FAO Fisheries Synopsis No. 85 FIRM/S85 (Distribution restricted) SAST - Green turtle - 5,31 (07) 005,02

SYNOPSIS OF BIOLOGICAL DATA ON IHE GREEN TUTLE Chelonia mydas (Linnaeus) 1758

Prepared by ftF. Hirth

FOOD AND AGRICULTURE OGANIZATION OF THE UNITED NATIONS ROME, 1971 DOCUMENTS OF THE FISHERY DOCUMENTS DE LA DIVISION DES DOCUMENTOS DE LA DIRECCION RESOURCES DIVISIONOF FAO RESSOURCES HALIEUTIQUES DU DE RECURSOS PESQUEROS, DEL DEPARTMENT OF FISHERIES DÉPARTEMENT DES PÊCHES DE DEPARTAMENTO DE PESCA DE LA LA FAO FAO

Documents whicharenot official Des documents qui ne figurent pas Esta Subdirección publica varias series FAO publications are issued in several parmi les publications officielles de la de documentos que no pueden conside- series.They are given a restricted distri- FAO sont publiés dans diverses séries. Ils rarse como publicaciones oficiales de la bution and this fact should be indicated font seulement l'objet d'une distribution FAO. Todos ellostienendistribución ifthey are cited.Most of them are restreinte, aussi convient-il de le préciser limitada, circunstancia que debe indicarse prepared as working papers for meetings, lorsque ces documents sont cités, Il s'agit en el caso de ser citados. La mayoría de or are summaries of information for use le plus souvent de documents de travail los títulos que figuran en dichas series of member governments, organizations, préparés pour des réunions, ou de ré- son documentos de trabajo preparados and specialists concerned. sumés d'informationàl'intentiondes para reuniones o resúmenes de informa- gouvernements des pays membres, ainsi ción destinados a los estados miembros, que des organisations et spécialistes inté- organizacionesy especialistas inte- ressés.Ces séries sontles suivantes: resados.

FAO Fisheries Report FIR IS (No.) FAO Fisheries Circular FIR ¡C (No.) FAO Fisheries Synopsis FIR IS (No.)

Special groups of synopses are iden- Des catégories spéciales de synopses Grupos especiales de sinopsis se dis- tified by symbols followed by classi- sontidentifiéesà l'aide de symboles tinguen con las siglas siguientes, seguidas fication numbers based on indexed code suivis des chiffres de classification basés por números de clasificación que se basan of "ASFA": sur le code d'indexation de la « ASFA»: en las claves de los indices de la «ASFA»:

SAST Data concerning certain species SAST Données sur certaines espèces et SAST Datos relativos a ciertas especies and fish stocks. populations de poissons, y poblaciones. MAST Information on methods and sub- MAST Renseignements sur des méthodes MAST Sinopsis sobre métodos y mate- jects. et des sujets. rias. OT Oceanographic data. OT Données océanographiques. OT Sinopsis sobre oceanografía. IT Limnological data. IT Données limnologiques. IT Sinopsis sobre limnología. and et y CART Information concerning fisheries CART Renseignements sur les pêcheries CART Información sobre los recursos and resources of certain countries et les ressources de certains pays acuáticos vivos de algunos países and regions (FID/S). et régions (FID/S). y regiones (FID/S).

FAO Fisheries Technical Paper FIRT/T (No.)

Special groups of TechnicalPapers Des catégories spéciales de documents Grupos especiales de documentos téc- are identified by: techniques sont identifiées à l'aide des nicos se identifican por las siglas siguien- symboles suivants: tes:

RE Indexed lists of experts and insti- RE Listes indexées d'experts et insti- RE Listas índices de expertos y de tutionsdrawnfromRegisters tutions tirées des registres tenus instituciones tomadas de los re- maintained by the Fishery Re- à jour par la Division des ressour- gistros que se llevan en la Direc- sources Division. ces halieutiques. ción de Recursos Pesqueros. CB Lists of periodicals, special sec- CB Listes de périodiques, des sections CB Listas de periódicos, secciones es- tions of "Aquatic Sciences and spéciales de la «Aquatic Sciences peciales de la « Aquatic Sciences Fisheries Abstracts (ASFA), and Fisheries Abstracts (ASFA) », and Fisheries Abstracts (AS FA) », specialbibliographiesandpa- des bibliographies particulières et bibliografías especiales y trabajos pers concerning documentation des articles sur les problèmes de relativos alosproblemas de problems. documentation. documentación. MFS Provisionaleditionsof FAO MFS Editions provisoires des « Manuels MFS Edicionesprovisionales delos Manuals inFisheries Science." FAO de science halieutique ». « Manuales de la FAO de Ciencias Pesqueras a.

Some documents also have another Certainsdocumentsportentparfois Algunos documentos tienen también identification, if, for example, they have d'autresnumérosd'identification,par otra identificaciónsi, por ejemplo, son been contributed to a meeting for which exemple s'ils ont été préparés pour une contribucionesa unareunión cuyos papers have been numbered according réuniondontles documents ont été documentos hansidomarcados con to another system, marqués àl'aide d'un autre système. arreglo a otros sistemas. FAO FttherioSynopir4 No85 FThM/585 jTJtBtribition retr1ctd) -J m'L1 5iit(O])OO5!P2

SYNOPSIS OF BIOLOGI(AL DATA ON PHIS GREEN IUBThE

Chelonia myda (Linnacue)1758

'repared by

Harold F. Hirth Department of Biology University of Utab Salt Lake City, Utah U.S,A.

FOOD AND AORICUTIIURE OHOANIZATION OF THE UNITED NATIONS Rome, December 1971 PREPPRATION 0F THIS SO?SIS

The present document which is the first FAO Species Synopsis on biological data of oommeroially important marino turtles, has bes prepared following the annotated outline for speoles synopais of data on species and stocks (FAO Fisheries Synopsis No.1 Revision 1, 1965), modified by the author to meet the special requirements of a synoptic review of turtle species.

The main purpose of this synopsis is to bring together the current information on the natural history of the green turtle, Chelonia nydas, and to draw attention to the major gaps in our knowledge of the species. In a few sections some of the older, classical .papera are cited end summarized in order to make the synopsis more comprehensive.

The author is Indebted to Mrs. Joan Hubbard, Interlibrary Loan Librarian and Mrs. Jearme Chapin, Life Science Librarian, University of Utah, for their help in tracking down rare manuscripts; and to Dr. John Hendrickson (University of Arizona) for allowing the author to peruse his personal library. Mrs. ICennie Harris, MrsBess Short and Mise Liwana Blau graciously and conscientiously typed several versions of the manuscript. The University of Utah Library and the Department of Biology financially aided the work through the purchase of inierlibrar'y loans.

Dr. Archie Carr (University of Florida) and Dr. H. Robert Bustard (The Australian National University) offered helpful suggestions after critically reading the manuscript and to them the author is indebted,

Distribution Bibliographic Hntry

FAO Department of Fisheries Rirth, H,F, (1971) FAO Regional Fisheries Offioors FAO Fish.Synop., (85):pag.var. Regional Fisheries Councils and Synopsis of biological date, on the Commissions green turtle Chelonia mydas (Linnaeus)1758 Selector Sï Author Nomenclature, taxonomy, morphology, biometrios, tagging, distribution - horizontal, hybrids, ecology, life history, nutrition, growth, behaviour, population dynamios, exploitation, fishery, legislation, habitht improvement, culture. Gron kitrile

G O N T E N T S

Pye No,

IBENTITY 1:1

1.1Nomenclatu.re I

1.11Valid nane i 1.12Objective aynonyTny I

1.2Thxononw i 1.21Affinities i 1.22Taxonomie status, and definition of size categories I 1.23Subspecies 1 1 .24Corsnon naInes 4

1.3)brpholoy 4

1.31Anatomy 4 1.32Cy-tomorphology 6 1.33Protein specificity 6

1.4Measuring and ta1ng turtlo 6

2 DISTRIBUTION 2:1 2,1Total area. i

2.2Differential distribution I

2.21Hatchlings, juveniles and sub-adults I 2.22Adulte 1

2.3Determinants of distribution chenes 8 2.4Hybridization 8

3 BIONO?&tCS A1 LIFE HISTORY 3: 1 3.1Reproduction 311Sexuality i 3.12Maturity 1 3.13Mating 1 3.14Fertilization 4 3.15Gonads 4 3.16Reproductive cycle, nesting behaviour and selection of nesting beach 4 3.17Eggs 3.2Embryonic and hatchiing phase 8 3.21Embryonic phase 8 3.22Hatehling period (behaviour, predators and survivorship) 9

3.3Juvenile9 sub-adult and adult phase 14 3.31 Longevity 14 3.32Hardineen 14 3,33Competitors 14 3.34Predrttort; 14 3.5 Paradies, 1ineasee, injures and abnormalities 14 IRM/S8'Green turtle Page No. 3.4Nutrition mid growth 3:14

3.41 Feeding and behaviour on the feeding pastures 14 3.42Food nnd feeding associates 15 3.43Growth rate 17 3.44Metabolism 20

3.5Posi-hatchiing movements and basking behaviour 21

3.51Developmental migrations 21 3.52 Dispersal and remigrations 21 3.53Navigation and orientation 23 3.54Basking 25 4 POPtJLA!FION 4:1 4.1Structure

4.11 Sex ratio 1 4.12Size composition i 4.2Abundance and density (of population)

4.21Average abundance i 4.22Changes in abundance I

4.3Natality and recruitment i 4.31Reproduction rates i 4.32Factors affecting reproduction 4.33Recruitment i 4.4 Mrtality and morbidity

4.41Mortality rates i 4.42Factors causing or affecting mortality 1 4.43Factors affecting morbidity 2 4.44Relation of morbidity to mortality rates*

4.5Dynoiijcs of populations (as a whole) 2

4.6The population in the community and ecosystem 2

5 XXPLOITATION 5:1

5.1Fishing equipment 1

5.11Gears and methods 1 5.12Boats 1 5.2Fishing areas

5.21General geographic distribution 1 503Fishing seasons 5.31General pattern of seasons 5.32 Dates of beginning, peak arid end of seanona 1 5.33Variation in date or duration of season 3

5.4Fishing operations and results 3

no information wae available to the author, this item has been omitted from the text FIRI4/S85 Green turtle iii

Pajc No.

PRYECTION AND MNAGEME}7P 6:1

6.1 Regulatoiy (legislative) mea8ure 1

6.11 Limitation or reduction of toti catch 1 6.12 Protection of portions of' populatione 6.13 Current status of the resource and tction progremme 2 6.14 Public education 2

6.2Control or alteration of physical features of the environment 2

6.3Control or alteration of chemical features of the environment 3

6.4Control or alteration of biological features of the environment 3

6.5Artificial stocking 3

7 Pu1FLE MABICUI/PURE 71

8 EFEIENGES 8 * I

FIR)/S83 Green tunjo

IDENTITY upiard-cu.rvthg marginais and mnali scales in the middle of -the front flippers.Further diffe- 11Nomenclature rences between the two species are pointed out by Williams, Grandison and Carr (1967), Bus-tard 1.11Valid naine (1969b), Bustard. and Liinpus (1969), and Cogger and Lindner (1969). Cholonia ntlas (Linnaous) 1758 1.22Taxonomie status and. definition 1.12Objective 1nonynr of size categories Testudo Linnaeus, 1758, p. 197, The green turtle is a morpho-species. Syst. Nat. (Ed. io) Stockholm.Type Because size categories of marine turtles are not locality, Ascension Island.. standardized, the four categories used in this report are defined, na follows: haichlingstill Chelonia m.yd.as Brongniart, 1800, Bull, Sci. bearing conspicuous umbilical scar; first few Soc. Philom., 2:89 weeks of post-hatching life; juvenile . umbilical soar inconspicuous or absent and carapace length Some authors consider the generic up to 40 cm (41) cm is ábout one-half the desiiation by Brongniar-t a nomen nudun.For carapace length of a reproductively mature example, Deraniyagala (1939) recoiizes the irid.iwidual); sub-adult post-juvenile but not following: yet reproductively mature (carapace length 40 -to 80 cm); adult reproductively mature; Chelcnia rnydas Latreilie, 1802, Hist. Nat. carapace length greater thau 80 cm (the minimal Rept, i, p. 22 sizealwhich females lay eggs is about 80 orn; for convenience it is assumed that males mature 1.2Taxono at about the sane time).Carapace lengths are straiit-line distances.A "yearling" is a one- 1.21Affinities year-old turtle. - Suprageneric 1.23Subspecies Kingdom Anirnalia It is best to use -the binomial, jjnia Subkingdom Metazoa mydas, for all green turtles until a detailed Phylum Chordata tazonornic study is made.However, several sub- Subphylum Vert ebrata species have been described, and no doubt othór Superclass Tetrapoda races will be named in the future.In fact, Class Reptilia the mydas complex may be one circuinglobal Subclass Anapsida "Rassenkreia" bu-t with significant gaps between Order Teatudina-ba the eastern Pacific and western Atian-tio-- Suborder Cryptodira Caribbean populations and between the East Superfamily Cholonioidea African and West African populations.Sub- Family Cheloniidae specific names have bean based largely on coloration, al-thout Smii;h and Taylor (1950) Generic state that a common arrangement restricts ndas to -the Atlantic andassizii to the Chelonia Brongeiart, Bull. Sci. Soc. Pacific Ocean either as species or subspecies. Philom. Vol. 2, 1800, p. 89.Genotype:Testudo At least sorne of -the subspecific terminolor nwdas Linnaeus. has been perpetuated in checklists anduide- books by Schinidt (1953), Conant et ai. Ç1956), The following generic identity is given by Conant (1958), Fukada (1965), S-bobbins (1966) Car (1952): "One pair of prefrontal scales; and Keiner and Wilson (1969), among others. tomium of lower jaw coarsely toothed, that of the upper jaw with strong ridges on its inner The Atlantic green turtle, Chelonia mydas surface; lateral laminas, 4."Longer descriptions ndas (Linnaeus) is found in the Atlantic Ocean, can be found in Deraniyagala (1939),aith sind Caribbean Sea, Gulf of Mecicond the Hedi- Taylor (1950) and Loveridge and. Williams (1957). terranean Sea,It has a predominantly brownish carapace, sometimes with olive or dark brown Specific blotches and streaks, and -the dorsal surfaces of the head., flìppera and tall are also Chelonia mydas is easily differentiated. predominantly brownish.The spaces between from its oongener, the flatback turtle, the scales on the -top and sidec of the head depressa Garmazi, by its more highly arched are yellowish; the plastron is usually whitish. carapace, sloping marginais and large scales in The shell of the malo is more elongate -than that the middle of the front flippers (Fig. i).In of the fornaio.A detailed. desoriptien of the contrast, C. depressa has a depressed carapaces subspecies cari be foundinCarr (1952). i2 FIRM/585 Green- urt1e

I F1R14J385 Green turtle Compared to the Caribbean population, adult region, notably Deraniyagala (1939), Suvatti females in the Ascension Island population have a (1950), Harrisson (1950-1969), and Hendrickson more angular emargination of the shell over the (1958) have also deferred from identifying the neck (Carr and Hirih, 1962), subspecies. The East Pacific green turtle, Chelonia la Taylor (1920) describes Chelonia mydas aassizii Bocourt, is found, along the Pacific ,iaponica from the Philippines as follows: coasts of Central and $outb America.The carapace above rusty reddish brown, each shield streaked. is basically greenish or olive-brown but sometimes with amber; head shields distinctly reddish, strongly flecked. with black, and. generally more each edged. with black; shields on sides of heed highly arched and narrower than that of an dark, with yellow along sutures; shields ou le Atlantic green turtle.The carapace is sometimes with black centres; plastron yellow,Taylor markedly indented above the hind flippers-this is (1970) briefly describea aduli japonicafrom usually lacking in individuals from the Atlantic Thailand as greenish to greyish brown, some (Carr, 1952).Much earlier, Baur (1890) stated individuals with dark rays; and. the plastron "there is no doubtwhatever that the Chelonia yellow.He goes on to say that the young are from the Pacific is a different species from the also olive to dark brown with some yellow Atlantic; the skulls are the same, but there are markings on the limbs; venter yellowish; and great differences in the form of the carapace". dark areas on flippers. One of the most easily recognizable forms in There are probably several races scattered the entire mydas complex is the Northeastern throughout the Pacific Ocean and the Indian Pacific gr en turtle, Chelonia mydacarrinegra Ocean and as already stated, the situation Caldwoll,This subspecies is characterized by warrants a careful systematic stuL3.y.The books black pigmentation on both dorsal and ventral by Deraniyagala (1953), }uroda et al. (1958), surfaces.The carapace is highly arched in large and A.non (no date) include colour illustrations females while it is low in large males.It occurs of green turtles from the Indo-Pacif io region. in large numbers throughout the Gulf of California A coloúred picture of the venter of a green and on the outer coast of Baja California. turtle from Indefatigable Island, Galpagoe, Cald.well (1962a) believes that all references to appears in the article by Johnson and Johnson green turtles off California and northward along (1959), the coast should be considered as indicating earrinegra and he gives a detailed account of this According to Schmidt (1945), becatLee øf subspecies.However, the problem of distinguishinghistorical zoogoograpby, one would expect the this subspecies from black agassizii along the marine turtle populations on the Pacific coast- Mexican and Guatemalan coasts still remains to be of the Americas to be more closely related. to solved.If this cannot be done, the proper name those on the Atlantic and, Caribbean coasts than for the black turtle will be Chelonia nylas to. iioe in the western Pacific.On the other agassizii (Hirth and Carr, 1970). hand, evidence is accumulating (see colour descriptions in preceding paragraphs and below) According to local inhabitants, two fair:Ly which suggests just the opposite. well marked stocks of Chelonia (or a variably coloured sinlepopulatïon) occur in the Galpagos The following accounts of green turtles in area (Carr, 1964).The turtle situation here the South Pacific (Hirth, MS) aro given as a warrants immediate study. start toward a descriptive classification of all turtles in the Pacific region (although it should Carr (1964) has noted some carrinegra be noted that there may be significant colour characteristics in sorno turtles in the Hawaiian variations among individuals of the sane pops-- Archipelage.The concurrent distribution of lation and certainly between different sizes), carrinejrar-liice and mydas-like forms in the In French Polynesia, a juvenile green turtle Central Pacific may indicato a tendency toward with a carapace length of 30.5 cui has a carapace polymorphism within a single population (Hirth and predominantly copper-brown with grey-black rays; Carr, 1970), the top of the head is black with well-defined sutures and the sides of the head are blackish- Chelonia nwìas ,japonica (Thunberg) is the namebrown; the flippers are black above and. white given to the Western Pacific green turtle (seo Carr,below but with the distal half below mottled 1942 for discussion).The type locality is Japan. yellow-black-brown,The throat and plastron aro Wex'muth and. Mertens (1961) give the range as white. Pacific and Indian Oceans.Okada (1938) lists japonica from Horisyn, Sikoku, ICyusya, Bonin A sub-adult female, carapace length of Islands, Okinawa and Taiwan. The Hawaiian Islands 68.5 cm, from French Polynesia, has a carapace probably represent the eaaternmost outpost of this streaked with copper, yellow, black and. bro:m: subspecies (Oliver and Shaw, 1953).Honegger the top of the head has a ground colour of black, (1967) and Gaymer (1968), but not Hirth and Carr but most scales have distinct copper spots; the (1970), use thia trinomial for the Seychelles sides of the head are blac)çish; the dorsuin of population,Other workers in the Indo-Pacific front flippers are black but with distal one- 1:4 FIRM 28Green turtle third edged with copper: the dorsum of hind (Hirth and Carr, 1970) is mottled with yellow, flippers are predominantly black; venter of all brown, black and green.The top of the head is flippers are yellowish with black ori distal third;spread with black, groen and brown spots, and yellowish-white plastron. there are well-marked yellowish seams between the scales.Adult males are similar to females Examination of ten adults(eight males; two with the exception that males have less yellow females) from Scilly, French Polynesia showedno and brown mottling on the carapace and top of significant colourdifferences between the sexes. the head.Whether these pigmontation differences Nine had marked indentations above the hind flippers, between the sexes are consistent throughout this All had post-ocular counts of 4/4. A population is unknown.Deraniyagala (1939) noted typical turtle with a carapace length of 100 cm some colour differences between the sexes around has a greenish brown carapace mottled with white, Ceylon.In general, the pigmentation of the yellow, olive and black; the top of the head is South Yemeni green turtle population is similar to the carapace in colourations the throat and plastron are white. consistent with Deraniyagala's (1939) descriptions of Ceylonese turtles of the same sizes. He describes eight pigmentation phases of the green A sub-adult green turtle (carapace length 45 turtle in which the carapace passes from clark cm) from Western Samoa hasa brown-green carapace greenish-bronze in the hatchiing to a brownish- streaked with black; top of head is brawn and, red phase, then a variegated phase, and ultimate).y sidos of head black; foui' flippers aboveare -tq the olive tint with black and, brown streaks black with some brown interspersed end beloware caarac-teristic of adults.Boulenger (1890) variegated yellow-black-brown; the plastron is describes young green turtles from Indian Ocean white. area as "dark brown or olive above, the limbs margined with yellow; yellow beneath, with a An adilt female (carapace length 96.5 cm) large dark brown spot on the hand and foot, from Neste.'n Samoa has a dark brown carapace Carapace of adult olive or brown, spotted or abundantly covered with li.t green, light brown marbled with yellowish". and yellow spots and, blotches; top of head is spotted bxown and green; sides of head basically Hoofien and Yaron (1964) describe the yellow with large black spots in middle of each carapaces of two adult turtles from the Dahiac scale; top of flippers spotted black-brown-green Archipelago as uniform ochraceous brown,They while flippers underneath are yellowish-white also mention the lack of distinct marginal with few black spots in middle of each flipper; indentations above the hind limbs. chin and plastron are yellowish. Common names An adult female (carapace length 101 cm) 1.24 from Tonga has a light green carapace heavily Some common names of the green turtle in mottled with black, dark brown and light brown; varioue areas of the world are given in Table f. top of head is basically greenish-black with The name "green turtle" is derived from the colour blotches of yellow and li.t brown; sides of the of i-'e fat. head are basically yellow but with black spots in scales; flippers above are dark copper-brown with The Raroians (Dariielsson, 1956) have names black spots and below are whitish-yellow witha for the various stages in the ontogeny of sea few black centredscales: the plastron is turtles: egg (tuo vaki), hatchiing (karëa), yellod eh. three classes of juveniles (torara, kpue, mkui), adult (tYfai). Sub-adult turtles (carapace lengths 58.4 and 60 cm) from the Fiji Islands and New Caledonia Sorno Tonga fishermen also have nanee for are very similar in thai the carapace is basicallythe colour phases and various sizes and sex. brownish-green with distinct copper, yellow arid green streaks; top of head is black with large copper spots and sides of head are blackish, 1.3Morphology fading into white near the lower jaw; flippers above are blackish with copper on forward edge 1.31Anatomy and tips; venter of flippers are white-yellow with some black scales on trailing edge; throat An annotated bibliography of some aspects and plastron are whitish-yellow. of the morphology of marine turtles is found in Ingle and Smith (1949).More recently, Bellairs The green turtle isanoccasional visitor in (1970) has written about the general anatomy and New Zealand and adults have been described by physiology of the groen turtle.An early account McCann (1966'o) as being olive or brown, the of the anatomy of Chelonia was given by Hoffhrimn shields with more or less distinct brown rays; (1890).Among the classical works on anatomy and, yellow beneath. are those of Pilliet and Bignon (1885) on the lacrimal gland; Keateven (1911) on cranial Basically, the carapace of a mature female from the Gulf of Men and the Seychelles Islands FIRN/585 Groen turtle

TABLE I

Common names of the green turtle. The geographical area or language ia followed by the vernacular.

