emp s Ridley Sea Turtle Lepidochelys kempii ead tart P I N SAndrés Her-rera lent a copy of his...

Herpetological Conservation and Biology 10(Symposium):309–377.Submitted: 7 May 2012; Accepted: 20 February 2015; Published: 28 June 2015.

Kemp’s Ridley Sea Turtle (Lepidochelys kempii) Head-Startand Reintroduction to Padre Island National Seashore, Texas

Charles W. Caillouet, Jr.1, Donna J. Shaver2 and André M. Landry, Jr.3

1Marine Fisheries Scientist-Conservation Volunteer, 119 Victoria Drive West, Montgomery, Texas77356-8446 USA, e-mail: [email protected]

2National Park Service, Padre Island National Seashore, P.O. Box 181300, Corpus Christi, Texas 78480-1300USA, e-mail: [email protected]

3Departments of Marine Biology, Wildlife and Fisheries Sciences, and Marine Sciences, Texas AM Universityat Galveston, P.O. Box 1675, Galveston, Texas 77553 USA, e-mail: [email protected]

Abstract.— Kemp’s Ridley (Lepidochelys kempii) is the most endangered sea turtle and is found inthe Gulf of Mexico and North Atlantic Ocean. It nests in greatest numbers near Playa de RanchoNuevo (RN), Tamaulipas, Mexico. Historically, nesting also occurred on beaches that, in 1962, becamepart of Padre Island National Seashore (PAIS) near Corpus Christi, Texas, USA. Kemp’s Ridley washeaded toward extinction when the Mexican government began protecting clutches of eggs (i.e., nests)and hatchlings at RN in 1966, but the population continued to decline. In 1974, the U.S. NationalPark Service (NPS) proposed reintroduction of Kemp’s Ridley to PAIS. Further planning by NPS,U.S. Fish and Wildlife Service (FWS), National Marine Fisheries Service (NMFS), Texas Parks andWildlife Department (TPWD), and Mexico’s Instituto Nacional de Pesca (INP) ensued in 1977 and ledto the bi-national Kemp’s Ridley Restoration and Enhancement Program (KRREP) implemented inJanuary 1978. Its goals were restoration of Kemp’s Ridley through enhancement of nesting successand survival at RN, and reestablishment of a breeding population at PAIS. At the time, head-start(i.e., captive-rearing to sizes thought capable of avoiding most natural predators at sea) was consid-ered essential to the second goal. Tagging and marking were necessary for identification after release,and mass-tagging hatchlings was not feasible. The NMFS Galveston Laboratory, Galveston, Texas,and collaborators head-started, tagged, and released the turtles into the Gulf of Mexico. NPS andcollaborators documented nestings. We review head-start and its relationships to the KRREP andreintroduction of Kemp’s Ridley to PAIS.

Key Words.—captive-rearing; conservation; head-starting; Lepidochelys kempii; mark-recapture; release; tagreturns; translocation; transport

Introduction

Kemp’s Ridley (Lepidochelys kempii), also re-ferred to as Atlantic Ridley, is the most endan-gered of the sea turtles. It has existed as a speciesfor 2.5–3.5 million y (Bowen et al. 1991). Its geo-graphic range is encompassed by the Gulf of Mex-ico (Pritchard and Márquez M. 1973; Zwinenberg1977; Márquez et al. 2004; Guzmán-Hernándezet al. 2007; Pritchard 2007; Ernst and Lovich2009) and North Atlantic Ocean (Brongersma

1972, 1982; Tomás and Raga 2007; Witt et al.2007; Insacco and Spadola 2010). According toNational Marine Fisheries Service (NMFS) etal. (2011), most nesting takes place on beachesbordering the west-central Gulf of Mexico. Thenesting epicenter is Playa de Rancho Nuevo (RN)near the Municipio de Aldama, in the State ofTamaulipas on the northeastern coast of Mexico(Fig. 1).

Nesting also occurs on other beaches inTamaulipas, in Texas, and southward to Veracruz;

Copyright c© 2015. Chares W. Caillouet, Jr.All Rights Reserved. 309

mailto:[email protected]

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Caillouet, Jr. et al.—2010 Head-starting Symposium: Head-starting and reintroduction of Kemp’sRidley Sea Turtle in Texas.

Figure 1. Western Gulf of Mexico showing locations of Galveston, Padre Island National Seashore, andRancho Nuevo (map prepared by Cynthia Rubio, National Park Service).

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nesting is sporadic elsewhere along Gulf of Mex-ico coasts of the U.S. and Mexico, and rare on theU.S. Atlantic coast (Márquez-M. 2001; NMFS etal. 2011). Werler (1951) was the first to report aKemp’s Ridley nesting (in 1948) in Texas, withinthe area later established in 1962 as Padre IslandNational Seashore (PAIS) near Corpus Christi,Texas (Fig. 1).

On 18 June 1947, Andrés Herrera recorded onmovie film an enormous arribada (Spanish forarrival by sea) of Kemp’s Ridley nesters near RN,representing the earliest recorded mass nestingfor this species (Carr 1963, 1967; Hildebrand1963; Burchfield and Tunnell 2004). Hildebrand(1963, 1982) estimated the size of the arribadaat 40,000 and 42,000 nesters, respectively. Carr(1967) explained how Hildebrand (1963) esti-mated the 40,000 nesters. The 40,000 estimatelater became a major benchmark among crite-ria established for Kemp’s Ridley population re-covery (U.S. Fish and Wildlife Service (FWS)and NMFS 1992; NMFS et al. 2011). Hildebrand(1963) noted that human exploitation of eggs atRN was a threat to survival of Kemp’s Ridleyarribadas and recommended that conservationmeasures be promulgated to prevent their extinc-tion.

Beginning in 1966, Kemp’s Ridley populationstatus was indexed by annual numbers of nests(clutches laid) at RN (Heppell et al. 2005, 2007;Márquez-M. et al. 2005; NMFS et al. 2011; Gall-away et al. 2013). Annual numbers of nests ontwo more beach segments, Tepehuajes and PlayaDos-Barra del Tordo, were later added to thepopulation index (Turtle Expert Working Group(TEWG) 2000; Márquez-M. et al. 2005; NMFSet al. 2011; Burchfield and Peña 2013; Gallawayet al. 2013).

Mexico’s Instituto Nacional de InvestigaciónesBiológico-Pesqueras, which later became the In-stituto Nacional de Pesca (INP; Guzmán del Próo2012), began protecting nests and hatchlings atRN in 1966 (Chavez et al. 1968), but annual nestscontinued to decline (Wauer 1978; FWS andNMFS 1992; NMFS et al. 2011). The decline re-

versed in 1986 (Caillouet 2006, 2010, 2014), withnests later increasing exponentially to more than20,000 in 2009 (NMFS et al. 2011; Burchfieldand Peña 2013; Gallaway et al. 2013; Caillouet2014). Unanticipated drops in combined annualnest numbers at the three index beach segmentsin Tamaulipas occurred in 2010, interrupting thepre-2010 exponential increase (Caillouet 2011,2014; Crowder and Heppell 2011; Allen 2013;Gallaway et al. 2013).

Our description of early attempts to reintro-duce Kemp’s Ridley to the lower coast of Texasis based on sources listed in Table 1. Andrés Her-rera lent a copy of his movie to Fred Lockettwho showed it at a meeting of the Valley Sports-men Club, Brownsville, Texas, in 1962, and thefilm was discussed during subsequent meetings.Club member Grover Singer began promoting theidea of starting a Kemp’s Ridley nesting colonyon South Padre Island (SPI) as an attraction forresidents and tourists. When Brownsville build-ing contractor Dearl Adams became president ofthe club, Grover Singer visited him frequentlyand further promoted the idea. Dearl Adams be-came more interested in Kemp’s Ridley afterreading the first chapter of Carr (1956). Laterhe talked with Henry Hildebrand whom he hadheard was transplanting Green Turtle (Cheloniamydas) hatchlings on the north end of Padre Is-land. Information provided by Henry Hildebrandwas enough to get Dearl Adams interested froma conservation standpoint.

In 1963, 98 Kemp’s Ridley eggs were ob-tained by Dearl Adams from Francis McDonaldwho operated a fishing camp called Campo An-drés at Barra del Tordo south of RN (see Fig. 2in Márquez-M. et al. 2005; see also Fig. 3 inBurchfield and Peña 2013). They were flownto Brownsville and transported to SPI, where91 were reburied on the beach and seven werekept in a tub of sand by Breuer (1971). Someeggs started development, but no hatchlings wereproduced. During 1964–1967, Dearl Adams andothers collected more than 5,000 eggs at RNand reburied them at SPI (Phillips 1989; Size-

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Table 1. Sources covering early attempts to reintroduce Kemp’s Ridley (Lepidochelys kempii) to the lowercoast of Texas.

Adams (1966, 1974) Phillips (1989)Breuer (1971) Sizemore (2002)Zwinenberg (1977) Burchfield and Tunnell (2004)Francis (1978) Burchfield (2005)

more 2002; Burchfield 2005). Among those as-sisting Adams were Kavanaugh Francis, Earland Olive Lippoldt, Ila Loetscher (who laterbecame recognized as the Turtle Lady of SPI)and others (Sizemore 2002; Burchfield 2005).The eggs were transported by vehicle to SPIor flown to Brownville then transported by ve-hicle to SPI. During 1964–1967, 1,227 hatch-lings were released into the Gulf of Mexicofrom SPI, with most (1,102 or 90%) released in1967 (Zwinenberg 1977; Phillips 1989; Sizemore2002; Burchfield 2005). At that time, Kemp’s Ri-dley was thought to reach maturity in 5.5–8 y(Márquez 1972; Pritchard and Márquez 1973;Francis 1978).

Volunteers set up camp in 1973 and begandaily patrols of the SPI beach from mid-Aprilthrough June to search for evidence of adult fe-males, and this continued annually through 1976.One Kemp’s Ridley nesting was documented in1974 and two in 1976. During 1974–1976, ad-ditional evidence of adult females in the areaincluded sightings of nesters, turtle tracks on thebeach, strandings of dead adult females (somemutilated), hatchling emergences, and one adultfemale caught (escaped unharmed) in a gill net.Francis (1978) believed the nestings resultedfrom the 1967 release of hatchlings, stating thatthere were no Kemp’s Ridley conservation pro-grams on SPI prior to 1967; he apparently dis-counted the combined releases of 125 hatch-lings (i.e., 1,227-1,102 = 125) from SPI during1964–1966. However, the observed nestings andother evidence of adult females in the SPI areaduring 1974–1976 could have been unrelated tothe hatchling releases by Dearl Adams and others(Donna Shaver, pers. obs.); e.g., they might have

involved surviving adult females from the pre-1966 residual population, or young adult femalesfrom early conservation efforts at RN.

Kemp’s Ridley Restoration and EnhancementProgram Planning Documents and Planners

Documents and planners.—Beginning inAugust 2010, one of us (Caillouet) obtainedinformation via e-mail exchanges, telephoneconversations, or both with six of the majorparticipants in the early planning (Table 2).We and Shaver and Caillouet (2015) alsoexamined unpublished planning documentsarchived at PAIS headquarters (copies havesince been archived at the National Park Service(NPS) Technical Information Center, Denver,Colorado):(1) NPS. 1974. Natural Resources ManagementPlan for Padre Island National Seashore. Pre-pared by PAIS Staff and Southwest Region Officeof Natural Science. Division of Natural Sciences,Southwest Region, NPS, Department of Interior,23 December 1974. Roland H. Wauer (ChiefScientist) recommended this 5-y plan, JohnW. Henneberger (Associate Regional Director,Professional Services) concurred, and T. R.Thompson (Acting Regional Director) approvedit; all were in the NPS Southwest Region, SantaFe, New Mexico.(2) Campbell, H.W. 1977. Feasibility Study:Restoration of Atlantic Ridley Turtle (Lepi-dochelys kempii) as a Breeding Species on thePadre Island National Seashore, Texas. Prelimi-nary Report (USNPS Order # PX7029-7-0505),Gainesville Field Station, National Fish andWildlife Laboratory, Gainesville, Florida. 2

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Table 2. Six of the major participants in early planning of the Kemp’s Ridley (Lepidochelys kempi) Restorationand Enhancement Program who were contacted by email, telephone, or both for information. (participant’saffiliations in 1977 are included).

Roland (“Ro”) H. Wauer, Chief, Division of Natural Resources Management, NPS Southwest Region, SantaFe, New Mexico, U.S.A.Jack B. Woody, Chief, Endangered Species, FWS Southwest Regional Office, Albuquerque, New Mexico,U.S.A.Edward F. Klima, Director, NMFS Galveston Laboratory, Galveston, Texas, U.S.A.James P. McVey, Chief, Aquaculture Research and Technology Division, NMFS Galveston Laboratory,Galveston, Texas, U.S.A.Peter C. H. Pritchard, Vice President for Science and Research, Florida Audubon Society, Maitland, Florida,U.S.A.Réne Márquez-M., National Sea Turtle Program Coordinator, INP, Mexico City, Mexico

November 1977. 23 p. including AppendicesI-III. Prepared for NPS Southwestern Region,Santa Fe, New Mexico.(3) NPS, FWS, NMFS, and Texas Parks andWildlife [Department; TPWD]. 1977. ActionPlan Restoration of Atlantic Ridley Turtle asa Breeding Species on Padre Island NationalSeashore, Texas 1978–1988. December 1977.36 p. including Appendices I-III.(4) NPS, FWS, NMFS, TPWD, and INP. 1978.Action Plan Restoration and Enhancementof Atlantic Ridley Turtle Populations Playade Rancho Nuevo, Mexico and Padre IslandNational Seashore, Texas 1978–1988. January1978. 30 p. including Appendices I–III.(5) NPS. 1978. Environmental Assess-ment/Review/Negative Declaration, Restorationand Enhancement of Atlantic Ridley TurtlePopulations Playa de Rancho Nuevo, Mexicoand Padre Island National Seashore, Texas1978–1988. NPS Southwest Region, February1978. 9 p. Roland H. Wauer (Chief, NaturalResources Management, Southwest Regionand Coordinator, Action Plan) recommendedthis assessment, and John E. Cook (RegionalDirector, NPS Southwest Region) concurred.

Correcting misconceptions.—None of us par-ticipated in the early planning or early years ofexecution of head-start or reintroduction. DonnaShaver became involved in the reintroduction ef-

forts by NPS at PAIS in May or June 1980. In Oc-tober 1981, Charles Caillouet became involved inhead-start efforts conducted by the NMFS Galve-ston Laboratory (hereinafter referred to as Galve-ston Laboratory). André Landry and his graduatestudents at Texas A&M University Galveston col-laborated in research related to head-start withthe Galveston Laboratory beginning in July orAugust 1984.

Upon close examination of the early planningdocuments, we became aware of misconceptionsand misrepresentations within the literature aboutthe planning for reintroduction, head-start, andtheir roles within the Kemp’s Ridley Restora-tion and Enhancement Program (KRREP; NPSet al. 1978 op. cit.). Unfortunately, we perpetu-ated some of the misconceptions and misrepre-sentations in our own publications. The literaturecontains varying accounts of early planning ofreintroduction and head-start, as well as varyingopinions regarding their adequacy as experimentsor conservation methods (see EVALUATIONS).In addition, many important details of early plan-ning have been overlooked, ignored, or forgottenuntil now. Fortunately, the unpublished planningdocuments were preserved and provide impor-tant details of early planning that can now becompared to published accounts in the literature.We focus needed attention on these historicaldocuments and the planners, because togetherthey provide necessary background and perspec-

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tive essential to evaluation of reintroduction andhead-start. They show that early planning wasscience-based, thorough, ambitious, optimistic,and remarkably detailed.

Most of the literature covering Kemp’s Ridleyhead-start and reintroduction treat them as if theyare equivalent. They certainly were related butdefinitely not equivalent. Although head-startingof the 11 year-classes (1978–1988) that were pu-tatively imprinted at PAIS was essential to rein-troduction, reintroduction was not essential tohead-starting the 23 year-classes (1978–2000)that were putatively imprinted at either PAIS orRN. It is essential that readers keep this distinc-tion in mind, because most publications to whichour review refers do not make this distinction. At-tempts at imprinting were based on a working hy-pothesis that imprinting was essential to the goalof reintroducing Kemp’s Ridley to PAIS. Knowl-edge of the mechanisms of imprinting, naviga-tion, and homing in sea turtles was limited at thetime of early planning and initiation of Kemp’sRidley head-start and reintroduction (Allen 1981).Eggs and hatchlings taken for head-start and rein-troduction were intentionally exposed to environ-mental conditions that were expected to imprintthem to PAIS or RN. It also was expected that atleast some surviving females would later matureand return to nest at PAIS or RN, where eggs orhatchlings had been putatively imprinted. Hereinwe use the terms imprint, imprinting, and im-printed only for convenience. They are not meantto imply that the turtles actually imprinted toPAIS or RN, or that surviving females from thesetreatment groups were thereby predisposed toreturn to nest at the beaches where they wereputatively imprinted.

In addition, we acknowledge that conditionsto which the turtles were exposed during head-starting in captivity at the Galveston Laboratorymay have altered their behavior, performance,and survivability following release. We empha-size that head-start and reintroduction were notdesigned or conducted to test hypotheses aboutimprinting, navigation, or homing.

Head-start and reintroduction to PAIS havebeen referred to in the literature as experimen-tal, experiments, operations, projects, programs,and even ranching (Table 3). Kemp’s Ridleyhad previously nested at PAIS (Werler 1951;Hildebrand 1963; Carr 1967; NPS 1974 op. cit.;NPS et al. 1978 op. cit.; Wauer 1978, 2014).Most International Union for Conservation ofNature (IUCN) Species Survival Commission(SSC) guidelines for reintroductions (IUCN/SSC1998) were fulfilled during Kemp’s Ridley rein-troduction to PAIS. Nevertheless, characteriza-tion and adequacy of head-start and reintroduc-tion as experiments have been challenged (seeEVALUATIONS). We emphasize that head-startand reintroduction were not planned or executedas hypothesis-testing experiments (Klima andMcVey 1982; Woody 1986; Eckert et al. 1994;Pritchard and Owens 2005; Shaver and Caillouet2015), although necessary research and exper-imentation occurred in conjunction with both.Head-start and reintroduction were largely op-erational and highly manipulative (Meylan andEhrenfeld 2000), although scientific considera-tions influenced their planning and guided theirexecution, oversight, and evaluation.

Reintroduction and head-start are generallythought to have been planned as ancillary orsubsidiary parts of the KRREP, which was notthe case (Wauer 1978). In 1974, NPS developedthe first plan for reintroduction of Kemp’s Ridleyto PAIS (NPS 1974 op. cit.; Wauer 1978), andthe combination of reintroduction and head-startwas the primary focus of planning in 1977 and1978 (Campbell 1977 op. cit.; NPS et al. 1977op. cit., 1978 op. cit.). Lest this early focus bemisconstrued, recovery priorities later shiftedappropriately toward greater focus on protectingnesting beaches, nesters, eggs, and hatchlings inTamaulipas, developing turtle excluder devices(TEDs) for shrimp trawls, and promulgatingand enforcing regulations requiring TEDs inshrimp trawls (FWS and NMFS 1992; NMFS etal. 2011). Although the early planners includedhead-start as an essential part of reintroduction,

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Table 3. Sources that referred to Kemp’s Ridley (Lepidochelys kempii) head-start and reintroduction to PadreIsland National Seashore as experimental, experiments, operations, projects, programs, or ranching.

Pritchard (1976) Donnelly (1994)Wauer (1978, 2014) Ross (1999)Woody (1981, 1986, 1989, 1990, 1991) Meylan and Ehrenfeld (2000)Klima and McVey (1982) Márquez M., R. (2001)Magnuson et al. (1990) Márquez M. et al. (2005)U.S. Fish and Wildlife Service and National MarineFisheries Service (1992)

National Marine Fisheries Service et al. (2011)

they made no recommendations or commitmentsin the early plans to test head-start as a separateconservation method or management tool.

Controversy and conflict.—From their begin-ning in 1978, Kemp’s Ridley reintroduction andhead-start were controversial within the sea turtlescience and conservation communities, despitethe good intentions of the early planners, unani-mous agreement among them and their agencies,and advice, concurrence, encouragement, andoversight of well-known and respected sea turtleauthorities. Bowen and Karl (1999) described seaturtle conservation as war in the sense that deeptensions exist between science and advocacy, andsuggested that scientific findings are sometimesmisused to promote conservation goals. Tensionsdeepened when sea turtle science and advocacybegan to impinge on the shrimping industry. Iron-ically, over the years some proponents becameopponents and vice versa.

Head-start and reintroduction were drawn intothe conflict over incidental capture of sea turtlesin shrimp trawls (Magnuson et al. 1990) andNMFS regulations requiring TEDs in shrimptrawls (Condrey and Fuller 1992; Iversen et al.1993; Yaninek 1995; Epperly 2003). Magnusonet al. (1990) concluded: “Of all the knownfactors, by far the most important source ofdeaths was the incidental capture of turtles(especially loggerheads and Kemp’s ridleys)in shrimp trawling. This factor acts on the lifestages with the greatest reproductive value forthe recovery of sea turtle populations.” Life

stages with the greatest reproductive value arelarge subadults and adults (NMFS et al. 2011),but all post-pelagic Kemp’s Ridley life stages arevulnerable to incidental capture in shrimp trawls.U.S. government responses to the controversyand conflict influenced the direction and durationof various components of reintroduction andhead-start, thereby adding to the challengesof carrying out these experiments. Sea turtleconservation and research require patience,resilience, and persistence.

Components of head-start andreintroduction.—Table 4 lists some sourcesthat reviewed the components of head-start andreintroduction. Most of these components wereelucidated in the video (TPWD. 2010. Sav-ing the Kemp’s Ridley Sea Turtle. Available fromhttp://www.youtube.com/watch?v=afgsYchpD_Q[Accessed 6 December 2012]). Operationalcomponents of reintroduction combined withhead-start, based on about 2,000 or fewer eggstaken annually from RN during 1978–1988,included: (1) collecting, transporting, and incu-bating clutches of eggs; (2) attempting to imprinteggs and hatchlings; (3) transporting hatchlingsto the Galveston Laboratory; (4) head-startinghatchlings at the Galveston Laboratory tosizes thought capable of avoiding most naturalpredators at sea; (5) tagging head-started turtlesin multiple ways at the Galveston Laboratory;(6) transporting head-started yearlings (7–15 moof age) to release sites in the Gulf of Mexico oradjoining bays; (7) releasing head-started year-

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http://www.youtube.com/watch?v=afgsYchpD_Q


Table 4. Sources that reviewed the components to Kemp’s Ridley (Lepidochelys kempii) head-start andreintroduction.

Wauer (1978, 2014) Phillips (1989)Klima and McVey (1982) Marquez-M (1994)Caillouet (1984, 1987, 2000) Shaver and Caillouet (1998, 2015)Fontaine and Caillouet (1985) Shaver (2001, 2005, 2006, 2007)Fontaine et al. (1985 1988b, 1989a, 1989b) Higgins (2003)Márquez M., R. (2001) Fontaine and Shaver (2005)Caillouet et al. (1986b, 1988, 1993, 1995a, 1995b,1995c, 1997a)

lings; (8) collecting, examining, and interpretingtag returns for head-started turtles; (9) trackingsome of the head-started turtles via radio, sonic,and satellite transmitters and receivers; and (10)documenting nestings of head-started turtles inthe wild.

