeding strategy on an organism that is selective pressures induced by coadapta- Mammalian Dispersal Patterns. University of Chicago Press, Chicago. largely panmictic and outcrossing in nation during captivity. Templeton, A.R., and B. Read. 1984. Factors ture, then the compromise one has to eliminating inbreeding depression in a captive make is to adapt the captive population to CONCLUSION herd of Speke's gazelle. Zoo Biol. 3:177-199. high levels of inbreeding. During the Templeton, A.R., H. Hemmer, G. Mace, U.S. Seal, W.M. Shields, and D.S. Woodruff. 1986. Local course of this adaptation the population The genetic management of captive adaptation, coadaptation, and population will suffer from an inbreeding depres- populations frequently requires compro- boundaries. Zoo Biol. 5:115-125. sion. Such inbreeding depressions can be mises between preserving genetic diver- Templeton, A.R., S.K. Davis, and B. Read. 1987. so severe that they may endanger the sity, altering the original genetic compo- Genetic variability in a captive herd of Speke's gazelle (Gazelfuspekeo. Zoo Biol. 6:305-313. success of the entire program. Hence, sition as little as possible, and practical Templeton, A.R., K. Shaw, E. Routman, and S.K. contrary eb the usual advice of no artifi- management constraints. Whenever Davis. 1990. The genetic consequences of cial selection, it is sometimes necessary such compromises need to be made, the habitat fragmentation. AM. Mo. Bot. Garden to deliberately manipulate the genetic first priority should be given to the pres- 77:13-27. fate of the population to ensure its sur- ervation of genetic diversity. Only if the vival under captivity. This was the situ- captive population has genetic diversity ation we encountered with Speke's ga- does it have the adaptive flexibility that it Alan R. Templeton is Professor of Biology and Genetics in the Department of Biology at Washing- zelle, whose captive population devel- needs to cope with an uncertain future. ton University, St. Louis, MO 63130. oped an extremely severe inbreeding This same adaptive flexibility allows the

Vol. 8 No. 1 Endangered Species UPDATE 17 Decision-Making About A Reintroduction: Do Appropriate Conditions Exist? by Devra G. Kleiman

The following example provides a alteration. There are captive populations in these forms historically. There is still framework for making decisions about of each species, but at different levels of significant ongoing deforestation within whether the appropriate conditions exist development. the ranges of L.chrysopygus and to recommend (or argue against) reintro- Table 2 lists ten necessary conditions chrysomelas, thus dictating against a ductions of captive-born animals or which should be met in order to recom- reintroduction at this time. It is question- translocations of wild individuals or mend a reintroduction/translocationpro- able whether the reasons for the decline groups. Recent reviews provide more gram. Additionally, it evaluates theposi- of rosalia are now fully under control. A reintroduction is not recommended without having protected habitat avail- Table 1, The ekiztmr of a swcassfui reintmduaioaprogram involving captive-bred mpmmals. able, It is likely that sufficient protected Reprinted from Klb(19891, with pemzissiori from BiaStknce 39(3):154.8 1989 American hstimte of Biological Sciences.] habitat is available for chrysopygus, but not chrysomelas. Protected habitat exists Captive population for rosalia, although in insufficient quan- Ongoing research in behavior, genetics, physioIogy, nutrition, rqmduction, and pathology tities for this species' future survival. Genetic and danqraphic management of the population Self-sustainingviable captive Mation It is preferable to reintroduce individuals or groups into unoccupied habitat Fletd Studies to prevent social disruption and disease Regular cursasea of the size, disuibutim, and genetics of hewild popularim transmission between wild and intro- Behavioral ecology studies [home range size, movements, habitat prefermas, social organizations, mating system, feeding, wd anti-predator adaptims) duced individuals. Thus, areas with Zocating existing suitable habitat containing critical resources for reintroduction small or no populations of wild tamarins need to be available. This condition Habitat I)rwrvation and management exists for rosalia, and probably for Protection of habitat from deforestation, degradation, and exploitation Rc~toMand ~~~anag~erttof degraded hahitats chrysomelas. However, the situation for hcmse ot mrfintmacc of the mmber of preservation areas chrysopygus is unknown at this time. Similarly, reintroductions should be Conservation education lor long-term support Professional uainjng tbmugh academic studies, workshops, internships, courses, and fellowships encouraged only when there is some Detedngthe; most appropriate public relations and educational strategies through sorveys certainty that the release of animals from Public rehtians educational &om using appropriate mass media (e.g., tdwkion, radio, different regions (both captive- and wild- magazines, and newspapers) born) will not jeopardize the existing LocaI both canmunity education, formal and infd native population through the transmis- Preparation an@reIntrgd4Ww d urimals sion of disease and/or social disruption. Goice of Earldidares and rrssessment of their characters for retrospective correlation with We do not have this confidence for any of pstdease sutvival the three forms of lion tamarins at this Training in swivdtechniques, including foraging and feeding, antipredator tactics, locomotion, and oti&on time, due to our limited knowledge of Adaption to Id wndions at release site {food, cIimate and temperatun, and disease) their biology and status. Release and longtern monitoring to evaluate causes of death and basis for survival The evaluation of the success of a reintroduction can only be accomplished by long-term monitoring, and must be detailed background (Griffith et al. 1989, tion of each form of lion tamarin with based on a thorough knowledge of a Kleiman 1989, Stanley-Price 1989); respect to each condition. Finally, a species' biology, distribution, and eco- components of a good reintroduction general recommendation is presented logical requirements. On a scale of one to program, as discussed in Kleiman concerning whether aprogram of reintro- five, with five being the best case sce- (1989), are presented in Table 1. duction is appropriate for each form at nario, I suggest that there is sufficient in- The lion tamarins (genus Leontopith- this time. formation available for rosalia, totally ecus) derive from the Atlantic coastal A reintroduction is unwarranted un- insufficient information available for rainforests of Brazil. All three species - less the causes for the initial reduction in chrysomelas, with chrysopygus some- L. rosalia (golden lion tamarin), L. species numbers have been removed. where in between. chrysomelas (golden-headed lion The major reasons for the decline of the Local support is essential to the suc- tamarin), and L. chrysopygus (golden- lion tamarins has been deforestation of cess of a reintroduction. A conservation maned lion tamarin) - are endangered, the Atlantic coastal rainforest, although education program in conjunction with a mainly due to habitat destruction and there has also been a thriving commerce reintroduction will attract and inform the

18 Endangered Species UPDA TE Vol. 8 No. l conservation programs for chrysomelas i5I reproductive groups, I suggest and chrysopygus are not yet sufficiently that we still con- developed, with respect to financial sup- sider reintroduc- port and the necessary infrastructure, to tion an experi- warrant a reintroduction effort. On the mental approach other hand, the rosalia program has a for this taxon. well-developed infrastructure and con- The success siderable resources to monitor the activi- of a reintroduc- ties of released animals. tion program can One major goal of a reintroduction only be evalu- program is to augment the numbers or ated through a genetic diversity of a population; rosalia comparison of currently needs such augmentation, the behavior and while chrysomelas does not. The situ- survivorship of ation for chrysopygus is not clear at this reintroduced vs. time. wild-born ani- Weighing the degree to which these mals. Thus, ac- necessary conditions are met for each local populace, and result in greater cess to the resources necessary to moni- species suggests that while reiniroduc- community support for the effort. Both tor theactivities of releasedanimals ises- tion efforts may be appropriate for rosalia and chrysopygus conservation sential for a reintroduction effort, espe- rosalia, they are not for chrysomelas or programs have strong educational com- cially since we have not yet perfected our chrysopygur at this time. Given the limi- ponents, while the education program for preparation and release techniques. The tations of personnel, money, time and re- chrysomelas is just sources for conserva- developing. tion activities, along A prerequisite to the re- with the limited num- introduction of animals ber of surplus individu- currently in captivity als available in popula- (whether captive- or wild- tions of imperiled spe- born) is a secure well-man- cies, it is imperative aged captive population, that the need for and with a long-term master- likely success of a plan and available surplus reintroduction be thor- animals. A reintroduction oughly evaluated be- should not jeopardize the fore it is attempted. genetic or demographic composition of the captive Literature Cited population. This condition Griffith, B.. J.M. Scott, J.W. is met in rosalia, but Carpenter, and C. Reed. not in chrysomelas and 1989. Translocation as a chrysopygus. An excep- species conservation tool: tion are some wild-born Status and strategy. Sci- ence 245477-480. confiscated specimens of Kleiman, D.G. 1989. Rein- chrysomelas in captivity in troduction of captive the Una Biological Re- mammals for conserva- serve in Bahia that cannot tion. BioScience 39:152- 161. be easily absorbed into the Stanley-Price, M.R. 1989. captive population. Animal Reintroductions: We still have much to The Arabian Oryx in learn about the methodolo- Oman. Cambridge Uni- gies of preparation, adap- versity Press, New York. tion, and release of lion tamarins. With so many Devra G. Kleiman is the unanswered questions Assistant Director for about the techniques that Research at the National will ensure success, e.g., Zoological Park, Smith- sonian Institution, Washing- for the injection of single ton, DC 20008. animals into established

Vol. 8 No. 1 Endangered Species UPDATE 19 Is Captive Breeding an Appropriate Strategy for Endangered Species Conservation? by Tony Povilitis

Is expanded use of captive breeding conservation and other safeguards for cies are isolated from natural communi- for species preservation in the best inter- species within nature. ties, coevolution of predators and prey, est of biological conservation? The an- In this paper, I argue that the lirnita- pollinators and flowers, biological mim- swer to this question depends on one's tions of captive breeding and other re- ics and models, etc., is curtailed. When view as to whether humans can change lated considerations severely undermine wild animals are denied the opportunity their environmentally destructive ways, the "last ark" strategy. However, captive to choose and compete for mates, sexual on how conservation is defined, and on a breeding if combined with far more selection ends. Moreover, for adaptive host of ethical and related concerns. comprehensive efforts to protect and evolution to occur, relatively large gene By and large, captive breeding propo- restore habitat can be of substantial pools, generally on the order of thou- nents tend to adhere to one of the follow- benefit in species conservation. sands of individuals in the case of verte- ing two views regarding the role of cap- brates (Soule 1987), should be subjected tivebreeding in conservation. According Biological Limitations to wild selection. to the "last ark" perspective, human Important behavioral changes in cap- behavior will not change and severe "Last ark" conservation might pre- tivity have been noted even over the short habitat collapse is inevitable (Foose serve some 1,000taxa of terrestrial verte- term, as in the case of altered foraging 1986, Cade 1988). Captive maintenance brates out of some 1,500 taxa estimated behavior in recently released Mississippi is thus needed to save as many species to become so endangered by the middle sandhill cranes (Zwank et al. 1988). as possible, with the hope that their re- of the next century that captive propaga- Special pre-release training is being developed for arboreal golden lion tamarins (Kleiman 1989), to improve ". . .the 'last ark' [captive maintenance of species] will locomotor and foraging skil1sbeforere- not be able to accommodate most endangered spe- lease into the wild. Because of uncer- tainty about the ability of captive-bred cies. It cannot foster the ecologically meaningful evo- animals to swive in some cao- lution of species. Nor has it room for higher elements tive have begun propagating of natural diversity, such as larger natural communi- animals in limited wild environments, ties and landscape ecosystems." such as the propagation of free-roaming red wolves on islands for later release in mainland reintroduction programs (Rees lease back into the environment might tion will be essential for their survival 1989). be possible when the human population (Foose 1986). However, little help is In sum, the "last ark will not be able stabilizes or declines - perhaps within expected for the vast majority of inverte- to accommodate most endangered spe- the next 200 to 1,000 years (Foose brates and plants which comprise most of cies. It cannot foster the ecologically 1986). As Conway (1988) pointed out, the world's vanishing biota. These taxa, meaningful evolution of species. Nor "it seems inevitable that most large which are among the most basic compo- has it room for higher elements of natural land vertebrates and many plants eventu- nents of ecosystems, nevertheless domi- diversity, such as larger natural commu- ally will survive only as wards of hu- nate the 4,000 to 6,000 species per year nities and landscape ecosystems. Fi- mans." conservatively estimated to be lost over nally, captive breeding may preclude the The alternative view regards captive the next decade from tropical deforesta- eventual restoration of some species to breeding as only a "hedge against tion alone (Wilson 1987, 1989). the wild by inadvertently promoting the extinction," an interim measure needed Even endangered species success- selection of traits suited to captivity. to retain species (Seal 1988), while fac- fully maintained in captivity face a pre- tors which threaten them (e.g., habitat carious future. In terms of the selective Drawbacks for Wild Populations destruction, hunting, pollution) are forces that shape a species,captive condi- brought under control. If habitat is lost tions cannot simulate wild environments. In some instances, endangered spe- entirely there is little point to species When species are removed from their cies in the wild may be harmed by in- preservation (Ehrlich and Ehrlich 1981). habitats, natural selection ends, and se- creased focus on captive breeding. Thus, the role of captive breeding falls lection for the captive environment be- Removal of wild individuals to establish squarely within the context of habitat gins (Lyles and May 1987). When spe- or stock captive populations may jeop-

20 Endangered Species UPDATE Vol. 8 No. 1 ardize the viability of seriously endangered species. For example, the recently developed Species Survival Plan for the Florida panther calls for removing ten individuals in 1990,and eight for at least the following two years (Seal and Lacey 1989), hma wild population currently numbering only an estimated 30 to 50 animals (Jordan 1990). Risks of losing individuals of more sensitive species during capture or in captivity, or by program-stimulated illegal removals could further stress some wild populations. Groups of endangered huemul (Andean deer), for example, are known by the author to have been repeatedly lost during capture attempts in the wild and through infectious diseases in captivity, with uncertain consequences for donor populations. Use of private populations to supply animals for formal breeding programs, such as that for the thickbilled parrot (Johnson et al. 1989) and angono- Maintaining 1,000 vertebrate taxa tat conditions deteriorate further, conser- lia (Burke 1990), could unwittingly would requue.maximal efficiency in use vationists face the specter of having to stimulateadditionaltaking from the wild. of zoo facilities (Foose 1986), or virtu- protect their "investments" through still Finally, wild populations of endan- ally all zoo spaces (Seal 1988). Given more intensive manipulation and care of gered species and other wildlife could be these limitations, Seal (1988) has both captive and released animals. The at risk from released captive-bred ani- characterized a management of high costs involved could make the mals not adequately screened for dis- endangered species as "a temporary whole endeavor of captive breeding and eases, and from maladaptive genes or haven for. . . those species that happen reintroduction prohibitive. behaviors acquired in captivity. Disease to gain a seat." Ultimately, a mas- outbreaks in wild desert tortoises, for Endangered sive "last ark" to Dre- example, may have begun with the re- species will inev- 11 . . . short-term succes- serve species lease of infected captive animals (U.S. itably be in direct depend on sustained Fish and Wildlife Service 1989). Also of competition for ses at captive propaga- socio~onomic ~ta- concern are disease outbreaks in wildlife limited zoo fiOn provide false reas- bilitv over centuries. originating from live vaccines used to space. Will pri- surance that the endan- ~~t 'such stability ap- protect released captive stock. ority be given to gered species problem is pears most unlikely species on the Solvable through Logistical and Financial basis of biologi- SitY structionwith and desta-de- Constraints cal needs or will measures=" bilization of the other factors, world's ecosystems. Some endangered vertebrates may be such as display lost from the "ark" because of breeding value and other institutional needs, pre- Political and Social failures or disease vulnerability in cap- vail? Will zoos vie for the most "charis- Consequences tivity. Endangered pmts, for example, matic" endangered species? As of 1986, are susceptible to both problems (Der- for example, there were at least 700 cap- Captive breeding as a strategy for pre- rickson and Snyder in press). Still other tive Prezwalski's horses, with the num- serving endangered species threatens to species will be excluded from captive ber growing by perhaps 100 individuals undermine habitat protection as the cen- programs for purely logistical and finan- per year (Smollar 1986). Moreover, tral focus of conservation. Promotion of cial reasons. large scale captive breeding programs, captive breeding politically, for example, Maintaining large numbers of endan- such as the controversial program cur- could misdirect public policies away gered vertebrates in captivity will be rently envisioned for the Florida panther from habitat conservation and divert very costly over the long term. Foose (Seal and Lacy 1989) could quickly fill funds that might otherwise be used for $J (1986), for example, estimated that some zoo spaces, leaving other endangered &JJ endangered species programs. At a $1,500 per individual animal per year is species in the lurch. recent hearing on legislation concerning needed for conventional breeding pro- As captive species become ever more the captive breeding of sea turtles, Repre- grams, exclusive of research needs. dependent on biotechnology and as habi- sentative WJ. "Billy" Tauzin (D-LA),

Vol. 8 No. 1 Endangered Species UPDATE 21 the bill's chief sponsor, indicated that the As Cade (1988) noted, "most human rect animal abuse. Wild animals are stocking of wild populations with cap beings continued to be captivated more sometimes held in overly restrictive tive-reared animals could mitigate losses by what engineers do than by what quarters, subjected to harassment while due to economic activities, as for in- ecologists say." Indeed, short-term suc- on public display, deprived of minimal stance, in the case of logging impacts on cesses at captive propagation provide sensory stimuli, sold as surplus animals the northern spotted owl. In essence, false reassurance that the endangered to unscrupulous dealers, or otherwise preservation by such means would re- species problem is solvable through g treated inhumanely (Fox 1990). Captive place conservation of self-sustaining & measures. breeding too often has more to do with populations. Will the impressionable public see justifying the existence of zoos than with Captive breeding already appears to endangered species in zoos merely as preserving endangered species (Grandy have undermined habitat conservation. curious rare objects or as living beings 1989). In the case of the Gila topminnow, for having intrinsic worth? Will it perceive Captive propagation of endangered example, stocking of artificially main- captive breeding technology as solely a species also subjects animals to undue tained habitats with captive-bred fish has tribute to human ingenuity, or as a moral stress and mortality when used to stock resulted in the reclassification of the obligation to temporarily offset the im- wild populations that are jeopardized species from endangered to threatened, pact of society's misguided destruction because of poor habitat conditions or despite continued habitat deterioration of habitat? adverse human activities (e.g., release of (Simons et al. 1989). Other species, like sea turtles where they are likely to be Attwater's greater prairie chicken (U.S. Ethics drowned by shrimp nets). Fish and Wildlife Service 1988) and From a biocentric view, captive several endangered river fishes (Hunt Captive breeding and manipulation of breeding of endangered species is ethi- 1989), continue to decline because of endangered wildlife raise important ethi- cally acceptable only if the animals or habitat problems while recovery pro- cal questions for both conservationists their descendants are released into rea- grams emphasize captive breeding. A and for society as a whole. For example, sonably secure habitats. But what if recovery program for the Florida panther captivity denies wild animals the oppor- conditions continue to deteriorate and there are no certain plans to reverse the situation? Is it ethical to support captive breeding when, in deference to purely

". . . captive breeding too often represents the path of least political resistance for government agencies -- a detour around contro- versies involving land and water use."

human (viz. economic) interests, society fails to take a firm and unequivocal stand in protecting critical habitats? Many people find value in free- proposes a captive population of some tunity to experience those habitats for living and unmanipulated wildlife. Yet 350 individuals which would count to- which they are, from evolutionary and conservation efforts that depend largely ward any future "recovered" population ecological standpoints, most suited; lost on captive maintenance risk "preserv- (Seal and Lacy 1989). Yet current recov- is the opportunity for them to engage in a ing" wildlife by taming it. The result ery efforts for the panther lack adequate range of behaviors in response to diverse could be eventual domestication for the habitat evaluations and conservation natural stimuli (e.g., foraging on a varied more "useful" captive species, and plans, while the wild population of 30 to diet, play activities, mate selection, flex- extinction of the others. In any event, 50animals continues to decline. In short, ible daily routine, etc.). Indeed, habitat captive breeding on a large scale may captive breeding too often represents the has value for individual animals as well well destroy our sense of the beauty, path of least political resistance for gov- as for species. mystery, spirituality, and dignity of spe- ernment agencies -- a detour around con- Captive maintenance, usually for rea- cies, and of what remains of the natural troversies involving land and water use. sons of economy, may also result in di- world.

22 Endangered Species UPDATE Vol. 8 No. 1 Consewation Ethos inn- nature. But so must advances in of captive breeding in conservation strategies. transportation and energy use systems, in Paga 141-165 in L. ICaufman and D. Mdor~, eds. The Last Extinction. The MIT Press, A basic tenet of conservation is to telecommunications, recycling technol- hbridge, MA. maintain in a safe and sound state the om, environmental education, birth con- Fox. M.W. 1990. Inhumane Societv, st.Martin's ecosystems upon which species depend, t2,and family, urban, and land use plan- Ass, New York. ning. After all, species preservation is Grandy, J.W. 1989. Captive breeding in ms. and to preserve basic ecological and evo- Humane Society News 34(3):8-11. lutionary processes. This tenet is central only one reason for preserving habitat. Hunt, C.E. 1989. Creating an endangered to the U.S. Endangered Species Act, and Others include a healthful and aestheti- ecosystems act. Endangered Species UP- perhaps has been best articulated in cally wholesome environment for DATE 6(3-4):1-5. people, and reduced risk of environ- Johnson, T.B., N.F.R. Snyder, and H.A. Snyder. Leopold's land ethic (Leopold 1949). 1989. The return of thick-billed pmts to The danger is in viewing captive mental catastrophe, such as that poten- Arizona. Endangered Species Technical Bul- breeding and related technologies as an tially caused by global warming. The letin 14(4):1,4-5. alternative strategy for conserving spe- broadest task is to integrate conservation Jordan, D. 1990. Mercury contamination: An- Florida cies. Conservation in captivity as op- values and techniques with economic other threat to the panther, Endangered Species Technical Bulletin 15(2):1,6. , posed to that in nature are two very dis- planning and development. Kleiman, D.G. 1989. Reintroduction of captive tinct things. In- mammals for amsewatim. BioScience 39: deed, the term 152-161. "conservation" Leopold, A. 1949. A Sand County Almanac. "To regard captive maintenance of a Ballantine Books, New York. as applied to a Lyles, M.L, and R.M. May. 1987. Problems in captivebreeding species as conservation is to commit the leaving the ark. Nature 326:245-246. strategy seemi conceptual error of seeing that species Noss, R.F. 1983. A regional landscape approach to maintaining diversity. BioScience wholly unwar- as somehow separate from its habitat." 33(11):700-706. ranted, consider- Povilitis, T. 1990. Where will wildlife live? ing the tiny frac- Humane Society News 35(2):8-13. tion of species Rees, M.D. 1989. Red wolf recovery effort inten- sifies. Endangered Species Technical Bulletin that can be protected through such ef- An appropriate goal for captive breed- 14(1-2):3. forts, the problem of adverse impacts of ing is to help preserve endangered spe- Seal, U.S. 1988. Intensive technology in the care captivity on behavior and gene selection, cies as part of comprehensiveplanning to of ex A populations of vanishing species. Pages 289-295 in E.O. Wilson, ed. Biodiver- and the fact that nature consists of far protect and restore their habitats. There sity. National Academy Press, Washington, more than species. To regard captive is scarce room for compromise. The DC. maintenance of a species as conservation focus must be on the expeditious return of sG, u.s., and R.C. Lacy. 1989. Florida Panther is to commit the conceptual error of see- endangered species ti the wild, so that Viability Andysis U.S. Fish & Wildlife Service, Gainesville, FL. ing that species as somehow separate adverse biological, social, political, and Simons, LH,, D.A. Hendrickson, and D. Pa- from its habitat. ethical consequences of captive breeding pUlias. 1989. Recovery of the Gila topmin- Confronted with a severe habitat cri- can be avoided. now: A success story? Cons. Biol. 3(1):11-15. sis, conservationists need not retreat to a Srnollar. D. 1986. ~et&to the wild. &&neered species Technical Bulletin Reprint 3(10):1-2. "last ark" ethos. But they should recog- Acknowledgements Soule, M.E. 1987. Where do we go from here? nize that the "preserve what we can" Pages 175-183 in M. Soule, ed. -viable Popu- approach to conservation has failed. I thank M. Fox, J. Grady, L. Landres, and T. lations for Conservation. Cambridge Univer- More explicit goals are needed. One goal Telecky for their kind review of an earlier draft of sity Press, Cambridge. this article. U.S. Fish & Wildlife Service. 1988. Regional might be to identify and conserve on a News. Endangered Species Technical Bulletin regional basis habitats essential to endan- Literature Cited 13(11-12):3. gered and declining wildlife populations, U.S. Fish & Wildlife Service. 1989. Emergency threatened natural community types, and Burke, R.L. 1990. Conservation of the world's Action Taken to Protect the Desert ~o&ise. Endangered Species Technical Bulletin 14(9- large landscapes that can still sdppon rarest tortoise. Cons. Biol. 4(2):122-124. Cade, T.J. 1988. Using science and technology to 10): 1,5-6. most native species (Noss 1983). An- reestablish species lost in nature. Pages 279- Wilson, E.O. 1987. The liule things that run the other might be the application of a no- 288 in E.O. Wilson, ed. Biodiversity. National world (the imponance and conservation of in- net-loss principle to all sensitive wildlife Academy Press, Washington, DC. vertebrates). Cons. Biol. 1(4):344-346. Conway, W. 1988. Can technology aid species Wilson. E.O. 1989. Threats to biodiversity. Sci- habitats (Povilitis 1990). Still another entific American 261 (3): 108-116. preservation? Pages 263-268 in E.O. Wilson, goal might be to limit our own numbers, ed. Biodiversity. National Academy Press, Zwank, P.J., J.P. Geaghan, and D.A. Dewhurst. for without limits there can be no politi- Washington, DC. 1988. Foraging differences between native and cally feasible limit to human use of land Demckson, S.R., and N.F.R. Snyder. In press. released Mississippi sandhill cranes: Implica- Potentials and limits of captive breeding in tions for conservation. Cons. Biol. 2(4):386- and water. Courageous and determined em parrot conservation. In S.R. ~eissin~er-and 'n~' conservation leadership will be needed N.F.R. Snyder, eds. New World Parrots in -despite institutional pressures and the conservation: Solutions fmm Captive Breed- Tony PoGlitis is Senior Scientist for Wddlife and ing. Smithsonian Institution Press. risk to jobs. Habitat Protectim with The Humane Society of the Ehrlich,P.R.,andA.H.Ehrlich. 1981. Extinction. United States, 21M1 NW, Captive breeding can play a key role Ballantine Books, New York. DC 20037. in any habitat-based strategy for preserv- Foose, T.J. 1986. Riders of the last ark: The role

Vol. 8 No. 1 Endangered Species UPDATE 23 Measures of the Value and Success of a Reintroduction Project: Red Wolf Reintroduction in Alligator River National Wildlife Refuge by Michael K. Phillips

Reintroduction is an important tech- away from the historic belief that wolves The red wolf is believed to have nique for recovering endangered and are a serious and consistent threat to evolved solely inNorth America (Nowak threatened species (Grifftth et al. 1989). human safety and a competitor with 1979). Thus, the species is an important Unfortunately, the technique is complex hunters for game. Since the first red part of the history and heritage of the and costly (Clark and Harvey United States. However, prior to 1988), and there are few if any reintroduction at ARNWR most accepted guidelines for defining U.S. citizens knew very little its value and success. Although about red wolves. The opportu- these concepts will be defined in nity to learn about free-ranging part on a species by species basis, red wolves might have been for- the task of identifying the vari- ever lost if not for the ARNWR ous values and successes of a reintroduction project. reintroduction project has im- The reintroduction project al- portant ramifications for deter- 1owedtheUSFWSto develop the mining the potential merit and red wolf into an effective "flag- effectiveness of reintroduction ship" species for conservation. programs in general. Quite honestly, the task was easy Since 1973, the endangered because wolves evoke strong red wolf (Canis rufus) has been emotions in people. Regardless the focus of a federal recovery of whether people are for or program (Carley 1975,U.S. Fish against wolf reintroduction, most and Wildlife Service 1984, are interested in the project. Parker 1988) (see case history). Thus, since 1986, the USFWS In 1987, the U.S. Fish and Wild- has been able to use the ARNWR life Service (USFWS) inten- red wolf project as a vehicle to sified recovery efforts by initiat- present information not only ing a reintroduction project at the about wolf restoration, but also Alligator River National Wild- about the plight of other endan- life Refuge (ARNWR) in north- gered species and environmental eastern North Carolina (Smith issues. and Phillips 1987, Phillips 1988, The red wolf reintroduction Phillips and Parker 1988, Meese program also portends the future 1989, Phillips 1990, Parker and for many species. As humankind Phillips in press). In this paper I continues to modify the land- discuss some interim measures scape, animal and plant species of the value and success of this will be squeezed into smaller and ongoing project. "Intensive management programs, smaller islands of suitable habi- like the one developed for red tat. Intensive management Pro- Measures of Value wolves in the refuge, will be neces- grams, like the one developed for red wolves in the refuge, will be The reintroduction of red sary if many endangered species are n,essq if many endangered wolves into ARNWR was ac- t~ persist and evolve." species are to persist and evolve. complished only after the The ARNWR project provides USFWS carried out an education conservationists with the oppor- program that prompted many people to wolves were released, the project has tunity to study and begin to perfect the change their attitudes toward other spe- been offered as badly needed proof process of ecological restoration. cies, even so-called "varmits" like that wilderness species and humans can The annual budget for the ARNWR wolves. The change represented a shift coexist. reintroduction project is about $160,000.

24 Endangered Species UPDATE Vol. 8 No. 1 Since most of this money is spent in perience is one of the most striking ex- ure the project's progress. Pups were northeastern North Carolina, the project amples of the project's value. born to free-ranging wolves during provides monetary benefits to citizens of In addition to stimulating individual spring 1988. In fact, four pairs of wolves this area However, in addition to gener- action, the reintroduction project producedlitters,ofwhich pups fromeach ating direct monetary benefits, the rein- prompted civic groups and private com- are still alive (Phillips 1989). Producing troduction has generated a great deal of panies to become involved with conser- and raising offspring in the wild is irrefu- free publicity for table evidence that red Dare County (where wolves can make the ARNWR is located). transition from de- Since 1986, a mini- "Producing and raising offspring in the wild is pendency on humans mum of 22 maga- irrefutable evidence that red wolves can make the for food to self-suffi- zinesand24newspa- transition from dependency on humans for food to ciency. However9the pen published sto- presence of wild-born ries about the proj- self-sufficiency. However, the presence of wild- wolves is just one ect. Regioml news- born wolves is just one component of a success- component of a sue- papers repeatedly fuI [reintroduction] program." cessful program. covered the project. Another measure The project was dis- of success is the bio- cussed during the logical information nightly newscasts of five. national and vation. For example, the North Banks gained through associated research and four regional television networks; Rotary Club (Kill Devil Hills,NC) devel- monitoring of the project. The backbone WVEC and WTKR, both based in Nor- oped a "conservation internship proof the project consists of radio-tracking folk, VA, and both with access to very gram" that consisted of a weekly stipend released individuals to monitor the re- large markets, repeatedly covered the of $50 to $100 provided to individuals sults of reintroduction efforts. To date, project. Three mini-documentaries were volunteering for the project. The pro- over 4,000 relocations have been reproduced, including one by the Austra- gram made a significant contribution to corded (U.S. Fish and Wildlife Service lian Broadcasting Company, and the red red wolf restoration, shielded the volun- unpubl. data), providing information wolf was featured in the nationally teer program from the vagaries of federal about red wolf home range characteris- broadcast "World of Audubon" docu- funding, and provided opportunities to tics, food habitats, activity patterns, mentary about restoration. Additionally, people interested in conservation. sociality, reproduction, and mortality. local radio stations presented informa- The red wolf reintroduction project This information is available to inter- tion about thereintroduction. All stories also prompted the Conservation Fund ested individuals simply by contacting about the ARNWR red wolf project de- (Washington, DC) to acquire approxi- the refuge office. This increase in picted Dare County as an area that has es- mately 47,000 ha of coastal plain habitat knowledge about red wolves and its ac- caped the trappings of the 20th century westof-. Theacquisitionwillbe cessibility to interested members of the and whose natural resources are still managed as a conservation area by the public is another indicator of the healthy and thriving - the same image USFWS and the North Carolina Wildlife program's success. which local businesses and politicians Resources Commission. This additional Although the fust three years of the portray in their advertising. Thus, there- acreage secures critical habitat for count- project presented some very difficult introduction project also indirectly bene- less wildlife species in addition to red management situations, the monitoring fits the local economy, since the fiscal wolves, and provides significant protec- program allowed the USFWS to stay health of the County depends almost tion to the Alligator River watershed and only a few steps behind the problems. solely on tourism. associated Albermarle-Pamlico estuar- Since the first wolf was released, 17 Media coverage effectively informed ine system. animals had to be recaptured on 26 millions of people about the reintroduc- occasions. In spite of our preparedness, tion project. Thirty-threepeoplebecame Evidence of Success recaptures took place under conditions so committed to the concept of restora- that were usually less than ideal. None- tion that they donated approximately Since the ARNWR red wolf reintro- theless, recaptures were carried out 10,000 ha to the project. In addition, duction was a fust, there was no accepted without inflicting significant long-term many volunteers were utilized, most of definition of success against which to damage to animals and with little whom were recent college graduates or compare the project's progress. The inconvenience to residents of the area. students completing degrees in wildlife technical proposal developed for the Successfully managing the wolves management. All vol~ntee~~received project defined successas the presence of helped to convince USFWS officials extensive training in red wolf restoration second generation wild-born pups in the and local citizens that wolf restoration and eventually made significant contri- refuge (Parker 1986) - a definition can be canied out in a controlled manner. butions. The tremendous opportunities developed mostly to provide USFWS This "track record" developed at afforded young biologists in need of ex- officials a yardstick with which to meas- ARNWR will be a tremendous aid to the

Vol. 8 No. 1 Endangered Species UPDATE 25 USFWS as it prepares and implements munities,. lamer- conservation efforts. and Parker. W.T. 1986. A Technical Proposal to Re- wolf restoration p&puns elseurhere. other imperiled species as well. ~eteii- esuush Ibe in the AU;gatOrRiver On the other hand, 15 of the 29 re- National Wildlife Refuge, North Carolina. U.S. nation of the wide range of possible Fir a %,,,ice, Asheville, NC 20 pp, leased wolves died during the first three benefits and successes of a project may Parker, W.T. 1988. The red wolf. Pages 596-607 years of the project. To some, 50% be useful in weighing the desirability of in WJ. Chandler, ed. Audubon wildlife Report mortality is unacceptable and evidence other captive breeding and reintroduc- pa~~$9~~~$~~;~~~;IF;::"A',"~;: thattheprogram isafailure. TheUSFWS tion initiatives in the future. cation of the experimental designation to recov- feels, however, that 15 deaths are not ery of endangered red wolves. Wildl. Soc. Bull. excessive. In fact, the mps,M.K. 1988. Progress USWS believes it is a of the red wolf restoration project in North Carolina. measure of the program's Pages 426-433 in R.O. Wag- success that all deaths ner, ed. Proc. of Annual were natural or acciden- "The values and successes of reintroduc- Meeting of the American tal, and not the result ofa fi0n projects often have the potential to Association Zoological Parks and Aquaria. Oglebay citizen acting irresponsi- extend beyond the immediate preservation Park, Wheeling, WV. blyoronsomeudounded of the reintroduced species, to positively Phiups. M.K 1989 Born in hatred for wolves. affect local citizens and communities, larger the wild. Wildlife in North Assessing the value Carolina 53:24-25. conservation efforts, and other imperiled mapsl M.K. 199a Media and success of an endan- and public involvement in gered species reintroduc- species as well." red WOE restoration. Pages tion program is not an 85-98 in B. Holaday, ed. Proceedings Arizona Wolf easy task. For the Symposium, March 23-24, ARNWR red wolf proj- 1990, Tempe, AZ ect, measures of the value and success of Literature Cited Phillips, M.K., and W.T. Parker. 1988. Red wolf reintroductionare varied* Some are recovery: A progress report. Cons. Biol. 2: 139- Carley, C.J. 1975. Adivities and fmdings of the 141. obvious and easily defined quantita- wolf recovery program from late 1973 to Smith , R.s., M.K. millips. 1987. Captive tively, whereas others are subtle and not July 1, 1975. U.S. Fish and Wildlife Service, breeding and ~introductionof red wolves in the Albuquerque, NM. 215 pp. wilds of North Carolina. Pages 82-90 in R.O. in monetary terms' Clark,T.W., and A.H. Harvey. 1988. Implement- less, each measure of value and success wagner, ed, pra, of ~~~~~l ~~~h~of the hg species recovery policy: American Association Zoological Parks and provides justification and evidenceof the ing as we go. End. Species UPDATE 535-42. Aquaria. Oglebay Park, Wheeling, WV. manageability of this landmark reStOra- Grifflth*BhI J.M. Scott, J.W. Carpenter, and C. U.S. Fish and Wildlife Service. 1984. Red wolf tion project upon which the very exis- Reed. lg89. Translocation as a cmser- recovery plan. U.S. Fish and Wildlife Service. vation tool: Status and strategy. Science ~~l~~~~,GA, 37 pp. tence of a species may depend. 245:477-480. The values and successes of reintr0- Nowak, R.M. 1979. North American quartemary duction projects often have the potential Can&. Monograph 6, Museum Natural History, Michael K, Phillips is a Biologist for the Red Wolf to University of Kansas. 154 pp. Recovery Project at the Alligator River National beyond the immediate preser- Meew, G.M. 1989. Intensified effom for red Wadlife Refuge, USws, Po Box 1969, Manta, ~ationof the reintroduced species, to wolves are paying off. End. Species UPDATE NC 27954. positively affect local citizens and com- 6:26-27.

