(Distribution restricted) SAST - Green turtle - 5,31 (07) 005,02

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

Prepared byftF. Hirth

FOOD AND AGRICULTURE OGANIZATION OF THE UNITED NATIONSROME, 1971

FAO Fisheries Synopsis No. 85 FIRM/S85

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

FAO Fttherio Synopir4 No85 FThM/585jTJtBtribition 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. HirthDepartment of BiologyUniversity of UtabSalt Lake City, Utah

U.S,A.

FOOD AND AORICUTIIURE OHOANIZATION OF THE UNITED NATIONSRome, December 1971

PREPPRATION 0F THIS SO?SIS

The present document which is the first FAO Species Synopsis on biological data ofoommeroially important marino turtles, has bes prepared following the annotated outlinefor 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 ofturtle species.

The main purpose of this synopsis is to bring together the current information on thenatural history of the green turtle, Chelonia nydas, and to draw attention to the majorgaps 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 andMrs. Jearme Chapin, Life Science Librarian, University of Utah, for their help in trackingdown rare manuscripts; and to Dr. John Hendrickson (University of Arizona) for allowingthe author to peruse his personal library. Mrs. ICennie Harris, Mrs Bess Short and MiseLiwana Blau graciously and conscientiously typed several versions of the manuscript. TheUniversity of Utah Library and the Department of Biology financially aided the work throughthe purchase of inierlibrar'y loans.

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

Distribution

FAO Department of FisheriesFAO Regional Fisheries OffioorsRegional Fisheries Councils andCommissionsSelector SïAuthor

Bibliographic Hntry

Rirth, H,F, (1971)FAO Fish.Synop., (85):pag.var.Synopsis of biological date, on thegreen turtle Chelonia mydas (Linnaeus) 1758

Nomenclature, taxonomy, morphology,biometrios, tagging, distribution -horizontal, hybrids, ecology, lifehistory, nutrition, growth, behaviour,population dynamios, exploitation,fishery, legislation, habitht improvement,culture.

Gron kitrile

G O N T E N T S

2 DISTRIBUTION 2:1

2,1 Total area. i

2.2 Differential distribution I

2.21 Hatchlings, juveniles and sub-adults I2.22 Adulte 1

2.3 Determinants of distribution chenes 8

2.4 Hybridization 8

3 BIONO?&tCS A1 LIFE HISTORY 3: 1

3.1 Reproduction

311 Sexuality i3.12 Maturity 1

3.13 Mating 1

3.14 Fertilization 43.15 Gonads 43.16 Reproductive cycle, nesting behaviour and selection of nesting beach 43.17 Eggs

3.2 Embryonic and hatchiing phase 8

3.21 Embryonic phase 83.22 Hatehling period (behaviour, predators and survivorship) 9

3.3 Juvenile9 sub-adult and adult phase 14

3.31 Longevity 143.32 Hardineen 143,33 Competitors 143.34 Predrttort; 143.5 Paradies, 1ineasee, injures and abnormalities 14

1.1

IBENTITY

Nomenclatu.re

Pye No,

1:1

I

iI

1.111.12

Valid naneObjective aynonyTny

1.2 Thxononw i

1.21 Affinities i1.22 Taxonomie status, and definition of size categories I1.23 Subspecies 1

1 .24 Corsnon naInes 4

1.3 )brpholoy 4

1.31 Anatomy 41.32 Cy-tomorphology 61.33 Protein specificity 6

1.4 Measuring and ta1ng turtlo 6

IRM/S8' Green turtle

Page No.

3.4 Nutrition mid growth 3:14

3.41 Feeding and behaviour on the feeding pastures3.42 Food nnd feeding associates3.43 Growth rate3.44 Metabolism

3.51 Developmental migrations3.52 Dispersal and remigrations3.53 Navigation and orientation3.54 Basking

4 POPtJLA!FION

4.1 Structure

no information wae available to the author, this item has been omitted from the text

3.5 Posi-hatchiing movements and basking behaviour 21

4.11 Sex ratio 1

4.12 Size composition i

4.2 Abundance and density (of population)

4.21 Average abundance i4.22 Changes in abundance I

4.3 Natality and recruitment i

4.31 Reproduction rates i4.32 Factors affecting reproduction4.33 Recruitment i

4.4 Mrtality and morbidity

4.41 Mortality rates i4.42 Factors causing or affecting mortality 1

4.43 Factors affecting morbidity 24.44 Relation of morbidity to mortality rates*

4.5 Dynoiijcs of populations (as a whole) 2

4.6 The population in the community and ecosystem 2

5 XXPLOITATION 5:1

5.1 Fishing equipment 1

5.11 Gears and methods 1

5.12 Boats 1

5.2 Fishing areas

5.21 General geographic distribution 1

503 Fishing seasons

5.31 General pattern of seasons5.32 Dates of beginning, peak arid end of seanona 1

5.33 Variation in date or duration of season 3

5.4 Fishing operations and results 3

14151720

21212325

4:1

7 Pu1FLE MABICUI/PURE 71

8 EFEIENGES 8 * I

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' populatione6.13 Current status of the resource and tction progremme 2

6.14 Public education 2

6.2 Control or alteration of physical features of the environment 2

6.3 Control or alteration of chemical features of the environment 3

6.4 Control or alteration of biological features of the environment 3

6.5 Artificial stocking 3

FIR)/S83 Green tunjo

IDENTITY

11 Nomenclature

1.11 Valid naine

Cholonia ntlas (Linnaous) 1758

1.12 Objective 1nonynr

Testudo Linnaeus, 1758, p. 197,Syst. Nat. (Ed. io) Stockholm. Typelocality, Ascension Island..

Chelonia m.yd.as Brongniart, 1800, Bull, Sci.Soc. Philom., 2:89

Some authors consider the genericdesiiation by Brongniar-t a nomen nudun. Forexample, Deraniyagala (1939) recoiizes thefollowing:

Chelcnia rnydas Latreilie, 1802, Hist. Nat.Rept, i, p. 22

1.2 Taxono

1.21 Affinities

- Suprageneric

Kingdom AnirnaliaSubkingdom MetazoaPhylum ChordataSubphylum Vert ebrataSuperclass TetrapodaClass ReptiliaSubclass AnapsidaOrder Teatudina-baSuborder CryptodiraSuperfamily CholonioideaFamily Cheloniidae

Generic

Chelonia Brongeiart, Bull. Sci. Soc.Philom. Vol. 2, 1800, p. 89. Genotype: Testudonwdas Linnaeus.

The following generic identity is given byCar (1952): "One pair of prefrontal scales;tomium of lower jaw coarsely toothed, that ofthe upper jaw with strong ridges on its innersurface; lateral laminas, 4." Longer descriptionscan be found in Deraniyagala (1939), aith sindTaylor (1950) and Loveridge and. Williams (1957).

Specific

Chelonia mydas is easily differentiated.from its oongener, the flatback turtle,depressa Garmazi, by its more highly archedcarapace, sloping marginais and large scales inthe middle of the front flippers (Fig. i). Incontrast, C. depressa has a depressed carapaces

upiard-cu.rvthg marginais and mnali scales in themiddle of -the front flippers. Further diffe-rences between the two species are pointed outby Williams, Grandison and Carr (1967), Bus-tard(1969b), Bustard. and Liinpus (1969), and Coggerand Lindner (1969).

1.22 Taxonomie status and. definitionof size categories

The green turtle is a morpho-species.Because size categories of marine turtles are notstandardized, the four categories used in thisreport are defined, na follows: haichling stillbearing conspicuous umbilical scar; first fewweeks of post-hatching life; juvenile . umbilicalsoar inconspicuous or absent and carapace lengthup to 40 cm (41) cm is ábout one-half thecarapace length of a reproductively matureirid.iwidual); sub-adult post-juvenile but notyet reproductively mature (carapace length 40-to 80 cm); adult reproductively mature;carapace length greater thau 80 cm (the minimalsize al which females lay eggs is about 80 orn;for convenience it is assumed that males matureat about the sane time). Carapace lengths arestraiit-line distances. A "yearling" is a one-year-old turtle.

1.23 Subspecies

It is best to use -the binomial, jjniamydas, for all green turtles until a detailedtazonornic study is made. However, several sub-species have been described, and no doubt othórraces will be named in the future. In fact,the mydas complex may be one circuinglobal"Rassenkreia" bu-t with significant gaps betweenthe eastern Pacific and western Atian-tio--Caribbean populations and between the EastAfrican and West African populations. Sub-specific names have bean based largely oncoloration, al-thout Smii;h and Taylor (1950)state that a common arrangement restrictsndas to -the Atlantic and assizii to thePacific Ocean either as species or subspecies.At least sorne of -the subspecific terminolorhas been perpetuated in checklists and uide-books by Schinidt (1953), Conant et ai. Ç1956),Conant (1958), Fukada (1965), S-bobbins (1966)and Keiner and Wilson (1969), among others.

The Atlantic green turtle, Chelonia mydasndas (Linnaeus) is found in the Atlantic Ocean,Caribbean Sea, Gulf of Mecico nd the Hedi-terranean Sea, It has a predominantly brownishcarapace, sometimes with olive or dark brownblotches and streaks, and -the dorsal surfacesof the head., flìppera and tall are alsopredominantly brownish. The spaces betweenthe scales on the -top and sidec of the headare yellowish; the plastron is usually whitish.The shell of the malo is more elongate -than thatof the fornaio. A detailed. desoriptien of thesubspecies cari be found in Carr (1952).

i 2

I

FIRM/585 Green- urt1e

F1R14J385 Green turtle

Compared to the Caribbean population, adultfemales in the Ascension Island population have amore angular emargination of the shell over theneck (Carr and Hirih, 1962),

The East Pacific green turtle, Chelonia laaassizii Bocourt, is found, along the Pacificcoasts of Central and $outb America. The carapaceis basically greenish or olive-brown but sometimesstrongly flecked. with black, and. generally morehighly arched and narrower than that of anAtlantic green turtle. The carapace is sometimesmarkedly indented above the hind flippers-this isusually lacking in individuals from the Atlantic(Carr, 1952). Much earlier, Baur (1890) stated"there is no doubt whatever that the Cheloniafrom the Pacific is a different species from theAtlantic; the skulls are the same, but there aregreat differences in the form of the carapace".

One of the most easily recognizable forms inthe entire mydas complex is the NortheasternPacific gr en turtle, Chelonia myda carrinegraCaldwoll, This subspecies is characterized byblack pigmentation on both dorsal and ventralsurfaces. The carapace is highly arched in largefemales while it is low in large males. It occursin large numbers throughout the Gulf of Californiaand on the outer coast of Baja California.Cald.well (1962a) believes that all references togreen turtles off California and northward alongthe coast should be considered as indicatingearrinegra and he gives a detailed account of thissubspecies. However, the problem of distinguishingthis subspecies from black agassizii along theMexican and Guatemalan coasts still remains to besolved. If this cannot be done, the proper namefor the black turtle will be Chelonia nylasagassizii (Hirth and Carr, 1970).

According to local inhabitants, two fair:Lywell marked stocks of Chelonia (or a variablycoloured sinlepopulatïon) occur in the Galpagosarea (Carr, 1964). The turtle situation herewarrants immediate study.

Carr (1964) has noted some carrinegracharacteristics in sorno turtles in the HawaiianArchipelage. The concurrent distribution ofcarrinejrar-liice and mydas-like forms in theCentral Pacific may indicato a tendency towardpolymorphism within a single population (Hirth andCarr, 1970),

Chelonia nwìas ,japonica (Thunberg) is the namegiven to the Western Pacific green turtle (seo Carr,1942 for discussion). The type locality is Japan.Wex'muth and. Mertens (1961) give the range asPacific and Indian Oceans. Okada (1938) listsjaponica from Horisyn, Sikoku, ICyusya, BoninIslands, Okinawa and Taiwan. The Hawaiian Islandsprobably represent the eaaternmost outpost of thissubspecies (Oliver and Shaw, 1953). Honegger(1967) and Gaymer (1968), but not Hirth and Carr(1970), use thia trinomial for the Seychellespopulation, Other workers in the Indo-Pacific

region, notably Deraniyagala (1939), Suvatti(1950), Harrisson (1950-1969), and Hendrickson(1958) have also deferred from identifying thesubspecies.

Taylor (1920) describes Chelonia mydas,iaponica from the Philippines as follows:above rusty reddish brown, each shield streaked.with amber; head shields distinctly reddish,each edged. with black; shields on sides of heeddark, with yellow along sutures; shields ou lewith black centres; plastron yellow, Taylor(1970) briefly describea aduli japonica fromThailand as greenish to greyish brown, someindividuals with dark rays; and. the plastronyellow. He goes on to say that the young arealso olive to dark brown with some yellowmarkings on the limbs; venter yellowish; anddark areas on flippers.

There are probably several races scatteredthroughout the Pacific Ocean and the IndianOcean and as already stated, the situationwarrants a careful systematic stuL3.y. The booksby Deraniyagala (1953), }uroda et al. (1958),and A.non (no date) include colour illustrationsof green turtles from the Indo-Pacif io region.A coloúred picture of the venter of a greenturtle from Indefatigable Island, Galpagoe,appears in the article by Johnson and Johnson(1959),

According to Schmidt (1945), becatLee øfhistorical zoogoograpby, one would expect themarine turtle populations on the Pacific coast-of the Americas to be more closely related. tothose on the Atlantic and, Caribbean coasts thanto. iioe in the western Pacific. On the otherhand, evidence is accumulating (see colourdescriptions in preceding paragraphs and below)which suggests just the opposite.