Table 5 lists sources covering precautions takento prevent the eggs for the reintroduction exper-iment from coming in contact with RN sand.Eggs were carefully placed in PAIS sand withinStyrofoamTM boxes and transported to PAISwhere they were incubated in the boxes. Hatch-lings that emerged were allowed to crawl downthe PAIS beach to the surf; after swimming ashort while they were scooped up in dip nets, putin cardboard or plastic boxes, and transportedto the Galveston Laboratory to be head-started.Putative imprinting at PAIS was terminated in1989 with release of the 1988 year-class of hatch-lings (Woody 1990, 1991), but components (8)and (10) of reintroduction continued and are on-going (Shaver and Caillouet 2015). Components(1) and (2) were conducted whenever Kemp’sRidley nestings were documented at PAIS andother Texas beaches, whether laid by head-started(including PAIS-imprinted and RN-imprinted tur-tles) or wild individuals (Shaver and Caillouet2015).

Head-start also included all components, but itwas conducted on 23 year-classes (1978–2000)by the Galveston Laboratory and its collaborators.For the 1989–1992 year-classes, about 2,000 orfewer hatchlings per year were transferred di-

Table 5. Sources covering precautions taken to pre-vent clutches of Kemp’s Ridley (Lepidochelys kempii)eggs, collected for the Padre Island National Seashorereintroduction experiment, from coming in contactwith Rancho Nuevo sand.

Fontaine et al. (1985 1988b, 1989a, 1989b)Fontaine and Shaver (2005)Higgins (2003)Klima and McVey (1982)Shaver (2001, 2005, 2006, 2007)Shaver and Caillouet (2015)Wauer (1978, 2014)Woody (1986)

rectly from RN (where they were putatively im-printed) to the Galveston Laboratory for head-start. In 1993, the annual number of hatchlingsreceived by the Galveston Laboratory from RNwas reduced to about 200 or fewer per year (Byles1993; Williams 1993; Caillouet 2005, 2006;Shaver and Wibbels 2007). Because head-startmethods were applied to year-classes 1993–2000,we consider these year-classes to have been head-started. However, they are referred to by NMFSas captive-reared for research purposes (Ben-jamin Higgins pers. comm.). Some individualsof the 2000 year-class were not released in 2001,but were kept in captivity for use in a PIT tag mi-gration experiment (Wyneken et al. 2010) then re-leased in 2003 (Benjamin Higgins, pers. comm.).In addition, 100 Kemp’s Ridley hatchlings col-lected from SPI in 2013 were captive-reared atthe Galveston Laboratory, then survivors were

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used in TED tests and released offshore of SPIin 2014 (Benjamin Higgins, pers. comm.). Thesearch for Kemp’s Ridleys that were head-startedor captive-reared by the Galveston Laboratoryand released into the wild has continued (Shaverand Caillouet 2015), whether or not they wereassociated with the reintroduction experiment.

All KRREP activities proposed by NPS et al.(1978 op. cit.) were to be conducted for 11 y(1978–1988), including the work at RN, PAIS,and the Galveston Laboratory (Woody 1986,1990, 1991). However, the emphasis on reintro-duction was clear. Under heading “III. ProgramActivities” (NPS et al. 1978 op. cit.), items Aand B were related to reintroduction; item C cov-ered beach monitoring to protect nesters and eggsat RN, as well as collection of 2,000 eggs forPAIS reintroduction and up to 2,000 hatchlingsfor head-start; item D covered research at RN;and items E–G covered care of hatchlings at RNand PAIS, other operations at PAIS, transportof hatchlings to the Galveston Laboratory, head-start, release, and monitoring.

With regard to accomplishments and evalua-tions of head-start and reintroduction, our reviewfocuses on activities carried out by the Galve-ston Laboratory and its collaborators. We also re-view captive breeding, because some head-startedKemp’s Ridleys from year-classes 1978, 1979,1982, and 1984 were distributed among marineaquaria for development of a captive brood stock(Table 6). Shaver and Caillouet (2015) coverKemp’s Ridley reintroduction, related conserva-tion and research on the turtles imprinted at PAISand RN, and the labor-intensive nester searchand detection program on-going at PAIS and else-where along the Texas coast.

Details of Early Planning

The first plan.—After we learned aboutRoland Wauer’s involvement in the earlyplanning, one of us (Caillouet) found out,serendipitously, that Wauer was scheduled topresent a seminar on birds at a plant nursery

Table 6. Sources covering distribution of head-startedKemp’s Ridleys (Lepidochelys kempii) from year-classes 1978, 1979, 1982, and 1984 among marineaquaria for development of a captive brood stock.

Balazs (1979)Brongersma et al. (1979)Caillouet (2000)Caillouet and Revera (1985)Caillouet et al. (1986b, 1986c, 1988)Márquez-M et al. (2005)Mrosovsky (1979)

(Martha’s Bloomers) in Navasota, Texas on 14August 2010. He contacted Wauer and attendedthe seminar to meet and interview Wauer, whograciously gave him a copy of Wauer (1999).Wauer (1999) described early planning of aproject “to provide increased protection tonesting Atlantic Ridley turtles, and to restorea nesting population to Padre Island, Texas”;the Galveston Laboratory’s participation in theproject was also described, but not referredto as head-start. However, Wauer (1978) hadalready referred to it as head-start. Wauer (pers.comm.) indicated that a Resources ManagementPlan for PAIS contained a project statementrelevant to reintroduction (Wauer 1978). Anotherof us (Shaver) located this plan (NPS 1974op. cit.) in the PAIS headquarters archives; itcontained Project No. 9 entitled Atlantic RidleyTurtle Reestablishment. She found out fromRobert Whistler (Donna Shaver, pers. comm.)that he drafted NPS (1974 op. cit.), as ChiefNaturalist for PAIS. Robert Whistler (pers.comm.) admitted that he knew little about seaturtles at the time, but learned from HenryHildebrand who influenced development ofProject No. 9, described as follows (verbatimfrom NPS 1974 op. cit.):

Project: Atlantic Ridley TurtleReestablishment

"Padre Island was once a major nestingsite of the Atlantic Ridley turtle. Accounts

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from old-time residents relate how theytraveled by wagons along the beach andhad to wait while turtles traveled fromtheir nests to the water. This speciesis no longer known to nest within theUnited States. It does nest on the coastof Mexico where it is protected, buteggs are quickly collected and sold whenauthorities are absent. The species may beendangered and should be considered forreintroduction onto Padre Island whichmay be its only protected nesting site.

Action (Research): Initiate research,and proceed with actions deemed con-ducive to the reintroduction of the species.First, vehicles must be restricted froma section of beach; vehicles should notbe permitted to drive over the nests anddestroy the eggs. At the present time,there is no feasible section. However,the Master Plan proposes closing LittleShell Beach to vehicular traffic, andthis beach could be used as a nestingarea. Investigate the desirability of acooperative agreement with Mexico forthe collection and transportation of turtleeggs. Establish liaison with other agenciesand individuals who have shown interestin the population dynamics of the Ridleyturtle. Encourage the use of Padre Islandas a natural laboratory.

Research: This study could best beundertaken by a Park Service biologist;investigations should be initiated soon ifthis species is to be saved from extirpa-tion. Research activities should include:(1) comparative habitat evaluation, (2)feasibility of collection of turtle eggs fromMexican beaches and introduction on aprotected beach of Padre Island NationalSeashore, and (3) a monitoring program.”

Oddly, Robert Whistler was not mentioned inNPS (1974 op. cit.), but Roland Wauer (pers.comm.) confirmed that Robert Whistler draftedit. Project No. 9 appears to be the earliest writ-ten proposal for reintroduction of Kemp’s Ridleyto PAIS. It provided succinct rationale and ex-plicit aims for reintroduction of Kemp’s Ridleyto PAIS and called for research that could beconsidered pre-monitoring, elements that werelater thought to have been absent in the planning(Mrosovsky 2007). A chapter drafted by RobertWhistler (Whistler, R. 2005. Padre Island Admin-istrative History Chapter Eight: Natural ResourceIssues. Available from http://www.cr.nps.gov/

history/online_books/pais/adhi8.htm [Accessed6 December 2012]); Donna Shaver pers. comm.)explained that he and Henry Hildebrand “com-bined efforts to propose the turtle project in 1974”and sought support and possible funding fromNPS and other federal agencies, but support andfunding were not forthcoming. NPS (1974 op.cit.) scheduled the reintroduction project to be-gin in the fifth year of the plan; we assume thatwas to be 1979, but it was implemented one yearearlier (NPS et al. 1978 op. cit.; Wauer 1978).

Wauer (1999) indicated that Clyde J. Jones (Di-rector, National Fish and Wildlife Laboratory,National Museum of Natural History, Washing-ton, D.C.) and he “. . . had developed the idea,initiated a feasibility study that was done by[Howard W.] Duke Campbell, and had invitedseveral sea turtle scientists to participate” as mem-bers of “the initial advisory group” that includedDrs. Archie Carr, Henry Hildebrand, and RenéMárquez. Roland Wauer (pers. comm.) clarifiedthat “the idea” was the decision made in Decem-ber 1976 to pursue further planning and imple-mentation of Project No. 9 (NPS 1974 op. cit.).Woody (1989) also confirmed that “. . . NPS pro-posed discussions of a project whose goal wouldbe to establish a nesting population of [Kemp’sRidley] sea turtles at the Seashore” (i.e., at PAIS);he also confirmed that “The possibilities of such aproject were discussed in 1976 and 1977 betweenregional representatives of FWS and NPS. . . ”

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NPS et al. (1977 op. cit.) listed members ofa Science Advisory Board (SAB), includingArchie Carr (University of Florida), HenryHildebrand (Texas A&I University), RenéMárquez-M. (University of Mexico), and PeterPritchard (Florida Audubon Society), who wouldserve as consultants. Roland Wauer and RobertWhistler participated in further planning during1977 and 1978, so it is surprising indeed thatneither NPS (1974 op. cit.) nor its Project No.9 were mentioned in any of the 1977–1978planning documents we examined. It is alsonoteworthy that NPS (1974 op. cit.) did notmention head-start or enhanced protection atRN, both of which were later recommended byCampbell (1977 op. cit.) of FWS.

Resumed planning.—Interestingly, Pritchard(1976) recommended head-starting as well asperfection and wide spread adoption of a trawlnet equipped with a wide-mesh guard that wouldkeep turtles out, to improve survival prospectsfor Kemp’s Ridley and to supplement ongoingMexican beach protection in Tamaulipas. Herecognized that captive-reared turtles mightbecome dependent on artificial feeding, losetheir proper fear of man, or fail to navigateproperly to their nesting beach once theyreached maturity; nevertheless, he consideredhead-start worth trying as an experiment. InMay 1977, Carr (1977) called for drastic ac-tion to prevent Kemp’s Ridley from disappearing:

“The species is clearly on the skids, andif present conditions continue it willshortly - in two years perhaps, or three,or five - be gone. The dramatic dropduring the 1950’s was caused by over-exploitation combined with very heavynatural predation pressures. The terminaldecline now in progress has been broughtabout by incidental trawler catch. Whenridleys were many and shrimping was lessintensive this factor was negligible. Todayit is wiping out the species.”

Table 7. Additional sources of information on earlyplanning of the Kemp’s Ridley (Lepidochelys kempii)Restoration and Enhancement Program.

Klima and McVey (1982)Woody (1986, 1989)Fletcher (1989)Dodd and Seigel (1991)Godfrey and Pedrono (2002)Caillouet, C.W., Jr. circa 1999. MarineTurtle Newsletter articles on status of theKemp’s Ridley population and actionstaken toward its recovery. Available fromhttp://www.seaturtle.org/mtn/special/kemps.shtml[Accessed 4 December 2014].

This desperate situation apparently promptedNPS, FWS, NMFS, TPWD and Mexico’s INPto begin planning the KRREP in 1977, and toimplement it in 1978 (NPS et al. 1978 op. cit.;Wauer 1978). The KRREP’s stated goals wererestoration of Kemp’s Ridley through enhance-ment of nesting success and survival at RN, andreestablishment of a breeding population at PAIS.Head-start was considered essential to the secondgoal. Early planning for the KRREP was alsodiscussed by sources listed in Table 7.

Roland Wauer played the central role inguiding and facilitating the planning during1977 and early 1978. He compiled the draftaction plan (NPS et al. 1977 op. cit.) and thefinal Action Plan (NPS et al. 1978 op. cit.) withinput provided by participating individuals andagencies (Wauer 1978). During the 1978 nestingseason, he traveled to RN (Pritchard and Gicca1978), assisted in collecting and packing eggs inPAIS sand for transfer to PAIS (Wauer 1978), andaccompanied the resulting hatchlings on theirflight from PAIS to Galveston (Roland Wauer,pers. comm.). We also learned that R. BruceBury accompanied Roland Wauer to RN in 1978to participate in oversight and implementationof efforts to ensure that the eggs to be moved toPAIS did not contact RN beach sand (R. BruceBury, pers. comm.). According to Pritchard

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and Gicca (1978), scientific visitors that werepresent for various periods during the first seasonof work associated with the KRREP includedPatrick Burchfield (Brownsville coordinator forthe project), Bruce Bury, Howard Campbell,Thomas Fritts, Roland Wauer, and Jack Woody(Project Coordinator).

Feasibility study.—The feasibility study(Campbell 1977 op. cit.) provided rationale,explicit aims, and pre-monitoring for reintro-duction (see Mrosovsky 2007). It encompassedbiological justification as well as mechanicaland political possibility for reintroduction.This study progressed in simultaneous phasesincluding: (1) documentation of Kemp’s Ridleyas a native breeding species on the PAIS; (2)review of mechanical and biological problemsassociated with moving sea turtle eggs from thenatal beach to establish new or reintroducedcolonies; and (3) contacts with all agencies andindividuals required for an operation of suchcomplexity, to determine their willingness andpotential to contribute to the effort.

The study began in May 1977 with a literaturereview and contacts with authorities expectedto possess unpublished data, including ArchieCarr, Henry Hildebrand, Frank Lund (Universityof Florida, Gainesville, Florida), Réne Márquez-M., and Peter Pritchard; all of them contributedcollectively and significantly to this data gather-ing effort. Early participation by such highly re-spected sea turtle scientists who were particularlyknowledgeable of Kemp’s Ridley is noteworthy.Ila Loetscher (Sizemore 2002) and Earl Lippoldtalso provided valuable unpublished data. In July-August 1977, physical suitability of the PAISbeach was examined by land and from the air andcompared to that at RN. Contacts also were madewith representatives of TPWD, NMFS, FWS (Of-fice of Endangered Species), and various aca-demic and conservation organizations to discusstheir interest in and potential contributions to theprogram, and the challenges of the program.

Campbell (1977 op. cit.) contained results andrecommendations for consideration in furtherplanning and commitment of resources. His wasthe first planning document to mention head-start,among those we examined. Campbell (1977 op.cit. Appendix II) recommended that head-start becombined with transplanting eggs and hatchlingsto reintroduce Kemp’s Ridley to PAIS. Clearly,tagging of head-started turtles was needed to pro-vide the essential means of linking them to head-start and reintroduction after their release. Oth-erwise, it would have been necessary to tag andrelease hatchlings to make it possible to evalu-ate reintroduction and head-start. At that time,the technology for mass-tagging Kemp’s Rid-ley hatchlings had not been developed (Pritchard1979; Allen 1981; Fontaine et al. 1993; Higginset al. 1997).

The literature review and correspondence con-ducted by Campbell (1977 op. cit.) produced noevidence of large-scale breeding aggregations ofKemp’s Ridley at PAIS, but indicated that individ-ual nestings had occurred with some regularityat PAIS over the preceding decade, not only byKemp’s Ridley but also by other sea turtle species.Hatching success of Kemp’s Ridley and otherspecies showed that eggs were fertile and phys-ical characteristics of the beach were suitablefor incubation. No obvious physical problemswere discovered regarding the PAIS beach, andstrong similarities were revealed between nestingbeaches at PAIS and RN. Beach slope and profile,sand grain size, and other physical characteristicswere essentially the same. Differences in air andwater temperatures at the two nesting sites wereminimal during the nesting season and thereforeconsidered unimportant. Mechanical and biolog-ical problems associated with transplanting seaturtle eggs were considered resolved over manypreceding years, and the process of transplant-ing eggs was considered routine for experiencedpersonnel. Moving and incubating eggs, hatch-ing them, and rearing young sea turtles had beenaccomplished throughout the world and were con-tinuing. The only area of uncertainty identified

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by the feasibility study was that of “actually es-tablishing new colonies on new beaches.” Noclearly successful transplant had previously beenachieved. Apparent failure of previous transplantattempts was attributed to poor understandingof imprinting of hatchlings to their natal beach,enormous first-year mortality of hatchlings, andlack of a suitable method for tagging hatchlingsso they could be recognized later when they wereadults. There also was concern that the popu-lation could not support any removal of eggsfrom RN for reintroduction to PAIS. Recogniz-ing that this was subjective and difficult to assess,Campbell (1977 op. cit.) recommended an ini-tial approach designed to be compensatory andminimize this potential problem. For reintroduc-tion, 2,000 eggs would be transferred from RN toPAIS and the resulting hatchlings transferred tothe Galveston Laboratory for head-start; in addi-tion, 2,000 hatchlings would be transferred fromRN to the Galveston Laboratory for head-start.Campbell (1977 op. cit.) expected that “captiverearing of as many hatchlings from the Mexicanbeach as are removed to Texas should return moreadults to Mexico than would the natural recruit-ment from the eggs removed to Texas if the “headstarting” concept has any validity at all.” How-ever, 2,000 eggs were not equivalent to 2,000hatchlings (i.e., hatch rate was less than 100%).In any case, Campbell (1977 op. cit.) apparentlyexpected hatch rate as well as survival rate dur-ing head-starting to be higher than had these eggsor hatchlings been left at RN. In retrospect, aslong as enough head-started turtles survived andreturned to reintroduce nesting to PAIS, it didnot matter whether or not head-starting increasedsurvival from hatchling to nester life stages tothe extent expected by Campbell (1977 op. cit.).It is also possible that Campbell (1977 op. cit.)included such ambitious expectations to elicitagreement from all parties. A related questionis whether Campbell (1977 op. cit.) expectedhigher survival rates in head-started Kemp’s Ri-dleys after their release than those of wild coun-terparts of the same sizes and ages, but there

is no way to answer this question. Regardless,head-starting was essential to grow the turtles tosizes at which they could be safely tagged, sothey could be linked (by their tags, after releaseinto the Gulf) to experimental reintroduction andhead-start upon recapture or nesting. Interest inand concern for Kemp’s Ridley was high amongall individuals and agencies contacted by Camp-bell (1977 op. cit.), and support among the agen-cies for the proposed reintroduction was unani-mous.

Campbell (1977 op. cit. Appendix II) listedfive options (paraphrased, but in their originalorder) for recovery of the Kemp’s Ridley popula-tion: (1) protection of the known nesting beach,(2) reduction in incidental kill by fisheries opera-tions, (3) establishment of breeding populationsin protected areas in Texas, (4) enhancement ofrecruitment into the Mexican population, and (5)establishment of a captive breeding population.Options (3) and (4) constituted the core of thereport’s recommendations. Interestingly, option(5) was considered highly desirable and perhapscritical for egg production if the natural breedingstock were lost before full recovery. Option (1)was recognized as ongoing and critical to long-term survival of Kemp’s Ridley, but in need ofemphasis. Option (2) was in progress and con-sidered necessary for survival of most sea turtlespecies, but unlikely to be realized within theexpected 2–10 y remaining for Kemp’s Ridley.Campbell (1977 op. cit.) noted that PAIS repre-sented a potentially secure breeding ground forKemp’s Ridley where disturbance to breedingand nesting could be controlled with minimumenforcement effort.

Campbell (1977 op. cit.) listed the followingfactors as the minimum necessary for potentialsuccess of the proposed reintroduction, recog-nizing that some of them were too new to befully evaluated before they were attempted: (1)incubation of eggs in sand from PAIS to avoidpossible chemical imprinting to RN sand, andessentially natural orientation exposure of hatch-lings to the natal beach and offshore waters at

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PAIS, (2) captive-rearing for 0.5–1 y to a sizeat which mortality due to predators is reduced,(3) adequate technique for marking young tur-tles to allow their recognition as adults, and (4)release of captive-reared turtles of a given year-class in an area and habitat consistent with theirage, place, and time of naturally occurring (i.e.,wild) young of the same year-class. All these fac-tors considered necessary for potential successwere operational, and much less stringent than themore elaborate evaluation criteria that were lateradded as reintroduction and head-start efforts pro-gressed (see EVALUATIONS). The above list offactors also suggests that establishing more strin-gent evaluation criteria to measure success beforereintroduction was attempted would have beenpremature, because the immediate and primarygoal was saving the declining Kemp’s Ridley pop-ulation from extinction (see Mrosovsky 2007).

Roland Wauer, Howard Campbell, and JohnC. Smith (Non-Game Project Leader, TPWD,Rockport, Texas) presented results and recom-mendations of the feasibility study to the JointSecretary’s and Southwest Regional AdvisoryBoards at Padre Island on 26 September 1977(Campbell 1977 op. cit. Appendix I). Weassume these “Advisory Boards” represented theSecretaries of the U.S. Department of the Interior(for NPS and FWS) and the U.S. Departmentof Commerce (for NMFS, within the NationalOceanic and Atmospheric Administration,NOAA), and the Southwest Regional Directorsof NPS and FWS; however, the NMFS SoutheastRegional Director was not mentioned. Afterthe 26 September 1977 presentation, TPWDexpressed interest in conducting negotiationswith government agencies in Mexico. AppendixI in Campbell (1977 op. cit.) was a veryimportant memorandum from Theodore R.Thompson (Deputy Regional Director, NPSSouthwest Regional Office) to the Directorof the NPS [whose name was not shown, butWilliam J. Whalen held the position], dated7 October 1977. Its subject was “Restorationof the Atlantic Ridley Turtle as a Cooperative

Interagency Project”; it summarized the meetingof 26 September 1977 and stated that: “The Stateof Texas already has talked with Mexico aboutobtaining turtle eggs. The Texas Department ofParks and Wildlife will host a meeting in lateNovember for agency personnel to develop aplan of attack and a timetable. I believe thatthis gathering should heavily involve Regionallevel personnel, who will follow through onactivities already started. It is particularlyimportant that the National Park Service, Fishand Wildlife Service, and the National MarineFisheries Service cooperate fully. Each agencymust play a significant role if the project is tobe successful.” This memorandum confirmedthat the State of Texas negotiated the transferof Kemp’s Ridley eggs from RN to PAIS forreintroduction, and it also mentioned the meetingthat TPWD would later host in Austin, Texason 17 November 1977, as scheduled in Camp-bell (1977 op. cit.) and NPS et al. (1977 op. cit.).