26 Endangered Species UPDATE Vol. 8 No. 1 Ethical Perspectives on Captive Breeding: Is It For the Birds? by Gary E. Varner and Martha C. Monroe

Introduction rhetoric of environmental politics.] A third ethical perspective fre- Someone who argues that endangered quently is called the "animal rights" On the surface of it, what could be species shouldbe preserved because they position, but this label is misleading. So- more ethical than struggling to preserve contribute to the welfare of present and called "animal rights advocates" usually an endangered species through a captive future generations of human beings do not defend the rights of all animals, but breeding program? The dedication of the would evaluate programs from an only "higher" animals like mammals and many individuals who have sacrificed to aa t hropocen trb establish captive propagation facilities perspective. Phi- across the country, and the thousands of losopher John volunteers who help make them work, is Passmore (1974) ". . . today's captive breeding pro- undoubtedly laudable. However, a adopted this per- grams can only be viewed in a strongly closer look at the various ethical justifi- sp&tive in an's positive light from a very narrow an- cations which might be offered for cap for thropocentric perspective, a perspective breeding reveals a more complex Nature, the first tive which matches neither the rhetoric picture. book-length treat- Our thesis is that, given the limita- merit of environ- nor the intuitions of those in the fore- tions of captive breeding technology and mental ethics by a front of the environmental movement." the financial limitations of the species professional phi- preservation movement, today's captive IuJCIpIIGI, a5 UIU breeding programs can only be viewed in economist William a strongly positive light from a very nar- Baxter (1974) in his widely-cited book, birds. This is because, from the animal row anthropocentric perspective, a per- People or Penguins: The Casefor Opti- rights perspective, what matters is the spective which matches neither the mal Pollution. From these authors' welfare of "sentient" beings (that is, rhetoric nor the intuitions of those in the purely anthropocentric perspective, en- conscious, experiencing beings), and forefront of the environmental move- dangered species should be preserved presumably many "lower" animals are ment. We do not mean to suggest that all only if and insofar as doing so is justified not sentient. [The "animal rights" label is captive propagation ought to be aban- on economic, scientific, recreational, also misleading insofar as many of those doned, but we do argue that sometimes, religious, or other grounds tieddirectly to who argue from this perspective (includ- when a choice must be made between human interests. ing Peter Singer) are not advocating what saving a particular species through A very different perspective, and a philosopher would recognize as rights captive breeding and acquiring one espoused by many environmentalist, for non-human animals. See Singer important habitat for in situ preservation is the holistic perspective championed by 1978, pp. 119-25, and Regan 1983, sec- of a whole biotic community, we ought Aldo Leopold in A Sand County Alma- tions 6.4 and 6.5.1. The most widely-read to choose the latter and knowingly nac. In his summary statement of the account of the sentientist perspective is consign a species to extinction. Such "land ethic," Leopold defined morally Peter Singer's Animal Liberation (1975), choices will be difficult, but our exami- correct actions as those which "tend to but the most philosophically rigorous is nation of the ethics of captive breeding preserve the integrity, stability, and Tom Regan's The Case for Animal suggests that current priorities need some beauty of the biotic community" (Leo- Rights (1983). Although Singer's and rearranging. pold 1948, pp. 224-225). Leopold'sprin- Regan's accounts differ greatly in detail, ciple is holistic, because it subordinates they are both (as we will put it) sentien- Three Ethical Perspectives the interests of individual members of the & Sentientists share the anthropocen- biotic community - be they deer or trists' emphasis on individuals, rather Captive breeding programs can be human beings - to the welfare of the than ecological systems. Both find in- evaluated from three very different ethi- community as a whole (Callicott 1989). trinsic value in individual organisms, cal perspectives. [Other perspectives are Those who argue that all wild species and only instrumental value in ecologi- possible, but these three represent those ought to be preserved, even if doing so cal systems. Sentientists simply find most commonly encountered in the lit- would not benefit human beings, usually intrinsic value in a wider range of erature of environmental ethics and the are arguing from such a holistic stance. individuals.

Vol. 8 No. 1 Endangered Species UPDATE 27 In practice, environmentalists com- representation and prevent inbreeding mammalian taxa, captive breeding pro- monly argue from a complex mixture of depression, including euthanization of grams can receive little or no support these three ethical perspectives. Al- offspring with overrepresented genes, from the sentientist perspective. though anthropocentric considerations embryo transfers between species, injec- From the holistic perspective, these predominate in the policy arena, environ- tions to superovulate females, and same resource limitations markedly limit mentalists commonly claim that environ- double-clutching, the value of captive breeding. The holis- mental problems will not ultimately be From a sentientist perspective, even tic perspective places a premium on pre- solved until we adopt and act on a non- if a species is going extinct in the wild so serving the integrity of natural anthropocentric ethic. For instance, ecosystems. But it is impossible to pre- Denis Hayes, national coordinator of the serve the integrity of an ecosystem with- first Earth Day, described the goal of the out preserving all (or at least most) of its event in these terms: "We hopedit would ". . . from the holistic component species. To the degree that lead to a new kind of ideology, a new perspective, today's the holistic perspective is truly ecosys- value system based on ecology and a captive breeding pro- temic, it will find relatively little value in reverence for [all] life" (New York programs which, like today's captive Times, 4/16/90). Similarly, in 1980, the grams are a rear- breeding programs, tend to preserve only United Nation's World Conservation guard action in a war the large, impressive mammalian and Strategy declared that: which we are still avian species and consigns the far more "Ultimatelythe behavior of entire losing." numerous endangered species of mol- societies towarch the biosphere lusks, insects, plants, and fungi to must be transformed if the extinction. achievement of conservation ob- The value of captive breeding pro- jective~is to be assured. A new that captive breeding is the only possible grams is further limited, from the holistic ethic, embracing plants and ani- way to preserve it, it is still difficult or perspective, by the degree to which suc- mals as well aspeople, is required impossible to justify captively breeding cessful reintroduction is thwarted by for human societies to live in har- the remaining individuals. Singer and continuing habitat destruction and the mony with the natural world on Regan both acknowledge this,andRegan maladaptive influences of captivity. If which they dependfor survival and labels the differential treatment of mem- the success of a captive breeding pro- well-being " (World Conservation bers of endangered species "environ- gram is measured in terms of the likeli- Strategy 1980). mental fascism" (Regan 1979, sec. 9.3; hood of eventually restoring the organ- In such statements, the sentientist and see also Singer 1979). However "good" isms to a natural system, then captive holistic perspectives tend to become a program may be when seen from the breeding must be seen as a way to buy blurred together, but what is clear is that perspective of the species, a captive time for effective habitat preservation environmentalists commonly place a breeding program cannot plausibly be and enhancement. From the holistic heavy emphasis on non-anthropocentric said to be good when viewed from the perspective, this must be at the core of a ethical perspectives. perspective of the individual animals successfulprogram. Arelatedproblemis involved. Where the individuals in- that prolonged captivity tends to erase Evaluating Captive Breeding volved are not sentient (plants, for in- learned information about feeding and stance, or mollusks or insects), aprograrn nesting sites and learned behaviors re- But how do captive breeding pro- is not sullied from thesentientistperspec- lated to reproduction. It is also a concern grams look from these non-anthropocen- tive. However, to the extent that eco- that after several generations in captivity, tric perspectives? From the perspective nomic limitations force us to focus re- genetic selection inevitably begins to of individual sentient animals involved sources primarily on the higher avian and favor traits suited to captivity - traits in a program, captive breed- which are unlikely to be ing is a moral atrocity. Eco- adaptive upon return to the nomic constraints prevent wild. confinement systems from All of this suggests allowing birds and mam- ". ..when a choice must be made between that only when captive mals adegreeof freedom of saving a particular species through cap- breeding is complemented movement comparable to five breeding and acquiring important by an aggressive Program the wild - a condition habitat for in gitu preservation of a whole 0f habitat preservation which must be especially does it look good from the distressing to first genera- biotic community, we ought to choose holistic perspective. Yet, tion breeders who have the latter and knowingly consign a Spe- the cause of spe- known freedom. ~1~0,very cie~to extinction." cies endangerment still invasive techniques must be remains habitat loss. So used to maintain full genetic from the holistic perspec-

28 Endangered Species UPDATE Vol. 8 No. 1 tive, today's captive breeding programs sources would lead us to preserve evolv- late for this strategy to work. These are are a rear-guard action in a war which we ing ecosystems. But as we have already the cases in which a balance must be are still losing. seen, Erom such an ecosystemic perspec- struck. Where resources are scarce, it Hence, it is only from the human- tive, a captive breeding program can be cannot plausibly be claimed that captive breeding is always warranted. For instance, it is not clear that we should spend millions of dollars to breed a species like "Enlightened anthropocentrists find instrumental the California condor, which has a thriv- value in ecosystems, whereas holists find intrinsic ing functional correlate (the turkey vul- value there, but neither perspective can endorse ture), if the money could be redirected to habitat acquisition which would benefit captive breeding without habitat preservation." other specieshh. "Flagship" projects, like the California condor recovery program, may ultimately generate a great deal of public support, but if we want to centered perspective of anthropocen- only as successful as a concurrent pro- be true to the holistic ideals of the envi- trism that great value can be placed on gram of aggressive habitat preservation ronmental movement, we must openly today's captive breeding programs. and enhancement. Enlightened anthro- question the wisdom of captive breeding Even here, however, it makes agreat deal pocentrists find instrumental value in in such cases. of difference which human interests ecosystems, whereas holists find intrin- captive breeding programs are expected sic value there, but neither perspective Literature Cited to serve. can endorse captive breeding without If our only goal is to preserve habitat preservation. Baxter, W.F. 1974. People orpenguins: The Case unusual strains of food crops or medici- for Optimal Pollution. Columbia University Press. New York, NY. nal plants as genetic resources, then al- Conclusion Callicott, J.B. 1989. Animal liberation: A triangu- though resource constraints severely re- lar affair. In In Defense of the Land Ethic. strict the number of species that can be We conclude that, given the limita- Statue University Press of New York, Albany. (For comparison, aho see "Animal liberalion thus preserved, captive breeding can be tions of captive breeding technology, and and environmental ethics: Back together seen as a highly valuable technology. As given the financial limitations of the again," in the same volume.) long as we have no interest in in sit~ current species preservation movement, Lapold, A. 1948. A Sand County Almanac. preservation, captive breeding looks captive breeding programs can only be Oxford University Press, London. New York Times. April 16, 1990. "Veteran of good from an anthropocentric perspec- viewed in a strongly positive light from a Earth Day 1970 Looks To A New Day." (Philip tive. very narrow anthropocentricperspective Shabecoff.) However, from a more enlightened emphasizing species' resource value to Norton, B. 1987. Why Presewe Natural Variety? anthropocentric perspective, captive present and future human beings. With- Princeton University Press, Princeton, NJ. Passmore, J. 1974. Man's Responsibility for breeding programs score poorly for the out meaning to trivialize this value, we Nature. Charles Scribner's Sons, New York. same reasons they scoredpoorly from the emphasize that it does not represent the Regan, T. 1979. Case for Animal Rights. Section holistic perspective. In Why Preserve kind of non-anthropocentric perspective 9.3 in K.E. Goodpaster and K.M. Sayre, eds. Natural Variety?, philosopher Bryan that so many environmentalists believe Ethics and Problems of the 21st Century. Uni- versity of Notre Dame Press, Notre Dame. Norton (1987) defends such an enlight- should guide our thinking. Regan, T. 1983. The Case for Animal Rights. ened anthropocentrism. He argues that, How should environmentalists re- University of Callomia Press, Berkeley. even if only one in 10,000 species has solve this impasse? One way would be to Singer, P. 1975. Animal Liberation. Avon Books, direct value to human beings, all species abandon their non-anthropocentric New York. Singer, P. 1978. The fable of the fox and the have indirect value, because diversity rhetoric. Yet, as Norton's argument unliberated animals. Ethics 88:119-25. begets diversity: increasing diversity shows, even the more enlightened an- Singer, P. 1979. Not for humans only: The place increases opportunities for niche spe- thropocentrists cannot support captive of non-humans in environmental issues. Pages cialization, which in turn produces more breeding without effective habitat pres- 191-2lX in K.E. Goodpaster and K.M. Sayre, eds. Ethics and Problems of the 21st Century. diversity, and so on, in an ascending ervation. The upshot is that both environ- University of Notre Dame Press, Notre Dame. spiral which produces those relatively mentalists arguing from a holistic per- World Conservation Strategy. 1980. Gland, Swit- rare species that are of direct use to us. spective and conservationists arguing zerland: IUCN, UNEP, WWF. And in a period of rapid biological im- from an enlightened anthropocentric poverishment, each species' value in- perspective should push for an intelligent Gary Vamer is Visiting Assistant Professor of creases dramatically, because loss of balance between habitat preservation and Philosophy, Texas A&M University, College Sta- diversity similarly begets further loss of captive breeding. While sometimes the tion, TX 778434237, Manha C. Monroe is a Ph.D. candidate at the School of Natural Re- diversity in adescending spiral. Thus, an preservation of a particular endangered sources, University of Michigan, Ann Arbor. MI enlightened understanding of our interest species can be accomplished through 48109-1115. in preserving species as genetic re- habitat preservation, all too often it is too

Vol. 8 No. 1 Endangered Species UPDA TE 29 Captive Propagation and the Conservation of Species: A U.S. Fish and Wildlife Service Perspective

James Tate, Jr.

Intent of the Endangered basic goal of conserving functioning sial unless a vigilant watch is kept on Species Act ecosystems. other threats to the species' survival.

In writing the Endangered Species Success Stories Questionable Programs Act (the Act), Congress specifically stated that the Act's purpose is to con- It is, of course, true that some species Unfortunately, for most species, such serve the ecosystems upon which endan- can be raised in captivity and returned a combined storehouse of historical ex- gered and threatened species depend. successfully to the wild. The best experience and environmental knowledge This concept - that it is preferable to ample may be the success of the captive simply does not exist. For example, conserve species by conserving rearing and reintroduction program for captive propagation "headstarting" ef- ecosystems - underlies the forts being undertaken by entire Act. Yet in the public- - several government agen- ity that often attends last- "Congress specifically stated that cies for the recovery of sea ditch efforts to save a spe- the [Endangered Species] Act's turtles, in particular cies, the central importance Kemp's ridley sea turtle of ecosystem conservation is purpose is to conserve the (Lepidochelys kempii), do often forgotten. What is left ecosystems upon which endangered not appear to have materi- in the mind of the public is and threatened species depend." ally advanced the recovery the misleading and danger- of these species. ous idea that the Act's pur- The original agreement pose might be to save species one by one, the peregrine falcon. This cooperative for this headstarting program was among in isolation. And possibly worse, that the effort of the public and private sectors of the USFWS, National Park Service success of the Act should be measured by the conservation community was made (NPS), National Marine Fisheries Serv- the success of such efforts. possible in part by the fact that nearly five ice (NMFS), the Texas Parks and Wild- Even though the U.S. Fish and Wild- centuries of recorded history exist on life Department, and Mexico's Secretary life Service (USFWS) conducts research techniques for returning hand-reared of Fisheries, with the USFWS serving as on ways to raise plants and animals in birds to the wild after a period of loose the coordinating agency. [This agree- captivity and release them into suitable control by humans. This process (called ment involved far more initiatives than habitat, the USFWS uses captive propa- "hacking"), although developed over the headstarting experiment, and al- gation and release only as a technique of centuries by trial and error, has only re- though not discussed here, success of last resort for recovering a species from cently been used successfully and sys- other efforts must not be overlooked.] In the brink of extinction. Examples of tematically toreturn such species as pere- the headstarting effort, the USFWS in co- species for which theUSFWS is attempt- grine falcons and bald eagles (Haliaeetus operation with Mexico, moved approxi- ing this approach include Knowlton's leucocephalus) to the wild. mately 2,000 Kemp's ridley eggs annu- cactus (Pediocactus knowltonii), Successful reintroduction of these ally during 1978-1988 from Rancho Kearney's bluestar (Amsonia kearney- avian species, however, was possible Nuevo, Mexico (the only site where they ana), desert fishes, peregrine falcon only because other major threats to their nest), to the NPS's Padre Island National (Falco peregrinus), California condor survival were being addressed at the Seashore. They were turned over to NPS (Gymnoglyps californianus), and black- same time. It was a combination of the personnel who incubated the eggs until footed ferret (Mustela nigripes) (see United States' ban on chlorinated hydro- they hatched. The hatchlings were re- related case studies). The use of captive carbon-based pesticides, a concerted leased briefly on Padre Island in an at- propagation with respect to any species captive-rearing effort, and the applica- tempt to imprint the baby turtles on the depends on many things, including tion of hacking techniques that allowed beach and adjacent waters. They were the feasibility of other less costly, or peregrine falcons and bald eagles to in- then transferred to NMFS facilities in more effective approaches. But funda- crease. These efforts have been success- Galveston, Texas, where they were mental to implementing the Endangered ful enough that it may even be possible to "headstarted" (maintained in tanks) for a Species Act is an understanding of this reclassify these species before long from period of months. In theory, the head- fact: maintaining species only in "endangered" to "threatened." However, started turtles then would be large captivity actually detracts from the Act's even reclassification will be controver- enough to avoid as high a mortality rate as

30 Endangered Species UPDATE Vol. 8 No. 1 the vulnerable hatchlings experience. almost every other effort, has been sus- The hatchery concept has great ap- The headstarted turtles were released pended. There has been no evidence that peal. It sounds ideal: if you need more into the Gulf of Mexico, usually in the the state's long-term efforts, "or any turtles, raise them in a hatchery -just waters off Padre Island. The original other headstarting efforts, have resulted like trout, quail, pheasants, or parrots. agreement among all the parties involved in a single headstarted animal surviving However, the hatchery concept fails to was to conduct the Padre Island experi- and entering a breeding population," address the problems that have brought ment for ten years. It was extended to 11 according to Jack Woody, the USFWS's many species to their present sad state. years, then terminated after the 1988 sea turtle coordinator. Until problems in their environment are transfer of 1,000 eggs to Padre Island. recognized and corrected, we are doing Wnder a separate five-year agreement, Need To Pursue Other Strategies little more than supporting a "put-and- 2,000 Kemp's ridley hatchlings are now take" operation. We are attempting to being moved annually from Rancho Like many other species we are study- treat the signs of the problem, but not the Nuevo to the NMFS facilities at ing, relatively little is known of the life actual cause of the problem. Unfortu- Galveston for continuation of experi- history or ecology of any of the several nately, there are no quick fries for sea mental headstarting.] species of sea turtles. Although we are turtles or any other species on the brink of The USFWS continues to cooperate learning rapidly, the need is to find recov- extinction. fully in the effort to conserve sea turtles, ery answers. Several avenues usually' Similar examples can be found in but it no longer advocates the head- need to be followed before we go to the other hatchery-produced species. The starting project. In his testimony before ultimate, last-ditch stand of captive introduction of non-native ring-necked the House Merchant Marine and Fisher- propagation. Research needs to continue pheasants to previously unoccupied ies Committee, Conley Moffett, Deputy on captive propagation, but not to the habitat throughout central North Amer- Assistant Director for Fish and Wildlife exclusion of other basic studies. ica was successful only because of re- Enhancement for the USFWS, stated that The example of the American alliga- peated introductions of previously wild " . .. at this time, with the data available, tor (Alligator mississippiensis) shows birds to the new habitat. Most hatchery the Servicedoesnot endorse headstarting that captive propagation may not be nec- personnel assume that the vast majority or any form of hatcheries for raising and essary for recovery. This species has of hatchery-reared birds released into a releasing sea turtles into the wild as a been brought back from the brink of new area will be dead within a year. recovery action or an attempt to maintain extinction. Under strict state oversight, Pheasant populations that are constantly present numbers of reproducing marine alligators are now actually being com- supplemented by hatchery-reared birds turtles in the wild." He made it clear that mercialized in certain areas. What the usually collapse soon after the cessation the recovery of Kemp's ridley sea turtle of releases. Similarly, hatchery officials has not been materially advanced by the know that only a tiny fraction of the Program. millions of trout and salmon raised and One would think that such an effort- released annually will survive to the by the agencies of two nations, private ". . . maintaining species point of reproducing successfully in the conservation organizations, and hun- only in captivity actually wild. It took even two tries to introduce dreds of volunteers - should make a detracts from the Act's the highly adaptable starling (Sturnus difference. It would appear to be a sound basic goal of con- vulgaris) into North America, one conservation measure. But the bad news serving functioning hundred years ago this year. is that there is no evidence that head- Many other examples can be found of started sea turtles can be imprinted suc- ecosystems." species not likely to benefit significantly cessfully to a natal beach. If everything from a propagation program, but which worked according to theory, the turtles would benefit from other conservation would return to the beach from which measures, such as habitat conservation they were hatched when they reach ma- alligator needed for recovery was the and control of take. Although in extreme turity. So far, however, no one knows same thing the sea turtles need: strict situations it may well be necessary, the what cues it takes or for how long a sea protection of nesting sites, strict protec- practice of raising species in captivity turtle must be imprinted in order to pro- tion from killing by humans, and long- must not be allowed to overshadow the vide the information it needs to return term management programs to prevent stated purpose of the Endangered Spe- many years later to a specific site. extinction. With such protection, the cies Act to conserve the ecosystems Like hacking hawks to the wild, head- USFWS believes the Kemp's ridley and upon which endangered and threatened starting turtles is not new. But it has been other sea turtle species will begin return- specie$ depend. attempted in many areas around the ing to safer population levels. If this world only for a few decades, with one of protection is not provided, then sea James Tate, Jr. is a Wildlife Biologist in the Divi- the largest and longest programs canied turtles, which have been around for 100 sicn of Endangered Species for the U.S. Fish and Wildlife Service, 1849 C Street, 452 ARLSQ, out by the State of Florida for over a 30- million years or so, may be lost to the Washington, DC 20240. year period. But Florida's program, like world.

Vol. 8 No. 1 Endangered Species UPDATE 3 1 The California Condor: Current Efforts for its Recovery by - Michael Wallace

Reasons for Decline ronment with enhancement from young Captive Breeding produced by a few captive pairs. Sus- The California condor (Gymnogyps pected causes of the decline prior to this Condors are large, long-lived birds californianus) was probably never an study included lack of reproduction that reach sexual maturity only after five abundant species. The fossil record in- due to pesticide contamination or due to six years of age. Their rate of repro- dicates, however, that 11,000 years ago to an insufficient food supply as a duction is slow in the wild, with one chick during the Pleistocene, the California result of an obvious reduction in being produced every two years. A spe- condor once had an extensive range; it foraging area. While these factors may cies with these reproductive parameters occupied the west coast extending from certainly have . contributed to the pre- must have a low mortality rate in order for British Columbia to Baja Califomia, as cipitous decline, field studies con- the population to be somewhat stable; for well as the southem-most states in the ducted during the early 1980s indicated condor populations to be maintained, United States across to Florida, and even that reproduction was close to normal. annual mortality rates have been calcu- hadapresence in upper New York state. Consequently, direct mortality alone lated to lie between five to seven percent Current hypotheses cite massive could account for the population (Verner 1978, Temple & Wallace 1988) extinctions of the "mega fauna," the reduction. Yet, from the early to mid 1980s, condor's food supply, during the lat- 23 to 40% annual mortality was ter Pleistocene as the reason for the documented in the wild condor popu- species' reduction and restriction in lation. This excessive and poorly un- range to only the West Coast (Emslie derstood rate of death eventually 1987). convinced the USFWS to capture the Carrion found along the Pacific remaining wild condors for their own beaches provided by the still-abun- safety, and on April 19,1987,the last dant marine animal populations al- bird was brought into captivity. All lowed the scavenger's survival into hope for the species' future rested recent times. Over the last few centu- with 27 individuals housed at two fa- ries, loss of foraging habitat, en- cilities: the Los Angeles Zoo and the croachment on nesting territories, San Diego Wild Animal Park. The and direct mortality - such as colli- program continues to rely entirely on sions with human-made structures, captive propagation and the eventual ingestion of poison-contaminated release of genetically surplus young carcasses, and shooting - are sus- to the wild. pected causes of the condor's de- In 1987, the population consisted cline. By the early 1980s, only a of 14 wild-caught juvenile and adult remnant population of about two condors and 13 young that were arti- dozen birds remained. ficially hatched from eggs taken from Intensive field studies conducted wild nests. The ability of female by the U.S. Fish and Wildlife Service condors to lay a replacement egg if (USFWS) and others on the remnant the first or even second egg is re- An adult California condor at the breeding facilities in the population in the early 1980s in- Los Angeles Zm Photo by M. Wallace moved during the breeding season volved an innovative photographic provides an opportunity to increase censusing technique and the mark- the reproductive rate up to six fold. ing of most individuals with wing Compared to birds of prey, con- tags and radio transmitters (Snyder ". .. on April 19,1987, the last dors adapt to captivity with relative and Snyder 1989, Snyder and bird was brought into ease. The first signs of pair formation Johnson 1985). During this period it captivity. All hope for the occurred in 1987,with two adults that was hoped that by quickly discover- species' future rested with 27 had been in captivity for about one ing the causes of the decline and im- year. In 1988, this same pair laid one mediately addressing the problems, individuals housed at two fertile egg that successfully hatched. the wild population could be saved facilities. . ." In 1989, five pairs produced seven and maintained in the natural envi- eggs, four of which were fertile and

32 Endangered Species UPDATE Vol. 8 No. 1 hatched healthy chicks, raising the popu- southern California, other ar- lation to 32 birds. In the 1990 season, eas both within and outside nine pairs produced 15 eggs. Eleven of the state of California are these were fertile and eight successfully being carefully considered. hatched healthy individuals. As with The list includes the Grand several other species in captivity, eggs Canyon in Arizona and large produced by first-time breeding condors, tracts of land owned by The even if fertile, do not necessarily hatch, Nature Conservancy in although their subsequent eggs have southwest New Mexico. been viable. If all goes well, production The Recovery Team has may be as high as 13 or 14 young in the recommended that the spe- 1991 season. cies be considered for Two events seem to be occurring downlisting to "threatened" reproductively with the captive flock at when two wild, disjunct this point. Most of the wild-caught birds populations numbering at are now relaxed enough in captivity to least 100 individuals each breed, and the young of the "wild" eggs have been established. hatched in the early to mid '80s have now matured and show no reluctance to Surrogate Work With immediately begin breeding. Some have Andean Condors even shown reproductive activity as early as four and five years of age. For many years, work with Andean condors (Vultur Release Criteria and gryphus) has served as a Population Goals model for the California condor program. Although If the current breeding success contin- endangered as well, popula- ues, releases to the wild can be expected tions of Andean condors to take place within the next few years. found the length of the Andes mountains In anticipation of success with captive The timetable for future releases is set by still number in the thousands. It is similar propagation of California condors, An- criteria recommended by the California in size, behavior and ecology to its north- deans were used as surrogates to develop Condor Recovery Team, a panel of em cousin. Because Andean and Califor- methods for releasing captive-raised biologists which advises the USFWS on nia condors are allopatric, lack of direct condors to the wild. Based on successful program direction. The four genetic and competition likely helped maintain these releases of black vultures (Coragyps behavioral requirements for release species as ecological equivalents with atratus) and turkey vultures (Cathartes candidates are: similar social, foraging and nesting char- aura) in Florida (Wallace and Temple 1) at least 96% of the heterozygosity of acteristics (Brown and Arnadon 1968). 1983), 11 Andean condors were released each of the nine founding lines of the The close similarities between the two in northern Peru. Seven survived and population must be retained in captivity condor species have thus far ensured that integrated successfully into the wild before subsequent progeny can be con- techniques developed on Andean con- population over a two-year period (Wal- sidered for release (this amounts to five dors have been equally applicable for use lace and Temple 1987). young per genetic line); on Californians. The climate, topography, food disper- 2) at least three pairs of founders are Many zoos throughout the world have sion and human activity levels are drasti- breeding; had success in breeding Andean condors cally different between the area that 3) candidates must be physically and (Cade 1986, Lint 1%0). Artificial incu- Andean condors were released in Peru behaviorally releasable; and bation and captive management tech- and the southern California environment 4) there must be at least three birds in niques used on the California condors at in which California condor releases are any one release group. the Los Angeles Zoo and the San Diego planned. While the captive California Under these guidelines the reproduc- Wild Animal Park have been based on condor flock increases over the next few tive pace of the birds themselves deter- extensive experience accumulated by years to a level where young are available mine the timetable in which releases will zoos working with eggs and young of for release to the wild, Andean condors occur. A few lines are close to meeting Andean condors (Kuehler and Witman are again being used in the recovery ef- these criteria; if reproduction in the 1988). Radiotelemetry transmitters and fort to tailor condor release methods coming season is exceptional, releases tracking techniques were also developed developed in Peru to the conditions that could commence in 1991, however, it is with Andean condors, as well as trapping exist in the southern California moun- more likely to occur in 1992. Although and field handling methods, prior to their tains. the first releases of California condors use on California condors (Wallace and Since 1988, nine zoos have contrib- are scheduled to occur in sites tested in Temple 1987). uted 29 fertile Andean condor eggs to the

Vol. 8 No. 1 Endangered Species UPDA TE 33 slopes. The local oil companies that use the power lines have consented to bury the offending section and others similar in position to help prevent future inci- dents with California condors. The remaining six condors of this first group were returned to captivity so they could not adversely influence six others subsequently released in a more isolated region of the range. As hoped, the second group chose a more secluded slope to practice soaring, and continued to respond much less positively than the first group to human activity when they later encountered it. Lead poisoning by ingestion of bullet fragments from unretrieved deer carcasses during the hunting season has been identified as a mortality factor in the original free-flying population of Calit black bears. Calf carcasses are acquired f& frqm neighhgdairies. Photo by M. Wallace fomia condors. A major question about the behavior of released condors is program -their entire production for the must be carefully chosen by considering whether their feeding activity and forag- 1988 and 1989 breeding seasons. Ac- the management logistics of maintaining ing pattern could be controlled in a way companied by zoo keepers, eggs were visual isolation while caring for the that would minimize their exposure to flown in portable incubators to the Los chick's physical needs, that the immedi- this hazardous situation. By feeding the Angeles Zoo and San Diego Wild Ani- ate topography must afford protection released Andeans on mountain peaks mal Park for the 56 to 62-day incubation from large mammalian competitors and where soaring flight for the inexperi- period. The chicks were reared in isola- provide adequate ridge lift to encourage enced buds is easy but where natural tion from human contact using hand flight, and also that the degree of human carcasses are scarce, the young condors puppets modeled after adult condors. activity in the surrounding landscape is have fed exclusively on food we offer, This strict isolation was maintained in the an important factor. even though they traverse over 100 miles field release pens as well, when the nest- of wild habitat. Training the birds to a lings were transferred at three to five Managing Causes of Mortality safer foraging pattern in this way is likely months of age to the areas in which they to help reduce mortalities in the first few would eventually be released. Main- In search of good flying conditions, years or possibly decades after California tained in groups of three to four at various buds in our first group were attracted condor releases have begun, but is not sites, seven Andeans were released in soon after release to particularly windy likely to eliminate lead poisoning as a 1988-89 and six in the 1989-90 season. slopes over a mile from the release area. mortality factor for this species in the These experimentalreleases were funded These slopes were also heavily used by long term. by the USFWS and the California De- workers from the petroleum field located The best alternative to lead bullets partment of Fish and Game (CDFG), and there. It appears that the close exposure being investigated at this time is copper. continue to be jointly supervised by Los to the high level of human activity, A pure copper slug in all the typical size Angeles Zoo and USFWS personnel. power lines and vehicular traffic over the loads for deer is currently being used in The importance of these "test runs" in first months of their fledging experience various parts of the country, and its release programs cannot be overly em- habituated the birds to those features in manufacturer claims good reception by phasized. The zoo production and man- their environment. When they eventu- the hunting public because of superior agement of the eggs and young in this ally ranged over a 100-milearea, they re- ballistics and competitive price. If cop- program afforded the opportunity to as- sponded to the presence of people and per bullets prove non-toxic to the birds sess the logistics of egg transport, im- human-made structures with far less fear and their use can be encouraged in areas prove incubation parameters, and revise than we would expect or like to see. Early where condors and deer hunters coexist, chick rearing methods in view of later in the releases, an inexperienced Andean then one major source of mortality can be release results. Important refinements in condor died as the result of a collision reduced. condor release techniques are being with a power line in the same area on a Andean condors may be used to teach made to make future releases of Califor- prominent ridge. The incident under- the first release group of California con- nia condors more efficient and safe. For scored the necessity of selecting sites dors a successful routine for life in the instance, we found that the release site with a sufficient buffer of unobstructed wild. If the current six Andeans are still

34 Endangered Species UPDA TE Vol. 8 No. 1 free and behaving appropriately when represent relatively safe areas for con- understanding learned with each circum- California condors are ready for release, dors to live. The agencies that preside stance, the causes can be dealt with on an there is an opportunity to capitalize on the over these important habitats, the individual basis. It appears that we can experience of the Andeans by having USFWS, the CDFG, the Bureau of Land expect a reasonable amount of success in them show the foraging pattern, roost Management, the U.S. Forest Service establishing populations - at least for sites and drinking pools t the newly and others, must becommended forcon- the short term. What cannot be deter- fledged Californians before the Andeans tinuing their protection of these lands mined, at this point, is the long-term are again taken captive. Utilizing the even while lacking the presence of the viability of a population recovering from experience of the Andeans in this way birds. such a tight genetic bottleneck. It will be could significantly reduce the chances of many decades before we really know the the newly fledged Californians acquiring success or failure of these efforts. behaviors that are maladaptive, or at least, speed their adjustment to a wild Cited environment. "We are fortunate to have a test surrogate for the Brown, L., and D.A. Amadon. 1968. Eagles, Habltat Protection for California condor such as Hawks and F~~CO~Sof the World. Country Life Reintroduced Condors the Andean [condor], but Bds. Cede, TJ. 1986. Propagating diurnal raptors in the methods developed wti~~:A review. Lnternational Year- One concern expressed by many book 24/25: 1-20. people when all of the California condors "Ing "'ltur have 'lso Emae, S.D. 1987, Age and diet of fossil ,for- were brought into captivity, was whether been applied to its Own nia condors in the Grand Canyon, Arizona. condor habitat would be lost to develop Science 237:768-770. recovery as well." Kueh1er.C.M.. and P.N. Witman. 1988. AM~cial ment and expansion of the human popu- incubation of California condor, Gymnogyp lace during the interim before condors californianw, eggs removed from the wild. Zoo could be released again into the wild. Biology 7:123-132. This concern begs the important ques- Lint, K.C. 1960. Notes on breeding Andean condors at the San Diego Zoo. International Zoo tion: exactly what is condor habitat? Conclusion Yearbook, 2:82. Their foraging habitat varies widely. Snvder. N.F.R.. and E.V. Johnson. 1985. Photo- One could safely say that it is wherever we are fortunateto have a test SUffO- iraphic cen*using of the 1982-1983 California condor population. Condor 87:l-3. they can find carrion and are not dis- gate for the condor such as the Snyder,N.F.R., and H. Snyder. 1989. Biology and turbed while eating it. At one time this Andean, but the methods developed US- conservation of the California condor. Pp. 175- included the California beaches and the ing Vultur have also been applied to its 267 in M.P. Dennis, ed. Current Ornithology. flatlands and open slopes around the San Own recovery as well. In order to VO~6. "es~,New YO& and London. Temple, S.A., and M.P. Wallace. 1988. Survivor- Fernando Valley - even the Los Ange- ' Over the possibility of Andeans ship patterns in a population of Andean condors les Basin itself. The foothills of the San proliferating in the wilds of North Amer- vdtw gryph~.Pp. 247-251 ~r B.U. Meyburg Joaquin Valley have been most recently ica, only females were allowed to be and ~.~.Chanceior,eds. Rapton in thelodl used extensively by the birds for forag- released. The 14 male Andeans that em World. WWGBP: Berlin. London, Paris. Verner, J. 1978. The California condor: Status of ing. Yet this area consists of a mosaic of hatched during the Califomia experi- the recovery effort. Gen. Tech. Rep. PSW-28, privately owned land that is being con- mental program were also reared in isola- U.S. Forest Senice, Washington D.C. tinuously converted from ranching to tion. However, they were then shipped to Wallace,M.P., and S.A. Temple. 1983. An evalu- agriculture at a rate that is unrelated to the Columbia, South America, and released ation of techniques for releasing hand-reared vultures to the wild. 4.400-423 in S.R. Wilbur condor's presence or absence. Condors in areas were their specieshas declined or and J.A. Jadtson, eds. Vulture Biology and will feed wherever food and opportunity no longer exists. When the California Management. California Press. are available - if establishment of safe experimental release program has been Wallace,M.P., and S.A. Temple. 1987. Releasing feeding areas is deemed as a necessary captive-reared Andean condors to the wild. J, completed, the females will be flown to Wildlife Management 5 1(3):541-550. management activity, then this will need Columbia and released into the wild to to be done as necessary. join the males. Both condor species are However, the "condor habitat" more directly benefitting from these programs. critical to the future survival of the spe- If captive propagation of California cies is the nestingiving area found in the condors succeeds to the degree we antici- Michael Wallace is Curator of Birds at the Los mountains of southern California. Fortu- pate, adequate numbers of young should Angeles Zoo (5333 Zoo Drive, Los Angeles, CA nately, huge core mountainous areas be available for annual releases. With 90027), and is also a member of the California currently are under protection by one basic release methods in hand, we are Condor Recovery Team. governmental agency or another; the likely to witness significant increases in Sespe Condor Sanctuary, Hopper Moun- populations as we attempt to establish tain National Wildlife Refuge, and the them in selected sites. Some mortalities Los Padres and Angeles National Forests will occur, but hopefully, with increased

Vol. 8 No. 1 Endangered Species UPDA TE 35 The Decline and Restoration of the Guam Rail, Rallus owstoni by Gregory J. Witteman, Robert E. Beck, Jr., Stuart L. Pimm, and Scott R. Derrickson