West orn Hnisphere Eastern Heniaphere

North America English green turtle, green sea turtle, green turtle, edible turtle edible turtle Gerran Danish West Indies, Trinidad, Tobago SuppenschildkrSt e green turtle, edible turtle, french greenback tortue franche, tortue de mer, Caribbean Spanish tortue verte tortuga, tortuga blanca, Portuguese tortuga verde -t art aruga, t art aruga-do-mar, MEi :ico tartaruga vendo la panama, caguama, ceguama OO4flufl, caguma negra, tortuga prieta, Ghana caguama prieta, mestiza, anjwaapuhulu, apuhuru, hala tortuga negra, sacacillo Swahili kas sa Guyana bettia Mozambique Suninam i-tat aruca, riruvi, risa, pat eri, krape casa, icaha Tupi in Brazil Bajuxil of Lamu muruaxid taza Hawaiian and. Tahi-tian Southern Yemen honu bissa, biasat al bahr, humsa Western Samoa Creol. on Aldabra Atoll laumei torLie do mer Tonga Islands Maldive Islands forni tu'a'uli, vol a form tuaku1a, Sjnhal eso form tuapolata, gal ksbNva, mas ksbMra, -tuai fonu vail ksbva Fiji IslaMs Tamil vomi damn, parrznai pl mai vonu loa, Malay on Cocos Islands mako loa, penyu ika damm Malaysia pum, sisek, penyx, penYu agar, penyu pulau Borneo pinu Philippine Islands pudno, tuntuga Chinese lu -tau-hai Japanese ao umigamo 1:6 FIRM/85 Oreen turtle morphologr; Nick (1913) on the development of the (Carr, 196Th).Notching marginal soutes is the skull; Ruches (1929a, b) on osteologr; Deraniyagala most common method used today by turtle blob-. (1939) on general ontogeny and. scalation; Villiers giste, but the long-term success of this method (1958) ori general anatomy; and. Parsons (1959) on has yet to be established.Identifying the nasal cavities.A study of respiratory muscles individuals through -immunological reactions has in nydae has been rende by Shah (1962).The total been suggested by Hendrickson (1969).Hughes lack of the lung muscle, surmoularie ritriatum (1970) in experimontm with loggerhead (Caretta pulmonale, is notable.Mlynarski (1961) has carotta) hatchlingm is implanting a piece of described the structure of soutes arid. Brongeraina mtainlesa steel wire under selected carapace (1968b) described some variations in vertebral scales in hopes of identifying individuals soutes ot' Surinais turtles. later-recaptured by x-rsying them (seo also Anon, 1969k).Pough (1970) has recently Parsons (1958) examined the choanal papillas suggested a method of marking hatchiinge asing of green turtles but was unable to give it a a plastic plug. function.Later, Smith (1961) proposed that the ohoanal rakers in adult green turtles are correlated with its herbivorous diet. 1.32Cytomorpholor According toIaicirio (1952) the female green turtle has ono lesa chromosome than the male. Plorking with Chelonia ,japonica from the Bonda Islands, he found, through histological examination, that the diploid number was 56 and. 55 in the males and females respectively. Characteristics of thehaemnoglobìnof C. nWtles have been studied by Dozy et al. (1964). Stephenson (1966) relates how the cells of mayùas have been used in cell culture studies, 1.33Protein specificity The serum protein concentration in Ohelonia re. mydais 1.8% (Frair, 1964).In most turtles it is between 2 and 6%.Serological studies suggest a close relationship among the five genera Fig. 2Measuring the plastron length of a of sea turtles (Frair, 1969). South Yemeni green turtle.The use of "giant" calipers gives rtraight-line 1.4Meanurin' and tagging turtles measurements.The mean on the right holds a pair of tagging pliars The etsndax'd measurements of sea turtles (all of which are straight-line distances) are length and width of carapace, length of plastron and width of head (see Fig. 2).The lengths of plastron and carapace are anteriormnost to posterloneost extensionm of each.The width of the carapace is the distance between itlateral margins at the widest point.The head measurement is taken at the groateot bony width. The moat common method of tagging marine turtles employs the use of a stainless steel tag clamped onto the trailing edge of a front flipper (see Fig. 3),The tags are numbered on one face and on the other side are inscribed instructions for the finder in various languages. Adult females are usually the only ones tagged because they can be easily caught on the nesting beachOne of the problems in sea turtle biolo im to invent a tag A tagc'd Wr.tern Srnnan p-rr'n tcrtl. that can be placed on or in a hatchling and uhich The stainless eteel. tags, numb"red. ant will identify i-t some years later,Attempts -to inscribed in various ianguaven lave mark or tag hatchlings by branding, -tattooing, and been adopted by marine turtle hioioçtnte use of reageotic tags have not been successful throughout the world FIRM/585 Green turtle 2,1 2 DISThIBiPPION for food; construction of dwellings, roads) have eliminated soma good nesting sites and, hence the 2.1Total area distribution of green turtles has been reduced (see section 5). Green turtles are found throughout the tropical seas end as stragglers in a far more 2.2Differential distribution extensive area.The major breeding and feeding areas, and hence the majority of green turtles, 2.21Hatchlings, juveniles and sub- are found between the northern and southern 200G adults isotherme (i.e., average temperature of surfac' water in coldest month la above 200G). It is postulated that the hatoblings leaving Ascension Island drift downstream (i.e., toward On the American side of the Atlantic Ocean, Brazil) wi-tb the South Equatorial Current (see green turtles have been captured or sighted as section 3.53), far north as New England (Lamson,1935;Pope, 1939; Dexter, 1947; Bleakney, 1965).Carr (1952) There is a dearth of information on 'the sars that individuals have been reported seen in bionomica of hatchliags and juveniles.In fact, Newfoundland waters but no specimens have been the first year of life of all marine turtles has taken.The southern locality record of the green been popularly labelled "the lost year" by marine turtle in the western Atlantic is 1ar dei Plata turtle biologists (see Brongersma,1970). and Necochea, Argentina (Barran and Freiberg, 1951; Freiberg, 1967). Developmental migrations of juveniles and sub-adults are discussed in section 3.51. 0m t1e other aide of the Atlantic, green turtles h.ve been recorded from the English Channel 2.22Adults (Hellmich1962) -to South Africa (Loveridge and WilliazUB, 1957; Rose, 1962; Hughes, 1970). The distribution of adults la determined to a large extent by -the location of their nesting r2he northernmost records for green turtles beaches and their feeding grounds.James J, in the Pacific are Washington State and British Parsons (1962) has written an elegant book in Columbia (Carl, 1955; Logier and Toner, 1961; which he documents and describes many of the Slater, 1963), BoniIslands (Stejneger, 1907), nesting sites around the world.The important Japan (Wang and Wang, 1956), and Korea (Shannon, still extant siteø, na well as recently 1956).They have been reported as far south as discovered nesting beaches, are summarized in- the the ICermadec Islands (Olìver, 1910), New zealand following sections. (McCann, 1966a, b) and Chiloe, Chile (Donoso- Barros, 1966). The well-known nesting areas are cited in Tab1II and Figs, 4 and 5.All the major Brongersma (1961) believes that the dead greennesting beaches are within 260M or S of the turtles that have been found on the Dutch coast equator.The majority of the beaches are on were released from passing ships.In 1952 a l:Lve islands.- Heron Island and Europa Island are the juvenile Chelonia (carapace length 36 cm) was two moat southerly major breeding sites of the stranded near Fetten on the coast of the green turtle; beaches on the coast of West Netherlands, but it is unhaown whether this too Pakistan, Baja California and In the Hawaiian was released by man. Islands are the moa-t northerly rookeries. Few Chelonia have been sighted off the western Some lesser-known green -turtle rookeries are coasts of South P.inerica sad Africa south of the cited in Table III.!'bat cf these are also on Tropic of Capricorn, and none have been seen southislands. of 40°S latitude on the eastern coast of South America.The lack of records in these southern In addition to the rookeries cited in Tablee latitudes may be due to the fact that the dominantII and III, nesting has been recorded in various surface currents in these regions (Fern or Humboldtother places in the tropics.These nesting Current, Faikland Current, Benguela Current) are locales are given below,In some of these places cold currents, the season or density of nesting is unknown and in other cases the beaches are probably no longer This synopsis does not include references to used by green turtles. strictly random sightings of green turtles. How- ever, such records should be pooled and analysed Distribution in the Pacific Ocean and in order to determine if cryptic patterns of adjacent seas. movement and distribution do exist.For exanp]e, the sightings of turtles in the shipping lares of Green turtles have been sighted or caught the ITorth Atlantic warrant plotting and study. off the Pacific shore of Panama (Evens, 1947), Peru (Murphy, 1925) and Chile (Ysnez, 1951) but Nan's activities and depredations on green nesting beaches on these shores are unknown. turtle nesting beaches (taking adults aM eggs Nesting on Islas de Revillagigedo and Baja 2:2 FIRM/$85 Green turtle

TABLE II Some major green turtle nesting sites

Locality Nesting SeasonPeak of Nesting Authority

WSTEHN HEMISPJIE1?E

Michoacari Coast, Mexíco May-S ep-t ember Peters, 1954, 1956-57 Duellman, 1961 Montoya, 1969 Tortuguero, Costa Rica Jl2neJov. late July-early Sept.Carr and. Ogrem, 1960 Shell Beach, Guyana March-August probably June-July Pritchard, 1969 Bigi Santi, Suririam Feb-Augusb April-May Pritchard, 1969 Schulz, 1969 isla Aves March-Doc. probably July Parsons, 1962 Lazell, 1967 Marice1958 Hirth MS)

EAZTERN EBIMISPHERE Ascension Island Doc,-June Feb.-April Carr and Hirth, 1962 Southern Yemen probably year one peak in Hirbh and Carr, 1970 around November Hawk Bay, year aróund. late June-early Nov. Hatt, 1957 West Pakistan Miìiton, 1966 Aldabra Atoll year around February4ay Hornoll, 1927 Europa Island. year around G.R, Hughes (MS) J.Y. Cousteau (film) Parsons, 1962 ro IOwan, Thailand year around May-July Penyapol, 1958 Nalaysia year around July-August Harriason, 1951, 1952, 1965a Hendrickson, 1958 de Silva, 1969a Indonesia probably year Somadikarta, 1968 around Philippine year around August Domantay 1952-53, 1963 "Turtle Islands" Heron Island, Australia Oct .-Maroh January-February Moorhouse, 1933 (and other Capricorn Bustarc11967a, (Ms) Group Islands) Bountiful Io., Australia June-Feb. Bustard (MS) (and. other islands in Parsons, 1962 Gulf of Carpentaia) FIltM/S85 Green turtle 2:

Fig. 4Some of the important green turtle nesting beaches (numbered oiroles) and feeding grounds (shaded areas) in the western hemisphere.Modified from Parsons (1962) and Carr (1965). 1 Ujalang, 2 D'Entrecaateaux, 3 F'unafutio, 4 = Vatoa,5 Palmereton, 6 Rakahanga, 7 Scilly, 8 Rarola, 9French Frigate Shoal, 10 = Galápagos, 11 Clarion, 12 Michoacan coast, 13 Bahía Magdelena, 14 Bahia San Bartolome, 15 = Quintana Roo, 16 = Tortugtiero, 17 Little magna, 18 Mona, 19 Aves, 20 Guyana beaches, 21 Surinam beaches, 22 Marajo, 23 Rio Doce, 24 « Trindde 2:4 FIR!fS Green turtle

Fig. 5 Some of the importent green turtle nesting beaches (numbered circles) and feeding grounds (shaded areas) in the eastern hemisphere. Modified from Parsons (1962) and Carr(1965), 1 = Cape Verde islanda, 2 = Ascension, 3 = Turopa, 4 = Comorea,5 Astove, 6 = Aldabra, 7 Southern Temen beaches, 8 = Masira,9= Hawkee Bay, 10 Lacoadivos, 11 = Naldives, 12 e Diamond, 13 = Cocos (Keeling) Is., 14= Gulf of Thailand beaches, 15 = Sonhla, 16 = West Malaysia Islands, 17= Sarawak turtle islands, 18 = Philippine turtle islands, 19 Schildpad, 20 = Baroeng, 21 = Flying Foam Is., 22= Gulf of' Carpentaria Islands, 23 = Torres Strait Islands, 24 = Heron Island,25= Central Caroline Islands, 26 = Oroluk FIRN/S85 Green turtle 2:5

TABLE III

Season of neting at some lesser-known and minor green turtle rookeries

Nesting (by monthi) Locality Authority J F N AM N D

Pacific Ocean and adjacent seas

Chiapas, Mexico XX XX X X Alvarez del Toro, 1960 Galpagoe In. X X X X P. Pritchard (MS) RoseAtoll X X Sachet, 1954 French Polynesia X X X Chabonis, L, said F., 1954 Hirth (MS) Tonga Islands X X X X Hirth (MB) New Ca].e'lonia X X Hirth (MS) Bikar Atoll, Marsh.11 Is, X Fosberg, 1969 Jano Is] and, Marshall Is. X Fosberg, 1969 Houtman Abrolhos, Australia X X Storr, 1960

Atlantic Ocean and adjacent seas

Yucatan Peninsula X X X X R. Marquez, pers. comm. West Indies XX XXX Underwood, 1951 Coast of West Africa X X X X X Gadow, 1909 Senegal X X X Cadenat, 1949 Villiers, 1958 FernandoP00: Gulf of Guinea XX X X Eiserrtraiut,1964

Indian Ocean

Naziwi Is., Tanzania X X X X X X X Anon, 1969b Dahlac Archipelago X Anon, 1969a; Urban, 1970 Antove and Comsoledo in Seychelles Is. XX X Mirth and Carr, 1970 Jurayd Is., Persian Gulf X X E, Hoag, pers. comm.