Draft action plan.—A draft “Restoration Planfor the Atlantic Ridley Turtle” was distributedby letter dated 28 November 1977 from RolandWauer to “all of the Austin participants” (Table8); the reference to “Austin participants” appar-ently referred to the 17 November 1977 meet-ing held in Austin, Texas (Campbell 1977 op.cit.; NPS et al. 1977 op. cit.). Roland Wauer didnot refer to this draft as an action plan, but hementioned head-start. Interestingly, the “Austinparticipants” represented NPS, FWS, NMFS,and TPWD, but not Mexico. Roland Wauer in-dicated that he had corresponded with “Archie,Peter, Henry and Rene” (i.e., Carr, Pritchard,Hildebrand, and Márquez-M., respectively) ask-ing them to serve on the “Science Board” (i.e.,the SAB), but did not mention whether theywere sent copies of the draft plan. We assumethey were sent copies, because Wauer’s letterrequested that recipients of the draft plan add de-tails on what was planned for the RN phase ofthe program so that he could prepare the finalplan, and he expressed hope that the plan would

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Table 8. List of all Austin participants to whom Roland Wauer sent copes of the Draft Restoration Plan forthe Atlantic Ridley Turtle (Lepidochelys kempii) via his letter 28 November 1977.

Bill Brownlee, Texas Parks and Wildlife (TPWD), Austin, Texas, U.S.A.Dr. Duke Campbell, U.S. Fish and Wildlife Service (FWS), Gainesville, Florida, U.S.A.Ted Clark, TPWD, Austin, Texas, U.S.A.John Dennis, National Park Service (NPS) Washington Support Office (WASO-550), Washington, D.C.,U.S.A.Dr. Don Eckberg, National Marine Fisheries Service (NMFS), St. Petersburg, Florida, U.S.A.Hal Irby, TPWD, Austin, Texas, U.S.A.Dr. Clyde Jones, FWS, Washington, D.C., U.S.A.Carol Justice, FWS. Washington, D.C., U.S.A.Dr. Ed Klima, NMFS, Galveston, Texas, U.S.A.Dr. James P. McVey, NMFS, Galveston, Texas, U.S.A.Floyd E. Potter, Jr., TPWD, Austin, Texas, U.S.A.John Turney, NPS PAIS, Corpus Christi, Texas, U.S.A.Robert G. Whistler, NPS PAIS, Corpus Christie, Texas, U.S.A.Jack Woody, FWS, Albuquerque, New Mexico, U.S.A.

be finalized before mid-December 1977.NPS et al. (1977 op. cit.) probably was the

draft plan mentioned in Roland Wauer’s letterof 28 November 1977 because it did not includedetails of “research to be conducted at RanchoNuevo.” Item C on p.14 of NPS et al. (1977 op.cit.) suggested that adding such research detailswould be the responsibility of Howard Campbell.The primary focus of this draft action plan(NPS et al. 1977 op. cit.) was reintroduction ofKemp’s Ridley to PAIS. Although the sectionof NPS et al. (1977 op. cit.) entitled “RanchoNuevo Beach Monitoring and Egg Collection”included protection of nesting turtles and theireggs from native and human predators during thenesting season at RN, it also included collectionof eggs at RN for reintroduction to PAIS. NPSet al. (1977 op. cit.) named Jorge Carranza(Director of INP) as a member of the AgencyCoordinating Committee (ACC), along with Don[Donald] Ekberg (Director, NMFS SoutheastRegional Office, St. Petersburg, Florida), Hal[Harold] Irby (TPWD, Austin, Texas), Ro[Roland] Wauer, and Jack Woody. NPS et al.(1977 op. cit.) indicated that members of theSAB would serve as consultants throughout thecourse of the program, and that NPS would serve

as coordinating agency for the SAB. Permitrequirements were identified, including a permitfrom Mexico, a [U.S.] Endangered SpeciesPermit and a Texas permit. This draft planemphasized that “the key to the entire programis Mexico’s participation and support” and that“Don Ekberg and a Texas representative willmeet with Mexico’s Jorge Carranza regardingthe program on December 7–9, 1977.” It statedthat “Jack Woody will initiate a request forauthorization to handle and transport eggsand turtles immediately upon completion ofthis plan,” and that “Bill Brownlee [TPWD]will initiate a request for authorization of theState of Texas upon receipt of a copy of theEndangered Species Permit application.” Agencyresponsibilities were designated as follows: (1)beach monitoring and egg collections at RNwould be the responsibility of Mexico and theFWS, “with special assistance by National ParkService personnel,” (2) incubation of eggs atPAIS and exposure of hatchings to the beachand surf at PAIS would be the responsibilityof the PAIS Superintendent, and (3) rearingfrom hatchlings until the head-started turtleswere tagged and released, as well as monitoringthereafter, would be the responsibility of NMFS.

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This early assignment of responsibility to NMFSfor monitoring the turtles following their releaseis important, especially in the context of itslater reiteration by NMFS’ National Sea TurtleCoordinator: “NMFS will place special emphasison detecting tagged turtles in the wild” (Williams1993). Beach monitoring at RN apparentlyreferred to protection of nesting turtles, eggs,and hatchlings from natural predators and humantake during the nesting season. However, itmay also have encompassed counting nests,eggs, and hatchlings. Special assistance by NPSpersonnel referred to the collection of 2,000eggs at RN, depositing them in PAIS sandwithin StyrofoamTM boxes, and transportingthem to PAIS. Another 2,000 eggs were to becollected and incubated at RN. Descriptionsof the “Rearing Project” and “Release andMonitoring” by the Galveston Laboratory wereincluded. However, other than indicating thatthe turtles would be tagged with numbered tagsand selected individuals would receive sonic tagsfor tracking, NPS et al. (1977 op. cit.) gave noadditional details regarding monitoring of theturtles after release. Also absent from NPS etal. (1977 op. cit.) were planned activities aimedat documenting head-started Kemp’s Ridleynestings at PAIS, RN, or elsewhere, althoughnestings were essential to accomplishing thestated goals of head-start and reintroduction(Shaver and Caillouet 2015). Documentingnestings of head-started Kemp’s Ridleys remainsessential to evaluating reintroduction andhead-start, together as well as separately.

Final action plan.—The finalized Action Plan(NPS et al. 1978 op. cit.) reiterated most of thedraft action plan (NPS et al. 1977 op. cit.). Withregard to the requirement that Jack Woody initiatea request for authorization to handle and transporteggs and turtles under an Endangered SpeciesPermit, a “Convention Permit” was added, appar-ently referring to the Convention on InternationalTrade in Endangered Species of Wild Fauna andFlora (CITES). No mention was made of the need

for personnel at the Galveston Laboratory to bepermitted, but they certainly were required tocarry copies of all necessary permits while en-gaged in head-starting and related activities. Thesame was true of all who collaborated with theGalveston Laboratory; e.g., NPS personnel alsohad permits from Texas and FWS. In addition,when head-started Kemp’s Ridley releases tookplace off Florida, Florida Department of NaturalResources (FDNR) permits were required.

The following responsibilities and require-ments were specified (they are paraphrased here)for the Galveston Laboratory within NPS et al.(1978 op. cit.): (1) hatchlings received fromPAIS and RN will be placed in flow-throughsystems in tanks and raceways, (2) each groupof hatchlings (PAIS and RN) will be maintainedseparately, (3) turtles will be fed a diet ofchopped, boneless, scaleless fish, shrimp, andother marine foods available, at rates of 5–8%of body weight per day or at whatever volumeof food they will consume in 3–4 h, (4) afterNovember 15, turtles will be placed in specialheated containers (with water temperatureapproximately 22◦ C) equipped with the bestdesigns of water treatment facilities to handlewastes produced by the turtles, (5) water willbe exchanged whenever it is determined tobe below quality for the turtles, (6) turtleswill be maintained for periods up to one yearthen released, (7) turtles imprinted at RN andthose imprinted at PAIS will be released atperiods up to one year, at locations and timesdetermined to be the most logical for youngturtles to occur, (8) turtles will be placed inStyrofoamTM transport boxes, kept moist, cool,and ventilated, and moved by boat to releaselocations, (9) turtles will be released on grassflats off Florida’s west coast and lower Gulf ofMexico, and other areas where yearlings havebeen observed, (10) all turtles will be taggedwith numbered tags and selected individuals willalso be tagged with sonic tags and tracked, and(11) in the second year (i.e., 1979), some 1 yold turtles will be tagged with radio transmitters

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and tracked from aircraft and satellites. TheAction Plan (NPS et al. 1978 op. cit.) gave nofurther details regarding monitoring head-startedturtles after release, or searching for and docu-menting their nestings at PAIS, RN, or elsewhere.

The captive breeding option.—Establishmentof a captive breeding colony was amongoptions considered by Campbell (1977 op.cit. Appendix II), but it was not among theactions recommended by NPS et al. (1978op. cit.). However, after NPS et al. (1978op. cit.) was implemented, Brongersma et al.(1979) recommended establishment of a captivebreeding colony of Kemp’s Ridleys at CaymanTurtle Farm Ltd. (CAY), Cayman Island, BritishWest Indies, “to ensure preservation of thisgenetic entity, if efforts to preserve the speciesin the wild should fail.” They recommendedusing existing aquarium specimens, accidentallycaught individuals, and hatchlings of the 1978year-class at the Galveston Laboratory. Balazs(1979) recommended further that the reservoirof breeding Kemp’s Ridleys be establishedthrough dissemination of hatchlings of the 1978year-class to responsible and consenting aquaria,oceanaria, and appropriate zoological facilities inthe U.S., Mexico, and other countries (Table 6).

The mysterious meeting.—Also mentionedin the literature (e.g., Klima and McVey 1982;Woody 1989), and by some of the early plannerswho were contacted in 2010 (by Caillouet), wasa multi-agency planning meeting purportedlyheld in January 1977 in Austin, Texas. Wewere unable to corroborate that such a meetingtook place at that time, based on the planningdocuments we examined. Perhaps the meetingin question was the one hosted by TPWD on 17November 1977 (Campbell 1977 op. cit.; NPS etal. 1977 op. cit.), or the one held in January 1978during which the Action Plan (NPS et al. 1978op. cit.) was approved by attendees. We areunaware of any additional planning documentsthat might corroborate a January 1977 meeting.

If it did take place, it may have been informal(Jack Woody, pers. comm.), and therefore notdocumented in a planning document or report.

Oversight.—We stress that agencies andindividuals that participated in various com-ponents of reintroduction and head-start didnot proceed without direction, approval, andoversight of their work. Each agency had internalprocedures to approve, control, and evaluateits work. NPS et al. (1978 op. cit.) requiredannual reviews to evaluate past, current, andfuture phases of the KRREP. Initially, the ACCand SAB provided oversight and guidance to allagencies participating in the KRREP. Additionaloversight and guidance were provided by aSea Turtle Working Group established in 1977within the MEXUS-Gulf Program, a formalfisheries agreement between INP and the NMFSSoutheast Fisheries Science Center in Miami,Florida (Berry 1987). References to a Kemp’sRidley Working Group (KRWG) can also befound in the literature (e.g., NMFS et al. 2011;Burchfield and Peña 2013). In any case, afterthe KRREP was initiated, a consortium ofrepresentatives of NPS, FWS, NMFS, TPWD,and INP (or its successor agencies) met annuallyto review progress and discuss and approve plansfor the next year’s work. At these meetings,oral presentations and recommendations weremade, proposals were distributed, results wereevaluated, and proposed work was discussed,modified as needed, and approved. Scientistsfrom non-government organizations (NGOs) anduniversities occasionally participated in thesemeetings. If agreement on a proposal was notunanimous, that did not necessarily prevent itsimplementation. However, participants wereusually in agreement and worked cooperativelyin making decisions and providing guidance.In some cases, annual reports were preparedand distributed (e.g., Pritchard and Gicca 1978;Pritchard 1980; Caillouet 1997; Shaver 2012;Burchfield and Peña 2013). All participatingagencies involved in the KRREP had permitting

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Table 9. Previous reviews of major accomplishments of Kemp’s Ridley (Lepidochelys kempii) head-start,reintroduction, and research activities, and reviews of Kemp’s Ridley head-start at the Galveston Laboratory.

Reviews of Major Accomplishments Reviews of Head-startCaillouet et al. (1995c) Wauer (1978, 2014)Caillouet (2000) Woody (1981, 1986)Shaver (2001, 2005, 2006, 2007) Caillouet (1984, 1987)Higgins (2003) Caillouet and Koi (1985)Fontaine and Shaver (2005) Fontaine and Caillouet (1985)Shaver and Wibbels (2007) Fontaine et al. (1985, 1988b, 1989a, 1989b,

1990a)Caillouet et al. (1986b, 1986c, 1988, 1993,1995c, 1997a)Manzella et al. (1988b)Duronslet et al. (1989)Leong et al. (1989)Shaver (2001, 2005, 2006, 2007)Shaver and Caillouet (2015)

authority over some aspect of the work. Ifsomething objectionable to a given agencywas proposed, and that agency had authorityto withhold a necessary permit, it could haveprevented implementation simply by refusingto issue the permit. A good example is FWS’prevention of annual importation of 2,000Kemp’s Ridley hatchlings into the U.S. forhead-start at the Galveston Laboratory in 1993,simply by not issuing the necessary FWS permit(Byles 1993; Williams 1993). Therefore, variouscontrols were in place to reconsider and modifygoals, objectives, and direction of the KRREP atany time.

Public awareness.—High public profiles ofreintroduction and head-start greatly enhancedawareness of the plight of Kemp’s Ridley as wellas the need for conservation of all sea turtles.Especially effective in this regard were Kemp’sRidley conservation advocacy efforts of Carole H.Allen (Allen 2013), Chairperson of the organiza-tion she founded and managed (Help EndangeredAnimals-Ridley Turtles; i.e., HEART), a stand-ing committee of the non-profit Piney WoodsWildlife Society of Houston, Texas. Carole Allenand HEART not only had strong educational fo-

cus involving children and parents, but also ad-vocated use of TEDs, raised funds, and providedfunding and other support for Kemp’s Ridleyconservation, as well as head-start and relatedresearch at the Galveston Laboratory, PAIS, RN,and Texas A & M University in College Stationand Galveston. Availability of sea turtles in cap-tivity at the Galveston Laboratory provided op-portunities for public viewing during scheduledtours and NOAA open houses in which Allenand others representing HEART sometimes par-ticipated (off site). Also effective was public ex-posure to releases of hatchlings produced fromclutches laid by head-started and wild Kemp’sRidleys at PAIS (Shaver and Caillouet 2015).

Accomplishments

The challenges of rearing marine turtles in cap-tivity are so great that Tonge (2010) consideredthem unsuitable even for laboratory usage andgave them little attention in his discussion ofmaintenance and husbandry of aquatic reptiles.Mass rearing Kemp’s Ridleys for 7–15 moin captivity was much more challenging andexpensive than rearing laboratory specimensor rehabilitating live-stranded individuals that

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are injured or ill. Sources summarizing majoraccomplishments and contributions of theNMFS Galveston Laboratory, NPS Padre IslandNational Seashore, and their collaborators tohead-start, reintroduction, and related researchare listed in Table 9. Kemp’s Ridley head-start atthe Galveston Laboratory was also previouslyreviewed by sources that provided details about(1) housing, raceways, and rearing containers,(2) seawater source, storage, quality, heating,monitoring, and replacement, (3) hygiene, (4)yolk absorption in hatchlings, (5) measuringand weighing turtles, (6) natural foods andcommercial feeds and feeding, (7) survival,(8) growth, and (9) prophylaxis, diagnosis andtreatment of diseases, all of which improved overtime (Table 9).

Collaborative research.—Availability of largenumbers of Kemp’s Ridleys in captivity, aswell as fewer Loggerhead (Caretta caretta)and Olive Ridley (L. olivacea) turtles, affordedopportunities for collaborative research at theGalveston Laboratory (Caillouet et al. 1986b,1988, 1995c; Fontaine et al. 1990a; Caillouet1997, 2000). The Galveston Laboratory col-laborated with individuals in federal and stateagencies, universities, and NGOs, as well as withveterinarians including Drs. Joseph Flanagan(Houston Zoo, Houston, Texas), Richard Hen-derson (Galveston, Texas), and Elliott Jacobson(University of Florida, Gainesville, Florida).Collaboration provided part-time employment,research experience, and career development forgraduate students, especially those associatedwith the Texas Sea Grant College Program,Texas A & M University, the University ofTexas Medical Branch, and the Louisiana StateUniversity Sea Grant College Program. TheGalveston Laboratory also participated in the SeaTurtle Stranding and Salvage Network (STSSN),providing additional part-time employment forgraduate students. Collaborative research topicsincluded hatchling yolk-absorption, naturalfoods and commercial feeds and feeding, wastes

from sea turtles and excess feed in seawater,diseases and treatments, necropsy, behavior,growth, survival, morphometrics, compositionof Rathke’s gland secretions, genetics, swim-ming performance, physiology (reproductive,metabolic, and respiratory), tags and tagging,radio-, sonic-, and satellite-tracking, geographicdistribution, strandings, rehabilitation, threatsat sea, captive breeding, TED testing andcertification, incubation temperature dependentsex-ratios, olfactory imprinting, and trace metals.Caillouet and Landry (1989) chaired the FirstInternational Symposium on Kemp’s Ridley SeaTurtle Biology, Conservation and Management,held in October1985 at Texas A & M Universityat Galveston. For copies of Galveston Laboratorypublications and reports related to head-start,reintroduction, and the KRREP, see Caillouet(1997) and (NOAA Fisheries, Galveston Labo-ratory, Galveston Publications. Available fromhttp://www.galvestonlab.sefsc.noaa.gov/publications/ [Accessed 7 December 2014]).

Initial work with Loggerhead.—Klima (1978)clearly intended that the Galveston Laboratorytest practicality of a rearing and release pro-gram as a viable conservation and managementtool, not only for Kemp’s Ridley but also forother sea turtle species including Loggerhead,Leatherback (Dermochelys coriacea), Hawksbill(Eretmochelys imbricata), and Green Turtle (seealso Bullis 1978). However, consultation withArchie Carr, Peter Pritchard, Henry Hildebrand,and Galveston Laboratory staff led to a decisionto conduct the initial head-start trial on Logger-head, a threatened species (Klima 1978; EdwardKlima, pers. comm.; Jack Woody, pers. comm.).Leong et al. (1980, 1989), Klima and McVey(1982), and Clary and Leong (1984) describedcaptive-rearing of Loggerheads at the GalvestonLaboratory during 1977. Loggerhead was clas-sified as a threatened species, and its captive-rearing allowed the staff to gain experience be-fore attempting to head-start Kemp’s Ridleys.

In September 1977, 1,060 Loggerhead hatch-

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lings were received from FDNR, Jensen Beach,Florida. They were reared in communal groupswithin large tanks and fed mostly raw fishthat had been frozen. Initially, seawater wasrecycled, but recycling was replaced later on bycontinuous, gradual flow-through. Almost allthe hatchlings became ill at one time or another.Sick turtles were removed from communal tanks,isolated individually in 10-liter plastic bucketscontaining seawater replaced daily, and providedexperimental chemotherapy (Leong 1979; Leonget al. 1980, 1989; Clary and Leong 1984). Whentheir condition improved they were returned tocommunal tanks. Overall survival was only 9%after 10 mo of rearing.

Kemp’s Ridley eggs and hatchlings forhead-start.—In 1978, the number of live Kemp’sRidley hatchlings received by the GalvestonLaboratory included 1,226 from RN and 1,854from PAIS (an additional hatchling from PAISwas dead on arrival) (Caillouet et al. 1986b,1987; Manzella et al. 1988b; Duronslet et al.1989; Fontaine et al. 1989b, 1990a). Thereafter,fewer hatchlings were received annually (exceptfor 2,025 received in 1990) by the GalvestonLaboratory to be head-started (ibid.; Caillouet1995b; Shaver and Wibbels 2007; BenjaminHiggins, pers. comm.). Within the period1978–1992, the number of Kemp’s Ridleyeggs taken from RN for the reintroductionexperiment (22,507 eggs), plus the estimatednumber (13,518 eggs) that produced hatchlingsat RN for head-start, represented 2.8% (or22,507 + 13,518 = 36,025 eggs) of the totalproduction of 1,286,900 eggs at RN during sameperiod (Caillouet 1995b). The annual take ofeggs that produced hatchlings for head-startduring 1993–2000 was ≈ 1/10 the annual takeof eggs during 1978–1992. Hatchlings fromyear-classes 1978–1988 were received by theGalveston Laboratory from PAIS as part of thereintroduction experiment (Shaver and Caillouet2015), and others were received directly fromRN (year-classes 1978–1980, 1983, 1989–2000)

and CAY (year-classes 1987–1988) (Caillouet1995b; Caillouet et al. 1995b). Hatchlings fromRN were transported by U.S. Coast Guard(USCG) or TPWD aircraft to Galveston. Inretrospect, the takes of eggs and hatchlings fromRancho Nuevo for Kemp’s Ridley head-startduring 1978–1985 did not prevent the populationdecline from reversing in 1986, although it mayhave prevented an earlier reversal.

Rearing turtles in groups.—During July-August 1978, Kemp’s Ridley hatchlings receivedby the Galveston Laboratory were initiallyreared in communal groups in large fiberglassraceways with seawater replaced thrice weekly,and they were fed a commercial, pelletized feed(Clary and Leong 1984; Leong et al. 1989).Every turtle contracted one or more diseases orinfections, some similar to those exhibited earlierby Loggerhead as well as other maladies thathad not been observed in Loggerhead. Kemp’sRidley hatchlings were very aggressive andbit or scratched each other causing injuriesthat led to further complications (Leong 1979;Klima and McVey 1982; Clary and Leong 1984;Fontaine et al. 1985). Sick individuals wereisolated and treated successfully in bucketsby techniques developed for Loggerhead, butmaladies recurred when successfully treatedturtles were returned to communal rearing.Further details concerning diseases in Kemp’sRidleys reared at the Galveston Laboratory, aswell as prophylaxis and treatment, can be foundin sources listed in Table 10.

Rearing turtles separately.—The practice ofcommunal rearing of Kemp’s Ridleys wasstopped after six months and replaced by rear-ing individuals separately, one per container (ini-tially in the same kind of buckets used earlierfor isolation and treatment of sick turtles); multi-ple containers were suspended side by side, par-tially submerged, in seawater within the race-ways. Bottoms of the buckets were perforatedwith holes that allowed seawater exchange and

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turtle excrement and debris from uneaten feedto escape. Various sizes of individual containers,all with perforated bottoms, were later used toaccommodate the increase in size of turtles (Ta-ble 11). Some containers were purchased; plasticflower pots used for hatchlings and plastic buck-ets used after the turtles grew too large for theflower plots. Larger containers were fabricatedfrom purchased materials, to accommodate fur-ther increase in size of the turtles. All containerswere made of resilient, corrosion-resistant (ex-cept for some metal fasteners), non-toxic materi-als. The containers had smooth inside surfaces toprotect the turtles from abrasion and to preventtheir biting off pieces, since they would ingestnon-feed items if available. The location of eachcontainer within the raceway was coded for pur-poses of record keeping. Suspension of contain-ers in groups within raceways facilitated thrice-weekly draining of seawater, removing sea turtlewastes and debris from uneaten feed, cleaningraceways, containers, and turtles, and refillingraceways with clean seawater. When a racewaywas drained, the turtles remained out of water onthe bottoms of their suspended containers untilthe raceway was cleaned and refilled.