The following accounts of green turtles inthe South Pacific (Hirth, MS) aro given as astart toward a descriptive classification of allturtles in the Pacific region (although it shouldbe noted that there may be significant colourvariations among individuals of the sane pops- -

lation and certainly between different sizes),In French Polynesia, a juvenile green turtlewith a carapace length of 30.5 cui has a carapacepredominantly copper-brown with grey-black rays;the top of the head is black with well-definedsutures and the sides of the head are blackish-brown; the flippers are black above and. whitebelow but with the distal half below mottledyellow-black-brown, The throat and plastron arowhite.

A sub-adult female, carapace length of68.5 cm, from French Polynesia, has a carapacestreaked with copper, yellow, black and. bro:m:the top of the head has a ground colour of black,but most scales have distinct copper spots; thesides of the head are blac)çish; the dorsuin offront flippers are black but with distal one-

1:4

third edged with copper: the dorsum of hindflippers are predominantly black; venter of allflippers are yellowish with black ori distal third;yellowish-white plastron.

Examination of ten adults (eight males; twofemales) from Scilly, French Polynesia showed nosignificant colourdifferences between the sexes.Nine had marked indentations above the hindflippers, All had post-ocular counts of 4/4. A

typical turtle with a carapace length of 100 cmhas a greenish brown carapace mottled with white,yellow, olive and black; the top of the head issimilar to the carapace in colourations the throatand plastron are white.

A sub-adult green turtle (carapace length 45cm) from Western Samoa has a brown-green carapacestreaked with black; top of head is brawn and,sidos of head black; foui' flippers above areblack with some brown interspersed end below arevariegated yellow-black-brown; the plastron iswhite.

An adilt female (carapace length 96.5 cm)from Neste.'n Samoa has a dark brown carapaceabundantly covered with li.t green, light brownand yellow spots and, blotches; top of head isspotted bxown and green; sides of head basicallyyellow with large black spots in middle of eachscale; top of flippers spotted black-brown-greenwhile flippers underneath are yellowish-whitewith few black spots in middle of each flipper;chin and plastron are yellowish.

An adult female (carapace length 101 cm)from Tonga has a light green carapace heavilymottled with black, dark brown and light brown;top of head is basically greenish-black withblotches of yellow and li.t brown; sides of thehead are basically yellow but with black spots inscales; flippers above are dark copper-brown withblack spots and below are whitish-yellow with afew black centred scales: the plastron isyellod eh.

Sub-adult turtles (carapace lengths 58.4 and60 cm) from the Fiji Islands and New Caledoniaare very similar in thai the carapace is basicallybrownish-green with distinct copper, yellow aridgreen streaks; top of head is black with largecopper spots and sides of head are blackish,fading into white near the lower jaw; flippersabove are blackish with copper on forward edgeand tips; venter of flippers are white-yellowwith some black scales on trailing edge; throatand plastron are whitish-yellow.

The green turtle is an occasional visitor inNew Zealand and adults have been described byMcCann (1966'o) as being olive or brown, theshields with more or less distinct brown rays;and, yellow beneath.

Basically, the carapace of a mature female fromthe Gulf of Men and the Seychelles Islands

FIRM 28 Green turtle

(Hirth and Carr, 1970) is mottled with yellow,brown, black and green. The top of the head isspread with black, groen and brown spots, andthere are well-marked yellowish seams betweenthe scales. Adult males are similar to femaleswith the exception that males have less yellowand brown mottling on the carapace and top ofthe head. Whether these pigmontation differencesbetween the sexes are consistent throughout thispopulation is unknown. Deraniyagala (1939) notedsome colour differences between the sexes aroundCeylon. In general, the pigmentation of theSouth Yemeni green turtle population isconsistent with Deraniyagala's (1939) descriptionsof Ceylonese turtles of the same sizes. Hedescribes eight pigmentation phases of the greenturtle in which the carapace passes from clarkgreenish-bronze in the hatchiing to a brownish-red phase, then a variegated phase, and ultimate).y-tq the olive tint with black and, brown streakscaarac-teristic of adults. Boulenger (1890)describes young green turtles from Indian Oceanarea as "dark brown or olive above, the limbsmargined with yellow; yellow beneath, with alarge dark brown spot on the hand and foot,Carapace of adult olive or brown, spotted ormarbled with yellowish".

Hoofien and Yaron (1964) describe thecarapaces of two adult turtles from the DahiacArchipelago as uniform ochraceous brown, Theyalso mention the lack of distinct marginalindentations above the hind limbs.

1.24 Common names

Some common names of the green turtle invarioue areas of the world are given in Table f.The name "green turtle" is derived from the colourof i-'e fat.

The Raroians (Dariielsson, 1956) have namesfor the various stages in the ontogeny of seaturtles: egg (tuo vaki), hatchiing (karëa),three classes of juveniles (torara, kpue,mkui), adult (tYfai).

Sorno Tonga fishermen also have nanee forthe colour phases and various sizes and sex.

1.3 Morphology

1.31 Anatomy

An annotated bibliography of some aspectsof the morphology of marine turtles is found inIngle and Smith (1949). More recently, Bellairs(1970) has written about the general anatomy andphysiology of the groen turtle. An early accountof the anatomy of Chelonia was given by Hoffhrimn(1890). Among the classical works on anatomyare those of Pilliet and Bignon (1885) on thelacrimal gland; Keateven (1911) on cranial

FIRN/585 Groen turtle

TABLE I

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

North Americagreen turtle, green sea turtle,edible turtle

Danish West Indies, Trinidad, Tobagogreen turtle, edible turtle,greenback

Caribbean Spanishtortuga, tortuga blanca,tortuga verde

MEi :icola panama, caguama, ceguama OO4flufl,caguma negra, tortuga prieta,caguama prieta, mestiza,tortuga negra, sacacillo

Guyanabettia

Suninamkrape

Tupi in Brazilmuruaxid

Hawaiian and. Tahi-tianhonu

Western Samoalaumei

Tonga Islandsforni tu'a'uli,form tuaku1a,form tuapolata,-tuai fonu

Fiji IslaMsvomi damn,vonu loa,mako loa,ika damm

Englishgreen turtle, edible turtle

GerranSuppenschildkrSt e

frenchtortue franche, tortue de mer,tortue verte

Portuguese-t art aruga, t art aruga-do-mar,tartaruga vendo

Ghanaanjwa apuhulu, apuhuru, hala

Swahilikas sa

Mozambiquei-tat aruca, riruvi, risa, pat eri,casa, icaha

Bajuxil of Lamutaza

Southern Yemenbissa, biasat al bahr, humsa

Creol. on Aldabra AtolltorLie do mer

Maldive Islandsvol a

Sjnhal esogal ksbNva, mas ksbMra,vail ksbva

Tamilparr znai pl mai

Malay on Cocos Islandspenyu

Malaysiapum, sisek, penyx, penYu agar,penyu pulau

Borneopinu

Philippine Islandspudno, tuntuga

Chineselu -tau-hai

Japaneseao umigamo

West orn Hnisphere Eastern Heniaphere

1:6

morphologr; Nick (1913) on the development of theskull; Ruches (1929a, b) on osteologr; Deraniyagala(1939) on general ontogeny and. scalation; Villiers(1958) ori general anatomy; and. Parsons (1959) onthe nasal cavities. A study of respiratory musclesin nydae has been rende by Shah (1962). The totallack of the lung muscle, surmoularie ritriatumpulmonale, is notable. Mlynarski (1961) hasdescribed the structure of soutes arid. Brongeraina(1968b) described some variations in vertebralsoutes ot' Surinais turtles.

Parsons (1958) examined the choanal papillasof green turtles but was unable to give it afunction. Later, Smith (1961) proposed that theohoanal rakers in adult green turtles arecorrelated with its herbivorous diet.

1.32 Cytomorpholor

According to Iaicirio (1952) the female greenturtle has ono lesa chromosome than the male.Plorking with Chelonia ,japonica from theBonda Islands, he found, through histologicalexamination, that the diploid number was 56 and.55 in the males and females respectively.

Characteristics of thehaemnoglobìnof C. nWtleshave been studied by Dozy et al. (1964).Stephenson (1966) relates how the cells of mayùashave been used in cell culture studies,

1.33 Protein specificity

The serum protein concentration in Oheloniare. myda is 1.8% (Frair, 1964). In most turtlesit is between 2 and 6%. Serological studiessuggest a close relationship among the five generaof sea turtles (Frair, 1969).

1.4 Meanurin' and tagging turtles

The etsndax'd measurements of sea turtles (allof which are straight-line distances) are lengthand width of carapace, length of plastron andwidth of head (see Fig. 2). The lengths ofplastron and carapace are anteriormnost toposterloneost extensionm of each. The width ofthe carapace is the distance between it lateralmargins at the widest point. The head measurementis taken at the groateot bony width.

The moat common method of tagging marineturtles employs the use of a stainless steel tagclamped onto the trailing edge of a front flipper(see Fig. 3), The tags are numbered on one faceand on the other side are inscribed instructionsfor the finder in various languages. Adult femalesare usually the only ones tagged because they canbe easily caught on the nesting beach One of theproblems in sea turtle biolo im to invent a tagthat can be placed on or in a hatchling and uhichwill identify i-t some years later, Attempts -tomark or tag hatchlings by branding, -tattooing, anduse of reageotic tags have not been successful

(Carr, 196Th). Notching marginal soutes is themost common method used today by turtle blob-.giste, but the long-term success of this methodhas yet to be established. Identifyingindividuals through -immunological reactions hasbeen suggested by Hendrickson (1969). Hughes(1970) in experimontm with loggerhead (Carettacarotta) hatchlingm is implanting a piece ofmtainlesa steel wire under selected carapacescales in hopes of identifying individualslater-recaptured by x-rsying them (seo alsoAnon, 1969k). Pough (1970) has recentlysuggested a method of marking hatchiinge asinga plastic plug.

FIRM/85 Oreen turtle

Fig. 2 Measuring the plastron length of aSouth Yemeni green turtle. The use of"giant" calipers gives rtraight-linemeasurements. The mean on the rightholds a pair of tagging pliars

A tagc'd Wr.tern Srnnan p-rr'n tcrtl .

The stainless eteel. tags, numb"red. antinscribed in various ianguaven lavebeen adopted by marine turtle hioioçtntethroughout the world

FIRM/585 Green turtle

2 DISThIBiPPION

2.1 Total area

Green turtles are found throughout thetropical seas end as stragglers in a far moreextensive area. The major breeding and feedingareas, and hence the majority of green turtles,are found between the northern and southern 200Gisotherme (i.e., average temperature of surfac'water in coldest month la above 200G).

On the American side of the Atlantic Ocean,green turtles have been captured or sighted asfar north as New England (Lamson, 1935; Pope,1939; Dexter, 1947; Bleakney, 1965). Carr (1952)sars that individuals have been reported seen inNewfoundland waters but no specimens have beentaken. The southern locality record of the greenturtle in the western Atlantic is 1ar dei Plataand Necochea, Argentina (Barran and Freiberg,1951; Freiberg, 1967).

0m t1e other aide of the Atlantic, greenturtles h.ve been recorded from the English Channel(Hellmich 1962) -to South Africa (Loveridge andWilliazUB, 1957; Rose, 1962; Hughes, 1970).

r2he northernmost records for green turtlesin the Pacific are Washington State and BritishColumbia (Carl, 1955; Logier and Toner, 1961;Slater, 1963), Boni Islands (Stejneger, 1907),Japan (Wang and Wang, 1956), and Korea (Shannon,1956). They have been reported as far south asthe ICermadec Islands (Olìver, 1910), New zealand(McCann, 1966a, b) and Chiloe, Chile (Donoso-Barros, 1966).

Brongersma (1961) believes that the dead greenturtles that have been found on the Dutch coastwere released from passing ships. In 1952 a l:Lvejuvenile Chelonia (carapace length 36 cm) wasstranded near Fetten on the coast of theNetherlands, but it is unhaown whether this toowas released by man.

Few Chelonia have been sighted off the westerncoasts of South P.inerica sad Africa south of theTropic of Capricorn, and none have been seen southof 40°S latitude on the eastern coast of SouthAmerica. The lack of records in these southernlatitudes may be due to the fact that the dominantsurface currents in these regions (Fern or HumboldtCurrent, Faikland Current, Benguela Current) arecold currents,

This synopsis does not include references tostrictly random sightings of green turtles. How-ever, such records should be pooled and analysedin order to determine if cryptic patterns ofmovement and distribution do exist. For exanp]e,the sightings of turtles in the shipping lares ofthe ITorth Atlantic warrant plotting and study.

Nan's activities and depredations on greenturtle nesting beaches (taking adults aM eggs

2,1

for food; construction of dwellings, roads) haveeliminated soma good nesting sites and, hence thedistribution of green turtles has been reduced(see section 5).