Manira Is. X Parsons 1962 FIRM/585 Oreen turtle California is mentioned by Beebe (1937), Brattetrom nesting in the Bonin Islands; and in 1968 (1955) end Parsons (1962).These areas should be Sampson mentioned hunting "sea close roconrxoitred again, with special, emphasis on to shore.If still extant, this would assessing nesting aggregations. constitute one of the most northerly breeding sites of Chelonia. In former years green turtles were readily caught in waters off the Hawaiien Islands espe- On the western perimeter of the Pacific cially near Galia, Mololcai and Maui,Today the Ocean, green turtles have been reported from majority of nesting is on French Frigate Shoal Amami-oshima in the Loo Choo Islands (.'Ryukyu) in the Hawaiian Lewards.Especially large (Koba, 1959) and, from Taiwan (Gee, 1929-30), concentrations of basking turtles have been seen Pope (1935) 'cites references pertaining to on Pearl and. Hermes Reefs and somewhat smaller green turtles laying eggs on islands near Hong concentrations on Limianeki Atoll anti Layson Atoll, Kong in July and August.However, nesting here all In the Hawaiian Leewards.Hendrickson (1969) now seems very doubtful. and Northshleld (1969) report green turtles are still found in the Leewards, but are scarcer than The density of nesting today on Pulnu they used, to be.Now that the Leeward Islands are Berhala off Sumatra is unknown, but Mohr (1927) wider the guadianship of the U.S. Bureau of Sport noted oviposìtion every month with a peak from Fisheries and Wildlife, the turtle population may November to January.Do Rooij (1915) listed. stabilize or even increase.In any case, whatever. m,as as occurring on the following islands of is left of the original population, the Hawaiien the Indo-Aus'tralian Archipelago; Sumatra, Leewards reriain the biggest green turtle rookery Biliton, Borneo, Java, Madura, Floree, Celebes, in the United States. Ambon, Benda Obi, Aru and New Guinea. Pangumbalian, a nesting site on the south coast Parsons (1962) end Wiens (1962) have docrnjnentedof Java, s-there a large number of eggs are theoccurrence of green 'turtles in Melanesia, harvested, is described by Rekeowardojo (1961). Micronesia end Polynesia.In earlier days Chelonia Sornadikarta (1962) lists some nesting beaches noeted, in fairly large numbers on Uli-thi, Ifaluk, in Indonesia which are rented out for their Gaferut, Hiato, Olimaran, Fanning Atoll, Christmas eggs.Some, no doubt, are green turtle rookariea- Atoll, D'Entrecastoaux Reefs (July through October),Parsons (1962) describes nesting near Baroeng nerican Samoa (August and September), 'Oro].uk Atoll,in Java.A thorough 'taxonomic and ecologic Raroia (June through September) and Vaton.The study should be made of the green turtieth density of nesting on Palmerston Atoll now is un- Indonesia.Especially cinco the Archipelago known, although Cari' (1965) liste it as a major occupies a unique position between the Indian nesting beach.There may still be a large nesting Ocean and South China Sea. colony on Palrnerston and, the situation warrants immediate attention.Preliminèry attempts at In addition to the large Sarawak rookeries assessing turtle stocks in the South Pacific have in i'ìalaysia (seo Table Ii) green turtles also been made by Anon (1969f) and Bustard (1969a). nest on small coastal islands in West Malaysia. The moat important of which are the Porhentinu Green turtles nest occasionally on the sandy and Redsñg Islands off the coast of lCslanten beaches of Arno Atoll in the Marshall Islands hut and Trengganu (Hendrickson, 1961). they are scarce and of no commercial importance (Hiatt, 1951),They aro now rarely seen around Besides the well-kncwu Philippine ItTurtle Kapinganarangi Atoll in the Caroline Islands where Islands", (see Table II)other islands in the they once were an important source of food, (Nierin,Sulu Sea where some nesting occurs include 1963).They are reported to neat in the Helen's Cavi li, Arena, Lumbucan, Bancoran and. San Reef area about 480 km southwest of Palau and on a Miguel (Martin, 1952-53). small island north of Palau and just off the island of Kayangel, all in the Caroline Islands Cockoroft (1968) stated that green 'turtles (Wilson, 1969).Carr (1965) cites nesting on are common in wa,rm waters off Papua, and Burley Ujelarig in the Central Carolines and on Punafutio (1924) briefly described nesting on a small and Rakahanga in the South Pacific.Fehl (1958) island in the Torres Strait. provides a few references to early accounts of green turtles in the western Pacific and. Indian Copulating pairs have been seen In Oceans.Dr. John Hendrickson (University of September and October off the beaches in Arizona) is currently studying the marine -turtle Northern Territory, Australia, and reliable populations in Micronesia. sources indicate largo nesting aggregations ori islands in the Gulf of Carpentaria (Cogger As, far as is known, the extent of nesting in and Lindner, 1969; Bastard, MS).Evidently the Ogasawara Islands (Bonin Islands) has not there is large-scale nesting on the western been elaborated on since (1931) account coast of Australia If the following quote is which included, statements that mating takes piace reliable: "All alone the northern beaches of from 1'larch to May and eggs are laid from May the mainland coast (of western Australia) and through August.Mmkino (1952) also referred. to selected off-shore ielaxidn, thouaandc of feniale FIRN/S8Green turtle 2r green turtles leave their natural environjnentthe (carapace lengih 30.4 cre) captured off Madeira. sea, te make a difficult and. exhausting journey on-. to the lend to lay their eggs" (Anon, 1969e). The northwest bulge of Africa Is one of Another article reports nestingnear Onslow in the least known areas as far as marine turtles WesternAustralia (Anon, 1970e). Mating is report- areconcerned,-C. nwdae appears in a glossary ed off the Great Barrier Reef in Octoberand prepared by Osorio de Castro (1954) and. is November (Roughley, 1951). simply listed in -the fisheries of coastal Portugal, Azores, Madeira, Morocco and Senegal. Distribution inAtlanticOcean and adjacent Pasteur and. Bone (1960) state that it is pro- seas bable that at least stray green turtles can be found offMorocco. Oreen turtles nest on Papaya Beach, Turtle Beach, Leguan Beach, Tigsr Island Beach,Sudxlie Dïs-bnjbutjon in the Indian Oceanand Beach, Zeelandia Beach, l4ahaica-Mazaicong Beach Mediterranean Sea and. Shell Beach (1arch through August) in Guyana; on Silebache BeachinFrench Guiana;and. on Bigi. Green turtles are reported as very common SantiBeachand on the strand. near the mouth of in Mozambique waters and need. better protection the Narowijne fiver inSurinain (Pritchard,1969). on the nesting beaches (Anon, 19701).Today Frairand Pral(1970)saw many greenturtles there la only scattered nesting on the coast of nesting in May.in Surinazn. Kenya (C. Ray, pers. comm.). oviposition has been observed.onthe Somali coast (Cozzolino, In addition to the references given in Pable 1938; Travis, 1967). II somepe,-tinentaccounts of the rookery on Aves Island canbefoundin articles by Zuloaga (1955) The photograph in the Arameo Handbook LAra- and. PhelpsandPhelps(1957). Aves is the only bina American Oil Company, 1968) with the legend above-wate partof the so-called "Aves Swell". "Giant Sea Turtles Breed on Islands in the Arabian The island is almostsurrounded. bya reef and, the Gulf" is probably agreen turtle. There aro also riiaxìmuinelevation above sea level is3.5m. scattered. reports of green turtles off Iran in the The island is also a well-known ternrookery. Persian Gulf, The fact that the island is sinking raises the question cf what returning, oviferoumturtles will Green turtles were reported asverycommon do when the-re is not enough beach left for nesting. in the Maldives some forty yearsago -. Monco(1958)states that it Is possible -be capturo (Deraniyagala, 1956). Nesting on Gomeros Islands, from 150 to 200 fecales in a few nights during June Diamond Island and Cocos Island is documented by and July on Aves Island. - Parsons (1962), The current statue of these islands as nesting' sites isunknown. The Ascension rookery has been briefly mentioned byPacker(1968) besides the authorities - - Shockley (1949) along with HatL (1957) and cited in Pable II. In addition to its remote Minton (1966) describes nesting in West Pakistan location, thi colony is different from some of (see Table ii), the other major rookeries inthatthe incubating eggs are rarelytakenby any predators but Several people (Bodenheimer,1935;.Hans, entrance tosomeof the beaches ismade hazardous 1951;Howella,1956)have reported the presence by the combination ofheavy seas androckyshore- of Chelonia on the Israeli coast. The area line. between El Anish and Hedera is sometimes named am a nesting site; however, it is highly unlikely Some of the much lesser-known rookeries in the that turtlesnest there now. A couple of Caribbean and. western Atlantic arelisted in Chelonia wore caught near Malta some forty years Parsons(1962). These include Isla Mujeresand ago (Despott, 1931). G. mydas is reported as other coral islands off thecoastof the Yuca-ben unoonmon in the Adriatic Sea (Redi, 1963). The Peninsula (see alsoFoeberg, 1962;Mexico, 1966) greenturtle along with loggerhead. andhawknbill MonaIsland; Maraj6 Island; beaches near Rio Doce turtles has been observed off southern Prance and Cabo Pio on the Brazilian coastandTrindade. (Knoepffler, 1961). Barth (1962) hasdescribed the nesting beaches on Trindade, hut there is a dearth of information The onlyknownbreeding colony of any size about thebreedingcolony -there. Green turtles in the Mediterranean is l;hs rookery on the have aleo beenseen off the shore ofQusimada southern coast of Turkey near Marsin and. Grande, about 70 kin southwest of Santos, Brazil Yasrortalik (lu-ion, 1967a). According to Mr. I. (Mertens,1955). - Sela of Israel, the colony is rapidly diminishing due io pverexploiiation. The rookery is now Parsons(1962)cites nesting in the Cape Verde being investigated by Dr. R.W. Hathaway Islands.Brongerema (1968a) quotes the work of (University of Uai), Sarmento(1948)which mentions the presence of juvenilegreenturtles around the Madeira Islands In some places in the Mediterranean the andBrongerema himself describes a juvenile reports of Chelonia are questionabl because some reporters have had. difficulty in 2:8 FIRM/585 Green turtle distinguishing the groen turtle from the loggerhead and March near Sunday Island in the Kermadec turtle. Group (Oliver, 1910).They do not breed in the islands but are reported to go north for mating. Green turtles are vry rare in the Black Sea. Individuals have been seen feeding on the bottom One was caught near Sozopol in Bulgaria about TO in shallow waters off Northern Territory, years ago (Valkanov, 1949; F\Thn and Vancea, 1961). Australia (Cogger and Lindner, 1969; Bustari, MS), and. ndas is reported as a staple item in Ication of feeding pastures The diet of a small number of coastal aboriginals still living a tribal way of life. Some of the principal feeding pastures are: upper west coast of Florida; northwoetern coast of In the Fiji Islands green turtles can be Yucatan peninsula; south coast of Caba; Mosquito seen feeding in the. passage between Taveuni coast of Nicaragua; Caribbean coast of Panama; and Vacua Levu (Bustard1970b).Hiz"th (MS) scattered areas along Caribbean coast of Colombia; has found feeding pastures around Viti Levu in scattered areas off Brazilian coast; west African the Fiji. Group as well as around Tongatapu in coast from Mauritania to Nigeria (see Villiers, the Tonga Islands. 1958, 1962 for descriptive accounts); Mozambique Channel; Bajun Islands and neighbouring coast of Very little is known about the population. Somalia; Khor Umaira in the Gulf of Men; Maldive dynamics, productivity or activities of green and Laccadive Archipelagoes; around Ceylon and turtles on the feeding pasturesthis in spite Krusadai Island (see Deraniyagala, 1939, and of the fact that a turtle probably spends most Kuriyan, 1950); scattered areas in the South China ofita life there. Sea and ii the Gulf of Thailand; northern sector of the Gr:at Barrier Reef; Torres Strait; Gulf of 2.3Determinants of distribution chaest Carpentaria; Flying Fbam Islands; Hawaiian Islands; see sections 2.1, 2.2, 3.16, 3.22 and around scme of the Fiji Islands and. Tonga Islands; 3,5 Pacific coast of Baja California; Gulf of California; Galapagos Islands, and parta of the coasts of 2.4Hybridization Ecuador and Peru.Figs. 4 and 5 indicato the approximate liriits of some of the better known Nothing is known of hybridization in feeding areas.As is to be expected, the most nature. The author suspects, however, that extensive pastures are along continental shelves. hybridization would occur if turtles are able to pass through the newly proposed croacPanama There are severa], lesser known feeding areas. Canal; and, if captive turtles escape from Green turtles are reported grazing between January turtle farms. P1RW585 Green turtle

3 }3IONOMICS AND LIFT HISTORY larger breeding areas (i.e., Tortuguero, Sarawak Islands, Bountiful Island, Aldabra Atoll) 3.1Reproduction one female is usually attended by several males and males show an interest in any female,This 3.11Sexuality suggests that polyandry may be the ffLQCIU ooerandi, but it may also be a reflection of the Green turtles are heterosexual.Adult malc3 fact that females, but never males, are exploited have a very long, somewhat prehensile tail reaciiIn on the nesting grounds.Or it may reflect on as far as or beyond the extended hind flippers. the fact that all males rernigrate (= periodic The tail of the adult f emale rarely extends beyond movement oui from and back to the original area) the margin of the carapace,tiales also have longerto the nesting ground. every year while most claws on their front flippers than do females. females have a biennial or triennial cycle.How- Some investigators have had difficulty in deter- ever, it seems unlikely that males do romigrato mining the sex of hatchlings, but Idakino (1952) each year because of the bioenergetic require- states that the sexes of even embryos can be mente of such extended travel.On the other determined by histological examination.According hand, the preponderance of males may not be real to him, the embryonic ovary of G.m. ,jaionica is at all but merely a reflection of the male's easily distingaishable from the testis by having very intense activity during the breeding season. a thick and elongated cortical layer of primordial It is also possible that the sex ratio may germ celle which show active proliferation.The indeed be in favour of males at the start of the testis is a little shorter in length but mach breeding season but that i-t shifts to ari equal thicker thn the ovary of the same age.He goes sex ratio as the season progresses due io arrival on to say that the testis is further characterized of more females or due to males leaving.It is by the fac t that the primordial germ cells develop noteworthy thai at Tortu1piero, nesting continuos as solid cords of irregular shape towards the after mating activity has ceased and males have medullary region of the gonads.Farther histo- virtually vanished from might. logical stadien are now in progress in the laboratories a-k the University of Utah. 0f' course, the preponderance of adult malee off some nesting beaches may also be explained 3.12I1aturity by an unequal sex ratio at hatching or a sexual advantage during ontogeny (see seckion 4.11). Investigators are not certain at what age green turtles reach maturity.Estimates are foui' The question of when the eggs of virgin to six years (Hendrickson, 1958), at least six females are fertilized remains sii enigua.It is years (Cari', 1967b), at least seven years hypothesized that the young females accompany- (Harrisoon, 1962d), five to seven years (Ehremfeld,the older turtles to the nesting grouird, mate 1970) and eight io thirteen years (Caldwell, the"e and return to home pastures without laying 1962o).The age, or sise at which sexual maturity (Carr and. Hirth, 1962).Also, according to Carr is reached may of courae vary between individuals (1965), "It seems unlikely that any of the eggs of the same population and also between of' the current season are fertilized as a result individuals of different populations.For example,of current mating.The Encounter probably Carr and Carr (1970b) have discussed the variation servos to fertilize eggs for the next nesting in maturation siso at the Tortuguero rookery. season, two or three years ahead".This storage Evidently this population inc1udetwo kinds of of spermatozoa may account for the infertility big turtles: some that grew big after reaching of large numbers of oggi: (see section 3.16). sexual maturity, and some that reached maturity at -k'. Montoya of Mexico reports (pers. coma.) that a large size. fertility in C.. agassizii eggs is higher in the first clutch of the season than in later The smallest nesting females recorded on the clutches. Tortuguero and Southern Yemen breeding beaches had. carapace lengths of 69.3 and. 78.7 cm respec- Some aspects of courtship and. copulation tively (Carr and Ogren, 1960; Hirth and Cari', off the Tortuguero rookery are given by Carr and 1970),Sizes of nesting females at some of the Giovarmoli (1957).They describe how pairing is better kxzown nesting beaches are given in Fig. 6. a strenuous and clumsy operation during which Corresponding weights can be ascertained from the female is often geashed and scraped..The Fig. 7. enlarged claw on the front; flippers of the male aids him in grasping the foro edge of the shell 3.13Eating of his mate. Copulation usually occurs just off the neat- It is plausible that some mating takea ing beaches, usually within one kilometre of shore.place at the feeding grounds or at other areas It is unknot-.'n whether the males accompany the away from the nesting beaches, although thie has females from feeding pastures to nesting grounds yet to be documented. or whether they make a synchronized rendezvous with the females at the rookery.At some of 'the 3:2 FI 58Green turtle

Nz225 SOUTHERN YEMEN

N2OO SARAWAK

N=200 COSTA RICA

N2OO ASCENSION

N 6OSURINAM

70 80 90 lOO f10 120 130 f40 CARAPACE LENGTHS OF MATURE FEMALES [IN CENTIMETERS]

Fig. 6 Sizes of nesting females at some major nesting sites0 The length of the horizontal line indicates the range and the vertical line the mean0 Data on the Sarawak population are taken from a graph in Hendrickson(1958). Other data are from Carr and Hirth(1962), Priichard (1969)ami Hjrth and. Carr(1970) F1RW385 Green turtle 3:3

200

175

150 (I)

o: e, -Jo zI0o H !275 w

50

25

75 25 50 lOO L$O CAfl/ PACE LENGTH IN CENTiMETERS

Fige7 Node). showing weight and carapacelength of greenturtles (both sexes), Adapted from HirthondCarr 197O)andbased mainly upon turtles caught inthe Gulf ofMen feoclingpastures tRM 38Green turtle 3.i4 Fertilization (i) newly matured individuals, (2) reburnin individuals nesting on a biennial cycle, (3 Fertilization is intrnal,The question of roturithg individuals nesting on a triennial delayed fertilization is discussed in the precedingcycle1 and (4) individuals exuerioncing a phase- section. shift in their reproductive cycle.Oviferous turtles may converge on the rookeries from 3.15Gonads different feeding pastures, como of which may be long distances from the breeding beach. See following section. Meat nesting takes placo at night, altlion 3.16Reproductive cycle, nesting daytime nesting does occur occasionally on behaviour and selection of nesting uninhabited Pacific atolls and has been seen in beach Colombia: (Medem, 1962) and Surinam (Pritchard, 1969).The time of year when nesting begins and. Men is most familiar with two relatively shortthe duration of the nesting period vary from one phases of the life history of the green turtle - region to another (Tables II and III).In some the nocturnal egg-.laying behaviour of the adult places, such as Malaysia, there is year-round female, and the subsequent two months of incubationnesting but with specific peak months, while in and, hatching. other places such as Ascension Island, nesting is restricted to a few months of the year.At Green turtles nest every four years in eastern.Tortugnero, a mainland beach where nesting is Australia (Bustard and Togxiotti, 1969), every three seasonal, the turtles appear in large numbers years in the South China Sea (Harrisson, 195Gb; rather suddenly, but tend to trickle away at the Hendricksoi&, 1958) and every two or three years in end of the season. the Caribban and South Atlantic, but with the triennial cycle predominant (cari' arid Ogrexi, 1960; Most green turtles lay between three and Cari', 1965).Recently Carr and Carr (1970a) have seven times each season at about ten to fifteen accumulated evidence which indicates that a four.. day intervals.Hendrickson (1958) stated that year reproductive cycle may also exist in the Costaabout 96% of 5,748 nesting green turtles in the Rican population.&z'eover, they have shown how, three Sarawk Islands returned -to the same island although an individual usually maintains a constantfor renesting during the reproductive season cycle, modulation may occur and the change may be even though two of the islands are only about either from a three to two year cycle or vice versa.500 metres apart. It is postulated that the phase-shift reflects ecological conditions on the feeding grounds,Thus, Renestinge on the Toz'tuguero beach aro individuals nesting in any one given year on some usually within 0.4 to 1.2 km of the preceding beaches around the Atlantic represent four aouroes nest_ng emergence (Cari' and Ogren, 1960), e-ven

TABLE IV Basic reproductive data a-t some major nesting beaches. The rango is given in parenthesus following the mean.

No. eggs Incubation period. Renost Ing interval Locality per nest in daye* in days Authority

Tortuguero 110 55,6 12.5 (18-193) (4870) (9-16) Cari' and Hirth, 1962 Surinain 142 13-14 (87226) Pritchard, 1969 Ascension 115.5 59.5 14.5 (53-181) (58-62) (10-17) Carr aiid Hirth, 1962 Sarawak 104.7 57-10 10,5 (3-134) (depends on season) (8-17) Hendrickson, 1958 Aunt ra.i ia 110 68,7 14 ioorhouse, 1933 (50-200) (65_72) Bastard, 1967a

* Incubation period is defined an the poriod between oviposition and anerence of the largest number of ha-tchlings on the surface Grejurtle thou the beach topography changes from year to ori the beach arid the littoral predators during year.Apparently, the males roam about just off.- the first few critical hours in the water, shore during the mating season, (2) metabolic heating of the nest by a mass of eggs, (3) social facilitation in the climb to In order to find out what females do between the surface, arid (4) group orientation in the nestingaa couple of mature individuals at crawl to the sea.Carr also speculates that Tortuguoro were rigged with helium balloons and hatchiinga may regroup somewhere after entering kept under visual observation (Cari', 196m), One the sea and somehöw take more advantage from individual that tias allowed to nest was tracked being a group again. the next day swimmthg around off the nesting beach. Another individual that was not permitted to nest Metabolic heating of the nest has been struck out in a southeasterly course for about six studied in several places.In Sarawak, hours, but the following night returned and nested Hendrickson (1958) found a progressive tempera- near the point where she was released,Further turo rièe (from about 28° to 35°C) in nests of studios on this subject using turtles rigged with developing eggs arid postulated that this might transmitters are now being conducted at Tortuguero be an important factor in influencing the per-. (Carr, pers. comm.) cent hatch.Carr and Hir-th (1961) found that the temperature of a nest on Ascension increased Table IV compares some basic reproductive data from 27.9 to 29.9° over a 61-day incubation at eome of the major nesting beaches.It appears period,In Australia, Bustard, and Greenhain that in addition to being among the biggest green (1968) found that the temperature within a turtles in the world, the females nesting in natural nest of eggs is initially about 25-26°C Surinam hav3 the greatest fecundity.Banks (1937) end that there is a rise in temperature to about studied ovposition in the Sarawak turtle islands 31°C during the latter part of incubation due to between 19:2 and 1935 and found that the average metabolic heating. number ofggs laid by each turtle, based upon thousand cf records, varied from 106 to 118,The In addition to the incubation data for some average number (and range) of eggs per nest at some major rookeries given in Table IV, additional of the other nesting grounds are as follows: information is available for lessor known Indonesia, 118 (48.-175) (Mohr, 1927); Sabak, 105.6 beaches.Dornantay (1952-53) and Penyapol (1958) (72-144) (}iarrisaon, 1965a) and Southern Yemen, report average incubation periods of 51-53 and 106 (70-130) (Hirth and Carr, 1970).Herriason 48-50 days in the Philippine Islands and in (1962e) describes the monthly laying cycles in Thailand, respectively; Thiksda (1965) states Sara'.îak.Bustard (1967a) found that clutch size that it is 63 days in the Ogasawara Islands; increases with age or size of the turtle On the andHirth and Cari' (1970) registered 48-49 days other hand, Domantay (1952-53) states that younger in Aden. turtles lay more eggs than older ones. The percent hatch (defined, as the number Cari' (196m) states that at Tortuguero the of hatchlings, out of the total egg complement, female lays more than 100 eggs on her mid-season which successfully roach the surface) varies trip ashore and fewer on her first and, last trips. from season to season in different parts of the 0L Ascension Island the first and second egg world (Table y).Pritchard(1967) states that complements of each turtle are usually bigger than "the fertility of groen turtle eggs is low, a the third (Cari' and Hirth, 1962).In the Ogasawara 50% hatching being about average".He also Islaxidn,Fukada (1965) reports that each turtle J.ays states that egg complements which hava been dug between one and five times each season and that up and. reburiecl take longer to hatch than other- each clutch consists of about 70 to 150 eggs. He wise, but the fertility does not appear to be observed four individuals through four nestings and substantially affected. no sigoificant correlation was found between the sequence of nestings and number of eggs per clutch Balasingham (1967), in an interesting study except that the last nesting had the fewest eggs with leatherback (Dermocbelys coriacea) nests, in each case.Domantay (1952-53; 1968) has observedfound that ho cou,J,d increase the percentage of several old Philippine turtles go through the entirehatohlinga from transplanted nests if eggs were nesting repertoire and them not 1ay eggs.Evidentlyburied in clutches of about 50.The enigaa of some turtles do not retain their reproductive capa- why the leatherback usual].y lays between 85 and city throughout life.Further studies on this 90 eggs per nest thus becomes more puzzling in subject are needed, especially in view of the view of Balasingham'a work.Similar studies practical importance this would have in the manage- should be conducted with green turtle clutches. ment of the species. The nesting behaviour of the (-reen turtle Carr (1967b) has lucidly discussed how the can be divided into eleven stages (Carr and number of eggs laid by a creen turtle at each Ogren, 1960): oviposition (i.e., about 100) has adaptive value for the species,Some of these advantages include i) Stranding, testing of stranding site, and (i) escape from predation by swamping the predators emergence from wave wash 3:6 YIR1/S85 Green trle

TABLE V

The percent hatch of green turtle eggs. All are reburied. clutches in egg hatcheries unless otherwise indicated.