Isolation rearing was confining, but providedadequate space for the turtles to swim around,submerge completely, and rest on the bottom oftheir individual containers. However, there wasearly concern that such confinement reduced the

Table 10. Sources covering diseases in Kemp’s Ri-dleys (Lepidochelys kempii) reared at the GalvestonLaboratory, as well as prophylaxis and treatment.

Leong (1979)McLellan and Leong (1981, 1982)Fontaine et al. (1983, 1985, 1990b)Clary and Leong (1984)Caillouet et al. (1986b)Leong et al. (1989)Robertson and Cannon (1997)Higgins (2003)Jacobson (2007)

turtles’ physical fitness. Physical fitness of head-started juveniles was tested in captivity, by ex-perimentally exposing a sample of turtles, oneat a time, to a controlled-current exercise reg-imen within a laminar flow tank (Stabenau etal. 1988, 1992). Each turtle in the sample wassubjected to this regimen 1 day each week overa six month period; the turtles swam againstthe current and their fitness showed signs of im-provement over time (Stabenau et al. 1988, 1992;Valverde et al. 2007). However, this exercise reg-imen could not be adopted as a routine practicefor all head-started turtles, because of limited per-sonnel, equipment, and funds, as well as the largenumber of turtles being reared.

Seawater temperatures were maintained withinan approximate range of 20–30C, mostly near26–30C, and rarely fell below 20C (Caillouet etal. 1986b; Fontaine et al. 1988b, 1989b; Cail-louet 2000; Higgins 2003). Forced-air heatersand heated seawater controlled air and seawa-ter temperatures during winter. Forced-air venti-lation controlled air and seawater temperaturesduring summer. Clean and warm seawater wereessential to successful rearing and disease pre-vention.

Malone and Guarisco (1988) analyzedseawater containing Kemp’s Ridleys, theirwastes and uneaten feed, and results were usedthereafter to guide seawater management duringhead-starting. They assessed seawater qualitybased on biochemical oxygen demand, totalKjeldahl nitrogen, ammonia nitrogen, nitritenitrogen, suspended solids, volatile suspendedsolids, and settleable solids. Wang (2005)

Table 11. Sources covering containers used to head-start Kemp’s Ridleys (Lepidochelys kempii) at theGalveston Laboratory.

Fontaine et al. (1985, 1988b, 1989b, 1990a)Caillouet et al. (1986c, 1988)Manzella et al. (1988b)Caillouet (2000)Higgins (2003)

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studied trace metal accumulation in Kemp’sRidleys during head-starting. Various indicatorsof seawater quality were routinely monitoredduring head-starting (Caillouet et al. 1986b;Fontaine et al. 1988b, 1989b; Duronslet et al.1989; Higgins 2003).

Artificialities of captive-rearing.—The turtleswere reared out of view of each other, but wereable to see their surroundings and caretakersabove their containers. Rearing exposed them toadditional artificialities of feeds, feeding, light,temperatures, sounds, and other physical andchemical characteristics of their captive environ-ment that may have predisposed them to maladap-tive post-release behaviors and survivability ascompared to their wild counterparts. Post-releasebehaviors considered abnormal or aberrant werereported for some head-started Kemp’s Ridleysobserved after release into the Gulf of Mexico(Fontaine et al. 1989a; Meylan and Ehrenfeld2000; Shaver 2007; Shaver and Wibbels 2007).Tag returns and short-term tracking by sonic, ra-dio, and satellite methodologies made it possibleto link post-release behaviors to head-start andreintroduction. However, the potential for suchdetrimental effects (Pritchard 1976; Woody 1990,1991) did not dissuade the early planners or theiragencies from approving and implementing head-start and reintroduction (NPS et al. 1978 op. cit.).

Conditions of captive care may also affect wildKemp’s Ridleys taken from their natural environ-ment for medical treatment and rehabilitationafter being found live-stranded, injured, or ill.Exposure of wild Kemp’s Ridleys to extendedhuman care and artificial conditions in captivityis not an issue when the turtles have permanentdisabilities, infirmities, or infectious diseasesthat prevent their return to the wild. However,when wild Kemp’s Ridleys are temporarilyheld in captivity for medical treatment andrehabilitation, human care may also predisposethem to maladaptive, abnormal, or aberrantbehaviors following their return to the wild.This potential apparently has not discouraged

medical treatment, rehabilitation, and releaseof such turtles (Caillouet 2012b; NOAA, U.S.Department of Commerce. 2010. NOAA’sOil Spill Response: Rehabilitated Kemp’sRidley Sea Turtles Released. Available fromhttp://www.noaa.gov/factsheets/new%20version/

releasing_kemp’s_ridley_turtles.pdf [Accessed7 December 2014]; Wider Caribbean SeaTurtle Conservation Network. 2008. SeaTurtle Care & Medicine. Available fromhttp://www.widecast.org/What/Regional/Medicine.html [Accessed 7 December 2014]). In fact, tagreturns and satellite tracking of rehabilitated wildKemp’s Ridleys can be useful in determiningtheir behavior following release (Lyn et al. 2012).Once released into the wild, head-started Kemp’sRidleys that survived to the calendar yearfollowing the one in which they were releasedwere assumed to have adapted to living in thewild (Caillouet et al. 1995a, 1995b). However,such short term survival was not proof that theirbehavior and survivability in the wild had beenor would be natural or normal for the species.

Duration of captive-rearing.—Caillouet et al.(1995a, 1995b, 1997a) considered Kemp’s Rid-leys reared in captivity for 7–15 mo before re-lease into the wild to be typical head-started tur-tles. Typical head-started Kemp’s Ridleys werereferred to as yearlings (age group 1) because

Table 12. Sources covering sizes and putative agesof wild Kemp’s Ridleys (Lepidochelys kempii) thesurface-pelagic, post-hatchling-early juvenile lifestage.

Zwinenberg (1977)Ogren (1989)Collard and Ogren (1990)TEWG (1998, 2000)Snover and Hohn (2004)Snover et al. (2005, 2007)Putman et al. (2010, 2013)NMFS et al. (2011)Witherington et al. (2012)

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they were released in the calendar year followingthe calendar year in which they were receivedas hatchlings (age group 0). More than 80% ofthem were 9–11 mo old when released (ibid.).Their sizes and ages at release into the wild (Cail-louet et al. 1995a, 1995b, 1997a) were withinthe size and putative age ranges of wild Kemp’sRidleys in the surface-pelagic, post-hatchling-early juvenile life stage (Table 12). Shaver andWibbels (2007) indicated that sizes of head-started Kemp’s Ridleys at release were compara-ble to those of the late pelagic stage or early post-pelagic stage of wild Kemp’s Ridleys, attributingthis to Ogren (1989), who also suggested thatwild Kemp’s Ridleys grow faster in the Gulf ofMexico than along the U.S. Atlantic coast. With-erington et al. (2012) estimated that the wild,surface-pelagic, juvenile Kemp’s Ridleys theyobserved were 1 or 2 y old. Typical head-startedKemp’s Ridley yearlings exhibited the distinc-tive coloration pattern described by Marquez-M.(1994) for 1 y old juveniles; viz. black carapace,almost white plastron and undersides of the neck,beak, upper eyelids, tail, and proximal parts ofthe tail, and a narrow, dorsal, white border alongthe periphery of the carapace.

The habitat of wild, surface-pelagic Kemp’sRidleys is near the ocean’s surface, where theturtles drift passively, dive, feed, and apparentlyrest while being dispersed by surface circulation(Table 13). The wild, surface-pelagic Kemp’s

Table 13. Sources covering the habitat of wild,surface-pelagic Kemp’s Ridleys (Lepidochelys kem-pii).

Zwinenberg (1977)Ogren (1989)Collard and Ogren (1990)TEWG (1998, 2000)Snover and Hohn (2004)Snover et al. (2005, 2007)Putman et al. (2010, 2013)NMFS et al. (2011)Witherington et al. (2012)

Ridleys observed by Witherington et al. (2012)were strongly associated with the surface-pelagicSargassum macroalgae community. Manzellaet al. (2001) observed two head-started Kemp’sRidleys of the 1987 year-class floating inSargassum after release in 1988, one 4.8 moafter release and the other 14.8 mo after release.During an offshore study conducted in southTexas in October-November 1988 and February-June 1989, no sea turtles were observed bydivers beneath Sargassum mats, and none werecaptured along with Sargassum in surface trawlhauls, nor did stomach contents of pelagic fishescaught near the mats contain hatchling turtles orparts thereof (Fontaine et al. 1990a).

Feeds and feeding.—Experimentation withvarious feeds and feeding methods led to routineuse of commercially produced, pelletized,dry, floating feed fed twice daily, in earlymorning and late afternoon (Fontaine et al.1985; Caillouet et al. 1986b, 1989b). Initialfeeding of hatchlings was delayed to allow timefor absorption of yolk (Fontaine et al. 1985;Fontaine and Williams 1997). Thereafter, thedaily total weight of feed received per turtlewas adjusted monthly, based on its percentageof average (arithmetic or geometric) body massof a monthly sample of turtles (Fontaine et al.1985, 1988b; Caillouet et al. 1986b, 1988, 1989;Caillouet 2000; Higgins 2003). The twice-dailyportions of feed were measured and distributedvolumetrically, based on the volume-massrelationship of the feed. Under this feedingregimen, daily weight of feed per turtle increasedexponentially during head-starting, while weightof feed as a percentage of body mass declinedlogarithmically (Higgins 2003). Problems withinsect invasion of pelletized feed held in drystorage led to its routine frozen-storage (Fontaineet al. 1985).

Growth and survival in captivity.—Growthin body weight and survival of Kemp’s Ri-dleys during head-starting were examined by

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sources listed in Table 14. Caillouet and Koi(1985) showed that the range in body weightwithin samples of Kemp’s Ridleys being head-started increased with age, based on year-classes 1981–1983 (i.e., variance in individualweight was heterogeneous). Therefore, they rec-ommended application of the geometric mean asa measure of central tendency of weights of tur-tles of a given age during head-starting (see alsoCaillouet et al.1986b, 1989).

Caillouet et al. (1986c) determined survivaland growth in weight year of turtles in year-classes 1978–1983, for PAIS and RN imprintgroups, and by clutches, during head-starting.They characterized growth in weight with a linearregression of the natural logarithm of weight (ing) on the square root of age (in days). Manzellaet al. (1988b) used the model to compare growthof turtles of the 1986 year-class reared in twodifferent sizes of containers, and detected no sig-nificant difference in growth within the two sizesof containers. Caillouet et al. (1997a) modifiedthis model for application in estimating growthin weight during head-starting of year-classes1978–1992, by expressing weight (w) in kg andage (t) in years, and estimating slope d (thegrowth index) and intercept ln(c) using linearregression:

ln(w) = ln(c) + d(t1/2) (1)

Table 14. Sources covering growth in body weightand survival of Kemp’s Ridleys (Lepidochelys kem-pii).

Klima and McVey (1982)Caillouet et al. (1986b, 1987, 1989, 1997a)Fontaine et al. (1985, 1990a)Manzella et al. (1988b)Duronslet et al. (1989)Landry (1989)Fontaine and Williams (1997)Fontaine and Shaver (2005)Shaver and Wibbels (2007)

Caillouet et al. (1997a) also estimated the rela-tionship between variance, s2, and arithmeticmean, w, of the combined year-classes; a scat-ter plot of s2 on w showed greater variability inw at t in year-classes 1978–1985 than in year-classes 1986–1992. Caillouet et al. (1997a) sug-gested that variability of w at t in year-classes1986–1992 may have been lowered by control-ling incubation temperature to produce female-dominated sex ratios (Shaver et al. 1988; Cail-louet 1995a) and by improving rearing facilitiesand methods. The slope of the relationship be-tween ln(s2) and ln(w) was very close to 2, indi-cating that the distribution of w at t was ln-normalwithin the range of t for head-started Kemp’s Rid-leys.

The relationship between w and straight cara-pace length (SCL) was determined from pairedw and SCL observations from year-classes1978–1992 combined (Caillouet et al. 1997a):

w = a(SCL)b (2)

Parameters ln(a) and b were estimated by linearregression of ln(w) on ln(SCL), as follows:

ln(w) = ln(a) + b(ln[SCL]) (3)

where ln(a) = 8.438, b = 2.920, n = 53,317,and adjusted r2 = 0.994. This large data setcould have contained paired observations forsome turtles weighed and measured more thanonce (i.e., at different ages) during head-starting.Equation (3) also was rearranged, withoutchanging its parameter estimates, and used toestimate SCL from w (Caillouet et al. 1997a).

Rearing more than 15 mo.—Kemp’s Ridleystransferred from the Galveston Laboratory toCAY and other marine aquaria to develop acaptive breeding stock were reared in captiv-ity for more than 15 mo, and were referred toas extended head-started or super head-started(Fontaine and Caillouet 1985; Fontaine et al.1985, 1988b; Caillouet et al. 1995a, 1995b). This

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distinction from typical head-started Kemp’s Rid-leys is important, because the extended captive-rearing to which super head-started turtles wereexposed may have conditioned them to greaterdependency on human care than typical head-started turtles. Turtles of the 2000 year-class thatwere released in 2003 are considered super head-started. We assume that the longer head-startedKemp’s Ridleys are exposed to human care andother artificial conditions of captivity, the greaterthe risk of their developing maladaptive, abnor-mal, or aberrant behaviors that could be detrimen-tal to their performance and survival in the wild.The same may be true for sick or injured wildKemp’s Ridleys brought into captivity for treat-ment and rehabilitation, then released. Therefore,we believe that evaluation of head-start shouldnot include super head-started individuals be-cause of their extended exposure to potentiallyirreversible effects of captive-rearing. However,we would not exclude super head-started indi-viduals from an evaluation of reintroduction; anesting by a super head-started Kemp’s Ridleyhas been documented at PAIS (Shaver and Cail-louet 2015).

Over a period of almost 10 y, the growth-in-weight trend for 10 super head-started Kemp’sRidleys of the 1978 year-class, at Sea-AramaMarineworld in Galveston, was asymmetricallysigmoid (Caillouet et al. 1986b; Duronslet etal. 1989; Fontaine et al. 1989b). This indicatedthat the inflection point on the growth curve hadbeen passed and that growth rate was decliningwith age, suggesting that the turtles were eitherapproaching maturity or had reached maturity.McVey and Wibbels (1984) reported growth incarapace length and weight in super head-startedKemp’s Ridleys at Sea-Arama Marineworld andMiami Seaquarium, Miami, Florida over a periodless than 20 mo.

Tags and tagging.—Witzell (1998) empha-sized that tagging sea turtles should be basedon a sound research plan and legitimate researchgoals, and these requirements were met by Camp-

Table 15. Some examples of sources emphasizingthat tags were essential to distinguish head-startedKemp’s Ridleys (Lepidochelys kempii) released intothe Gulf of Mexico from wild conspecifics.

Campbell (1977 op. cit.)NPS et al. (1978 op. cit.)Klima and McVey (1982)Caillouet (1984, 1987)McVey and Wibbels (1984)Fontaine and Caillouet (1985)Fontaine et al. (1985, 1990a, 1993)Caillouet et al. (1986b, 1987, 1995c, 1997b)Manzella et al. (1988a)Higgins et al. (1997)

bell (1977 op. cit.) and NPS et al. (1978 op. cit.).Without tags, head-started Kemp’s Ridleys re-leased into the Gulf of Mexico could not havebeen distinguished from wild conspecifics (Table15). Tags on head-started Kemp’s Ridleys linkedtheir recapture data to previously recorded in-formation on year-class, imprint group, releaselocation, and hatchling emergence dates (fromwhich their age was calculated; see Caillouet etal. 2011).

In most cases, each type of tag was tested onsmall numbers of head-started Kemp’s Ridleyswhile they were in captivity, to ensure the tagwas safe for application to larger numbers tobe released. Tags were applied to head-startedKemp’s Ridleys well before their release to allowhealing of wounds caused by tagging. Prior torelease, each turtle was weighed and SCL wasmeasured. Each was also examined to make sureits tags were properly applied and retained beforerelease. An approach of applying more than onetag to head-started turtles was tested on samplesof turtles of year-classes preceding the 1984 year-class (Fontaine et al. 1993; Benjamin Higgins,pers. comm.). Multiple-tagging was expected toincrease chances that at least one tag would beretained and recognized when the turtles wererecaptured.

External metal (MonelT M or InconelT M) tagswere applied to the trailing edge of a foreflipper

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(usually the right) of all turtles to be releasedfrom each year-class, about one month beforetheir release (Fontaine et al. 1990a, 1993; Cail-louet et al. 1995a; Fontaine and Shaver 2005;Shaver and Wibbels 2007). These tags were ini-tially issued by the NMFS Miami Laboratory,Miami, Florida, then later by the CooperativeMarine Turtle Tagging Program (CMTTP) thatcoordinates the distribution of sea turtle tags,manages tagging data, and facilitates exchangeof tag information for projects conducted in theAtlantic Region (Archie Carr Center for Sea Tur-tle Research, University of Florida. 2014. Coop-erative Marine Turtle Tagging Program. Avail-able from http://accstr.ufl.edu/resources/tagging-program-cmttp [Accessed 7 December 2014]; Pe-ter Eliazar, pers. comm.). Each tag bore a uniquealphanumeric code and inscription requesting re-porting the turtle’s recapture to the NMFS MiamiLaboratory.

Living tags (Hendrickson and Hendrickson1981, 1983; Bowman 1983) were first applied ex-perimentally to samples of turtles of year-classes1978 (only one turtle), 1980 (180 turtles) and1982 (436 turtles), then to most or all survivingindividuals of year-classes1983–2000 (Fontaineet al. 1993; Caillouet et al. 1997b; Fontaine andShaver 2005; Shaver and Wibbels 2007; Ben-jamin Higgins, pers. comm.). The living tag isa permanent, light-colored mark (autograft) onthe darker carapace. As applied to a head-startedKemp’s Ridley, it was formed by excising a smallpiece of light-colored plastron tissue and implant-ing it into a darker-colored carapace scute cho-sen to identify the turtle’s year-class (Bowman1983; Fontaine et al. 1985; Caillouet et al. 1986a,1997b; Fontaine et al. 1988a, 1993). Such scutetissue implants grow in size as the turtle grows.Living tags were applied 6–8 week prior to re-leasing turtles (Benjamin Higgins, pers. comm.).

We used internal coded-wire tags(Northwest Marine Technology Inc. 2014.Coded wire tags (CWT). Available fromhttp://www.nmt.us/products/cwt/cwt.shtml[Accessed 7 December 2014]) experimentally

to small numbers of turtles from year-classes1978 (9 turtles) and 1982 (12 turtles; Fontaine etal. 1993; Fontaine and Shaver 2005; BenjaminHiggins, pers. comm.). They were applied tomost turtles of year-classes 1984–1997, but toless than half of the turtles in year-classes 1998and 1999 (ibid.). CWTs were applied to onlyfour year-classes (1985–1988) of the elevenimprinted to PAIS. None were applied to turtlesof year-class 2000. CWTs were magnetizedeither before or after being implanted, whichwas determined by the type of hypodermicinjector used (Higgins et al., 1997; BenjaminHiggins, pers., comm.). They were implantedinto the flesh in the phalanges area of a rightor left foreflipper, and can be detected in X-rayimages by trained personnel (Fontaine et al.1993). Trained personnel can also detect them byscanning both foreflippers with a magnetometer,if they were magnetized before implantation. Ifthey were not magnetized before implantation,they can be magnetized upon recapture bypassing a magnet over both foreflippers, whichcan then be scanned with a magnetometer.Some turtles were also successfully tagged,experimentally, with CWTs in both the rightand left hind flippers in 1991, but tagging in theforeflippers was the preferred routine (BenjaminHiggins, pers. comm.). In some cases, flippersof dead-stranded Kemp’s Ridleys have beenarchived in frozen storage for later examinationfor CWTs.

Internal passive integrated transponder (PIT)tags, also called radio frequency identification(RFID) tags, were applied to samples of turtlesin year-classes 1978, 1982, and 1984–1989, thento most turtles in the 1990–2000 year-classes(Fontaine et al. 1993; Fontaine and Shaver 2005;Benjamin Higgins, pers. comm.). PIT tags wereinserted with a hypodermic injector into the fleshof the pectoral muscle in the axial area of a fore-flipper of each turtle, around two months beforeturtles were released (Fontaine et al. 1993; Ben-jamin Higgins, pers. comm.). In turtles ≥ 30 cmSCL, PIT tags were inserted with a hypodermic

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http://accstr.ufl.edu/resources/tagging-program-cmttp

http://www.nmt.us/products/cwt/cwt.shtml


injector into the flesh of a foreflipper, in a loca-tion posterior to the fibula (Benjamin Higgins,pers. comm.). In some cases, flippers of dead-stranded Kemp’s Ridleys have been archived infrozen storage for later examination for PIT tags.PIT tag migration in Kemp’s Ridley has beeninvestigated by Wyneken et al. (2010).

Flipper tags were easily recognized, but notpermanent. Living tags were permanent, but notalways recognized as tags by untrained observers.CWTs were permanent but required a magne-tometer or X-ray image for detection by trainedpersonnel; their binary codes could be deter-mined only after the tags were excised, whichwas prohibited on live animals but not on deadspecimens (e.g., found stranded). PIT tags werepermanent but required well trained and experi-enced personnel using compatible detectors todetect them and read their alphanumeric codes(Manzella 1988); X-ray images also can be usedto detect them. PIT tag detection was sometimesprevented by incompatibilities between PIT tagsand PIT tag readers from different manufactur-ers, or by use of PIT tag-compatible readers ofdifferent detection strengths from the same manu-facturer (Donna Shaver, pers. comm.). Scars leftby foreflipper tags that were lost for various rea-sons can indicate previous tagging, but they aloneare not sufficient evidence to distinguish head-started Kemp’s Ridleys from wild conspecificswhen flipper tags are lost and tagging scars re-main (Caillouet et al. 1997b). In our view, proofthat a recaptured, stranded, nesting, or otherwiseobserved Kemp’s Ridley was head-started cancome only from detection of external or internaltags which clearly identify it as head-started. Un-fortunately, some observers used unexpected lo-cations, ease of capture, or other putatively aber-rant behavior of Kemp’s Ridleys in the wild, toconclude they were head-started, in the absenceof proof from retained external or internal tagsto confirm it (e.g., Woody 1991; Bowen et al.1994).

In addition to the four types of tags alreadymentioned, radio- or sonic-tags were attached to

Table 16. Sources of information on transportinghead-started Kemp’s Ridleys (Lepidochelys kempii).

NPS et al. (1978 op. cit.)Fontaine et al. (1985, 1989a, 1989b)Caillouet et al. (1986b, 1987)Manzella et al. (1988a)Duronslet et al. (1989)Higgins (2003)

Table 17. Sources of information on head-startedKemp’s Ridley (Lepidochelys kempii) release sites.