2.2 Differential distribution

2.21 Hatchlings, juveniles and sub-adults

It is postulated that the hatoblings leavingAscension Island drift downstream (i.e., towardBrazil) wi-tb the South Equatorial Current (seesection 3.53),

There is a dearth of information on 'thebionomica of hatchliags and juveniles. In fact,the first year of life of all marine turtles hasbeen popularly labelled "the lost year" by marineturtle biologists (see Brongersma, 1970).

Developmental migrations of juveniles andsub-adults are discussed in section 3.51.

2.22 Adults

The distribution of adults la determined toa large extent by -the location of their nestingbeaches and their feeding grounds. James J,Parsons (1962) has written an elegant book inwhich he documents and describes many of thenesting sites around the world. The importantstill extant siteø, na well as recentlydiscovered nesting beaches, are summarized in- thefollowing sections.

The well-known nesting areas are cited inTab1 II and Figs, 4 and 5. All the majornesting beaches are within 260M or S of theequator. The majority of the beaches are onislands. - Heron Island and Europa Island are thetwo moat southerly major breeding sites of thegreen turtle; beaches on the coast of WestPakistan, Baja California and In the HawaiianIslands are the moa-t northerly rookeries.

Some lesser-known green -turtle rookeries arecited in Table III. !'bat cf these are also onislands.

In addition to the rookeries cited in TableeII and III, nesting has been recorded in variousother places in the tropics. These nestinglocales are given below, In some of these placesthe season or density of nesting is unknown andin other cases the beaches are probably no longerused by green turtles.

Distribution in the Pacific Ocean andadjacent seas.

Green turtles have been sighted or caughtoff the Pacific shore of Panama (Evens, 1947),Peru (Murphy, 1925) and Chile (Ysnez, 1951) butnesting beaches on these shores are unknown.

Nesting on Islas de Revillagigedo and Baja

Michoacari Coast, Mexíco

Tortuguero, Costa RicaShell Beach, GuyanaBigi Santi, Suririam

isla Aves

Ascension IslandSouthern Yemen

Hawk Bay,West Pakistan

Aldabra AtollEuropa Island.

ro IOwan, ThailandNalaysia

Indonesia

Philippine"Turtle Islands"

Heron Island, Australia(and other CapricornGroup Islands)

Bountiful Io., Australia(and. other islands inGulf of Carpentaia)

Jl2neJov.March-August

Feb-Augusb

March-Doc.

Doc,-Juneprobably yeararound

year aróund.

year aroundyear around

year aroundyear around

probably yeararound

year around

Oct .-Maroh

June-Feb.

TABLE II

Some major green turtle nesting sites

WSTEHN HEMISPJIE1?E

May-S ep-t ember

late July-early Sept.probably June-JulyApril-May

probably July

EAZTERN EBIMISPHERE

Feb.-Aprilone peak inNovember

late June-early Nov.

February4ay

May-July

July-August

August

January-February

Peters, 1954, 1956-57Duellman, 1961Montoya, 1969

Carr and. Ogrem, 1960

Pritchard, 1969Pritchard, 1969Schulz, 1969Parsons, 1962Lazell, 1967Marice 1958Hirth MS)

Carr and Hirth, 1962Hirbh and Carr, 1970

Hatt, 1957Miìiton, 1966Hornoll, 1927G.R, Hughes (MS)J.Y. Cousteau (film)Parsons, 1962Penyapol, 1958Harriason, 1951, 1952, 1965aHendrickson, 1958de Silva, 1969aSomadikarta, 1968

Domantay 1952-53, 1963

Moorhouse, 1933Bustarc1 1967a, (Ms)

Bustard (MS)Parsons, 1962

2:2 FIRM/$85 Green turtle

Locality Nesting Season Peak of Nesting Authority

Fig. 4 Some of the important green turtle nesting beaches (numbered oiroles) andfeeding grounds (shaded areas) in the western hemisphere. Modified fromParsons (1962) and Carr (1965). 1 Ujalang, 2 D'Entrecaateaux,3 F'unafutio, 4 = Vatoa, 5 Palmereton, 6 Rakahanga, 7 Scilly,8 Rarola, 9 French 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

FIltM/S85 Green turtle 2:

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. Modifiedfrom 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 ofThailand 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 CarolineIslands, 26 = Oroluk

FIRN/S85 Green turtle 2:5

Locality

TABLE III

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

Nesting (by monthi)J F N A M N D

Authority

Pacific Ocean and adjacent seas

Chiapas, Mexico X X X X X X Alvarez del Toro, 1960

Galpagoe In. X X X X P. Pritchard (MS)

Rose Atoll X X Sachet, 1954

French Polynesia X X X Chabonis, L, said F., 1954Hirth (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 X X X X X Underwood, 1951

Coast of WestAfrica X X X X X Gadow, 1909

Senegal X X X Cadenat, 1949Villiers, 1958

Fernando P00:Gulf of Guinea X X X X Eiserrtraiut, 1964

Indian Ocean

Naziwi Is.,Tanzania X X X X X X X

Dahlac Archipelago X

Antove and Comsoledoin Seychelles Is. X X X

Jurayd Is.,Persian Gulf X X

Manira Is. X

Anon, 1969b

Anon, 1969a; Urban, 1970

Mirth and Carr, 1970

E, Hoag, pers. comm.

Parsons 1962

FIRM/585 Oreen turtle

California is mentioned by Beebe (1937), Brattetrom(1955) end Parsons (1962). These areas should beroconrxoitred again, with special, emphasis onassessing nesting aggregations.

In former years green turtles were readilycaught in waters off the Hawaiien Islands espe-cially near Galia, Mololcai and Maui, Today themajority of nesting is on French Frigate Shoalin the Hawaiian Lewards. Especially largeconcentrations of basking turtles have been seenon Pearl and. Hermes Reefs and somewhat smallerconcentrations on Limianeki Atoll anti Layson Atoll,all In the Hawaiian Leewards. Hendrickson (1969)and Northshleld (1969) report green turtles arestill found in the Leewards, but are scarcer thanthey used, to be. Now that the Leeward Islands arewider the guadianship of the U.S. Bureau of SportFisheries and Wildlife, the turtle population maystabilize or even increase. In any case, whatever.is left of the original population, the HawaiienLeewards reriain the biggest green turtle rookeryin the United States.

Parsons (1962) end Wiens (1962) have docrnjnentedthe occurrence of green 'turtles in Melanesia,Micronesia end Polynesia. In earlier days Chelonianoeted, in fairly large numbers on Uli-thi, Ifaluk,Gaferut, Hiato, Olimaran, Fanning Atoll, ChristmasAtoll, D'Entrecastoaux Reefs (July through October),nerican Samoa (August and September), 'Oro].uk Atoll,

Raroia (June through September) and Vaton. Thedensity of nesting on Palmerston Atoll now is un-known, although Cari' (1965) liste it as a majornesting beach. There may still be a large nestingcolony on Palrnerston and, the situation warrantsimmediate attention. Preliminèry attempts atassessing turtle stocks in the South Pacific havebeen made by Anon (1969f) and Bustard (1969a).

Green turtles nest occasionally on the sandybeaches of Arno Atoll in the Marshall Islands hutthey are scarce and of no commercial importance(Hiatt, 1951), They aro now rarely seen aroundKapinganarangi Atoll in the Caroline Islands wherethey once were an important source of food, (Nierin,1963). They are reported to neat in the Helen'sReef area about 480 km southwest of Palau and on asmall island north of Palau and just off theisland of Kayangel, all in the Caroline Islands(Wilson, 1969). Carr (1965) cites nesting onUjelarig in the Central Carolines and on Punafutioand Rakahanga in the South Pacific. Fehl (1958)provides a few references to early accounts ofgreen turtles in the western Pacific and. IndianOceans. Dr. John Hendrickson (University ofArizona) is currently studying the marine -turtlepopulations in Micronesia.

As, far as is known, the extent of nesting inthe Ogasawara Islands ( Bonin Islands) has notbeen elaborated on since (1931) accountwhich included, statements that mating takes piacefrom 1'larch to May and eggs are laid from Maythrough August. Mmkino (1952) also referred. to

nesting in the Bonin Islands; and in 1968Sampson mentioned hunting "sea closeto shore. If still extant, this wouldconstitute one of the most northerly breedingsites of Chelonia.

On the western perimeter of the PacificOcean, green turtles have been reported fromAmami-oshima in the Loo Choo Islands (.'Ryukyu)(Koba, 1959) and, from Taiwan (Gee, 1929-30),Pope (1935) 'cites references pertaining togreen turtles laying eggs on islands near HongKong in July and August. However, nesting herenow seems very doubtful.

The density of nesting today on PulnuBerhala off Sumatra is unknown, but Mohr (1927)noted oviposìtion every month with a peak fromNovember to January. Do Rooij (1915) listed.m,as as occurring on the following islands ofthe Indo-Aus'tralian Archipelago; Sumatra,Biliton, Borneo, Java, Madura, Floree, Celebes,Ambon, Benda Obi, Aru and New Guinea.Pangumbalian, a nesting site on the south coastof Java, s-there a large number of eggs areharvested, is described by Rekeowardojo (1961).Sornadikarta (1962) lists some nesting beachesin Indonesia which are rented out for theireggs. Some, no doubt, are green turtle rookariea-Parsons (1962) describes nesting near Baroengin Java. A thorough 'taxonomic and ecologicstudy should be made of the green turtiethIndonesia. Especially cinco the Archipelagooccupies a unique position between the IndianOcean and South China Sea.

In addition to the large Sarawak rookeriesin i'ìalaysia (seo Table Ii) green turtles alsonest on small coastal islands in West Malaysia.The moat important of which are the Porhentinuand Redsñg Islands off the coast of lCslantenand Trengganu (Hendrickson, 1961).

Besides the well-kncwu Philippine ItTurtleIslands", (see Table II) other islands in theSulu Sea where some nesting occurs includeCavi li, Arena, Lumbucan, Bancoran and. SanMiguel (Martin, 1952-53).

Cockoroft (1968) stated that green 'turtlesare common in wa,rm waters off Papua, and Burley(1924) briefly described nesting on a smallisland in the Torres Strait.

Copulating pairs have been seen InSeptember and October off the beaches inNorthern Territory, Australia, and reliablesources indicate largo nesting aggregations oriislands in the Gulf of Carpentaria (Coggerand Lindner, 1969; Bastard, MS). Evidentlythere is large-scale nesting on the westerncoast of Australia If the following quote isreliable: "All alone the northern beaches ofthe mainland coast (of western Australia) andselected off-shore ielaxidn, thouaandc of feniale

FIRN/S8 Green turtle

green turtles leave their natural environjnentthesea, te make a difficult and. exhausting journey on-.to the lend to lay their eggs" (Anon, 1969e).Another article reports nesting near Onslow inWestern Australia (Anon, 1970e). Mating is report-ed off the Great Barrier Reef in October andNovember (Roughley, 1951).

Distribution in Atlantic Ocean and adjacentseas

Oreen turtles nest on Papaya Beach, TurtleBeach, Leguan Beach, Tigsr Island Beach, SudxlieBeach, Zeelandia Beach, l4ahaica-Mazaicong Beachand. Shell Beach (1arch through August) in Guyana;on Silebache Beach in French Guiana; and. on Bigi.Santi Beach and on the strand. near the mouth ofthe Narowijne fiver in Surinain (Pritchard, 1969).Frair and Pral (1970) saw many green turtlesnesting in May.in Surinazn.

In addition to the references given in PableII some pe,-tinent accounts of the rookery on AvesIsland can be found in articles by Zuloaga (1955)and. Phelps and Phelps (1957). Aves is the onlyabove-wate part of the so-called "Aves Swell".The island is almost surrounded. by a reef and, theriiaxìmuin elevation above sea level is 3.5 m.The island is also a well-known tern rookery.The fact that the island is sinking raises thequestion cf what returning, oviferoum turtles willdo when the-re is not enough beach left for nesting.Monco (1958) states that it Is possible -be capturofrom 150 to 200 fecales in a few nights during Juneand July on Aves Island. -

The Ascension rookery has been brieflymentioned by Packer (1968) besides the authoritiescited in Pable II. In addition to its remotelocation, thi colony is different from some ofthe other major rookeries in that the incubatingeggs are rarely taken by any predators butentrance to some of the beaches is made hazardousby the combination of heavy seas and rocky shore-line.

Some of the much lesser-known rookeries in theCaribbean and. western Atlantic are listed inParsons (1962). These include Isla Mujeres andother coral islands off the coast of the Yuca-benPeninsula (see also Foeberg, 1962; Mexico, 1966)Mona Island; Maraj6 Island; beaches near Rio Doceand Cabo Pio on the Brazilian coast and Trindade.Barth (1962) has described the nesting beaches onTrindade, hut there is a dearth of informationabout the breeding colony -there. Green turtleshave aleo been seen off the shore of QusimadaGrande, about 70 kin southwest of Santos, Brazil(Mertens, 1955). -

Parsons (1962) cites nesting in the Cape VerdeIslands. Brongerema (1968a) quotes the work ofSarmento (1948) which mentions the presence ofjuvenile green turtles around the Madeira Islandsand Brongerema himself describes a juvenile

2r

(carapace lengih 30.4 cre) captured off Madeira.