Locality Percent hat ch Authority

Tort uguero, Costa Rica 50 Carr and Hirth, 1962 Ascension Island 54 (natural nests) Carr and Hirih, 1962 Malaysia 47 Hendrickson, 1958 30-79 Harrisson, 1961, 1962a 53-55 Balasinghein, 1965 Anon, 196Th Sabah 70-90 de Silva, 1969a Heron Is., 67 Bus-tard and. Greenhaxn, 1968 Australia 55 (naural nests) Noorhouse, 1933

Selecting of course and crawling from surL to it may involve sensory appraisal of the beach, nest mite Evidently this trait is lacking in the Pacific forms (Bust ard and Cire enham, 1969). Selecting of nest site The tendency to make "half-moons" (. non- Clearing of nest premises egg laying emergence in which the trail up and down the beach looks like some form of a Excavating of body pit parabolic curve) and trial holes may be signs of appetitive behaviour, although they may also Excavating of nest hole merely forms of reproductive unreadiness. Hendickson (1958) believes that abandoned nests Oviposition in advanced stages are due to vibrations in the substrate caused by nesting turtles nearby. Filling, covering and packing of nest hole Phis certainly would be of. adaptive value on crowded beaches by spacing out the nests (see Filling of body pit and concealing of site of section 4.5). nesting The females are very shy when ascending io) Selecting of course and locomotion back to sea the beach. At this time they can be easily soared (i.e., by barking dogs, lighta etc.) ii) Re-entering of wave wash and traversal of the back into the sea. Hendrickson (19583 describes surf the ascent up the beach to the high beach plat- form in some detail. When -the turtle is digging Some detailed aspects of the nesting repertoirethe nest hole and egg hole, she io much less of the Atlantic green turtle are given in Carr end sansitive to alarming stimuli. Once the consum- Cijovannoli (1957); Carr and Ogren (1960) and Carr matory act of egg-laying has commenced, virtually and Hirth (1962). Nesting of the Indo-Pacific form nothing will canse the turtle io stop. It is at is described in varying degrees of detail by Beebe this stage that large beach predators can easily (1937?; Hendrickson (1958); Bustard and Greenhain kill a turtle. The complete nesting repertoire, (1969); and Hirth and.Carr (1970). i.e., from water's edge back to surf, takes about two to three hours. "Sand-smelling" or "sand-nuzzling" by the newly stranded female on the beach has been At some rookeries there appears to be no observed by several people in different localities direct correlation between phase of the moon or (carD and Giovannoli, 1957; Carr and Ogren, 1960; stage of tide aM nesting, but around some of Carr and Hirth, 1962; Hirth and Carr, 1970). The the Pacific atolls a high tide does aid the exact function of this mannerism is unknown, but turtle in reaching the beach and most nesting t11N/385 Green turtle io at this time.In other places indigenous Green turtles nesting in Suman appear to poople will claim that most nesting taken place be very plastic in their beach preferences since at certain phases of the lunar cycle. the beaches there are continuously ehaging in physical structure and location (Schulz, 1964, Short of imparting a tendency to pack solid, 1969), aerate too freely, or to waterlog, the physico- chemical characteristics of good turtle beachen Some of -the beaches in -the Philippine are not very well known.However, one charaoter- Turtle Islanda are becoming unfi'b for meeting due intio common -to most rookeries is the presence of -to excess accumulation of logs, uprooted. trees, a beach platform well above flood tide.It is on roo-be of coonu-b palms and other debris cast such a platform that most nesting taken place. ashore (Domentay, 1952-53).Similarly, the Hirth and Carr (1970) found that the texture and strand vegetation is encroaching upon some of composition ofhe sand on some major nesting -the nesting beaches in Western Samoa (Hlrth, MS). beaches vary widely.For example, -the sand on the big rookery a-t Sharma (Southern Yemen) is composed. The cues that green turtles use in select- of particles chiefly in the 0.5 to 0.25 mm olass - ing their nesting beaches are e-bili unknown, that is, medium grain sand.The beaches at It is well established, however, that the same Ascension and. Aven Islands are mainly composed of nesting beaches are used sear after year.For coarse sands and. the beach at Tortuguero is made example, no adult turtle tagged at Tortuguero up of fine sand.Other features of -tha nesting has been re-taken nesting anywhere else except beach sand, vary as much as particle size does. at Toriuguero.It is also possible that Golcur, for instance, varíen from, olive grey at hatchlings are somehow imprinted with some Tor-tuguero to white on ¡tidabra; soluble salt sensible essence of the beach on which -they content on Sharma is about one third that at hatch (Koch, Cari', Ebrenfeld, 1969) end that ib Tor-tuguero; organic content a-b Sbarma is less than is detection of' -this that guides them back to half thai at Aldabra and Aven Islands.Pritchard the same beach when they return as reproductively (1966) reports -that green turtles iay eggs on a mature females ready to nest.This imprinting beach in Guyana that is componed enbirely of trait is also the rationale behind the restocking broken sea shells.Ponyapol (1958) han stated programme in the Caribbean (seo seotion 6.5). that green turtles nest on different types of sandy beaches in Thailand.On the other hand, One of the biggest nesting concentra-tiene, Hendrickson and Balaninghaxn (1966) found that in of green turtles In the world and the site of Malaysia green turtles prefer beaches of fine sand. extensive ethoecoiogical research is the beach Nesting beach preferences, or lack thereof, in near Tortuguero on the Caribbean coast of Costa importent information for the turtle farmer who dica.The nesting beach is approxima-boly 38 must get his turtles to deposit eggs on man-made kilometres in ox-bent and has been described by beaches, Cari' (1956), Ca.rr and Giovennoli (1957), Cari' and. Ogren (1959, 1960), Cari' and Hix'th (1962), Green turtles have laid eggs on completely }Iirth (1963), Carr, Hix'th and. Ogren (1966), cwtficial beaches in the Seychelles Islanda, Cari' (196Th) and Stubbn (1970). Hauaiian Islands and on Grand Cayinan during the course of pilot farming experiments.Details of 3.17Eggs tho type of sand used are lacking. Green turtle eggs are spherical and. white. The presence or absence of vegetation is not During oviposition -the eggs are usually dropped a universal prerequisite for green turtle nesting one, two or three at a timo.Mast eggs are beaches.Some parts of the rookery at Tortuguero covered. with mucous when they leave -the cloaca. are almost covered with Ipomea, $osu-rium, and Egg laying takes about -ten minutes.Freshly Sporobolus, and the green turtles nesting in laid eggs in Sarawak weIgh 36 g each (range eastern Australia seem to prefer the older, more 28.6-44,7) and -those In Southern Yemen weigh vegetated cayo.The beaches on the Sarawak Turtle 40.4 g (range 30-44).The average diameter of Islands are devoid of vegetation1 while on the green turtle eggs ranges from 40 an in Sarawak west Malaysian Islands pioneer plants do invade (Hendrickson, 1958) to 54.6 mm on Aacennion the nesting area.There are some plants on the Island (Carr and Hir-th, 1962).Eggs on the Aves Island strand, but only scattered clumps of Tortuguero and Southern Yemeni beaches average vegetation on the big Southern Yemen rookeries., between 42 and. 46 ram (Cari' and Hirth, 1962; There aro virtually no plants on the nesting Hirtb and Cari', 1970).A few abnormally small beaches on Ascension Island.In Australia, Buatard. or non-spherical eggs are usually found. in each and Greenham (1969) found that shrubs and trees on clutch. the nesting beaches may servo as a cue to the initiation ofhole-digging since a majority of If we assume -that the average green turtle turtles studied commenced digging a body-pit in weighn about 125kg(wet weight), that she, clone proximity to these forms of vegetation.This lays 550 eggsper seasonand thateachegg behaviour has adaptive value since the presence of weighs 38 g,then wo canconcludothat the roc-blets and/or moisture prevents sand slippage average adultfornaio produoeaabout21 kg during the construction of the egg chamber. of eggs per season, or an amount equivalent to 3:8 FIR14/G85 Green turtle about 17% of her own bod,y weight. predator. Penyapol (1958) has analysed the nutritive Rascona are a terrible menace to loggerhead valua of green turtle eggs.He found that ash eggs on the Florida coast and may be preventing accounted for 1.11% of 100 g of turtle eggs. rehabilitation of green turtle rookeries there Based upon the same sample size, he recorded the (Carr, pers. comm.). following major constituents: moisture (79.03 g), protein (11.8 g), phosphorus (227,85 mg), calcium On high-density nesting beaches, such as (93.58 mg) and iron (1.95 mg).The cabrio content Heron Island, Europa Island, and the Sarawak of the 100 g sample was 121.36 (compared to 146 in Turtle Islands, nesting females may destroy hen's egg and 189 in duck's egg),Vitamina B1 and incubating clutches as a oonsequenoe of digging B0 registered 453 and 442 micrograms, respectively. their own nests (see section 4.5) The total edible portion of the sample was figured to be 92.8%. 3.2 nbz7onio and hatohling phase Nan eats the eggs of green turtles chiefly for 3.21 hibryonio phase their protein content; occasionally local demand is based on a belief that they are an aphrodisiac. Domantay (1968) has doouniented. the embryobor of the green turtle from the time the Green turtle es are eaten by a variety of e leaves the ovary to the day of hatching. predators (Table VI).As the table indicates, His observations were made in the Philippine feral dogs take a large number of turtle eggs. Islanda where incubation lasts about 50 days. umani of course, take more eggs than any other According to Domantay, the early stages of

TABLE VI Predation on Green Turtle Eggs

Locality Predator Authority

WESTERÑ }MISPIIHRE Tor'tuguero, Costa Rica dogs, buzzards, opossums, pigs, white-lipped pecoary, coatimundi Carr, 1956, 196Th Galapagos Is, feral dogs, pigs Schmidt and Inger, 1957 P. Prit chard (MS)

EASTERN IIEISPIIERE Lldabra Atoll sand. crabs (Brachyura sp.) robber crabs.rgus latro) ibis, dogs,rata Honeggor, 1967 Southern Yemen feral dogs, foxesbeach Hirth and Carr, 1970 crabs Anon, 19670 West Pakistan feral dogs, jackals, crows1 monitor lizards Minton, 1966 Malaysia ghost crabs (0cyod.e sp.) monitor lizards (Varanus ap.) rat s Hendrickson, 1958 East Indies monitor lizards, pigs (Sus barbatus) Raven, 1946 Philippine Islands dogs, monitor lizards Domantay, 1952-53 FIR48r Orcen turtle 3:9 gastru].ation are completed before the egg is laid; A rather oldbut comprehensive monograph by the end of the fifth day the neural tube is on embryology, especially that of the head, has differentiated into the brain and spinal cord; been written by Parker (1880). differentiation of the heart and gut bogin about the eighth day; the nervous, digestive and Agassiz (1857) described in detail the oirculatory systems aro fully distinguishable by embryology of various turtles, mostly Chrysemys the eleventh day; at about the seventeenth day and Chelydra.It would be interesting -to the eye is differentiated with a lens and iris compare the embryology of green turtles With and the anlage of the carapace is present and those described in Agassis's classic work. differentiating; the scales are fully formed and pjnented by the thirty-first day; two days before Bustard, Sisidsa, Jonkìns and Taylor (1969) hatching the yolk meanures about 30 mm in diameter.have shown that the eggshell is the major source According to Domantay it takes the hatohlings at of calcii.un for developing embryos. least three days to reach the surface. The new hatchiing cuts through its eggshell Penyapal (1958) found that embryos from the with the aid of a horny protuberance, sometimes Thai land rookeries have a carapace length of called the egg-oarunole, on the tip of its Snouio about 5, 11.2 and 36 mm on days 14, 20 and 36, This caruncle disappears after a fow days. respectively; that over the same period the width of the carapace varies from 4 to 31 mm; and, that 3.22Hatohling period (behaviour, the forelimba grow faster than the hindlimbs after predators and curvivorship) day 16. The dorsum of the haichling is black or Deraniyagala (1939) recorded that a 16.-day- dark brownish black or dark greenish black and old Ceylonese embryo has a carapace length of 11 the edges of the carapace and flippers are mm and a carapace width of 6 mm.At the same usually white; the venter is white except the time, the lengths of the fore flippers and hind. ends of -the flippers which are black bu-t edged flippers are 3.5 and 3.0 mm, respectively. with white (see Fig. 8).Sizes of haichlings Between 29 and. 36 days of age, the lengths of are given in Table VII.A picture of a the carapace and flippers almost double, and by hatohling just breaking through the eggshell is the thirty-sixth day the carapace is pißmented provided by Carr (1967o).Albino haichlings (dark olive).According to I'orhouse (1933) all have been reported from Sarawak (Harrisaan, 1963) traces of the umbilicus disappear ab about three and from Costa Rica (Carr, 1967b). weeko of age.

TABLE lITI Sizes of one-day-old haichlings from three different nesting populations(from Carr and. Hirth, 1962;and Hirth and Carr, 1910). Values represent the mean end range and are given in millimetres, A hatchling from Southern Yemen weighs about 23 g (range 20-28 in sample of 20).

Tortuguero Ascension S. Yemen N=100 N1 00 N=20

Length of carapace 49,7(46,0-56.0) 51,7(49,1-55,0) 46,9(44.0-48,4) Width of carapace 38,5(35,0-48,2) 41.6(38,0-48.0) 33,5(31,2.-37,5) Length of plastron 40.0(33.5-46.1) 42,6(38,2-48,1) 36,8(32.8-40,6) Width of head 15,2(13,0-17.2) 17,0(15,5-20,5) 14,2(12,5-15,6) 3:10 Fmnu4/s85 Greco 'turtle Ing only when nightfall brings lower sand surface temperatures,Bustard (1967b) found that the majority of Australian hatchlinge also emergo from the underground. nest at night and he suggests that inactivity during the day of the "lead" turtles near the surface has a dampening effect on the activity of those belowthus ensuring that emergence does not take place when air or sand surface tempera- tures could be lethal.Nroaovsky (1968) concluded thermal inhibition of activity is a major factor in limiting the emergence of hatchling green turtles to nocturnal hours, Activity is slowed down when sand. temperatureo rise above 28.5°C, The advantages of emerging at night are chiefly two: (i) it eliminates exposure to hot sand surface -tesperatures which could be lethal or which could slow down the turtle in its crawl to the sea and. thus lengthen its exposure to predators(2) it eliminates I'ig. 8One-hour old. green turtle hatohlizigs, each exposure to diurnal predators, about 23 g.The dark carapace and. flippers, edgeddth white, are typical of the Hatcblingo crawl along the sand. surface apciea. using their four flippers in typloal reptilian fashion (front member moved in conjunction with The movements of hatoblinga can conveniently hind, member on opposite side), while adults be divided into the following categories (from on shore heave themselves along using some- Carr and. Ogren, 1960): (i) from nest to surface, times only the front flippers simultaneously (2) from nest -to wave-wet sand, (3) across wave- and sometimes ali four limbs simultaneously. wet sand, (4) through wave-wash, (5) through breakers, and (6) travel by hatohlings and. Carr and Ogre:a (1960) conducted a series immature stages (also see sections 3.51 and 3.53). of tests -to elucidate the orientation (= the Phases (j) mid (2) above have been the most taking up of a direction in relation to a intensively studied, while virtually nothing is stimulus) of the hatchlings from the nest to known about the last phase listed. above. the sea,They concluded, that -the fundamental guai sense involved visual stimuli, and that The movement from the nest to the surface is the response is a modified phototaxis,The a combination of negative geotaxis and allolomiinetie "o annosa òf outlook" of a sea-mlnj horizon behaviour, Hendrickson (1958), Carr and Ogren will draw them away from a sun or moon when (1960) and Carr and Hir-th (1961) provide detailed. either is over the land.But this relation- accounts of the complex series of activities that ship between intensity and area is evidently characterize the emergence process - the salient reversed in the case of very bright light. feature being that turtles in a nest are not Morhouse (1933), for example, stated that completely independent individuals, but are sort he could en-tice the hatchlings back onto -the of a super-organism, composed of collaborating beach after they had entered the sea by hatchlings whose combined. efforts carry the shining a bright light.Carr and Ogren (1959) ful" of individuals to the surface.The climb to found that they could draw hatchling leather- the surface usually takes between three and seven backs back -to shore after they had entered days.Emergence from the sand usually takes place the surf by setting a kerosene lamp on the at night.Sometimes the entire complement emerges shore or by shining a flashlight on them, at the saine time, while on other occasions the And,. evidently the combined effects of hatchiinge emerge in small lots, over a period. of illumine-ted sky over cities and. bright street

24-72 hours.This is probably due to subdivision lights account for -the destruction of logger-- of the group by differences in hatching schedules. heads (Caret-ta caret-ta) on the coastal highways Turtles hatching singly have reduced prospects of' of Florida (McParlane, 1963),?csovaky emerging from the neat, and even less chance of (1970) has stated that any kind of visual reaediing the sea, stimulus has some effect on -the seafinding orientation of hatchlings and -that under some Hendrickson (1958) believes that some innate cir':uma-tances the sun or moon could distract mechanism prevents hatchiinga from emerging on the hathhlings from the shortest route to -the sea surface during the heat of the day.Ho postulates and thus increase exposure -time to predators. that if they encounter sand temperatures much Although the problem, is not completely resolved, above 33°C, they cease activity and resume climb- Nrosovskzy (1970) e-ta-tes that the distraction In/8 Groen Lcr io 3:11 to very bright light is counterbalanced by a Lab-roared yearlings retain the ability to behavioural mechanism kuown an tropotaxia find the sea and, even cross complex terrain to (= simultaneous comparison of intensities on the reach it, and adult males evidently possess the two sides, and orientation according to the same ability, although once reaching -the surf balance thus struck) which ensures that hatchlings as hatchlingn, they nover return to a terrestrial will i'oach the water wider almost all conditions. environment except perhaps to bask (C5rr and Hirth, 1962).After conducting a variety of The sea-finding ability of hatohlings am well experiments with G.m. carrinegra in the Gulf of as adult females has been further studied on the California, Caldwelr and Caidwell (1962) Tortuguero beach by Ehronfeld and Carr (1967) md concluded that the sea approach ability is Ehrenfeld (1967, 1968).Spectacles containing present in both sexes of all ages. special filters were fitted to the heads of adulte, and hatchlings were tested in a circular arena As already mentioned, the habits of the where the view of the horizon was unobstructed or hatehling after it enters the sea are completely where it was blocked by a low wall.They unknown.Caer (196m, 1969o) postulates that substantiated the claim of others that the watèr- the batchlings are pelagic for some months finding orientation is primarily a visual process because observations on captive hatchiingo (based on brightness rather than colour) and. that suggest that they have -trouble feeding on the it involves an appraisal of beach topography.Theybottom in deep water and that their counter- also concluded that there was no innate compass shading (i.e., dark dorsum: white venter) direction based on celestial cues, although Ftsoherresembles pelagic fish.Many marine organisms (1964) did find a fixed direction preference are pelagic during the early stages of their life, related, to the position of the sun in hatchlings Cari' suggests that the Sa,rgasso Sea might be one tested in a water arena. place to look for young -turtles, especially sinos -they have not been seen in shore-waters anywhere. In regard to the visual acuity of marine It is also postulated that hatehlings are turtles, Walls (1963) stated that "the female carnivorous (in captivity they readily oat a marino turtle is in a bad way visually when she variety of animal food). Theoretically, the sea.- comes ashore to lay her eggs at night, for her air interface would be an optimal niche for a aerial vision mut 'be hazy and dim even if the small, weak-swimming, air-breathing, carnivorous moon is bright".Walls describes the anatomy, animal.Hughes (1969a) believes that loggerhead accomodation, optic axes and. other aspects of the hatehlings are pelagic and that their movements cheionia.n eye.Ebrenfeld (1967) established that are regulated to a large extent by dominent green turtles aro very myopic when their eyes are surface currents. out of water,Nevertheless, as described above, vision is indispensable for orientation on land. Bustard (1970a) states that -the black carapace plays en important role in elevating Retinal oil globules in G.m. nWdas have been the hatchling' s body temperature when it floats examined by Gronda and Haden t1970).The areal on the surface of the sea.He also recorded distribution shows a mean total o. 7,803 for that hatchlings start to dive regularly at about orange, 8,460 for yellow and 4,132 for colourless. two to three months of age when ihy weigh about The apparent lack of arealdifferentiation is 1,200 g. correlated with its narine niche. Mrosovsky and. Ehettleworth (1968) suggest Mrocovsky and Cari' (1967) examined light that the temperature of currents off the nesting preferences of' hatchlinga in a simple apparatus on grounds may partly de-termine to what extent the natural nesting beach and discovered that whoa hatohlings swim or remain planktonic. gtven a two-choice situation, they prefer blue and. green stimuli over red.Light preferences wore Cald.well (1969) cites records of hatchiinge analyzed further by Mrosovsky (1967) and by being dipneti;ed off their rookeries in the Mroaovar and Shcttleworth (1968), and. it was foundRevillagigedo Islands and off the coast of that while wavelength preferences of hatohlings aroMicthoaean, Mexico,How long hatchlingo remain controlled largely by brightness, the existence of in these offshore waters is unknown, sorne colour preference cannot be ruled out.Ernst and, Hamilton (1969) poritulate -that turtles with Predators of hatohlings are given in Table some nocturnal habits prefer the shorter wavelengthsVIII.As the table shows, hatohlings are taken of light and that diurnal apodes prefer the longerby a variety of avian, terrestrial and marine wavolongilis. predators. In oummary, it has been shown that a bright- A hypothetical surv±imrship curve (Fig. 9) neon preference (at the level cL' the horizon), a indicates the greatest mortality takes place in tropotactic reaction to light and the association the very early months of life.It has been of thoperi horizon with the seaward direction estimated that only 2-3% of the haichlings could explain thc eca-finding orientation of survive to athilthocd (Banks, 1937).It may be liatchiino, at leant on the Caribbean beacliec. as low an 3:12 FI1ilJ/s85 Oreen tut1ç