Klima and McVey (1982)McVey and Wibbels (1984)Wibbels (1984)Fontaine and Caillouet (1985)Manzella et al. (1988a, 1990)Fontaine et al. (1990a)Caillouet et al. (1995a)Fontaine and Shaver (2005)Shaver (2005)Shaver and Wibbels (2007)

limited numbers of head-started Kemp’s Ridleysfor purposes of tracking them following release(Timko and DeBlanc 1981; Klima and McVey1982; Wibbels 1984; Manzella et al. 1990).Renaud et al. (1993) conducted an experimentin which they attached imitation satellitetransmitters to carapaces of 12 Kemp’s Ridleysthat were 32-mo old (i.e., super head-started), todetermine their retention over a 16 mo periodin captivity. Shaver and Rubio (2008) trackedhead-started nesters by satellite after attachingplatform transmitter terminals to their carapaces.

Transport and release.—Table 16 lists sourcesof information on transporting head-startedKemp’s Ridleys, via ground vehicles, aircraft,and vessels. Release dates for year-classes1978–1992 are given in Caillouet et al. (1995a).Most turtles of the 1978 and 1979 year-classeswere released off the west coast of Florida. Only113 turtles of the 1978 year-class and six of1979 year-class were released along the Texascoast. Because of the threat of incidental capture

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in shrimp trawls, most releases in Texas werecarried out during periods and in areas closedtemporarily to shrimping. For example, releaseof some of the 1978 year-class turtles off PadreIsland in 1979 took place during the periodwhen Texas offshore territorial waters, from the7.3 m depth contour out to nine nautical miles,were closed to shrimping (Griffin et al. 1993).After the 1979 year-class was released, releasesalong the Texas coast usually took place duringthe Texas Closure, an annually recurring andcoordinated NMFS-TPWD closure to shrimptrawling in offshore federal and state watersduring a period of varying duration in May-July,which allows Brown Shrimp (Farfantepenaeusaztecus) to grow to larger, more valuable sizesbefore harvest (Griffin et al. 1993; Matlock2010). Depending on the release sites chosen(Table 17) and the availability of vessels eachyear, head-started Kemp’s Ridleys were releasedfrom vessels operated by NOAA, USCG, Uni-versity of Texas Marine Science Institute, TexasA & M University, TPWD, Florida Departmentof Environmental Protection (FDEP), or ARCOOil and Gas Company. Yearlings were usuallytransported by vehicles to the docking locationsof release vessels (Table 16). However, onoccasion some were transported by vehicle toGalveston’s Scholes Airfield, where they wereloaded onto USCG aircraft that flew them toairports near vessel docking locations; they werethen taken by vehicle to the docking locations.All turtles were released one to a few at atime from the stern of the vessel while it wasunderway at relatively slow speed.

Table 18 lists sources of numbers of head-started Kemp’s Ridleys released annually intothe Gulf of Mexico; the numbers releasedreflected variation in numbers of hatchlingsreceived by year-class and imprint group (PAIS,RN, and CAY), as well as survival during head-starting. The overall total number released fromyear-classes 1978–2000 was 23,967, including14,776 from year-classes 1978–1988 (i.e., those

associated with the reintroduction experiment),7,800 from year-classes 1989–1992, and 1,391from year-classes 1993–2000 (Benjamin Higginspers. comm.). Super head-started Kemp’sRidleys were included in these numbers (ibid.).We were unable to reconcile the 23,967 total withthe 23,987 total reported by Shaver and Wibbels(2007), but the difference is only 20 turtles. Mostturtles released from year-classes 1978–1988were in the PAIS imprint group, with fewer inthe RN and CAY imprint groups (Caillouet etal. 1995a; Fontaine and Shaver 2005; Shaverand Wibbels 2007). All turtles released from the1989–2000 year-classes were in the RN imprintgroup (Shaver and Wibbels 2007; BenjaminHiggins pers. comm.). The major head-starteffort involved the 1978–1992 year-classes, afterwhich numbers were reduced by about 1/10 foryear-classes 1993–2000 (Shaver and Wibbels2007). Some head-started Kemp’s Ridleys fromthe 1980 year-class (197 turtles) were released inCampeche Bay (Caillouet et al. 1995a; Shaverand Wibbels 2007). Most from the 1981–2000year-classes were released along the coast ofTexas. Although releases off west Florida wererecommended by NPS et al. (1978 op. cit.), theywere discontinued and replaced by releases in thewestern Gulf of Mexico to reduce the probabilitythat the turtles would escape into the Atlantic(Fontaine et al. 1989a). An exception was therelease of 29 head-started Kemp’s Ridleys of

Table 18. Sources of information on numbers of head-started Kemp’s Ridley (Lepidochelys kempii) releasedannually.

Caillouet et al. (1986b, 1995a, 1995b, 1997a)Manzella et al. (1988a)Duronslet et al. (1989)Fontaine et al. (1989a)Caillouet (1995b, 2000)Shaver (2005, 2007)Fontaine and Shaver (2005)Shaver and Wibbels (2007)Shaver and Caillouet (2015)

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the 1992 year-class off Panama City, Florida in1993, after their use in TED testing (Shaver andWibbels 2007; Benjamin Higgins, pers. comm.).

Time lapse to nesting.—During years1966–1977 that preceded reintroduction andhead-start, 280,000 hatchlings were releasedthrough conservation efforts at RN (TEWG2000). Young (i.e., putative neophyte) Kemp’sRidley nesters began appearing at RN in 1976,10 y after hatchling recruitment was restoredby INIBP (Marquez-M. 1994; Pritchard 1997).During 1978–1988, when eggs for reintroductionwere being transferred from RN to PAIS, 548,420hatchlings were released at RN (TEWG 2000).Annual numbers of nests at RN declined from5,991 in 1966 to a low of 740 recorded in 1985before the decline reversed in 1986 (TEWG2000). In comparison, the first documentednesting of a PAIS-imprinted and head-startedKemp’s Ridley occurred in 1996, 17 y afterthe first head-started turtles were released in1979 (Shaver 1996; Shaver and Caillouet 2015).During the 23 y of head-starting, 2,106,641hatchlings were released through conservationefforts in Tamaulipas (TEWG 2000; data foryears 1999 and 2000 were provided by JaimePeña pers. comm.); 1,558,221 of these hatchlingswere released during 1989–2000, when onlyRN imprinted hatchlings were transferred tothe Galveston Laboratory. Shaver and Caillouet(2015) reported the number of hatchlingsreleased at PAIS from nestings by head-startedKemp’s Ridleys, most of which came from eggslaid at PAIS.

Tag returns and tracking.—Tag returns wereamong the most important and essential data re-quired for evaluation of head-start and reintro-duction (Campbell 1977 op. cit.; NPS et al. 1978op. cit.; Pritchard 1979; Allen 1981). Tag returnsand tracking results were necessary to evaluateperformance of head-started turtles following re-lease and to determine overall effectiveness ofreintroduction and head-start. Not only was it es-

Table 19. Sources of information on head-startedKemp’s Ridley (Lepidochelys kempii) tag returns andtracking results.

Timko and DeBlanc (1981)Klima and McVey (1982)Wibbels (1983a, 1983b, 1984)McVey and Wibbels (1984)Manzella et al. (1988a, 1990, 1991)Fontaine et al. (1989a)Shaver (1991, 1998, 2007)Manzella and Williams (1992a, 1992b)Caillouet (1994)Cannon et al. (1994)Werner (1994)Werner and Landry (1994)Caillouet et al. (1995a, 1995b, 1997b, 2011)Fontaine (1995)Fontaine and Schexnayder (1995)Schmid (1998)Shaver and Caillouet (1998, 2015)Kenyon et al. (2001)Metz (2004)Fontaine and Shaver (2005)Landry et al. (2005)Shaver and Wibbels (2007)Seney and Landry (2008)Shaver and Rubio (2008)Snover et al. (2008)Shaver and Caillouet (2015)

sential to the head-start and reintroduction experi-ments that significant numbers of head-started fe-males survived to maturity and reproduced in thewild, but also that significant numbers were doc-umented by tag returns as having nested (Shaverand Wibbels 2007; Caillouet et al. 2011; Shaverand Caillouet 2015). Tag returns provided bio-logical information on age, survival, movements,dispersal, distribution, habitat utilization, growth,feeding ecology, trace metals in blood, adapta-tion, behavior, maturation, and reproduction ofhead-started Kemp’s Ridleys recaptured or oth-erwise observed after release into the Gulf ofMexico.

Table 19 lists sources of information on head-started Kemp’s Ridley tag returns and tracking

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results. Tag returns were received through theSTSSN, from deliberate searches for nesters atPAIS and other areas of Texas, and serendip-itously (Caillouet et al. 1995a; Fontaine andShaver 2005; Shaver and Wibbels 2007; Shaverand Caillouet 2015). The STSSN was the ma-jor source of tag returns for head-started Kemp’sRidleys; they were recorded incidentally to docu-mentation of sea turtles strandings (Fontaine andShaver 2005). Although conservation efforts inTamaulipas required tagging and examination ofnesters for tags (NPS et al. 1978 op. cit.; NMFSet al. 2011; Shaver and Caillouet 2015), only sixhead-started Kemp’s Ridleys were reported tohave nested in Mexico; one was PAIS-imprintedand the other five were RN-imprinted (Shaverand Wibbels 2007; Donna Shaver, pers. comm.).

Identification of a Kemp’s Ridley as head-started can be challenging. A head-startedKemp’s Ridley could be erroneously identifiedas wild if its foreflipper tag or tags were missingand if those who recaptured or found the turtlewere unfamiliar with living tags and internal tags,did not have appropriate, functioning equipmentto detect and decode internal tags, or were inade-quately trained in proper use of such equipment.Nevertheless, overall tag return rate for head-started Kemp’s Ridleys was about 4% (Shaverand Wibbels 2007). Interestingly, tag return ratewas 4% for 1,859 wild nesters tagged at RN dur-ing 1966–1978, based on 82 tag returns reportedduring 1966–1979 (data from Table 2, p. 161 inMárquez, M. et al. 1982). Similarity between theoverall tag return rates of head-started Kemp’sRidleys (yearlings through adults, including bothsexes) and wild nesters at RN is surprising, be-cause the estimated annual survival rates of wildsubadults (85.0–93.5%, ages 6–11 y) and adults(85.0–93.5%, ages ≥ 12 y) are higher than thoseestimated for small wild juveniles (60.7–81.5%,ages 2–5 y) (NMFS et al. 2011). Estimated an-nual survival rates of head-started Kemp’s Rid-leys released into the wild (Caillouet et al. 1995a)were generally lower than those estimated fortheir wild counterparts over similar ranges in size

and age (NMFS et al. 2011), but many factors in-fluence tag returns besides survival. Therefore, acomparison of tag returns for head-started turtleswith those of wild nesters at RN is not strictlyvalid.

Fontaine and Shaver (2005) reported 958 tag re-turns from head-started year-classes 1978–1992.For year-classes 1993–2000, the only tag returnsreported by Shaver and Wibbels (2007) werefrom one individual from the 1993 year-classthat nested twice in 2003. Since then, additionaltag returns for head-started Kemp’s Ridleys havebeen documented (Shaver and Caillouet 2015),and more may be documented in the future, butnumbers of head-started survivors are diminish-ing. The most recent nesting by a head-startedKemp’s Ridley was documented in 2013 (DonnaShaver, pers. comm.). Recovery methods weredetermined from tag returns by Caillouet et al.(1995a) and Fontaine and Shaver (2005), andshowed that head-started Kemp’s Ridleys werevulnerable to anthropogenic threats similar tothose to which wild Kemp’s Ridleys were ex-posed (see also Fontaine et al. 1989a; Manzellaand Williams 1992a, 1992b; Shaver and Wibbels2007; NMFS et al. 2011). We combined tag re-turns (data from Fontaine and Shaver 2005) forsome of the recovery methods into fewer cate-gories, and recalculated percentages after elimi-nating entries with unknown and not reportedmethods that together represented 115 tag re-turns. Based on the remaining 843 tag returnsfor which a method was reported, 420 strandings(live, dead, and floating dead) represented 49.8%,161 incidental captures in shrimp trawls repre-sented 19.1%, and 209 recaptures by all otherfishing gears or methods combined (commercial,recreational, and sampling) represented 24.8%.For nesters tagged at RN, Marquez-M. (1994)reported that 71.2% of the tag returns were fromshrimp trawling in the Gulf of Mexico.

The threat of incidental capture of Kemp’s Ri-dleys in shrimp trawls was high (Yaninek 1995;Epperly 2003; Lewison et al. 2003) when reintro-duction year-classes 1978–1988 were released in

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1979–1989, respectively. Federal regulations re-quiring seasonal use of TEDs in shrimp trawls inoffshore waters were first implemented in 1987,but they were not very effective in that year dueto legal challenges and federal government sus-pensions of their enforcement (ibid.). Temporal-spatial coverage of federal TED regulations ex-panded thereafter, until TEDs were required inshrimp trawls at all times and in all areas of thesoutheast U.S. in 1994 (ibid.; Sheryan Epperly,pers. comm.). Even after TEDs were required inshrimp trawls at all times and in all areas, seaturtle strandings continued to be correlated withshrimp trawling in the Gulf of Mexico (Caillouetet al. 1996; Lewison et al. 2003; Finkbeiner etal. 2011; Caillouet 2012b). Only the 1993–2000year-class turtles were released in an environ-ment in which the threat of incidental capturein shrimp trawls was substantially reduced com-pared to years preceding 1993.

Releasing most head-started turtles along theTexas coast, and mostly during periods andin areas temporarily closed to shrimping, gavethem short-term protection from interactions withshrimp trawling, but not during periods open toshrimping before or after such closures (Shaver1998, 2005; Lewison et al. 2003). Interestingly,shrimp trawling effort in the northern Gulf ofMexico began to decline in the late 1980s or early1990s (Caillouet et al. 2008; Nance et al. 2010),which could have contributed along with TEDsto reduction in the threat of incidental capture inshrimp trawls to head-started and wild Kemp’sRidleys (Caillouet 2010, 2014).

After release, head-started Kemp’s Ridleysdispersed and some apparently adapted, survived,and grew (Caillouet et al. 1995a, 1995b, 2011).Some females also matured and produced viableoffspring in the wild (Fontaine and Shaver 2005;Shaver and Wibbels 2007; Shaver and Caillouet2015). This occurred, despite their exposureto artificial conditions and manipulations incaptivity (from the egg stage to their release asyearlings) and to multiple anthropogenic andnatural threats in the wild. A viable nesting

colony of Kemp’s Ridley now exists at PAIS (seeFigure 2 in Shaver and Caillouet 2015), consist-ing of head-started and wild nesters (Fontaineand Shaver 2005; Shaver and Wibbels 2007;Shaver and Caillouet 2015), some of which maybe offspring of head-started nesters (Caillouet2005). Hatchlings produced from clutcheslaid by head-started Kemp’s Ridley nestershave been indistinguishable (so far) from thoseproduced from clutches laid by wild nesters,because hatchlings produced from clutcheslaid by head-started Kemp’s Ridleys have notbeen marked or tagged at PAIS. However, it ispossible that genetic studies conducted at PAIS(Frey et al. 2014) may eventually link some ofthe wild nesters to head-started mothers (Shaverand Caillouet 2015).

Growth and maturity after release.—Somaticgrowth and maturity in head-started Kemp’sRidleys in the wild have been estimated byCaillouet et al. (1995b, 2011) and Snover et al.(2008). The most definitive analysis of somaticgrowth in subadult and adult life stages ofhead-started Kemp’s Ridleys was Caillouet etal. (2011). They fitted a von Bertalanffy growthcurve to post-release SCL (cm) versus knownage (t, in y) of 82 head-started Kemp’s Ridleyrecaptures. The data set was limited to turtles ≥40 cm SCL at recapture, which included turtlesin transition from subadults to adults, as well as49 that were known nesters (i.e., known adultfemales); all were in the declining phase ofgrowth. Juveniles < 40 cm in SCL, consideredto be in the exponential phase of growth, wereexcluded. Use of this size-constrained data setavoided complications related to fitting the vonBertalanffy growth curve to data from juvenilesgrowing exponentially (Snover et al. 2008;Caillouet et al. 2011). The resulting growthcurve (Caillouet et al. 2011) was:

SCL = SCLasym[1 − e−k (t−t0)] (4)

where the estimated asymptote SCLasym =

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64.08 cm, estimated k = 0.1817, estimated t0 =

3.7011, and e is the base of natural logarithms;adjusted r2 = 0.898, and n = 82. Applyingequation (4) and assuming an SCL at maturityof 58 cm (Márquez M. 1994), we estimatedage at maturity at t = 9.2 y. Assuming SCLat maturity is 60 cm (Snover et al. 2007), weestimated the time lapse to maturity at t = 11.5y; recent demographic modeling of the wildKemp’s Ridley population assumed it to be 12y (NMFS et al. 2011; Gallaway et al. 2013).Age at maturity estimated from somatic growthcurves increases by increasing increments as theassumed size at maturity increases. Chaloupkaand Musick (1997), Chaloupka and Zug (1997),and Snover et al. (2007; Bjorndal et al. 2014)reviewed age, growth, and maturity in wild andhead-started Kemp’s Ridleys.

Captive breeding.—To establish a captivebrood stock, 264 head-started Kemp’s Ridleyswere distributed among CAY and numerous ma-rine aquaria in the U.S. (Caillouet and Revera1985; Caillouet et al. 1986b, 1988; Duronslet etal. 1989). The Kemp’s Ridleys at CAY providedadditional research opportunities (Rostal 2005,2007; Holder and Holder 2007). Twelve Olive Ri-dley hatchlings were obtained from Ross Witham,FDNR, Jensen Beach, Florida in September 1984and were head-started; the six survivors weretransferred to Miami Seaquarium in May 1985(Caillouet et al. 1986b).

CAY was successful in breeding Kemp’s Rid-leys in captivity (Wood and Wood 1984, 1988,1989). Two 5 y olds produced eggs in 1984. Onereleased seven eggs in a seawater tank but nonedeveloped, and the other deposited 62 eggs in anest but only three hatchlings emerged and alldied. Four 7 y olds nested in 1986, laying a to-tal of 535 eggs from which 78 viable hatchlingswere produced. The 184 hatchlings producedfrom nestings at CAY in 1987 and 1988 weretransferred to the Galveston Laboratory and head-started (Caillouet et al. 1995b); 130 yearlingswere released from year-class 1987, and 14 from

year-class 1988 (Caillouet et al. 1995a). Produc-tion of Kemp’s Ridley hatchlings through cap-tive breeding at CAY was terminated after 1988(Duronslet et al. 1989; Márquez-M. et al. 2005).Surviving super head-started adults that had beenhead-started at the Galveston Laboratory and sentto CAY were returned by FWS to the Galve-ston Laboratory and released into Galveston Bayin 1992. The remaining brood stock, includingsurvivors originally sent as hatchlings to CAYfrom RN, and others produced by captive breed-ing, were transferred to XCaret Eco Park, Quin-tana Roo, Mexico (Caillouet 2000; Márquez-M.et al. 2004, 2005; Ana Cecilia Negrete, pers.comm.). Two viable hatchlings were produced in1986 from a nesting named Little Fox at MiamiSeaquarium (David Owens, pers. comm.). LittleFox was the only survivor among five Kemp’sRidley hatchlings from RN that were given toIla Loetscher by Henry Hildebrand in 1971 (Size-more 2002); this female had been reared in captiv-ity by Ila Loetscher, Sea Turtle, Inc., and flown asan adult to Miami Seaquarium where she nested.Bentley (1989) described unsuccessful attemptsin 1984 and 1985 at captive breeding Kemp’sRidleys at Miami Seaquarium; one of the threeadult females involved was Little Fox.

Evaluations

The December 1981 issue (No. 19) of MarineTurtle Newsletter contained articles suggestinghow head-start might be evaluated, including sev-eral that applied to Kemp’s Ridley (Hendricksonand Hendrickson 1981; Pritchard 1981; Woody1981). Table 20 lists additional evaluations ofKemp’s Ridley head-start, and we may haveoverlooked others. It is noteworthy that most ofthese evaluations took place before the first docu-mented nesting of a head-started Kemp’s Ridleyoccurred in 1996 (Shaver 1996).

Two major evaluations of Kemp’s Ridley head-start were initiated by NMFS, both of them exter-nal to this agency. The first was conducted byWibbels et al. (1989. Blue Ribbon Panel Review

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Table 20. Sources of additional evaluations ofKemp’s Ridley (Lepidochelys kempii) head-start.

Pritchard (1979, 1997, 2006)Ehrenfeld (1982)Dodd (1985)Mortimer (1988, 1995)Allen (1990, 1992)Magnuson et al. (1990)Wibbels (1990, 1992)Woody (1990, 1991)Dodd and Seigel (1991)Frazer (1992, 1997)FWS and NMFS (1992)Shaver and Fletcher (1992)Taubes (1992a, 1992b)Donnelly (1994)Eckert et al. (1994)Heppell and Crowder (1994, 1998)Caillouet et al. (1995c)Heppell et al. (1996, 2007)Heppell (1997)Ross (1999)Seigel and Dodd (2000)Godfrey and Pedrono (2002)Spotila (2004)Caillouet (2005, 2006)Fontaine and Shaver (2005)Mrosovsky (2007)Shaver and Wibbels (2007)Kemp’s Ridley Recovery Team (2009)NMFS et al. (2011)Shaver and Caillouet (2015)

of the National Marine Fisheries Service Kemp’sRidley Headstart Program. Available fromhttp://www.galvestonlab.sefsc.noaa.gov/publications/pdf/930.pdf [Accessed 8 December 2014]);see also Wibbels 1990). The second was con-ducted in September 1992 (Eckert et al. 1994).

Blue ribbon panel review.—Wibbels et al.(1989 op. cit.) and Wibbels (1990) evaluatedhead-start based on its potential to contribute sig-nificantly to Kemp’s Ridley recovery, rather thanon its originally planned role as part of reintro-duction (NPS et al.1978 op. cit.). They recog-

nized that considering head-start as a potentialconservation method was controversial amongscientists. However, such an evaluation was con-sistent with the expectations for head-start ex-pressed by Klima (1978). FWS’ termination ofPAIS imprinting after 1988 (Woody 1990; JackWoody, pers. comm.) had already disconnectedhead-start from reintroduction, leaving head-startto be evaluated on its own merits and perfor-mance. The panel concurred with the decisionto end what it referred to as artificial imprint-ing at PAIS, noting that imprinting was simplyone unproven hypothesis to explain how sea tur-tles choose nesting beaches. Moreover, the panelstated that artificial imprinting to PAIS added ex-tra variables to the head-start experiment, all withpotential to interfere with its effectiveness. Thisview was especially ironic, because head-startwas essential to tagging turtles of year-classes1978–1988 to provide the means for evaluat-ing the reintroduction experiment. Likewise, itwas essential to tagging turtles of year-classes1989–2000 to provide the means for evaluatingthe head-start experiment. The panel did not de-scribe the extra variables or how they might in-terfere with effectiveness of head-start.