The northwest bulge of Africa Is one ofthe least known areas as far as marine turtlesare concerned, -C. nwdae appears in a glossaryprepared by Osorio de Castro (1954) and. issimply listed in -the fisheries of coastalPortugal, Azores, Madeira, Morocco and Senegal.Pasteur and. Bone (1960) state that it is pro-bable that at least stray green turtles can befound off Morocco.

Dïs-bnjbutjon in the Indian Ocean andMediterranean Sea

Green turtles are reported as very commonin Mozambique waters and need. better protectionon the nesting beaches (Anon, 19701). Todaythere la only scattered nesting on the coast ofKenya (C. Ray, pers. comm.). oviposition hasbeen observed. on the Somali coast (Cozzolino,1938; Travis, 1967).

The photograph in the Arameo Handbook LAra-bina American Oil Company, 1968) with the legend"Giant Sea Turtles Breed on Islands in the ArabianGulf" is probably a green turtle. There aro alsoscattered. reports of green turtles off Iran in thePersian Gulf,

Green turtles were reported as very commonin the Maldives some forty years ago -.

(Deraniyagala, 1956). Nesting on Gomeros Islands,Diamond Island and Cocos Island is documented byParsons (1962), The current statue of theseislands as nesting' sites is unknown.

- - Shockley (1949) along with HatL (1957) andMinton (1966) describes nesting in West Pakistan(see Table ii),

Several people (Bodenheimer, 1935;. Hans,1951; Howella, 1956) have reported the presenceof Chelonia on the Israeli coast. The areabetween El Anish and Hedera is sometimes named ama nesting site; however, it is highly unlikelythat turtles nest there now. A couple ofChelonia wore caught near Malta some forty yearsago (Despott, 1931). G. mydas is reported asunoonmon in the Adriatic Sea (Redi, 1963). The

green turtle along with loggerhead. and hawknbillturtles has been observed off southern Prance(Knoepffler, 1961).

The only known breeding colony of any sizein the Mediterranean is l;hs rookery on thesouthern coast of Turkey near Marsin and.Yasrortalik (lu-ion, 1967a). According to Mr. I.Sela of Israel, the colony is rapidly diminishingdue io pverexploiiation. The rookery is nowbeing investigated by Dr. R.W. Hathaway(University of Uai),

In some places in the Mediterranean thereports of Chelonia are questionabl becausesome reporters have had. difficulty in

distinguishing the groen turtle from the loggerheadturtle.

Green turtles are vry rare in the Black Sea.One was caught near Sozopol in Bulgaria about TOyears ago (Valkanov, 1949; F\Thn and Vancea, 1961).

Ication of feeding pastures

Some of the principal feeding pastures are:upper west coast of Florida; northwoetern coast ofYucatan peninsula; south coast of Caba; Mosquitocoast of Nicaragua; Caribbean coast of Panama;scattered areas along Caribbean coast of Colombia;scattered areas off Brazilian coast; west Africancoast from Mauritania to Nigeria (see Villiers,1958, 1962 for descriptive accounts); MozambiqueChannel; Bajun Islands and neighbouring coast ofSomalia; Khor Umaira in the Gulf of Men; Maldiveand Laccadive Archipelagoes; around Ceylon andKrusadai Island (see Deraniyagala, 1939, andKuriyan, 1950); scattered areas in the South ChinaSea and ii the Gulf of Thailand; northern sectorof the Gr:at Barrier Reef; Torres Strait; Gulf ofCarpentaria; Flying Fbam Islands; Hawaiian Islands;around scme of the Fiji Islands and. Tonga Islands;Pacific coast of Baja California; Gulf of California;Galapagos Islands, and parta of the coasts ofEcuador and Peru. Figs. 4 and 5 indicato theapproximate liriits of some of the better knownfeeding areas. As is to be expected, the mostextensive pastures are along continental shelves.

There are severa], lesser known feeding areas.Green turtles are reported grazing between January

and March near Sunday Island in the KermadecGroup (Oliver, 1910). They do not breed in theislands but are reported to go north for mating.Individuals have been seen feeding on the bottomin shallow waters off Northern Territory,Australia (Cogger and Lindner, 1969; Bustari,MS), and. ndas is reported as a staple item inThe diet of a small number of coastalaboriginals still living a tribal way of life.

In the Fiji Islands green turtles can beseen feeding in the. passage between Taveuniand Vacua Levu (Bustard 1970b). Hiz"th (MS)has found feeding pastures around Viti Levu inthe Fiji. Group as well as around Tongatapu inthe Tonga Islands.

Very little is known about the population.dynamics, productivity or activities of greenturtles on the feeding pasturesthis in spiteof the fact that a turtle probably spends mostofita life there.

2.3 Determinants of distribution chaestsee sections 2.1, 2.2, 3.16, 3.22 and3,5

2.4 Hybridization

Nothing is known of hybridization innature. The author suspects, however, thathybridization would occur if turtles are ableto pass through the newly proposed croacPanamaCanal; and, if captive turtles escape fromturtle farms.

2:8 FIRM/585 Green turtle

P1RW585 Green turtle

3 }3IONOMICS AND LIFT HISTORY

3.1 Reproduction

3.11 Sexuality

Green turtles are heterosexual. Adult malc3have a very long, somewhat prehensile tail reaciiInas far as or beyond the extended hind flippers.The tail of the adult f emale rarely extends beyondthe margin of the carapace, tiales also have longerclaws on their front flippers than do females.Some investigators have had difficulty in deter-mining the sex of hatchlings, but Idakino (1952)states that the sexes of even embryos can bedetermined by histological examination. Accordingto him, the embryonic ovary of G.m. ,jaionica iseasily distingaishable from the testis by havinga thick and elongated cortical layer of primordialgerm celle which show active proliferation. Thetestis is a little shorter in length but machthicker thn the ovary of the same age. He goeson to say that the testis is further characterizedby the fac t that the primordial germ cells developas solid cords of irregular shape towards themedullary region of the gonads. Farther histo-logical stadien are now in progress in thelaboratories a-k the University of Utah.

3.12 I1aturity

Investigators are not certain at what agegreen turtles reach maturity. Estimates are foui'to six years (Hendrickson, 1958), at least sixyears (Cari', 1967b), at least seven years(Harrisoon, 1962d), five to seven years (Ehremfeld,1970) and eight io thirteen years (Caldwell,1962o). The age, or sise at which sexual maturityis reached may of courae vary between individualsof the same population and also betweenindividuals of different populations. For example,Carr and Carr (1970b) have discussed the variationin maturation siso at the Tortuguero rookery.Evidently this population inc1ude two kinds ofbig turtles: some that grew big after reachingsexual maturity, and some that reached maturity ata large size.

The smallest nesting females recorded on theTortuguero and Southern Yemen breeding beacheshad. carapace lengths of 69.3 and. 78.7 cm respec-tively (Carr and Ogren, 1960; Hirth and Cari',1970), Sizes of nesting females at some of thebetter kxzown nesting beaches are given in Fig. 6.Corresponding weights can be ascertained fromFig. 7.

3.13 Eating

Copulation usually occurs just off the neat-ing beaches, usually within one kilometre of shore.It is unknot-.'n whether the males accompany thefemales from feeding pastures to nesting groundsor whether they make a synchronized rendezvouswith the females at the rookery. At some of 'the

larger breeding areas (i.e., Tortuguero,Sarawak Islands, Bountiful Island, Aldabra Atoll)one female is usually attended by several malesand males show an interest in any female, Thissuggests that polyandry may be the ffLQCIUooerandi, but it may also be a reflection of thefact that females, but never males, are exploitedon the nesting grounds. Or it may reflect onthe fact that all males rernigrate (= periodicmovement oui from and back to the original area)to the nesting ground. every year while mostfemales have a biennial or triennial cycle. How-ever, it seems unlikely that males do romigratoeach year because of the bioenergetic require-mente of such extended travel. On the otherhand, the preponderance of males may not be realat all but merely a reflection of the male'svery intense activity during the breeding season.It is also possible that the sex ratio mayindeed be in favour of males at the start of thebreeding season but that i-t shifts to ari equalsex ratio as the season progresses due io arrivalof more females or due to males leaving. It isnoteworthy thai at Tortu1piero, nesting continuosafter mating activity has ceased and males havevirtually vanished from might.

0f' course, the preponderance of adult maleeoff some nesting beaches may also be explainedby an unequal sex ratio at hatching or a sexualadvantage during ontogeny (see seckion 4.11).

The question of when the eggs of virginfemales are fertilized remains sii enigua. It ishypothesized that the young females accompany-the older turtles to the nesting grouird, matethe"e and return to home pastures without laying(Carr and. Hirth, 1962). Also, according to Carr(1965), "It seems unlikely that any of the eggsof' the current season are fertilized as a resultof current mating. The Encounter probablyservos to fertilize eggs for the next nestingseason, two or three years ahead". This storageof spermatozoa may account for the infertilityof large numbers of oggi: (see section 3.16).-k'. Montoya of Mexico reports (pers. coma.) thatfertility in C.. agassizii eggs is higher inthe first clutch of the season than in laterclutches.

Some aspects of courtship and. copulationoff the Tortuguero rookery are given by Carr andGiovarmoli (1957). They describe how pairing isa strenuous and clumsy operation during whichthe female is often geashed and scraped.. Theenlarged claw on the front; flippers of the maleaids him in grasping the foro edge of the shellof his mate.

It is plausible that some mating takeaplace at the feeding grounds or at other areasaway from the nesting beaches, although thie hasyet to be documented.

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 lengthof the horizontal line indicates the range and the vertical linethe mean0 Data on the Sarawak population are taken from a graphin Hendrickson (1958). Other data are from Carr and Hirth (1962),Priichard (1969) ami Hjrth and. Carr (1970)

3:2 FI 58 Green turtle

F1RW385 Green turtle

200

175

150

(I)

o:e,o-J

I0ozH!275w

50

25

25 50 75 lOO L$O

CAfl/ PACE LENGTH IN CENTiMETERS

Fige 7 Node). showing weight and carapace length of green turtles (both

sexes), Adapted from Hirth ond Carr 197O) and based mainlyupon turtles caught in the Gulf of Men feocling pastures

3:3

3.i4 Fertilization (i) newly matured individuals, (2) reburninindividuals nesting on a biennial cycle, (3

Fertilization is intrnal, The question of roturithg individuals nesting on a triennialdelayed fertilization is discussed in the preceding cycle1 and (4) individuals exuerioncing a phase-section. shift in their reproductive cycle. Oviferous

turtles may converge on the rookeries from3.15 Gonads different feeding pastures, como of which may be

long distances from the breeding beach.See following section.

3.16 Reproductive cycle, nestingbehaviour and selection of nestingbeach

Men is most familiar with two relatively shortphases of the life history of the green turtle -the nocturnal egg-.laying behaviour of the adultfemale, and the subsequent two months of incubationand, hatching.

Green turtles nest every four years in eastern.Australia (Bustard and Togxiotti, 1969), every threeyears in the South China Sea (Harrisson, 195Gb;Hendricksoi&, 1958) and every two or three years inthe Caribban and South Atlantic, but with thetriennial cycle predominant (cari' arid Ogrexi, 1960;Cari', 1965). Recently Carr and Carr (1970a) haveaccumulated evidence which indicates that a four..year reproductive cycle may also exist in the CostaRican population. &z'eover, they have shown how,although an individual usually maintains a constantcycle, modulation may occur and the change may beeither from a three to two year cycle or vice versa.It is postulated that the phase-shift reflectsecological conditions on the feeding grounds, Thus,individuals nesting in any one given year on somebeaches around the Atlantic represent four aouroes

TABLE IV

Meat nesting takes placo at night, altliondaytime nesting does occur occasionally onuninhabited Pacific atolls and has been seen inColombia: (Medem, 1962) and Surinam (Pritchard,1969). The time of year when nesting begins and.the duration of the nesting period vary from oneregion to another (Tables II and III). In someplaces, such as Malaysia, there is year-roundnesting but with specific peak months, while inother places such as Ascension Island, nestingis restricted to a few months of the year. AtTortugnero, a mainland beach where nesting isseasonal, the turtles appear in large numbersrather suddenly, but tend to trickle away at theend of the season.

Most green turtles lay between three andseven times each season at about ten to fifteenday intervals. Hendrickson (1958) stated thatabout 96% of 5,748 nesting green turtles in thethree Sarawk Islands returned -to the same islandfor renesting during the reproductive seasoneven though two of the islands are only about500 metres apart.

Renestinge on the Toz'tuguero beach arousually within 0.4 to 1.2 km of the precedingnest_ng emergence (Cari' and Ogren, 1960), e-ven

tRM 38 Green turtle

* Incubation period is defined an the poriod between oviposition and anerence of thelargest number of ha-tchlings on the surface

Basic reproductive data a-t some major nesting beaches. The rango is given in parenthesusfollowing the mean.

Locality No. eggsper nest

Incubation period.in daye*

Renost Ing intervalin 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

Grejurtlethou the beach topography changes from year toyear. Apparently, the males roam about just off.-shore during the mating season,

In order to find out what females do betweennestinga a couple of mature individuals atTortuguoro were rigged with helium balloons andkept under visual observation (Cari', 196m), Oneindividual that tias allowed to nest was trackedthe next day swimmthg around off the nesting beach.Another individual that was not permitted to neststruck out in a southeasterly course for about sixhours, but the following night returned and nestednear the point where she was released, Furtherstudios on this subject using turtles rigged withtransmitters are now being conducted at Tortuguero(Carr, pers. comm.)