INTRODUCTION found rails restricted to two small dis- cillatus) (Beck 1988b). Unfortunately, continuous populations in northern the bridled white-eye, rufous-fronted The Guam rail, Rallus owstoni, is a Guam (Aguon 1983). A repeat of the fantail, and Guam flycatcher were so rare large rail (200 to 300gms)known histori- same survey in 1983 found an 80% .de- by mid-1983 that captive programs for cally only from Guam, with no closely cline from the previous year. By 1984, these species had to be discontinued related forms nearby in Micronesia only approximately 20 rails remained (Beck 1988b). (Baker 1951, Ripley 1977). Baker in northern Guam, with the possibility of (1951), however, speculates that the a few scattered isolated individuals CAPTIVE BREEDING species may have had a wider range in elsewhere (Beck 1988a). Consequently, former times. R. owstoni was distributed the Guam rail was listed on the U.S. Propagation efforts for the king- over much of Guam in all habitats except Endangered Species List in 1984 fisher and rail were subsequently accel- wetlands (Baker 1951, Beck and Savidge (USFWS 1984a,b). The last confirmed erated, and a cooperative captive breed- in press), although Jenkins (1979) con- ing program was organ- sidered both savanna and mature mixed ized under the auspices forest to be marginal habitat. On Guam, of the American ASSO- the rail was an opportunistic omnivorous "The story of the decline of Guam's ciation of Zoological feeder that appeared to prefer animal forest birds . . . is one of the most Parks and Aquariums over vegetable matter; it was known to spectacular examples of the damage (AAZPA). initial par- eat gastropods, skinks, geckos, insects, caused by an introduced species." ticipants included the and carrion, as well as seeds and palm DAWR, the Philadel- leaves (Jenkins 1979). phia Zoological Garden The Guam rail was a year-round (Shelton 1986a,b,c), the ground nester with a nesting peak occur- observation of a Guam rail in the wild Smithsonian Institution's National Zoo- ring during the rainy season. Both sexes occurred in 1987 (Beck 1988a). logical Park (Derrickson 1986a,b), and shared in the construction of a shallow The story of the decline of Guam's the New York Zoological Society's nest made of loose grasses and in the forest birds (see particularly Savidge Bronx Zoo (Sheppard 1985). Between incubation of eggs which lasted about 19 1987) is one of the most spectacular February 1983, and September 1986, days (Jenkins 1979). The typical clutch examples of the damage caused by an 21 rails were taken from the wild as eggs, consisted of three to four eggs, but the introduced species (Pimm 1987). The chicks, immatures, or adults for propaga- number of clutches per year on Guam spatial and temporal distribution of the tion purposes. was unknown (Jenkins 1979). The 172 introduced brown tree snake (Boiga ir- Because sexual maturity occurs at broods observed in the wild ranged in regularis) was well-correlated with the only four months of age and breeding is size from one to four, with an average of decline of Guam's native birds. Forest year-round, the captive population has two. One family group that was observed bird communities containing up to ten grown rapidly despite pairing difficul- foraging in the wild consisted of two species lost all of those species, with ties caused by high levels of aggression. adults, several irnmatures and three population declines occurring sequen- The first successful breeding of the rail chicks (R. E. Beck, Jr. and E. S. Morton, tially in the central, southern, and north- as a part of the captive breeding program unpublished observations). em portions of the island. When it be- occurred at the DAWR's facility on came apparent in 1982 that many, if not April 17, 1984 (Beck 1984). The first DECLINE OF THE GUAM RAIL all, of Guam's native forest birds would breeding of the rail at a mainland U.S. soon be extirpated, DAWR personnel zoo occurred in September 1984, at the Prior to the 1960s, the Guam rail was made the decision to develop captive National Zoological Park (Derrickson abundant island-wide with its population breeding programs for Guam's five re- 1986a,b), There are presently over 180 estimated to be in the tens of thousands maining endemic species: the Guam rail, rails in captivity in 14 U.S. zoos and in (Lint 1968). By 1982, a survey con- Guam Micronesian kingfisher (Halcyon Guam, derived from 13 of the original ducted by the Division of Aquatic and c. cinnomomina), ~fo~s-frontedfantail 2 1 potential founders. If three additional Wildlife Resources of the Department of (Rhipidura rufifrons uraniae), Guam surviving wild-caught birds breed suc- Agriculture (DAWR), making use of flycatcher (Myiagra freycinen), and cessfully, the number of founders can be tape recorded play-back of rail calls, bridled white-eye (Zosterops c. conspi- raised to 16. Unfortunately, all attempts

36 Endangered Species UPDATE Vol. 8 No. 1 to pair and breed these three hand- THE ROTA INTRODUCTION the Guam rail (USFWS 1989). The pres- raised birds have failed thus far due to ence or absence of ecologically similar social and sexual incompatibility. Choice of Location species is known to influence the success Fortunately, the captive population has of introductions on oceanic islands continued to be characterized by an even In planning the restoration of any (Moulton and Pimm 1985). sex ratio. species, one must answer the obvious Since detailed information on the Although captive populations can questions of where the introduction rail's historical habitat preferences is obviously serve as important species should take place, how many individuals lacking, we intend to release birds in a reservoirs, they necessarily lack a num- should be released, and how many re- number of locations and habitats on Rota. ber of important features of wild popula- leases will be required to establish a vi- By following the birds after release and tions. Genetic variation can be quickly able population. Because there is no determining their habitat prekrences and depleted in small populations as a result known way to control or eradicate the survival, we will beable to better identify of genetic drift and inbreeding (Foose brown tree snake on Guam at present potential sites on the island for future and Ballou 1988). Furthermore, behav- (Savidge 1987, Fritts 1988), it was de- releases. ioral adaptations necessary for survival cided to establish the rails on other is- in the wild may be progressively lost in lands in the Marianas (Beck 1987, Beck Number of Releases and Birds the altered captive environment and Savidge in press). Released (Kleiman 1980). In short, re-establishing The island of Rota was subsequently and maintaining a wild population selected for initial reintroductions for a In the case of the Guam rail, there should be a high priority for any captive variety of reasons. First, the island is is necessarily a limit to the number of breeding effort (Conway 1980). located only 50 km north of Guam, and its individuals that can be taken from the Any reintroduction should attempt to proximity and access by commercial air- captive population for release. The cur- provide as broad a cross-section of origi- line would minimize logistical problems. rent size and productivity of the captive nal genetic variation as possible in the Second, the island has only about 1,500 population suggest that we will soon be released animals (Hedrick et al. 1986). inhabitants, which would allow for an able to produce between 75 to 90 rails Because the pedigree of the captive effective public education program. per year for release, without jeopar- population of Guam rails is relatively Third, Rota's climate and plant and ani- dizing either the genetic or demographic shallow (Derrickson 1986a,b), current mal communities are nearly identical to composition of the captive population. genetic and demographic management is Guam, and contains no predators not Yet even with this number of birds, aimed at maximizing founder genome found on Guam. Finally, there are no many questions remain concerning the equivalents -a strategy that effectively other rails or ecologically similar species optimal number of birds per release, maximizes genetic diversity in both the on Rota that might impact, or be ad- the frequency of releases, and the number captive and the reintroduced populations versely affected by, the introduction of of release locations. Fortunately, both (Haig et al. 1990). The potential loss of crucial behavioral adaptations is, of course, more difficult to predict. However, recent analyses involving birds and mammals (Grif- fith et al. 1989) and gamebirds (Wit- teman and Pimm in prep.) have demonstrated that reintroductions of wild- caught animals are much more likely to succeed than reintroductions of captive- raised animals. While a genetic explana- tion for these results cannot be ruled out, it seems likely that behavioral deficits in captive-bred animals are the likely cause of this difference. Although the captive breeding program for the rail has attempted to avoid the loss of wild behavior by maintaining birds in natural settings and minimizing inadvertent selection for tameness, progressive behavioral change through time in captivity is inevitable. Therefore, it is extremely prudent to proceed with re- establishing a wild population as soon as possible. Guam rail (Railus owsfon3 Photo by Jessie Cohen, National Zoological Park

Vol. 8 No. 1 Endangered Species UPDATE 37 theoretical insights and empirical than demographic accidents, and below possible with the number of birds we studies can help us make these decisions. seven pairs, the reverse was true. now have. Both environmental disturbances The second approach examined the In short, we estimated that the best and demographic accidents appear to be success of several hundred introductions way to introduce the Guam rail would be important in determining the fate of small of gamebirds into the United States to split the large number of individuals wild populations (Pirnm et al. 1988). At (Witteman and Pimm in prep.). We available for release into smaller release extremely small numbers, breeding may considered only those species that were groups, but not so small in size that the produce only individuals of one sex, all eventually successfully introduced. In- numbers fall to levels where demo- the individuals may die in the same year terestingly, even for these species and graphic accidents are the principal cause from independent causes, or individuals even when large numbers of individuals of extinction. On the other hand, if only may never even find a mate. These are were released, most introductions failed. very few individuals were available, the demographic accidents, and by releasing This result suggests that the severity of release would have to be postponed until more individuals, we minimize the environmental effects varies considera- a larger number of individuals become chance that they will prevent successful bly, so choosing the right place and time available. The more releases attempted, introductions. Environmental distur- for an introduction is important. On the better the chance of hitting the right bances (for instance, a major typhoon), average, about seven releases were re- time and place. As indicated above, result in many individuals dying of the quired before one was successful. captive breeding efforts are such that we same cause. As such, they can causeeven The relationship between success expect to be able to release 75 to 90 per very large populations to become extinct, and the numbers of individuah intro- year. This number provides enough indi- or a very large introduction to fail. The duced was logistic, that is, with increas- viduals for several releases, and is close obvious way to minimize such effects is ing numbers, success first accelerated to the optimum release number providing to introduce individuals to different and then reached a plateau. The inflec- that the birds remain near each other and places. For instance, it is likely that some tion point is where the per individual form a breeding population. parts of the islands will fare better than cost of becoming successfully estab- others (in the example of the typhoon, lished is minimized (around 75 individu- First Rota Release some parts of the island may be better als for the gamebirds), and the maximum sheltered). This is a strategy of spreading success rate was about 15%. This rela- We released the first Guam rails on risks, and it also helps to minimize the tionship also suggests a strategy for Rotabetween December 1989, and early problems associated with our lack of optimizing the chance of a successful in- February 1990. On Rota the birds were detailed knowledge about the rail's habi- troduction. If several hundred individu- held for several days at a holding facility tat preferences and requirements. als are available for release, several at the release site. During this time they The difficulty is obvious. With a smaller releases in different places and were weighed, measured, and fitted with fixed number of birds available for re- at different times can have a higher color leg bands and harness-mounted lease, we cannot release large numbers of probability of establishing at least one radio transmitters. The transmitters were of both solar and battery types and population.. - When the total number of indi- weighed between four to seven percent ". . . progressive behavioral change viduals falls of the tails' body weight. The rails were through time in captivity is inevitable. below 75, the fed a reduced portion of the feed used in Therefore, it is extremely prudent to release into smaller the Guam breeding facility and a supple- proceed with re-establishing a wild ones decreases the ment of live-caught insects. Supplemen- population as soon as possible." overall probability of tal feeding began on Guam, and was in- success. tended to enhance the foraging abilities The optimal num- of the rails after release into the wild. birds (to minimize the risk of demo- ber for gamebird releases must be about It became clear that use of radio trans- graphic accidents), to large numbers of 75 individuals, for reasons already ex- mitters would be critical in ascertaining places (to minimize the risk of environ- plained. This is roughly comparable to the birds' disposition; because of their mental disturbances). So what is the the optimum result based on the island secretive behavior, there were only two optimal solution and how do we find it? study, that is, seven breeding pairs. confirmed sightings of rails one week We have used two approaches. The However, seven pairs (14 individuals) after the release. Unfortunately, eight first (Pimm et al. 1988) examined the fate must be on the low side for an optimal (36%) of the 22 birds released carried of several hundred small populations of release, for there would certainly be transmitters that failed, so we have no about a hundred species of buds on is- some non-breeding birds in the popula- idea of their fate. Obtaining reliable lands off the British coast. Using an tion. At the very least, these estimates transmitters is now a major concern. Of indirect and technical argument that (between 14 to 75 individuals) suggest the birds we could track, there seemed to space prevents our repeating here, we that we have enough rails to attempt an be little evidence of site fidelity, destroy- estimated that populations above seven introduction. If the optimal size for an ing any hopes that we could establish a breeding pairs were more likely to fail introduction were an order of magnitude viable population in the area we chose. because of environmental disturbances larger, for example, it would not be The birds often moved along roads fol-

38 Endangered Species UPDA TE Vol. 8 No. 1 lowing their release, suggesting that they University of Tennessee, the Smithsonian Institu- Rail. Condor 81(4):404-408. were seeking more open habitats than the tion, Section 6 Endangered Species Aa grants to Kleiman, D.G. 1980. The sociology of captive the Guam Division of Aquatic and Wildlife Re- propagation. Pages 243-261 in M.E. Soule and one into which we released them. In sources, and numerous member zoos of the B.A. Wilcox, eds. Conservation Biology, Sin- choosing roads, the birds were vulner- American Association of Zoological Parks and auer Associates, Inc., Boston, MA. 395 pp. able to traffic, and three birds were killed Aquariums. Lint, K.C. 1968. A rail of Guam. Zoonooz 41(5):16-17. by vehicles. The other certain major Moulm,M.P.,andS.LPimm. 1985. Theextent cause of mortality was cat predation. Literature Cited of competition in shaping an experimental Aguon, C.F. 1983. Survey andinventory of native avifauna. Pages 80-97 in J. Diamond and T. landbirds on Guam. Pages 143-156 in Research Case, 4s. Community Ecology. Harper and Projea Report, Guam Aquatic and Wildlife Row. New York, NY. 'The more releases Resources, FY 1983 Annual Rep* Depart- Pimm, S.L. 1987. The snake that ate Guam. attempted, the better ment of Agriculture, Guam. Trends in Ecology and Evolution 2:293-295. Baker, R.H. 1951. lheavifauna of Micronesia, its Pimm,S.L, H.L. Jones,andJ. Diamond. 1988. On the chance of hitting origin, evolution and distribution. University of the risk of extinction. American Naturalist the right time and Kansas Publication,Museum of Natural History 132:757-785. 3(1):1-359. Ripley, S.D. 1977. Rails of the world. '~odine, place." Beck, R.E., Jr. 1984. Survey and inventory of the Boston, MA. 406 pp. native landbirds on Guam. In Research Project Savidge, J.A. 1987. Extinction of an island forest Report, Guam Aquatic and Wildlife Resources, avifauna by an introduced snake. Ecology Although the initial mortality was FY 1984 Annual Report, Depament of Agri- 68(3):660-668. culture, Guam. Shelton, LC. 1986a. An update on the plight of discouraging, it was by no means unex- Beck, R.E., Jr. 1987. Captive breeding of the Guam's avifauna. Philadelphia Zoo Review pected. Sources of mortality, such as Guam Rail and other native birds. In Research 2(2): 1-5. vehicular kills and cat predation, can be Project Report, Guam Aquatic and Wildlife Shelton, L.C. 1986b. Captive propagation of the Resources, FY 1987 Annual Report, Deprt- Micmesian Kingfisher. Philadelphia Zoo prevented with appropriate measures. ment of Agriculture, Guam. Review 2(2):28-31. Currently, efforts to control feral cats are Beck, R.E., Jr. 1988a. Survey and inventory of the Shelton, L.C. 1986~.Endangered birds of Guam. being made, and alternative sites are native landbirds on Guam. In Research Project Pages 223-225 in American Association of being considered that have less traffic. It Report, Guam Aquatic and Wildlife Resources Zoological Parks and Aquariums 1985 Annual FY 1988 Annual Report, Department of Agri- Proceedings. is less obvious what measures are neces- culture, Guam. Sheppard, C. 1985. Silence on Guam. Animal sary to create a wild population in which Beck, R.E., Jr. 1988b. Captive breeding of the Kingdom 88(4):35-38. encounters between individuals will be Guam Rail and other native bids. In Research U.S. Fish and Wildlife Senice. 1984a. Endan- frequent enough for mating. Our plans Project Report, Guam Aquatic and Wildlife gered and threatened wildlife and plants; Deter- Resources, FY 1988 Annual Report, Depart- mination of endangered status for the Guam rail. call for additional releases of Guam rails ment of Agriculture, Guam. Federal Register 49(71):14354-14356. on Rota over the next several years in Be&, R.E., Jr., and J. A. Savidge. In press. Recov- U.S. Fish and Wildlife Service. 1984b. Endan- both the dry and rainy seasons until a ery plan for the endangered native forest birds of gered and threatened wildlife and plants; Deter- breeding population is established. The Guam and Rota, Mariana Islands. U.S. Fish and mination of endangered status for seven birds Wildlife Service, Honolulu, Hawaii. and two bats of Guam and the northem Mariana experience gained from these releases Conway, W.G. 1980. An overview of captive Islands. Federal Register 49(167):33881- will accomplish several important goals: propagation. Pages 199-208in M.E. Soule and 33885. the Guam rail will have been returned to B.A. Wilcox, eds. Conservation Biology. Sin- U.S. Fish and Wildlifeservice. 1989. Endangered the wild at the earliest possible moment auer Associates, Boston, MA. 395 pp. and threatened wildlife and plants; Determina- Derrickson, S.R. 1986a. A cooperative breeding tion of experimental population status for an to a snake-free environment; a source of program for the Guam Rail (Rallm owstoni). introduced population of Guarn Rails on Rota in wild rails for possible future transloca- Pages 233-240 in American Association of the Commonwealth of the northem Mariana tion to Guam will be established on Rota; Zoological Parks and Aquariums 1985 Annual Islands. Federal Register 54(208):43%6- and release techniques perfected on Rota Proceedings. 43970. Demckson, S.R. 1986b Captive propagation of Witteman, G J. and S.L. Pimm. In prep. The risk will be available for future use on Guam the Guam Rail - An update. Philadelphia Zoo of extinction: lessons from introduced species. when Boiga irregularis is controlled Review 2(2): 19-23. there. Foose, TJ., and J.D. Ballou. 1988. Management of small populations. Int. Zoo Yearbook 27%- 41. Gregory J. Witteman is a Ph.D. student in the Acknowledgements Fritts, T.H. 1988. The brown t~esnake, Boiga Graduate Program in Ecology at the University of irregularis, a threat toPacific Islands. U.S. Fish Tennessee, Knoxville, Tennessee, U.S.A. 37996- and Wildlife Service, Biol. Rep. 88(31):1-36. 1610; Robert E. Beck, Jr. is a Wildlife Biologist in The Guam Rail Introduction Projea is a joint Gtiffith, B., J.M. Scott, J.W. Carpenter, and C. the Aquatic and Wildlife Resources Division, project of the Guam Aquatic and Wildlife Re- Reed. 1989. Translocation as a species conser- Department of Agriculture, P.O. Box 2950, Agana, sources Division, the American Association of vation tool: Status and strategy. Science Guam, U.S.A. 96914;StuartL Pimm is a Professor Zoological Parks and Aquariums, the Graduate 245:477-480. in the Department of Zoology and the Graduate Program in Ecology at the University of Tennes- Haig, S.M., J.D. Ballou, and S.R. Demckson. Program in Ecology at the University of Tennes- see, the Commonwealth of the Northem Mariana 1990. Management options for presening see, Knoxville, Tennessee, U.S.A. 37996-1610 Islands Department of Natural Resources, the U.S. genetic diversity: Reintroductionof Guarn Rails and at the Centre for Population Biology, Imperial Fish and Wildlife Service, and the Conservation to the wild. Conserv. Biol. 4(3):290-300. College at Silwood Park. Ascot, Berks SLS 7PY, and Research Center of the Smithsonian Institu- Hedrick,P.W.,P.F. Brussard,F.W. Allendorf, J.A. England; and Scott R. Demckson is Curator of tion. It has been supported by funds from Wildlife Beardmore, and S. Onack. 1986. Protein Ornithology at the National Zoological Park, Conservation International, the World Wildlife variation, fitness, and captive propagation. Zoo Conservation and Research Center, Fmt Royal, Fund, Intemational Council for Bird Preservation, Biology 5:91-99. Virginia, U.S.A. 22630. Wildlife Preservation Trust International, the Jenkins, J.M. 1979. Natural history of the Guam

Vol. 8 No. 1 Endangered Species UPDATE 39 Peregrine Falcon Recovery

by Tom J. Cade

associated with badly over- (However, a single pair apparently held grazed sheep pastures on right through the 1970s in southern (Ratcliffe 1980). It was, Quebec.) By 1970, peregrines had also however, the heavy use - disappeared from the northern Great beginning in the late 1940s Plains of the Dakotas, Montana, and -of a new, potent class of Alberta, where a sizeable population had chemicals, the organo- nested on steep cut-banks along major chlorine pesticides, that rivers. A few pairs held onto their eyries soon resulted in the un- and continued to breed at a greatly re- precedented declines in duced rate in the Rocky Mountains of peregrine populations re- Colorado, southern Utah, western New corded in the 1950s Mexico, and northern Arizona, although throughout Europe, the they vanished from thenorthem Rockies. United States, and Canada In California, numbers were reduced - (Hickey 1969, Cade et al. especially along the coast -to about ten 1988). percent of the original population of The two main culprits some 200 or more pairs by the early were contaminants in the 1960s. Acomparable declineoccuned in prey eaten by falcons: di- Oregon, again along the coast and along eldrin (HEOD) acting di- the Columbia River where 13 known rectly to increase mortality, eyries were abandoned. Events in Wash- and DDT (DDE) acting in- ington were not well chronicled except in directly through its effects the Okanagan Valley where nesting pere- on eggshells and conse- grines apparently vanished prior to the quent decrease in reproduc- use of DDT (Nelson 1969). Further south tion. The relative irnpor- in Baja California, peregrines disap- tance of these two factors peared entirely from the Pacific side, but Species Decline apparently varied from region to region, they remained at their eyries on islands in and their respective roles in the conti- the Gulf of California. Most cosmopolitan and most admired nent-wide declines in Europe and North The Peale's peregrines (F. p. pealei) of birds, the wide-ranging peregrine fal- America have been much discussed (see in the maritime habitats of the Pacific con (Falco peregrinus) experienced various papers in Cade et al. 1988). Northwest Coast from Washington north major losses in population during the The end result of the combined effects and west through the Aleutian Chain 1950s, 1960s, and early 1970s,primarily of organochlorine toxicity was wide- were never reduced much by pesticides in Eurasia andNorth America (Cade et al. spread abandonment of historical falcon except in Washington, but the decline 1988). Earlier there had been some local eyries. In Europe, the loss of breeders which affected thecontinental peregrines and regional reduction in the number of ranged from complete extirpation for eventually extended northward into the breeding pairs owing to climatic or other some regional populations (e.g., the taiga and tundra regions of Canada and environmental influences, e.g., the loss tree-nesting peregrines in Germany, Po- Alaska. By the early 1970s, local to of a sparse tree-nesting population along land, and the Baltic states), to reductions regional declines amounting to 50 to 75% the Mississippi River prior to 1900 of 50 to 75% or more of original numbers of the original number of breeders had (Hickey 1942, 1969), the disappearance in the British Isles and other parts of been recorded in northern districts right of peregrines and their replacement by Europe. across the continent (see various papers prairie falcons (Falco mexicanus) in the In North America, nesting peregrines in Cade et al. 1988). intermountain region of the Pacific completely disappeared from the Missis- The low point in the number of breed- Northwest in association with a pro- sippiRivereastward to the Atlantic in the ers remaining in North America was longed period of dry years beginning in United States, and from most of contigu- reached around 1975 (Kiff 1988). It can the 1920s (Nelson 1969). and the loss of ous southern Canada- in a region where be conservatively estimated at 30 to 50% about 100 eyries in the western Scottish an estimated 400 to 450 pairs formerly of the pre-DDT population of approxi- highlands owing to a reduced prey base nested (Hickey 1942, Kiff 1988). mately 10,000 adult pairs (see Figure 1).

40 Endangered Species UPDATE Vol. 8 No. 1 Captive Breeding and the next ten years the following private sity of Saskatchewan, Saskatoon (Lynn Reintroduction Efforts breedersmadeirnportantcontributionsto Oliphant). In addition, two major pro- the development of breeding methodol- grams also became established in Eu- In response to these unprecedented ogy in North America: Frank Beebe, rope: one in West Germany under the population changes, the U.S. Fish and Lany Schrarnm, Heinz Meng, James direction of Christian Saar, in association Wildlife Service (USFWS) first listed the Enderson, John Campbell, R. Wayne with the University Falkenorden; and anatum subspecies as endangered in Nelson, Robert Berry, and Lester Boyd. one in Sweden coordinated by Peter 1969 under the old act; the tundn'us sub- In the same period the following institu- Lindberg at the University of Goteborg, species, newly named by White (1968), tional operations got underway: the in association with the Swedish Society was listed as endangered in 1970. The Canadian Wildlife Service facility at Fort for the Conservation of Nature. entire species has also been listed on Wainwright, Alberta, under the direction Falcon breeders worked out the main Appendix I of CITES since 1975. In of Richard Fyfe; the USFWS's endan- techniques for propagating peregrines addition, use of DDT was greatly cur- gered species effort with beach-trapped from 1965 to 1975 (Cade andFyfe 1978, tailed in the United States in 1972, fol- peregrines at Patuxent, Maryland; and Cade 1980, Weaver and Cade 1983). It lowed soon afterward by dieldrin. The Peregrine Fund, Inc., first at Cornell was learned, for example, that wild- Interest in breeding peregrines in University under the direction of Tom caught immature or adult falcons could captivity as a tool for their preservation Cade, later also atFort Collins, Colorado, seldom be induced to breed in captivity, and reintroduction developed from the in cooperation with the Colorado Divi- while hand-reared nestlings often could Madison Peregrine Conference in 1965, sion of Wildlife, and finally permanently be, either by the mating of paired males when it was fust realized that the species located at the World Center for Birds of and females or with the aid of artificial was in serious trouble in both North Prey in Boise, Idaho, under the direction insemination, depending on how the fal- America and Europe (Hickey 1969). A ofWilliam Burnham. Someotherinstitu- cons were treated during their formative post-conference meeting of falconers tional programs joined in later: the Santa periodof socialization. Doubleand triple and biologists resulted in the formation Cruz Predatory Bird Research Group at clutching or the sequential removal of of the Raptor Research Foundation, Inc., the University of California, Santa Cruz, single eggs proved to be effective methods for greatly increasing the productivity of captive females. Artificial incubation became the standard method for the development and hatching of eggs. A ' variety of feeding methods proved effective in rearing the chicks to maturity, but most commonly lab technicians feed them artificially by hand for the first ten to fourteen days, after which they are placed in the care of parent falcons for the remainder of their development and

= EUU~united smtel growth to flying age. About 90 to95% of all chicks hatched can be successfully ' bhlMomairu/Gat BuW Gnml Merdco/ reared by this method. Pacific SouJlwat By 1975, sufficient numbers of cap-

= &la ~lif~/ tively produced peregrines were being c~plh4a~nlud MC~CO reared so that experimental releases could begin in order to learn the best methods for re-establishing these pere-

FlGURE 1. bdrmted numbera d historical ~edne ~roria(uarr)ing grines in the outdoors. That summer, The c~dnl")lnd cumnc (I~CC1985) breeding Peregrine pin in Nonh Peregrine Fund released 16 young by Amcrla. H-historial rerritoria (11-7000-10.000); C-cment con. Bdpin (n-1153t). hacking at five sites in the eastern United Reprinted with permission of Cade et al. 1988. States. Twleve of these birds were re- sighted at or near their release sites a vear as an umbrellaorganization to coordinate under the direction of Brian Walton and la(a (Cade and Dague 1975,1976). he efforts to obtain suitable falcons for closely associated with The Peregrine number of released falcons steadily inbreeding, to establish bonafide breeding Fund; the Raptor Center at the University creased to the current level of several projects, and to disseminate information of Minnesota (Patrick Redig and Hani- hundred per year for all of North Amer- quickly on methods of breeding (Cade son Tordoff), the Macdonald Raptor ica, about 200 of which are released by 1988). From the outset it was clear that Research Center of McGill University The Peregrine Fund, Inc. this effort would involve both institu- (David Bird), and the Saskatchewan Co- In all, more than 3,500 peregrines tional programs and private projects. In operative Falcon Project at the Univer- have now been released in the United

Vol. 8 No. 1 Endangered Species UPDATE 4 1 States and about 1,000 in Canada. Three I I methods have been used: hacking; fostering to wild peregrine parents; and, to a Status of Peregrine Falcon Recovery 1989 lesser extent,cross-fostering, mostly into the nests of prairie falcons (Sherrod et al. 1981, Barclay and Cade 1983, Cade et al. 1988, Linthicum 1989). Each method has proved to be about as equally effective as the others, although there are some reservations about cross-fostering.

Species Recovery

Peregrine populations in North Amer- ica have been increasing dramatically since the late 1970s,both by natural processes and by artificial restoration (Cade et al. 1988). The efficacy of captive propagation and reintroduction must be evaluated within the overall context of this recovery (see Figure 2). wd~baudma&b*n~at~mum~olo~rmyoccu In North America, northern boreal I and arctic nesting pop~lations have Reprinted with permission of the Peregrine Fund. shown the greatest natural increases since the low point reached around 1975. In the western United Srates, there are peregrines extending down the Labrador Some long-studied populations are now currently (1990) about 500 occupied coast suggests the likelihood that there about twice as large as they were in for- eyries, but the rate at which new pairs are will soon be an exchange of individuals mer decades. For example, along the being found each year makes this figure between this wild population and the Colville River in Arctic Alaska where very ephemeral. It is uncertain to what released falcons which have been ex- 32 to 36 pairs nested in the 1950s, now extent this number reflects the real in- panding slowly up along the Gulf of St. there are more than 50 pairs; along the crease as opposed to the discovery of Lawrence. upper Yukon in Alaska where 15 to 16 long-used sites previously unknown. pairs nested in earlier years, there have However, no one doubts that an increase Uniqueness of Re1nt rod uced been around 30 pairs in recent years has been underway for a decade or more. Populatl~ns (Ambrose et al. 1988); and the entire Releasedperegrines, identifiedby bands, reach of the Yukon in Alaska had more occur at 100 to 150of theseeyries,mostly Thus, approximately one-fourth of than 125 pairs in 1990 (P. Bente, in California, Colorado, northern Utah, the original nesting population has been per. comm.). The population nesting andtheGreater YellowstoneEcosystem, restored to this eastern region of North around Sondre Stromfjord in western where they have contributed substan- America through the use of captively Greenland has steadily increased from tially to peregrine recovery. Indeed, produced peregrines for reintroduction. five to six pairs found at 21 known therewerenoknown breedingpairsin the As Cade, Redig, and Tordoff (1989) eyries in 1974-1975, to 43 pairs at 67 Greater Yellowstone Ecosystem when havediscussed,therestoredpopulationis cliffs in 1987 (Mattox and Seegar 1988), The Peregrine Fundbegan reintroduction not an exact ecological or genetic and the population has continued to in- therein 1980. Yetby 1990, there were28 duplicate of the original. In particular, crease since (7'. Maechtle, per, comm.). occupied eyries (W. Heinrich, per. the reintroduced peregrines so far have Similar increases have occurred in Can- comm.), and it is likely that there are been unable to reclaim the formerly ada, and the overall increase in the others to be discovered in this rugged, used nesting habitat along the major, breeding populations of boreal and arctic mountainous region. lowland rivers such as the Mississippi regions has been especially well- In eastern North America south of the and Susquehana because of predation reflected by the number of fall migrants boreal forest, there are now more than by great homed owls (Bubo virginianus). counted and trapped at Cedar Grove, 100 re-established pairs of peregrines in On the other hand, the released falcons Wisconsin, Assateague Island, Mary- the Maritime Provinces of Canada, have learned to use a variety of novel lanwirginia, and Padre Island, Texas southern Quebec, and Ontario, in 11 nesting biotypes that were seldom used (see relevant papers in Cade et al. 1988). states from Maine to South Carolina and by the former "duck hawks" - these These records indicate that the northern in five mid-western states. As far as is include special nesting towers in migratory populations nesting above known, these are all released falcons or coastal salt marshes, bridges, de- 55 degrees N must be approaching their progeny, although the recent dis- commissioned ships, and skyscrapers in 10,000 pairs. covery ofa substantial number ofnesting major cities.

42 Endangered Species UPDATE Vol. 8 No. 1 Urban environments seem to be espe- program to repopulate the northern retum to the wild. Int. Zoo Yearbook 20:23-35. Cade, T.J. 1988. The breeding of peregrines and cially favored by released peregrines, Great Plains. other falcons in captivity: an historical sum- and the number nesting in cities has in- There can be no doubt that restricting mary. Pages 539-548 in TJ. Cade et al., eds. creased rapidly (Cade and Bird in press). the use of organochlorine pesticides has Peregrine Falcon Populations: Their Manage- In 1990, at least the following cities had been the most important action taken to ment and Recovery. The Peregrine Fund. Inc.. Boise, Idaho. 949 pp. Cade, TJ., and D.M. Bird. In press. Peregrine falcons nesting in an urban enviament: A review. Canadian Field-Naturalist. ". . . nowhere within its global range has the peregrine Cade, T.J., and P.R. Dague, eds. 1975. The been able to re-establish breeding populations by natu- Peregrine FundNewsletter3:l-6. Laboratory of Ornithology, Cornell University. ral processes (dispersal of surplus birds) where it was Cade, T.J., and P.R. Dague, eds. 1976. The totally extirpated by pesticides in regions measuring Peregrine Fund Newsletter 4: 1-12. Laboratory of Ornithology, Comell University. from hundreds of thousands to millions of square kilo- Cade, TJ., and R.W. Fyfe. 1978. What makes meters. .. It is precisely in such regions where captive peregrine falcons breed in captivity? Pages 25 1- 262 in S.A. Temple, ed. Endangered Birds: breeding and reintroduction can play their most impor- Management Techniques for Preserving tant role in species restoration." Threatened Species. University of Wisconsin, Madison. Cade. TJ., J.H. Enderron, C.G. Thelander, and C.M. White, eds. 1988. Peregrine Falcon nesting or territorial pairs established: benefit the recovery of this species - Populations: Their Management and Recovery. New York topped the list with nine to both in North America and in Europe. The Peregrine Fund, Inc., Boise, Idaho. 949 pp. Cade, TJ., P.T. Redig, and H.B. Tordoff. 1989. ten pairs, then Minneapolis-St Paul Captive breeding and reintroduction, Peregrine falcon restoration: Expectation vs. with five pairs, followed by Los however, have also played important reality. Loon: 61: 160-162. Angeles, San Francisco, Long Beach, roles in restoring populations in regions Hickey, J.J. 1942. Eastem population of the duck San Diego, Nampa, Calgary, Edmon- where peregrines were totally extirpated, hawk. Auk 59:176:204. Hickey, JJ., ed. 1969. Peregrine Falcon Popula- ton, Winnipeg, Saskatoon, Ottawa, Que- and in speeding the recovery of remnant tions: Their Biology and Decline. University of bec City, Montreal, Chicago, Milwau- populations. Similar conservation initia- Wisconsin Press, Madison. kee, Detroit, Toledo, Cleveland, Pitts- tives can be widely applied to other birds Kiff, L 1988. Commentary - Changes in the burgh, Boston, Springfield, Philadel- of prey as needed, and indeed, to many status of the peregrine falcon in North America: An overview. Pages 123-140 in TJ. Cade et al., phia, Atlantic City, Baltimore, and Nor- other kinds of birds and animals as well. eds. Peregrine Falcon Populations: Their Man- folk. Single birds occur in many other As a consequence of the greatly im- agement and Recovery. The Peregrine Fund, cities. proved status ofpopulations,theUSFWS Inc.. Boise, Idaho. 949 pp. Linthicum, J., ed. 1989. Peregrine Falcon Moni- downlisted the tundrius subspecies from toring, Nest Management, Hack Site, and Cross- Conclusion endangered to threatened in 1984, and it Fostering Efforts. Santa CNZ Predatory Bird should soon be a candidate for delisting. Research Group. University of California, Peregrines still remain absent from The time has also come to consider Santa CNZ Mattox, W.G., and W. Seegar. 1988. The Green- one large region of North America: the downlisting or delisting anatum popula- land peregrine falcon survey, 1972-1985, with northern Great Plains of Montana and tions on aregional basis in western states, emphasis on recent population status. Pages 27- Alberta, where they once nested in con- particularly those in California, Arizona, 36 in TJ. Cade et al., eds. Peregrine Falcon siderable numbers on the cut-banks of New Mexico, Utah, and Colorado. It is Populations: Their Management and Recovery. The Peregrine Fund, Inc., Boise, Idaho. 949 pp. major rivers such as the Missouri and indeed gratifying to all who have worked Nelson, M.W. 1969. The status of the peregrine in Saskatchewan and their tributaries. It is so diligently on behalf of peregrine re- the Northwest. Pages 61-72 in JJ. Hickey, ed. interesting to note that nowhere within its covery during the past 20 years to see the Peregrine Falcon Populations: Their Biology global range has the peregrine been able objective of the Endangered Species Act and Decline. University of Wisconsin Press, Madison. to re-establish breeding populations by reaching fulfillment in the case of this Ratcliffe, D.A. 1980. The Peregrine Falcon. T. & natural processes (dispersal of surplus species. A.D. Poyser. Calton, England. birds) where it was totally extirpated by Sherrcd, S.K., W.R. Heinrich, W.A. Bumham, pesticides in regions measuring from J.H. Barclay,and T.J. Cade. 1981. Hacking: A Literature Cited Method for Releasing Peregrine Falcons and hundreds of thousands to millions of Other Birds of Prey. The Peregrine Fund, Inc., square kilometers (e.g., eastern North Ambrose, R.E., R.J. Ritchie, C.M. White. P.F. Fon Collins, Colorado. America, the tree-nesting region of Schempf, T. Swem, and R. Dittrick. 1988. Weaver, J.D., and TJ. Cade, eds. 1983. Falcon Changes in status of peregrine falcon popula- Propagation: A Manual on Captive Breeding. Germany' poland:andbe Bal- tions in Alaska. Pages 73-82 in TJ. Qde et al., The Peregrine Fund, Inc., Ithaca, New York. tic states). It is precisely in such regions eds. Peregrine ~al~Pmlations: Their White, C.M. 1968. Diagnosis and relationships of where captive breeding and reintroduc- ana an em kt and ~ecovew. The Peregrine" the North American tundra-inhabiting pere- tion can play their mos~impomtrole in ~und~ic.,Boise, Idaho. 949 pp. grine flacons. Auk 85179-191. Barclay, J.H., and TJ. Cade. 1983. Restoration of species restoration. The last major effort theperegrinefalconin united States. Tom J. Cade is Director of the Raptor Research and involving peregrines in North America ~bdConservation 1:3-37. Technical Assistance Center, Boise State Univer- should be a joint U.S.-Canadian Cade, TJ. 1980. The husbandry of falcons for sity, 1910 University Drive, Boise, ID 83725.