TABLE VIII Predation on Green Turtle Hatohlings

Locality Fred at or Authority

WESTERN IISPHERB Tortuguoro, Costa Rica buzzards, dogs, jackfish, kingfish, snook, sharks Carr, 1956, 196Th Gulf Coast of Maxioo dogs, hogs, crabs, birds, fish, sharks Mexico, 1966 Pacific Coast of Mexico raccoonshogs, dogs, crabs, fish, sharks Mexico, 1966 01PO9 Is. feral oats, nigh-b herons, frigate birds, ghost crabs, groupera P. Pritchard (MS) Mopelia, Frenoh Polynesia hermit crabs, sea birds, sharks Eggleston, 1953 Marshall Is. Polynesian rat (Rattus eculana) red hermit crabs (Coeriobita penata) Fosbeng, 1969 Fiji Is. mongoose Bustand, 1970b

EASTERN HEMISPHERE Ascension Is. feral cats, rock cod, common jack fish, sharks Cam and Hirbh, 1962 Aldabra Atoll hermit crabs, robber crabs, rats, frigate bird (Fregata minor aldabrensis), flamingoeni- ruber ant iquorum), grey heron (Ardea cinerea), black and whit o crow (Conia aihus), vara-vara (Lutianus bohar) caraxigue-1 ec-dent s (Caranx iobilis) Horne]J., 1927 sharks, barracuda Honegger, 1967 Soirthorn Yemen beach crabs, birds, groupers, sharks Hirth and Carre 1970 West Pakistan crabs, birds, mammals Minton, 1966 Cocos Is. ghost crabs Gibson-Hill, 1950 Malaysia monitor lizard, rats, ghost crab (Ocypode oeratopthalma), snakes (Boiga dendro hUa, Python ret iculatus , fish, sharks Hendrickson, 1958 Heron Is, "host crabs, rats, silver gull Australia (Larus novae-.hollandiae), black.-tipped reef shark Noorhouse, 1933 (Carcharinuspaiianzani) Buatard, 1967a 1'i1*l/385 Green turfle 3:13

1

ÑuY

I t t t t I t

's I O) I o t t w m

GREEN TURTLE

T

AGE IN YEARS

rig. 9Hy-pothetical surv1.vrship curvos of green turtle hatohlins a and man.Plotted on the bae of survivors per thoumaM (log scale) and age in relative units 3:14 FIRM/S8Green turtle 3.3Juvenile, sub-rtult .nd adult phase of the turtle's upper eyelidsHundreds of barnacles (Platylepas hexastylos) encrusted the 3.31Longevity dorsuni and plastron of a sub-adult female taken in Chincoteague Bay, Ilarylend (Schwartz, 1960). Little is known about the life span of green Gupta (1961) took five trematodas from the turtles in nature.The fact that reproductive intestine of a green turtle from Trinidad maturity may be reached after anywhere from four to (Cricccephalus albus, Pleurogonius mehrai, 13 years; that some females have been recapturec Neoctangium travassosi, Deuterobaris chelonei several times, after two to three years'absence, on and Schizarnphiztomoides chelonei Caballero y their nesting beach; together with observations that Caballero (1962) found the trematode, Orchidasma predation on adults is minimal (except 'for man), amphiorchis, in the intestine of a turtle and that epizootics are unknown, and that food, is Simha and Chattopadhyaya (1969) record a new plentiful, all together suggest that adult turtles blood fluke from the heart of a green turtle, more than 20 years of age may be comon (except in places where they are exploited by man). Smith nd Coatee (1938) describe fibra- epithelial growths in the skin of etydas and they Pope (1939) reported that a green turtle from discuss the microscopio structure,The growths the Pacific Ocean lived for 15 years in the New in wild turtles occur on the neck, eyelids, York Aquarium. axillae and groin. 3.32Hardinees:(see sections 3.13, 3.16, Nothing is known of epizootios among green 3,4) turtles if, indeed, there are any. 3,33Competitors Some individuals may get hopelessly wedged between rocks in 'their approach to nesting The only known vertebrate competitors of any beaches and thereby perish (Oa,rr and Hirth, importance are dugongs, manatees and some herbi- 1962). vorous fish (see section 3.42) 3.4 Nutrition and growth 3.34Predators 3.41Feeding and behaviour on the Sharks prey upon large green turtles feeding pastures (MDorhouse, 1933; Tinker, 1941; Travis, 1959, 1967; Hendrickson, 1969; Hirth and Carr, 1970),About 4% Most adults, and to some extent sub-adults of adult females observed on Nalaysian nesting feed chiefly on their grazing pastures (see beaches showed evidences of shark damage section 3.42), although in some regions of the (Hendrickson, 1958).Groupers are capable of world turtles may feed en route between breeding taking individuals up to 4.5 kg (Tinker, 1941), A and feeding sites,As far as is known, no 22.7 kg green turtle was found inside the stomach feeding, or very little, takes place off the of a tiger shark caught near Clarion Island (Beebe, nesting grounds. 1937). The feeding habits of juveniles are not Large terrestrial mammals such as ferai dogs, well-known, although it is commonly believed tigers and jaguars may take a few adult females as that they are carnivorous for their first few they deposit their eggs on beaches, but such pro- months of 'life, dation is probably minimal, It is assumed -that some time after the 3.35Parasites, diseases, injuries and turtle passes one year of age or when it weighs abnorirali-ties between I and 4 kg, it becomes mainly herbivorous (see section 3.42).Whether this dietary shift Ingle and Smith (1949) provide a list of is gradual or seasonal is unknown,This change references pertaining to the parasites of marine in' food preference is not uncommon, since it has turtles,Their list includes: Distonuan constricturn,been reported in other reptiles, especially Ozobranchus branchiatus, Stornato lepas transversa, fresh-water turtles and lizards, iThytidodoides intestinalia, R, similis and Platylepas loe,Nora recently, other Sizes of grazing males and females on the investigators have reported on parasites.Carl feeding pastures in the Gulf of Men are given (1955) recorded barnacles (Balanus crena.tus) on the in Table IX,As is to be expected, the female carapace of a sub-adult green turtle found on the 'feeding population includes smallér individuale coast of British Columbia,Hendrickson (1958) than the nesting population. records barnacles (Chelonibia testudinaria, Ste hanole as muricat) and leechea (Ozobronchus Cari' and Carr (1970a) suggest that condi- branchiatuson turtles in the South China Sea. tians on the feeding ground may affect the inter- He aleo observed mosquitoes biting the soft akin val between reproductive cycles (see- section 3,15), P[R!4 58Green turtle

TABLE IX Sizes (in centimetres) of green -Lurtlee caught on the feeding pas-hures near Khor Ulnaira, Southern Ymhen (from Mirth and Carre 1970)..

1JLE$, n 178 MALES, 11=112 Menge Mean Range

Carapace length 48,3 - 111,8 90,4 71,1 - 104,1 Carapace width 35,6 -.88,9 68,6 50,8 -81,3 Plastron length 38.1 -86,4 72,9 58,4 -81.3 Head width 8..9 -17.8 14,2 10,2 -17,8

Travis (1967) observed green turtles on -theirbest turtle grass pastures aro within this unit. feeding pastures off the east coast of Somalia end discovered that they "slept" in 30 to 50 minute A brief discussion of some of -the physiolo- bouts usually wedged under natural overhangs..He gical problems encountered by diving turtles is also desci ibes catching sea turtles with remoras. discussed in Gans and Gaunt (1969).In laboratory tests Berkson (1966) was able to show that Carr and Ogren (1960) n-tate that the green metabolic rates are much reduced in prolonged turtles ir. the Caribbean usually spend the night ondiving and -that the green turtle must be very the bottom, or among rock or coral crags, and that resistant to asphyxiai distress.Later, Borkson uitahle sleeping sites may be located some distance(1967) found that when the green turtle is away from their feeding pasture. subjected to hydrostatic pressure a-s. groat as 19 atm., as long as nine minutes may e).apse between Parrish (1958) observed that, in captivity, a heart beats.He also measured nitrogen tension resting turtle curfacee far air every 10-56 ninutes..in carotid artery blood and in tracheal air and Active turtles surface every30secondsta ton found that equilibrium conditions aro never minutes.Surfacing for air at night usually occursattained during a prolonged deep dive, but that about every 35-45 minutes.Green turtles slept enough nitrogen enters -the blood io render the resting on the bottom of the seaquariuxn and sleepinggreen turtle susceptible to gas emboli in the began at sundown and lasted all night,Walker bran after emergence. (1959) observed that nostrils of green turtles are olosed when they rest in an aquarium and. he suggests MoCutcheon (1947) reported that the specific -that the closure is an adaptation to prevent wateroxygen affinity of an adult green turtle is 19,0, from éntering or air leaving the respiratory compared to 28.5 for that of a loggerhead and to passages when the animal is resting.. 12.0 for that of a box turtle (Terrapene).The differences in the unloading capacity of oxygen Observations were made on several large G.m. are correlated with behaviour and habitat. carrinegra in captivity (Caldwell and Caldwell, 1962) and it was recorded that they slept on the bottom of the aquarium with the eyes tightly shut 3.42Food and feeding associa-tes during the night, but nighttime sleep was interrupted by fairly frequent (at least once an hour) trips to Some food items of adult green turtles are the surface to breathe.Presumably the sarao listed in Table X,As the list indicates, adult behaviour takes place in nature on the feeding turtles are mainly herbivorous bu-t they aro not pastures.However, turtles have been reported averse to esting animals.In fact, captive feeding at night in the grazing pastures in-the Gulfturtles are commonly fed animal food (section of Men (Mirth and Cai-r, 1970). 3.43). Sleeping habits of the hatchlings are quite Other Investigators have reported on -the different.They usually float on the surface with feeding habits of the green turtle but their their front flippers drawn in along each side of theaccounts are -of a general nature or they have not carapace.Some float with their front flippers given the exact feeding site or they have extended at right angles to the carapace. reported on only one or t» individuale.Theeò works follow: Zostera along the Atlantic coast Carr (1967e) postulates that green turtles (Kingsley, 1885); algae end Zostera (Gadow, 1909); rarely dive deeper than 12 fm. Mostofthe q,vinoctocoa, Thalassia, Zostera and Halophila 16 FIR1 58Gree12.turle

TABLE X Food of Mature Green Turtles

Locality Food Authority

LTHN llEMISPtERE Bermuda Zost era Babcook, 1937 Florida Vallianeria, 1eill, 1958 Thalassia, (ymodocea, Cari' and Caldwell, jellyfish, 1956 West Indies Thalassia, molluscs, crust acea Underwood, 1951 Gulf Coast of Mexico Cymodocea, Thalassia Mexico, 1966 Monquito Cay, icaragua Gymodocea, Thalassia Cari', 1954 Ciaras Brazil algae (Rhodophyoeae, 0hlorophyceae Phaeophyceae), Deplanther, molluscs, osoidians, sponges, bryozosais, crustaceans, echinodorms Ferreira, 1968 Pacific Coast of Mexico algae (GclidiuxnSargassun, Rhoc1?menia, Gracilaria, Cari' 1961 Griffitsia, LiaRon, tJlva Anon, 1966 Grat elon,,), odocea H. Marquez, Perse Thalascia COITUS0 Chile Zostera?, alga, sponges Vanes, 1951 DonosoBarros, 1966 Galapagos algae mangrove root e and shoot s F. Pritchard (MS) Maui, Hawaii green algae Mirth (MS) Tonga Is. Halodule, Halophila, yringodian Hirisli (MS) Fiji Is. red and green algae, Syringodiun Mirth (MS) Kextnadeo Is, Pterocladia capillacea Oliver, 1910

EAZ'1MHI HEMISFItSE

Seychelles Is, odo Horiell, 1927 Gulf of Aden Posidonia (or Enhalus), 1aloaule, brown and red Mirth and Carr, aigLe 1970 Kruaadai Is. algae (Gracillaria, Sargas sun), Cymodocea Kuriyan, 1950 Ceylon ymodocea, Thalassia, Doraaiiyagala, Halophila, algae 1939, 1953 Sulú Sea green algue and Sargasaum Domazitay, 1952-53 Grest Barrier fleef sea grasses Yonge, 1930 1?IRN/585 Green turtle 3:17 (Ingle and Smith, 1949); plant matter, crabs, ascidisne end barnacles in gut of one individual taken off South Africa (Hughes, Pass and Mentis, 1967); algae in the South Seas (Roedelberger and Groschoff, 1967); eelgrauealgae, crabe and snails (Schwartz, 19673; and, animal food such as sponges, Pteropods and jellyfish (Brongersma, 1967). Pieces of plastic have been found in the stomachs of turtles taken in the South Pacific (Hirth, MS) and in turtles captured off the Pacific coast of Costa Rica (Carr, pere, comm.). One of the best described feeding pastures is found near Thor Umaira, Southern Yemen (FAo, 1968; Hirth and Carr, 1970).Here the densitis of turtle grass reach 2,500 leaves per m The stomach of six mature turtles packed solidly with two species ofturtle grassweighed between Fig. 10Underwater photo of Syringodium 1.8 and 2,5 kg.Unfortunately, information on isoetifoliurn, a marine angiosperm, near assimilation is lacking.Ricker (1969) estimates the Fiji Islands.This is a favourite that the K value for marine herbivores would food, of herbivorous sub-adult and adult rarely exceed i%. K is defined as the growth green turtles in some areas of the coefficient, or the trles annual increment of Southwest Pacific Ocean weight divided by the quantity of food it ham con seized, pastures, and competition between these groups In terms of primary organic production, the Is probably minimal,In the West Indies, Western Indian Ocean is a region of contrast, Randall (1965) found parrotfishes (Scarus and containing some of the richest and some of the nestSparisoma), surgeon'ish (Acanthurus37'Ïinoids infertile waters in the world,Although measure (techinus, Tripneinstes and Diadema), and the mente have not been made along the Somali coast green conch (Strombus gigas) competing with the where good green turtle pastures are found, the greca turtle for marine grasses (Thalassia and region has been assigned a moderately high level ymoìocea).Stomach analyses of balahoo fishes of production (.26,50 gCm2 daj) and it is Hemiraniphus brasilienais, in the West Indies possible that much higher levels occur (Rydher et showad their main food to be leaves of Thalassia al,, 1966),Primary and secondary productivity (Bu:rkliolder, llurkholcler and Rivero, 1959). studies are greatly needed, especially on the Marner and Randall (1952) wrote about the in- feeding grounds, bat it may be that feeding vertebrate life in a turtle grass area around an experiments in captivity are the only practical atoll in the Gilbert Islands and found that even way to elucidate enerr flow in the relatively invertebrates were scarce. According to Wiens simple mun-grass--turtle food chain, (1962) the turtle grass (Thalassia) around some of the Pacific atolls rarely grows over 0.3 s Barkholder, Burkholder and Rivero (1959) high hut it makes dense stands over surfaces of obtained some quantitative and qualitative data on sand in which itcreeping rhizomes form an turtle grass, Thalassia testudinurn, in the West interwoven mat. He also found that turtle grass Indies. Among other things, they reported a bio- may be the dominant vegetation in lagoons from mass (dry weight) of 5.66 kg/m2, of which 80 to zero tide level to a depth of 1.6 fm. 90% was underground; and, that the leaves contained about 13% protein, 25% ash, 36% carbohydrates, 3.43Growth rate 16% crude fibre and 0,5% fat,Nere studies like this are needed, Rates of growth under natural conditions are relatively un1nown, although estimations C. den Hartog (1970) has written an interest- have appeared in the published literature,As Ing monograph on the sea grasses of the world,In long ago as 1916, Schmidt stated in regard to many Instances the distribution of green turtles the Paribbean turtle fisheries that "the most coincides with the distribution of sea grasses, important facts to be determined are the rate and. certainly a century ago the fit was even of growth of turtles and their migrations". closer (see Rig. lo), While some good information on movements of turt:Los in some parts of the Caribbean is now Marine turtles and sirenians are probably theavailable, there is still a dearth ogood only big vertebrates graz±ng the extensive marine empirical data on growth rates.Duringthe 3W FIRM $8Green turtle course of his pioneer studies on growth rates in of similar lengths in the South China Sea; and Caribbean turtles, Schmidt found that nine turtles while the weights of the larger females from initially weighing between 2.3 and 20 kg showed Yemen are similar to the Caribbean green turtles, weight increases of from 138 to 430 g per month they weigh somewhat less than the Atlantic green after 3.5 to 11 months in the ocean.His findings turtles on Ascension Island.A generalized also suggest that green turtles weighing aboix2.3, weight-length curve is provided in Fig. 7. 5.5 and 9.1 kg are about 1-1.5, 2-2.5 and 3-3.5 Generally speaking, the weight of a green turtle years old, respectively.He further postulated at reproductive maturity will be five thousand that a one-year-old green turtle is about 27 cm in times greater than its weight at hatching. length and a two-year-old is about 35 cm long. Worhouse (1933) estimated that one-year--old The weights of mature females in Surinais turtles are 20.3 cm long.Ingle andith (1949) with carapace lengths of from 100 to 122 cm state that growth rate in early years is about vary from 121 to 224 kg (Pritchard, 1969),The 3.2 kg or 17.8 cm in carapace length per year. average weight of a mature female from One-year-old turtles weigh between 1.8 and 3.6 kg Tortuguero, Costa Rica, is about 114 kand the according to Parsons (1962).'Carr (1967e) believes maximum is about 159 kg (Parsons, 1962). that turtles weigh between 0.9 and 2.3 kg when they are one-year-old.Australian green turtles weigh Caidwell (1962b) noted' a tendency for about 22 g at birth end about 1 200 g two to three males of C.m. carrinegra to be lighter than months later (Bustard, 1970a). females beginning at a carapace length of about 76,2 cm.He also described the relationship Cari- and Caidwell (1956) relate how an between carapace length (L) in inches and weight Atlantic green turtle weighed 20 kg at time of (w) in kg as follows: log W = -2.14 + 2.60 log L. capture and then weighed 2.7 kg more when it was recaptured 102 days later.Other gains were 0.03 As Fig. 6 indicates, the mean lengths of cm per day in carapace length and 0.01 ein per day nesting turtles in the Gulf of Men, China Sea in width of carapace. and Caribbean Sea are similar, while the Guiana and Ascension turtles are probably the world's Uithazn and Carr (1968) report that a yearling largest. released when weighing about 830 g was recaptured 30 months later some 65 nautical miles from the The heaviest green turtle reported in the release site and weighed 6,350 g.This recapture literature weighed 386.4 kg (Carr, 1952).The was of further significance because the individual heaviest reported in recent years was a 241 kg was a pen-reared turtle and the fact that it was female on Ascension (Carr and Hirth, 1962). recaptured is some evidence that pen-reared animals may be able to adapt to the normal ecologic regimen Almost half of the wet weight of a green of the species after a year in captivity. turtle represents edible protein.Deraniyagala (1939) and Schroeder (1966) respectively state Hendrickson (1958) postulated that there is that approximately 41." and 40% of the live no marked decrease in rate of growth until the weight is edible meat. animal reaches maturity and that maturity is attained in four to six years.On the other hand, Ingle and Smith (1949) found that the Caldwell (1962e) theorized that there is a marked flippers and meat of a 119 kg male and a 96 kg decrease in growth rate after about the first three female constituted 47 and 52% of the total years and that tropical turtles require at least weight respectively.The rest of the weight of eight years to reach maturity.i.kre recently Cari- the female was distributed as follows: head, and Goodman (1970) have ahown that after reaching 2.4%; tail, 0.24%; neck, 1,7%; kidney, 0.23%; reproductive maturity, female green turtles at liver, 2.3%; heart, 0.24%; calipee, 6.7%; Tortuguero, Costa Rica grow only about 0.25 cm per carapace, 15.5%; and entrails, 10.5%.The year in carapace length and width (see Fig. ii). weights of 45 and 68 kg females were also According to Carr and Goodman, "It now appears that distributed in a similar fashion. some green turtles mature at small, and others at large sizes, and that once they mature-that is, A hatchling from the Maldive Islands, fed once they have made their first trip to the nestingon fish and bread, weighed 4.4 kg and had a beach- their growth becomes negligible". carapace length of 320 mm and a carapace width of 270 mm after about13monthsin These data suggest that the growth of green captivity (Deraniyagala, 1939).Gibson-Hill turtles is similar to most other animals, i.e., (1950) reports that the Malay on the Cocos characterized by a sinoid growth curve. Islands raise green turtles in captivity and that they reach a length of about 25 cm in one Working in the Gulf of Men, Birth and Cal-r year.Hatchlings kept in semi-captivity in a (1970) found no significant differences between lag000ri in the South Pacific had carapace weights of females and males of the saine length. lengths of from 15 to 20 cm after ten months The same authors compared the weights of turtles a diet of kitohen scraps, fish and thellfih from different populations and found that female (Wiens, 1962),Three green turtles raised in an turtles from Southern Yemen are heavier than those aquarium grew from about half a kg to about American