Table IV compares some basic reproductive dataat eome of the major nesting beaches. It appearsthat in addition to being among the biggest greenturtles in the world, the females nesting inSurinam hav3 the greatest fecundity. Banks (1937)studied ovposition in the Sarawak turtle islandsbetween 19:2 and 1935 and found that the averagenumber of ggs laid by each turtle, based uponthousand cf records, varied from 106 to 118, Theaverage number (and range) of eggs per nest at someof the other nesting grounds are as follows:Indonesia, 118 (48.-175) (Mohr, 1927); Sabak, 105.6(72-144) (}iarrisaon, 1965a) and Southern Yemen,106 (70-130) (Hirth and Carr, 1970). Herriason(1962e) describes the monthly laying cycles inSara'.îak. Bustard (1967a) found that clutch sizeincreases with age or size of the turtle On theother hand, Domantay (1952-53) states that youngerturtles lay more eggs than older ones.

Cari' (196m) states that at Tortuguero thefemale lays more than 100 eggs on her mid-seasontrip ashore and fewer on her first and, last trips.0L Ascension Island the first and second eggcomplements of each turtle are usually bigger thanthe third (Cari' and Hirth, 1962). In the OgasawaraIslaxidn,Fukada (1965) reports that each turtle J.aysbetween one and five times each season and thateach clutch consists of about 70 to 150 eggs. Heobserved four individuals through four nestings andno sigoificant correlation was found between thesequence of nestings and number of eggs per clutchexcept that the last nesting had the fewest eggsin each case. Domantay (1952-53; 1968) has observedseveral old Philippine turtles go through the entirenesting repertoire and them not 1ay eggs. Evidentlysome turtles do not retain their reproductive capa-city throughout life. Further studies on thissubject are needed, especially in view of thepractical importance this would have in the manage-ment of the species.

Carr (1967b) has lucidly discussed how thenumber of eggs laid by a creen turtle at eachoviposition (i.e., about 100) has adaptive valuefor the species, Some of these advantages include(i) escape from predation by swamping the predators

ori the beach arid the littoral predators duringthe first few critical hours in the water,(2) metabolic heating of the nest by a mass ofeggs, (3) social facilitation in the climb tothe surface, arid (4) group orientation in thecrawl to the sea. Carr also speculates thathatchiinga may regroup somewhere after enteringthe sea and somehöw take more advantage frombeing a group again.

Metabolic heating of the nest has beenstudied in several places. In Sarawak,Hendrickson (1958) found a progressive tempera-turo rièe (from about 28° to 35°C) in nests ofdeveloping eggs arid postulated that this mightbe an important factor in influencing the per-.cent hatch. Carr and Hir-th (1961) found thatthe temperature of a nest on Ascension increasedfrom 27.9 to 29.9° over a 61-day incubationperiod, In Australia, Bustard, and Greenhain(1968) found that the temperature within anatural nest of eggs is initially about 25-26°Cend that there is a rise in temperature to about31°C during the latter part of incubation due tometabolic heating.

In addition to the incubation data for somemajor rookeries given in Table IV, additionalinformation is available for lessor knownbeaches. Dornantay (1952-53) and Penyapol (1958)report average incubation periods of 51-53 and48-50 days in the Philippine Islands and inThailand, respectively; Thiksda (1965) statesthat it is 63 days in the Ogasawara Islands;and Hirth and Cari' (1970) registered 48-49 daysin Aden.

The percent hatch (defined, as the numberof hatchlings, out of the total egg complement,which successfully roach the surface) variesfrom season to season in different parts of theworld (Table y). Pritchard(1967) states that"the fertility of groen turtle eggs is low, a50% hatching being about average". He alsostates that egg complements which hava been dugup and. reburiecl take longer to hatch than other-wise, but the fertility does not appear to besubstantially affected.

Balasingham (1967), in an interesting studywith leatherback (Dermocbelys coriacea) nests,found that ho cou,J,d increase the percentage ofhatohlinga from transplanted nests if eggs wereburied in clutches of about 50. The enigaa ofwhy the leatherback usual].y lays between 85 and90 eggs per nest thus becomes more puzzling inview of Balasingham'a work. Similar studiesshould be conducted with green turtle clutches.

The nesting behaviour of the (-reen turtlecan be divided into eleven stages (Carr andOgren, 1960):

i) Stranding, testing of stranding site, andemergence from wave wash

Selecting of course and crawling from surL tonest mite

Selecting of nest site

Clearing of nest premises

Excavating of body pit

Excavating of nest hole

Oviposition

Filling, covering and packing of nest hole

Filling of body pit and concealing of site ofnesting

io) Selecting of course and locomotion back to sea

ii) Re-entering of wave wash and traversal of thesurf

Some detailed aspects of the nesting repertoireof the Atlantic green turtle are given in Carr endCijovannoli (1957); Carr and Ogren (1960) and Carrand Hirth (1962). Nesting of the Indo-Pacific formis described in varying degrees of detail by Beebe(1937?; Hendrickson (1958); Bustard and Greenhain(1969); and Hirth and.Carr (1970).

"Sand-smelling" or "sand-nuzzling" by thenewly stranded female on the beach has beenobserved by several people in different localities(carD and Giovannoli, 1957; Carr and Ogren, 1960;Carr and Hirth, 1962; Hirth and Carr, 1970). Theexact function of this mannerism is unknown, but

TABLE V

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

Carr and Hirth, 1962

Carr and Hirih, 1962

Hendrickson, 1958

Harrisson, 1961, 1962a

Balasinghein, 1965

Anon, 196Th

de Silva, 1969a

Bus-tard and. Greenhaxn, 1968

Noorhouse, 1933

it may involve sensory appraisal of the beach,Evidently this trait is lacking in the Pacificforms (Bust ard and Cire enham, 1969).

The tendency to make "half-moons" (. non-egg laying emergence in which the trail up anddown the beach looks like some form of aparabolic curve) and trial holes may be signsof appetitive behaviour, although they may also

merely forms of reproductive unreadiness.Hendickson (1958) believes that abandoned nestsin advanced stages are due to vibrations in thesubstrate caused by nesting turtles nearby.Phis certainly would be of. adaptive value oncrowded beaches by spacing out the nests (seesection 4.5).

The females are very shy when ascendingthe beach. At this time they can be easilysoared (i.e., by barking dogs, lighta etc.)

back into the sea. Hendrickson (19583 describesthe ascent up the beach to the high beach plat-form in some detail. When -the turtle is diggingthe nest hole and egg hole, she io much lesssansitive to alarming stimuli. Once the consum-matory act of egg-laying has commenced, virtuallynothing will canse the turtle io stop. It is atthis stage that large beach predators can easilykill a turtle. The complete nesting repertoire,i.e., from water's edge back to surf, takesabout two to three hours.

At some rookeries there appears to be nodirect correlation between phase of the moon orstage of tide aM nesting, but around some ofthe Pacific atolls a high tide does aid theturtle in reaching the beach and most nesting

3:6 YIR1/S85 Green trle

Locality Percent hat ch Authority

Tort uguero,

Costa Rica 50

Ascension Island 54 (naturalnests)

Malaysia 47

30-79

53-55

Sabah 70-90

Heron Is., 67Australia 55 (naural

nests)

t11N/385 Green turtle

io at this time. In other places indigenouspoople will claim that most nesting taken placeat certain phases of the lunar cycle.

Short of imparting a tendency to pack solid,aerate too freely, or to waterlog, the physico-chemical characteristics of good turtle beachenare not very well known. However, one charaoter-intio common -to most rookeries is the presence ofa beach platform well above flood tide. It is onsuch a platform that most nesting taken place.Hirth and Carr (1970) found that the texture andcomposition of he sand on some major nestingbeaches vary widely. For example, -the sand on thebig rookery a-t Sharma (Southern Yemen) is composed.of particles chiefly in the 0.5 to 0.25 mm olass -that is, medium grain sand. The beaches atAscension and. Aven Islands are mainly composed ofcoarse sands and. the beach at Tortuguero is madeup of fine sand. Other features of -tha nestingbeach sand, vary as much as particle size does.Golcur, for instance, varíen from, olive grey atTor-tuguero to white on ¡tidabra; soluble saltcontent on Sharma is about one third that atTor-tuguero; organic content a-b Sbarma is less thanhalf thai at Aldabra and Aven Islands. Pritchard(1966) reports -that green turtles iay eggs on abeach in Guyana that is componed enbirely ofbroken sea shells. Ponyapol (1958) han statedthat green turtles nest on different types ofsandy beaches in Thailand. On the other hand,Hendrickson and Balaninghaxn (1966) found that inMalaysia green turtles prefer beaches of fine sand.Nesting beach preferences, or lack thereof, inimportent information for the turtle farmer whomust get his turtles to deposit eggs on man-madebeaches,

Green turtles have laid eggs on completelycwtficial beaches in the Seychelles Islanda,Hauaiian Islands and on Grand Cayinan during thecourse of pilot farming experiments. Details oftho type of sand used are lacking.

The presence or absence of vegetation is nota universal prerequisite for green turtle nestingbeaches. Some parts of the rookery at Tortugueroare almost covered with Ipomea, $osu-rium, andSporobolus, and the green turtles nesting ineastern Australia seem to prefer the older, morevegetated cayo. The beaches on the Sarawak TurtleIslands are devoid of vegetation1 while on thewest Malaysian Islands pioneer plants do invadethe nesting area. There are some plants on theAves Island strand, but only scattered clumps ofvegetation on the big Southern Yemen rookeries.,There aro virtually no plants on the nestingbeaches on Ascension Island. In Australia, Buatard.and Greenham (1969) found that shrubs and trees onthe nesting beaches may servo as a cue to theinitiation of hole-digging since a majority ofturtles studied commenced digging a body-pit inclone proximity to these forms of vegetation. Thisbehaviour has adaptive value since the presence ofroc-blets and/or moisture prevents sand slippageduring the construction of the egg chamber.

Green turtles nesting in Suman appear tobe very plastic in their beach preferences sincethe beaches there are continuously ehaging inphysical structure and location (Schulz, 1964,1969),

Some of -the beaches in -the PhilippineTurtle Islanda are becoming unfi'b for meeting due-to excess accumulation of logs, uprooted. trees,roo-be of coonu-b palms and other debris castashore (Domentay, 1952-53). Similarly, thestrand vegetation is encroaching upon some of-the nesting beaches in Western Samoa (Hlrth, MS).

The cues that green turtles use in select-ing their nesting beaches are e-bili unknown,It is well established, however, that the samenesting beaches are used sear after year. Forexample, no adult turtle tagged at Tortuguerohas been re-taken nesting anywhere else exceptat Toriuguero. It is also possible thathatchlings are somehow imprinted with somesensible essence of the beach on which -theyhatch (Koch, Cari', Ebrenfeld, 1969) end that ibis detection of' -this that guides them back tothe same beach when they return as reproductivelymature females ready to nest. This imprintingtrait is also the rationale behind the restockingprogramme in the Caribbean (seo seotion 6.5).

One of the biggest nesting concentra-tiene,of green turtles In the world and the site ofextensive ethoecoiogical research is the beachnear Tortuguero on the Caribbean coast of Costadica. The nesting beach is approxima-boly 38kilometres in ox-bent and has been described byCari' (1956), Ca.rr and Giovennoli (1957), Cari'and. Ogren (1959, 1960), Cari' and Hix'th (1962),}Iirth (1963), Carr, Hix'th and. Ogren (1966),Cari' (196Th) and Stubbn (1970).

3.17 Eggs

Green turtle eggs are spherical and. white.During oviposition -the eggs are usually droppedone, two or three at a timo. Mast eggs arecovered. with mucous when they leave -the cloaca.Egg laying takes about -ten minutes. Freshlylaid eggs in Sarawak weIgh 36 g each (range28.6-44,7) and -those In Southern Yemen weigh40.4 g (range 30-44). The average diameter ofgreen turtle eggs ranges from 40 an in Sarawak(Hendrickson, 1958) to 54.6 mm on AacennionIsland (Carr and Hir-th, 1962). Eggs on theTortuguero and Southern Yemeni beaches averagebetween 42 and. 46 ram (Cari' and Hirth, 1962;Hirtb and Cari', 1970). A few abnormally smallor non-spherical eggs are usually found. in eachclutch.

If we assume -that the average green turtleweighn about 125kg (wet weight), that she,lays 550 eggs per season and that each eggweighs 38 g, then wo can concludo that theaverage adult fornaio produoea about 21 kgof eggs per season, or an amount equivalent to

3:8

about 17% of her own bod,y weight.

Penyapol (1958) has analysed the nutritivevalua of green turtle eggs. He found that ashaccounted for 1.11% of 100 g of turtle eggs.Based upon the same sample size, he recorded thefollowing major constituents: moisture (79.03 g),protein (11.8 g), phosphorus (227,85 mg), calcium(93.58 mg) and iron (1.95 mg). The cabrio contentof the 100 g sample was 121.36 (compared to 146 inhen's egg and 189 in duck's egg), Vitamina B1 andB0 registered 453 and 442 micrograms, respectively.The total edible portion of the sample was figuredto be 92.8%.