Vol. 8 No. 1 Endangered Species UPDATE 43 Captive Breeding and Reintroduction Programs

Nene or Hawaiian goose (Ncsockn sondvicenris) Photo by Jeff Black

This photographi is taken of a young male Hawaiian (Nene) goose in Haleakala National Park, recently fitted with a new plastic leg band - inscribed with the letters "W-that can be read with a telescope from 25h. The bird, which is the pgeny of released parents, was cap&ed by State Forestry & Wildlife and National Pa* Service biologists in August as part of an increased initiative in the recovery program. The new initiative includes research and management programs to improve Nene habitat by enhancing feeding opportunities and reducing predators. The major thrust of the program, which includes a public information campaign, is due to begin in 1991 when initial funding is anticipated. . Contads: Dr. Jeff Black, The Wildfowl & Wetlands Trust, Slimbridge, Gloucester, GL2 7BT, United Kingdom; or Dr. Carol Terry, DLNR,Division of Forestry and Wildlife, 1151 Punchbowl St, Honolulu, HI %813, USA.

Arabian oryx (Ory* leucoryx) Photo: The Living Desen

The Arabian oryx originally ranged throughout most of the Near East, i.e., the Arabian Peninsula north to the Syrian Desert and Mesopotamia. Towards the middle of the 19th century, the oryx began to disappear from the northem parts of its range, aided by increased human activity in the desert and the availability of modem firearms. Automobiles and auto- matic weapons were the fmal straw, and wild populations succumbed to severe hunting pressure. Thus, in the 1960s the Fauna Preservation Society, in cooperation with the IUCN Species Survival Commission, initiated a captive breeding program with wild-aught individuals and a few ux, specimens. All oryx presently living in North America and European collections are descendants of these ten founder animals. Captive propagation has been quite successful.

44 Endangered Species UPDATE Vol. 8 No. 1 Prmwalski's horse (Eqw pnewlskii) Photo: Zoological Society of San Diego

Przewalski's horse is the only true species of wild horse. Extinct in the wild, approximately 1,000 animals now exist in captivity. Although capable of interbreeding with domestic horses, the Pnewalski's horse is a distind species having 66 chromosomes (domestic hones have 64). A global management plan group exists that is coordinating the efforts on behalf of zoological parks to cooperate with nations within the historic range of the species to reintroduce it to its historic range in China, Mongolia, and Central Asia.

Keamey's blue star (Amsonia kearneyana) Photo by Frank Reichenbacher

Individuals from private and governmental organizations collaborated to reintroduce Keamev's blue star (Amonia kearneyana) to a site withi its historical range in south central Arizona. Continuing threats to its survival include over-grazing, insect predation, and floods. When the U.S. Fish and Wildlife Service proposed Keamey's blue star for listing as endangered in 1988, only eight individuals in one population remained; today the reintroduced population numbers 136.

Vol. 8 No. 1 Endangered Species UPDATE 45 Captive Propagation in the Recovery of Whooping Cranes by James C. Lewis

Reasons for Decline "K-selected" species is slow; by 1965 the serve as a source of birds to start other wild population was only 65 birds. As a wild populations. The whooping crane (Grus ameri- result, the species was federally listed as Development of a captive flock began cana) probably was most abundant dur- "endangered" in 1967. in the spring of 1966, when a single male ing the Pleistocene. They were found was moved to Patuxent Wildlife Re- from the Arctic Sea to the high plateau of Captive Breeding Program search Center (Patuxent). Earlier studies central Mexico, and from Utah east to in Canada had indicated that a single egg New Jersey and Florida. In the 19th State and provincial agencies are might be removed from each two-egg century, the principal known breeding important contiibutors to recovery ac- clutch without significant harm to the range was in parts of the prairie pothole tions within their areas of responsibility. wild population. Fifty eggs were moved region. A non-migratory population However, the Canadian Wildlife Service from Wood Buffalo National Park to existed in southwestern Louisiana until (CWS) and the U.S. Fish and Wildlife Patuxent (1967- 1974), to start the popu- the late 1940s. Service (USFWS) are the primary agen- lation. Another 56 eggs were transferred Principal factors contributing to the cies implementing the recovery action. (1982-1989) to improve genetic compo- species' decline in recent history are The main concern of the two agencies is sition and to further build the captive believed to be wetland destruction, hunt- the single self-sustainingwild population flock. ing, and shooting for museum speci- which winters along the Gulf Intercoas- Difficulties of captive propagation mens. An estimated 500 to 700 birds tal Waterway, feeding mainly on blue have made development of the Patuxent survived in 1870, but they had disap- crabs and soft-shelled clams. The water- flock a long-term project. Starting with peared from the principal United States way is one of the busiest in the world. A eggs meant waiting five to six years for breeding range by the 1890s. The last barge accident could release petroleum sexual maturity, which was further de- known nesting in southern Canada oc- or chemical products, destroy the food layed by conditions of captivity. Hatch- curred in 1922. The only self-sustaining base, and kill many birds. As a conse- ability was best when eggs were incu- populationnow nests in Northwest Tem- quence, a captive flock was needed to bated by sandhill cranes rather than in tories, at Wood Buffalo National Park. ensure the survival of the species and mechanical incubators. There were These birds winter in and problems formulating adequate diets for near Aransas National the chicks, preventing parasite and dis- Wildlife Refuge on t9e ease infections, providing water and central Texas Gulf Coast space, controlling light and temperature, where the population low preventing access by predators, and de- was 16 birds in 1941. veloping suitable animal husbandry tech- Life history character- niques (Carpenter and Gabel 1984, Der- istics contributed to the rickson and Carpenter 1987). Many of decline and continue to the husbandry techniques developed for hamper recovery of the sandhill cranes were inadequate for the species. Whooping more fragile whooping cranes. cranes become sexually The problems mentioned have been mature as early as age solved to a large degree, but until they three, but on the average were solved, they contributed to losses of begin breeding at age eggs, chicks, or adults to behavioral and five (Kuyt and Goossen physical abnormalities. The biggest 1987). The clutch size is single disease loss was the eastern equine two eggs, but a majority encephalitis outbreak in 1984, when of the successful pairs seven adult and subadult birds died, in- arrive on the winter cluding several of the best breeders. grounds with only a The first eggs were produced by a single young. Longevity captive-reared female in 1975. Since in the wild is believed then the number of females producing to be up to 25 years. eggs has ranged annually from zero to Recovery of such a five. All females have been artificially

46 Endangered Species UPDATE Vol. 8 No. 1 inseminated. In the 1980s, reproductive foster-reared whooping cranes learned cross-fostering, the USFWS is now plan- females on average produced 3.2 fertile the migration routes and stopover sites ning to use captive-reared whooping eggs and fledged 1.4 chicks. The flock traditionally,used by their foster parents cranes to start a second self-sustaining has produced 177 fertile eggs since 1975. @rewien et al. 1989). wild population. The captive-reared Sixty-one fertile eggs were moved to No cross-fostered whoopers, how- birds would be conditioned for release in Grays Lake National Wildlife Refuge for ever, have paired and produced eggs. the wild. Special rearing methods in- use in the attempt to start a wild popula- Since 1975, cross-fostered females of clude rearing the birds in isolation from tion (1976-1984). To illustrate the diffi- age four through ten years have passed humans and acquainting them with natu- culty in raising whooping cranes, the 166 through a spring nesting season on 26 ral foods and conditions (Horwich 1989). fertile captive-produced eggs retained at occasions without breeding. Males and This technique has been successful in Patuxent and the 71 Canadian eggs which females appear to be associating together supplementingthe wild population of the hatched resulted in only 65 birds surviv- frequently enough, during winter and on endangered Mississippi sandhill crane ing in fall 1990. The average mortality, (Grus canadensispulla)which is nonmi- rate among fledged birds has been eight gratory. percent annually. "Reintroducing an endan- The captive flock was split in 1989, and 22 birds moved to International Cross-Posterlng and gered species is a particu- Crane Foundation (ICF) in Baraboo, Reintroduction Efforts larly challenging experi- Wisconsin. This action was prompted mental process when the aftera series of health problems seriously Reintroducing an endangered species species must develop threatened the entire flock at Patuxent. is a particularly challenging experimen- migratory patterns." Flock division also provided an opportu- tal process when the species must de- nity to utilize the crane avicultural exper- velop migratory patterns. Using an inno- tise at ICF. The captive flock now totals vative research approach, Drs. Roderick 65 birds. Drewien and Elwood Bizeau of the Uni- spring migration stopover,to permitpair- The Director of the USFWS and the versity of Idaho suggested across-fostering at the times when pairing occurs in the Director General of CWS have approved ing experiment to restore whooping natural flock. The absence of breeding fieldwork in Florida to determine the cranes to the northern Rockies. They may be due to improper sexual imprint- feasibility of establishing a non-migra- suggested use of the abundant greater ing among the females. tory whooping crane population. Sev- sandhill cranes (Grus canadensis tabida) Dr. Edward 0.Garton, biornetrician enty-seven percent of the deaths or disap- nesting in Grays Lake National Wildlife at the University of Idaho, working with pearances of the fledged cross-fostered Refuge in Idaho as foster parents for Dr.RodDrewien andothers (Garton et al. birds occurred during spring or fall mi- whooping cranes. Preliminary studies 1989),modelled the cross-fosteredpopu- gration. A non-migratory flock would had shown that the typical two-egg clutch lation to predict when it might become not be exposed to migration hazards. could be removed and another single egg self-sustaining. In the model they as- Another advantage is that a non-migra- substituted without the parent sandhill sumed: 1) the cross-fostered females tory population would be more concen- cranes abandoning the nest. would be breeding at the same rate as the trated year-round in a smaller geographic Project objectives included determin- females in Canada, and 2) survival of area. The more restricted movements ing if: 1) sandhill pairs could success- birds in their first year would be similar to would increase the opportunity for asso- fully hatch, rear, and fledge whooping that of first year birds in Canada. Despite ciations between birds that would be crane young from eggs; 2) foster-reared these optimistic and unrealized assump- compatible mates, and increase the op- whoopers would accompany their par- tions, with the future transfer of 30 eggs portunity for egg production. ents to traditional wintering sites and per year, the population would only reach If whooping cranes are released in return to their natal area; and 3) whoop- six breeding pairs after 50 years. "It is Florida, they will be reared in captivity ing cranes would survive and reproduce obvious from all scenarios modelled that and specially conditioned for release in in these habitats. The project began in egg transplants of less than 30 eggs per the wild. Modelling studies suggest that 1975, and egg transfers from Canada year will not suffice to establish a self- a goal of releasing 20 cranes annually for (216) and the captive flock (73) contin- sustaining population in a reasonable at least ten years will increase theproba- ued through 1988 (Drewien et al. 1989). period of time. Natural breeding will be bility of successfully establishing a wild Two hundred and ten eggs hatched, and essential to establish a self-sustaining population. As Demckson (1987) noted, 85 chicks survived to attain flight age. population" (Garton et al. 1989). " . . . the most significant historical factor Mortality rates were excessively high in Fieldwork in the cross-fostering project limiting production has been the small all age categories in the cross-fostered is scheduled to end in summer 1991, size of the captive flock." To correct that population in comparison to the natural when project personnel will concentrate problem, theUSFWS has been endeavor- population which nests in Canada (Gar- on finishing their final contract report. ing since 1988 to build the captive flock ton etal. 1989). The population peakedat The self-sustaining wild whooping with captive-produced eggs and eggs 33 birds in 1985. Thirteen of these birds crane population now contains about 150 from the Canadian population so captive still survive, including four females. The birds. As a consequence of the results in pairs can support the release planned to (continued on bottom of next page)

Vol. 8 No. 1 Endangered Species UPDATE 47 Conservation of Crocodilians: The Release of Captive-Reared Specimens

by F. Wayne King

Reasons for Decline prey to herons, otters, fish, turtles, and The recovery process can be speeded mature crocodiles. From this variety of up somewhat by incubating wild eggs ar- During the 1950s and 1960s, popula- causes, between 50 to 90% of young tificially and rearing the young in captiv- tions of crocodilians all over the world crocodilians are lost during their first two ity for later release back into the wild were depleted by indiscriminate hide to three years. In addition, as young when they are large enough to resist most hunting, killing as vermin, and habitat crocodilians begin to mature, they often natural predators. Such "head starting" loss. During the late 1960s and 1970s, are killed by large males attempting to eliminates the large, early loss to preda- programs were launched in many coun- drive subadults out of their temtories. tors and reduces the overall loss to the ten tries to conserve these depleted and en- percent or less caused by disease. Terri- dangered populations (Bustard 1969, torial males will still attack subadult Cott and Pooley 1972). Some, like the "Under normal circum- crocodiles as they approach mature sizes, American alligator, Alligator mississip- stances probably less but more individuals will reach that size piensis, were sufficiently abundant that class and presumably more will survive. when killing was prohibited, populations than 1 to 2% of wild recovered. However, other species had [crocodilian] hatchlings Reintroduction of the G ha rial become so endangered that simply end- survive to sexual maturity." ing the killing was not enough. The few The gharial, Gavialis gangericus, was adult specimens that remained were so the subject of the first large-scale pro- scattered that breeding was almost non- Under normal circumstances probably gram of captive rearing and reintroduc- existent, and much of what little repro- less than one to two percent of wild hatch- tion for an endangered crocodilian spe- ductive output did occur was lost. lings survive to sexual maturity. Large cies. Up until the 1950s and early 1960s, Each generation a small number of crocodilians require eight to fifteen gharial were abundant in the large rivers young are lost to disease, but many more years to reach sexual maturity, so once a of the Indian subcontinent: the Ganges, are lost to predators. Crocodilian nests population is severely depressed, recov- Brahmaputra, Indus, and Mahanadi riv- are often raided by predators that feed on ery is slow; years pass before the few ers of India, Pakistan, Nepal, Bhutan, and eggs: carnivorous lizards, mongoose, young that survive predators and temto- Bangladesh, as well as the Kaladan and raccoons, bears, vultures, crabs, and ants. rial males can in turn breed and contrib- Irrawaddy rivers of Burma. Because of Hatchlings and young crocodilians are ute to the recovery. indiscriminate killing for their hides, as

(continuedfrom preceding page) begin in Florida in the mid 1990s. Proceedings 1984 Conference on Avian Medi- Cranes at Gray Lakes NWR. Final Report by Aviculture techniques for whooping cine, Association of Avian Veterinarians. Fish and Wildlife Depament, University of Demckson, S.R. 1987. Captive propagation of Idaho. Prepared for U.S. Fish and Wildlife cranes are still being refined. Although whooping cranes, 1982-1984. Pages 377-386 in Service, Albuquerque, NM. 47 pp. ow knowledge is incomplete, captive- J.C. Lewis and J.W. Ziewitz, eds. Proceedings Horwich, R.H. 1989. Use of surrogate parental reared birds are our only practical source 1985 Crane Workshop, Platte River Whooping models and age periods in a successful release of of individuals to release in establishing a Crane Maintenance Trust, Grand Island, NE. hand-reared sandhill cranes. Zoo Biology Derrickson, S.R., and J.W. Carpenter. 1987. 8:379-390. new population. I am confident that Behavioral management of captive cranes - Kuyt, E., and J.P. Goossen. 1987. Survival, age captive propagation and conditioning of Factors influencing propagation and reintroduc- composition, sex ratio, and age at first breeding birds for wild release can ultimately re- tion. Pages 493-51 1 in G.W. Archibald and R.F. of whooping cranes in Wood Buffalo National Pasquer, eds. Proceedings 1983 International Park, Canada. Pages 230-244 in J.C. Lewis and sult in additional self-sustaining popula- Crane Workshop, International Crane Founda- J.W. Ziewitz, eds. Proceedings 1985 Crane tions of non-migratory and migratory tion, Baraboo, WI. Workshop, Plane River Whooping Crane Main- whooping cranes. Drewien, R.C., W.M. Brown, and E.G. Bizeau. tenance Trust, Grand Island, NE. 1989. Whooping Crane Cross-Fostering Ex- periment. Unpublished interim report prepared Literature Cited for the U.S. Whooping Crane Recovery Team. James Lewis is the Whooping Crane Coordinator 10 pp. for the U.S. Fish and Wildlife Service, PO Box Carpenter, J.W., and R.R. Gabel. 1984. Establish- Garton, E.O., R.C. Drewien, W.M. Brown, E.G. 1306, Albuquerque, NM 87103. ment and management of a breeding program Bizeau, and P.H. Hayward. 1989. Survival for captive endangered birds. Pages 237-265 in Rates and Population Prospects of Whooping

48 Endangered Species UPDATE Vol. 8 No. 1 vermin, and for use in folk medicine, as home range of the captive-reared ani- solely to the gharial. In 1975, India well as accidental drowning in fishing mals. With their sudden release into launched companion programs to con- nets, and collection of their eggs for food, totally unfamiliar, albeit natural, sur- serve its other two species of crocodiles: the gharial all but disappeared through- roundings, the young crocodilians set out the mugger crocodile, Crocodyluspalus- out most of its historical range. Prelimi- to find their familiar home range. In tris, and the saltwater crocodile, Croc- nary surveys in the early 1970s indicated marshes or lakes, they may simply wan- odylus porosus. Restocking newly es- that as few as 200 adult gharials might der away. In rivers, they may swim out tablished sanctuaries and suitable habitat survive in India, fewer then 20 in Bangla- into the current and be swept down- with young hatched and reared in captiv- desh, and certainly no more than 20 in Pakistan. An estimated 65 to 70 adults survived in Nepal. The status in Burma remains unknown. So few adults remained that simply protecting them would not soon return populations to their former numbers. Thus, in 1975, the government of In- dia, with technical assistance from the wildlife division of the United Nations Food and Agriculture Organization and funding from theU.N. Development Pro- gramme, launched an all-out effort to save the gharial. Sanctuaries were established to protect the few scattered adults. For example, the Satkosia Gorge Sanctu- ary was declared on the Mahanadi River in 1976, at which time only five gharials Young gharials (Gavialis gangeticus) at the Deori Rearing Centre, National Qlambal Sanctuary, India could be found there. Eggs were col- Photo by R.J. Rao lected from various remnant populations and hatched in government-operated stream. Some of the first gharials re- ity played a majur role in these programs hatcheries. By 1990, a total of 13 rearing leased in India were discovered later as well (Choudhury 1990). By 1990, a stations for gharial were in operation in many kilometers downstream. total of more than 2,000 young muggers India (Choudhury 1990). Young were This search for home is not limited to from 3 1rearing stations were released in reared in captivity for up to three years young crocodilians. In the late 1960s,the dozens of sites. The rearing stations still before being released back into the wild. New York Zoological Park shipped eight hold an additional 12,000juveniles. The By 1989, a total of 1,000 captive- adult American alligators, Alligator mis- species now is breeding in the wild at 10 reared gharials had been released in eight sissippiensis, to Florida for release into to 12 protected sites and in 25 rearing protected areas in India (Choudhury the Everglades National Park. Included stations and zoos. 1990). Half of that total, 550 captive- in the shipment were several animals Similarly, nine rearing stations pro- reared gharials, had been released into measuring about three meters in length duced over 1,200 saltwater crocodiles the Satkosia Gorge Sanctuary, which and weighing over 100 kg. Within two which were released into the Bhitarkan- increased the resident population to 25 weeks of their release, all but two of the ika Sanctuary, where the species now adults in 1988 (Kar 1989). Assuming gators had moved away from the release breeds, and 24 have been released in that the original five gharials resident in sites. Even these two later disappeared. other parts of the country. Relatively few the gorge were among the 25 recent resi- The chance of success can be in- saltwater crocodiles were found in dents, the survival rate for the released creased if the animals being released are Bhitarkanika's mangrove habitat when captive-reared gharials is four percent. maintained for several weeks in pens the sanctuary was declared, but a January More significantly, however, an adult constructed in the natural wetland at the 1988census of the sanctuarycounted433 female gharial began laying eggs in the release site. This allows the animals to saltwater crocodiles, suggesting a 40% Satkosia Gorge in 1984. adapt to a new home range prior to ven- survival rate up to that time. One of the problems hindering the turing out into the natural habitat when reintroduction of wild populations by the the surrounding fence is later breached. Crocodilian Reintroduction release of captive-reared crocodilians is Programs Worldwide the movement of animals away from the Other Crocodilian Reintroduction release site. This is a problem for all Programs in India Conserving crocodilians by captive crocodilians regardless of their habitats, rearing and restocking has not been con- but is especially critical for riverine spe- Since hatcheries and rearing facilities fined to India. Similar "head starting" cies. Prior to their release, the pens in can be used for more than one species at programs are a common feature of many which they were reared constituted the a time, India did not limit its attention crocodilian conservation programs:

Vol. 8 No. 1 Endangered Species UPDATE 49 -, -, Alligator mis- been raised in captivity, and a total of 394 diles were released into the newly de- sissippiensis: In Louisiana, U.S .A,, this gharials released into the Nayarani, clared CaAo Guaritico wildlife refuge program allows American alligator eggs hpti, Kali Gandaki, and Babai rivers (Ayanagiiena 1990). Another release is to be collected from the wild under state (Maskey 1990). A 1990 survey of the planned to supplement the small popula- license for supplementing production on Nayarani revealed 3 1 wild and 40 cap- tion in the Capanaparo River in the San- alligator farms. It also requires farmers tive-reared gharials, representing 18%of tos Luzardo National Park. to return to the state for release back into the wild 30% of the hatchlings or 17%of the four-foot alligators produced fiom those eggs (Joanen 1989, Joanen and "Ranching gives local people a justification for McNease 1990). tolerating the presence of large, aquatic carnivores American,Crocodylus acutus: In Venezuela, a small number of that may threaten humans and livestock, and a motive captive-reared American crocodiles for maintaining wetlands." were released into the Jatira reservoir. Plans are also underway for releasing American crocodiles, produced by the captive breeding program on the Masa- the captive-reared gharials released Phili~~inecrocodile, Crocodylus guaral ranch, into the Cuare National there. mindorensis: Silliman University Ma- Wildlife Refuge (personal communica- Pakistan is scheduled to receive rine Laboratory, Negros, is breeding tion from Andres Seijas and John gharial from the program in India so a Philippine crocodiles to restock pro- Thorbjamarson). captive breeding program can be initi- tected habitat. In a collaborative effort In Colombia, INDERENA, the gov- ated for restocking areas of protected with ~illirnan~niversity,thespecies also ernment agency responsible for wildlife habitat. is being bred in the Gladys Porter Zoo, conservation, requires the five farms li- J&xeletys crocodile, Crocodylus Brownsville, Texas, U.S.A. Offspring censed to breed American crocodiles to moreletii: ZQO Atlanta in Georgia, produced at the zoo will be returned to return five percent of their hatchlings to U.S.A., has been breeding Morelet's Silliman University for release. the government for restocking purposes. crocodiles for more than 15 years. Some In addition to these programs al- Black,Melanosuchus ni- of the captive-produced offspring have ready underway, restocking depleted ger: In August 1990, a total of 25 black been returned to Mexico and Belize for habitat with captive-bred animals could caimans from a captive group on El release into the wild. make significant contributions to the Dorado ranch were released into Nor- Weer crocodile, Crocodylus conservation of several species of croco- mandia lagoon adjacent to the Beni Bio- palustris: Pakistan is planning a pro- dilians: logical Station where they could be program for breeding mugger crocodiles in Cuban crocodile, Crocodylus rhom- tected (Thorbjarnarson 1990). captivity for restocking former habitat. bifer: Two government-operated farms Caiman lati- Nile crocodile, Crocodylus nilot- specialize in breeding Cuban crocodiles rostris: An experimental program is icus: The government of Zimbabwe is- to preserve the species, to serve as a underway in Argentina for captive breed- sues an annual quota for Nile crocodile tourist attraction, and to provide hides ing of broadsnouted caiman for possible eggs collected from the wild to supple- for the market (Ramos Targarona 1989). release. ment production on licensed crocodile The Cuban crocodile disappeared from Common,Caiman crocodi- farms. A percentage of the crocodiles the Lanier Swamp on Isla de la Juventud lus spp.: In Colombia, more than 30 hatched from the wild eggs are available (Isla de Pinos) following the introduc- farms have been established for captive to the government for restocking should tion of Caiman crocodilus (see Com- production of caiman. INDERENA, the they be needed. However, as long as mon Caiman below). If the feral caiman government agency responsible for wild- wild populations are stable or increasing are removed, the swamp could be re- life conservation, requires the farms to no return is required. stocked with Cuban crocodiles from the return five percent of their hatchlings to Orinoco crocodile, Crocodylus in- farms. the government for restocking purposes. termedius: Three centers in Venezuela Siamese crocodik, Crocodylus sia- m,Gavialis gangeticus: In are breeding and rearing the critically mensis: This species is virtually extinct 1978, Nepal established a "head start- endangered Orinoco crocodile in captiv- in Thailand and is extremely rare, if not ing" program in Royal Chitwan National ity for release into protected habitat; the extinct, in Indonesia. However, acaptive Park, site of the largest surviving wild centers at the Masaguaral ranch and the population of approximately 8,000 pure- gharial population. Eggs were collected El Frio Biological Station have been in blood Siamese crocodiles is held by the and hatched in a hatchery, and the young existence for more than a decade, and the Samu~rakanCrocodile Farm and Zoo, reared for later release at one to two center at the Llanos University outside Bangkok (Luxmooreet al. 1985). meters in length (Maskey 1989). By (UNELLEZ) for only about four years. If suitable habitat in Thailand can be 1990, approximately 1,000 gharials had In April 1990,31 young Orinoco croco- identified and adequately protected, the

50 Endangered Species UPDATE Vol. 8 No. 1 farm could supply the necessary animals eggs. However, under the PNG conser- importantly, in providing motivation to for reintroduction. vation program, crocodile ranchers pay governments and local peoples to pro- The release of crocodilians into the landowners the equivalent of U.S. $1.50 tect wild individuals and habitat. wild has not always been done for conser- for each crocodile egg collected on their vation. Some releases of exotic croco- land. In addition to the thousands of Ltterature Cited dilians into habitat outside their ranges dollars they earn from selling eggs, the have caused ecological problems for landowners are given one hen egg for Ayamgiiena, J. 1990. An update on the recovery program for the Orinoco crocodile. Crocodile native crocodilians: each crocodile egg collected. The money Specialist Group Newsletter 9(3):16-18. Common,Caiman crocodi- pays the landowners for protecting Bused, R. 1969. Cdilians of the world: [US: Although not part of a conservation crocodiles in the wild, and the hen eggs Summary of the present position. Pages 3 13- program, the deliberate release or escape replace crocodile eggs on the dinner 320 in World Wildlife Yearbook 1969. World Wildlife Fund, Morges, Switzerland. of pet caimans has resulted in the estab- table. Annual censusing indicates that Choudhury, B.C. 1990. Indian crocodile conser- lishment of feral populations of common the wildpopulation isincreasing,and that vation situation report: Action Plan for the caiman in south Florida, U.S.A., Puerto collecting crocodile eggs solely for 1990s. Pages 337-345 in Crocodiles: Proceed- Rico, and Cuba -see also Cuban croco- human consumption has all but disap- ings of the 10th Working Meeting of the Croco- dile Specialist Group, Vol. 2. IUCN, Gland, dile above. peared. Switzerland. 345 pp. bwGuinea, .. . Crocodylus Similarly, in the U.S.A., conservation Con, H., and A.C. Pdey. 1972. Crocodiles: The novaeguineae, and -, programs in Florida, Louisiana, and Status of Crocodiles in Africa. IUCN Publica- Crocodylus mindorensis: Japanese oc- Texas allow landowners to earn money tion New Series, Suppl. Pap. 33. IUCN, Morges, Switzerland. 98 pp. cupation troops during World War I1 from the sale of alligator eggs or hatch- Joanen, T. 1989. Louisiana alligator fanning introduced New Guinea and Philippine lings collected from their wetlands. program. Crocodile Specialist GroupNewslet- crocodiles into Palau (Kimura 1968). Where zoning and environmental laws ter 8(0ct-Dec):26-27. The introduced species now compete allow them to do so, landowners could Joanen, T., and L. McNease. 1990. Alligator farming programs in Louisiana. Pages 1-10 in with the small native population of salt- earn more money by turning their wet- Crocodiles: Proceedings of the 9th Woking water crocodile, Crocodylus porosus. lands into agricultural lands or into hous- Meeting of the Crocodile Specialist Group. ing developments. However, many land- IUCN, Gland, Switzerland. Vol. 2. 380 pp. Crocodilian Ranching owners enjoy the out-of-doors, and the Kar, S. 1989. Crocodile conservation programme in Orissa: An overview. Crocodile Specialist money they collect from allowing the Group Newsletter 8(bct-Dec):16-18. With few exceptions, "head starting" harvest of alligator eggs or hatchlings is Kimura, N. 1968. Bulletin of Crocodiles in the programs involving crocodilians are an sufficient excuse to maintain the wet- Palau area. Atagawa Tropical Garden and Al- adjunct to efforts to conserve wild popu- lands. ligator Farm Bulletin No. l. 49 pp. fi Japa- nese.] lations through ranching. Ranching in- This money that ranchers and land- Luxmoore, R., J. Bando, S. Bmad, and D. Jones. volves protecting adult breeders in the owners in PNG, the USA, and elsewhere 1985. A Directory of Crocodilian Farming wild and harvesting their eggs or hatch- earn makes them less tolerant of poachers Operations. NCN Wildlife Trade Monitoring who would overexploit the resource or of Unit, Cambridge, U.K. 204 pp. lings torear in captivity for later commer- Maskey, T. 1989. Movement and survival of cial slaughter. Money earned from the developers who would destroy the habi- captive-reared gharial, Gavial~gange~icus,in sale of hides and meat benefits the croco- tat. Crocodilians are rather like the the NarayaniRiver,Nepal. Univ. Florida Ph.D. dilerancher, and ranching in turn benefits mythical "goose that laid the golden Dissertation. 187 pp. Maskey, T. 1990. Update on gharial releases. crocodilian conservation. egg." In the past, crocodilians, like the Crocodile Specialist Group Newsletter As ranching depends on wild eggs or goose, were destroyed by poachers and 9(3):12. hatchlings for its stock, it can only be hunters who overexploited the resource. Ramos Targarona, R. 1989. Crocodile manage- sustained as long as wild adult breeders Ranching is changing that. Now in order ment in Cuba. Crocodile Specialist Group Newsletter 8(0ct-Dec):22. are sufficiently abundant to support the to exploit the resource on a sustainable- Thorbjamarson, J. 1990. Bolivian Melonosuchus harvest. This forces ranchers to be con- yield basis, ranchers and landowners adventure. Crocodile Specialist Group News- cerned about the wild populations. In must protect wild crocodilians and their lener 9(3):14-15. contrast, farming, which involves the habitat. hatching of eggs laid in captivity from Obviously, headstarting and captive matings of captive parents, is not depend- breeding of crocodilians for conservation ent on the wild population and does not purposes differgreatly in their objectives directly benefit the wild resource. from ranching for commercial profit. F. Wayne King is Deputy Chairman of the IUCNI Ranching gives local people a justifi- However, some of the same techniques SSC Crocodile Specialist Group at the Florida cation for tolerating the presence of large, apply, and all can be used to benefit wild Museum of Natural History. Gainesville, FL aquatic carnivores that may threaten populations. Thus, in the case of croco- 3261 1, U.S.A. humans and livestock, and a motive for dilian conservation, captive breeding maintaining wetland habitat. For ex- combined with economic incentives has ample, villagers along the Sepik River of been quite effective in actually increas- Papua New Guinea (PNG) eat crocodile ing wild populations, and perhaps more

Vol. 8 No. 1 Endangered Species UPDATE 5 1 Captive Breeding and Reintroduction: Recovery Efforts for the Virgin Islands Boa, Epicrates monensis granti by Peter J. Tolson

History (Pregill 1982), eustatic events and the These efforts were coupled with a com- subsequent genetic isolation of several prehensive reproductive research pro- The Virgin Islands boa, Epicrates populations, and the stochastic processes gram using congeners as models which monensis granti, was first discovered by associated with the fragmentation of a established the proximate environmental Major Chapman Grant on the island of once contiguous range have also doubt- and social factors critical to reproduction Tortola, British Virgin Islands (Grant less contributed to the decline of this in this species (Tolson andTuebner 1987, 1932). Erroneously described by Stull taxon (USFWS 1986, Tolson 1988). Tolson 1989). (1933) as a subspecies of the Puerto Rico The reproductive program has thus far boa, the snake was designated as endan- Captive reeding and been successful: 48 offspring have been gered under the U.S. Endangered Species Reintroduction produced by ten founders at the TZS, of Act listing for E. inornatus in 1973. which 40 have survived. Representatives Taxonomic work by Sheplan and In 1985, in cooperation with the of two distinct demes - La Cordillera, Schwartz (1974) later demonstrated that USFWS, the Departmento de Recursos PuertoRico,and St. Thomas,U.S. Virgin the snake was actually asubspeciesof the Naturales de Puerto Rico (DRN),and the Islands -are now maintained in captiv- Mona Island boa, E. m. monensis. In Division of Fish and Wildlife, U.S. Vir- ity. As each deme exhibits a distinct col- 1980, the legislative status of the snake gin Islands (USVIFW), the Toledo Zoo- oration, it was decided to maintain deme was finally clarified by the U.S. Fish logical Society (TZS) initiated a captive integrity in the captive snakes and man- and Wildlife Service r iage- them as two sepa- (USFWS), and it re- rate populations, i.e., ceived endangered spe- individuals captured cies protection under on La Cordillera are the correct nomencla- not crossbred with ture (USFWS 1980). stock originating from Apparently never St. Thomas. Unfortu- very common, its ex- nately, the two male tremely disjunct distri- and three female boas bution as a Puerto Rico collected from St. Bank endemic (Nellis Thomas have not yet et al. 1983, Mayer and reproduced; two of Laze11 1988) provides these females are evidence for a long his- sexually immature and tory of extirpation and the third expelled in- decline on numerous fertile ova after her islands and cays (low first reproductive at- islets of coral or sand) . . tempt in captivity. The hgin Islands boa juveniles born at the Toledo &logical ~ardens Photo by Art ~eber' of the Bank since the rarity of snakes on St. Pleistocene. It is believed that a variety breeding program for this species. The Thomas makes them very difficult to ob- of factors have contributed to the present focus of the TZS program is the produc- tain, and the actual size of the population dire status of this snake. Habitat destruction of snakes for reintroduction efforts. there is unknown. tion was among the earliest of perils, as The basis for a management strategy was To aid in captive management, the widespread areas of the Virgin Islands developed under the auspices of the TZS maintains a regional studbook for were denuded and periodically burned USFWS Recovery Plan (USFWS 1986), the species under the auspices of the for the cultivation of sugar cane. The in- and included two years of field research American Association of Zoological troduction of exotic predators such as the on the ecology and distribution of the Parks and Aquariums (AAZPA). In roof rat, Rattus rattus, the house cat, snake in Puerto Rico and the Virgin Is- addition, apetition to establish a Species Felis catus, and the Indian mongoose, lands. Data continues to accumulate SurvivalPlan for this snake has been sub- Herpestes auropunctatus, also had seri- from wild populations through an ongo- mitted to the AAZPA. (Species Survival ous adverse effects on boa populations. ing mark-and-recapture study of over Plans are cooperative management pro- Climatic changes during the Pleistocene 200 marked individuals on La Cordillera. grams in which several zoos pool re-