Herpetologists) and the Ichthyologists of Society by

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Copyriglrt, 1970;

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Goodman, and Cari' (from

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Numbers

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Rica. Costa at ortuguoro, -turtles green female, circles and Squares

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I Fig. INCHES .ø . I IN GROWTH _J - turtle Green jJ$85 3:20 IIH14 S85 Green turtle 23 kg in nine and a half years (í'lowor, 1931). from the Indo-Pacific region.The origin of Caidwell (1962c) reported the weight increases of turtle poison is unknown but most investigators four hatchiinga maintained in captivity for three who have studied the problem appear to be rather years as follows: the mean weight increase was consistent in their opinion that the toxin is 135 g the first year, 2,725 g the second yeax, derived from poisonous marine algae eaten by and 5 000 g the third year.In captivity, turtles (see Haistead, 1970 for review of hatchl.ing green turtles can grow up -to between literature; also Boulanger, 1890; Halsteasl, 22.5 and 25 cm in six months according to 1959, 1969; and Taylor, 1970).The symptons of Schroeder(1966).Evidently green turtles in chelonitoxication var-y with the amount Q1' flesh oaptivity readily eat fish and other meat (Bee ingested and with the person.Symptoms generally also Hornell, 1927; Lveridge, 1945; and Parrish, 1958). after eating the meat and initial symptoms include nausea, diarrhea, epigastric pain and People in the Tuamotu Archipelago have raised ver-ti'.There are no known antidotes for ha-tohlings on coconut meat and fish and they have cheloni-toxin.The treatment is symptomatic. obtained a length of 50 to 71 cm in 3 to 3- year. Further studies on chelonitoxication and iaheneen on various islands in the Kingdom of documentation of green turtle cases aro needed, Tonga sometimes keep adult turtles in kraals and feed thais on turtle grasawhichwashes up on the Aimnonia is the major end product in the beach (Birth, HZ), urine of mydaa as it is in marine teleo stesa fishes. The green turtle also excretes a large 3.44Metabolism amount of hippuric acid and this may be due to its herbivorous diet (Kha.lil, 1947). Appa Rao and Dutt (1965) examined -the chemical compos&-tion of muscle of an adult male green An interesting adaptation of Chelonia tur-tlo,Results indicate that the water content to its marine environment, pointed out by of turtle flesh is close -to that of fishes (i.e.,. Schmidt-Nielsen (1959) among others, is its 80 g water per 100 g muscle),The fat content ability to discharge excess salt by means of (0.76 g per 100 g muscle) is lower -than in most salt glands located near the eyes.In thi fishes,In 100 g of muscle -they found 15.35 g regard the green turtle is similar to the marine. protein, 510 mg phosphorus, 210 mg calcium, 34 mg iguana and sea snakes.When on land, a turtle iron, 0,21 mg copper, 0.20 mg cobalt and 0.20 mg discharging salt looks as if it is "crying". mi,ybdenuin. Carr and Ogren (1960) suggested that -the "crying" of nesting females also serves to keep the eyes ChIefly because green turtle oil is usfor olean of sand. therapeutic purposes in some parts of Mexico (used in treatment of tuberculosis, leprosy, and Holmes and McBean (1964) have clearly shoan as a source of vitamins), Girai and Cascajares that the "salt glanda in mydas is -the predoTsinatt (1948, Girai, Giral and Girai (1948) and Girai route of sodium and potassium excretion,They (1955) have begun a study of its chemical further hypothesize that mea turtles drink una properties.Some of the mont common fatty acids water not only to obtain osmotically free water in the 01]. are laurio, myristic, palmitio, atearicbut also -Lo enable the excretion of. potassium and. paimitoleic.The presence of large amounts of ingested as food,Such information say be launa and syria-tic acids may be characteristic of important for fu-tune turtle farmers in providing Chelonia mydam.The sterol of body oil in a green -the proper milieu for their capbives, turtle from New Guinea contained 96% cholesterol, 3% B-sitoeterol, 0,2% campeaterol and 0.15% Ellis and. Abel (1964a, b) describe how the stJnasterol (Nina-to and Otomo, 1969), tun-bis' s salt gland tissue is in some ways different from the salt-secreting epithelia of Biochemical composition and nutrient quality fish and birds,Abel and Ellis (1966) state of the flesh may vary among individuals of a that histologically the modified. lacrymal gland population depending on such things as age and of marine turtles may be classified. as compound sex, s-tmge in the reproductive cycle, kinda of branched tubular glands and that the salt- vegetation eaten on the feeding pasture, and secreting nasal glands of marine birds, the physiological stress induced by long-range rectal. glands of elasmobrancha and the lacrymal movements, glands of marine turtles have many common cytochemical and cytological features. Evidently, -the flesh (or fat or blood) of some Chelonia is -toxic to humans,Halatead (1970) The marine plants that -the turtles graze gives a lie-b of references pertaining to ses on probably have a salt concentration similar turtle poIsoning.The list contains only one to that of sea water and no doubtthe salt cane of poisoning by green turtles and this gland of the tun-tic operates in response to incident caused 18 deaths in Ceylon in 1840.How- -this electrolytic load. ever, the list also contains other instances of cheloni-toxica-tion in which the species was um- Thorson (1968) measured the body water in unknown,Most reports of cheloriitoxication are sub-adult . riwdas and C,. agasøijLand 58Greenurtle determined its apportionment among the major fluid been 175 international recoveries (Cari', 1967o)d compartments.Both subspecies were similar in Mst of the recaptures have been made on the having total body water account for about 64% of feeding pastures off the Nicaraguan coast.One body weight. individual was recovered in Trinidad, some 2,400 kai from the site of tagging. Two Pbr some aspects of diving physiolo'-, see individuals marked on the Tortuguero beach nero section 3.41. later taken about 800 kin away in Colombia,It was calculated that the minimum speed of travel 3,5Poa-t-hatchling movements and, basking for one individual over a 34-day period was behaviour 23.5 km per day and that the second turtle travelled.aminintum of 21.6 km per day over a 3.51Developmental migrations 37-day period (Cai'x' and Ogren, 1960).It now appears that two main fleets converge upon Developmental migrations may be a natural Tortuguero to nest - one comes from the south part of the life cycle of the green turtle, (i.e., from Panama and Colombia) and the other although little is known about the subject. A from the Miskito Coya and Nicaraguan coast non-breeding population ranging in weight from 5 (Cari', 196m),One of the most wnazing facts to 52 kg appears on the feeding pastures off the pieced together from tagging work is that the upper west coast of Florida in April and. disappearsTortuguero turtles return to the same senent in November (Carr and Ca],dwell, 1956).This of beach to nest, after absences of from two to population is made up chiefly of sub-adults and a four years (see section 3.16). few juveniles,The Florida pastures may be a stop in the developmental migration of turtles which Some 556 turtles have been tagged while hatch on the rookeries of Mexico and. the Yucatan nesting on Ascension Island and ten have Peninsula.Sub-adults are also taken between subsequently been recovered on the feeding Maroh and November in the state of Sonora, Nexic. pastures off Brazil, a dispersal of some 2,240 Carr (1961) draws a parallel between thisSonorem km (Cari', 1967c).These turtleB also show a colony and the fleet of young individuals whiöb propensity to return to specific, small beaches appear off the west coast of Florida.Schmidt on Ascension for'nesting (Cari', 1965), (1916) may have been describing a developmental migration when he said ,"Whether the green turtle, ve females tagged on Musa and Sharma the species most commonly cau.t, breeds in the beaches in Southern Yemen have been recaptured Danish. West Indios, or whether it first arrives on the east coast of Somalia (Hirth and Cari', there as specimen20-30 cm in length is, 1970).Assuming that the turtles swam with strangely enough, unknown, and it is a matter of the prevailing surface currents, the longest great importance". recapture would constitute a movement of about 3,200 ion (i.e.,'rom Sharma to Chisimayo around An itinerant population of sub-adult green the Gulf of Aden),Minimum rate of travel for turtles is found near Bermuda and is probably two Yemeni green turtles was computed at 22.4 carried there by the Gulf Stream (Iwbray and ion per day for 85 days. Caldwell, 1958).A juvenile and sub-adult (lengths 22.5 and 42,5 cm) have been recovered Pritchard (1969) reported that one en on the beaches of Long Island, N.Y. (Latham, turtle tagged in Surinaxn was recovered in Brasil 1969), but apparently were waifs. after hav'ingtravelled adistance of about 1,600 ka in six weeks.Schulz (1969) also disclosed 3.52Dispersal and remigrations the recovery of two Surinam turtles in Brasil. I- thus appears that the turtle grass pasturee 1bst adult green turtles customarily make off the coast of Brazil are the feeding grounds seasonal trips between feeding pastures and breed- for two distinct nesting populations, viz., the ing grounds, and some of these movements may Ascension Island nesters and the Surinas nestere. encompass thousands of km (see section 2 and 3.1). Fig. 12 shows some places where adult females have Harrisson (1960) stated that a female been recovered after having been tagged on their tagged at Talang Besar, Sarawakin1953 was nesting beaohes,All the arrows in the figure do recovered in 1959 in a net off Kimanis, North not indicate location of feèding pastures, or Borneo,This repreaerrted a dispersal of about routes of travel, but they do indicate the 600km. magnitude of the spread from three major nesting beaches and thus give an indication of the "total Hendrickson (1969) reported that two greene range" of at least some individuals in the tugged on French Frigate Shoals were later rèoop.- population.The two best studied reznigration -tured in the main Hawaiian Islands about 800km patterns are those of the Tortuguero and Ascension ESE; and that two individuals marked at Pearl and Island colonies, Hermes Reef were res ighted on French Frigate Shoals -again a movement of about 800 lun ESF. From 4,200 mature green turtles that have been tagged at the Tortuguero rookery, there have Green turtles tagged on Heron Island, :22 1IhN 53 Greon turtle

3Q0

20°-

loo-

00

loo-

2Q0

3Q0

Fig. 12Longdistance recoveries of green turtles.Arrows indicate spread from the nesting beach and are not intended to suggest routes. A = tagging site was Tortuguero, Costa Rica (adapted from Carr,196Th);B tagging site was Musa and Sharzna Beaches, Southern Yemen (adapted from Hirth aM Carr,1970); C = tagging site was Ascension Island, South Atlantic (adapted from Carr, 1967b) B'I 58Green turtle Queenslandhave been recaptured in Papua and New South Equatorial Current are quite fertile, At Caledonia b3ustard, 1969c). places where the current diverges, there would also be more nutrient cycling through upwelling. Montoya (1969) found a general movement north According to Carr (1967b) it would. take a along the Mexican coast after oviposition. batchling about -three weeks to t to Brasil from Ascension. It is postulated that the green turtles nesting on the southern atolls of the Seychelles The Ascension-Brazil pattern may not ve Islaids remigrate there from feeding pastures off originated under present physiographical the East African coast and from the Mozambique conditions, but nay be an artifact of' continental Channel, althouthere have never been any tag drift. Th± idea is discussed by Carr (1967a). recoveries to substantiate th.is.,Large numbers Ii this connection, Myers (1967) has found that of green turtles, including 5uvenileo, are seen the fish evidnce siipports -the concept that the off the beaches in Tongaland, but they do not breed. final parting of Africa and South,nerica, and. there (Hughes, 19691,). hence the origin of' the South Atlantic Ocean, took place at some time during the earlier half In some of the earliest tagging experiments, of' the Mesozoic. Schmidt (1916) released 65 marked green turtles of various sizes around St. Thomas and St. Jan 5.n Carr (1965) considered celestial navìtion the West Indies and recaptured 14% from three to as a possiblo guidance mechanism in long-range eleven months later. st recoveries were made movements of the green turtle, but at least bi- just, a few kg from sites of release.The longest coordinate star navigation no longer seemso bc movement recorded was about 80 km and this a realistic hypothesis in view of the fact that individual was recaptured. -ten months after being Ehrenfeld and Koch (1967) found the Atlantic tagged.Schmidt concluded that young individuals green turtle to be extremely myopic when its are rather stationary. eyes are ou-t of water.However, other types of navigation involving the sun or moon have not Carr and Sweat (1969) report that a yearling yet bean critically evaluated or -tested. green turtle tagged and released in the lower Florida Keys on 17 October 1967 was retaken on 15 Several -types of orientation and navigation, November 1968 off Cape Hatteras, North Carolina. with special reference to the Ascension-Brazil remigration, are discussed by Carr (1967b). Oliver (1955) made 23 observations on an These include orientation based upon olfactory adult green turtleas it cruised leisurely in a cues, light-compass sense, maetic sense, large salt water stockade and he reported a perception of Coriolis force and inertial cruising speed of 1.4 to 2.2 km per hour,Various guidance.He also discusses some of the problems swimming patterns of captive turtles are further a green -turtle would have to overcome in order to described by Parrish (1958). navigate using celestial information. 3.53Navigation and. orientation It would be informative to be able to track e. "fleet" of traen turtles in their movement froe The ability of green turtles to navigate (= the coast of Brazil to Ascension - assuming that ability to initiate and. maintain directed movement they do travel in concert.Future research may independently of learned landmarks) across vast show that swimming together improves accuracy of expanses of featureless ocean is one of the ,noøt orientatìou, especially when the goal is an intriguing aspects of their biology. island.Hamilton (1967) has already discussed this idea in relationship to avisa remigra-tione Working with six loggerheads displaced off theand has postulated that orientation of a flock Florida coast, CarD (1962a) was able to show that of birds is more accurato than individual at least the initial movements of some individuals orientation; and. that flocking may be helpful were non-random,In the same paper, Carr discussesto juveniles making the -trip for the firat timo. the value of telemetry in unravel1in the mys-teries of open-sea navi-tion. Now that some older theories of non-visual navigstion have boon resurrected (for review, The evolution anc3 ramifications of the seo Griffin, 1969), it uld be worthwhile to Ascension-Brazil pattern are discussed by Carr and examine a green turtle' a sensitivity to subtle Hir-bh (1962) and Carr (1962b, 1964, 1965, 1967a), gsOphya&Ql and oceanographic cues, i.e., It is hypothesized that the hatchlings are carried forces whiih separately or synergistically could to Brasil by the South Equatorial Current and that.provide a bi-coordinate grid.A step in this for adults returning to the island. the ehorteat direction has been taken by Koch, CarE' anti route would be directly eastward from the hump of Ehreuf old (1969).They postulate that olfactory Brazil against the same current.Evidently food. cues emanating frote Ascension Island could be would be available for the surface-swimming recouized by turtles off the Brazilian co hatchlings, since according to Walford (1958), the arid thus play a part in the Ascension-Brazil surface waters of at least some branches of the romigration.Homing studies aro now underw5y on 3:24 FIRM/S85 Green tu

Fig. 13A radioequipped green turtle.Such turtics may be useful in tracking movements a-t sea.In one modification, the turtle tows a float containing the transmitter (photo courtesy A, Carr and Scientific fimerican; see Carr, 1965) 3 2

Ascenajon Island. Preliminary results indicate Several hypotheses have heen offered to non-random movements by a turtle out of sight of explain the banking habit anoag all kiods of fixed landmarks (Carr, pers. comm.). turtles, Ongle(1950),Neill and Alien(1954) and Boyer (1965) suggest that basking might be a Ilicigvay et al, (1969) studied sensitivity to form of thermo-regulation, aid ecdypis, free the mouths in three sub-adult green turtles and. they turtle of ectoparasites or elimlnate epizoio found maximum sensitivity in the region of 300 to algae0 Pritchard and Greenhood (1968) suggest 400 Hz, They conclude that the oar is clearly a that basking aids in the synthesis of Vitamin D. low-frequency receptor with a useful span of The habit may also aid digestion or it may be a probably60to 1,000 Hz, and they further state prelude to oviposition (but in the Hawaiian that "both on land and in the sea, this ear no Leewards male green turtles bask too), In the doubt enables the animal to perceive many importent Gulf of Carpentaria female green turtles bank signals". Is it possible that green turtles possess during the mating season (Buntard, MS), It is some kind of sonar sense? also possible that banking serves a social function, It is most likely thatturtle navigation is a composite process employing different senses and In view of the ever-expanding human popula- based upon a multiplicity of cues, tion (and the quest for food) banking aggregations of green turtles are mow at a disadvantage in Preliminary attempts at tracking turtles for their struggle for survival, Banking has not short distances have been made with floats, been observed in Atlantic green. turtle populations. balloons end radio-telemetry (see Fig. 13). It If the habit wan not more widespread. in earlier mag well be that tracking turtles with earth- years, its occurrence today in sosie populations orbiting satellites will be the only way to follow and mot others may be just behavioural polymor- movements with any great precision. phism.

3,54Basking Green turtles, like all reptiles, are cota- therme, Hjrth(1962)found that the cloacal Some Pacific green turtles are known to bask temperatures cf adult, female, green turtles at or rest on shore (see Fig. 14). The basking habit Tortugilero, Costa Rica, upon emerging from the has been reported from such widely scattered sea for oviposition, were very similar to the Pacific localities as Islas Revillagigedo, Hawaiian sea water temperature (average cf turtles29,9C; Leeworda and. the Galpagos (Mellen,1925;Wetmore, range of sea water temperature27,5-28,5°C). 1925;Grant,1927;Carr,1952;Kenyon and Rice, 1959;Parsons,1962),

Fig. 14Basking green turtles in the Hawaiian Leewards (photo, courtesy E, Kridler), Only Pacific turtles are known to bask