Nan eats the eggs of green turtles chiefly fortheir protein content; occasionally local demand isbased on a belief that they are an aphrodisiac.

Green turtle es are eaten by a variety ofpredators (Table VI). As the table indicates,feral dogs take a large number of turtle eggs.umani of course, take more eggs than any other

Tor'tuguero, Costa Rica

Galapagos Is,

Lldabra Atoll

Southern Yemen

West Pakistan

Malaysia

East Indies

Philippine Islands

TABLE VI

Predation on Green Turtle Eggs

WESTERÑ }MISPIIHRE

dogs, buzzards, opossums,pigs, white-lipped pecoary,coatimundi Carr, 1956, 196Thferal dogs, pigs Schmidt and Inger,

1957P. Prit chard (MS)

EASTERN IIEISPIIERE

sand. crabs (Brachyura sp.)robber crabs.rgus latro)ibis, dogs,rataferal dogs, foxes beachcrabsferal dogs, jackals, crows1monitor lizardsghost crabs (0cyod.e sp.)monitor lizards (Varanus ap.)rat smonitor lizards,pigs (Sus barbatus)dogs, monitor lizards

FIR14/G85 Green turtle

predator.

Rascona are a terrible menace to loggerheadeggs on the Florida coast and may be preventingrehabilitation of green turtle rookeries there(Carr, pers. comm.).

On high-density nesting beaches, such asHeron Island, Europa Island, and the SarawakTurtle Islands, nesting females may destroyincubating clutches as a oonsequenoe of diggingtheir own nests (see section 4.5)

3.2 nbz7onio and hatohling phase

3.21 hibryonio phase

Domantay (1968) has doouniented. theembryobor of the green turtle from the time thee leaves the ovary to the day of hatching.His observations were made in the PhilippineIslanda where incubation lasts about 50 days.According to Domantay, the early stages of

Honeggor, 1967Hirth and Carr, 1970Anon, 19670

Minton, 1966

Hendrickson, 1958

Raven, 1946Domantay, 1952-53

Locality Predator Authority

FIR4 8r Orcen turtle

gastru].ation are completed before the egg is laid;by the end of the fifth day the neural tube isdifferentiated into the brain and spinal cord;differentiation of the heart and gut bogin aboutthe eighth day; the nervous, digestive andoirculatory systems aro fully distinguishable bythe eleventh day; at about the seventeenth daythe eye is differentiated with a lens and irisand the anlage of the carapace is present anddifferentiating; the scales are fully formed andpjnented by the thirty-first day; two days beforehatching the yolk meanures about 30 mm in diameter.According to Domantay it takes the hatohlings atleast three days to reach the surface.

Penyapal (1958) found that embryos from theThai land rookeries have a carapace length ofabout 5, 11.2 and 36 mm on days 14, 20 and 36,respectively; that over the same period the widthof the carapace varies from 4 to 31 mm; and, thatthe forelimba grow faster than the hindlimbs afterday 16.

Deraniyagala (1939) recorded that a 16.-day-old Ceylonese embryo has a carapace length of 11mm and a carapace width of 6 mm. At the sametime, the lengths of the fore flippers and hind.flippers are 3.5 and 3.0 mm, respectively.Between 29 and. 36 days of age, the lengths ofthe carapace and flippers almost double, and bythe thirty-sixth day the carapace is pißmented(dark olive). According to I'orhouse (1933) alltraces of the umbilicus disappear ab about threeweeko of age.

TABLE lITI

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

3:9

A rather old but comprehensive monographon embryology, especially that of the head, hasbeen written by Parker (1880).

Agassiz (1857) described in detail theembryology of various turtles, mostly Chrysemysand Chelydra. It would be interesting -tocompare the embryology of green turtles Withthose described in Agassis's classic work.

Bustard, Sisidsa, Jonkìns and Taylor (1969)have shown that the eggshell is the major sourceof calcii.un for developing embryos.

The new hatchiing cuts through its eggshellwith the aid of a horny protuberance, sometimescalled the egg-oarunole, on the tip of its SnouioThis caruncle disappears after a fow days.

3.22 Hatohling period (behaviour,predators and curvivorship)

The dorsum of the haichling is black ordark brownish black or dark greenish black andthe edges of the carapace and flippers areusually white; the venter is white except theends of -the flippers which are black bu-t edgedwith white (see Fig. 8). Sizes of haichlingsare given in Table VII. A picture of ahatohling just breaking through the eggshell isprovided by Carr (1967o). Albino haichlingshave been reported from Sarawak (Harrisaan, 1963)and from Costa Rica (Carr, 1967b).

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)

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

3:10

I'ig. 8 One-hour old. green turtle hatohlizigs, eachabout 23 g. The dark carapace and. flippers,edged dth white, are typical of theapciea.

The movements of hatoblinga can convenientlybe divided into the following categories (fromCarr and. Ogren, 1960): (i) from nest to surface,(2) from nest -to wave-wet sand, (3) across wave-wet sand, (4) through wave-wash, (5) throughbreakers, and (6) travel by hatohlings and.immature stages (also see sections 3.51 and 3.53).Phases (j) mid (2) above have been the mostintensively studied, while virtually nothing isknown about the last phase listed. above.

The movement from the nest to the surface isa combination of negative geotaxis and allolomiinetiebehaviour, Hendrickson (1958), Carr and Ogren(1960) and Carr and Hir-th (1961) provide detailed.accounts of the complex series of activities thatcharacterize the emergence process - the salientfeature being that turtles in a nest are notcompletely independent individuals, but are sortof a super-organism, composed of collaboratinghatchlings whose combined. efforts carry theful" of individuals to the surface. The climb tothe surface usually takes between three and sevendays. Emergence from the sand usually takes placeat night. Sometimes the entire complement emergesat the saine time, while on other occasions thehatchiinge emerge in small lots, over a period. of24-72 hours. This is probably due to subdivisionof the group by differences in hatching schedules.Turtles hatching singly have reduced prospects of'emerging from the neat, and even less chance ofreaediing the sea,

Hendrickson (1958) believes that some innatemechanism prevents hatchiinga from emerging on thesurface during the heat of the day. Ho postulatesthat if they encounter sand temperatures muchabove 33°C, they cease activity and resume climb-

Fmnu4/s85 Greco 'turtle

Ing only when nightfall brings lower sandsurface temperatures, Bustard (1967b) foundthat the majority of Australian hatchlingealso emergo from the underground. nest atnight and he suggests that inactivity duringthe day of the "lead" turtles near the surfacehas a dampening effect on the activity of thosebelowthus ensuring that emergence does nottake place when air or sand surface tempera-tures could be lethal. Nroaovsky (1968)concluded thermal inhibition of activity is amajor factor in limiting the emergence ofhatchling green turtles to nocturnal hours,Activity is slowed down when sand. temperatureorise above 28.5°C,

The advantages of emerging at night arechiefly two: (i) it eliminates exposure to hotsand surface -tesperatures which could belethal or which could slow down the turtle inits crawl to the sea and. thus lengthen itsexposure to predators (2) it eliminatesexposure to diurnal predators,

Hatcblingo crawl along the sand. surfaceusing their four flippers in typloal reptilianfashion (front member moved in conjunction withhind, member on opposite side), while adultson shore heave themselves along using some-times only the front flippers simultaneouslyand sometimes ali four limbs simultaneously.

Carr and Ogre:a (1960) conducted a seriesof tests -to elucidate the orientation (= thetaking up of a direction in relation to astimulus) of the hatchlings from the nest tothe sea, They concluded, that -the fundamentalguai sense involved visual stimuli, and thatthe response is a modified phototaxis, The"o annosa òf outlook" of a sea-mlnj horizonwill draw them away from a sun or moon wheneither is over the land. But this relation-ship between intensity and area is evidentlyreversed in the case of very bright light.Morhouse (1933), for example, stated thathe could en-tice the hatchlings back onto -thebeach after they had entered the sea byshining a bright light. Carr and Ogren (1959)found that they could draw hatchling leather-backs back -to shore after they had enteredthe surf by setting a kerosene lamp on theshore or by shining a flashlight on them,And,. evidently the combined effects ofillumine-ted sky over cities and. bright streetlights account for -the destruction of logger- -

heads (Caret-ta caret-ta) on the coastal highwaysof Florida (McParlane, 1963), ?csovaky(1970) has stated that any kind of visualstimulus has some effect on -the seafindingorientation of hatchlings and -that under somecir':uma-tances the sun or moon could distracthathhlings from the shortest route to -the seaand thus increase exposure -time to predators.Although the problem, is not completely resolved,Nrosovskzy (1970) e-ta-tes that the distraction

In/8 Groen Lcr io

to very bright light is counterbalanced by abehavioural mechanism kuown an tropotaxia(= simultaneous comparison of intensities on thetwo sides, and orientation according to thebalance thus struck) which ensures that hatchlingswill i'oach the water wider almost all conditions.

The sea-finding ability of hatohlings am wellas adult females has been further studied on theTortuguero beach by Ehronfeld and Carr (1967) mdEhrenfeld (1967, 1968). Spectacles containingspecial filters were fitted to the heads of adulte,and hatchlings were tested in a circular arenawhere the view of the horizon was unobstructed orwhere it was blocked by a low wall. Theysubstantiated the claim of others that the watèr-finding orientation is primarily a visual process(based on brightness rather than colour) and. thatit involves an appraisal of beach topography. Theyalso concluded that there was no innate compassdirection based on celestial cues, although Ftsoher(1964) did find a fixed direction preferencerelated, to the position of the sun in hatchlingstested in a water arena.

In regard to the visual acuity of marineturtles, Walls (1963) stated that "the femalemarino turtle is in a bad way visually when shecomes ashore to lay her eggs at night, for heraerial vision mut 'be hazy and dim even if themoon is bright". Walls describes the anatomy,accomodation, optic axes and. other aspects of thecheionia.n eye. Ebrenfeld (1967) established thatgreen turtles aro very myopic when their eyes areout of water, Nevertheless, as described above,vision is indispensable for orientation on land.

Retinal oil globules in G.m. nWdas have beenexamined by Gronda and Haden t1970). The arealdistribution shows a mean total o. 7,803 fororange, 8,460 for yellow and 4,132 for colourless.The apparent lack of areal differentiation iscorrelated with its narine niche.

Mrocovsky and Cari' (1967) examined lightpreferences of' hatchlinga in a simple apparatus onthe natural nesting beach and discovered that whoagtven a two-choice situation, they prefer blue and.green stimuli over red. Light preferences woreanalyzed further by Mrosovsky (1967) and byMroaovar and Shcttleworth (1968), and. it was foundthat while wavelength preferences of hatohlings arocontrolled largely by brightness, the existence ofsorne colour preference cannot be ruled out. Ernstand, Hamilton (1969) poritulate -that turtles withsome nocturnal habits prefer the shorter wavelengthsof light and that diurnal apodes prefer the longerwavolongilis.

In oummary, it has been shown that a bright-neon preference (at the level cL' the horizon), atropotactic reaction to light and the associationof th operi horizon with the seaward directioncould explain thc eca-finding orientation ofliatchiino, at leant on the Caribbean beacliec.

3:11

Lab-roared yearlings retain the ability tofind the sea and, even cross complex terrain toreach it, and adult males evidently possess thesame ability, although once reaching -the surfas hatchlingn, they nover return to a terrestrialenvironment except perhaps to bask (C5rr andHirth, 1962). After conducting a variety ofexperiments with G.m. carrinegra in the Gulf ofCalifornia, Caldwelr and Caidwell (1962)concluded that the sea approach ability ispresent in both sexes of all ages.

As already mentioned, the habits of thehatehling after it enters the sea are completelyunknown. Caer (196m, 1969o) postulates thatthe batchlings are pelagic for some monthsbecause observations on captive hatchiingosuggest that they have -trouble feeding on thebottom in deep water and that their counter-shading (i.e., dark dorsum: white venter)resembles pelagic fish. Many marine organismsare pelagic during the early stages of their life,Cari' suggests that the Sa,rgasso Sea might be oneplace to look for young -turtles, especially sinos-they have not been seen in shore-waters anywhere.It is also postulated that hatehlings arecarnivorous (in captivity they readily oat avariety of animal food). Theoretically, the sea.-air interface would be an optimal niche for asmall, weak-swimming, air-breathing, carnivorousanimal. Hughes (1969a) believes that loggerheadhatehlings are pelagic and that their movementsare regulated to a large extent by dominentsurface currents.

Bustard (1970a) states that -the blackcarapace plays en important role in elevatingthe hatchling' s body temperature when it floatson the surface of the sea. He also recordedthat hatchlings start to dive regularly at abouttwo to three months of age when ihy weigh about1,200 g.

Mrosovsky and. Ehettleworth (1968) suggestthat the temperature of currents off the nestinggrounds may partly de-termine to what extenthatohlings swim or remain planktonic.

Cald.well (1969) cites records of hatchiingebeing dipneti;ed off their rookeries in theRevillagigedo Islands and off the coast ofMicthoaean, Mexico, How long hatchlingo remainin these offshore waters is unknown,

Predators of hatohlings are given in TableVIII. As the table shows, hatohlings are takenby a variety of avian, terrestrial and marinepredators.