After 1988, hatchlings for continued head-starting were putatively imprinted in Tamaulipas,as using procedures similar to those employedat PAIS during 1978–1988. Standard proceduresfor collecting, transplanting, and incubating eggsas well as releasing emergent hatchlings intothe Gulf of Mexico from Tamaulipas beacheswere similarly manipulative, even though mostof the hatchlings were released (i.e., not col-lected for head-starting). The major differencewas that hatchlings for head-start, whether puta-tively imprinted at RN or PAIS, were collectedfrom the surf and sent to the Galveston Labora-tory. In Tamaulipas, most naturally laid clutcheswere collected and transplanted into artificial(prepared by humans) nest cavities within beachcorral hatcheries, but some were also placed inlocal sand within StyrofoamT M boxes (NMFSet al. 2011). Hatchlings that emerged were re-

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http://www.galvestonlab.sefsc.noaa.gov/publications/pdf/930.pdf

http://www.galvestonlab.sefsc.noaa.gov/publications/pdf/930.pdf


trieved, transferred in containers to release lo-cations, placed on the beach, and protected dur-ing their crawl to the surf (Márquez-M. et al.2005; Heppell et al. 2007). Hatchlings producedin nests left in situ (ibid.; Gallaway et al. 2013;Bevan et al. 2014) were not protected during theircrawl to the surf. Shaver and Wibbels (2007)reported an overall hatching success of 77.1%for eggs obtained from RN and incubated inStyrofoamT M boxes at PAIS (i.e., the 1978–1988year-classes), as compared to an overall hatchingsuccess of 63.1% for eggs incubated in corralsat RN (Shaver 2005). The main differences be-tween operations in Tamaulipas and PAIS werethe numbers of clutches transplanted and hatch-lings released, although there may have beenother differences involving methods of transport-ing eggs, transporting emergent hatchlings, anddistances crawled by the hatchlings to the surf.For head-start, hatchlings taken from Tamaulipasin 1978 and after 1988 were collected from thesurf and transported to the Galveston Laboratory,just as were those produced from eggs incubatedat PAIS. In other words, putative imprinting ofhead-started Kemp’s ridleys as hatchlings wasartificial in Tamaulipas as well as at PAIS.

Wibbels et al. (1989 op. cit.) lauded the Galve-ston Laboratory for its refinement of first-yearcaptive-rearing into what they called an exactscience, its having better facilities and staff thanelsewhere in the world for such purpose, and forits having increased public awareness of the sta-tus of sea turtles through the efforts of CaroleAllen and HEART. They noted that long termtag returns showed that head-started Kemp’s Ri-dleys could adapt and grow in the wild, and thatcaptive breeding studies indicated they could suc-cessfully reproduce in captivity. Wibbels et al.(1989 op. cit.) justified the cost of head-start bythe unprecedented opportunity to address its ef-fectiveness as a sea turtle conservation technique.In addition, peripheral research associated withhead-start was considered to have potential forproviding technologies and data that could en-hance sea turtle conservation in general (e.g., de-

velopment of permanent tagging techniques) andprovide life history information. Costs of reintro-duction and head-start did not appear to be majorconcerns of the early planners (NPS et al. 1978op. cit.), but they did not plan continuation ofhead-start beyond 11 y.

Wibbels et al. (1989 op. cit.) established thefollowing new criteria (paraphrased ) for assess-ing success of head-start: Provisional Criteria.(a) apparent competence of head-started turtlesat and after release, measured by their survival,growth, body weight, feeding behavior, orienta-tion, and reactions, as compared to those of wildKemp’s Ridleys, (b) ratio of recoveries (tag re-turns and strandings) of head-started to those ofnaturally occurring Kemp’s Ridleys, taking intoaccount the number of hatchlings produced at RNand the number of hatchlings head-started (i.e.,produced at PAIS), as well as possible biasedsampling due to presence of tags on head-startedturtles, and (c) comparison of recovery locationsof head-started and wild Kemp’s Ridleys; and Ul-timate Criterion. The proportion of nesting head-started females should increase gradually overa 5-y period relative to the proportion of wildnesting females. Recommendations by Wibbelset al. (1989 op. cit.) included (1) limiting head-starting to 2,000 hatchlings per year and ≤ 1 y,and releasing all healthy and normally developedindividuals (i.e., without obvious genetic defects)after about 8–12 mo of captive-rearing, with theexception of captive-rearing a minimal numberfor ≤ 2 y for use in TED testing, (2) using tagreturns and stranding data from head-started andwild Kemp’s Ridleys to provide insight regardingcompetence of head-started turtles in the wild ascompared to wild counterparts, (3) continuing de-velopment of practical means of assessing turtlefitness, using controlled feeding to prevent themfrom becoming too fat, and using physical andphysiological data to guide such procedures, (4)continuing research on tagging with conventionaltags, living tags, CWTs, and PITs, (5) continu-ing research to develop a practical and non-lethalsexing technique, especially for hatchlings, (6)

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continuing research on sex ratios of head-startedturtles with the aim of achieving a 50F:50M ratioor one with a slight female bias, but certainlynot one > 70% female, by changing nest loca-tion or choosing eggs early or late during thenesting season at RN, (7) continuing research to-ward developing radio- and sonic-tracking tech-nology on both head-started and wild Kemp’sRidleys, to generate baseline data on ecology, be-havior, and competence of the head-started turtlesin the wild, (8) continuing recording and compil-ing strandings and sightings of head-started andwild Kemp’s Ridleys, (9) subjecting internal andexternal research proposals concerning head-startto external review, and developing a specific pro-tocol for submitting research proposals regardinghead-started Kemp’s Ridleys, and (10) requir-ing that the entire head-start staff emphasize tothe public that head-start is an experiment, thatit should not be viewed as the means of savingKemp’s Ridley, and that the primary element ofKemp’s Ridley recovery is protection in the natu-ral habitat (e.g., with TEDs).

Wibbels et al. (1989 op. cit.) concluded that itwas impossible at that time to determine if head-started Kemp’s Ridleys recruited into the naturalbreeding pool, because shrimping-induced mor-tality rate of both wild and head-started Kemp’sRidleys was so high that few head-started turtleswere expected to reach sexual maturity. Theyemphasized that head-start should not be viewedas a panacea for sea turtle conservation or ameans of restoring the population. Otherwise,they warned, protection of the species in itsnatural habitat could be jeopardized. The panelrecommended that head-start be continued butnot expanded for 10 y following installation ofTEDs on all shrimping vessels in U.S. watersof the Gulf of Mexico and Atlantic. Thatrecommendation was based on estimates of thetime required to reach sexual maturity.

Eckert et al. (1994) panel review.—In August1992, three years after the Blue Ribbon PanelReview, Eckert et al. (1994) reviewed the experi-

mental design of head-start. Although imprintingat PAIS was mentioned as having been concur-rent with head-starting through 1988, after whichit was discontinued, it was given no further at-tention except to state that the imprinting pro-gram was subsequently focused on monitoringat PAIS for head-started nesters. In reality, puta-tive imprinting associated with head-start contin-ued; its location was simply shifted from PAIS toTamaulipas. Like Wibbels et al. (1989 op. cit.),Eckert et al. (1994) characterized head-startingas “controversial among sea turtle biologists al-most from its inception,” and cited numerous pa-pers supporting this characterization. They indi-cated that criticisms of head-starting tended to re-volve around concerns that head-started Kemp’sRidleys would not nest or integrate into the wildpopulation due to their confinement during thefirst year of life, and that head-starting acted tomollify resource policy makers into believing thatthe causes of the dramatic decline in the Kemp’sRidley population did not need to be addressed.They pointed out that proponents of head-startinggenerally responded to such criticisms by claim-ing not enough time had elapsed for the turtlesto reach maturity and nest, that marking tech-niques in the early years were inadequate, andthat tagged turtles did not remain tagged longenough to be recognized as head-started whenmature and encountered on the nesting beach.

Eckert et al. (1994) emphasized that head-starting was an experiment and not a mitigationmeasure or technique. Their review focused in-stead on whether the experimental design was ap-propriate, and on recommendations they thoughtwould improve it. Eckert et al. (1994) indicatedthat head-starting should be judged on economicand policy bases if and when proven feasible.Their stated goal was to improve the experimen-tal design so that it would provide scientific in-formation needed to determine whether or nothead-starting should be continued. They exam-ined head-starting based on four questions: (1) Isthere a testable hypothesis?; (2) Are sample sizesadequate and unbiased?; (3) Does the experiment

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have a control and an experimental group?; and(4) Does the project have an end point or finalgoal?

Eckert et al. (1994) concluded that, at its incep-tion, head-starting had no experimental hypothe-sis that could be tested to determine its effective-ness in supporting the wild population. They at-tributed this alleged lack to a focus on challengesof captive-rearing, to the extent that considera-tion of an experimental design was overlooked,and to an apparent assumption “that survival inthe marine environment was assured after hatch-lings were raised for a year.” They acknowledgedthat captive-rearing had become very success-ful, and that an experimental hypothesis or setof goals had emerged (i.e., Wibbels et al. 1989op. cit.) because of outside pressure. Eckert etal. (1994) generally agreed with and applaudedthe provisional criteria proposed by Wibbels et al.(1989 op. cit.) to guide head-starting, but consid-ered the ultimate criterion to be flawed. In 1992,the number of nestings by wild turtles was in-creasing, but no nestings by head-started turtleshad been documented. The panel accurately con-cluded that the only way the proportion of nest-ings by head-started turtles could increase rela-tive to that of wild turtles would be for nestings byhead-started turtles to increase more rapidly thanthose of wild turtles. Furthermore, the annualnumber of head-started Kemp’s Ridleys releasedinto the Gulf of Mexico decreased after 1993,and releases were sporadic after 2001, while wildhatchling releases increased exponentially (Gall-away et al. 2013; Caillouet 2014). Eckert et al.(1994) suggested a revised goal for the head-startexperiment, expressed in a two-part hypothesis:head-starting can produce Kemp’s Ridley juve-niles that are able to join the natural, wild popula-tions, find their way to nesting beaches, procreate,and hatch viable offspring of their own, and head-started turtles demonstrate equivalent or superiorbiological fitness (defined as equal or better sur-vival rates from egg to reproductive adult, andequivalent or better fecundity) when compared tothose of wild Kemp’s Ridleys. They did not spec-

ify particular nesting beaches under the first part,which implied that they were more concernedwith whether head-started Kemp’s Ridleys wouldfind any nesting beach and procreate, than withwhich beach the turtles might choose for nesting.The first part of their hypothesis has evidence tosupport it (Caillouet 1998; Fontaine and Shaver2005; Shaver and Wibbels 2007; Shaver and Cail-louet 2015). The second part has not been fullysupported, although fecundity of head-startednesters has been shown to be similar to that ofwild nesters (Shaver and Wibbels 2007; Shaverand Caillouet 2015).

With regard to the second part of the Eckertet al. (1994) hypothesis, we believe it is unreal-istic to evaluate head-starting on a basis of itsproducing nesters of biological fitness superiorto that of wild Kemp’s Ridleys. A more appro-priate, alternate hypothesis would expect head-starting of a given number of hatchlings to returnmore nesters to Tamaulipas than the same num-ber of hatchlings released there (see Campbell1977 op. cit.). Expecting superior biological fit-ness of head-started turtles implies that captive-rearing could somehow improve them in waysother than simply increasing short term survivalin captivity, as compared to their wild counter-parts of similar sizes and ages. Efforts directedtoward maintaining physiological fitness duringhead-starting were not fully successful. Althoughsurvival rate during head-starting (Fontaine et al.1989b; Fontaine and Shaver 2005; Shaver andWibbels 2007) was higher than that estimatedfor wild Kemp’s Ridleys of similar sizes andages (NMFS et al. 2011), other factors relatedto head-start may have offset this initial advan-tage after the turtles were released. Equivalentbiological fitness in head-started and wild turtleswould be a more reasonable null hypothesis forevaluating success, performance, or effectivenessof head-start as a potential recovery method. Weare unaware that any other Kemp’s Ridley conser-vation or management method has been expectedto produce turtles of superior biological fitnessas a criterion for declaring it effective. Regard-

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less, the second part has been partially fulfilledby determining the number of eggs per clutchfor head-started Kemp’s Ridley nesters (Shaverand Caillouet 2015). Head-start by itself, discon-nected from its important role in reintroductionof Kemp’s Ridley to PAIS, has not been demon-strated to be successful as a mitigation techniqueor conservation method. The likelihood that itmight eventually be considered successful dimin-ishes as the number of head-started Kemp’s Rid-leys in the population declines, even though addi-tional head-started nesters (ones not previouslydocumented) may be found in the future. Thislikelihood is further reduced by the apparentlylimited efforts directed toward searching for head-started Kemp’s Ridley nesters anywhere exceptat PAIS and other beaches in Texas (Shaver andCaillouet 2015). However, if offspring of head-started nesters can eventually be distinguishedgenetically from those produced by wild nesters,their contributions to the Kemp’s Ridley popula-tion may be estimable in the future.

Eckert et al. (1994) concluded that the con-trol group representing wild Kemp’s Ridleyshad been overlooked, leaving no comparisonby which to measure the head-start experiment.They pointed out that stranding data used to de-scribe distribution of the species in the Gulf ofMexico were biased by representing only tur-tles that died near the coast and washed ashore,while tagged turtles were more readily reported.Therefore, they recommended a rigorous mark-recapture program based on archival tagging ofwild Kemp’s Ridley hatchlings, and maintenanceof previously attached or implanted tags on head-started Kemp’s Ridleys, for purposes of determin-ing rates of growth, survival, tag retention, anda host of important life history characteristics inboth groups. They emphasized that valid compar-isons could only be based on turtles recapturedduring the proposed tag-recapture program, toavoid biases associated with comparisons of re-captures in pre-TED years with those in years inwhich TEDs were required.

Eckert et al. (1994) defined head-started turtles

as the experimental group and wild turtles as thecontrol group, then addressed the topic of samplesize required to detect effects of introducinghead-started turtles into the population. Forthe control group, they proposed mass-tagginghatchlings at RN with the same types of archivaltags (CWTs or PIT tags) that had been appliedto head-started Kemp’s Ridleys, so the twogroups could be distinguished from one another,especially on nesting beaches. By contrast,although NPS et al. (1978 op. cit.) called forcontinued tagging of Kemp’s Ridley nesters atRN, this tagging was not intended to evaluatesuccess of reintroduction or head-starting.Instead, permanent tagging of wild femalesnesting at RN was proposed as a continuationof ongoing work conducted by INP duringyears prior to initiation of the KRREP (MárquezM. et al. 1982). Interestingly, Marquez-M.(1994) noted that very little tagging of Kemp’sRidleys had been done outside of RN, andindicated that the increase in tagging followingimplementation of the KRREP was especiallythat of head-started turtles. In any case, a controlgroup of wild adult females tagged at RN hadalready been established and enhanced overthe years. In addition, tagging of wild Kemp’sRidleys of any life stage from hatchlings toadults was not necessary to distinguish themfrom multi-tagged, head-started Kemp’s Ridleys,unless all external and internal (archival) tagsapplied to head-started turtles were lost, evenwhen tagging scars remained. Assuming that ahead-started Kemp’s Ridley lost its foreflippertag(s) but retained a living tag, PIT, or CWT, itprobably would have been identified erroneouslyas wild if not examined for living tags, PITs, orCWTs. Therefore, the implant locations, codes,or other characteristics of internal archival tagsimplanted into wild Kemp’s Ridleys of any lifestage had to be different from those for internalarchival tags applied to head-started turtles, sothat internally tagged turtles in these two groupscould be distinguished by their tags (Fontaineet al. 1993; Caillouet et al. 1997b; Higgins et

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al. 1997). Nevertheless, it was potentially veryuseful to tag large numbers of wild hatchlingsat RN with archival tags that could link themby year-class to their RN origin, and to hatchdates from which their ages could be determinedupon recapture. Interestingly, this was similarto the expected utility of archival living tagsapplied to hatchling turtles, as envisionedby John and Lupe Hendrickson in the 1980s(Hendrickson and Hendrickson 1981, 1983;Mrosovsky and Godfrey 2003; Kishinami,C. 2003. John Roscoe Hendrickson biogra-phy: A daughter’s memories. Available fromhttp://www.cedointercultural.org/JRHtext.htm[Accessed 8 December 2014]).

Use of PIT tags on large numbers of hatchlingswas considered cost-prohibitive by the Galve-ston Laboratory, so only CWTs were consideredand used for mass-tagging hatchlings at RN, asrecommended by Eckert et al. (1994). Prior tothe mass-tagging, three laboratory studies wereconducted to determine the effects of tagginghatchlings with CWTs, the first on Loggerheadin 1993, and the other two on Kemp’s Ridleysin 1994 and 1995 (Higgins et al. 1997). Mass-tagging of Kemp’s Ridley hatchlings with CWTsat RN took place in 1996 (3,336), 1997 (10,002),1999 (10,010), and 2000 (20,537), for a total of43,885 hatchlings (Caillouet 1998; Snover andHohn 2004; Snover et al. 2007; Benjamin Hig-gins, pers. comm.). This 4-y total was more thantwice the 23-y total for head-started Kemp’s Ri-dleys tagged and released, but not all of thesehead-started turtles were tagged with CWTs (seeTags and tagging).

Fontaine et al. (1993) and Higgins et al. (1997)explained the proper protocols required to detectand decode CWTs, whether magnetized or notbefore release of the turtles. If these protocolsare not carefully followed, CWTs will not bedetected (Fontaine et al. 1993; Caillouet et al.1997b; Higgins et al. 1997; Benjamin Higgins,pers. comm.). More importantly, tagging of head-started year-classes had already been conductedin ways that distinguished them from the mass-

tagged hatchling year-classes, making it essentialthat the established detection and decoding pro-tocols be followed (ibid.).

Eckert et al. (1994) acknowledged that theirrecommendations included some protocols thatwould be considered impossible for most sea tur-tle species, but believed them possible for Kemp’sRidley because of its small population that nestedon a single primary beach. The ambitious, large-scale mark-recapture program they recommendedfor NMFS was aimed at gathering the followinginformation on head-started and wild Kemp’s Ri-dleys: (1) survival rate of hatchlings to maturity,(2) average survival rate of juveniles to maturity,(3) growth rates of juveniles, (4) behavior, in-cluding habitat selection, movement, and migra-tion patterns, (5) physiology, comparing physicalfitness of head-started and wild caught turtles,(6) sex ratios of in situ populations, (7) size fre-quency distributions of juvenile populations, and(8) age at maturity. They recommended furtherthat NMFS: (1) establish a large number of net-ting and capture areas around the Gulf of Mexicoand U.S. east coast for purposes of capturingand archival tagging Kemp’s Ridleys, and gath-ering data on size, sex, habitat characteristics,physiological parameters, and food preferences,(2) demonstrate that magnetized CWTs or PITtags do not adversely impact hatchlings, (3) tag(during two consecutive nesting seasons) a largenumber of wild hatchlings with archival tags thatidentified their year-classes, and release them atRN for the purpose of determining survival ratesfrom hatchling to maturity, (4) assess tag reten-tion rates for wild and head-started turtles at dif-ferent life stages, (5) double-tag all head-startedand wild-captured turtles with visible tags, andat least single-tag them with archival tags, and(6) organize and support the tagging program toensure its longevity and provide means to detectarchival tags by the wide range of groups thatmight encounter tagged turtles.

Eckert et al. (1994) recognized that detectionof tagged turtles (head-started or wild) on nest-ing beaches depended upon levels of searching

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http://www.cedointercultural.org/JRHtext.htm


for nesters and checking them for tags, as wellas the numbers of marked turtles available to bedetected; i.e., the smaller the numbers the greaterthe detection effort required and vice versa. Eck-ert et al. (1994) considered the relatively smallnumbers of head-started turtles disadvantageousbecause they required a high level of effort tobe detected, yet they also considered the rela-tively small numbers advantageous because ofwhat they indicated were fewer turtles that wereimpacted by the experimental technique of head-starting. We reiterate that annual numbers of eggsand hatchlings for reintroduction and head-start(Caillouet 1995b) were purposefully limited fromthe onset by NPS et al. (1978 op. cit.), and theselimits were controlled by the participating agen-cies. Eckert et al. (1994) acknowledged that thelevel of effort aimed at detecting head-startedKemp’s Ridley nesters at RN was not high, andcited Pritchard (1990) who estimated that only48.4% of Kemp’s Ridleys nesting at RN in 1989were seen during beach monitoring. This 48.4%detection rate is comparable to that in Texas(Shaver and Caillouet 2015), and it may be ashigh as can be expected given that it takes onlyabout 45 min from the time a nester leaves thewater to the time she returns after nesting. More-over, Eckert et al. (1994) stated that tagging tech-niques applied to head-started Kemp’s Ridleysvirtually guaranteed they would not be detected.However, tag returns reported prior to the Eck-ert et al. (1994) review and thereafter indicatedthat head-started Kemp’s Ridleys could indeed befound when sufficient effort was directed towardsearching for, finding them, and examining themfor tags (Shaver and Caillouet 1998; Fontaine andShaver 2005; Shaver and Wibbels 2007). Never-theless, we concur with Eckert et al. (1994) thatthe probability of detecting head-started Kemp’sRidley nesters was and continues to be muchlower than that of wild nesters (whether taggedor not). This is an expected result of the muchlarger and increasing (through 2009) numbers ofwild Kemp’s Ridley females in the population ascompared to the much lower and declining num-

bers of head-started Kemp’s Ridley females, com-bined with relatively low efforts toward searchingfor and detecting head-started nesters outside ofTexas.

The search for Kemp’s Ridley nesters in Texas,head-started and wild, has been much moreintense at PAIS, SPI, and Boca Chica Beach(near the mouth of the Rio Grande river) thanelsewhere in Texas (Shaver and Wibbels 2007;Shaver and Caillouet 2015). Coupled with thefact that most head-started Kemp’s Ridleys werereleased along the Texas coast, the greater ef-fort in searching for them as nesters on Texasbeaches probably increases the likelihood of find-ing them there than elsewhere in the Gulf of Mex-ico or North Atlantic Ocean. In addition, PAISwas the only area of the Texas coast selected theKemp’s Ridley reintroduction experiment (NPSet al. 1978 op. cit.); it continues to be the epi-center of nesting in Texas (Shaver and Caillouet2015).

Eckert et al. (1994) apparently did not antic-ipate that their proposed tagging of wild hatch-lings with archival tags at RN, if implemented,would provide a means of measuring the effort di-rected toward their detection and documentation.Because the hatchlings were tagged and releasedat RN, it was expected that surviving femaleswould return as adults to nest at RN, assuminghoming and fidelity to the natal beach. To ourknowledge, there have been no documented nest-ings by survivors of this control group at RN orelsewhere. If some females from each of the fouryear-classes in this group survived through the2014 nesting season, they would be 14, 15, 17,and 18 y old adults capable of nesting. Yet, therehave been relatively few documented tag returnsfrom any life stage in this group (Snover andHohn 2004; Snover et al. 2005, 2007; Dodge etal. 2008), suggesting that efforts to detect themhave been limited at best. This paucity of tagreturns could also be due in part to inadequacyof training of personnel to search for Kemp’sRidleys in this control group, lack of required de-tection equipment, or both (Higgins et al. 1997;

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Benjamin Higgins, pers. comm.). In addition, itis time-consuming to locate these tags, using rec-ommended procedures (Fontaine et al. 1993; seeTags and Tagging), and the detection equipmentis very expensive. Nevertheless, assuming thatsome survivors from this control group remain inthe population, they represent a highly valuableresource that could provide the kinds of informa-tion envisioned by Hendrickson and Hendrickson(1981, 1983) and Eckert et al. (1994), if theycould be found and their tags detected (Caillouetet al. 1997b). At the very least, archived flippersof dead specimens can be examined for CWTs aswell as other tags.