1_Oreen turtle 4:1

4 POPULATION 4.2Abundance end deiiity (of popultioj 4.1Structure 4.21Average abundance 4.11 Sex ratio It is estimated that about 10,000 females make up the entire population nesting on the Natural sex ratios in green turtle populationsSarawak Turtle Islands (Banks, 1937;Parsons, are virtually unknown.No studies have been made 1962;Honegger, 1968). Schulz (1969) estimate which elucidate the sex ratios in clutches of -that there aro about 2,500 females in the entire hatchlings and virtually nothing is known about Surinais population. Carr (1969b) calculates sex ratios in the juvenile and sub-adult popula-. that about 6,000 females nest annually at -tiens.It is only when larger individuals are Tortu,guero. Hir-th and Carr (1970) estimate that caught on the feeding and breeding grounds that less than 1,000 females nest annually on Aldabr%. biologists are able to glean something of the sex Dr. Bustard estimates the total breeding female ratios and these data are not entirely accurate. population of the three common species (green, flatback and loggerhead) in Queensland waters to Hirth and Cs.rr (1970) provide evidence that be not less than 75,000 (Anon, 1970g). females outnumber males on some of the feeding pastures off Southern Yemen.An overall sex ratio Relative sizes of other nesting popula-tiónW of 56% females and 44% males is based on a total can be ascertained by referring to references of 4,376 individuals collected over a period of given in Table II, seven years.The figures include sub-adults and adults. Cromie (1966) states that green turtles represent about 20% of all turtles found Calduell (1962b) found that in Baja Californiathroughout the warm seas of the world. waters, over four sampling periods in which the number of landed turtles varied from 104 to 465, 4.22Changes in abundance (see section females comprised between 68 and 92% of the catch. 5.4 for a discussion of the Presumably these were taken on the feeding pastures significant decreases in sizes of and were sub-adults and adults, turtle populations) Cax'r axai Giovannoli (1957) examined two 4.3Natality and recruitment oargoes of green turtles caught on the feeding pastures off Nicaragua between February and April, 4.31Reproduction rates end in one case there were 27 males and 66 females; in the second cargo were 105 males and 271 females, Details of egg production are provided in The sizes were not given, but all were probably seotion8 3,16, 3.17 and 3.2.Some of the sub-adults and adults. important aspects of reproduction pertaining to population studies are (i) a typical, adult Fxpert turtle fishermen in the Kingdom of female produces about 550 eggs each reproductive Tonga have advised Hirth (NS) that females out- season and. this usually is every two or three number males in the feeding pastures off Tongatapu.years, (2) some femaleamay not retain reproduc- tive capacity -throughout life, (3) at some major 11 is common to see more males than females, rookeries about half the eggs laid do no-t hatch, at least for part of the breeding seanon, off someand. (4) on high density nesting beaches some of the major nesting beaches,Reasons for this incubating eggs are destroyed by turtles nesting az's discussed in section 3.13. la-t er, 4.12Size composition 4.32Factors affecting reproduction: see sections 3 and 4.5 According to Galdwell and Caidwell (1969), the Iscarcity of records of small turtles seen in 4.33Recruitment (Some factors the ocean as compared with adult or sub-adult determining recruitment such as individuals is a most intriguing and puzzling growth and seasonal movements circumstance.While the apparent rarity maar result are discussed In sections 3.4 from not looking in the proper places, it may be and 3.5) that the lack of numbers is real.Considering the high mortality of the very young, the rapid growth 4.4ISrtalIty and morbidity of those that survive, and long life after reach- ing maturity, it is quite possible for the 4.41Nortality rates: see section population -to consist mostly of larger turtles". 3.22 and Fig.. 9 Data relative to the sizes of nesting females 4.42Factors causing or affecting are given in Fig. 6.Sizes of grazing males and mortality (Predation on eggs, females in the Gulf of Men are given in Table IX. young and adults has been F11U1JS85 Green turtle asoil in sections 3.11, 3.22 reptiles wefl adapted to their aquatic habitat, and 3,34,The direct and indirest but who periodically crawl onto dry land to lay effects of f jailing are discussed their eggs.At such times, when on their nest- in section 5) ing beach, they may be the dominant nocturnal vertebrate of the strand association.Their 4.43Factors affecting morbidity (Some crawling and digging activities at least tempo- parasites are listed in section rarily affect the beach topography and their 3.35 and chelonitoxication is eggs and hatohlinga are a source of food for a discussed in section 3.44) variety of predators (see sections 3.21 and 3.22). 4.5Dynamics of populations (as a who]e) The distribution of adult groen turtles in Bustard and. Tognetti (1969) have produced a the littoral and sublittoral zones of tropical model to show how nest destruction is dependent On seas is directly re].aied to their nesting population density and how this provides a mecha- beaches, feeding pastures and perhaps ocean ui.nm to regulate population size (see'ig. 15). currents.Long-ren:biennial or triennial The mechanism tends to keep a population within nesting remigrations across deep water are certain limits,They state that "at a low popu- typical for at leási some members of some popu- lation density its effects are negligible; however, lations, i.e., the population of green turtles if a population increased greatly, it would not in any given region may include two components, stay ai the new levo1 due ohia mec1iansm, which a migratory group, and a group that remains in tends to restoro the population to its original the area year-round and nests on the nearest level."They also postulate that the Barrier optimal beach (mee section 3.5). Reef region of Australia may be the last placo whore natural regulation of population size by the The relationship of green turtles With mechanism of density-dependent neat destruction competitors in the "turtle grass community" is can still be observed today. This is becaUse on dicused in section 3.42.most adults occupy many other nesting beaches the species w been the second trophic level in a simple predator seriously ovorexploited and the population density food chain, viz., plant-turtle.Hatchiinga and is below the level at which nest destruction by maall juveniles are chiefly carnivorous and turtles would operate as a regulatory mechanism, hence occupy at least the third trophic level in a predator food chain, viz., green plant- Bustard and. Tognetti's model would also seem herbivore-hatchling. to be operational for such sites as the Sarawak Turtle Islands where many eg, if not dug by man, The green turtle occupies a broad ecolo- would be destroyed by turtles nesting later. gical niche.This is partly due to the lack of' strong interspecific competition. 4.6 The poLatiOn in the communi tem Green turtles are large, egg-laying, marine FIRM/335 Green turtle 4:3

500 lOO 200 300 400 TURTLE POPULATION

Fig. 15Ì&del showing correlation of nest destruction with size of turtle population (from Bustard and Tognetti,1969;Copyright,1969,by the American Association for the Advancement of Science)

FI- SB Green turtle

5 EXPLOITATION South Pacific Islands. 5.1Fishing eQuipment Harpoons are widely usad to capture both sexes when they conregate off the nesting 5.11Gears and methods grounds.McCarthy (1955) describes, in a popu- lar way, how aboriginale harpoon turtles in the Indigenous methods of turtle fishing include Gulf of Carpentaria. diving for them and using harpoons, spearguns, traps, seines, auckerfish and decoys. Schmidt (1916) described how the fishermen in the Dutch West Indies used wooden decoys in Green turtles are usually caught on their the shape of a turtle, along with nets, to feeding pastures by- using nets and seines.Car-r capture green turtles.Wooden decoys are still and Caidwell (1956) describe how turtle fishermen used in several areas of Central America.They seeking juvenile and sub-adult individuals on the are especially useful in attracting males. grass pastures off the Gulf coast of Florida use tangle nets from about 97 to 194 m long, 2.4 to On the nesting grounds adult females aro 3 n deep, and with a 20.3 to 30.5 cm bar mesh. simply "turned on their backs",This can be done by one man,No special equipment is needed Duncan (1944) describes how Carman Islanders to locate and dig up the eggs on the beaches, capture turtles w-ith nets on the feeding pastures although a straight stick to probe for the e off Nicaragua, and Parsons (1962) further describes is useful. how turtlers on the Nicaraguan pastures "set their nets over marked rocks to which the turtles Mechanized turtle hunting is becoming more customarily retire at night and wait for them to widespread.Two companies operate in the feed- become entangled in the nets when they surface to ing pastures off western Australia and the blow".Huxley (1962) gives a very brief account following account describes their methods of turtling by the Cayman Islanders. (Anon, 1969e): "Each licensed freezer boat has several anali 16-ft scooter catcher boats, The beach seines used by the Yexneni in the powered by 40 hp outboard motors,These scooter Gulf of Men to catch sub-adults and adults vary boats operate within a one-mile radius of the from about 97 to 388 m in length and are up to mother freezer boat, in the relatively shallow 12 m deep with a 10 to 46 cm mesh (Hirth and Carr, water inside the offabare reefs, where the 1970).According to Tressler and Lemon (1951) the turtles graze on the brown and green algae of most popular form of gear for catching turtles is the rocky sea bed.When a turtle is located, the gill net, it is harpooned from the scooter boats as it races through water from 3 to 8 ft deep. On Vasa and Straatmans (1954) describe turtle attaining a full load of about 10 turtles, the netting on the pastures in the Tonga Islands as scooter boats unload their catch on board the follows: "A fisherman, having located an area of freezer boat fo:processing,Turtles are gutted, seaweed between the small, uninhabited islands, beheadeit, washed, drained and blast frozen, drops his net in a straight line at right angle to Each carcass weighs about 120 lbs dressed. When the flowing current and returns home.After two the freezer boal; attains a full load, usually days he goes back to retrieve his net and usually about 300 turtLes taken in about three days' finds turtles of all sizes caught in the not.In fishing, it returns t.o port to unload its a 50 by 12 ftnet, as many as ten turtles can be catch". expected and often more, depending upon the length and width of the net and the area where it is 5,12Boats: see preceding section dropped". 5.2Fishing areas Yonge (1930) has described how natives used to dive for sleeping turtles off the Great Barrier 5.21General geographic distribution Reef.Some of the ways of fishing for turtles in the Trust Territory are described in the Anthropo- Host green turtles are caught on their logical Working Papers (1957). feeding pastures, in shallow water just off the casting beaches, and on the nesting beaches Suckerfish or remera (Echeneis sp.) have been (see section 2). used to capture turtles in such widely scattered places as the South China Sea, northern Australia., 5,3Fishing seasons east coast of Africa and the Caribbean,Parsons (1962) gives an excellent account of this extrae- 5.31General pattern of seasons: see ordinary custom. sections 2 and 6 Scuba divers use Bpeargimsto get green 5.32Dates of beginning, peak and turtles of various sizes in such placesasthe end of seasons: see section 6 Seychelles Islands, Caribbean Islands, and some and Table XI 52 FIR1 E38 Green turtle

TABLE Xl Current regulations regarding the harvesting of green turtles and their eggs

Location Regulation

Pacific Ocean and adjacent areas Hawaiian Leewards complete protection for ail marine turtles and eggs (Anon, 1968b;Hendrickson, 1969;Laycock, 1970). 1iidwar Island protection for turtles less than 60 cm in carapace length. Trust Territory offull protection for turtles and, eggs on shore;and in the water for turtles the Pacific with carapace length lese than 61 orn (Wilson, 1969). Kingdom of Tonga following regulations now wait ing approval by Governmentcomplet e year-around protection for egand for turtles with carapace length more than 87.5 cm; protection for all turtles of all sizes from I Novmaber 'to 28 February;ban on the sale or export of anr turtle shell greater than 87.5 cm;complete protec- tion for the leatherback of all sizes at all tines, Fiji Islanda same regulations as for Tonga, awaiting Government approval. Rose Atoll, complete pLu(ection for turtles and eggs, Lu ericen Samoa Malarsia complete protection for nesting turtles;Government controls harvesting of eggs (about one million ennuall7);(arch is closed season for eggs in Sabah (Harrisson 1950_1969; Hendrickson, 1958;de Silva, 1968, 1969b). Queensland, full protection for all marine turtles end eggs (Bustard, 1969d), Australia Atlantic Ocean and adjacent areas Mexico full protection for eggapermit required to take turtles during open season; on Pacific ooaa-t closed season generally extends from 1 June through 31 October; on Gulf coast closed season extends from i May through 31 August, Costa Rica adults and eggs fully protected on nesting beaches;and protection for turtles within 4.8 km of nesting beaches (but harpooners can operate beyond this limit), Panama full protection for the green turtle (Myers, 1970), Surinam complete protection for nenti,ng 'turtles mid eggs on some of the major nesting beaches (see text), renoh Guiana protection for adults and eggs during May, June, July (Anon 19691; Pritchard, i969. Trinidad and protection for turtles and eggs from 1 June through 30 September, Tobago kscension Island full protection for turtles and eggs, Ipdian Ocean and adjacent areas British Indian complete protection for turtles end eggs (but see text). Ocean Territory and Seyoheiies Islands FIRM SB Green turtle 5:3 5.33Variation in date or duration of Parsons (1962) has documented the over- season: see Tables II, III and exploitation and demise of green turtles in section 6 various parts of the world, specifically in Bermuda, Dry Tortugas, the Caribbean Sea and in 5.4Fahinoperations and. ret],. the Indian Ocean,The decline of the green turtle population in the Seychelles has been Of the seven species of marine turtles, the commented on more recently by Honegger (1967), green -turtle is of the greatest economic value to Gayiner (1968) and. Hirth and. Carr (1970). man.Its flesh, eg:, and "calipee" (= cartila According to Acharji (1950), green turtles off used to maki soup) have been eaten for centuries. Ceylon are netted between November and March,- Now the growing market for ibs skin and oil, coup:Ledand the annual catoh is around 1,000, :ribhhe oven greater need for, animal protein especially in the tropics, places -this reptile in An estimate of the economica of the 'cen en especially important but precarious position., turtle fishery in some areas of the western Carr Ç1952) called the green turtle "the mosl hemisphere is provided by Ingle and Smi-.h (1949). valuable reptile in the world." Statistics show that about 2,000 turtles were oaugh-b annually in Costa Rica from about 1930 The numbers of turtles caught on -the feeding. bo 1948.Twenty-five men were en&ted in the pas-bures as well as the numbers of females taken fishery in 1948.These men caught turtles on from the nesting beaches have declined rapidly in the nesting beaches between June and. September. the current oen-bury.Parsons (1962) provides a The value of sea turtles exported from Costa comprehensive cultural and historical survey of Rica over a 15-year period. varied. from $9,161 man's attitudes toward the flesh arid eggs of the in 1937 to $900 in 1942. green turtle. Murphy (1925) reported that a turtle fish- In some countries bordering lhe South China ery was opera-ted from time to time out of Pisco, Sea, eggs rather than turtles are harvested.Banks Peru.Its existence today is unknown, (1937) reported that the Sarawak Turtle Islands yielded. 2 119 912 eggs in 1927 and theseh513a value Cald.well (1963) gives a historical summary of U.S. $3,O(0.Of these, 70% were consumed or of the turtle fishery in Baja California. sold locally,Ir: 1935, 98% of the 924,000 eggs Harpooning is the most common method of taking collected (value U.S. $7, 769) were consumed locally.turtles in that area and. the fishery operates year-round..Hubba (1960) also mentions the Harriason (1962b, 192d, 1965a, 196Th) has turtle fishery operating in that area. Townsend documented the decline of green turtles in Malaysia (1916) reported a catch of 162 green turtles in through an analysis of' egg-harvest records. Ho a single haul of a seine 90 m long in San points out that in the thirties a year with over Bartolome Bay in the Gulf of California on 11 two million eggs was commonplace, while in the April 1889.rfley were reported to nest in early and middle sixties a year with less than April and. Nay. three-quarters of a million (only 99,307 in 1966) was commonplace,He speculates that the reasons The average annual harvest of C.in. m,ydas for the decline to be both local (i.e., disturbance from the east coast of Mexico between 1963 and of nesting females; increased small boating 1967 was approximately 225 t (R, Marquez, pers. activity off nesting beaches) and. international comm.),The mean annual harvest of C.rn, (i.e., large steamers and. mechanized fishing fleets carrinegra off western Mexico between 1963 and in area; catching adults at sea). was about 5,300 t (R. Marquez, pers. come,), During the peak months of nesting on Langawan, The development and decline of the green one of the Philippine "Turtle Islands", about turtle fishery in the Gull' of Mexico, Caribbean 45,000 eggs are collected and sold each month arid British Honduras has been discussed by Ingle (Domantay, 1952-53).The same number of eggs, or arid Smith (1949), Westermann (1953), Carr (1954), more, are harvested on at least two other islands Smith (1954), Caldwelland. Cari' (1957), Cari' and in the group.Risher, Simon arid Vincent (1969) Ingle (1959), arid Neill and Allen (1959).Cari' estimate that one million eggs are collected (1954) stated that "more than any other dietary annually in the Philippines, factor, the green turtle supported the opening up of' the Caribbean". In recent years, two western Australia process-. in companies have undertaken commercial exploita- An indication of the former abundance of' ion of the green turtle off the coast of western green turtles around the Bahamas end Cuba is aralia (Anon, 1969c).The areas fished comprise given by Cateshy (1743). the territorial waters extending along the coast- line for approximately 160 km north from south The commercial catch of turtlea (almost all latitude 23°10'.In 1968 and 1967, respectively, green) In Hawaii in 1969 was 4,625 kg (value U.S. the export of processed turtle meat from western $2 320).These figures should be considered Australia was about 164,558 and. 135,922 kg. The minimal because many more are taken but not fishing season generally commences in June or July report ed.. and terminates in September or October. iThM/85 Green turtle

Between 1953 and. 1967 up to 262 green turtles reported catches of sea turtlos of all species have been sold annually in Papeete, Tahiti, increased from 20,000 metric tone in 1964 to36 and 33 thousand metric tons in1967 and 1968, The Eijian's fondness for green turtles la respectively. Whether these increases are real we lidocument ed in archaeological excava-t ions or due to better coverage in more recent years (Clifford,1951). is unknown. Japan and !4exioo together accounted for 95% of the landings (of ali species) in1968. Parsons (1962) estimated that between 15,000 The breakdown by species is not known, but greens and 20,000 green turtles are taken annually to ridleje and hawksbillø probably account for most supply coninercial markets. Reese(1917)mentioned of the landings. that in one year 15,000 were imported into England. All the available evidence indicates that Thecatches of marine turtles have inoreased green turtle populations are slow to recover from eigsificantly in recent years (FAO. 1970).The ovarexploitation. FIPJV'S85 Green turtle 6:1