A hypothetical surv±imrship curve (Fig. 9)indicates the greatest mortality takes place inthe very early months of life. It has beenestimated that only 2-3% of the haichlingssurvive to athilthocd (Banks, 1937). It may beas low an

WESTERN IISPHERB

Tortuguoro, Costa Rica buzzards, dogs, jackfish,kingfish, snook, sharks

Gulf Coast of Maxioo dogs, hogs, crabs, birds,fish, sharks

Pacific Coast of Mexico raccoons hogs, dogs, crabs,fish, sharks

01PO9 Is. feral oats, nigh-b herons,frigate birds, ghost crabs,groupera

Mopelia, Frenoh Polynesia hermit crabs, sea birds,sharks

Marshall Is. Polynesian rat (Rattus eculana)red hermit crabs (Coeriobitapenata)

Fiji Is. mongoose

EASTERN HEMISPHERE

feral cats, rock cod,common jack fish, sharkshermit crabs, robber crabs, rats,frigate bird (Fregata minoraldabrensis), flamingoeni-

ruber ant iquorum),grey heron (Ardea cinerea),black and whit o crow (Coniaaihus), vara-vara (Lutianus bohar)caraxigue-1 ec-dent s (Caranx iobilis)sharks, barracudabeach crabs, birds, groupers,sharkscrabs, birds, mammalsghost crabsmonitor lizard, rats, ghostcrab (Ocypode oeratopthalma),snakes (Boiga dendro hUa,Python ret iculatus , fish, sharks"host crabs, rats, silver gull(Larus novae-.hollandiae),black.-tipped reef shark(Carcharinus paiianzani)

Carr, 1956, 196Th

Mexico, 1966

Mexico, 1966

P. Pritchard (MS)

Eggleston, 1953

Fosbeng, 1969

Bustand, 1970b

Cam and Hirbh, 1962

Horne]J., 1927Honegger, 1967

Hirth and Carre 1970Minton, 1966Gibson-Hill, 1950

Hendrickson, 1958

Noorhouse, 1933Buatard, 1967a

3:12 FI1ilJ/s85 Oreen tut1ç

TABLE VIII

Predation on Green Turtle Hatohlings

Locality Fred at or Authority

Ascension Is.

Aldabra Atoll

Soirthorn Yemen

West PakistanCocos Is.Malaysia

Heron Is,Australia

O)

owm

T

1

AGE IN YEARS

ÑuY

IttttIt's

I

Itt

GREEN TURTLE

rig. 9 Hy-pothetical surv1.vrship curvos of green turtle hatohlins aand man. Plotted on the bae of survivors per thoumaM (logscale) and age in relative units

1'i1*l/385 Green turfle 3:13

3:14 FIRM/S8 Green turtle

3.3 Juvenile, sub-rtult .nd adult phase

3.31 Longevity

Little is known about the life span of greenturtles in nature. The fact that reproductivematurity may be reached after anywhere from four to13 years; that some females have been recapturecseveral times, after two to three years'absence, ontheir nesting beach; together with observations thatpredation on adults is minimal (except 'for man),that epizootics are unknown, and that food, isplentiful, all together suggest that adult turtlesmore than 20 years of age may be comon (except inplaces where they are exploited by man).

Pope (1939) reported that a green turtle fromthe Pacific Ocean lived for 15 years in the NewYork Aquarium.

3.32 Hardinees:(see sections 3.13, 3.16,3,4)

3,33 Competitors

The only known vertebrate competitors of anyimportance are dugongs, manatees and some herbi-vorous fish (see section 3.42)

3.34 Predators

Sharks prey upon large green turtles(MDorhouse, 1933; Tinker, 1941; Travis, 1959, 1967;Hendrickson, 1969; Hirth and Carr, 1970), About 4%of adult females observed on Nalaysian nestingbeaches showed evidences of shark damage(Hendrickson, 1958). Groupers are capable oftaking individuals up to 4.5 kg (Tinker, 1941), A22.7 kg green turtle was found inside the stomachof a tiger shark caught near Clarion Island (Beebe,1937).

Large terrestrial mammals such as ferai dogs,tigers and jaguars may take a few adult females asthey deposit their eggs on beaches, but such pro-dation is probably minimal,

3.35 Parasites, diseases, injuries andabnorirali-ties

Ingle and Smith (1949) provide a list ofreferences pertaining to the parasites of marineturtles, Their list includes: Distonuan constricturn,Ozobranchus branchiatus, Stornato lepas transversa,iThytidodoides intestinalia, R, similis andPlatylepas loe, Nora recently, otherinvestigators have reported on parasites. Carl(1955) recorded barnacles (Balanus crena.tus) on thecarapace of a sub-adult green turtle found on thecoast of British Columbia, Hendrickson (1958)records barnacles (Chelonibia testudinaria,Ste hanole as muricat) and leechea (Ozobronchusbranchiatus on turtles in the South China Sea.He aleo observed mosquitoes biting the soft akin

of the turtle's upper eyelids Hundreds ofbarnacles (Platylepas hexastylos) encrusted thedorsuni and plastron of a sub-adult female takenin Chincoteague Bay, Ilarylend (Schwartz, 1960).Gupta (1961) took five trematodas from theintestine of a green turtle from Trinidad(Cricccephalus albus, Pleurogonius mehrai,Neoctangium travassosi, Deuterobaris cheloneiand Schizarnphiztomoides chelonei Caballero yCaballero (1962) found the trematode, Orchidasmaamphiorchis, in the intestine of a turtle andSimha and Chattopadhyaya (1969) record a newblood fluke from the heart of a green turtle,

Smith nd Coatee (1938) describe fibra-epithelial growths in the skin of etydas and theydiscuss the microscopio structure, The growthsin wild turtles occur on the neck, eyelids,axillae and groin.

Nothing is known of epizootios among greenturtles if, indeed, there are any.

Some individuals may get hopelessly wedgedbetween rocks in 'their approach to nestingbeaches and thereby perish (Oa,rr and Hirth,1962).

3.4 Nutrition and growth

3.41 Feeding and behaviour on thefeeding pastures

Most adults, and to some extent sub-adultsfeed chiefly on their grazing pastures (seesection 3.42), although in some regions of theworld turtles may feed en route between breedingand feeding sites, As far as is known, nofeeding, or very little, takes place off thenesting grounds.

The feeding habits of juveniles are notwell-known, although it is commonly believedthat they are carnivorous for their first fewmonths of 'life,

It is assumed -that some time after theturtle passes one year of age or when it weighsbetween I and 4 kg, it becomes mainly herbivorous(see section 3.42). Whether this dietary shiftis gradual or seasonal is unknown, This changein' food preference is not uncommon, since it hasbeen reported in other reptiles, especiallyfresh-water turtles and lizards,

Sizes of grazing males and females on thefeeding pastures in the Gulf of Men are givenin Table IX, As is to be expected, the female'feeding population includes smallér individualethan the nesting population.

Cari' and Carr (1970a) suggest that condi-tians on the feeding ground may affect the inter-val between reproductive cycles (see- section 3,15),

P[R!4 58 Green turtle

Carapace lengthCarapace widthPlastron lengthHead width

TABLE IX

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

Travis (1967) observed green turtles on -theirfeeding pastures off the east coast of Somalia enddiscovered that they "slept" in 30 to 50 minutebouts usually wedged under natural overhangs.. Healso desci ibes catching sea turtles with remoras.

Carr and Ogren (1960) n-tate that the greenturtles ir. the Caribbean usually spend the night onthe bottom, or among rock or coral crags, and thatuitahle sleeping sites may be located some distance

away from their feeding pasture.

Parrish (1958) observed that, in captivity, aresting turtle curfacee far air every 10-56 ninutes..Active turtles surface every 30 seconds ta tonminutes. Surfacing for air at night usually occursabout every 35-45 minutes. Green turtles sleptresting on the bottom of the seaquariuxn and sleepingbegan at sundown and lasted all night, Walker(1959) observed that nostrils of green turtles areolosed when they rest in an aquarium and. he suggests-that the closure is an adaptation to prevent waterfrom éntering or air leaving the respiratorypassages when the animal is resting..

Observations were made on several large G.m.carrinegra in captivity (Caldwell and Caldwell,1962) and it was recorded that they slept on thebottom of the aquarium with the eyes tightly shutduring the night, but nighttime sleep was interruptedby fairly frequent (at least once an hour) trips tothe surface to breathe. Presumably the saraobehaviour takes place in nature on the feedingpastures. However, turtles have been reportedfeeding at night in the grazing pastures in -the Gulfof Men (Mirth and Cai-r, 1970).

Sleeping habits of the hatchlings are quitedifferent. They usually float on the surface withtheir front flippers drawn in along each side of thecarapace. Some float with their front flippersextended at right angles to the carapace.

Carr (1967e) postulates that green turtlesrarely dive deeper than 12 fm. Most of the

best turtle grass pastures aro within this unit.

A brief discussion of some of -the physiolo-gical problems encountered by diving turtles isdiscussed in Gans and Gaunt (1969). In laboratorytests Berkson (1966) was able to show thatmetabolic rates are much reduced in prolongeddiving and -that the green turtle must be veryresistant to asphyxiai distress. Later, Borkson(1967) found that when the green turtle issubjected to hydrostatic pressure a-s. groat as 19atm., as long as nine minutes may e).apse betweenheart beats. He also measured nitrogen tensionin carotid artery blood and in tracheal air andfound that equilibrium conditions aro neverattained during a prolonged deep dive, but thatenough nitrogen enters -the blood io render thegreen turtle susceptible to gas emboli in thebran after emergence.

MoCutcheon (1947) reported that the specificoxygen affinity of an adult green turtle is 19,0,compared to 28.5 for that of a loggerhead and to12.0 for that of a box turtle (Terrapene). Thedifferences in the unloading capacity of oxygenare correlated with behaviour and habitat.

3.42 Food and feeding associa-tes

Some food items of adult green turtles arelisted in Table X, As the list indicates, adultturtles are mainly herbivorous bu-t they aro notaverse to esting animals. In fact, captiveturtles are commonly fed animal food (section3.43).

Other Investigators have reported on -thefeeding habits of the green turtle but theiraccounts are -of a general nature or they have notgiven the exact feeding site or they havereported on only one or t» individuale. Theeòworks follow: Zostera along the Atlantic coast(Kingsley, 1885); algae end Zostera (Gadow, 1909);q,vinoctocoa, Thalassia, Zostera and Halophila

Menge Mean Range

48,3 - 111,8 90,4 71,1 - 104,135,6 -. 88,9 68,6 50,8 - 81,338.1 - 86,4 72,9 58,4 - 81.38..9 - 17.8 14,2 10,2 - 17,8

1JLE$, n 178 MALES, 11=112

Galapagos

Maui, HawaiiTonga Is.

Fiji Is.

Kextnadeo Is,

Seychelles Is,Gulf of Aden

Kruaadai Is.

Ceylon

Sulú SeaGrest Barrier fleef

TABLE X

Food of Mature Green Turtles

LTHN llEMISPtERE

Bermuda Zost eraFlorida Vallianeria,

Thalassia, (ymodocea,jellyfish,

West Indies Thalassia, molluscs,crust acea

Gulf Coast of Mexico Cymodocea, ThalassiaMonquito Cay,

icaragua Gymodocea, ThalassiaCiaras Brazil algae (Rhodophyoeae,

0hlorophyceae Phaeophyceae),Deplanther, molluscs, osoidians,sponges, bryozosais, crustaceans,echinodorms

Pacific Coast of Mexico algae (Gclidiuxn Sargassun,Rhoc1?menia, Gracilaria,Griffitsia, LiaRon, tJlvaGrat elon,,), odoceaThalascia

Chile Zostera?, alga, sponges

algaemangrove root e and shoot sgreen algaeHalodule, Halophila,yringodian

red and green algae,SyringodiunPterocladia capillacea

EAZ'1MHI HEMISFItSE

odo

Posidonia (or Enhalus),1aloaule, brown and redaigLealgae (Gracillaria,Sargas sun), Cymodocea

ymodocea, Thalassia,Halophila, algaegreen algue and Sargasaumsea grasses

Babcook, 19371eill, 1958Cari' and Caldwell,

1956

Underwood, 1951

Mexico, 1966

Cari', 1954

Ferreira, 1968

Cari' 1961Anon, 1966H. Marquez, Perse

COITUS0

Vanes, 1951DonosoBarros, 1966

Horiell, 1927

Mirth and Carr,1970

Kuriyan, 1950Doraaiiyagala,

1939, 1953Domazitay, 1952-53

Yonge, 1930

16 FIR1 58 Gree12.turle

Locality Food Authority

F. Pritchard (MS)Mirth (MS)

Hirisli (MS)

Mirth (MS)Oliver, 1910

1?IRN/585 Green turtle

(Ingle and Smith, 1949); plant matter,crabs, ascidisne end barnacles in gut ofone individual taken off South Africa(Hughes, Pass and Mentis, 1967); algae inthe South Seas (Roedelberger and Groschoff,1967); eelgraue algae, crabe and snails(Schwartz, 19673; and, animal food such assponges, Pteropods and jellyfish (Brongersma,1967).