Eckert et al. (1994) commented on the rele-vance of the Blue Ribbon Panel’s (Wibbels et al.1989 op. cit.) suggestion that few if any head-started Kemp’s Ridleys were expected to reachmaturity because shrimping-induced mortalityrates were so high. Eckert et al. (1994) agreedthat mortality rates associated with trawling werelikely high, and that TEDs should be required onall shrimping vessels, but suggested that coverageof the nesting beach was probably not adequateto detect any head-started turtles even if mortalitydue to shrimping were nil. They proposed thatbeach coverage at RN be increased to observeall turtles that nest there and that all field teamsbe outfitted to detect head-started turtles. Theysuggested that increased beach coverage shouldbe able to determine whether any head-startedturtles were currently nesting there. They alsowere uncomfortable with relying on living tagsto detect head-started turtles, suggesting thesetags could be easily misinterpreted and that re-search on their retention and detection rate in thewild had been inadequate. We disagree with theirassessment of the living tag, which has provenespecially useful in documenting head-startedKemp’s Ridley nestings, whether by itself or incombination with other retained tags (Shaver andCaillouet 2015).

Despite their detailed discussion of factorsthat reduced the probability of documentinghead-started Kemp’s Ridley nesters, Eckert et al.

(1994) suggested that numbers of head-startedturtles already tagged with CWTs, PIT tags, orboth, and released were adequate to representthe experimental group, and that it was unnec-essary to release any more head-started turtles.However, the Galveston Laboratory continued tohead-start and tag Kemp’s Ridleys (year-classes1993–2000) with foreflipper tags, living tags,CWTs, and PIT tags, and release them off

Galveston, albeit based on about 200 hatchlingsreceived per y. The mass-tagging of hatchlingsat RN provided a large and very valuableresource of tagged, wild Kemp’s Ridleys ofknown natal beach origin and approximateage, calculable from hatch and recapture dates.Unless greater efforts are directed towardsearching for, detecting, and documenting tagreturns of survivors in this control group, thisimportant resource will have been wasted. Inour view, survivors from these releases arepotentially more valuable toward providingimportant biological information about the wildpopulation of Kemp’s Ridleys, than as a controlgroup to test head-start. Recommendations byEckert et al. (1994) would also be applicable tomass-tagging (with CWTs or PIT tags) of wildKemp’s Ridleys along the U.S. Atlantic coast, toprovide a means of determining the proportionthat returns to nest at RN or other Gulf ofMexico beaches. It can safely be assumed thatmost Kemp’s Ridleys along the U.S. Atlanticcoast originated from nestings on beaches inthe western Gulf of Mexico, especially those inTamaulipas (NMFS et al. 2011). One Kemp’sRidley that nested at PAIS had been rehabili-tated in Massachusetts, tagged, and released.Published records documenting Kemp’s Ridleyreturns from the Atlantic to the Gulf of Mexicoappear to be fewer than 20, and all were reportednesting at or near RN (Schmid 1995; Chaloupkaand Zug 1997; Schmid and Witzell 1997; Witzell1998; Schmid and Woodhead 2000). Untilsubstantially more evidence is amassed to thecontrary, it will continue to appear that mostKemp’s Ridleys that nest in the Gulf of Mexico

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have not spent time in the Atlantic.

Evaluations via demographic modeling.—Heppell et al. (1996) emphasized that survivaland growth rates for both head-started and wildKemp’s Ridleys must be known before head-startcan be evaluated. Survival rate of head-startedKemp’s Ridleys from egg to reproductive adulthas not been determined per se. However,survival rates from eggs to hatchlings (Shaver2005; Shaver and Wibbels 2007; Shaver andCaillouet 2015) and hatchling to yearlings(Fontaine and Shaver 2005) in captivity, andfollowing release of yearlings into the Gulfof Mexico (Caillouet et al. 1995a) have beenestimated. For comparison, survival rates ofvarious life stages of wild Kemp’s Ridleys havebeen estimated (TEWG 1998, 2000; Heppell etal. 2005, 2007; NMFS et al. 2011; Gallaway etal. 2013). Based on matrix modeling, Heppelland Crowder (1994) predicted that head-startingcould not contribute significantly to the Kemp’sRidley population, even if head-started juvenileshad the same survival and growth rates as wildjuveniles. Crowder et al. (1998) stated that thebest use of this type of analysis was to eliminatemanagement alternatives unlikely to lead topopulation recovery, and gave Kemp’s Ridleyhead-start as an example. Using a series ofhypothetical matrix models, each with a differentage at maturity, Heppell and Crowder (1998)showed that only slight increases in the numberof nesting Kemp’s Ridley females might beexpected from a head-start program, but theseincreases were inconsequential compared to thepotential benefits of TEDs.

Discussion

Meylan and Ehrenfeld (2000) reviewed ac-tive intervention and management approachesthat promote sea turtle conservation and pre-vent sea turtle populations from declining. Theysuggested that Bjorndal (1982) remains a use-

ful guide that advocates least-manipulative tech-niques. Meylan and Ehrenfeld (2000) explainedthat an inherent problem with manipulative man-agement techniques is that once one is chosen,many decisions are required to implement it, andin most cases the information needed to guidethe decisions is inadequate. Nevertheless, Mey-lan and Ehrenfeld (2000) characterized Kemp’sRidley head-start as the world’s premier head-start project, and indicated that it complementedefforts to protect nests and release hatchlings atthe nesting beach in Mexico.

There can be no doubt that reintroduction andhead-start were highly manipulative. The earlyplanners were focused on reestablishing Kemp’sRidley nesting at PAIS, and proposed methodsthey thought would accomplish this goal, basedon limited information available at the time. Theexisting nesting colony at PAIS (Plotkin 1999;Lewison et al. 2003; Shaver and Caillouet 2015)is evidence that the early planning was sufficient.In December 2000, TPWD implemented regula-tions prohibiting shrimp trawling within 5 mi ofthe southern Texas coast from 1 December eachyear through 15 July of the next year (Lewisonet al. 2003). Lewison et al. (2003) implied thatthe second nesting population on Padre Islandwas worthy of protection by this seasonal closureto shrimping. They estimated that this closurewould likely reduce mortality of adult Kemp’sRidleys by as much as 39%, and “protect a sec-ond Kemp’s ridley nesting beach to safeguardthe population should a catastrophe affect the pri-mary nesting beach in Rancho Nuevo, Mexico.”

Inadequacy of information was also evident inthe decision to terminate imprinting at PAIS in1989, while allowing head-start to continue withhatchlings imprinted at RN. However, timing ofthis termination was consistent with the scheduleproposed by NPS et al. (1978 op. cit.). In addi-tion, there was concern that the reintroductionand head-start experiments were giving a falseimpression, within the U.S. and elsewhere, thatthey were proven sea turtle conservation methods,and this impression might undermine protection

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of the wild stock (Jack Woody, pers. comm.).Regardless, this decision not only reduced theprobability that reintroduction would achieve itsgoal, but it also separated head-start from rein-troduction, thereby raising the bar for evaluationof head-start as a potential stand-alone conser-vation measure. In retrospect, this decision didnot prevent reintroduction of Kemp’s Ridley toPAIS, and it allowed head-start to be evaluatedseparately, on its own merit and performance.

Although Kemp’s Ridley head-start involvedmanipulation of egg through yearling life stages,once the surviving yearlings were released intothe Gulf of Mexico they were left to fendfor themselves (Eckert et al. 1994; Caillouetet al. 1995a, 1995b; Caillouet 2005; Fontaineand Shaver 2005; Shaver and Wibbels 2007).The methods used at RN and PAIS to supplyhatchlings for head-start, combined with head-start methods used at the Galveston Laboratory,boosted survival rates of life stages from eggthrough yearling (Fontaine and Shaver 2005;Shaver and Wibbels 2007; Shaver and Caillouet2015) as compared to survival rates of wildKemp’s Ridleys of similar life stages. Operationsin Tamaulipas and PAIS that produced hatchlingsfor head-start were short-term compared to thosethat extended human control and manipulation atthe Galveston Laboratory.

It is important, especially in the context ofthe current situation facing Kemp’s Ridley (Cail-louet 2014), to consider that reintroduction ofKemp’s Ridley to PAIS might have been accom-plished without head-start, by annually trans-planting eggs to PAIS from RN, and releasingemergent hatchlings at PAIS instead of head-starting them. However, had hatchlings been re-leased at PAIS instead of being head-started,reintroduction might have taken longer than itdid when combined with head-starting, becausehatchling mortality is naturally high. Conversely,head-starting may have decreased survival andperformance after release as compared to wildKemp’s Ridleys of similar ages and sizes. Turtlesthat died during head-starting were the source

of sex ratio information that was used to deter-mine pivotal incubation temperature; without thisinformation, it would have taken longer to de-termine pivotal temperature for Kemp’s Ridleys.As it was, only four year-classes (1985–1988) ofPAIS-imprinted hatchlings were female-biased(Shaver and Caillouet 2015).

Finally, determining success of releasing hatch-lings at PAIS (instead of head-starting them), asan alternative means of reintroducing nesting toPAIS, would not have been possible without tag-ging hatchlings. Without head-start and tagging,any increased nesting at PAIS that might haveresulted from releasing hatchlings at PAIS prob-ably would have been attributed to other conser-vation efforts and not to reintroduction efforts.This underscores the need for better methods formass-tagging hatchlings (Allen 1981).

Manipulative management techniques wereand still are employed in Tamaulipas on a verylarge scale (Burchfield and Peña 2013). During1978–1992, 1,286,900 eggs were produced atRN as compared to an estimated 36,025 eggstaken for reintroduction and head-start during thesame period (Caillouet 1995b), a ratio of about36:1. During 2009 alone, 1,089,452 hatchlingswere released from Tamaulipas beaches (Burch-field 2009). The manipulative management pro-cedures employed in Tamaulipas gave eggs andhatchlings the substantial boosts in terrestrial sur-vival rate that saved Kemp’s Ridley from extinc-tion and helped put it on an exponential trajectory(through 2009) toward recovery (Table 21). It isalso possible that such procedures applied towardincreasing the Kemp’s Ridley population mighthave had some negative effects (Burke 1991;Dodd and Seigel 1991; Meylan and Ehrenfeld2000; Mrosovsky 2007).

Reintroduction was not planned as a means oftesting the imprinting hypothesis, but its opera-tions were designed with the intention of imprint-ing Kemp’s Ridleys to PAIS before head-startingthem (NPS et al. 1978 op. cit.). Imprinting wasseen as necessary to reintroduction and head-start.However, even though most wild Kemp’s Ridleys

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Table 21. Sources covering the manipulative man-agement procedures employed in Tamaulipas, andthe reductions in at-sea mortality due to use of tur-tle excluder devices (TEDs) and declining shrimptrawling effort, that helped put the Kemp’s Ridley(Lepidochelys kempii)population on an exponentialtrajectory (through year 2009) toward recovery.

Heppell and Crowder (1994, 1998)Heppell et al. (1996, 2005, 2007)Heppell (1997)TEWG (1998, 2000)Epperly (2003)Lewison et al. (2003)Caillouet (2005, 2006, 2010, 2011, 2014)Márquez-M. (2001)Márquez-M. et al. (2005)Crowder and Heppell (2011)NMFS et al. (2011)Gallaway et al. (2013)Caillouet (2014)

nest on Gulf of Mexico beaches of Mexico (espe-cially at RN), significant numbers nest in Texas,and some individuals have been documented tonest both in Mexico and Texas, suggesting thatnatal beach imprinting is not absolute (Shaverand Caillouet 2015). Nevertheless, reintroduc-tion and head-start can both be evaluated indi-rectly in the context of the imprinting hypothesis(Shaver and Caillouet 2015).

Putman and Lohmann (2008) hypothesizedthat Kemp’s Ridleys detect two elements of themagnetic field, inclination angle and total in-tensity, as Loggerhead turtles are known to do(Lohmann and Lohmann 1994, 1996). Putmanand Lohmann (2008) examined the hypothesisthat Kemp’s Ridley hatchlings at RN imprint onone of these elements of RN’s magnetic field anduse this information to navigate and return thereas adults to reproduce. As a first step, they con-ducted modeling under assumptions of 10 y and15 y at sexual maturity, and an inability of theturtles to compensate for gradual drift in mag-netic signatures of RN over time. Theoretically,if the turtles imprinted on inclination angle at

RN, they could return within an average distanceof 23 km from RN if absent 10 y, or within anaverage distance of 32 km if absent 15 y. If theyimprinted on magnetic field intensity, they couldreturn within an average distance of 89 km fromRN if absent 10 y, and within an average distanceof 132 km if absent 15 y. These results were con-sistent with the known precision of natal homingin Kemp’s Ridleys, and indicated that magneticimprinting could return turtles sufficiently closeto their natal region where they could then useother local cues to pinpoint particular nesting ar-eas. Simply stated, if turtles learn the magneticsignature of their home area before hatching, oracquire this information during their crawl to thesurf, and if magnetic navigation is primary inguiding sea turtles to their natal beach as adults,then head-started Kemp’s Ridley imprinted atPAIS should have homed on PAIS as adults andthose imprinted at RN should have homed onRN as adults, unless other factors prevented ordisrupted the effects of magnetic imprinting.

According to David Owens (pers. comm.), asimilar case can made for homing based on ol-factory imprinting to geological and faunal cues.However, a limitation of olfactory imprinting isthat odorants typically do not extend long dis-tances through the sea, and turtles approachingfrom directions that are not down-current pre-sumably cannot detect odorants from the natalarea until they are very near the target (Lohmannet al. 1999, 2013). For this reason, adult turtleshave been hypothesized to use two different setsof cues sequentially as they navigate to their na-tal beaches: magnetic information may be usedover long distances to guide turtles into the gen-eral vicinity of their nesting areas, and local cues(e.g., olfactory, visual, or both) may be involvedin selecting the final nesting sites (Lohmann et al.2008a, 2008b).

Putman et al. (2010) investigated the hypoth-esis that relative abundance of Kemp’s Ridleynests on beaches in seven coastal regions of thewestern Gulf of Mexico can be explained bythe location and characteristics of surface cur-

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rents that facilitate passive transport of pelagichatchlings and early juveniles (≤ 2 y old) tocoastal foraging grounds in the Gulf and westernNorth Atlantic. The seven regions were Texas,North of RN, RN, South of RN, Veracruz, SouthCampeche, and North Campeche. Computer sim-ulated particles (hypothetical hatchlings) werereleased 45–55 km offshore from each regionduring 1 June–20 July in 2004–2007, and theirpattern of passive dispersal over two years wassimulated. Annual results were averaged and theaverages used to rank the seven regions accord-ing to overall favorability of surface circulationin dispersing the turtles. Surface circulation wasmost favorable to turtles originating at RN wheremost nests occur, followed by South Campeche,Veracruz, North of RN, Texas, North Campeche,and South of RN. Highest to lowest rankings byestimated nest abundance were RN, Veracruz,South of RN, Texas, and South Campeche, withNorth Campeche and North of RN tied at zeronests. It is interesting that Texas placed fifth inranking by favorability of surface circulation,but fourth in ranking by estimated abundanceof nests. In other words, Texas ranked higherin estimated abundance of nests than did SouthCampeche, North Campeche, and North of RN,two of which (South Campeche and North of RN)outranked Texas in favorability of surface circu-lation. Especially significant is that Texas’ esti-mated nest abundance outranked that for North ofRN (i.e., between RN and Texas) as well as SouthCampeche, which had the second most favorablesurface circulation ranking. In addition, Texas’estimated nest abundance outranked that of NorthCampeche. Since most nesting in Texas occurs atPAIS, this suggests that reintroduction of Kemp’sRidley to PAIS took place despite Texas’ lessfavorable surface circulation ranking.

Because exponential growth in the Kemp’s Rid-ley population was interrupted in 2010, and nestsnumbers declined in 2013 and 2014 (Caillouet2014), it may be necessary to intensify the ma-nipulative conservation approaches in Tamauli-pas that contributed to exponential increase in

the population through 2009. We will not makespecific recommendations in this regard. How-ever, although not necessary for restoration ofpopulation growth, an improved lifetime tag, ap-plicable to large samples of hatchlings, wouldbe useful to identify their year-class and beachof origin (Hendrickson and Hendrickson 1981,1983; Eckert et al. 1994), as a means of evalu-ating these approaches. We suggest that a lessinvasive, non-surgical, living tag be developedfor this purpose, one applicable to marking morethan one scute per hatchling, since marking com-binations of scutes increases the number of year-classes that can be distinguished from one an-other. We also recommend that greater attentionbe paid to searching for and documenting recap-tures of tagged Kemp’s Ridleys, especially onnesting beaches.

How Should Success be Judged?

Ultimately, the general question arises as tohow success of Kemp’s Ridley head-start andreintroduction should be judged (Burke 1991;Dodd and Seigel 1991). According to Dodd andSeigel (1991), the goal of any conservation pro-gram is establishment (or enhancement) of a vi-able, self-sustaining population that is at leaststable. This implies that the KRREP and relatedconservation efforts would have to establish or en-hance such a Kemp’s Ridley population to provetheir success. Burke (1991) responded to Doddand Seigel (1991), indicating that (1) it is dif-ficult to determine at what point a populationbecomes self-sustaining and stable, (2) such pop-ulation characteristics are subject to variability,and (3) there are no guarantees they are sustain-able with or without human intervention. Thebi-national recovery plan (NMFS et al. 2011)provides criteria for assessing when the Kemp’sRidley population has recovered and can be re-moved from the list of endangered species (i.e.,delisted, after first being down-listed to threat-ened status). These criteria represent very highstandards for evaluating success of Kemp’s Ri-

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dley conservation efforts. Since Kemp’s Ridleyhas not yet been down-listed to threatened status,neither the KRREP nor any other conservationmeasures aimed at Kemp’s Ridley recovery havesingly or in combination proven successful bythe standards of Dodd and Seigel (1991). How-ever, it is safe to say that conservation effortswithin the KRREP and those outside the KRREP(e.g., TED regulations and enforcement), as wellas declining shrimp trawling effort, helped setthe Kemp’s Ridley population on an exponen-tial trajectory toward recovery (Table 21) thatlasted until it was interrupted in 2010 (Caillouet2011; Crowder and Heppell 2011; Gallaway et al.2013; Caillouet 2014). The exponential increase(through 2009) in numbers of clutches laid onTamaulipas beaches certainly was a highly pos-itive outcome, because it indicated that conser-vation efforts and some exogenous factors (e.g.,declining shrimping effort) worked in positiveways toward Kemp’s Ridley recovery. Protectingnesters, eggs, and hatchlings in Tamaulipas defi-nitely had a positive effect on the Kemp’s Ridleypopulation (ibid.). Use of TEDs in shrimp trawlsand declining shrimping effort in the Gulf of Mex-ico increased survival of post-pelagic Kemp’sRidleys at sea (Table 21).

Kemp’s Ridley nesting has been reintroducedto PAIS (Shaver and Caillouet 2015), and thenesting colony there was also growing exponen-tially through 2009. Some of the wild nesters atPAIS could be offspring of head-started nesters,but others could have been produced by nestingsof wild Kemp’s Ridleys in Texas, Mexico, andelsewhere. No doubt head-starting had a positiveeffect by allowing tagging of yearlings so theycould be identified as part of Kemp’s Ridley rein-troduction to PAIS. However, it may be impossi-ble to determine whether head-starting by itself,outside its context as part of the reintroductionexperiment, had a positive effect on the popu-lation, since numbers of head-started individu-als in the population are declining. Nevertheless,head-starting and reintroduction were successfulin producing a considerable body of useful bio-

logical information about Kemp’s Ridleys thatotherwise would not have been produced.

Cost-effectiveness of reintroduction and head-start eventually became a consideration in theirevaluation (Woody 1990; Shaver and Fletcher1992; Taubes 1992a, 1992b; Wibbels 1992; Eck-ert et al. 1994). NPS et al. (1978 op. cit.) didnot call for collection and archival of cost dataor assessment of cost-effectiveness of any partof the KRREP. As far as we are aware, noneof the participating agencies imposed such re-quirements later on, to assure that data wereavailable for quantitative assessments of cost-effectiveness of the KRREP as a whole or in itsseparate parts. Neither of the two Kemp’s Ridleyrecovery plans (FWS and NMFS 1992; NMFSet al. 2011) called for cost-effectiveness assess-ments of any conservation effort applied to date.Wibbels et al. (1989 op. cit.) justified the costs ofhead-starting simply by invoking the worldwideinterest in possible effectiveness of head-startingas a conservation technique, and the unprece-dented opportunity afforded by the GalvestonLaboratory to address this issue. Cost per hatch-ling released at RN and cost per head-startedyearling released into the Gulf of Mexico werecompared by Woody (1991), but he did not pro-vide details regarding how his cost estimates werederived. Shaver and Fletcher (1992) noted thatcosts of incubation and imprinting at PAIS weresignificantly lower than those of head-starting.This was to be expected, if for no other reasonthan shorter duration of operations, simpler andsmaller facilities, and smaller sizes of eggs andhatchlings at PAIS, as compared to longer du-ration of operations, more complex and largerfacilities, and larger turtles associated with head-starting. A further complicating factor in eval-uating cost-effectiveness of head-start and rein-troduction is that actual costs included “in kindcontributions” made by a variety of agencies, or-ganizations, institutions, and industries, the val-ues of which were not determined. In addition,there is a question whether costs of research con-ducted in support of reintroduction and head-start

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should be included in their evaluation.Any future assessment of cost-effectiveness of

reintroduction and head-start should include com-parable assessments of cost-effectiveness of pro-tecting nesters, eggs, and hatchlings in Tamauli-pas, the requirement for TEDs in shrimp trawls,and any other conservation methods aimed atKemp’s Ridley recovery. Such comparable as-sessments have not been conducted to our knowl-edge, with one exception. During 2000–2002,the National Center for Ecological Analysis andSynthesis (NCEAS) conducted a study (NCEAS,Biggest Bang for the Buck: Really Melding De-mographic Theory with Economics. Availablefrom https://www.nceas.ucsb.edu/projects/3560[Accessed 16 August 2014]) that attempted to de-termine cost-effectiveness of various Kemp’s Rid-ley conservation methods including experimentalhead-start (see Caillouet 2005, 2006, 2009, 2010).Its results have not been published as far as we areaware. However, the fact that the study was con-ducted suggests that cost data were available andused. If in the future a cost-effectiveness study isconducted to compare Kemp’s Ridley conserva-tion methods, we believe it should include costsin relation to all life stages from egg throughreproductive adult, as recommended by Eckertet al. (1994); it should also include costs of de-mographic modeling (see Heppell and Crowder1994, 1998; Heppell et al. 1996, 2005, 2007;Crowder et al. 1998). Because research has beenassociated with all conservation methods appliedto date, the value of these research contributionsshould logically be integrated into any future cost-effectiveness analysis. However, considering thecomplexities of determining and tracking cost-effectiveness of conservation and research overtime, and the paucity of available cost data, we be-lieve an accurate evaluation of cost-effectivenessof Kemp’s Ridley conservation approaches is notpossible at this time.