52 Endangered Species UPDATE Vol. 8 No. 1 sources to develop captive breeding some concern, as completely random cooperation in recovery efforts for this strategies for endangered wildlife.) matings within the captive population species. With continued commitment may be difficult to achieve due to the from these agencies, it is believed that Crltlcal Factors Affecting reluctance or inability of some potential future management actions will be suc- Reintroduction Success founder snakes to reproduce. Two males cessful as well. captured in 1986 from the wild on La Reintroduction is perceived to be a Cordillera have yet to produce offspring Acknowledgements reasonable strategy in the recovery of this in captivity, despite numerous reproduc- species because of the availability of tive opportunities. No genetic analysis The author wishes to acknowledge Conservation Pmjeu support from the Institute of Museum Serv- protected, relatively undisturbed cays exists for any of the wild populations; the ices from 1987 to the present (grants IC-70095-87 which exhibit a number of community degree of heterozygosity within popula- and IC-90453-89), which have made field research attributes identified by principal compo- tions and the degree of relatedness be- on Epicrates monensis possible. nents analysis as typical of habitat where tween isolated populations is completely this species occurs (Tolson 1988). unknown. Until these data are generated, Literature Cited However, any reintroductions to the wild genetic management will be haphazard. will involve only those individuals origi- But there are several attributes of this Chandler C.R., and PJ. Tolson. 1990. Habitat use by a boid snake, Epicrates monensis, and its nating from a source population which is program which should ultimately result anoline prey,AnoliF cristatellus. J. Herpeiology closest to the release site. A lack of in its eventual success. There is a strong 24(2):151-157. correlation between availability of cap- base of research on which to base man- Grant, C. 1932. Herpetology of Tortola; notes on tive-bred individuals from specific lo- agement decisions (see Tolson 1988, Anegada and Virgin Gorda, British Virgin Is- lands. J. Dept. Agr. Puerto Rico 16:339-346. calities and site preparation has been the Tolson and Tuebner 1987, Chandler and Mayer, G.C., and J.D. Lazell, Jr. 1988. Distribu- greatest barrier to immediate reintroduc- Tolson 1990). The environmental and tional records for reptiles and amphibians from tion attempts. Captive-born offspring social parameters necessary to induce the Puerto Rim Bank. Herpetological Rev. from St. Thomas founders are not yet reproduction can be duplicated fairly 19(1):23-24. Nellis, D.W., R.L Norton, and W.P. Mach. available for release even though there easily in captivity and the husbandry of 1983. On the biogeography of the Virgin Is- are two rat-free cays in the Virgin Islands the species is not difficult. Spacerequire- lands boa, Epicrates monensis grad. 3. Herpe- that could serve as reintroduction sites. ments are minimal, and the survivorship tology 17(4):413-417. In fact, in the opinion of the author, the of captive neonates is extremely high Pregill, G.K. 1982. Late Pleistocene herpetofau- - nas from Puerto Rib. Univ. Kansas Mus. Nat greatest single problem to overcome in exceeding 83% at the TZS. Although E. Hist Misc. Publ. No. 7 1. 72 pp. any release program for this species is the monensis reproduces biennially, longev- Sheplan, B.R., and A. Schwaltz 1974. Hispanio- eradication of exotic mammals from ity may exceed 20 years, lifetime female lan boas of the genus Epicrates (Serpentes, potential reintroduction sites. Rat andcat Boidae) and their Antillean relationships. Ann. reproductive output is potentially very Camegie Mus. 45(5):57-143. control programs have been initiated at high, and generation times can be ex- Stull, O.G. 1933. Two new subspecies of the two potential reintroduction sites on off- tended topreserve genetic diversity in the family Boidae. Occas. Pap Mus. Zool. Univ. shore cays, one near Puerto Rico, and captive population. In addition, demo- Michigan, No. 267:l-4. Tolson. PJ. 1988. Critical habitat, predator pres- another in the Virgin Islands. Snakes graphic research has demonstrated that sures, and management of Epicrates monenris cannot be released at these localities until this species can exist in high densities on on the Puerto Rico Bank: A multivariate analy- control programs have been proven to be small islands. With a probable high sis. U.S. Dept Agr. Forest Sew. General Tech. successful. Recolonization, deliberate or degree of inbreeding in several wild Rpt RM-166:228-238. Tolsm. PJ. 1989. Breeding the Virgin Islands accidental, of Rattus or Felis to reintro- populations, the genetic load of deletri- boa, Epicrates monensis grad, at the Toledo duction sites could doom any reintro- ous alleles is apparently not high. Zoological Gardens. Inter. Zoo Yrbk. 28:163- duced populations. A third suitable cay, Several additional activities are un- 167. Steven Cay, already cleared of rats by the derway to prepare for reintroduction at- Tolson. PJ., and V.A. Tuebner. 1987. Ihe role of social manipulation and environmental cycling USVIFW in 1983, is also suitable for tempts. Animals slated for release are fed in propagation of the bid genus Epicrates: reintroduction. The site identified by the live adult male crested anoles, Anolis Lessons from the field and laboratory. Pages author (Tolson 1988) on Cayo Icacos as a cristatellus, to insure the snakes are able 606-613 in Amer. Assoc. Zoological Parks and primary candidate for reintroduction was to capture and feed upon their natural Aquariums Reg. Conference Proc. A.A.ZP.A.,Wheeling, West Virginia, 1987. severely damaged by Hurricane Hugo in prey. Veterinary protocols for screening U.S. Fish and Wildlife Service. 1980. Status of September 1989, and an alternate site on of animals to be released are being devel- Virgin Islands boa clarified. Endangered Spe- La Cordillera had to be located. oped at the TZS. The initial releases will cies Tech. Bull. 512. probably involve radiotracking of indi- U.S. Fish and Wildlife Service. 1986. Virgin Islands TmBoa Recovery Plan, U.S. Fish and Evaluation of Reintroduction viduals implanted with transmitters to Wildlife Service, Atlanta, Georgia. 23 pp. Strategies assess to what degree captive-raised individuals duplicate the behaviors of There are other problems which must wild snakes. be overcome if reintroduction is to suc- Federal, Commonwealth, Tem torial, Peter J. Tolson is the Conservation Biologist at the Toledo Zoological Gardens, 2700 Broadway, ceed in this species. Genetic manage- and Society management personnel Toledo, Ohio 43609. ment of the population may also be of have a five-year history of successful

Vol. 8 No. 1 Endangered Species UPDATE 53 Release and Translocation Strategies for the Puerto Rican Crested Toad, Peltophryne lemur by Robert R. Johnson introduction Although the rarity of this species has stochastic events and population demo- been questioned since its rediscovery in graphics. In fact, skewed sex ratios, Amphibians, with both aquatic and 1967, no one has demonstrated that there reported by Rivero et al. (1980), may terrestrial stages in their life cycles, are exists any more than 25 individuals in the hasten the extirpation process. particularly sensitive to environmental northeast coastal plane near Quebradil- None of the northern troughs are pro- stresses, and to contaminants acquired in las, and 3,000 individuals in Guanica tected, and cattle still utilize them as their diet (herbivorous in the lanial stage Forest in the southeast. Although water sources. Each is highly eutrophic, and insectivorous as an adult) or through Moreno (1985) has described the and spraying of nearby pastureland for their permeable skin. As a result, there Guanica population as healthy, numer- cattle ticks may result in high levels of has been a recent concern over what is ous and stable, an examination of utilized pesticides in field runoff, which is the perceived to be a global decline in am- breeding sites illustrates the present pre- only source of water for the troughs. One phibian populations. carious status of this species. of the nearby pastures has even recently Release and translocation programs been utilized as a junkyard for the storage for amphibians, which have reproductive Northern Population of automobiles. The recent acquisition of strategies unlike most other vertebrate land for hotel development near Isabella species, are not well developed as of yet. In the north, Peltophryne lemur is has increased land value in this area - An examination of the progress of the known to have bred in four concrete, even small land holdings are for sale at Puerto Rican crest@ toad program may walk-in cattle troughs located near Queb- $6,000 per acre. be illuminating for those involved in or radillas. Each of the four sites flanks the Individual sightings of toads in this contemplating similar amphibian release same dry gully which carries storm run- area of development should result in programs. off a short distance before terminating at protection or enhancement of the breed- the sea. This ravine is utilized by toads as ing habitat as part of the development Current Status of Toad a refuge and as a migration route to the proposal. Unfortunately, development Populations cattle troughs. No more than 25 individu- may destroy breeding areas or essential als have been seen at any one breeding toad habitat before their significance is Perhaps the most significant factor event. If this represents the number of identified. The lack of field surveys and underlying the captive breeding and re- breeding adults, this population is not reliable field reports to confm the pres- lease of Puerto Rican crested toads is the large enough to sustain itself, given the ence of toads or toad habitat hinders ef- precarious state of wild populations. susceptibility of small populations to fective decision-making in this regard.

Southern Population

In 1984, Miguel Canals, the Guanica Forest Manager, discovered a large population of crested toads in a flooded parking area of Guanica Forest on the southern coast. This population consists of approximately 3,000 individuals which reproduce in one pond located 100 meters from the sea. Despite its protected location in the Tarnarindo sector of the Guanica forest, the pond is under threat of inundation by sea water during tropical storms. Indeed, a direct hit by a hurricane could result in permanent loss of the breeding pond if the small section of barrier beach is cut into by heavy seas. Furthermore, the beachside View of breeding pond in Guanica Forest. Toads live in the hills behind the beach. Posts were-~~ erected by the FGkst Manager to protect the breeding pond. Photo by B. Johnson. parking lot receive's heavy day use

54 Endangered Species UPDATE Vol. 8 No. I throughout the year. As the water in the after consultation with Puem Rico De- this site for subsequent breedings: the parking area is continuous with that in partment of Natural Resources @NR) ditch which each year drained the the breeding area, any contamination of staff ecologists, who requested the re- flooded parking area in which the toads soil or water might affect developing lease of as many toadlets as possible bred was dammed; a series of posts were toads. from captive-bred offspring still the planted to prevent cars from driving into When 13 inches of rain fell in a 24- property of the DNR as part of breeding the breeding area and destroying vegeta- hour period during Hurricane Gilbert in loan agreements. 1985, several ponds were breached as This northern site water rushed down from the surrounding was within the hills. One such pond, which sustained a historic range, did limited number of breeding pairs, was not have an exist- washed out. Miguel Canals witnessed ing toad popula- many pairs of toads washedout to sea and tion, and was char- has not observed any toads returning to acterized by dry this pond during the subsequent breeding forest and exposed events, In June 1990, while radiotracking limestone, consid- crested toads which had bred for the first ered to be an time in almost two years, we visited this essential compo- pond and observed Leptodactylus al- nent of the type biaris (white-lipped frog), but no Pelto- habitat. phryne lemur. In 1985, 20 The natural beauty of the beaches in toadlets were col- properties adjacent to the Guanica Forest lected from leaf Male Puerto Rican crested toad Photo by B. Johnson has increased pressures for development. litter surrounding Concerns have been expressed over the the Guanica breeding pond by represen- tion used by toads during egg laying; and impact of support infrastructures and the tatives from Metro Toronto Zoo and during the three-week breeding period need to improve accessibility,which will DNR. These toadlets formed the basis of the road to the piyking lot is closed to be necessary if hotel development oc- a southern founder population to be prevent water contamination. A public curs. Additionally, as with the northern managed under the AAZPA SSP pro- information campaign through local population, the known populations and gram. Breedings of these founders re- schools and newspapers has resulted in breeding habitat of this species in the sulted in the recommendation by the local acceptance of the changes which south cannot be considered secure. DNR for two additional releases of have occurred in this high-use area to toadlets into an area of Guanica forest ensure the survival of the toad. Thus, the History of Releases selected for the presence of water which survival of the species to date has been collected in borrow pits and because it largely dependent on the efforts of one In 1966, six northern toads were was considered to be outside the migra- individual. found - the first sightings since 1931. tory range of toads using the beach-side After a 1982 meeting at the Jardin Zool- breeding pond. The only evidence of Current Status of Release ogico de Puerto Rico to discuss the sup- survivorship has come from a 1989 dis- Program port role of captive propagation, several covery of two animals released as part of toads were found during a visit to one of a group of 640 in 1988. The long-term survival of the crested the cattle troughs. These individuals later Radio-tracking of 12 captive-raised toad is dependent upon protection of reproduced and their offspring were sent toads in 1989, and 12 wild, post-repro- existing breeding sites and establishment to the Buffalo and Indianapolis Zoos. ductive toads in 1990, has provided data of satellite populations. However, there The crested toad subsequently became on survivorship, home range, and habitat are no plans for further releases into the the first amphibian to be included in the use. The small size of released toadlets wild until suitable sites with release American Association of Zoological makes followup on the success of intro- ponds, either natural or human-made, Parks and Aquariums (AAZPA) Species ductions or releases difficult, but none- can be identified. Survival Plan (SSP). theless important. To date, the only In 1990, a Metro Toronto Zoo re- After a hormonally induced breeding measure of success has been the presence search team collected data on pH, salin- at the Buffalo Zoo in 1983, offspring or absence of toadlets at release sites. ity, calcium and temperature at Guanica were released into the cattle trough in to characterize the chemistry and eco- which their parents were discovered Conservation Ih Action logical characteristics of the breeding (Miller 1985). Northern toads bred at the ponds. This pond profile data is essential Buffalo and Toronto Zoos in 1984 and The importance of the discovery of to allow managers to maintain the char- 1985 were released into a small human- the Guanica population by Miguel Ca- acter of existing breeding ponds in case made pond in Cambalache Forest Re- nals cannot be underestimated, nor can of any sudden or long-term adverse serve. Cambalache forest was chosen the series of actions he initiated to secure changes, and to identify any other exist-

Vol. 8 No. 1 Endangered Species UPDATE 55 ing ponds suitable for crested toad repro- ing ponds must be characterized to test ern populations in the Quebradillas area duction. If no appropriate ponds are the suitability of future breeding sites seems imminent. The establishment of located, it will be necessary to construct (satellite ponds), and to monitor long- satellite populations may be an alterna- breeding ponds which include the essen- term changes which may negatively tive to augmenting existing populations tial components of the present pond. It is impact existing breeding ponds. from captive-bred stock. anticipated that new 6) If northern lines are exurpated, ponds, whether southern animals may be translocated to naturally formed the historic range. Studies of mitochon- during rains or con- drial DNA (Anna GoebeVGeoff Minton, structed to fill during University of Colorado at Boulder) are rain, would receive underway to determine the extent to tadpoles from two which northern and southern populations sources. Some tad- have diverged. poles would be 7) Genetic swamping can be antici- translocated from pated with the release of up to 5,000 the present breeding headstarted offspring from a single cap- pond to adjacent tive breeding event. For this reason, it is ponds outside the necessary to limit representation from migratory range of each founder and to utilize as many the toad, and a sec- founders as possible. This is especially ond pond would re- important if the wild population has suffered a severe reduction in numbers ceive captive-bred Photo by B. Johnson tadpoles returned to or there is evidence of bottlenecking. Puerto Rico for re- In areas where the existing population lease. has been extirpated, the release of These satellite such genetically related lines may be ponds would serve "The success of reintroduction pro- the only alternative, even if long-term asreservoirsoftoads gramsmustbeconsideredinatime environmental change or stress has in the event of frame of perhaps ten or more years, been a factor in the extirpation of the chastic destruction species. Genetic swamping may be re- of the present breed- and after a number of releases." duced if releases involve tadpoles, the ing pond. As well, lifestage during which mortality is the ponds to receive highest. tadpoles would al- 8) Given the high mortality rate in low the comparison of the success of 3) The establishment of new breeding wild amphibian populations, the translocations of tadpoles from the populations might be improved if the number of animals released should be Guanica pond with releases of captive- translocation of wild tadpoles and release based on the projected survival rate and bred animals. This data is important for of captive-bred toads was considered in the carrying capacity of the release envi- future releases of this and other imperiled tandem. The success of each approach ronment. species in order to make the most effec- would provide comparative data for fu- 9) It is preferable to keep all animals tive use of small populations in future ture release or uanslocation. intended for release in conditions which conservation programs. 4) Several founder populations are simulate the environmental stresses of required for the repeated release of num- the release environment. Short-term Strategies for Future Toad bers of unrelated lines. The status of holding in large outdoor enclosures Releases northern populations is of particular (preferably at the release site itself) concern given the limited number of would facilitate the initial orientation As a result of our experience to date, toads in captivity and known to exist in process and physiological adjustment to we suggest the following points be con- the wild. arid environments. The importance of sidered in future release programs: 5) It would appear that the number of map and compass orientation from and to 1) Given the need to locate a natal breeding animals presently known from natal ponds requires further testing. The pond for a breeding generation of toads, the north has fallen below the minimum controlled release of offspring to test the success in the short term should not be viable population size necessary to sus- effectiveness of introduction procedures anticipated. The success of introduction tain a population. Further study will and stock may be advisable before im- programs must be considered in a time indicate the need to augment populations plementation over larger areas. frame of perhaps ten or more years and centered at each of the four breeding 10) It may be necessary to monitor after a number of releases. ponds as an appropriate management and, if present in high densities, eliminate 2) The chemistry and ecology of exist- option, although the extirpation of north- mongoose, rat and marine toad popula-

56 Endangered Species UPDATE Vol. 8 No. 1 tions in breeding or release areas. Re- education must be integrated into the S~mmary leased toadlets may have to be protected conservation program. Other species for short periods of time from high densi- may benefit from increased awareness of The success rate for non-game ties of lizard and avian predators, at least the reasons for the decline of popular or translocations will be low until trials can until they have found rocky refuges. target species. For example, Metro be conducted and analyzed. A period of 11) Captive breeding and msloca- Toronto Zoo is producing a poster, to be relatively poor establishment must be tions must not be ends in themselves. circulated by DNR, which will relate the anticipated which should not negatively Protection and restoration of existing toad to its unique habitat in Guanica for- impact the consideration of species for habitat are essential components of con- est. It invites Puerto Ricans to identify fder introduction attempts. The reservation programs in- lease of captive-bred ani- volving releases of cap- mals and habitat restora- tive-bred animals. The tion are management natural colonization of "The exemplary successes of this tools which will increas- human-made or enhanced program may in fact create an ingly be the last resort for ponds, strategically placed environment in which the preserva- a number of species. The in patches extending away potential of success in from the prime breeding tion of lesser-known species can release programs must be sites, may be a valuable be anticipated." considered as only one adjunct to the existing management strategy population, rather than the among others in stem- removal and return of ani- ming the rapid and cata- mals. These satellite ponds may be colo- other populations of the toad, andappeals strophic loss of amphibian species nized much more rapidly than those for protection of essential habitat and. diversity. seeded with naive animals returned from other threatened species. captivity, and their value may be propor- 14) The AAZPA Puerto Rican Literature Cited and Other tional to their proximity to protected Crested Toad SSP Program and conser- Suggested Reading natal ponds. This is an exciting aspect of vation within, protected reserves in species conservation as it involves the Puerto ~icomust be integrated. We may new science of restoration ecology, and never fully understand the diversity of Johnson, R.R., and F.L Pahe. 1989. The release of hem Rian creaed toads: would require the enhancement or crea- relationships between toad and environ- bptivemanagement impliCBtionsand the cat- tion of habitat on an exwrimental basis. ment which will ensure the continued us connection. Paees 962-967 in AAZPA Re- 12) The demograpdic profde for in- survival of this island endemic. In light [email protected]. Miller, TJ. 1985. Husbandry and breeding of the lroductions will be weighted towards of our limited understanding of the pmc- hem (PeltoPhr>lne lemurl numerous tadpoles or newly metamor- esses of extinction of bee-ranging popu- ,-en, on its na~dhis,,. z, Biolwv-. phosed toadlets since, initially, these lations, we must continue to manage 4:281-286. &e the fodder upon which klective populations and habitat which may haie M'erJmd EL.Pahe~ds.1989. Sutusof the hemRican crested toad, Peltophryne lemur: processes occur. Some consideration been modified either intentionally or An ovetview, Crawshaw,G.,B. R. should be given to retaining a limited unintentionally by human impact. Lacv. C. Smith. and P. Tolson. Paees 53-58 in number of individuals in captivity until 15) With the establishment of hiiatimd Yearbook 28. - they are of reproductive a& before re- additional breeding sites, this could be- Moreno* 1984. UnpublishcdnOtesOn phryne lemur Cope. Internal document for the lease. This would eliminate the life come a short-term conservation project. hem Rim of ResOUKeS. stages in which highest mortality occurs The winding down of the conservation Paine, F.L., and J. DuvaL 1984. h search ofa rare before the actual release, but perhaps effort necessary at this time to ensure toad. Animal Kingdom Magazine 88(5):33-38. reduce the fitness of the animals since the the survival of the crested toad would Rivem, J.A., H. Mayow, E- Earemera, and 1. Izquierdo. 1980. Sobre el Bufo lemur (Cope). adults are naive to the home environment allow the allocation of limited captive J. Sci. 15:33-40. (Johnson and Paine 1989). Which of breeding resources for other species in these considerations are most relevant to need of intensive management. The the production of reproducing animals exemplary successes of this program has not yet been established. To release may in fact Create an environment in Bob Johnson is Curator of Amphibians and Rep- both lifestages would in effect be a hedg- which the preservation of lesser- tiles & the Metro Toronto PO Box 280, West ing for survival until reproduction can known species can be anticipated. The Hill, Ontario, CANADA MlE 4R5. occur. involvement of zoo-bred animals 13) The release of captive-bred ani- released to a protected environment mals along with protection and, if neces- ensures that captive space will be avail- sary, enhancement of existing breeding able for the continued survival of am- habitat can create publicity for the fate of phibian biodiversity globally and in this and other threatened species. Public Puerto Rico.

Vol. 8 No. 1 Endangered Species UPDATE 57 kpidochelys kempii hatchlings Photo by M. Graham Blake

Mexico has been studying the decline of Lepidochelys kempii at Rancho Nuevo, Tamaulipas, since the mid-l%Os. In 1978, Mexico, the U.S. Fish and Wildlife Service, and other U.S. agencies began a binational effort to save the species. For the last ten years, the U.S. Field Assistance effort has been administered by personnel of the Gladys Porter Zoo. In 1990, %7 nests were protected with 66,663 hatchlings released at Rancho Nuevo and an additional 2,025 shipped to the National Marine Fisheries labo- ratories at Galveston for headstarting. It is hoped that protecting the hatchlings from land living or aerial predators while crossing the beach will enhance the species chances for future recruitment.

58 Endangered Species UPDA TE Vol. 8 No. 1 Central Florida Scrub: Trying to Save the Pieces

Susan R. Wallace

Introduction onto protected land will Christman (1988) calls these areas decade. Along the Ridge, where rare be the only way to save many of Florida's "ancient scrubs" to distinguish them species may once have been common, rare endemic plants. As the scientific and from similar coastal scrub communities are now acres and acres of dead groves philosophical discussions continue, and which lack the same concentration of and ruderal vegetation. while the technology is still being devel- endemics. Some ancient scrub sites are Cattle ranching also has been respon- oped, time is running short. so rich that many rare species, both plant sible for clearing large portions of the and animal, occur together. On the one state. Additionally, human develop- The Ancient Scrub Community undisturbed site for the rare Florida ment, including homes, shopping malls, ziziphus, Ziziphus celata (the other two tourist attractions, and everything that Because of its unique geological his- sites are in "improved" pasture), there are goes with them, continues at arapid pace. tory, Florida is home to an unusually 23 endemic plant species (Delaney et al. Major highways travel along the crests of rich assortment of endemic plants and 1989),of which 11 are considered at risk. upland areas - Highway 27 atop the animals. An area of high ground Lake Wales Ridge and Highway 1 - I down the center of the state, the Lake along the Atlantic Coast Ridge - Wales Ridge, was a series of islands "Introduction onto protected right through some of the rarest plant during the Pleistocene interglacials. land will be the only way to habitat in the state. As sea levels rose and fell, and Flor- save many of Florida's rare Florida's initial efforts at land ida was shaped and re-shaped, the conservation and acquisition had isolated populations on these islands endemic plants." been directed towards wetland pres- evolved to produce one of the - - ervation. Now attention has turned to highest rates of endemism in the the remarkable species diversity of continental United States. However, most of these rare endemics the high ground areas. A state land-pur- On the crest of the Ridge are have extremely narrow ranges, even chasing program and The Nature Conser- remnants of the longleaf pine/wiregrass within the scrub communities of the vancy have recently set aside several community which was once common in Ridge (Christman and Judd 1990). There large tracts of pristine scrub habitat, and the Southeast. Here, however, it is punc- is no one site, nor even a group of sites, more sites are under consideration. tuated with rarities like the delicate an- which contains the entire range of hetero- But all of this may come too late for nuals: wide-leaf warea (Warea am- geneity along the Ridge. [Coastal scrub many rare species which exist now in plexifolia), Carter's mustard, (W. car- communities, particularly along the only a few remnant patches d scrub -in teri), and the dwarf plum (Prunus gen- Atlantic and northern Gulf coasts, also a vacant lot, along a fence row, in a ditch, iculata). contain unique scrub species, although behind a gas station. Many endangered At a slightly lower elevation, pre- not as many. And again, the ranges are species live only in small populations on sumably the site of ancient shorelines, generally very narrow.] land that is too urban to protect and too are patches of scrub habitat which repre- expensive to buy. These plants have no sent the state's richest flora and most en- Species Decline legal protection, no management, and dangered community. These areas are nowhere to go, surrounded by develop characterized by deep, dry sands of low Habitat destruction has been wide- ment and vulnerable everyday to bull- fertility, intense hot sun, a short surn- spread in Florida, and these areas of high dozers, cement trucks and herbicide mer rainy season, and long droughts in ground -the endemics' home ground- spray. winter. Each scrub patch is slightly dif- have been particularly hard hit. Two- For instance, Lakela's mint (Diceran- ferent, both biologically and aestheti- thirds of the ancient scrubs in central dra immaculata) is known only from a cally. Generally there is a cover of Florida have been destroyed (Christman ten-mile stretch of Highway 1, less than dwarfed sclerophyllous oaks, palmettos 1988). Much of the Lake Wales Ridge one mile wide, right in the middle of an (Serenoa repens, Sabel etonia), and has been cleared for citrus; the deep sands expanding urban area. Although some cactus (Opuntia spp.). Often, but not and (once) mild climate are ideal for seemingly robust populations still exist, always, there is an overstory of sand pine citrus and large portions of the area have they are on land with expensive highway (Pinus clausa). The soil may be yellow been planted in mile after mile of groves. frontage and "For Sale" signs. sand or the dazzling; snow-white sand Ironically, much of this citrus -- at least The related Garrett's mint (Diceran- found on many of Florida's present-day the northern half -- is now destroyed, dra christmanii) was known from five .beaches. wiped out by three major freezes in one sites, one of which was recently de-

Vol. 8 No. 1 Endangered Species UPDATE 59 stroyed. Other populations are quite where conditions are just right, appears just such a problem (Wallace 1990). small; the largest is on part of a defunct adequate, though never robust. Clearly Seedling generation for this genus has subdivision. Longspurred mint (Dicer- we don't understand what this plant been disappointing under greenhouse andra cornutissima) grows in fence rows needs, and its requirements are very ex- conditions (for reasons not yet clear), of an interstate highway and along road- acting. With incomplete knowledge, although it is frequently robust in the cuts in a nearby housing development. little time, limited resources, and very wild. We haveestablished an experimen- Wedge-leafed eryngium (Eryngium few seeds to work with, our intuition is to tal population of cutting-grown Scrub cuneifolim) is known from 20 sites along put the few seeds available on carefully mint (Dicerandrafrutescens) in the pro- the Ridge, ranging a distance of less than 20 miles (Chrisman and Judd 1990). One protected population grows at Arch- bold Biological Station. The other plants grow on private land and roadsides; a few forlorn remnants hang on in the sand behind a shopping center. Ziziphus celata is known from three sites, a total of five clones. Only one site is undisturbed. All are privately owned. This sad litany mentions only a few examples. Some species now have a few populations on protected sites - which seems reassuring when compared to the species which have no protection at all. But realistically we are still looking at numbers too few and areas too small to consider any of the species secure.

Propagation and Reintroduction - Efforts federally listed plant species. Photo by S. Wallace Bok Tower Gardens' Endangered Plant Program was begun in response to this critical situation. Four acres have "A state land-purchasing program and The Nature been set aside, away from the public area Conservancy have recently set aside several large of the Gardens, to house the a & col- tracts of pristine scrub habitat. . . But all of this may lection. A new greenhouse and a grid of come too late for many rare species which exist now in special planting beds have been constructed where genetically representa- only a few remnant patches of scrub - in a vacant lot, tive collections of the plants can be along a fence row, in a ditch, behind a gas station." propagated, grown, and monitored regularly. Now, with four years of experience chosen sites and let nature take over. tected buffer area sunounding the Gar- and 30 species accessioned into the col- Fortunately, not all the species we work dens. These were taken from a small lection (to one degree or another), the with are this humbling. population very near the Gardens, found limitations of & preservation have Other species can be maintained in in a vacant lot of a subdivision, and become clearer and the need for introduc- cultivation,but are so short-lived, with an planted on a site which seems to approxi- tion projects more imperative. attrition rate so high, that they require an mate the original. (The associates, how- Some species simply won't respond overwhelming commitment of man ever, are not exactly the same, thus one to cultivation at all. Scrub lupine (Lu- hours, greenhouse bench space, and land thing we expect to find out is how much pinus aridorum) is such an example. area to continually repropagate. This is this matters.) If this population prospers Few cuttings will root. Those that do root especially true when each clone has to be and seeds itself, we will look for pro- die immediately after transplanting, tracked individually in an effort to main- tected sites within the ranges of the other unable to tolerate any root disturbance. tain a genetically representative popula- species. Seeds will germinate, then dwindle and tion. A naturally occurring population of die within a few weeks, even under nu- The four Dicerandra species in our Warea amplexifolia grews within the merous permutations of cultural recollection, whose natural strategies rely wiregrass of Bok Tower Gardens' Pine gimes. Yet natural seedling generation, more on volume than longevity, present Ridge Preserve area. (Of the three other

60 Endangered Species UPDATE Vol. 8 No. 1 A grid of special planting beds houses the a& colledion of endangered plants at Bok Tower rare plant preservatibn, combining the Gardens. In the foreground is Chry~op~hfl~rida~. Photo by s.Wallace knowledge and resources of botanists, horticulturalists, land managers and populations of this species, one is on 16 plants lived to flower and set seed. public agencies. state park land, another in a vacant lot, This high attrition rate (from 600 to 16, Here in central Florida we are in a and the fourth in the path of a land-eating althoughthe rest lhe 2*000 seeds may desperate lastditch effort to "save the sand mine.) This fall-flowering annual come up next year) is not sqris-pises" after decades of thoughtless and crucifer, with dainty lavender blooms, 'g for an but it is cautiOn~ unintelligenttinkering with our environ- shows a marked variation in population for On seed as ment. Where habitat destruction has size from one year to the next, appar- a conservation tool. It is apt to take far been relentless and population numbers ently forming a fluchlating seed bank in more seed than we imagine to establish have dwindled to pathetic levels, bold the soil. (The related Carter's mustard, a PP~~~~~~.Only action is called for, even - W. carteri, has appeared on sites where 0' a scale large enough '0 reflect - with sometimes incomplete bowl- it had not been observed at all for several natural systems will provide the answers, edge and imperfen years previously (Bissett 1988).) There of which germination rates are only a appears to be a clear relationship to fire, small part of the story. It will be several Literature Cited with a winter burn stimulating an yearsbefore we can draw any morecon- increase in numbers and vigor the clusiOns about this Wares Bissett, N. 1988. TheNatives Nursery, Davenport, following year. although this fall we will seed another n.Personal communication. With some clues as to the species' site area with 10,OOo seeds. Christman, S.P. 1988. Endemism and Florida's preferences and management require- to Interior Scrub. Final Repod to Florida Game In contrast the above-mentioned and Freshwater Fish Commission. Tallahassee, ments, we have looked for ways to in- examples, however, gx sib preservation FL. crease our W. amplexijolia population has worked s~ectacularlvwell for other Christman. S.P.. and W.S. Judd. 1990. Notes on and establish new ones. Propagation species in o;r collectio~.Woody spe- Rants &d&c a Florida SC~.Biological Sciences. No. 1,1990. under nursery conditions yielded an -. ties like the pygmy fringe Wee, IW~~CV.K.R.. R.P. Wunkrlin, and B.F. HB~S~. impressive number of vigorous-looking Chionanthus pygmaeus, the four-pet- 1986.. ~ediscoveryof ziziphrrs cclnto (Rham- seedlings, all of which succumbed to aled ---uawuaw, Asimina tetramera, and naceae). SIDA 13(3):325-330. caterpillars, leaf miners, and fungal pa- the exceptionally rare zziPhus celaia, as walla~.SR. 1990. UP~and paved Over: Rare endemics find a new home in a Florida thogens before blooming. Repeated at- well as semi-woody perennials like the baranid Bkcal of the Lin- tempts to raise this plant with petunia- three scrub mint Conradina species -- C. ne, Society. press. growing technology failed to keep a plant brevifolia, C. glabra, and C. grandijlora alive long enough to flower. - are long-lived with more stable per- Direct seeding into our natural area sonalities. In each case, the horticul- (after a bum, on a site where Warea had tural is enough devel- Susan R. Wallace is Curator of Endangered Plants never been seen before) produced oped to produce the plants in large at the Bok Tower Gardens, PO Box 3810, Lake slightly better results. Two thousand numbers and keep the populations alive Wales, 33859-3810. seeds produced 600 seedlings, of which indefinitely.

Vol. 8 No. 1 Endangered Species UPDA TE 61 Down But Not Out: Reintroduction of the Extirpated Malheur Wirelettuce, Stephanomeria malheurensis by Robert L. Parenti and Edward 0. Guerrant, Jr.

INTRODUCTION outbreeding S. exigua ssp. coronaria. uted to the decline in S. malheuremis. Sympatric to the point of sometimes Possible alleopathic effects of cheatgrass Known only from a single site in having intermingling stems, S. are being investigated by the USFWS in southeastern Oregon, the herbaceous an- malheurensis occurs at the northern limit cooperation with Boise State University. nual, Stephanomeria malheurensis of its progenitor's range, which extends Preliminary studies show that cheatgrass (Malheur wirelettuce) was discovered in throughout much of California and the does inhibit germination, growth, and 1966, placed on the federal list of endan- west. Malheur wirelettuce, however, is development of selected species. gered species in 1982, and by 1985, had known to occur only at a single site: an apparently become extinct. Recovery areaon BLM land about 25 miles south of RECOVERYRESEARCH efforts begun in 1987 by the U.S. Fish Burns, Oregon, which is now designated and Wildlife Service (USFWS), along the South Narrows Area of Critical Experimental Design with the Bureau of Land Management Environmental Concern, (BLM), were possible only because vi- Recovery efforts have been compli- Because the ability of the sole origi- able seed had been stored offsite and was cated by the fact that the original habitat nal site to support Mglheur wirelettuce available for reintroduction. The seed was altered both by a fire in 1972, and appears to have been compromised, the had been cuefully maintained by Dr. subsequent invasion of cheatgrass recovery plan includes research intended Leslie Gottlieb, an evolutionary geneti- (Bromus tectorum) - a very aggressive to answer a number of questions about cist at the University of California at weed native to Europe. Never known to the biology of the species in order to Davis, who had discovered, described, number more than 750 plants, the decline determine what, if any, managerial op- and worked extensively with the plant. of the wireleuuce was first noted in 1980 tions exist for the BLM to re-establish Stephanomeria malheurensis is of -1981, and by 1985-1986, no individuals and maintain a viable population of considerable scientific interest in part could be found. Simultaneously, Malheur wirelettuce. To better under- because it belongs to one of he most cheatgrass coverage increased drarnati- stand the biology of Malheur wirelettuce well-studied parent-offspring relation- cally. What had been mostly bare soil and its competitive interactions with ships known between any two species. was replaced by nearly a 100% ch- cheatgrass and other associated species, The self-pollinating S. malheurensis is eatgrass cover, suggesting that inter- the BLM, in conjunction with the almost certainly arecent derivative of the specific competition may have contrib- USFWS, established a series of experimental plots at the original site in 1987. FOU; separate ploiwere established, each surrounded with rodent-proof wire mesh enclosures extending four inches below ground and 32 inches above. Each plot is dominated by one of four species common to the study area: the three natives, green rabbitbrush (Chrysothamnus vicidijlorus), big sagebrush (Artemisia tridentata var. wyomingensis), and Great Basin wildrye (Elymus cinereus);and the exotic cheatgrass. Three of the plots are square, 5m on a side, and the cheatgrass plot is twice that size, 5m by 10m. In 1987, all plots except the cheatgrass plot were completely weeded of cheatgrass. Into these plots were planted 1,000 seedlings of S. malheurensis seedlings, which had been grown by The Berry Botanic Garden of Portland, Oregon, from seed Robert Parenti at the field site pointing LO where the last naturally occurring wirelettuce plant was provided by Gottlieb. The plants were found (flags mark seedlings presumed to be offspring) Photo: LR. McMahan, Berry Botanic Garden thoroughly moistened on planting and

62 Endangered Species UPDATE Vol. 8 No. 1 thereafter for about four weeks until the seed set, and seed release are signifi- soil seed bank then this may not be a plants had become established. cantly later in plots which include the serious setback. In any case, consider- Survivorship and reproduction were exotic grass. In addition, survival and able quantities of stored seed now exist generally high. In 1988, with seed in the recruitment rates are lower in thoseplots. offsite. soil from plants grown there the year Also in 1990, all of the approximately before, the 5m by 5m plots were divided 250 seedlings from The Berry Botanic CONCLUSION