6 PROTECTION AND MANAGEMENT nesting grounds in Surinani were declared a nature reserve in 1969 (includes Ellenti, 6,1Regulatory (legislative) measures Baboensanti, Pruimenboom and Galibi beaches), Lecal Caribs, however, are allowed. to take a 6.11Limitation or reduction of total certain quota of eggs each year. catch: see Table XI and following sections Indian Ocean and adjacent areas 6,12Protection of portions of popu.- Legislation recently passed. by the lation Seychelles and. British Governments (August, 1968) provides complete protection for both There are striking differences among various sexes and. eggs -throuì.out the Seychelles Islands countries regarding protection of green turtles. and the British Indian Ocean Territory.Today There are no regulations at all pertaining to their there are no big, concentrated nesting places capture on the high seas.Current regulations, in the Seychelles Islands, although in the last that were available to the author, are set out in century A.ldabra was one of the biggest green Table XI,Various other recommendations and the turtle nesting grounds in the world.The philosophy behind some of the ordinances are general ecolor of Aldabra and its potential discussed in the following regional sub-sections. role as a site for a biological research station (in contrast to its proposed role as a military Pacific Ocean and adjacent areas landing strip), have been discussed. by Stod.dart (1967, 1968a, 1968b) and Beamish (1970).Brief It should be noted. that the new Tonga and Fiji descriptions of the Aldabran rookery are resolutions (see Table JOE) will provide protection provided by Abbott (1893), Keynes (1959) and for all eggs and for animals above a certain size Hortmaz(1967).As in other places where the rather than below a certain size.The philosophy green turtle exhibits pronounced. migratory behind these ordinances is geared to the belief patterns, the Seychellois are concerned. over than an adult turtle remains reproductively active the fate of their turtles which now receive full for a considerable number of years and protection protection on Seychelles beaches but which of adults and their eggs should in time produce an migrate elsewhere to feed and apparently get no increase in the population.Turtle populations in protection (Anon, 1970d.).The same situation Fiji and Tonga should be monitored after these new led recently -to the Tripartite conference among laws go into effect since the ordinances may serve Panama, Conta Rica and Nicaragua (see preceding as models for other areas in the South Pacific. section). (in Tonga and Fiji the carapace is measured along its arch; straight line measurement iould place In and. near the Malagasy Republic, the legal size a few centimetres below 87.5 cm). instituted. turtla reserves are located at Nosy Mambo, Nosy Iraxtja, Nosy Trozona, Nosy Va, As far as is known, the eggs are generally Chesterfield. IsleZLd and. Europa Island.On some eaten in Thailand, indonesia, Burma, Borneo and other beaches in. the Nalagasy- Republic it is the southern Philippine Islands; adults are forbidd.en to take sea turtles wheù they are preferred in New Guinea and Melanesia (Penyapol, laying, 1958; Parsons, 1962; Harrisson, 1962d; Somadikarta, 1968; and Anon, 1969d). In Southern Yemen, green turtle flesh is mainly a commodity for export.In view of a Atlantic Ocean and adjacent areas growing export trade and possibility of Over- exploitation, the Government of Southern Yemen In September and October 1969, representativeswas advised. to limit the annual catch to 1 000, from Costa Rica, Nicaragua and Panama met in San of which not more than half were to be females; Jose and agreed to suspend all turtling for a and., to provide completo protection for nesting period of three years after which reappraisal of females and. eggs on the major rookeries in the the status of the resource would be made and eastern provinces (FAO, 1968).Whether or not appropriate international controls could be these recommendations have been adopted by the established (Anon, 1970a; Carr, 197Oa).If Government remains unknown, successful, this will be the first sigaificant international agreement on sea turtles, Although there are no marine turtle con-. servation laws in West Pakistan, the green About half of the big rookery at Tor-tuguero, turtle rookery at Hawke' s Bay seems in no Costa Rica will be enclosed in the new Costa Rica imminent danger because local religious customs National Park (Anon, 1968o; Anon, 1970h). forbid. eating of turtle flesh and very few indigents eat turtle eggs. Complete protection is provided for adults as well as for eggs on some of the major nesting InSeptember, 1968,-the new African Conven- beaches in Surinam (Pritchard, 1969).On other tion for Conservation of Nature and Natural beaches in the country, adults and their eggs are Resources was sigeed by 38 heads of state or protected only during certain seasons.The Galib:L their representatives.All marino turtles are 6:2 PIFLM/S85 Green turtle placed in "Class A" which is defined as follows: The establishment of breeding sanctuaries "Class A species shall be totally protected through-is certainly one way to perpetuate the stocks. out the entire territory of the contracting states, Ascension in the South Atlantic, Europa and end the hunting, killing, capture or collection of Aldabra in the Southwest Indian Ocean, Rose specimens shall be permitted only on the authoriza.- Atoll in the South Pacific and French Frigate tion in each case of the highest competent authority Shoal in the Hawaiian Archipelago are good and only if required in the national interest or examples of such breeding reserves. for scientific purposes",Ourry-Lindahi (1969) describes the events leading up to this African A series of marine national parks could conservation convention. also be established in key places where marine turtles are especially abundant.Kenya, for 6.13Current status of the resource and example, has two marine parks ai-id is considering action programmes a third (see Randall, 1969, for discussion of marine parks). The green turtle is currently listed as a "depleted species" in the I.U,C,N, Red Data Book The problems manifest in green turtle (Honegger, 1968).Consequently, at least for the management and conservation are indeed complex. present time, the keeping of green turtles as pets Factors that must be considered include turtle should be discouraged (cf. Murphy, 1968).Chelonia biolo-; local customs and religions; demo- nzydas is listed as a "peripheral" species in the graphic trends; economics; federal regulations; 1968 edition of Rare and Endangered Fish and Wild- and, perhaps most important of all, interna-. life of the Thi-ted States (Anon, 1968a), and is tiorial cooperation.The fact that fishing included on Ziswiler's (1967) list of the most jurisdiction claimed by nations varies from gravely threatened animals. about 5 km to 321 km complicates any rational plan for the management of any marine turtle. The green turtle (and all other marine turtles) was mentioned repeatedly in conservation resolutions 6.14Public education adopted at the Second International Congress of the World Wildlife F\ind which met in London in November Because education plays a large part in 1970. any conservation and management programme, there should be more pictorial accounts like Many pleas for the wise management and conser- Jacques Cousteau's three film loops on the vation of green turtles have been made.Some of reproductive biolor and conservation of the the more recent are those of Anon (1959), Hillaby green turtle (available from Oceanography (1965), Pritchard (1966), Carr (1967b, 1969d, 1970b), Unlimited, Lodi, New Jersey) and more films Bertram and Bertram (1968), Duniont (1969), Harrisson like -the exoellent turtle conservation filni (1969b), Honegger (1969) and Crowo (1970), produced in Sabak and shown on television around the world (Bill Burred Television Co., In 1933 Moorhouse emphasized that much has Los Angeles, California). still -to be learned about green turtles, especially the sex ratios in the nest, survival rates, where "Solomon, tue Sea Turtle" is a semi-. young spend their lives, and growth rates.Forty documentary film which portrays the homing years later biologists still know very little about prowess of a green turtle (see Anon, 1969e), these four facets in the life of the green turtle, all of which are needed in order to establish 6.2Control or alteration of physìcal rational management programmes. features of the environment Major action programmes recommended by a group The encroachment of vegetation and the of sea turtle conservationists meeting in M,rges, accumulation of debris on some nesting beaches Switzerland (10-13 March 1969) included (i) increasedhas been mentioned in section 3,16. incubationand hatchiing programmes, using proven techniques, (2) study and analysis of world A study of the physical parameters of good. exploitation patterns, (3) a broad information turtle pasture is needed (see sections 2.22 and programme, (4) beach surveys where data are lacking, 3.42). followed up by expert advice as required, and (5) establishment of special sanctuaries under Bustard and Greenham (1968) found that on scientific management.The recommendations received the Heron Island beaches no-t all egg chambers much publicity (for example, Anon, 1969k; Anon, dug are successful. Failures are caused by 1970b; and Anon, 1970c), obstruction of the digging action of the turtle 1)7 large or thickly matted tree roots and by It is this author's opinion that in addition collapse of the egg chamber sides due to to the recommendations given above, major research insufficient support of the sand. thrusts should be directed-toward population dynamica, productivity and studies on the ontogeny of orientation, FIRJ'/S85 Green turtle 6:1 6. Control or alteration of chemical chiefly because of the increased hatohling yield. features of the environment At Toriuguero adults are taken 'for local consumption and eggs are seldom harvested. Aocord.into laboratory experiments of Bustaril and. Groenhaxn (1968), green turtle eggs are able to Although little is known regarding the hatch in chloride concentrations very much greater. numbers omales in a natural population, it may than those occurring in the natural egg chamber. be -that some of the hunting pressure on femaloc Chloride concentration is not, therefore, a could be relieved by cropping surplus maleo, limiting factor in incubation success except for those few nests which aro laid below the spring 6.5Artificial etockin hit tide mark.Bustard. and Greenham also declare that they have successfully incubated many turtle Mere than 100,000 ha.tchling green turtles eg in coral sand moistened only with distilled from Tor-tuguero have been distributed to 28 wat er. different localities in Central and South Morion, Mexico, Nest Indies, Bahamas, Florida end Texas 6.4 Control or alteration of biological by the Caribbean Conserva-tien Corporation (Dr, f satures of the environment Archie Carr, Technical Director) in hopes of replenishing turtle stocks.Mast of -the A variety of species management programmes releases are on sites where green turtles are in existence (see section 6.12). formerly nested in abundance.Batches of' hatchiinga are released with the hope that they Hendrickson (19589 1961) believes that go back to the place released. .-drawn there by exploitation of eggs has a much less adverse effectsome kind of sensory imprinting to 'baste, smell on turtle populations then slaughter of adult or feel.To date there are no indications of females and that the exploitation for eggs is more definite success; however, at some release sites profitable in terms of human nutrition than is it appears -that some of -the hatchuings are stay-. exploitation for flesh and fat,Hendrickson's ing on and. establishing resident populations strategr of harvesting turtle eggs rather than (Carr, 1967b).A pictorial account of the meat can be briefly summarized as follows: the restocking programme in the Caribbean is given adult female lias a high reproductive potential by Parsons (1962) and. Marquez (1966),Restock- and the species is adapted to sustain enormous ing programmes in Florida h-ave been discussed losses a-t the very early stages of it-s life.The in a popular fashion by Stephens (1968). harvesting of eggs constitutes exploitation at a stage when the species is adapted to withstand The "Brotherhood of -the Green Turtle" is a such losses.The individuals which do survive to loosely knit group whose efforts to s-ave the adulthood presumably remain reproductively active green turtle led to the founding of the Caribbean for a considerable number of' years,If the averageConservation Corporation. fesale experiences more than three breeding seasons in a lifetime, then the waight of protein available Turtle hatcheries also operate in Malaysia in the form of eggs would. exceed the weight and Australia.Harrisson (1950_1969) describes available in the form of flesh and. fat.The the long established hatcheries on the Ssrawak elegance of Hendrickson's schomeatleast in turtle islands,In 1968 more than 100,000 eggs Malaysia, revolves around the fact that it is also were dug up and reburied in the Sabeh hatchery. in harmony with local customs and religion. An egg hatchery has been in opera-tion for several years in Australia (&zs'tard and Greriham, Hendrickson and Alfred (1961) estimated that 1968), about two million eggs constituting something over 50 t of high grade protein were consumed each year The current practices are to release in Malaysia and -they also stated "preliminary ha-bchlings either on the beach or in the water studies have indicated that as little as 2 or 3% both in the day-time and at night,Some are of the nests, handled properly, could probably reared for several days or weeks and then provide sufficient recruitment to the wild popu- released in the sea just off the rookeries, lations to keep them at their present level, This latter procedure reduces mortality from provided other circumstances remain unchanged". littoral predators.However, what effects holding batchlings in captivity and then Since in some arena of the tropics about half releasing them at sea have upon their natural of the eggs which are laid do not hatch, it would behavioùr patterns are unknown. be of' immense value to devise a method of distingaishing fertile from nonfertile eggs, At the present time the best (and simplest) These nonfertile eggs could then be utilized for method of natural stocking is to provide human consumption. protection for mes-bing females, eggs and hatchlins on the rookeries.If predation on Caz'r (196Th) stated that afterover a decade eggs and/or hatchuings is high and cannot be of research at Tortuguero, Costa Rica, more green controlled, and in those few places ehero nesting turtles are nesting now than in previous years, density is so high that lato nesters destroy the 6:4 FIRN/585 Green turtle nests of their predecessors, then tile establishment or to release them from boats offshore, thereby of centralized egg hatcheries on the nesting beach eliminating littoral predation, this author is justified Hatchlinge produced in these favours the former because it most nearly hatcheries should be released as soon as possible approximates the natural behaviour of the animal. after emerging on the surface. Although, as previously mentioned, the methods of releasing The transplanting of eggs or hatelilings hatchllngs are debatable, i.e., whether to release outside their natural nesting areas should be them on the beach and let them crawl to the sea, discouraged until sorne evidence of success is obtained in the Caribbean restocking scheme. FIRM/S85 Green -turtle 7:1

7 TURTLE MARICULIPURE The feasibility of turtle ranching has been mentioned in a general way by Schroeder (1966). Oreen turtle ranching and farming are still Ehrenfeld (1970) postulates that turtle ranching in the pioneer stages of development.True turtle may have sorno promise if the sub-adults and farming implies that the unit is completely adults can be induced to stay on natural pastures independent of wilcistock (turtles live and mate near a farm without being penned. under controlled conditions and isy eggs on man- medo beaches),In turtle ranching the unit is ?'wbray (1965) reports that green turtles dependent upon ,ild popu:'.ationc for eggs end the mats regularly in the Waikiki Aquarium and that turtios arc kept in varying d.oeeo of oaptiv-lby eg'i are dropped in the water.He believes that (i.e.,, from freellving lndivduolo in feuoc1-.off they would deposit their egi on a suitable pcittu'oo to completo containment in oonoiotc poo:Lu) oartificial beach if one w provided. !Iost turtle biolorite believe the;t turtle ronchin, ii' aHiomptcd t'all, ohould be only o tempcu'nvy Paruouri (i 9ci2) ropox-te how ycn'ng South cohortio, the reaco» being that nalitu-al pop tiationo luiiericuu 'ivor ttu'tlou, Podoeuumiri ocrriiq, have iil1 not withritceid ooatiaucilpuaeitisatio&' cl' been lirunuplanbcclnio L'-ko Valenoiu in the Iiiü oge.Oar(1969a) pointu out another !lndoauhigulaiithi end arovowi1ig end clololieriouc effect of the p-cnature oprued of there,They uro »hic ubilicing artificial turtle noul'e and that io the rico of now rLesting bochocj that have been oonritruotcon merkotu und hih aioeu before the volume prodno.. uomo 0± the lobado of thu lake. tien n000euuy to oateíy demando and relievo prouutu'o on natuael popula-tionri io achieved0Necd. )ioiiiuniI. bueli: toapino have buon riurcd loso to coy, lilie ultimatu lu turtle inarisulturo io cn'0000riL'uhly in tue IloibIdtatoc miii methode riiug proven too}iuiqueri of agvi culture uced haro eiuy UOVO auaddou for cou turtles, cina animal huobenthy.iwlliial o.bbonlptc at turtle A brief modal l'or tur'apiu rouchuzig io Given by equacalliuro uhould be on a emuli coule and. should nglo cudmltui (1949). be conducted only if iioientifio expertisa io available0 borlcmr curjto. ulouu hava boon LLMCd for 17 yoaro in the Unitccl States (Han, 1970) Some of the foctoru whioli uuot be oonoidercd and. corno of the methode developed in l'arìing thri beforo initiating a turtle farei (or ranch) are the species might be adapted to oca tux't].oc, followings lon-torin financing, elimination of predators end compotibore in onclomtu'ss and on Pinchot (1970) disousoec the pocalbility of nesting beaches, protection from pollutants and. enriohing the lagoono of atolls and raising poachers, availability of veterinarians to advise various animal species therein,His ideas merit on disease problems, year-round availability of consideration and would be especially applioablo natural food (in nature, green turtle ha-bchlings in the South Pacifie reg-ion. are chiefly carnivorous and adults are herbivorous)., growth rates obtainable with commeroia:L feed, water It may even prove possible as well as and sanitary requirements, optimum density, economical to grow certain plants via hlroponics pollution abatement, transportation to processing for captive turtles,Several large European and plants, expertise on how to utilize all turtle American zoos have employed these methods to products (flesh, oil, skin and shell) and facilitiesproduce green grasc supplement for zoo animals. to conduct scientific experiment s concomitantly The zoo keepers believe that high nutrition with the commercial scheme. grasses may also stimulate reproductive processes and thus increase fecundity, Sholbourne (1970) describes some priorities am far as marine fish cultivation is concerned ami Preliminary experiments In Sarawak indicate the same would seem to apply to turtle culture. that the first six months are the most difficult In addition to some of the factors mentioned above,for rearing hatchiinge in captivity (Harrisson, he describes the provisions of "high quality" seed 1956a). stock and physi co-chemical requirement s. Significant pilot attempts at rearing Iversen (1968) describes some of the advantagesloggerhead turtles in captivity (and resulting of mariculture including the control of predation growth rates) have been described by Uehida and artificial selection for rapid growth, high (1967), fecundity and resistance to disease.He also discusses current techniques of milkfiah and. mullet A pilot turtle ranch has been established culture and mentions the green turtle as a candidateon Great Inagua Islzid in the Bahamas, under the for sea farming. ausp:Lces of the Caribbean Conservation Corporation. Here about 1,000 ha of turtle grass pasture have Some general principles of aquaculture along been walled in and hatchlings have grown up to with some of the problems and a few selective fish 3.6 kg in two years on a diet of conch (Gai-r, models have been discussed by Ba.rdaeh and ETther 196Th), (1968). *2 S8'5 Groen turtle

Five Email green tur-ble ranches ara noti in operation. Popular acoowita of Maricmlturo are operation in Parres Strait Australia under the provided by Gabier(1969),Anon(1969g)and direction of Dr, H.R, Bustard (Anon1970f Anon(1969j). Duatard, MS). According to scott(1970)there are several A pilot turUe ranch (or farm) is also major economic obstacles io the development of operating in Cuba. Details are lackin'. marine aquaculture, including (i) the absanoc of a strong demand for the high costproducta of At the present time a large turtle farn aquaculture (except for luxuries or freshwater (iculture, Ltd., with about 40,000 turtles of carp) in spite of the rising worldpopulation, various aizea) is in operation on Grand Cayman, and (2) the absence of property rights in natural Dritish West Indies. Initia], reportsindicate a watersand problems ofinternational jurisdiction hi.. degree of success in some aspeobs of the on the high seas. S so urtle -i1

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Schmidt, J, Marking experiments with turtles inthe Danish West Índiee. Nedd.Komia,Ho'vunders.(Ser. 1916 Fisk.), 5:1-26

Schmidt, H.P., Problems in the distribution of the marine turtles.Mar,Lif e 000aa.Pap,,(1)z7-1O 1945

A checklist of North American amphibians ' reptiles, Chi , University of Chi 1953 Press, 280p.

Sohmidt, H.P. and R.P. I.r, Living reptiles of theorld. New York, Doubl.,287p. 1957

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Schulz, 3.P.,, Zeeoohildpadden, deel II: Zeesehildpa&len in Suriname,Paromaribo, Dienst ' boabeheer 1964 Suriname, 44p.

National situation report re: Marine turtles in Surinais, IUC Publ Jew Ser.Suppl.Pap., 1969 (20):19-33 - Schwartz, P.3., The barnacle,Platylepashexast los, encrusting a green turtle, Chelonia-nydas 1960 from Chinooteague Bay, Maryla , Chesapeake Sci.,1(2):116-7 , Maryland turtles. University of Maryland, Natural Resources Institute,Publjo.79 1967

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Shannon, F.A., The reptiles and amphibians of Ko,.'ea,Ilerpetologica,12:22.-49 1956

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SYNOPSIS OF FISHERIES BIOLOGICAL DATA

This is one of a series of documents issued by FAO, CSIRO and USFWS concerning species and stocks of aquatic organisms of present or potential economic interest.The primary purpose of thisseriesis to make existing information readily available to fishery scientists according to a standard pattern, and by so doing also to draw attention togaps in knowledge.ltis hoped that synopses in this series will be useful to other scientists initiating investigations of the species concerned or of related ones, as a means of exchange of knowledge among those already working on the species, and as the basis for comparative study of fisheries resources.They will be brought up to date from time to time as further information becomes available either as revisions of the entire document or their specific chapters.

The relevantseries of documents are: FAO Fisheries Synopsis No. FR/S replacing, as from 1.1.63, FAO Fisheries Biology Synopsis No. FB/S CSIRO Fisheries Synopsis No. D FO/S and USFWS FAO Fisheries Synopsis No. NMFS/S

Synopses in these series are compiled according to a standard outline described in Fib/Si Rev. 1(1965). FAO, CSIRO and USFWS are working to secure the cooperation of other organizations and of individuai scien tists in drafting synopses on species about which they have knowledge, and welcome offers of help inthis task. Additions and corrections to synopses already issued will also be most welcome. Comments including suggestions for the expansion of the outline and requests for information should be addressed to the coordinators and editors of the issuing organizations. FAO: Fishery Resources Division Marine Biology and Environment Branch Food and Agriculture Organization of the United Nations Via delle Terme di Caracalia 00100 Rome, Italy USFWS: CSIRO: Chief, Scientific Publications Unit Scientific Editor National Marine Fisheries Service CSIRO Division of Fisheries and Oceanography Building 67, U.S. Naval Support Activity Box 21 Seattle, Washington 98115, U.S.A. Cronulla, N.S.W. 2230 Australia

Consolidadedlistsof species or groups covered by synopses issued to date or in preparation will be issued from time to time.Requests for copies of synopses should be addressed to the issuing organization.

The followingsynopses in this series have been issued since January 1970:

DFO/S4 Synopsis of biological data on the rainbow prawn, Parapenacopsis sculptiis (Heller, 1862) 1970 DFO/S5 Synopsisofbiologicaldataon the schoolprawn, Metapenaeus mac/say! (Haswell, 1879) 1970 BCF/S41 Synopsis of biological data on chum salmon Oncorhynchus keta (Walbaum) 1972 July 1970 FIRI/S80 Synopsis of biological data on the eel Anguilla anguilla (Linnaeus) 1758 October 1970 Rev. 1 * FIRM/S91 Synopsis of biological data on the common shrimp Crangon crangon (Linnaeus, 1758) October 1970 * FIRM/S92 Synopsis of biological data on the shrimp Panda/us montagui (Leach, 1814) October 1970 * FIRM/S93 Synopsis of biological data on the Jumbo tiger prawn Penaeus monodon Fabricius, 1798 October 1970 * FIRM/S94 Synopsis of biological data on the Indian prawn Penaeus indicus H. Mime Edwards, 1837 October 1970 * FIRM/S95 Synopsis of biological data on the prawn Panda/us platyceros Brandt, 1851 October 1970 * FIRM/S96 Synopsis of biological data on the penaeid prawn Solenocera indica Nataraj, 1945 October 1970 * FIRM/S97 Synopsis of biological data on the penaeid prawn Metapenaeus dobsoni (Miers, 1878) October 1970 * FIRM/S98 Synopsis of biological data on the penaeid prawn Metapenaeus affinis (H. Mime Edwards, 1837) October 1970 * FIRM/S99 Synopsis of biological data on the ocean shrimp Panda/us jordani Rathbun, 1902 October 1970 * FIRM/S100Sinopsis de datos bìológicaos sobre el camarón blanco Penaeus schmitti Bur- kenroad, 1936 October 1970 * FIRM/Si 01 Synopsis of biological data on the white shrimp Penaeus setiferus (Linnaeus, 1767) October 1970 '' FIRM/S102Synopsis of biological data on the brown shrimp Penaeus aztecas aztecus ives, 1891 October 1970 * F1RM/S103Synopsis of biological data on the pink shrimp Penaeus duorarum duorarum Burkenroad, 1939 October 1970 * FIRM/S104Synopsis of biological data on the penaeid prawn Metapenaeus tnonoceros (Fabricius, 1798) October 1970 * FIRM/Si05Synopsis ofbiologicaldataon the penaeid prawn Metapenaeus brevicornis (H. Milne Edwards, 1837) October 1970 * FIRM/5106 Synopsisofbiologicaldata on the penaeid prawn Parapenaeopsis stylifera (H. Milne Edwards, 1837) October 1970 * FIRM/S107Sinopsis del camarón nailon Heterocarpus reedi October 1970 Bahamonde, 1955

D FO/SS Synopsis of biological data on the greentail prawn Metapenaeus bennettae Racek and Dall, 1965 1970 DFO/S7 Synopsis of biological data on the eastern king prawn Penaeusplebejus Hess, 1865 1970 DFO/S8 Synopsis of biological data on the banana prawn Penaeus merguiensis de Man, 1888 1970 FIRM/S83 Synopsis ofbiologicaldata on Saccorhiza polyschides (Lightf.) Batt. October 1970 FIRM/S82 Synopsis of biological data on North Atlantic sandeels of the genus Ammodytes (A. tobianas, A. dub/us, A. americanas and A. marinus) November 1970 NMFS/S79 Synopsis of biological data on Pacific Ocean Perch, Sebastodes alutus (Gilbert), Cramer (1895) December 1970 FIRM/S38 Synopsis of biological data on knobbed wrack Ascop/iyllum nodosum (Linnaeus) Rev. 1 Le Jolis December 1970 FIRM/S84 Synopsis of biological data on haddock Melanogrammus aeg/efinus (Linnaeus, 1758) December 1971 FRm/587 Synopsis of biological data on Laminaria hyperborea (Gunnerus) Foslie December 1971 F1RM/S85 Synopsis of biological data on the green turtle Chelonia mydas (Linnaeus) 1758 December 1971

* Published in the Proceedings of the World Scientific Conference on the Biology and Culture of Shrimps and Prawns, Mexico, 12 to 21 June 1967.

Ht/C3466/I .72/E/I /900