Epilogue

Robert Whistler drafted NPS (1974 op. cit.),which contains the earliest recommendation toreintroduce Kemp’s Ridley to PAIS, and RolandWauer and other NPS officials approved it.Clyde Jones and Roland Wauer initiated furtherplanning in 1977 (Wauer 1999), beginning withHoward Campbell’s feasibility study (Campbell1977 op. cit.), and they established a boardof professional advisors (i.e., the SAB) whoprovided valuable input and oversight during theplanning. Reintroduction of Kemp’s Ridley toPAIS was the primary focus of early planning(NPS 1974 op. cit.; Campbell 1977 op. cit.;NPS et al. 1977 op. cit.; NPS et al. 1978 op.cit.; Wauer 1978). Roland Wauer and NPSplayed leadership and coordinating roles in theearly planning with input from participatingindividuals and agencies. Reintroduction,head-start, and protection and enhancement atRN were incorporated into the Action Plan (NPSet al. 1978 op. cit.) that was implemented in1978 as the KRREP. Wauer (1978) and Woody(1986) elucidated the original importance ofreintroduction of Kemp’s Ridley to PAIS, andWoody (1991) reinforced it:

“The original concept was to attempt to establisha second nesting population at Padre Island asinsurance against the possibility that the RanchoNuevo core group might be lost, as well as todevelop the techniques necessary to maintainthe species in captivity should all else fail. Theconcept made sense in 1978, and I supported theidea then.”

This concept still makes sense, especially in con-texts that a nesting colony now exists at PAIS,and pre-2010 expectations regarding populationrecovery (NMFS et al. 2011) have changed (Cail-louet 2010, 2011, 2014; Crowder and Heppell2011; Gallaway et al. 2013).

The KRREP went as planned for 11 y, dur-ing which many unanticipated problems were

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https://www.nceas.ucsb.edu/projects/3560


identified and attempts made to solve them. Forexample, the problem of male-biased sex-ratioswas solved in 1985, when sufficient data had ac-cumulated to determine pivotal incubation tem-perature; incubation temperatures were then con-trolled in ways that produced female-biased sexratios in the 1985–1988 year-classes imprintedat PAIS (Mrosovsky and Godfrey 2010; Cail-louet 2012a; Shaver and Caillouet 2015). InTamaulipas and elsewhere, incubation tempera-tures during the nesting season must also be gen-erally favorable to producing females, becausesex ratios in the Kemp’s Ridley population arefemale-biased (Coyne and Landry 2007; Wibbels2007; Mrosovsky and Godfrey 2010; Caillouet2012a). PAIS imprinting was terminated in 1989(Wibbels et al. 1989 op. cit.; Wibbels 1990), buthead-starting continued with 1989–2000 year-class hatchlings imprinted at RN. In 1993, thenumber of hatchlings received annually from RNwas reduced to about 200 or fewer; in 1994, theturtles from the 1993 year-class were used inTED testing and certification, and year-classesthereafter were used for research on tags and tag-ging until year-class 2000 was released. Searchesfor and documentation of head-started nestersand compilation of tag-returns continue to thisday, especially in Texas and mainly at PAIS. De-spite all changes and challenges associated withreintroduction and head-start, nesting of Kemp’sRidleys was reintroduced to PAIS.

Putman and Lohmann (2008) indicated thatmagnetic imprinting occurs in hatchlings on theirnatal beach, and that adults use this informa-tion to guide them sufficiently close to their na-tal beach so that additional local cues can helpthem find it. This implies that hatchlings alsosense, store, and use information about mag-netic fields they experience during dispersal topelagic habitats as hatchlings and during longand repetitive migrations to and from foraginggrounds as they survive and grow toward matu-rity. If similar imprinting occurred in hatchlingsthat were head-started, and if their captivity andlocations of release as yearlings did not interfere

with their ability to navigate and return as adultsto their natal (i.e., imprinting) beach, then sur-vivors imprinted at PAIS should have returned asadults to PAIS and those imprinted at RN shouldhave returned to RN. If head-starting altered themagnetic memory, survivors might be expectedto home as adults to Galveston beaches; head-started Kemp’s Ridleys have nested there (Shaverand Caillouet 2015). Head-started yearlings wereexposed to different magnetic fields at their re-lease sites and during dispersal from those sites.If, as Putman et al. (2010, 2013) suggest, sur-face circulation influences distribution and sur-vival of young Kemp’s Ridleys and where theyeventually return to nest, and if this applies aswell to head-started yearlings, they might be ex-pected to return to nest near their release sites.It is also possible that moving eggs and hatch-lings, head-starting, and releasing yearlings invarious locations disrupts natural, magnetic mem-ory mechanisms to the extent that most if not allsuch turtles are destined to unnatural, aimlesswanderings, thus reducing their chances of everreproducing. Yet, head-started Kemp’s Ridley fe-males found beaches, nested on them, and pro-duced viable eggs and hatchlings in Texas andMexico (Shaver and Caillouet 2015). Therefore,some head-started Kemp’s Ridleys survived tomaturity, found mates, copulated, nested, andtheir eggs produced viable hatchlings. Appar-ently, nesting at PAIS was sufficient to reestablisha nesting colony there. It remains to be deter-mined whether this colony will continue to growand become sustainable. It continues to be sup-ported by manipulative conservation methods, asare the nesting colonies in Mexico (NMFS etal. 2011). The combined influences that reducedthe probability of reintroducing Kemp’s Ridleynesting to PAIS were insufficient to prevent it,and this makes the reintroduction all the moreremarkable.

Clearly, the amounts of effort applied in search-ing for head-started and wild nesters on beachesin the U.S. and Mexico influence numbers ofnesters found and documented (Byles 1993;

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Williams 1993; Eckert et al. 1994; NMFS et al.2011; Shaver and Caillouet 2015). The numberof nesters examined also influences the chance offinding a head-started Kemp’s Ridley. The factthat relatively few head-started Kemp’s Ridleynestings have been documented on beaches any-where other than PAIS implies that reintroduc-tion worked, but it also could have been affectedby the high level of searching for nesters, bothhead-started and wild, at PAIS. More nestingsof head-started Kemp’s Ridleys may have oc-curred than have been documented anywhere.Had search and tag detection efforts aimed atdocumenting head-started Kemp’s Ridley nestersbeen equally intense in Tamaulipas and Texas,then a valid comparison could be made betweennestings of PAIS and RN imprint groups. If, onthe other hand, the location of release of head-started Kemp’s Ridleys played a dominant role inwhere they returned to nest, a completely differ-ent distribution of their nesting might be expected(Shaver and Caillouet 2015). No nestings by wildKemp’s Ridleys tagged with CWTs as hatchlingsat RN have been documented anywhere despitethe 43,885 released during 1996–2000 at RN.

In our opinion, the results of reintroduction arevery promising. The current level of nesting atPAIS greatly exceeds that documented histori-cally (Hildebrand 1963; NPS 1974 op. cit.), al-though nesting levels were poorly documentedbefore 1996 (Shaver and Caillouet 2015). Thegoal of reintroducing Kemp’s Ridleys to PAISwas achieved, but there are no guarantees thecolony will continue to increase and becomesustainable. Incidentally, the same is true fornesting colonies in Tamaulipas and elsewhereon Mexico’s coast of the Gulf of Mexico, andinterruption of the exponential increase after2009 is of great concern in this regard. The needfor a secondary nesting colony has not ended,and the Kemp’s Ridley population is still endan-gered (NMFS et al. 2011). Further efforts will beneeded to monitor, sustain, and enhance nestingcolonies in Tamaulipas and Texas until Kemp’sRidley is delisted (NMFS et al. 2011).

Head-starting contributed to reintroduction, be-cause it provided tagged turtles from year-classes1978–1988 that could be linked to reintroduction.It also increased survival during the first year oflife. However, reintroduction through transplan-tation of eggs and release of hatchlings mighthave worked had it been tried, but mass-taggingof hatchlings would have been required to pro-vide a means of assessing success of such anapproach.

One rationale given in NPS et al. (1978 op. cit.)for head-starting hatchlings from RN in 1978 wasthat it was expected to compensate for takingeggs for reintroduction. This could be interpretedas an expectation on the part of the planners thathead-starting would be an effective conservationor management tool in and of itself (Klima 1978;NPS et al. 1978 op. cit.). However, we wouldnot recommend it for this purpose. Head-startingof Kemp’s Ridley has not shown promise as amanagement tool outside its connection to rein-troduction. Perhaps head-starting worked to thedegree it did because the turtles were in theirpelagic phase, after which they were ready phys-iologically and behaviorally for a new diet anda benthic life style (David Owens pers. comm.).Most were reared 9–11 mo before release, and thepelagic phase has been assumed to last two years(TEWG 1998, 2000; Heppell et al. 2005, 2007;NMFS et al. 2011; Witherington et al. 2012).

Recommendations made by Eckert et al.(1994), that NMFS initiate a major effort tosearch for wild Kemp’s Ridleys CWT-tagged ashatchlings at RN, did not materialize. Although43,885 hatchlings were tagged with CWTs at RN,few tag returns have resulted, suggesting thatefforts aimed at detecting these turtles were inad-equate. Without the required effort to find theseCWT-tagged turtles as nesters in Tamaulipas orelsewhere, we cannot determine whether releaseof head-started Kemp’s Ridleys returned morenesters to RN than would have been returned hadthe hatchlings been released at RN, even thoughhead-started nesters have been documented nest-ing at RN, PAIS, and elsewhere. Another obvious

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question is why some head-started Kemp’s Rid-leys imprinted to RN have nested in Texas andelsewhere outside of RN (Shaver and Caillouet2015). It is possible that they nested in Tamauli-pas in greater numbers but were not examined fortags (external and internal) and therefore werenot documented.

The numbers of nesters (head-started and wild)at PAIS are significant and were increasingthrough 2009. Also important to consider is thatsome of the thousands of female hatchlings pro-duced by head-started nesters at PAIS likelysurvived, nested, and produced more offspring.Therefore, over time, reintroduction and head-start may have had, and may continue to have,multiplicative effects on the nesting colony atPAIS as well as nesting elsewhere (Caillouet2005). Although hatchlings produced by head-started nesters have not been tagged before re-lease, ongoing genetics work might be able tolink some of them to head-started nesters (Shaverand Caillouet 2015).

Continued head-starting of Kemp’s Ridleyswas recommended by Wibbels et al. (1989 op.cit.), Wibbels (1990), and Eckert et al. (1994).Unfortunately, the mass-tagging and releasesof hatchlings at RN, recommended by Eckertet al. (1994), produced few returns, none ofwhich were documented nesting at RN. However,mass-tagging of hatchlings with CWTs wasshown to be practicable. The lack of documentednestings at RN from these releases suggests thatlittle if any effort was expended toward theirdetection. However, if none survived to maturity,the result would be similar.

Acknowledgments.—We are grateful to all whocontributed to planning, funding, execution, over-sight, and evaluation of Kemp’s Ridley reintro-duction and head-start, as well as to research andother activities related to reintroduction and head-start. They are far too numerous to list here, butmost if not all are mentioned in the text and citedliterature of this review, along with their con-tributions. Clark Fontaine, Theodore Williams,

Benjamin Higgins, and others at the GalvestonLaboratory who were directly involved in rear-ing, tagging, and releasing turtles, and manag-ing tagging and tag-return data, deserve specialrecognition for their dedicated efforts under diffi-cult circumstances. Special thanks go to RolandWauer, Robert Whistler, Jack Woody, EdwardKlima, James McVey, Peter Pritchard, and RenéMárquez-M. for their contributions to early plan-ning, to their support of experimental head-startand reintroduction, and for their generous re-sponses to inquiries about their contributions. Weare also grateful for the dedication, influence,and advice of Henry Hildebrand, Archie Carr,and Peter Pritchard, who helped inspire the KR-REP. We thank Russell Burke, Roland Wauer,Carole Allen, Benjamin Higgins, David Owens,Nathan Putman, Kenneth Lohmann, KennethDodd, Pamela Plotkin, and Theresa Caillouetfor reviewing various versions of the manuscriptand offering supportive and helpful comments,corrections, and suggestions. This work couldnot have been accomplished without funding andpermitting by NPS, NMFS, FWS, TPWD, andINP (and its successor agencies; Burchfield andPeña 2013). CITES, FWS, and TPWD permit-ted importation of eggs and hatchlings to Texasfrom Mexico. FDNR permitted releases of head-started turtles in Florida waters. Eggs and hatch-lings were supplied by René Márquez-M. (INP),Peter Pritchard (Florida Audubon Society, Mait-land, Florida), Patrick Burchfield (Gladys PorterZoo, Brownsville, Texas), and their staffs.

We are especially indebted to Carole Allen,HEART, and the Piney Woods Wildlife Societyfor steadfast support of head-start, reintroduction,and the KRREP, and dedication to educating thepublic about the importance of conservation ofKemp’s Ridley and other sea turtles. We thankJoseph Flanagan, Richard Henderson, and ElliottJacobson for veterinarian services and collabo-ration. We thank Edward Klima, James McVey,Jack Woody, Milford Fletcher, Peter Pritchard,René Márquez-M., Patrick Burchfield, and otherswho provided oversight, guidance, and direction.

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NOAA, USCG, the University of Texas, TexasA & M University, TPWD, FDEP, and ARCOOil and Gas Company provided vessels for tur-tle releases. USCG and TPWD provided aerialtransport of eggs from RN to PAIS, or hatchlingsfrom RN to Galveston; on occasion, USCG pro-vided aerial transport of yearlings from Galvestonto airports near distant release sites. Importantdocuments, relevant literature, data or other in-formation were provided by Shauna Ertolacci,Roland Wauer, Benjamin Higgins, Jack Woody,Edward Klima, Peter Pritchard, René Márquez-M., Jaime Peña, James McVey, Mike Ray, ElliottJacobson, Sheryan Epperly, Nathan Putman, Ken-neth Lohmann, David Owens, Selina Heppell,Brian Bowen, Kenneth Dodd, Nicole Yin, Pe-ter Eliazar, Karen Bjorndal, Nicholas Mrosovsky,Matthew Godfrey, Anne Meylan, Joseph Flana-gan, Pamela Plotkin, Helen Lawrence, WilliamTydeman, Thane Wibbels, Blair Witherington,Kara Dodge, Ana Negrete P., Don Frels, KenLawrence, Bruce Bury, and Natalie Wiest. Thesenior author thanks his wife, Nancy Laird Cail-louet, for her support and patience over the yearsof preparation and revision of the manuscript. Hededicates his contribution to this paper to her andhis children, Suzanne R. Caillouet, Theresa E.Caillouet, Michelle M. C. Bailey, and C. Christo-pher Caillouet.

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Charles W. Caillouet, Jr. has a B.S. in Forestry(1959) and M.S. in Game (Wildlife) Management(1960) from Louisiana State University, and Ph.D.in Fishery Biology (1964; minors Statistics andPhysiology) from Iowa State University. His doctoralresearch focused on the relationship between forcedexercise and blood lactic acid in Channel Catfish(Ictalurus punctatus). During 1964–1967, while anAssistant Professor at University of SouthwesternLouisiana (now University of Louisiana Lafayette), heinvestigated abundance and distribution of postlarvalBrown Shrimp (Farfantepenaeus aztecus) andWhite Shrimp (Litopenaeus setiferus) in VermilionBay, Louisiana. During 1967–1972, as AssociateProfessor at Rosenstiel School of Marine andAtmospheric Science, University of Miami, he helpedestablish a commercial marine fisheries statisticscollection program for Puerto Rico’s Departmentof Agriculture, conducted statistical analyses andevaluation of sport fishery catch rates in EvergladesNational Park, and directed and conducted Sea Grantaquaculture research on Pink Shrimp (F. duorarum).He was to first to induce maturation in femalepink shrimp by eyestalk ablation. As SupervisoryFishery Biologist (Research) and Chief of variousdivisions and branches at the Galveston Laboratoryduring 1972–1998, he supervised and conductedresearch related to biology, population dynamics, andaquaculture of Penaeid shrimp, and head-startingKemp’s Ridley; he also managed environmentalimpact studies related to petroleum explorationand production, and Strategic Petroleum Reserve

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salt dome brine disposal. In October 1985, he andDr. André Landry chaired the First InternationalSymposium on Kemp’s Ridley Sea Turtle Biology,Conservation and Management, at Texas A&MUniversity Galveston. In 1988, Dr. Caillouet assistedin drafting IUCN guidelines for use by CITESin reviewing and evaluating sea turtle ranchingproposals. In November 1995, he and his staff

received a U.S. Department of Commerce BronzeMedal Award for superior federal service related tosea turtle research and related activities. He servedon the TEWG (1998), and peer reviewed the draft5-Yr Review for Kemp’s Ridley published in 2007.In 2004, he participated in Stakeholders meetingsassociated with development of NMFS et al. (2011),peer reviewed an early draft, and commented ona later draft during the public comment period.To date, Dr. Caillouet has authored or coauthored137 publications and reports. He is an EmeritusMember/Fellow of the American Institute of FisheryResearch Biologists, and a member of the IUCN/SCCMarine Turtle Specialist Group. Following retirementin June 1998, he remained engaged in Kemp’s Ridleyand Penaeid shrimp research as a volunteer. http://www.gulfbase.org/person/view.php?uid=ccaillouet.Photographed by Olan Mills.

Donna J. Shaver received her B.S. in WildlifeBiology (1981) from Cornell University, M.S. inBiology (1984) from Texas A&I University, andPh.D. (2000) in Zoology from Texas A&M University

in College Station, Texas. Her doctoral researchfocused on the distribution, residency, and seasonalmovements of the Green Sea Turtle in Texas. Dr.Shaver has worked with sea turtles since 1980. Sheworked with the National Park Service at PadreIsland National Seashore from 1980–1993. From1993–2003, she was Director of the U.S. GeologicalSurvey Padre Island Field Research Station. Since2003, she has been Chief of the Division of Sea TurtleScience and Recovery for the National Park Serviceat Padre Island National Seashore. Dr. Shaver is theTexas Coordinator of the U.S. Sea Turtle Strandingand Salvage Network. She is a member of the Kemp’sRidley Sea Turtle Recovery Team, KRWG, andIUCN Species Survival Committee Marine TurtleSpecialist Group. Dr. Shaver oversees a variety of seaturtle research and conservation projects conductedin Texas, collaborates with other researchers inthe U.S. and Mexico, and provides training andleadership to biologists and volunteers working withsea turtles in Texas and Mexico. Her largest andmost long-term effort has been the Kemp’s RidleySea Turtle Restoration and Enhancement Project atPadre Island National Seashore. She has conductedresearch on unhatched Kemp’s Ridley eggs, sex ratiosand incubation temperatures for Kemp’s Ridley, seaturtle nesting and strandings in Texas, Kemp’s Ridleyforaging ecology, and movements and habitat useby adult Kemp’s Ridley and juvenile Green turtles.Her work has been recognized with awards fromseveral organizations including most recently the U.S.Fish and Wildlife Service 2013 Endangered SpeciesRecovery Champion Award for Agency Partner, andHarte’s Heroes 2014 Legends of the Gulf Award. Todate, she has authored or co-authored more than 110publications and reports dealing with sea turtles. Shehas been interviewed by numerous media outlets,was featured as ABC World News Tonight’s Personof the Week on July 29, 2005, and was namedthe Corpus Christi Caller Times Newspaper 2011Newsmaker of the Year on January 1, 2012. http://www.gulfbase.org/person/view.php?uid=dshaver.Photographed by National Park Service.

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Andre M. Landry, Jr. holds a Ph.D. degree inWildlife and Fisheries Sciences from Texas A&MUniversity and has been involved in sea turtle researchsince 1984. He served as Director of the Sea Turtleand Fisheries Ecology Research Laboratory at TexasA&M University where he taught sea turtle coursesand conducted sea turtle research. Dr. Landry’s initialsea turtle research efforts began in 1984 by first col-laborating with the NMFS Galveston Laboratory inresearch related to Kemp’s Ridley head-start, thenshifting in 1985 to documenting sea turtle strand-ing dynamics along the Louisiana and Texas coasts.He developed a standardized stranding survey pro-tocol carried out by his graduate students workingas NMFS surveyors. Landry and his students necrop-sied over 250 stranded sea turtle carcasses to docu-ment possible cause of death and generate natural his-tory data on species composition, length-frequency,sex ratios, reproductive condition, and food habits.Landry’s work on stranding dynamics enabled fed-eral contracts to assess mortalities in sea turtles un-related to shrimping and characterize toxicology ofsea turtles comprising a mass stranding event alongthe upper Texas coast in July 1991. His involvementwith head-starting Kemp’s Ridley sea turtles at theNMFS Galveston Laboratory assessed the effect ofdifferent feeding regimens on growth of head-startedhatchling and juvenile Kemp’s Ridleys and the effectof exercise on swimming performance and staminaof captive-reared Kemp’s Ridleys. Dr. Landry’s live-animal work involved numerous state, federal andprivate grants/contracts to document various aspectsof natural history of sea turtles and anthropogenic im-pacts on these organisms through in-water capture andnesting-beach patrol operations in the northwestern

Gulf of Mexico. As a result of this research, Landry’scapture of 1,026 sea turtles (including 660 Kemp’sRidleys) from waters of Texas and Louisiana facili-tated (1) an in-water Kemp’s Ridley monitoring pro-gram at Sabine Pass during 1992–2002, (2) the largestprogram of directed in-water capture of sea turtles(80% being Kemp’s Ridley) in the Gulf of Mexicofor research and stock assessment purposes, (3) re-lease of 972 tagged (with flipper tags and PITs) indi-viduals available for subsequent recapture and study,and (4) development of a multidisciplinary facultyteam whose access to these in-water capture and re-capture data produced new findings, research initia-tives and biotechnology critical to sea turtle recovery.Landry’s patrols of upper Texas coast beaches dur-ing 2007-2012 provided invaluable data on Kemp’sRidley nesting dynamics and migration behaviors.The aforementioned activities afforded Landry, his re-search collaborators, 30 graduate students, dozens ofundergraduate interns and hundreds of citizen volun-teers unprecedented opportunities to describe Kemp’sRidley population dynamics based on abundance,size composition, ecology, sex ratio, physiology, re-production, nesting activity, migration, food habits,health, pollutant bioaccumulation, and fate of wildand head-started conspecifics. Landry was a memberof NMFS’ TED Certification Advisory Panel, anda regular contributor to the Kemp’s Ridley WorkingGroup. In 2004, he participated in Stakeholders meet-ings associated with development of the second re-vision of the bi-national Kemp’s Ridley RecoveryPlan (NMFS et al. 2011) and provided commentsand input to the Kemp’s Ridley Recovery Team. Dr.Landry retired in August 2012. http://www.gulfbase.org/person/view.php?uid=landry (Photographed byLee Ann Day Landry)

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