This ongoing recovery effort, begun with the planting of seedlings in 1987, has been extremely important in establishing research and management direc- tions that must be taken in the future. A focus of continued study is clarification of the effects of native and exotic species on wirelettuce. Data collected over a period of years will be necessary to provide information about their interactions and allow investigators to determine whether changing environmental conditions alterthoseinteractions. Without the experimental plantings, we would have little to go on to determine the survivability of S. malheurensis. This project is an excellent example of the cooperation paired-quadrant 50%/100% regime was Garden were planted outside of the plots between the USFWS, BLM, academic maintained in the cheatgrass plot. Due to in order to: 1) ascertain the effects of scientists, and private organizations to small mammal depredation of Malheur native herbivores, and 2) establish ex- enhance the survival of a species with wirelettuce in the sagebrush plot in 1988 perimental plots to examine the longev- such scientific importance. -a recognized natural threat to the spe- ity and germination patterns of naturally The primary goal of continued study cies - that plot had to be re-established occurring soil seed banks. Native herbi- is to determine management options for in 1989. It was weeded of all cheatgrass, vores, primarily the blacktailed jackrab- protection and perpetuation of Malheur and 80 greenhouse-establishedseedlings bit, destroyed all of the seedlings that wirelettuce. In retrospect, the existence from The Berry Botanic Garden were were not protected and watered for the of artificially "banked" seeds maintained planted. Thus, there were no contrasting soil seed bank experiment. This may not offsite appears to have made the differ- treated/untreated quadrants in the sage- have been a "fair test" because the year ence between extinction and continued brush plot in 1989. In 1990, the ch- was so dry, and therefore these plants existence for S. malheurensis. eatgrass plots were prepared in the same were conspicuous forage. manner as the other three plots, i.e., por- The replicated soil seed bank study is OTHER INFORMATION SOURCES tions were divided into "treated" and designed to follow the fate of a single "untreated" quadrants. cohort of seeds: what proportion of seeds Gonlieb, LD. 1973. Genetic differentiatim, will geminate either in the fall or spring of sympatric speciation, and the origin of a diploid species of Slephammeriamalhewensis. Amer. Results the first year, and for how many years J. Bot. 60(6):545-553. will seedlings from a single year's seed Gottlieb, L.D. 1977. Phenotypic variation in Data collected through 1989 show crop emerge thereafter? Populations of Stephanomeria exigua ssp. wronaria (Can- that S. malheurensis plants are influ- annual plants rely for survival on recruit- positae) and its recent derivative species "malheurensis." Amer. J. Bot. 64(7):873-880. enced differently by the various species. ment from the supply of viable seed in the Gottlieb, LD. 1979. The origin of phenotype in a Wildrye appears neither to affect nega- soil. The pattern of S. malheurensis seed recently evolved species. Pages 264-286 in tively the vigor of Malheur wirelettuce, survivorship in the soil has taken on more O.T. Solbrig, S. Jain, G.B. Johnson,P.H.Raven, eds. Topics in Plant Population Biology. Co- nor alter its phenology. Rabbitbrush, on than an academic interest because .there lumbia University Press, New York. the other hand, does appear to have some appears to have been no seedling recruit- Gottlieb. L.D.. and J.P. Bennett. 1983. Interfer- negative effect. The effects of sagebrush ment at all in 1990. It was a very dry and ence between individuals in pure and mixed cannot yet be determined because of the cold spring, which apparently did not cultures of Stephanomeria malheurensis and its herbivory and subsequent necessity to provide the very narrow temperature progenitor. Amer. J. Bot. 70(2):27&284. reestablish that population. The most window in the upper 50s (degrees Fahr- Robert L. Parenti is the Tri-state Plant Manager for profound effects on Malheur wirelettuce enheit), known to stimulate germination. the U.S. Fish and Wildlife Service, Boise, Idaho. were associated with cheatgrass: plants This may very well have been a blessing, Edward 0. Guermnt, Jr. is Conservation Diredor grow more slowly than in its absence, and because the rest of the growing season and Seed Bank Curator at The Beny Botanic Gar- den, Portland, OR. the phenological events of flowering; was dry as well. If there remains a viable

Vol. 8 No. 1 Endangered Species UPDA TE 63 Reintroduction of the Texas Snowbell, (Styrax texana) by Paul Cox

Few other areas in the United States gered in 1984. Today there are currently Building upon the early groundwork can boast the ecological diversity found fewer than 40 individuals known from laid by the RPSC, we soon found that the in Texas. A combination of vastly differ- seven widely scattered populations. Fur- Texas snowbell took to cultivation with ent edaphic and climatic factors give rise thennore, some of these populations only a few minor problems. Thus it did to no less than ten popularly recognized consist of only a single specimen. not take long to develop a fair-sized vegetational regions. Rainfall across the The Texas snowbell is actually quite captive population of about 150 plants. state varies from as much as 50 inches in an ornamental shrub. Away from the We determined that the best germination the eastern pine hardwood forests to as steep cliffs it stands about ten feet tall rate resulted from stratifying the seed for with handsome heart-shaped 60 days in moist sphagnum. By satisfy- leaves that are dark green above ing these production and subsequent and whitish below. In mid to cultivation requirements, we met our first late April the plants are deco- goal of establishing a captive population. rated with beautiful 1/2 to 314 inch-long, bell-shaped white Reintroduction Efforts flowers. These are followed later by 114 to 1L? inch hard, Encouraged by this success, we began round fruits usually containing to explore the possibility of reintroduc- a single seed. Unfortunately for ing these plants into protected habitats in the fruits, the parent plants are the wild. An Institute of Museum Serv- almost always found leaning ices grant helped pave the way for this out from cliffs where the seeds endeavor, but the project developed in little as five inches in the western deserts fall into creeks or dry streambeds and are earnest when Charlie MacDonald, with and mountains. swept away by the next good rainfall. the U.S. Fish and Wildlife Service One of the most significant areas is This situation, together with the fact that (USFWS), approached us and proposed a the Edwards Uplift, a limestone plateau the plants are eagerly eaten by herbi- three-year grant to fund Texas snowbell located in central Texas, locally known vores, has combined to push the Texas reintroduction efforts. as the Texas Hill Country. This fairly snowbell to the brink of extinction. In order to carry out this task it was rugged region consists of rolling hills our good fortune to team up with Toney deeply cut by steep-sided creeks and Captive Propagation Keeney, a professor at Southwest Texas ravines. Most of the Hill Country area Junior College in Uvalde. Toney has has been severely abused during the last The Texas snowbell has long been spent many years studying the snowbell, century. Coupled with a warming and recognized as. Texas' most seriously knows the populations thoroughly, and drying trend, overgrazing has devastated threatened native plant species. Pioneer long ago gained the trust of the some- much of the natural flora. Not only is the propagation work on this and many other times wary landowners. Rather than native white-tailed deer population at an endangered Texas species was per- resent the interference from outsiders, all time high, but the proliferation of formed by the now defunct Rare Plant Toney welcomed us with open arms. The exotic game animals such as the axis deer Study Center (RPSC) of theuniversity of reintroduction of the Texas snowbell was and aoudad sheep have added to the Texas at Austin. In 1985, sometime after something he had long contemplated but plight of native vegetation. The wide- the demise of theRPSC, the San Antonio lacked the resources to implement. He spread increase of sheep and goats has Botanical Center became a participating had already picked out a site and garnered only exacerbated the degradation. Large institution with the Center for Plant support of one landowner and the ranch areas have been overgrazed to the point Conservation (CPC), an organization foreman. Gaining such support can be a that the survivability of some plant spe- dedicated to preserving our nation's difficult task as ma1 Texans are often cies is seriously in question. endangered flora. One of the CPC's intensely resentful of programs which Enter here the Texas snowbell goals is to establish secure captive they view as government interference. (Styrax texana) located deep within the populations as insurance for some of However, for this job Toney Keeney is heart of the Hill Country. This obscure the more critically endangered species. well-suited, with a polite laid-back and shrub grows exclusively on cliffs associ- The first plant brought into the San sincere approach that is very disarming. ated with the deeply cut waterways of the Antonio Botanical Center's collection A second landowner was secured and the area. Due to low population numbers, the under this program was the Texas project was ready for implementation. species was listed as federally endan- snowbell. The effectiveness of these efforts illus-

64 Endangered Species UPDATE Vol. 8 No. 1 trates the importance of working closely began to put on new growth when the secure individuals, it must be questioned with local authorities who are invaluable cooler fall weather arrived in the fall. whether or not these individuals can to such a complex project. After three years the project results become a successful regenerating popu- Overall, the selected reintroduction have been very encouraging. The Mare lation under present circumstances. sites are good reflections of actual snow- Creek site has 18 survivors that now Herbivorous animals exert a great pres- bell habitat, i.e., steep banks near reliable average43.2 cm in height. To accommo- sure on virtually all Hill Couny plants; water. As muchas was possible under the date this growth, new cages were con- the Texas snowbell is noteworthy only circumstances, careful consideration structed for eight plants and installed in because it already had such relatively low was given to every important aspect of February, 1990. At Cypress Creek, population numbers. site selection. The plants had to be con- mortality was lower but growth was Secondly, the work by Texas Parks fined to areas within their historic range, slower. Only one of the original 24 plants and Wildlife has shown that protected and kept as far as possible from popula- was lost and the remainder have added populations seem to be capable of sexual tions of the closely related, somewhat more than 24 cm in height. While they are reproduction, but only within the con- more eastern sycamore-leaf snowbell not growing fast, these plants look fines of their enclosure. However, there (Styraxplatanifolia)to prevent the possi- healthy and appear in stable condition. is, at present, insufficient funding or en- bility of hybridization between these two This phase of the project thus seems to be thusiasm for a project that would exempt species whose ranges do not normally fairly successful. The protected plants herbivores fiom an area large enough to overlap. Other important aspects tden are growing satisfactorily within the accommodate the need for an expanding into account include available moisture, confines of their wire cages, although Texas snowbell population. soil type, exposure, and similar associ- they have yet to reach breeding size. ated plant species. We eventually chose The ability of these plant. to regener- Conclusion two reintroduction sites, called Mare ate under the present conditions is ques- Creek and Cypress Creek. tionable. The bulk of the Hill Country is Our part of the project has been Planting the progeny had to be per- so seriously overgrazed by both native largely successful to date. Propagation formed while perched upon eight foot and exotic herbivores that protection is and cultivation techniques are refined to ladders carried into the field. Once in- necessary on a community level, not just the point that the Texas snowbell can be stalled in their new homes on the bluffs for individual plants. Recent work by the grown in captivity with confidence. adjoining the creek, the plants had to be Texas Parks and Wildlife Department Reintroduced plants have survived the watered andmeasured regularly through- found that a fenced population of Texas crucial first few years of establishment out their early years. This was heroically snowbell demonstrated successful pre- and are growing well. carried out by Toney Keeney and re- liminary regeneration. Although the snowbell may not actu- cruits, who had to use the same precari- These combined efforts indicate two ally be the most threatened plant species ous procedures as in the planting expedi- things. First, the Texas snowbell can be in Texas, it is probably the most well- tions. To protect the young plants from reintroduced into protected native habi- known and longest-studied of our na- marauding herbivores and rodents, the tats, provided it is defended against ani- tives. Preliminary results are encourag- seedlings were covered by small chicken mal predation. While the project offers ing. With increased education and public wire cages. Toney's ongoing mainte- awareness, success of the project may nance and record keeping is entirely prove to be a turning point in the manage- funded by the USFWS grant. ment of Texas endangered species. In 1987, we planted a total of 25 seedlings at the Mare Creek location and 24 Additional Sources of Information at the Cypress Creek site. The Mare Creek seedlings, planted on May 21, Cory, V.L. 1943. ?he genus Sfyrax in central and averaged approximately 14 cm in height, western Texas. Madrono 7(4):110-115. Consoulin, GJ. 1974. A revision of Styrar (Styra- while Cypress Creek seedlings, installed caceae) in North America, Central America, and on July 13, averaged 16 cm. the Caribbean. Side 5(4):191-258. As fate would have it, during the last Mahler, W.F. 1981. Status report on Styrax [ex- week in May and early June, record rain- ma. USFWS, Albuquerque, NM. Pde,J.M., and D.H. Riskind. 1987. Endangered, fall fell on the Mare Creek location. Most Threatened, or Protected Native Plants of of the plants there were either destroyed Texas. Texas Parks & Wildlife Dept, Austin. or damaged by flood waters and associ- U.S. Fish and Wildlife Senice. 1984. Final rule to determine Sfyrax tera~(Texas snowbell) to be ated debris. Consequently, on July 17, an endangered species. Fed. Reg. 49:40036- 24 replacements were replanted, but on 40038. the limestone ledge above Mare Creek. U.S. Fish and Wildlifeservice. 1987. Texas Snow- The Cypress Creek "population," on the bell (Styrar twnu) Recovery Plan. USFWS, other hand, was unaffected by the heavy Albuquerque, NM. 53 pp. rains, although one plant was lost to Paul Cox is at the Supervisor of the San Antonio San drought stress during the hot summer Planting Styrax texanu on limestare bluffs at the Botanical Center, 555 Funston. Antonio, TX months. Many plants, at both locations, Mare Creek site Photo by C. Mchald 78209.

Vol. 8 No. 1 Endangered Species UPDATE 65 Transplantat ion of an Otherwise Doomed Population of Barrett's Penstemon, Penstemon barrettiae by Edward 0.Guerrant Jr.

Unlike other projects reviewed in this Translocation Efforts plant, although not all separate plants are Special Issue, this work does not address necessarily different genetic individuals a formal recovery effort for a listed spe- In May 1986, Ms. Julie Kierstead, then because the plants can spread naturally cies. Rather, it describes the first phase of The Berry Botanic Garden, took mul- by the rooting of shoot fragments. of an opportunistic experiment involv- tiple cuttings of vegetative shoots from as An initial planting in the spring of ing the transplantation of rootedcuttings many plants as she could reach - an 1987, which was undertaken in a con- taken from a newly discovered but amount equalling somewhat less than half spicuous location near the visitor center doomed population of Penstemon bar- the plants visible - either from the for publicity purposes, was done at the rettiae, a candidate (C2) for listing, to a ground or in a "cherry picker" supplied by wrong time of year. Consequently, this nearby site. (Candidate 2 species are the Corps. The cuttings were taken to the planting resulted in 100% mortality the imperiled species deemed by the U.S. Garden wherean attempt was made to root first summer. However, in the spring of Fish and Wildlife Service to have insuf- them. Results were generally encourag- 1988,70 rooted cuttings from 21 differ- ficient data for listing.) ing, but varied widely from complete ent parent plants were planted in two With large bluish-green leaves and success with some clones (9 of 26 clones), more appropriate habitats nearby, both showy tubular rose-colored flowers, to complete failure in others (3 of 26 already occupied by P. barrettiae. As of the small shrub, Penstemon barrettiae, clones). The term "clone" is used here to May 1990, at least half of these plants is an attractive plant. Until this popula- refer to all cuttings taken from a single were still alive (some loose tags were tion was discovered in 1985, the approximately dozen known populations of the species were limited to about a five mile stretch along both banks of the Co- lumbia River in Washington about 50 miles east of Bonneville dam, with scattered populations to the north along the Klickitat River. It is a colonizing species that grows on rocky cliffs and gravelly hillsides, often along highways "While wholesale and railroads where it is vulnerable to transplantation of indiscriminate collecting and herbicide use. endangered popu- This disjunct population was discov- lations is clearly ered shortly before the Army Corps of not the method of Engineers was scheduled to begin con- choice to preserve struction of a new navigational lock at genetic diversity, in the Bonneville Dam. Although at the time not legally required to protect this this case the alter- "Candidate 2" species (even though native was certain its habitat was slated for certain de- destruction." struction), the Corps helped support a cooperative project to salvage cuttings, and facilitated their transplantation at two nearby sites. While wholesale transplantation of endangered populations is clearly not the method of choice to preserve genetic diversity, in Ed Guerrant and Jack Poff inspecting a site for introduction this case the alternative was certain de- of Peastemon barrettiae at Bonneville Dam. struction. Photo by L.R. McMahan

66 Endangered Species UPDA TE Vol. 8 No. 1 found in the most recent survey, which despite an initial appearance of success. markably straightforward attempt has may belong to surviving plants), and In this case, success has been achieved succeeded. three are beginning to produce flowers. thus far only in locations already occu- A strength of this strategy of translo- The existing shoots of several plants have pied by the species - a situation not cating threatened populations is that ide- largely died back, but they are producing always available, and not necessarily ally, given enough lead time before a population's destruction, all of the genetic information in the population can be salvaged -at least temporarily. This "From a strategic or policy perspective, another information can then be housed at several at once to minimize the chance of weakness [of translocating populations] is the ap mortality. Even in this nearly ideal situ- parent ease of the method; where land usses con- ation, though, some clones were lost. flict, simply moving the offending occupants to Nevertheless, the Garden still has some another suitable site should not be offerred as an of these clones, and additional out- early or preferred option." planting is planned in the future. Seeds were also collected along with the original cuttings, and are currently housed in The Berry Botanic Garden's Seed Bank for Rare and Endangered Plants of the vigorous new shoots from the base of the even desirable. All of the plants origi- Paci.fic Northwest. These offer further plant. Of the 2 1 clones planted out originally at Bonneville before construction options for future planting. nally, 15 (7 1%) are still represented by at may well have been one breeding popula- Ultimately, when construction on the least one living individual. Much of the tion anyway, so moving "aliens" into an new lock is finished, there will be new mortality appears to be due to soil ero- already occupied site (the only suitable apparently suitable habitat available, and sion. habitat available nearby) probably did ideally we will attempt the ultimate goal not contaminate the indigenous gene of this project: to recolonize the Evaluation of Strategy pool. But genetic contamination remains unoccupied cliff face habitat with the an important unanswered question. original occupmts, or their progeny. Simply moving a threatened popula- There are other weaknesses as well: This project is, I believe, best viewed as tion by transplanting cuttings superfi- genetic variation may well be lost, and an opportunistic experiment- perhaps a cially offers an appealing, pragmatic success cannot be determined for a con- preview of things to come if global option for conserving genetic diversity siderable length of time. Not all plants warming occurs. It is too early really to -at least in theshort term. However, the yielded viable cuttings, and not all clones evaluate the results with any confidence, method involves many uncertainties and show equal survivorship. Consequently, other than to say that some genetic diversity has apparently been lost, but that some remains where otherwise there might have been none. "This [translocation] project is, l believe, best viewed as an opportunistic experiment Additional Sources of Information - perhaps a preview of things to come if global warming occurs." Kientead, J. 1986. Barrett's penstamon: A story. Plant Conservation 1 (2):l. Schwartz, A. 1988. Banking on seeds to avert extinction. Audubon 22-27. may not represent a generally desirable the genetic composition of the resulting alternative. The most obvious shortcom- population is probably different and less ing is illustrated by the fact that all plants diverse than the original population. Edward 0.Guemt, Jr, is Conservation Director rapidly died in the first planting because From a strategic or policy perspective, and Seed Bank Curator at the The Beny Botanic Garden, 11505 SW Summerville Ave., Portland, the habitat was apparently not adequate, another weakness is the apparent ease of OR 97219 even for this easily propagated colonizer. the method; where land uses conflict, The limited number of "surplus" indi- simply moving the offending viduals in rare populations makes such occupants to another suitable site should failed experiments potentially disastrous not be offered as an early or preferred for the survival of an imperiled species. option. Another problem with trans- However, failure also can be pro- plantation is that it may take many tracted over a much longer time span, years before we know if even this re-

Vol. 8 No. 1 Endangered Species UPDATE 67 Colorado Squawf ish Reintroduction Efforts in the Lower Colorado River Basin by Buddy L. Jensen

INTRODUCTION Unfortunately, these activities have re- Hatchery and Technology Center sulted in the decline of many native fishes (NFHTC)in Dexter, New Mexico, from Dam and irrigation diversion con- throughout the country. This paper dis- rearing sport fish to holding, studying struction have extensively modGed most cusses the demise of, and some of the re- and culturing imperiled native fishes of major U.S . rivers. Additionally, rivers covery efforts for, one of the unique na- the Southwest. This was a major step and streams commonly were treated with tive fishes impacted by development forward in the effort to conserve native fish toxicants when projects were com- activities in the Colorado River basin of fish species. pleted in order to remove "trash" fish the Southwest. Dexter NFHTC serves as a refuge for prior to dam closure. Most native fishes The rivers arid springs of the Ameri- imperiled fishes, provides opportunities were considered expendable in the inter- can Southwest, along with their native for research and development projects est of carrying out these development aquatic fauna, changed dramatically using captive-reared fish, and produces projects or enhancing sport fishing pro- during the 20th century (Miller 1%1). selected species for recovery programs grams. These projects, when combined Over time, the activities described above identified in recovery plans or other in- with other habitat modifying activities, resulted in a continuous and progressive teragency documeny (Jensen 1983, i.e., pumping of ground water, over- decline in the distribution and abundance Rinne et al. 1986). The Dexter NFHTC grazing, and poor logging practices, had of the unique endemic ichthyofauna of currently maintains 14 species of imper- a deleterious effect on the natural quality the region (Johnson and Rinne 1982). iled fishes. Since the inception of the of our river systems and their fauna. The voices raised in opposition to these current program, more than 25 species Non-native sport fish species were practices were a select few, those with have been held at Dexter for one or more stocked extensively with little or no con- insights far beyond the majority of fish- purposes. The Colorado squawfish, sideration given to potential impacts on ery managers of the time. Twenty-two Ptychocheilus lucius, a federally listed native fishes or other aquatic organisms. years ago in Death Valley, some of these endangered species (Federal Register foresighted indi- [32(43):40001] March 11, 1967 and viduals were in- [39(3): 11751 January 4, 1974) is one of strumental in es- the species being maintained and pro- tablishing the duced at Dexter NFHTC for reintroduc- Desert Fishes tion purposes. This account briefly ex- Council - a amines the captive breeding program for group of con- Colorado squawfish at Dexter NFHTC, cerned profes- and reintroduction efforts for the species sionals whose pri- in the lower Colorado River basin in mary goal was to Arizona. conserve, protect, and perpetuate DESCRIPTION AND CURRENT native desert RANGE fishes and their habitats. Twelve The Colorado squawfish is North years ago, the America's largest, native minnow. The combined efforts term "lucius" means "pike-like," and of the Desert squawfish do resemble northern pike in Fishes Council, body form: individuals are elongate, the academic fish- compressed dorso-ventrally, and have a eries community, long flattened head. Adults have a dark and key U.S . Fish olivaceous-colored back which lightens and Wildlife Serv- along the sides to a whitish belly. Al- ice (USFWS) per- though a true minnow (Family Cyp- sonnel led to the rinidae), it historically attained lengths changing of the and weights approaching six feet (183 Figure 1. Past and present distribution of the Colorado squawfish in the Colorado River system (from Colorado Squawfish Recovery Plan 1990). role of the Dexter cm) and 100 pounds (45 kg) (Miller National Fish 1961). In recent years, however, indi-

68 Endangered Species UPDATE Vol. 8 No. 1 viduals over 30 inches (76 cm) and 15 Historically, the Colorado River was pers. comm.). Perhaps more impor- pounds (6.8 kg) have been difficult to savage and unpredictable, known simply tantly, exotics comprise an estimated find. by its one dominating color - el Rio 99% of the total fish biomass present in Like the other big native fishes of the Colorado, the Red River of the West the system. In addition to being greatly Colorado River -- razorback sucker (U.S. Fish and Wildlife Service 1990). reduced in numbers, the Colorado (Xyrauchen texanus), bonytail chub Seventeen hundred miles (2,700 km) squawfish now occupies only about 25% (Gila elegans), and humpback chub long and dropping more than 2.0 miles of its former range, and has been (Gila cypha) -- Colorado squawfish are (3.2 krn) in its journey from the high extirpated from the entire lower basin long-lived. However, squawfish grow mountains of Wyoming, Utah and Colo- (Figure 1). and achieve sexual maturity much slower than the other three species; captive- reared razorback sucker and bonytail chub can reproduce at two years of age, while the squawfish must be six years old (Jensen 1983,Rinne et. al. 1986). Recent studies of adult bonytail chub and razorback sucker revealed that both speCies live for more than 40 years (McCarthy and Minckley 1987, Minckley et. al. 1989). Although data are limited, this author estimates that historically, Colo- rado squawfish lived to be 60 to 80 years of age and perhaps older. Sixteen year- old squawfish maintained at Dexter NFHTC average about 25 inches (63.5 cm) in length and 5.7 pounds (2.6 kg) in rado to the Sea of Cortez, the river RECOVERY ACTIVITIES weight. Thus it is easy to speculate that a courses through high canyons 90% of the six-foot, 100-pound fish would be ex- way. In this turbulent, silty river evolved Because of existing conditions, re- tremely old, perhaps over 100 years old. one of the most remarkable assemblages covery of the Colorado squawfish will Native only to the expansive Colo- of American freshwater fishes, uniquely requireamajoreffort. Between 1973 and rado River basin of the western United adapted to survive some of the most tur- 1979,59individuals were collected from States and northwestern Mexico, the bulent aquatic conditions on earth. the wild and transferred to Willow Beach "white salmon of the Colorado" was well Beginning with the construction of NFH, Arizona. From these fish, several known by early settlers for its white, Roosevelt Dam on the Salt River in 1913, year classes were produced (Toney 1974; flaky, sweet meat and its migratory and continuing with more than 20 addi- Hamman 1980, 1981). Representatives spawning habits (Minckley 1973). This tional mainstream and tributary struc- of two of these classes (1974 and 1981) voracious predator was historically com- tures completed since then, humans have are currently held at Dex ter NFHTC. The mon in the main river channels through- dramatically altered the character of el 1974 year class has been spawned for ten out the Colorado River basin (Figure 1). Rio Colorado. No longer flowing freely consecutive years to produce young-of- An indication of their former abundance to the ocean, the River has become a the-year for research and reintroduction was noted by Miller (1961) who stated, series of large impoundments over much purposes. Genetic diversity of the cap- "Until about 1911, the species was so of its course, connected by cold, clear tive broodstocks was maximized to en- abundant in the lower Colorado that indi- waters uncharacteristic of the former sure genetically fit offspring for reintro- viduals got into the irrigation ditches and river. Someof the Arizona tributaries are duction (Ammerman & Morizot 1989). were pitchforked out onto the banks by now dry in their lower reaches. For the hundreds for use as fertilizer." example, over 200 miles (320 km) of the Spawning and Culture Gila River flows only seasonally during SPECIES DECLINE intense summer thunderstorms or in Willow Beach and Dexter hatchery the spring following uncommonly wet managers developed and refined squaw- The decline of the Colorado squaw- winters. fish spawning techniques. To induce fish is probably related to a combination These disruptions to the physical riv- ovulation, females receive an intraperi- of factors, the most important being: 1) erine environment, coupled with intro- toneal injection of carp pituitary as they direct loss of habitat; 2) blockage of duction of numerous exotic fishes, have approach final maturation of ova (Ham- spawning and seasonal migration routes caused a steady decline in the abundance man 1981,1986;Inslee 1983). Egg pro- by dams and diversions; 3) changes in of the native fish fauna. There are 35 fish duction averages about 18,000llb flow and temperature regimens below species native to the Colorado River, (39,690kg) of body weight; one ten- dams; and 4) interactions with non-na- but more than 50 introduced fishes have pound female at Dexter NFHTC gave tive, introduced fishes. become established (W.L. Minckley, over 200,000 eggs during a single spawn.

Vol. 8 No. 1 Endangered Species UPDATE 69 Ovulation occurs 18 to 24 hours fol- lations must be included in upper basin been monitored up to three days immedi- lowing injection. Ovulated eggs are strategies, their recovery/reintroduction ately following stocking to evaluatepst- hand-smpped into porcelain pans con- efforts have progressed more slowly. stccking dispersal of, and predation on, taining sperm diluent solution, Milt from Reintroduction and monitoring of stocked fish. Generally, the stocked fish two or more males is expressed into the Colorado squawfish in the Verdeand Salt move rapidly downstream and are preyed spawning pans directly from the fish. River systems of central Arizona began on by both native and exotic predators. Water is then added to the mixture to in 1985 (Johnson 1985). activate the milt. During the spawning Individuals stocked and process, the entire mixture is stirred with populations established a feather to ensure adequate mixing of under this recovery ef- sexual products, and to prevent the eggs, fort are classified as which are highly adhesive, from sticking "nonessential experi- to the pan. Following fertilization, a mental" under Section slurried bentonite clay solution is added 106) of the Endangered to the mixture to reduce the adhesiveness Species Act, as of the eggs prior to placing them in float- amended (Federal Reg- ing egg baskets. The eggs are washed ister Vol. 50(142): gently to remove the clay solution and 30188-30195) (Brooks allowed to water-harden for 30 minutes 1986). (Nonessential prior to enumeration. populations are notcon- Eggs are enumerated gravimetrically sidered tobe essential to and placed in incubators receiving a flow the survival of the spe- Photo: B JensenDNFHTC of aerated, 70" Fahrenheit water. Hatch- cies and thus are not ing begins at about 96 hours and peaks at afforded protection under Section 7 of Additional monitoring has been carried about 108 hours. Sac fry are transferred the Act.) This approach to reintroduction out a few weeks to six months following to tanks where swimup occurs at approxi- was necessary because water develop- stocking. Based on information col- mately % hours post hatching. Swimup ment and consumptive-use interests lected during the latter surveys, survival fry are stocked in earthen rearing ponds at feared that water use might be restricted of introduced squawfish appears low. a density of 125,00O/surface acre. or controlled due to the presence of a Recovery of stocked fish has occurred on Swimup fry are also shipped live in plas- federally protected species. numerous occasions, but primarily tic bags containing water and oxygen, via Since 1985, a total of 488,117 Colo- within a few weeks following stocking; air freight, to other cooperating agencies rado squawfish fingerlings from Dexter only a few individuals that have been in and institutions. Colorado squaw fish NFHTC have been stocked into central the system for more than six months have attain a length of three inches (7.6 cm) Arizona waters, along with an additional been recovered. Although results are during their first growing season (June - 301 adult, surplus broodfish. The Dexter inconclusive because of difficulty in October) at Dexter NFHTC. Fingerlings facility also provided thousands of fry sampling the large, remote river habitats, and adults are transported in standard fish and fingerlings to Page Springs State they are nevertheless indicative of the distribution units. Fish Hatchery in Cornville, Arizona, for situation, and support the premise that rearing and stocking. Between the two successful recovery of Colorado squaw- Stocking stations, a total of 518,258 fingerling or fish will occur neither quickly nor easily. adults have been reintroduced into his- Protecting surviving wild stocks of toric habitats in the lower basin. Al- SUMMARY AND DISCUSSION Colorado squawfish is irnportant,sorein- though these numbers seem significant, troduction efforts have been approached they number less than the ova from three The progressive aspect of the Colo- differently between the upper basin ten-pound females,and are probably infi- rado squawfish reintroduction program states (Wyoming, Utah and Colorado) nitely small compared to the numbers of has been the successful development of where wild populations stillexist,and the young fish produced under historic con- spawning and culture techniques for the lower basin states (New Mexico, ditions. species; large quantities of fish can be Arizona, Nevada and California) where produced for reintroduction purposes Colorado squawfish were extirpated at Monitoring upon request. The difficulties encoun- least 25 years ago (Minckley and Deacon tered have been in the attempts to suc- 1968). The primary objectives guiding Within the constraints of personnel, cessfully reintroduce fingerling fish. early reintroduction efforts in the lower funding, and time constraints, biologists The reasons why reintroductions have basin included attempts to locate suitable from the Arizona Game and Fish Depart- been minimally successful cannot be habitat for reintroduction, setting up ment, USFWS, and Arizona State Uni- precisely determined, but probably in- stocking protocols for a listed species, versity have evaluated the success of include a combination of some or all of the and conducting post-stocking dispersal troductions annually. River reaches re- following unranked factors: 1) dams and and survival studies. Since extant popu- ceiving reintroduced squawfish have other habitat alterations; 2) presence of

70 Endangered Species UPDATE Vol. 8 No. 1 numerous exotic species that interact ing structures. Instead, the most apparent cult. A continued major commitment to with and prey on reintroduced fish; 3) problem affecting successful reintroduc- recovery of the species is required if it is native species predatorlcompetition in- tion is the presenceof exotic fishes which to be saved in the upper basin and suc- teractions brought about by altered habi- both prey upon and compete with squaw- cessfully reintroduced in the lower basin. tats; 4) reintroduction of comparably fish for available food and space re- Time alone will reveal the final chapter in small numbers of fish in large, diverse sources. Reservoirs impounded by dams the saga of the Colorado squawfish -the habitats; 5) sampling inefficiencies due harbor large populations of exotic spe- native white salmon of el Rio Colorado. to personnel and habitat constraints; 6) cies, thus providing a constant source of life history parameters (Colorado squawfish which move upstream to occupy ACKNOWLEDGMENTS fish are long lived and grow slowly, so habitat needed and historically utilized one cannot expect immediate results by native fishes. Additionally, non-na- The following individuals reviewed the manu- from the efforts made thus far); and 7) tive fishes continue to be stocked directly script and provided many useful editorial and homing. into the rivers. Yet, the physical riverine constructive canments: Dr. W.L. Minckley, Dr. Tyus (1985) and Tyus et al. (1985) habitat above mainstream reservoirs James E. Johnson, Dr. Donald C. Hales, James E. Brooks and Atha Sharon Coats. reported homing behavior and spawning appears adequate to support the Colorado migrations for wild Colorado squawfish squawfish and other extirpated species as in the upper basin. Tyus cautioned about well. Assuming that existing habitat LITERATURE CITED stocking hatchery-reared fish into habi- conditions are maintained in these areas, tats containing wild fish because their exotic fishes appear to represent the prin- Ammeman, L.K.,and D.C. Morizot. 1989. Bio- inability to home could possibly impact cipal roadblock to successful establish- chemical genetics of endangered Colorado extant populations of the species. If ment of Colorado squawfish in the lower squawfish populations. Trans. Am. Fish. Soc. 118(4):435-440. Tyus's conclusions are correct, this could Colorado River Basin. Establishment Bmks, James E. 1986. Reintroduction and moni- also be a problem in the lower basin even alone does not equal recovery, however, toring of Colorado squawfish (Ptychocheilus though natural populations have already and once established, the role of migra- Iucius). Annual Report, Nongame Branch, been extirpated. If squawfish imprint tion and homing may become important Arizona Game and Fish Department, Phoenix. 14 PP. during early life stages, migrate back to factors in the maintenance of self-sus- Hamman, R.L. 1980. Spawning and culture of spawning grounds to reproduce, and will taining populations. Colorado squawfish, humpback chub and bony- spawn only at those locations, hatchery- Other efforts that would enhance re- tail chub during 1980 at Willow Beach Nalional reared fish likely will not reproduce in the Fish Hatchery. Pages 137-144, 159-166. and covery possibilities of Colorado squaw- 169-175 in Final Report of the U.S. Fish and wild since they imprinted at the hatchery fish in the lower basin include the follow- Wildlife Service and U.S. Bureau of Reclama- which is out of basin. However, observa- ing ideas: 1) renovation of river reaches tion, Part 3. Colorado River Fishery Project, tions made at Dexter NFHTC suggest to remove or at least reduce numbers of Salt Lake City, Utah. Hamman, R.L. 1981. Spawning and culture of that squawfish do not necessarily require non-native species, 2) mechanical re- Colorado squawfish in raceways. Progressive spawning migrations and homing to suc- moval (electrofishing,etc.) of non-native Fish Culturist 43(4):173-177. cessfully reproduce: fish hatched and fishes, 3) strict baitfish laws, 4) commer- Hamman, Roger L 1986. Induced spawning of reared in the Colorado River basin have cial harvest of riverine catfish popula- hatchery-reared Colorado squawfish. Progres- sive Fish Culturist 48(1):72-74. spontaneously reproduced in ponds at tions, 5) liberalization of bag limits for Inslee, Theophilus D. 1983. Spawning and hatch- Dexter NFHTC. Although recruitment sport fishes in sections of selected river- ing of the Colorado squawfish (Ptychocheilus has been low, it is specuiatedthatit would ine habitats, 6) dedication of one or more lucius). Station Report. Dexter National Fish be much higher if adequate substrate and rivers or streams to native fish only, and Hatchery, Dexter, New Mexico. 20 pp. Jensen, Buddy L 1983. Culture techniques for current were provided to simulate a natu- 7) public education. Whether politics seleded Colorado River imperiled fishes. Pm- ral spawning bed. This does not mean will ever permit the manipulation of large ceedings Nolthwest Fish Culture Workshop that spawning migrations and homing are river reaches to remove non-native fishes XXXIV:130-142, Moscow, Idaho. not important to the continued survival of in order to re-establish extirpated native Johnson, J.E. 1985. Reintroducing the natives: Colorado squawfish and woundfin. Proceed- wild individuals, only that successful fishes, remains to be seen. The Colorado ings Desert Fishes Council XVII:118-124. reproduction can occur in the absence of squawfish perhaps represents a unique Johnson J.E., and J.N. Rime. 1982. The Endan- these factors. It remains obvious that opportunity in this area since both the gered Species Act and Southwest fishes. Fish- much more work is needed to fully under- states of Arizona and Colorado would eries 7(4):1-8. McCanhy, M.S., and W.L Minckley. 1987. Age stand the role that homing and migration like to establish this species as a sportfish estimation for razorback sucker (Pisces: Ca- play in the life history of this species. with controlled take. Although this des- tostornidae) from Lake Mohave, Aha-Ne- Although the physical presence of ignation would do little to emphasize the vada. Joumal of the Arizona-Nevada Academy of Sciences 21337-97. dams must have played a major role in the need to recover and conserve a species, it Miller, Robert R. 1961. Man and the changing fish demise of the Colorado squawfish may be a step toward public acceptance fauna of the American Southwest. Papers of the throughout its historic range, they may and support of the efforts needed to re- Michigan Academy of Science, AN, and Let- not be directly affecting the outcome of covery this and other important native ters XLVI 365-404. Minckley, W.L. 1973. Fishes of Arizona. Arizona early reintroduction efforts in the lower fish species. Game and Fish Department. Phoenix. basin since all stockings are taking place Obviously, much work remains to be Minckley, W.L., and J.E. Deacon. 1968. South- in riverine habitats upstream from exist- done, and the task that lies ahead is diffi- western fishes and the enigma of "endangered (continued on bottom of next page)

Vol. 8 No. 1 Endangered Species UPDATE 71 A Fish Faunal Conservation Program: The Lake Victoria Cichlids by Les Kaufman

Introduction faunas are particularly suspect. Like invertebrates and plants, fishes are so In general, efforts to protect the nu- "Lake represents piose that only a few fragments of a merous s~eciesof imwriled fish in the the first time an entire few faunas could wssiblv be maintained United &tes and in -the world lag far faunal assemblage has under an SSP a&xh ;sing available behind initiatives to preserve furry and been listed as endan- space and resources. However, even if feathered "charismatic megaverte- gered by the IUCN, al- wise decisions could be madeconcerning brates." For instance, only one marine which of the many species to preserve, fish species - the totoaba (Cymscion though convincing argug there are numerous challenges to first macdonaldz) -is even listed for protec- merits could be made to overcome in animal husbandry, and in tion under the Endangered species Act. list many other[s] . . . A the scientific study of genetics, epigenet- The 1988 IUCN Red Data Book lists 5% complete list of threat- ics, microevolution, and developmental imperiled freshwater fish species, nearly ened and endangered biology. The Lake Victoria hap- half of them from a single body of water lochromines provide a potent case his- -- Lake Victoria in East Africa, bordered fishes. . . would probably tory of the nature of these many and by Kenya, Uganda, and Tanzania. Lake include more than 10% of complex challenges. Victoria represents the first time an entire the 22,000 or so known faunal assemblage has been listed as en- fishes." Lake Victoria Haplochromines dangered by the IUCN, although convincing arguments could be made to list Lake Victoria, the world's largest many other assemblages, such as most lake, contains an endemic radiation of desert fishes of North and Central Amer- in 1987 an IUCN Captive Breeding Spe- over 350 haplochromine cichlid fish ica, the stream fshes of the southern cialist Group (CBSG) was established to species which are known for their re- Appalachians, the ephemeral killifishes develop propagation and educational markable diversity of feeding mecha- of eastern South America, the orestines programs for endangered aquatic spe- nisms and corresponding morphological of the Andes, the crater lake fishes of the cies, especially fishes (see Kaufman and behavioral specializations. In addi- Cameroons, and the rainforest fish fau- 1987, 1989; Brown 1987). Three fish tion, the Lake is inhabited by two en- nas of Madagascar, Sri Lanka, and Aus- faunas were chosen for Species Survival demic and three introduced tilapiine tralia A complete list of threatened and Plan (SSP) programs: the Lake Victoria cichlids, along with about 40 other fish endangered fishes, based only on pres- haplochromines, Appalachian stream species from nine other families. Thepri- ently available information and infer- fishes, and North and Central American mary cause of the Victorian cichlids' pre- ence, would probably include more than desert fishes. Of these, the Lake Victoria cipitous decline is the successful intro- 10% of the 22,000 or so known fishes. SSP has progressed the farthest to date. duction of a large alien predator, the Nile In recognition of this crisis, and the The restoration and preservation of a perch (Lutes niloricus), although deoxy- limited but important role that captive single endangered species is a monumen- genation and overfishing have certainly propagation might play in forestalling it, tal task, thus proposals to rescue entire contributed to cichlid decline and failure

(continuedfiom preceding page)

species." Science 159:1424-143 1. April 3,1985. Fish Culture Section and Fisher- Rivers, Colorado and Utah, 1981. Proceedings Minckley, W.L., D.G. Buth, and R.L Mayden. ies Management Section of the American Fish- Desert Fishes Council Xm:102-108. 1989. Origin of broodstock and allozyme vari- eries Society, Bethesda, Maryland. U.S. Fish and Wildlife Service. 1990. Colorado ation in hatchery-reared bonytail, and endan- Toney, Donald P. 1974. Observations on the Squawfish Recovery Plan. Colorado River gered North American cyprinid fish. Trans. propagation and rearing of two endangered fish Fishes Recovery Team, Denver, Colorado. Am. Fish. Soc. 118(2):131-137. species in a hatchery environment Proceedings Rinne, J.N., J.E. Johnson, B.L. Jensen, A.W. of the Annual Conference of the Westem Asso- Ruger, and R. Sorensen. 1986. The role of ciation of State Game and Fish Commissioners Buddy L Jensen is the Director of the Dexter hatcheries in the management and recovery of 54:252-259. National Fish Hatchery and Technology Cen~er, threatened and endangered fishes. Pages 27 1- Tyus, Harold M. 1985. Homing behavior noted for 285 in Richard H. Stroud, ed. Fish Culture and Colorado squawfish. Copeia 1985 (1):213-215. U.S. Fish and Wildlife Service, PO Box 219, Dexter, New Mexico 88230. Fisheries Management, Proceedings of a Sym- Tyus, H.M., EJ. Wick, and D.L Skates. 1985. A posium on The Role of Fish Culture in Fisheries spawning migration of Colorado squawfish Management, Lake Ozark, Missouri, March 3 1- (Ptychockilus lucius) in the Yampa and Green

72 Endangered Species UPDATE Vol. 8 No. I to rebound in areas where Nile perch ness, rates of endemism, and taxonomic ing species-oriented conservation ef- themselves have been overfished. The composition (Echelle and Komfield forts. Distinguishing individual species actual rate and severity of cichlid 1984, Greenwood 1984). Thus, some within major species groups is very extinction is difficult to ascertain, as data argue that given the similarity in the three difficult, and requires comprehensive, is being collected much more slowly than faunas, it wouldbe better to focus limited carefully curated specimen collections the ecosystem is changing. Damage to resources for research and conservation such as exist only at the University of fish fauna in certain areas is obvious, upon those faunas which are still rea- Leiden, the British Museum of Natural albeit complicated: haplochromines are sonably intact and undisrupted. This History in London, and to a lesser extent, nearly gone from some areas, nile perch view is untenable, however, as the faunas Harvard University in Massachusetts. from others, perch and haplochromines are largely independent of one another, Many species are still undescribed and live together in some places where and thus offer a replicated evolutionary may remain so long after they are extinct fishing is off limits, and neither is doing experimenr the loss of one lake would in the wild. very well where fishing pressure is greatly weaken the strength of evolution- The coloration of living, sexually high. Despite initial skepticism regard- ary hypothesis-testing (Kaufman 1989, active males is one of the most im- ing original reports that a mass Avise 1990, Meyer et al. 1990). portant taxonomic characters for distin- extinction was occur- guishing haplo- ring, it now appears 7 71chromines (e.g., that these reports were Hoogerhoud et al. not exaggerated; at 1983), but these colors least 40% of the fauna change when a speci- is extinct (Lake Victo- men is preserved. Con- ria Research Team in sequently, researchers prep.). have begun to develop color transparencies to Significance of the accompany specimens. Cichlids However, there are perhaps only five to six The endemic cich- Haplochromis chilotes (Paralabidochromischilotes) from Lake Victoria experts in the world lids have far-reaching Photo by L. Kaufman who are currently quali- significance as a com- fied to identify large ponent of the Lake numbers of species ecosystem. ". . .there are serious ethical implications about from color vansparen- For the people of the cies; proper training of region they were a di- allowing the at least partially preventable new investigators de- rectandindirectsource extifl~ti~nof several hundred species. Lake uwn the of protein: native ti- Victoria's endemic fishes, and history's judge- bf these'individuals to la~iinecichlidsarestill mentconcerninghumanresponsetotheirplight, catalogathoro~ghcol- themostdesirablefood will unquestionably become an important case lection. fish in the lake; and the Although the color introduced nile perch, study in environmental ethics." of preserved specimens now the most impor- is a useful identifica- tant fishery, depended tion tool, color patterns on haplochromines for food. Aside from Perhaps more importantly, however, of individuals within a species vary their food value, many of the endemic there are serious ethical implications widely. To further complicate matters, haplochromines feed upon snails that about allowing the at least partially pre- observations indicate that coloration of serveasvectorsforschistosomiasis.The ventable extinction of several hundred sexually active males changes elimination of these fish species is thus species. Lakevictoria'sendemic fishes, ontogenetically (i.e., during develop- speculated to have potentially severe and history's judgement concerning ment). Molecular systematics, while impacts on Victorian riparian peoples human response to their plight, will un- showing promise as an identification (Slootweg 1986,1987). questionably become an important case technique, has not provided the hoped- From a scientific standpoint, Lake study in environmental ethics. for panacea; electrophoresis often fails to Victoria's fish fauna represents a classic resolve forms that differ markedly in re- example of "explosive speciation" - of Systematic Characterization of productive coloration, ecology, and particular interest to evolutionary scien- the Fauna form. Although progress is being made tists. However, cichlid radiations are in the use of mitochondria1DNA (Meyer characteristic of lakes in the region, and The complexity of haplochromine et al. 1990), this technique is not yet re- within Lake Malawi and Tanganyika are relationships has been a major obstacle to liable in distinguishing haplochromine even comparable in terms of speciesrich- understanding their evolution and focus- species.

Vol. 8 No. 1 Endangered Species UPDA TE 73 Phenotypic Plasticity and be the best means of managing hap by the British Museum, the Hap- Artificial Selection lochromines. lochromine Ecology Survey Team, and the Lake Victoria Research Team sug- One of the most puzzling aspects of Pre~e~ingSpecies integrity gest that the situation in Lake Victoria is evolutionary differentiation in tooth and probably deteriorating very rapidly, jaw form of haplochromines is that, de- Establishing the correct identity and but in at least some parts of the lake, spite their morphological specializa- relationships of haplochromines is only shallow-water inshore cichlid assem- tions, many of these species also exhibit the first step in their conservation. Pre- blages do not appear to have been a remarkable degree of plasticity in the same characters. For example, the mor- phology of Astatoreochromis alluaudi varies between snail-eating populations, which exhibit massive pharyngeal jaws "In the case of Victorian haplochromines, the informa- with molariform teeth, and non-snail tion base is wholly inadequatefor the task of prioritizing eaters, which have proportionately captive propagation programs, and quite far from the smaller, more gracile jaws with papilli- stage where planning of enhancement or restoration of form dentition. Wild-caught individuals wild populations might be considered." of this species exhibited hypertrophied jaws and teeth characteristic of the Lake Victoria population, but their progeny, raised principally on soft foods, exhibited the hypertrophic condition, as did serving the integrity of species distinc- severely disrupted. The multinational wild-caught individuals maintained on tions in captivity may be even more dif- collaboration recently established by the soft foods for several months. Hooger- ficult. There is little evidence of inter- National Oceanic and Atmospheric houd has since confirmed epigenetic specific hybridization of hap- Administration's (NOAA) Large Lakes plasticity in this.species through elegant lochromines within Lake Victoria, but of the World Program has renewed experiments and functional analyses preliminary observations strongly indi- long-term ecological research on Lake (Hoogerhoud 1984,1986). cate that hybridization is both possible Victoria. In a strict Darwinian sense, acquired and probable within the confines of aq- characters are not heritable, and therefore uariums less than several thousand liters Action Plan for Victorian such phenotypic plasticity should not in volume. Interfertility among hybrids Haplochromines pose problems for captive breeding pro- appears to be common, and bizarre hy- grams. Unfortunately, certain individu- brids between morphologically disparate In hopes of consolidating conserva- als are often better suited, for one reason haplochromine species have been ob- tion efforts by European, North Ameri- or another, to the aquarium environment, served in hobbyists' aquariums. Simi- can, and Kenyan institutions, the CBSG and thus grow faster, monopolize breed- larly, entirely distinct Malawiian cichlids has drafted a five-year plan for the devel- ing encounters, and are, by accident or that have never been known to hybridize opment of a formal SSP program for the design, selected for subsequent breeding in nature, do so readily in aquariums; entire Victorian endemic fish fauna or exhibition. The extreme plasticity of such "homogenization" of Malawi (Aquarium Action Plan 1989). The ob- haplochromines ensures that at least mbuna stocks is a common and familiar jectives of the program are: some individuals will deviate from natu- problem in large cichlidexhibitsat public 1) To establish exhibits and pro- ral patterns of trophic development. aquariums. gramsexplaining freshwater extinctions, Since these individuals grow faster than the importance of conserving native their brethren, they may be preferentially Insufficient Data fishes, and the relationship between fish selected, leading to the eventual canali- conservation and habitat preservation; zation or "assimilation" of characters One of the most serious problems 2) To stabilize,conserve, and propa- that were originally acquired on an facing nascent fish conservation efforts gate haplochromine stocks presently in individual basis, through epigenesis is the extremely limited database on dis- North America and Europe for research (i.e., a Baldwin effect). Consequently, it tribution and abundance of threatened and education; is extremely important to assess the species in the wild. In the case of Victo- 3) To assess the status of Victorian criteria by which individuals are chosen rian haplochromines, the information cichlids in the wild, determine conserva- for breeding, as well as the possible need base is wholly inadequate for the task of tion priorities, and incorporate up to 36 of for a captive environment that will prioritizing captive propagation pro- the estimated 300 haplochromine species foster the development of wild pheno- grams, and quite far from the stage where into a faunal SSP program as is feasible types. Cryopreservation of the bulk of planning of enhancement or restoration and appropriate. stock, with genes funneled through a of wild populations might be considered. Additional key elements of the plan small number of breeding adults, may What is known from recent expeditions include:

74 Endangered Species UPDATE Vol. 8 No. 1 Networking of the major organization biology. For this reason alone, a Ress, Orono, Maine. tions to be involved in CBSG activities, portion of the fauna shouldbe maintained Greenwood, P.H. 1984. African cichlids and evolutionary theories. Pages 141-154 in A.A. including the American Association of in the wild, and as a last result through Echelle and I. Komfield, eds. Evolution of Zoological Parks and Aquariums, the captive propagation. Species Flocks. University of Maine Press, IUCN, the U.S. Fish and Wildlife Serv- The creation of an SSC captive breed- Orono, Maine. ice, and corresponding agencies in other ing specialist group and initiation of field Lake Victoria Research Team (Haplochmmine Ecology Swey Team). In prep. countries; laboratory research on Lake Victoria Hoogerhoud, RJ.C. 1984. A taxonanic reconsid- Acquisition of all stock of known fishes are important steps. We're grate- eration of the haplochromine genera Gauro- origin now in captivity; and ful for the support of NOAA, the Na- chromir Greenwood, 1980 and Labrochrornis Regan, 1920 (Pisces, Cichlidae). Neth. Jour. Development of a 35mm slide and tional Science Foundation, the Institute Zool. 34539-565. specimen reference collection at the for Museum Sciences, the Environ- Hoogerhoud, R.J.C. 1986. Taxonomic and eco- Harvard Museum of Comparative Zool- mental Protection Agency, and the Pew logical aspects of morphological plasticity in om. Scholars Program for Conservation and mollusdvorous haplochromines (Pisces, Cich- lidae). Pages 131-134 in M.D. Craponde Proposed research activities in- the Environment for their generous sup Caprona and B. Fritzsch, eds. Proceedings of clude: long-term field studies in collabo- port for our field and laboratory research the 3rd European Workshop on Cichlid Biol- ration with Kenya, Tanzania, and and captive breeding program. This ogy. Ann. Kon. Mus. Mid. Afr., Zod. Weten- Uganda to assess the changing status of project is a collaboration of scientists sch. 251. Hoogerhoud, R J.C., F. Wine, and C.D.N. Barel. endemic species in the lake, and explore from Kenya, Uganda, Tanzania, the 1983. The ecological differentiation of two possibilities for long-term on-site con- United States, Holland, Great Britain, closely resembling Haplochromis species fm servation measures; a survey of the na- and Israel, and involves over 30 partici- Lake Victoria (H. iris and H. hiarm; Pisces, ture and scope of developmental plastic- pating museums, universities, and public Cichlidae). Neth. Jour. Zool. 33(3):283-305. Kaufman, L.S. 1987. Caught between a reef and ity in a variety of haplochromine species; aquariums. Yet, aquariums cannot hope a hard place: Why aquariamust investin captive the developmentofprotocols for measur- to serve as arks for more than a very small propagation. Pages 352-368 in Proceedings of ing genomic diversity in captive and wild proportion of threatened freshwater fish the National Meeting of the American Associa- stocks; and the exploration of cryopre- diversity. In the long term, the goals of tion of Zoological Parks and Aquaria. Kaufman, LS. 1989. Challenges to fish faunal servation of gametes as a management the Captive Breeding Specialist Groups conservation programs as illustrated by the tool for cichlid stocks. must eventually turn back to the fostering captive biology of Lake Victoria dchlids. Pages of good environmental stewardship - 105-120 in B.L Dresser, R.W. Reece, and EJ. Conclusion through education, through the transfer Mamska, eds. 5th World Conference on Breed- ing Endangered Species in Captivity, Cinnan- of technologies that foster renewable nati, OH; 1988. To gain perspective, one should con- exploitation of native species, and Meyer. A., T.D. Kocher, P. Basasibwaki, and A.C. sider multiplying the scope of the pro- through continuing basic research. The Wilson; 1990. Monophyletic origin of Lake Victoria cichlid fishes suggested by mitochon- drial DNA sequences. Nature 347550-553. Slootweg, R. 1986. Optimal prey size selection by two molluscivorous cichlid species and its ll. . . aquariums cannot hope to serve as arks for more implications for snail control. Pages 57-60 in M.D. Craponde Caprona and B. Fritzsch, eds. than a very small proportion of threatened freshwater Proceedings of the 3rd Eumpean Workshop on fish diversity. In the long term, the goals of the Captive Cichlid Biology. Ann. Kon. Mus. Mid. Afr., Zool. Wetensch. 251. Breeding Specialist Groups must eventually turn back Slootweg, R. 1987. Prey selection by molluscivo- to the fostering of good environmental stewardship.. ." rous cichlids, foraging in a schistosomiasis vector snail, Biomphaluria glabrala. Oecolgia 74:193-202.

Note: This paper is a modified and updated gram outlined above by the number of tasks before us are daunting - their version of a paper pblished in the proceedings freshwater fish faunas likely to become of the 5th World Conference on Breeding En- contemplation is a depressing task in it- dangered Species in Captivity in Cincinnati, endangered in coming decades. The self. But we have begun. Ohio, in 1988, edited by B.L. Dresser, R.W. probability of maintaining intact hap- Reece, and E.J. Maluska (see Kaufman 1988). lochtomine communities in Lake Victo- Literature Cited ria is extremely low, and the odds of restoration or effective conservation Avise, J.C. 1990. Flocks of African fishes. Nature management are highly uncertain. 347512-513. Les Kaufman is Chief Scientistatthe New England However, the research program neces- Brown, N. 1987. Conservation: A plea for a Aquarium (Central Wharf, Boston, MA 021 10) sary to conserve a portion of this assem- balanced view. Pages 199-202 in Proceedings and an Associate Professor in Ichthyology at Har- blage is of general theoretical interest, of the National Meeting of the American Asso- vard University. ciation of Zoological Parks and Aquaria. and is likely to yield important results for Echelle, A.A., and I. Kornfield, eds. 1984. Evolu- evolutionary, molecular and conservation of Species Flocks. University of Maine

-- - -

Vol. 8 No. 1 Endangered Species UPDATE 75 Apache Trout Culture: An Aid to Restoration by Bob David

The Apache trout, Oncorhynchw Arizona native trout) once inhabited Protection Efforts apache, formerly Salmo apache, histori- most perennial streams above 1,827 cally inhabited streams in the White meters within its historical range. The preservation of the Apache trout Mountains af east central Arizona, in- However, beginning in the late 19th was initially undertaken by the White cluding the headwaters of the Salt, San century, human interactions began to Mountain Apache Tribe in the late 1940s. Francisco, and Little Colorado Rivers. have a marked effect on populations. At that time it was believed that the Although the species has been noted in Grazing, logging, road construction, remaining pure populations in existence the literature since the late 1800s, it was and other habitat impacts altered were present in only a few streams on the not until 1972 that Robert R. Miller of watersheds, causing increased tempera- Fort Apache Indian Reservation. In

gill covers, and life Service fins. Dorsal, pel- (USFWS),which vic, and anal fins led to a better are edged in white "The production and distribution of large numbers of unders,ding of and occasionally Apache trout, however, is not the final answer in the the status of the display orange tips, recovery of this threatened species. Artificial propa- species. ~t was

Apache trout also fish from Ord displays the largest Creek, a popula- dorsal and adipose fins of any other tures and siltation, which are tion located on the Fort Apache Indian trout (Oncorhynchus spp.). Spots are detrimental to the spring-spawning Reservation, to its Sterling Springs State normally scarce and frequently Apache trout. Stocking of exotic Fish Hatchery. This was thebeginning of outlined with a noticeable pale halo. trouts, which began as early as 1917, captive rearing efforts designed to assist The light yellow belly may also harbor also contributed to the decline of the in the efforts to recover the species. a few spots and often exhibits blotches species. These highly efficient exotic The Apache trout was originally listed of orange in both sexes, particularly competitors have included the rainbow, as "endangered" under the Endangered during the spawning season. A yellow brown, brook, and Yellowstone Species Act of 1973. However, follow- "cutthroat" slash is dso present in the cutthroat trouts, of which the ing the recommendation of the Recovery gular folds, however, the ancestry of rainbow and cutthroat trouts are able to Team, the listing was changed to "threat- this species is linked more closely to hybridize with the Apache trout. This ened" in order to assist management and the rainbow trout than that of the cut- hybridization, along with competition, culture efforts. In addition, the Recovery throat. is attributed as a major factor in the Team also set a goal for the restoration of Researchers generally believe that extirpation of the Apache trout from 30 discrete populations of the species the Apache trout (also known as the many streams which it formerly within its historic range, as a condition Arizona trout, Arizona golden trout, inhabited. for delisting.

76 Endangered Species UPDA TE Vol. 8 No. 1 Captive Breeding Efforts targeted 30 streams prior to delisting of ceded by habitat improvement projects the species. designed to restore watersheds to a con- Attempts at culturing Apache trout by State-of-the-art techniques were used dition conducive to reproduction and the State of Arizona in the early 1960s in developing culture methods for survival of native species. and 1970s were largely unsuccessful due Apache trout. These included the use of In addition, we must be concerned to the conditions and methods applied. oxygen injection into the female body with the genetic integrity of the popula- ~uituretechniques currently in use on domestic rainbow trout strains were tried on wild Apache trout with very poor results. ~ighmortalitiesand the inability "Stocking of a native species in historical waters does of the species to utilize commercial trout diets eventually led to the disbanding of nothing to repair damaged habitat, nor will its eggs be the program in 1981. able to survive the onslaught of silt common to pres- Initial research on controlled, ent spring run-offs. Stocking must be preceded by cial propagation of Apache trout by the habitat improvement projects. . .9t USFWS began at the Alchesay-Williams Creek National Fish Hatchery Complex in May of 1983. Wild Apache trout were spawned on-site from the East Fork of the White River. Spawning personnel were cavity to expel eggs (air spawning). Milt tions.we choose to propagate. Artificial airlifted into the remote site, and fertil- was collected from anesthetized males culture should not impair the ability of a ized eggs delivered to the Williams by aspiration, subjected to short-term species to re-establish itself in a wild Creek Station by helicopter. Eggs were storage, andclassifiedas to viability prior state. Thiscan be ensured through the use. hatched, and fry were subjected to trials to fertilization. Resulting fry were fed of biochemical systematics to monitor designed to determine suitability of in- in trials using the latest high-perform- the continuity of a polymorphic gene troductory feeds. Out of 2,715 eggs col- ance diets designed for larval develop- pool. Unfortunately, pure populations of lected, only 240 hatched and survived to ment of warm water species found Apache trout remain only in isolated be used as future broodstock. During difficult to culture, such as walleye and headwater regions, cut off from other May 1984, wild Apache trout were again muskellunge. Trials resulted in the populations, where they have evolved spawned on site from the East Fork of the selection of a specific semi-moist diet, into relatively homozygous entities. White River. Out of 1,869 eggs collected introduced to the fry by use of remote, There is some question now concerning 1,204 were hatched, and 704 two-inch mechanical feeders controlled by their ability to adapt to other waters with fish were able to be retained as brood- programmed timers. conditions different from that in which stock. These two-year classes of wild The use of these, and other culture they currently exist. This is another rea- fish formed the basis for all future culture techniques led to the initial spawning and son why it is important to preserve what work at the Williams Creek Station. culture of 1,200 Apache trout in 1986, little polymorphism remains in the spe- 22,000 trout in 1987, 90,000 in 1988, cies by attempting to avoid genetic loss Reintroduction Program 175,000 in 1989, and 550,000 in 1990. due to selection during artificialpropaga- Apache trout production in 1990 will tion. One strategy to ensure this integrity Herein began a rather ambitious goal result in the accomplishment of the initial is the periodic introduction of genetic to replace all rainbow trout stocking on goal of replacing all rainbow trout stock- material from the wild into captive ppu- the Fort Apache Indian Reservation with ing with Apache trout by 1991. lations. In the case of Apache trout, similar programs using only the Apache alteration of spawning times between trout. The rainbow trout program had Evaluation of Captive Breeding wild and captive populations has made involved the stocking of approximately and Reintroduction Strategy this difficult. It is hoped that cryopreser- 500,000 catchable (20 cm), subcatchable vationofgametes from wild stock may be (15 cm), and fingerling (8 cm) rainbow The production and distribution of used to assist in this process. trout annually in waters consisting of large numbers of Apache trout, however, Currently, research is being con- over 1,200 hectares of lakes and 650 is not the final answer in the recovery of ducted on the survival of reintroduced kilometers of streams. This goal is not this threatened species. Artificial propa- Apache trout. Marked fish have been entirely a recovery effort, but a program gation is only a single tool among many placed in both lake and stream environ- designed to enhance current sport fishing in the overall plan to delist the species. ments in order to determine their ability enterprises while complimenting recov- Stocking of a native species in historical to adapt to different environmental con- ery plans. In addition to providing for a waters does nothing to repair damaged ditions and their ability to compete with sport fishery, the program also supplies habitat, nor will its eggs beable to survive other species. Only preliminary data fingerling Apache trout to the AGFD for the onslaught of silt common to present have been gathered, however, there are their use in restoring a portion of the spring run-offs. Stocking must be pre- some indications that winter SUI-viva1by

Vol. 8 No. 1 Endangered Species UPDATE 77 Apache trout in stream environments remains unavailable to fishermen by of individual gene pools, culture of en- may exceed that exhibited by exotic spe- stocking numbers approaching carrying dangered or threatened species can cies such as the rainbow trout. Survival capacity. In this way we are able to gain greatly enhance a recovery effort. in lake environments may favor intro- public support and possibly proceed in ducedspecies. The temperature, oxygen, restoring a site that may not otherwise be Additional Sources of Information considered due toadversepublic Behke, RJ., and M. Zun. 1976. Biology and opinion. management of threatened and endangered "Restoration of streams through re Another area mu&. U.S. Dept of Agric. For. Sew. Gen. in which M~- Tech. Rep. RM-28. establishment of historical species Miller, R.R. 1950. Notes on the cutthroat and propagation rainbow trouts with heciescripticn of anew spe- is not always a popular may prove to be cies from the GhRiver, New M&co. &as. management objective. In many a valuable tool is POP. MUS. ~ool.Univ. Mich., Ann Abr. in the 529: 1-42. cases, sportsman groups are op- Miller, R.R. 1972. Classificatim of the native technique posed to the loss of existing fisheries ~UISof A-na with the description ofa new composed of exotic species gene swamping. species, hlmo apchc. Copeia 1972:401-422. . . ." This involves the Novy, J. 1985. Forgotten waters: Arizana Wild- life. AZ Game and Fish Dept. hbl. 14-17. of large U.S. Department of Interior. 1979. Recovery Plan numbers of en- for the Arizona trout, Salmo apuche Miller, dangered spe- 1972. U.S. Fish and Wildlife Service, Al- and chemical changes inherent in lakes cies on top of existing exotic populations. hquequel NM. of Intenor' 19" Apche taut favor a species with a broad, polymer- While hybrid populations may result,it is implementation plan Unpubl. manusaipt. phic gene pool. Thus, Apache trout, hypothesized that natural attrition, U.S. Fish and Wildlife Service, Pinetop. AZ. which evolved in conditions of high through several generations, may pro- 15 pp. oxygen concentrations and relatively duce a relatively stable chemical balances, may not beable pure population. to compete in these environments. This application Restoration of streams through re- would be re- establishment of historical species com- stricted to resto- "Artificial propagation may be used to position is not always a popular manage- ration of non- shorten the time a stream remains ment objective. In many cases, sports- critical popula- unavailable to fishermen by stocking man groups are opposed to the loss of tions. numbers approaching car4ng capac- existing fisheries composed of exotic Other uses of ity. In this way we are able to gain species which are providing a significant artificial propa- sport fishery. Past experiences have seen gation in the re- public support . . ." these sites used in endangered species covery of endan- restoration, only to be closed to sport geredtroutshave fishing for extended periods while small yet to be defined. reintroduced numbers of fish expand B i o l o g i s t s their populations to a size capable of sup- should be enc0LUaged to explore further Bob David is Assistant Project Leader at the Alche- porting a fishery. Artificial propagation possibilities for the use of this manage- sa~-W*ams Creek National Fish Hatchery* PO may be used to shorten the time a stream ment tool. Through careful management Box 398, Whiteriver, AZ 85941.

I Subscription Information I I To subscribe to the Endangered I I Species UPDATE, enclose a check I 1 Mail subscription to: or money order for $23 ($18 for I students and senior citizens). Make I Name checks payable to The University of I I Michigan. I I organization I I Return with payment to: I I Address Endangered Species UPDATE 1 I School of Natural Resources I I CitylState/Lip University of Michigan I I AM Arbor, MI 48109-1 115 I I I

78 Endangered Species UPDA TE Vol. 8 No. 1 The Red Wolf: Recovery of an Endangered Species by Michael K. Phillips

Species Decline est from the zoo community. The Point red wolves, and the annual federal Defiance Zoo and Aquarium in Tacoma, budget for captive breeding increased to Red wolves (Canis re)ranged Washington, initially was the only zoo- about $200,000. As of October 1,1990, throughout the southeastern United logical facility involved. If not for the 131 red wolves existed in captivity. The StatesbeforeEuropeansettlement of that commitment of a handful of people and revised Red Wolf Recovery Plan (U.S. region. However, by 1980, the species the Point Defiance Zoo, the red wolf Fish and Wildlife Service in press) calls was considered extinct in the wild would have slipped into oblivion. for increasing the popula- (McCarley and Carley tion to 330 animals. 1979). Demise of the red The reintroduction proj- wolf was due to many ect in northeastern North factors: human persecu- Carolina occurred within tion of wild canids and de- the confines of the 63,636 struction of habitat forded ha Alligator River Na- the last few red wolves to tional Wildlife Refuge use marginal habitat in (ARNWR) and adjacent Louisiana and Texas Department of Defense where they bred with land covering an additional coyotes (Canis latrans) 20,454 ha. The areas con- and suffered heavy para- sist of marshes, nonriverine site infestation (Carley swamp forests, pocosins, 1975). Although the and agricultural fields. Lo- plight of the red wolf was cal climate is characterized recognized in the early by hot summers, mild win- 1960s (McCarley 1962), ters, and high humidity. the species was not listed The areas are bisected by as endangered until 1967, numerous logging roads in and did not receive "prior- "By the mid-1 970~,the USFWS realized it various stages of develop- ity treatment" until pas- was not possible to preserve the species in mentlrepair; many are only sage of the Endangered the wild, and uded that iecovery season all^ passable with a species Act in 1973' In could only be achieved through captive 4-wheel drive vehicle or on that year the U.S. Fish and Wildlife Service breeding and reintroductions." ,, Before release, each (USFWS) initiated a re- wolf was acclimated.to the covery program. refuge. The length of accli- By 1985, six zoological facilities held mation varied from a few weeks to 2t Captive Breeding and red wolves and the captive population years. Duringacclimation weminimized Reintroducation Efforts had grown to 65 individuals. With the human contact, hoping to reduce the species secure, the USFWS intensified wolves' tolerance of humans, varied the By the mid-1970s, the USFWS real- recovery efforts by initiating a reintro- feeding regime to expose the animals to ized it was not possible to preserve the duction project in northeastern North feast or famine, weaned the wolves from species in the wild, and concluded that Carolina. Because the reintroduction dog food and fed them an all meat diet, recovery could only be achieved through represented the first attempt in history to and provided the opportunity to hone captive breeding and reintroductions. By restore a carnivore species that was deter- predatory skills by giving them live prey. the late 1970s, the captive breeding pro- mined to be extinct in its former range Before the wolves were released, each gram was established utilizing 17 pure (McCarley and Carley 1979), the project was given a health check and fitted with red wolves that were captured in the generated tremendous interest about the a motion sensitive radio-collar. Nine of Texas and Louisiana refugium (USFWS prospects of recovering the species. the first ten wolves released were im- in press). Initially the program lan- From the fall of 1987 through October planted with radioactive tags (Crabtreeet guished due to a limited federal budget 1990, 14 more zoological facilities al. 1989). The tags allowed us to assign (about $30,000 per year) and little inter- committed themselves to maintaining collected scats to individual wolves. Ad-

Vol. 8 No. 1 Endangered Species UPDATE 79 ditionally, the first nine wolves were irn- planted with abdominal transmitters. These transmitters were placed as back- ups to the radio-collars, Pups which were too small to.weararadio-collar at the time of release were also implanted with abdominal transmitters.

Species Response

Between September 1987, to mid- October 1990, 29 captive-born wolves (19 adults and 10 pups) were released on 13 occasions (Smith and Phillips 1987, Phillips 1988, Phillips and Parker 1988). In addition, a minimum of six pups born in the wild during 1988 and 1990 were monitored by USFWS personnel (Phil- lips 1989). Wild-born offspring are irrefutable evidence that captive-born- and-reared adults can make the transition from captivity to life in the wild. As of mid-October 1990, a minimum of 19 wolves were free-ranging in northeastern North Carolina. After varying lengths of time, most lodged in his trachea, one drowned after consultation between the USFWS and released wolves settled into home ranges being captured in a leghold trap set by a interested groups. The USFWS briefed that varied in size from 50 km2to 100 fur trapper, and four drowned after appar- representatives of environmental organi- km2, small home ranges were situated in ently trying to cross a wide expanse of zations in Washington, DC, the North agricultural areas, whereas large home water north of ARNWR. The USFWS Carolina Congressional Delegation, the ranges were situated in forested regions. believes it is a measure of the program's North Carolina Department of Agricul- Mated pairs actively defined home success that all the deaths were natural or ture, the Governor's office, local offi- ranges through scent-marking and howl- accidental, and apparently not the result cials, and local landowners. The U.S. Air ing. These behaviors, along with intraspecific strife, suggest that reintroduced red wolves will occupy exclusive home in part to enlist support from local sportsmen, the ranges. As aresult, sociality is expected ". .. to significantly affect the size of the rein- USFWS decided to permit hunting and trapping of troduced population. other game species in the reintroduction area, even Reinvoduced wolves were crepuscu- though such activities may result in the accidental lar and more active at night than during 'take' of a red wolf." the day. Analysis of 1,100scats indicated that white-tailed deer (Odocoileus vir- ginianur), raccoon (Procyon lotor), and marsh rabbits (Sylvilaguspalustris) were of a citizen acting irresponsibly, moti- Force and Navy were briefed because important food items. Because wolves vated by some unfound hatred for they conduct training missions on 17,716 were released in areas containing no or wolves. ha adjacent to the refuge. Numerous few coyotes, biologists are uncertain how Clearly, vehicles are an important personal contacts were made with local these two species may interact in the source of mortality. To alert motorists to citizens, especially hunters and trappers, future. the presence of wolves along highways, in preparation for four public meetings During the first three years of the the North Carolina Department of trans- held during February 1986. reintroduction effort, one animal was portation erected red wolf crossing signs returned to captivity and not re-released, and local radio stations began to air pub- Experimental Population Status and 15 wolves died: five animals were lic service announcements. killed by vehicles, two by other wolves, The reintroduction of red wolves At the briefings/meetings, consider- one of a uterine infection, one of pleural would not have been possible without able effort was spent explaining the effusion and internal bleeding, one suffo- public support, which was cultivated in significance of the decision to consider cated after a raccoon kidney became part through considerable pre-release reintroduced wolves as members of a

80 Endangered Species UPDA TE Vol. 8 No. 1 "experimental/nonessential" population wolves have been released on three is- cal obstacles to human coexistence with (Parker and Phillips in press). The ex- lands off the southeast coast. From 1988 wild wolves. perirnentalJnonessentia1designation was through October 1990, ten animals were The revised Red Wolf Recovery Plan promulgated under the 1982 amend- released on Bulls Island, a component of (U.S. Fish and Wildlife Service in press) ments to the Endangered Species Act the Cape Romain National Wildlife Ref- calls for maintaining 330 animals in (Public Law No. 97-304), and provided captivity and 220 animals in the wild at a the USFWS with the ability to relax re- - minimum of three different mainland strictions of the Act to encourage coop- locations. Because of captive breeding eration with reintroduction projects from ". .. insufficient habitat and and subsequent reintroductions, the red those likely to be affected by them. For habitat destruction per se wolf recovery project enjoys tremendous example, in part to enlist support from are not the limiting factors local, regional, and national support. local sportsmen, the USFWS decided to for red wolf recovery . . . USFWS personnel working on the proj- permit hunting and trapping of other ect are confident that current momentum game species in the reintroduction area, The problem is that much will thrust the species to the edge of even though such activities may result in of the available suitable recovery within 10 to 20 years. the accidental "take" of a red wolf. The habitat in the southeast USFWS decided that prosecution would U.S. is privately owned. Literature Cited not be pursued when taking of a red wolf Landowner support is was unavoidable, unintentional, or did Carley, C.J. 1975. Activities and findings of the not result from negligent conduct -pro- therefore required. . ." red WOK recovery program fmm late 1973 to July 1, 1975. fish and Wildlife Service, vided that the incident was reported Albuquerque, NM. 215 pp. immediately to the refuge manager (or Crabtree, R.L., F.G. Burton, T.R. Garland. D.A. other authorized personnel). The uge- off the coast of South Carolina; cur- DaCatalo, and W.H. Rickard. 1989. Slow- USFWS further decided that wolves release radio isotope implants as individual rently two animals inhabit the markers for carnivores. Journal of Wildlife could be taken in defense of human life, During July 1989, nine wolves were re- M~~~~~~~~~57:949-954. but not to prevent or reduce depredations leased on Horn Island, a commnent of ~cCarley,H. 1%2. The taxonomic status of wild (e.g., of livestock, chicken, or pets). In the Gulf Island National seashores off cab- (Ca'dae) in the southcentral United States. Southwest Naturalist 7:227-235. of instances of depredations, citizens are re- the coast cmentl~five Mc&dey, H,and C.J. &ley. 1979. Recent quired to contact USFWS or state conser- animals inhabit the island. During Au- changes distribution and status of wild red vation officers authorized to institute gust 1990, four wolves were released on wolves (Canis r@). End. Species Rep. No. 4. control measures. Fortunately, to date no the St. Vincent National Wildlife Ref- UrS. Vsh and Wildlife Service. Albuquerque, NM. 38 pp. depredations have occurred. 'gey an off the coast of Parker. W.T.. and M.K. Phillips. In press. Appli- Florida; all four are still free-ranging. cation of the ex~erimentalmulation. . desiena- tion to recove; of endangered red wokes. Program Success The island proiects'- primary. objective" is to provide young wild-reared wolves for Society Phillips, M.K. 1988. Progress of the red wolf Not surprisingly, the reintroduction mainland reintroduction sites. Currently ,,,, tion pmjea in North Pages project attracted considerable interest the USFWS is planning to initiate a sec- 426-433 in R.O. Wagner, ed. Proceedings from the media and private citizens and mainland project in the Smoky Annual Meeting American Association Z& (Phillips 1990). Since the fall of 1986, a Mountains National Park during August Parks and Aquaria. Oglebay Park, -- - Wheeline. WV. minimum of 22 magazines and 24 news- 1W1. ~hillips,M.K. 1989. Bom in the wild. 'Wildlife in papers published stories about the proj- Because red wolves are "generalists" North Carolina 53%-25. ect. In addition, the project was dis- (ex., in their diet reauiremenk. thev can Phillip. M.K. 1990. Media and plblic involve- r. . cussed during the nightly newscasts of in wide'variety of habitats* ment in red wolf restoration Pages 85-98 in B. flooksha Holada, ed. Proceedings Arizona Wolf Sympo- five national and four regional television Hence, insufficient habitat and habitat sium,~~~~h 23-24, 1990, T~~~,A~ networks, and was the focus of four tele- destruction per se are not the limiting Phillips, M.K., and W.T. Parker. i988. Red wolf vision documentaries. factors for red wolf recovery; there is RcOVeV: a Progress rePo*. Comervation Biology 2:139-141. Thirty-three private citizens donated of plenty meet the Smith, R.S., and M.K. Phillips. 1987. &pive approximately 10,000hours of volunteer population recovery objectives as out- breedine and reintroduction of--~~ red wolves-- in the~--- time to the project. Local civic groups lined in the USFWS recovery plan. The wilds ofl~orthCamlina. Pages 82-90 in R.O. helped with fund-raising. Three land- problem is that much of themavailable Wagneh *. Prodings -~nnualMeeting owners entered into agreements with the suitable habitat in the southeast U.S. is American Association Zoological Parks and Aouaria. OelebavPark.-. . WheelinemWV. USFWS that provided wolves access to privately owned. Landowner support is U.S. iish and Wildlife Service. In ;is. Revised an additional 20,000 ha adjacent to the therefore required for successful reintro- red wolf recovery plan. U.S. Fish and Wildlife refuge. The restoration project now duction initiatives. Thus, unlike most Se~ce* GA. covers approximately 106,000 ha of fed- endangered species, recovery of the red Michael K. Phillips is a Wildlife Biologist for the eral and private land. wolf is not so much dependent on the Red Wolf Recovery Project at the Alligator River Success at ARNWR spawned several setting%ideofUndistutbedhabitat as it is National Wildlife Refuge, USFWS, PO Box 1969, ancillary projects. For example, red on overcoming the political and logisti- Manteo, NC 27954.

Vol. 8 No. 1 Endangered Species UPDATE 8 1 The Conservation Program for the Golden Lion Tamarin, Leontopithecus rosalia by Devra G. Kleiman, Benjamin B. Beck, Andrew J. Baker, Jonathan D. Ballou,------Lou Ann Dietz, and James M. Dietz

The Golden Lion Tamarin Conserva- captive population to preserve genetic forestation prior to the creation of the tionbgram's (GLTCP) ultimate goal is variability; Reserve, only an estimated 3,000 hec- the survival of the golden lion tamarin 3) the development of a public educa- tarescurrently offers habitat suitable for and the preservation of Atlantic Coastal tion program involving the local and na- tamarins. Although there have been no Rainforest habitat. This is a unique mul- tional community to gain support for the known cases of poaching of wild tamar- tidisciplinary effort, involving pure re- conservation of species and natural habi- ins in the Reserve since 1983, tarnarins search and applied conservation. The tats, and the training of Brazilian nation- are still being poached and are entering als in the area of conservation biology the illegal animal trade in Brazil. Defor- and wildlife management to meet future estation, however, has been reduced sig- needs in Brazil; nificantly in recent years. 4) the protection, management, and Because it was created from several expansion of the Poco das Antas Bio- cattle ranches, the Poco das Antas Re- logical Reserve and surrounding habitat serve is only about 40% forested. Forest areas, and research on restoration tech- regeneration was retarded due to annual niques for rehabilitatation and reforesta- fires during the dry season. However, the tion of degraded tropical habitats; construction of firebreaks in the mid- 5) the development and refinement of 1980s exerted some control over the techniques for reintroducing captive- spread of fires until February 1990, born golden lion tamarins into the wild, when an especially dry wet season, to- and translocating wildborn tamarins into gether with strong winds, resulted in a protected habitats; and severe fire that destroyed regenerating 6) biological studies of species sympa- vegetation in an area covering over 25% tric with golden lion tamarins in order to of the Reserve. Studies by biologists understand better the community ecol- from the University of Georgia are docu- ogy of the Atlantic Coastal Rainforest, to menting forest regeneration and the evaluate the condition of available habi- effects of the fue. tats for tamarin translocations and rein- The GLTCP also supports studies of troductions, and to increase our under- other species endemic to the Atlantic standing of the life histories of other rare, Coastal Rainforest of Brazil, partly to endangered or endemic fauna of this increase our knowledge of some of these region. rare and endangered forms, but also to understand better the animal and plant The Wild Population community in which golden lion tamarins live. Colleagues have conducted Historically, golden lion tamarins projects on the endangered maned sloth were distributed along the lowland (Bradypus torquatus) and the little- coastal forest of Rio de Janeiro State, known bamboo rat (Kannabateomysam- possibly extending into southern Espirito blyonyx). Santo. Deforestation in this area for ag- The current population estimate of riculture, timber, and charcoal has re- golden lion tamarins in the Poco das duced available golden lion tamarin Antas Reserve is 290. This estimate has separate components of the program habitat to isolatedforest patches in five risen steadily since the first estimate of include: municipalities within Rio de Janeiro 75 in the late 1970s. This trend 1) studies of the demography, popula- State. Few of these patches exceed 1,000 probably reflects both better information dynamics, behavioral ecology, ge- hectares in size. The only officially pro- tion and a real population increase netics, physiology, nutrition, reproduc- tected area containing golden lion tamar- over the past decade. The current tion and social behavior of golden lion ins is the Poco das Antas Biological estimate of the population outside the tamarins in the wild; Reserve in the municipality of Silva Jar- Reserve, excluding reintroduced 2) biological studies of captive golden dim. The Reserve is approximately animals, is 550, distributed among lion tamarins and the management of the 5,300 hectares in area, but because of de- two government-owned areas of 800

82 Endangered Species UPDA TE Vol. 8 No. 1 and 1,500 hectares, and a number of Between 1972 and 1975, the popula- commercial trahsactions. In the same privately-owned areas. tion grew slowly, but demographic year, the golden lion tamarin was in- Studies initiated in 1983 on the basic analyses suggested that it was still cluded as one of the first species in the biology of Leontopithecus rosalia have headed for extinction. However, re- American Association of Zoological resulted in extensive information on search focusing on the behavior, nutri- Parks and Aquariums' Species Survival habitat requirements, territory size, and tion and husbandry of the species finally Plan. social structure. This information has began to have an impact. Major advances By 1983, the captive population had been critical to identification of potential were made when findings revealed that grown to over 370 individuals. Efforts habitat (e.g ., for reintroduction and reproduction is most successful when were begun to slow the population translocation), estimates of currentpopu- animals are maintained in monogamous growth (which was as high as 25% per lations, formulation of survey tech- pairs and offspring are kept in the family year) by reducing the number of breed- niques, and planning strategies for inte- groups to assist parents in rearing ing pairs, and controlling the number of gration of native and reintroduced popu- younger siblings. In addition, the high young produced by each pair through the lations. Also, through these studies, we protein requirements of tarnarins became use of contraceptive implants. have established that the Poco das Antas clear, resulting in a major diet change Today, the captive population of population is demographically healthy, for the zoo population. golden lion tamarins is the most inten- with a high birth rate and an infant sur- With a better understanding of the sively managed global population of any vival rate higher than that seen in the cap husbandry requirements of the species, species in captivity. The population tive population. However, the consists of over 560 animals distributed ~ese&eis not large enough to in 119 institutions worldwide. Popula- maintain a genetically viable tion management, based on demographic population of tarnarins over and genetic analyses of the population, the next 50 to 100 years with- determine who is to be paired with out inbreeding problems. Currently, the Reserve population is probably at or "Today the captive near the carrying capacity of population of golden lion the habitat. tamarins is the most History of the Captive intensively managed Population global population of any species in captivity." The collaborative management of the captive population of golden lion tamarins began whom, how many offspring they are to in 1972, when the Wild Ani- produce and when. mal Propagation Trust con- vened an an international con- The Reintroduction ference to evaluate the status of the golden lion tamarin in Once the breeding of the captive captivity and the wild. At that population had achieved some degree of time, there were fewer than 80 regularity and control, discussions were individuals in captivity, and initiated in 1982 with the Brazilian gov- colonies at different institutions were reproduction increased. In 1981, an In- ernment (IBDF), the Rio de Janeiro Pri- managed independently of each other. ternational Management Committee was mate Center (CPRJ-FEEMA), and other Mortality and natality rates indicated formed to coordinate the global manage- interested parties concerning the possi- that the population was not self-sus- ment of the captive population and assure bility of starting a reintroduction pro-

I taining. that it was managed according to sound gram using captive-born animals that Following the conference, staff from genetic and demographic guidelines. were surplus to the needs of the captive the National Zoo initiated an intensive Strict husbandry and management proto- population. These institutions and their research program on captive individuals cols were developed for institutions staffs have worked and continue to col- to determine the cause of the mortality holding golden lion tamarins. Any insti- laborate closely with us. and reproductive problems plaguing the tution interested in acquiring the species Between 1984 and September 1990, population. At the same time, an Interna- was required to'apply to the Committee we have reintroduced 75 zoo-born and tional Studbook was developed by for approval, and sign a Cooperative six wild-born golden lion tamarins. The Marvin Jones to track information on the Research and Management Agreement. wild-born tamarins had been captured pedigree and life-history data for each An essential element of the Agreement illegally as pets and subsequently confis- individual in the population. was that no tamarins were to be used in cated by Brazilian authorities. Twenty-

Vol. 8 No. 1 Endangered Species UPDATE 83 "Population management, based on demographic and genetic analyses of the population, determine who is to be paired with whom, how many offspring they are to produce and when."

Zoo Keeper Gene Maliniack at the National Airport with crated tamarins Photo: Jessie Cohen, Natiobal Zoological Park, Smithsonian Institution seven of the zoo-borns and four of the Publlc and Professlonal population, resources, and setting for our wild-borns still survive. There have Education activities -- a process somewhat like been 39 births in the wild to reintro- market research. To understand the duced parents; 26 infants survive. One of the major goals of this program population better, we conducted inter- Reintroduced tamarins and their off- is the training of a cadre of professional views about local knowledge, and atti- spring live in 15 groups with reproduc- Brazilian conservation biologists. We tudes and behavior towards the forest, tive potential in the Poco das Antas Bio- have been recruiting students to the local wildlife, and the Reserve. We also logical Reserve and on seven adjacent GLTCP on a regular basis and have made determined that the best methods for privately owned ranches. Some live to- our facilities available for several wild- reaching the local community were tally without provisioning; others are life management and conservation biol- through the mass media, i.e., television visited and fed by observers daily. ogy training programs. We also are col- and radio. To build a positive relation- Losses of reintroduced tarnarins have laborating with the Universidade Federal ship with the community, we encouraged resulted from starvation, exposure, de Minas Gerais in Belo Horizonte. community involvement in the planning disease, predation, bee sting, snakebite, In addition, we have, as an important and implementation of project activities, wounding in intraspecific aggression, part of our activities, a community educa- and began developing interactions with and theft by humans. Most losses tion program. Without the collaboration all sectors of the community, including were due ultimately to lack of of the people in the region where golden teachers, politicians, private land own- recognition of natural foods, lack of lion tamarins still exist, all other efforts ers, businessmen, etc. recognition of non-avian predators and will have limited results for the long-term We then selected educational mate- dangerous non-predatory animals, conservation of the species. This com- rials and activities which interested the deficient locomotion, and poor spatial munity education program is developing local leaders and which seemed most orientation. a model for the solution of conservation likely to have the widest results for the Pre-release training in such skills, problems locally, nationally in Brazil, least cost. We produced printed material which was very labor-intensive, con- and internationally. for schools, did audio-visual and live ferred only a transient advantage and was The education program is best un- presentations, and developed a variety of deemed not cost-effective. Instead, derstood in terms of a systems model; the materials after first testing prototypes. tamarins are now released shortly after model helps to focus efforts on priority We worked with a single community shipment to Brazil with little training or problems, systematically develop solu- initially to test the effectiveness of our acclimation. They are provisioned and tions, and ensure thecontinuedeffective- programs, and then expanded ow reach. otherwise supported intensively for 18 ness of the solutions. We first defined the We have, to date, conducted activities months as they learn to forage, locomote priorities by addressing local conserva- with schools, farmworkers, Reserve and orient in the forest itself. tion problems. Specific objectives guards, and the press, to name a few Survival is inversely correlated with which we set include: reducing defores- specific groups, as well as the entire age. Intact family groups with two to tation, assuring the permanent conserva- community through larger events such four subadults survive longer than adult tion of privately-owned remnant forests as parades. pairs. Pairing a previously reintroduced in the region, reducing fire in the Poco The education program sought to subadult with an opposite-sexed new das Antiis Reserve, reducing the illegal evaluate the effectivenessof the separate subadult reintroductee increases survival commerce of golden lion tarnarins, and components of the program by a second and accelerates natural foraging and lo- reducing illegal hunting in the Reserve. set of questionnaires after two years of comotion. We then identified and evaluated the community activities. Results suggested

84 Endangered Species UPDA TE Vol. 8 No. 1 that there were significant changes in the the Friends of the National Zoo, World Wildlife Forman, L, D.G. Kleiman,R.M. Bush, J.M. Dietz, attitudes and knowledge of local Brazil- Fund, National Science Foundation, Wildlife Pres- J.D. Ballou, L Phillips, A.F. Coimbra-Filho, ervation Tmst International, National Geographic and SJ. O'Brien. 1986. Genetic variation ian adults. For example, to support a Society, the Frankfun Zoological Society, and the within and among lion tamarins. Amer. J. Phys. species, a public needs to be able to US Fish and Wildlife Service. We also thank the Anthro. 71:l-11. identify it. There was a significant in- local, state and federal offices of IBAMA; the Rio Kleiman, D.G. 1977. Progress and problems in & Janeiro hateCenter (CPRJ-FEEMA); the lion tamarin, Loontopirhecus rosalia rosalia, c-e in the percentage of people who Brazilian Foundation for Nature Conservation reproduction. Int. Zoo Yearbook 17:92-97. recognized and could identify a photo- (FBCN); and the US Consulate in Rio de Janeim. Kleiman, D.G. 1977. Characteristics of reproduc- graph of a golden lion tarnarin two years We are especially grateful to Adelmar Coimbra- tion and sociosexual interactions in pairs of lion after the initiation o'f our educational Filho and A1cide.s Pissinatti (CPRJ-FEEMA) and tamarins (Loontopithecur rosalia)during the re- Dionisio Pessamilio, Director of the Poco das produdive cycle. Pages 181-190 in D.G. program. We are now in the stage of Antas Biological Reserve, and to the many indi- Kleiman, ed. The Biology and Conservation of altering our programs in keeping with our viduals who havecontributed theirtime and energy the Callitrichidae. Smithsonian Press, Wash- survey results to improve the cost-effec- to this program. ington, DC. tiveness of our activities, expanding our Kleiman, D.G. 1981. Lcontopithcus rarcllia. Additional Sources of information Mammalian Species 148:l-7. activities to other sites, and training indi- Kleiman, D.G. 1982 Cooperative research and viduals who can begin to develop compa- management agreement for the golden lion Ballou,J.D. 1985-1988. 1983-1987. International rable programs in other regions of Bwil. L. tamarins. AAZPA Newsletter 2316. Studbock for the Golden Lion Tamarin, Kleiman, D.G. 1983. The behavior and conserva- rosalia rosalia. National Zoological Park, DC. tion of the golden lion tamarin, leontoplhcus r. Conclusion Washington, raroliia. Pages 35-53 in M. %ago de Mello, ed. Golden Ballou, J.D. 1989. 1988 International A Primatologia No Brasil. Sociedade Brasileira Lion Tamarin Studbook. National Zoological de Primatologia, Brasilia, DF. In the case of the GLTCP,we feel that Park, Washington, DC. Kleiman, D.G., ed. In press. Conservation Biology the combination of activities and the in- Ballou, J.D. In press. Small population manage- of the Golden Lion Tamarins: A Case Study. ment: Contraception of golden lion tamarins. In teraction among the separate compo- Smithsonian Institution Press, Washington, U.S. Seal and W. Amand, eds. Contraception in nents of the program have resulted in DC. Wildlife. Kleiman, D.G., J.D. Ballou, and R.F. Evans. 1982. much greater success towards our ulti- Beck, B.B., J.M. Dietz, D.G. Kleiman, M.I. Castro, mate goal of preserving golden lion Luz. Pro- An analysis of recent reproductive trends in R.M. Lemos de Sa, and V.L 1986. captive golden lion tamarins, Lcontopirhecus r. jecto mico-leao. IV. Reintroducao de micos- tamarins and the Atlantic Coastal Rain- rosalia, with comments on their future demo- forest than if we had concentratedon only leoes dourados (Lcontophhecus rosalia, Lin- graphicmanagement Int. Zoo YIMr. 2294-105. naeus, 1776) (Callitrichidae, Primates) de ca- Kleiman, D.G., B.B Beck, J.M. Dietz, LA. Dietz, one or two aspects. We also believe that tiveiro par seu ambiente natural (Reintrodudion J.D. Ballou, and A.F. Coimbra-Filho. 1986. this multidisciplinary effort is a model of golden lion tamarins into their natural habi- Cmservation program for the golden lion tat), Pages 243-249 in M. Thiago de Mello, ed. worth following for any organizations tamarin: Captive research and management, A Primatologia no Brasil, 2. Sociedade Brasil- intending to pursue a captive breeding/ ecological studies, educational strategies, and eira de Primatologia, Brasilia, DF. reintroduction. Pages 959-979 in K. Be- reintroduction effort in future. Beck, B.B., and M.I. Castro. In press. Environ- nirschke, ed. Primates: The Road to Self- menu for endangered primates. In E.F. Gib- Sustaining Poplations. Springer-Verlag, NY. bons, ed. Naturalistic Captive Environments Acknowledgements Kleiman, D.G., RJ. Hoage, and K.M. Green. for Animal Behavior Research. State Univer- 1988. The lion tamarins, genus Loontopirhecus. isty of New Yo& Press, Albany, NY. This program has been supported by the Na- Pages 299-347 in R.A. Mittermeier, A.B. Ry- Bronikowski, E.J., Jr., B.B. Beck, and tional Zoological Park, Smithsonian Institution, lands, A.F. Coimbra-Filho, and G.A.B. da Fon- M. Power. 1989. Innovation. exhibi- sea, eds. Ecology and Behavior of Neotwpical tion, and conservation: Free-ranging Primates, Vol. 2. World Wildlife Fund, Wash- tamarins at the National Zoological ington, DC. Park. Pp. 540-546 in 1989 AAZPA Kleiman, D.G., B.B. Beck, J.M. Dietz, and L.A. Annual Conference Proceedings. Dietz In press. Costs of reintroduction and Dietz, LA. 1985. Captive-bom golden criteria for success: Accounting and accounta- lion tamarins releasd into the wild: A bility in the Golden Lion Tamarin Conservation report from the field. Primate Conser- Program. In J.H.W. Gipps, ed. Beyond Captive vation 6:21-27. Breeding: Reintroducing Endangered Species Dietz, J.M., A.F. Coimbra-Filho, and to the Wild. Oxford Univeristy Press, Oxford. D.M. Pessamilio. 1986. Projecto mico-leao. I. Um modelo para a con- servacao de uma especie ameacada & extincao (The golden lion tamarin Devra G. Kleiman, Assistant Director for Re- project as a model for the wnserva- search*; Benjamin B. Beck, Associate Director for tion of an endangered species). Pages Biological Programs*; Andrew J. Baker, Cum- 217-222 in M. 'Ihiago de Mello, ed. A riai Intern*,$*; Jonathan D. Ballou, Population Primatologia no Brasil, 2. Sociedade Manager+; Lou Ann Dietz, SeniorProgram Officer Brasileira de Primatologia, Brasilia, for Brazile**; and James M. Dietz, Assistant Pro- DF. fessor** work for the following institutims: *Na- Diet& L.A.;and E.Y. Nagagata. 1986. tional Zoological Park, Smithsonian Institution, Community conservation education Washington, DC 20008; ** Department of Zool- program for the golden lion tamarin. ogy, University of Maryland, College Park, MD. Pages 8-16 in J. Atkinson, ed. Build- 20740; *** Wodd Wildlife Fund, 1250 24th St. ing Support for Conservation in Rural NW,Washington, DC 20037. Areas - Worltshop Proceedings, Vol. 1. QLF-Atlantic Center for the Environment, Ipswich. MA.

Vol. 8 No. 7 Endangered Species UPDATE 85 Black-Footed Ferrets On The Road To Recovery by Tim W. Clark

Introduction History of Recent May 1985, state and federal agency Restoration Efforts administrators decided to take a few fer- The black-footed ferret restoration rets into captivity later in the year if the efforthas been long, problematic, and at Discussion about how to restore ferret wild population was large enough times tenuous. However, currently the populations was part of the Meeteetse (Thome and Oakleaf in press). Unfortu- program is on the right road for full spe- ferret studies from the beginning. In nately, by the time six ferrets were cap- cies recovery. This paper outlines some 1983, Richardson et al. (1986) first for- tured, nearly all of the wild population historical milestones of the recent ferret mally examined recovery options and had died of canine distemper. After this recovery effort. Much of the progress is strongly recommended the captive-rear- tragic start to captive breeding, another summarized in an upcoming annotated ing/translocation option, and called for six ferrets were successfully taken into bibliography on the black-footed ferret an interagency meeting soon thereafter captivity; between then and February which lists 118 articles published be- to decide on a strategy. In 1984,Carpen- 1987, when the last wild ferret was cap- tween 1986 and 1990 (Reading and ter (1985: 12.1) summarizedefforts in the tured, a total of 18 wild ferrets were taken Clark in press). previous decade to captive breed ferrets into captivity. Today's ferret population The black-footed ferret (Mustela ni- and concluded that "it would appear comes from these few wild founders. gripes) is a medium-sized mustelid of that a successful captive breeding pro- The fate of the species now rests on the Great Plains that appears to be an gram . . . can be realized as soon as a captive breeding and restoration. obligatory predator on prairie dogs (Cynomys spp.) (Clark 1989). Because of their presumed competition with agriculture and livestock, these rodents have been exterminated by widespread poisoning, shooting, and other means for nearly a century and have been reduced by 70 to 90+% in both numbers and area over their entire original range. These legally mandated extermination programs pose a continuing limitation on planned ferret reintroductions. About 1980, the ferret was considered extinct by many people. But discovery of a small population near Meeteetse, Wyoming, in 1981 brought hope for species recovery. Nearly 300 ferrets were studied over the next few years, with a peak population of 129 in 1984. Catastrophe struck in 1985 when naturally contracted canine distemper killed most ferrets, leading to a species low of about ten individuals in 1986. The ferret is extinct in the wild today and only exists in three captive popula- founder population . . . can be taken into Plans for Ferret Restoration tions totalling about 181 animals. Con- captivity." In 1984, Clark et a]. tinued increases in captive populations, (19857.1 1) noted that theUsingle[Meet- The ferret is federally listed under the reintroduction, and successful manage- eetse] ferret population is currently Endangered Species Act, thus the U.S. ment - including restoration and long- highly vulnerable to extinction" and that Fish and Wildlife Service (FWS) is re- term management of the prairie dog "captive breeding/translocation efforts quired to ensure its survival. In early ecosystem (Clark et al. 1989, Miller et . . . should begin this fall." Later, Forrest 1982, the FWS requested that the Wyo- al. in press) - offer the only road for et al. (1985) outlined ferret management ming Game and Fish Department restoration. and reintroduction considerations. In (WGFD) assume "lead agency" status

86 Endangered Species UPDATE Vol. 8 No. 1 for ferret recovery in Wyoming. As a Center in Front Royal, Virginia. In 1989, physiology, and captive rearing are de- result, the state controlled the pace and 25 litters and 62 surviving young were scribed in Seal et al. (1989). direction of the national recovery effort. produced. Additional ferrets were sent to By 1987, Wyoming produced a Strategic these two zoos from Wyoming. At the Relntroductlon Plan, the 199111992 objectives of which time of this writing, about 70 young have included: "Initiate experimental reintro- been born and more are possible. In late Reintroducing the Meeteetse ferrets duction of black-footed ferrets by 1991 to several new sites was part of the first and afull scale reintroduction program of conservation and research plans for the more than 50 black-footed ferrets per newly discovered ferrets (Clark 1981), year by 1992" (WGFD 1987:5). The and was formally recommended in 1983 1996 objective was to: "Reestablish at and 1985, with the reports of Richardson least 2 black-footed ferret wild popula- et al. (1986) and Forrest et al. (1985). tions in Wyoming and maintain these Reintroductionbecame formalized as the populations with a total of at least 80 basic recovery strategy in government breeding-aged adults" (WGFD 19875). plans in 1987 and 1988 (WGFD 1987, The 1988 FWS Recovery Plan objec- FWS 1988). tive is: "To ensure immediate survival of As the basis for reintroductions, the the black-footed ferret by: 1) Increasing Meeteetse ferret environment was de- the captive population of black-footed scribed, a habitat suitability model was ferrets to a census size of 200 breeding devised (Houston et al. 1986) and later adults by 1991; 2) Estiblishing a pre- revikd (Minta and Clark 1989), and breeding census population of 1,500 management guidelines were produced free-ranging black-footed ferret breed- (Clark 1986). Habitat models were first ing adults in 10 or more populations with applied to potential translocation sites in no fewer than 30 breeding adults in any Montana (Clark et al. 1987). Since then, population by the year 2010; and 3) the FWS has developed another habitat Encouraging the widest possible distri- model (Biggins et al. 1989), and applied bution of reintroduced black-footed fer- it to several sites under the auspices of the ret populations" (FWS 1988:19). Ferret Black-Footed Ferret Interstate Coordi- increases in captivity and plans for rein- nating Committee established in 1986. troduction are close to these schedules. The Committee meets at least once 1990, additional ferrets may be sent to yearly ,and otherwise communicates Captive Breeding Efforts several more zoos in Kentucky, Colo- through a FWS Secretariat. rado, Arizona, and Canada. Future in- Several reintroduction sites have been Captive breeding efforts got off to a creases in captive populations are ex- located throughout the western United frightening start, but have since been pected. States. One of the biggest is in northcen- very encouraging. In 1985, the first six Much of the success of the captive ual Montana, estimated to be capable of ferrets taken into captivity at a WGFD breeding effort is due to advice and assis- supporting about 500 ferret families. facility died of canine distemper con- tance provided by the Captive Breeding Restoring ferrets to the wild is now a tracted in the wild, in part because all Specialist Group (CBSG) of the Intema- matter of developing successful reintro- animals were held in close proximity to tional Union for the Conservation of duction techniques and subsequent man- one another. Unfortunately, two of the Nature and Natural Resources' Species agement procedures. ferrets canied fatal distemper and trans- Survival Commission. In late 1985, Reintroduction considerations have mitted it to the remaining four. The CBSG was invited by WGFD and FWS been the subject of much discussion and second six ferrets captured in 1985 were to aid in ferret recovery. "Members of several meetings over the past few years. individually quarantined and survived. CBSG provided expertise in genetics, For example, Miller (1990) described The 1986 breeding period produced no management of small populations, hus- studies using closely related Siberian young ferrets. The last 12 ferrets in the bandry, reproductive biology and small ferrets as models to teach black-footed wild were added to captivity in 1986 and population biology" (Thorne and ferrets to avoid ground and aerial preda- early 1987. In 1987, two litters with Oakleaf in press: 14). In addition, CBSG tors and to capture prey. Beck and Miller seven surviving young increased the assisted Wyoming andFWS "in prepara- (in press) reviewed some implications captive population to 25. In 1988, 13 tion of Wyoming's strategic plan and the for ferret recovery from experience with litters and 34 surviving young were pro- revised recovery plan" and in many other the golden lion tamarin reintroduction duced. That year the population was ways (Thorne and Oakleaf in press: 14). effort. To develop a successful reintro- subdivided,with eight young going to the In 1988, Wyoming and FWS terminated duction paradigm, Reading (1990) is Henry Doorly Zoo in Omaha, Nebraska, their formal relationship with CBSG, studying a host of variables -scientific1 and seven young to the National Zoologi- feeling that their mission had been met. technical considerations, authority rela- cal Park's Conservation and Research Numerous aspects of ferret biology, tions among those involved, organiza-

Vol. 8 No. 1 Endangered Species UPDATE 87 tional aspects, and normative features- tise need to be brought in early on in the Black-Footed Fenet Workshop Proceedings, all part of reintroduction efforts. His process. Sept. 18-19, 1984, Laramie, WY. Wyoming Game and Fish Department Cheyenne, WY. work is directed at providing managers Clark, T.W., J. Grensten, M. Gorges, R. Crete, and with a readily accessible framework for The Future J. Gill. 1987. Analysis of black-footed ferret species' reintroductions, both expediting translocation sites in Montana. Prairie Natural- the reintroductions and rendering the Successful reintroduction and man- ist 19.43-56. Clark, T.W., D. Hinckley, and T. Rich, eds. 1989. process more effective. agement of the black-footed ferret is an ?he prairie dog ecosystan: Managing for biological diversity. Montana Bureau of Land Management Wildlife Tech. Bull., No. 2:l-55. Fish and Wildlife Service (FWS). 1988. Black- footed ferret recovery plan. U.S. Fish and Wild- life Service, Denver, CO. 154 pp. Forrest, S.C., T.W. Clark. L Richardson, and T.M. Campbell El. 1985. Black-footedferret habitat: Some management and reintroduction considerations. Wyoming Bureau of Land Manage- ment Wildlife Tech. Bull., No. 21-75. Houston, B., T.W. Clark, and S. Minta. 1986. Habitat sustainability index model for the black- footed ferret: A method to locate transplant sites. Great Basin Naturalist Memoirs 8:99-114. Miller, B. 1990. "Boot camp" for Siberian pole- cats. Conservation and Research Center News- letter 2(1):5-7. Many details remain to be worked out lookers alike hope ferret restoration will Miller, B., C. Wimmer, D. Biggins, and R. Read- before ferrets can be restored. Adequate go smoothly and quickly. ing. In Press. A prcposal to conserve black- footed ferrets and the prairie dog ecosystem. acclimatization of ferrets to release sites Environmental Management. will be employed. Many individual fer- Acknowledgements Minta, S.C., and T.W. Clark. 1989. Habitat suita- rets will be released at each reintroduc- bility analysis of potential translocation sites for tion site. How much pre- and post-re- Appreciation is extended to the World Wildlife black-footed ferrets in northcentral Montana. lease training is needed for reintroduced Fund-U.S., Montana Department of Fish, Wildlife Pages 29-45 in T.W. Clark, D. Hinckley,andT. and Parks. Montana Bureau of Land Management, Rich, eds. The Prairie Dog Ecosystem: Manag- ferrets is an open question. Considerable Fanwood Foundation, Nu Lambda Trust, New- ing for Biodiversity. Montana Bureau of Land assessment and observation of released Land Foundation. Lost Arrow Corporation, Cathy Management. Billings, MT. animals is imperative. These and other Patrick Foundation, and the Chicago Zoological Reading, R.P. 1990. Proposal to develop an Society. endangered species reintroduction paradigm: A details need attention. case study - the black-footed ferret. Yale Since the research and captive breed- CT. Literature Cited University, New Haven, Unpublished. 49 ing attempts of South Dakota ferrets in pp. + appendices. the 1960s and '70s, it has been widely Reading, R.P., and T.W. Clark. In press. Black- Beck, B.B., and B. Miller. In press. Implications footed ferret annotated bibliography, 1986- acknowledged by the scientific and con- for black-footed ferrets from the reintroduction 1990. Montana Bureau of Land Management, servation community that captive breed- of golden lion famarins. In Proceedings of the Wildlife Tech. Bull. No. 3. Black-Footed Fern Reintroduction Workshop. Richardson, L, T.W.Clark,S.C. Forrest,and T.M. ing and reintroduction were necessary March 29-31, 1990, Laramie, WY. components of ferret recovery. With this Campbell ID. 1986. Black-footed ferret recov- Biggins, D., B. Miller, B. Oakleaf, A. Farmer, R. ery: A discussion of some options and consid- background and the early assessment of Crete, and A. Dood. 1989. A system for erations. Great Basin Nat. Man. 8:169-184. the Wyoming ferret population status evaluating black-footed ferret habitat. Report Seal, U.S., E.T. Thome, M.A. Bogan, and S.H. Prepared for the Black-Footed Ferret Interstate and conservation needs, Richardson et al. Anderson, eds. 1989. Conservation biology Coordinating Committee. U.S. Fish and Wild- and the black-footed ferret Yale University in 1983 (1986), Clark et al. in 1984 life Service, Fort Collins, CO. 25 pp. Press, New Haven. CT. 302 pp. (1985), and Forrest et al. (1985) ada- Carpenter, J.W. 1985. Captive breeding and Thome, E.T., and R. Oakleaf. In press. Species mantly called for captive breeding to be management of black-footed ferrets. Pages rescue for captive breeding: Black-footed ferret 12.1-12.13 in S. H. Anderson and D. B. Wey, initiated early in the restoration effort. It as an example. Page 6 in J.H.W. Gipps, organ- eds. Black-Footed Ferret Workshop Proceed- izer. Abstracts From the Symposium on Be- seemed clear then and now that captive ings, Sept. 18-19, 1984, Laramie, WY. Wyo- yond Captive Breeding: Reintroducing Endan- breeding and reintroduction were the ming Game and Fish Dep. Cheyenne. WY. gered Species to the Wild, Nov. 24-25, 1989. only strategies open for species restora- Clark, T.W. 1981. The Meeteetse black-footed The Zoological Society of London, The Mam- ferret conservation studies: A proposal. Box tion. One lesson to be learned from the mal Society, and The Primate Society of Great 2705, Jackson, WY. 78 pp. Britain. 32 pp. ferret experience is that government offi- Clark, T.W. 1986. Some guidelines for manage- Wyoming Game and Fish Department (WGFD). cials controlling the pace and direction of ment of the black-footed ferret. Great Basin 1987. A strategic plan for the managanent of endangered species programs need to Naturalist Memoirs 8: 160-168. black-footed ferrets in Wyoming. Wyoming Clark, T.W. 1989. Conservation biology of the Game and Fish Dept Repon, Cheyenne. 70 pp. understand that captive breeding is a black-footed ferret. Mvsrelo nigripes. Wildlife legitimate conservation tool and needs to Preservation Trust, Special Scientific Report Tim W. Clark is President of the Northem Rockies be used in a timely manner. A second No. 3. 175 pp. Conservation Cooperative, Box 2705, Jackson, Clark, T.W., S.C. Forrest, L. Richardson, T.M. WY 83001, and an adjuna professor at the School lesson is that both captive breeding and Campbell, D.E. Casey, and K.A. Fagerstone. reintroduction can be technically diffi- of Forestry and Environmental Studies, Yale Uni- 1985. Black-footed ferret prey base. Pages 7.1 - versity, New Haven, CT 065 11. cult, thus pertinent research and exper- 7.14 in S.H. Anderson and D.B. Inkley, eds.

88 Endangered Species UPDATE Vol. 8 No. 1 Expanding the Range of Species Conservation Information

In 1976, The U.S. Fish and Wildlife Service Office of Endangered Species began publishing the Endangered Species Technical Bulletin to keep agencies, private organizations, industry, and concerned individuals abreast of developments in the federal endangered species program, at that time, the Bulletin was distributed free of charge to all who requested it. In 1981, however, federal budget cuts forced the Fish and Wildlife Service to limit distribution to federal employees and federally spon- sored researchers. This meant that the public could no longer receive up-to-date information on the listing, recovery, and status of endangered species and related federal actions and programs. - In resDonse to the cutbacks, the School of Natural Resources at the University of Michigan initiated a reprint program in 1983. Since then, the UPDATE has grown into a uniaue forum for ideas and infor-mation on endan- $. hl gered species protection. Each month, the regular version of the UPDATE combines two publica- tions in one: a reprint of the Endangered Species Technical Bulletin and a cover section produced by the School of Natural - 4 Resources. In this way, the UPDATE blends reports on the status of individual species with articles and editorials discussing a broad range of species conservation issues. To keep this important source of information available, we depend on the support and partici- pation of subscribers. Our annual subscription fee is $23 ($18 for students and senior citizens). This covers the cost of printing and mailing. We hope you decide to support our efforts and become part of the UPDATE forum.

Subscription Information

To subscribe to the Endangered Mail subscription to: Species UPDATE, enclose a check or. money order for $23 ($18 for Name students and senior citizens). Add $5 for postage if outside the U.S. Make checks payable to The Organization University of Michigan).

Return with payment to: Address Endangered Species UPDATE School of Natural Resources The University Of Michigan City / State /Zip Ann Arbor, MI 48109-1 115 ?*,, s - 'b C 4 * - * -*. -* % ,+ - -3." #$$*-<* i ,.: JlCBCI P * 8-y"- -- "The first law P%-.4", " , ,. * - aa: of intelligent .+,-* * J tinkering is to save all the pieces." -- Aldo Leopold

_ _._-______-I_(----

Non-Proftt Endangered Species UOrgan~zatron S POSTAGE PAID Ann Arbor, Mi UPDATE Permit No 144 School of Natural Resources The Untversity of Michigan Ann Arbor, MI 481 09-111 5