Pieces of plastic have been found inthe stomachs of turtles taken in the SouthPacific (Hirth, MS) and in turtlescaptured off the Pacific coast of CostaRica (Carr, pere, comm.).

One of the best described feedingpastures is found near Thor Umaira,Southern Yemen (FAo, 1968; Hirth and Carr,1970). Here the densitis of turtle grassreach 2,500 leaves per m The stomach ofsix mature turtles packed solidly with twospecies of turtle grass weighed between1.8 and 2,5 kg. Unfortunately, information onassimilation is lacking. Ricker (1969) estimatesthat the K value for marine herbivores wouldrarely exceed i%. K is defined as the growthcoefficient, or the trles annual increment ofweight divided by the quantity of food it hamcon seized,

In terms of primary organic production, theWestern Indian Ocean is a region of contrast,containing some of the richest and some of the nestinfertile waters in the world, Although measuremente have not been made along the Somali coastwhere good green turtle pastures are found, theregion has been assigned a moderately high levelof production (.26,50 gCm2 daj) and it ispossible that much higher levels occur (Rydher etal,, 1966), Primary and secondary productivitystudies are greatly needed, especially on thefeeding grounds, bat it may be that feedingexperiments in captivity are the only practicalway to elucidate enerr flow in the relativelysimple mun-grass--turtle food chain,

Barkholder, Burkholder and Rivero (1959)obtained some quantitative and qualitative data onturtle grass, Thalassia testudinurn, in the WestIndies. Among other things, they reported a bio-mass (dry weight) of 5.66 kg/m2, of which 80 to90% was underground; and, that the leaves containedabout 13% protein, 25% ash, 36% carbohydrates,16% crude fibre and 0,5% fat, Nere studies likethis are needed,

C. den Hartog (1970) has written an interest-Ing monograph on the sea grasses of the world, Inmany Instances the distribution of green turtlescoincides with the distribution of sea grasses,and. certainly a century ago the fit was evencloser (see Rig. lo),

Marine turtles and sirenians are probably theonly big vertebrates graz±ng the extensive marine

3:17

Fig. 10 Underwater photo of Syringodiumisoetifoliurn, a marine angiosperm, nearthe Fiji Islands. This is a favouritefood, of herbivorous sub-adult and adultgreen turtles in some areas of theSouthwest Pacific Ocean

pastures, and competition between these groupsIs probably minimal, In the West Indies,Randall (1965) found parrotfishes (Scarus andSparisoma), surgeon'ish (Acanthurus37'Ïinoids(techinus, Tripneinstes and Diadema), and thegreen conch (Strombus gigas) competing with thegreca turtle for marine grasses (Thalassia andymoìocea). Stomach analyses of balahoo fishes

Hemiraniphus brasilienais, in the West Indiesshowad their main food to be leaves of Thalassia(Bu:rkliolder, llurkholcler and Rivero, 1959).Marner and Randall (1952) wrote about the in-vertebrate life in a turtle grass area around anatoll in the Gilbert Islands and found that eveninvertebrates were scarce. According to Wiens(1962) the turtle grass (Thalassia) around someof the Pacific atolls rarely grows over 0.3 shigh hut it makes dense stands over surfaces ofsand in which it creeping rhizomes form aninterwoven mat. He also found that turtle grassmay be the dominant vegetation in lagoons fromzero tide level to a depth of 1.6 fm.

3.43 Growth rate

Rates of growth under natural conditionsare relatively un1nown, although estimationshave appeared in the published literature, Aslong ago as 1916, Schmidt stated in regard tothe Paribbean turtle fisheries that "the mostimportant facts to be determined are the rateof growth of turtles and their migrations".While some good information on movements ofturt:Los in some parts of the Caribbean is nowavailable, there is still a dearth o goodempirical data on growth rates. During the

course of his pioneer studies on growth rates inCaribbean turtles, Schmidt found that nine turtlesinitially weighing between 2.3 and 20 kg showedweight increases of from 138 to 430 g per monthafter 3.5 to 11 months in the ocean. His findingsalso suggest that green turtles weighing aboix 2.3,5.5 and 9.1 kg are about 1-1.5, 2-2.5 and 3-3.5years old, respectively. He further postulatedthat a one-year-old green turtle is about 27 cm inlength and a two-year-old is about 35 cm long.Worhouse (1933) estimated that one-year--oldturtles are 20.3 cm long. Ingle and ith (1949)state that growth rate in early years is about3.2 kg or 17.8 cm in carapace length per year.One-year-old turtles weigh between 1.8 and 3.6 kgaccording to Parsons (1962). 'Carr (1967e) believesthat turtles weigh between 0.9 and 2.3 kg when theyare one-year-old. Australian green turtles weighabout 22 g at birth end about 1 200 g two to threemonths later (Bustard, 1970a).

Cari- and Caidwell (1956) relate how anAtlantic green turtle weighed 20 kg at time ofcapture and then weighed 2.7 kg more when it wasrecaptured 102 days later. Other gains were 0.03cm per day in carapace length and 0.01 ein per dayin width of carapace.

Uithazn and Carr (1968) report that a yearlingreleased when weighing about 830 g was recaptured30 months later some 65 nautical miles from therelease site and weighed 6,350 g. This recapturewas of further significance because the individualwas a pen-reared turtle and the fact that it wasrecaptured is some evidence that pen-reared animalsmay be able to adapt to the normal ecologic regimenof the species after a year in captivity.

Hendrickson (1958) postulated that there isno marked decrease in rate of growth until theanimal reaches maturity and that maturity isattained in four to six years. On the other hand,Caldwell (1962e) theorized that there is a markeddecrease in growth rate after about the first threeyears and that tropical turtles require at leasteight years to reach maturity. i.kre recently Cari-and Goodman (1970) have ahown that after reachingreproductive maturity, female green turtles atTortuguero, Costa Rica grow only about 0.25 cm peryear in carapace length and width (see Fig. ii).According to Carr and Goodman, "It now appears thatsome green turtles mature at small, and others atlarge sizes, and that once they mature-that is,once they have made their first trip to the nestingbeach- their growth becomes negligible".

These data suggest that the growth of greenturtles is similar to most other animals, i.e.,characterized by a sinoid growth curve.

Working in the Gulf of Men, Birth and Cal-r(1970) found no significant differences betweenweights of females and males of the saine length.The same authors compared the weights of turtlesfrom different populations and found that femaleturtles from Southern Yemen are heavier than those

of similar lengths in the South China Sea; andwhile the weights of the larger females fromYemen are similar to the Caribbean green turtles,they weigh somewhat less than the Atlantic greenturtles on Ascension Island. A generalizedweight-length curve is provided in Fig. 7.Generally speaking, the weight of a green turtleat reproductive maturity will be five thousandtimes greater than its weight at hatching.

The weights of mature females in Surinaiswith carapace lengths of from 100 to 122 cmvary from 121 to 224 kg (Pritchard, 1969), Theaverage weight of a mature female fromTortuguero, Costa Rica, is about 114 k and themaximum is about 159 kg (Parsons, 1962).

Caidwell (1962b) noted' a tendency formales of C.m. carrinegra to be lighter thanfemales beginning at a carapace length of about76,2 cm. He also described the relationshipbetween carapace length (L) in inches and weight(w) in kg as follows: log W = -2.14 + 2.60 log L.

As Fig. 6 indicates, the mean lengths ofnesting turtles in the Gulf of Men, China Seaand Caribbean Sea are similar, while the Guianaand Ascension turtles are probably the world'slargest.

The heaviest green turtle reported in theliterature weighed 386.4 kg (Carr, 1952). Theheaviest reported in recent years was a 241 kgfemale on Ascension (Carr and Hirth, 1962).

Almost half of the wet weight of a greenturtle represents edible protein. Deraniyagala(1939) and Schroeder (1966) respectively statethat approximately 41." and 40% of the liveweight is edible meat.

Ingle and Smith (1949) found that theflippers and meat of a 119 kg male and a 96 kgfemale constituted 47 and 52% of the totalweight respectively. The rest of the weight ofthe female was distributed as follows: head,2.4%; tail, 0.24%; neck, 1,7%; kidney, 0.23%;liver, 2.3%; heart, 0.24%; calipee, 6.7%;carapace, 15.5%; and entrails, 10.5%. Theweights of 45 and 68 kg females were alsodistributed in a similar fashion.

A hatchling from the Maldive Islands, fedon fish and bread, weighed 4.4 kg and had acarapace length of 320 mm and a carapace widthof 270 mm after about 13 months incaptivity (Deraniyagala, 1939). Gibson-Hill(1950) reports that the Malay on the CocosIslands raise green turtles in captivity andthat they reach a length of about 25 cm in oneyear. Hatchlings kept in semi-captivity in alag000ri in the South Pacific had carapacelengths of from 15 to 20 cm after ten monthsa diet of kitohen scraps, fish and thellfih(Wiens, 1962), Three green turtles raised in anaquarium grew from about half a kg to about

3W FIRM $8 Green turtle

jJ$85

Green

turtle

- _J

GR

OW

TH

IN INC

HE

S

.ø .

I I t

o

Fig.

11 Grow

th

of mattu'e

female,

green

-turtles

at ortuguoro,

Costa

Rica.

Squares

and

circles

show

mean

growth

for

the

corresponding

growth

interval.

Num

bers

indicate

size

of samples

represented

by means.(from

Cari'

and

Goodm

an,

1970;

Copyriglrt,

1970,

by the

Am

erican

Society

of

Ichthyologists

and

Herpetologists)

23 kg in nine and a half years (í'lowor, 1931).Caidwell (1962c) reported the weight increases offour hatchiinga maintained in captivity for threeyears as follows: the mean weight increase was135 g the first year, 2,725 g the second yeax,and 5 000 g the third year. In captivity,hatchl.ing green turtles can grow up -to between22.5 and 25 cm in six months according toSchroeder (1966). Evidently green turtles inoaptivity readily eat fish and other meat (Beealso Hornell, 1927; Lveridge, 1945; and Parrish,1958).

People in the Tuamotu Archipelago have raisedha-tohlings on coconut meat and fish and they haveobtained a length of 50 to 71 cm in 3 to 3- year.iaheneen on various islands in the Kingdom ofTonga sometimes keep adult turtles in kraals andfeed thais on turtle grasa which washes up on thebeach (Birth, HZ),

3.44 Metabolism

Appa Rao and Dutt (1965) examined -the chemicalcompos&-tion of muscle of an adult male greentur-tlo, Results indicate that the water contentof turtle flesh is close -to that of fishes (i.e.,.80 g water per 100 g muscle), The fat content(0.76 g per 100 g muscle) is lower -than in mostfishes, In 100 g of muscle -they found 15.35 gprotein, 510 mg phosphorus, 210 mg calcium, 34 mgiron, 0,21 mg copper, 0.20 mg cobalt and 0.20 mgmi,ybdenuin.

ChIefly because green turtle oil is us fortherapeutic purposes in some parts of Mexico(used in treatment of tuberculosis, leprosy, andas a source of vitamins), Girai and Cascajares(1948, Girai, Giral and Girai (1948) and Girai(1955) have begun a study of its chemicalproperties. Some of the mont common fatty acidsin the 01]. are laurio, myristic, palmitio, atearicand. paimitoleic. The presence of large amounts oflauna and syria-tic acids may be characteristic ofChelonia mydam. The sterol of body oil in a greenturtle from New Guinea contained 96% cholesterol,3% B-sitoeterol, 0,2% campeaterol and 0.15%stJnasterol (Nina-to and Otomo, 1969),

Biochemical composition and nutrient qualityof the flesh may vary among individuals of apopulation depending on such things as age andsex, s-tmge in the reproductive cycle, kinda ofvegetation eaten on the feeding pasture, andphysiological stress induced by long-rangemovements,

Evidently, -the flesh (or fat or blood) ofsome Chelonia is -toxic to humans, Halatead (1970)gives a lie-b of references pertaining to sesturtle poIsoning. The list contains only onecane of poisoning by green turtles and thisincident caused 18 deaths in Ceylon in 1840. How-ever, the list also contains other instances ofcheloni-toxica-tion in which the species was um-unknown, Most reports of cheloriitoxication are

from the Indo-Pacific region. The origin ofturtle poison is unknown but most investigatorswho have studied the problem appear to be ratherconsistent in their opinion that the toxin isderived from poisonous marine algae eaten byturtles (see Haistead, 1970 for review ofliterature; also Boulanger, 1890; Halsteasl,1959, 1969; and Taylor, 1970). The symptons ofchelonitoxication var-y with the amount Q1' fleshingested and with the person. Symptoms generally

after eating the meat and initial symptomsinclude nausea, diarrhea, epigastric pain andver-ti'. There are no known antidotes forcheloni-toxin. The treatment is symptomatic.Further studies on chelonitoxication anddocumentation of green turtle cases aro needed,

Aimnonia is the major end product in theurine of mydaa as it is in marine teleo stesafishes. The green turtle also excretes a largeamount of hippuric acid and this may be due toits herbivorous diet (Kha.lil, 1947).

An interesting adaptation of Cheloniato its marine environment, pointed out bySchmidt-Nielsen (1959) among others, is itsability to discharge excess salt by means ofsalt glands located near the eyes. In thiregard the green turtle is similar to the marine.iguana and sea snakes. When on land, a turtledischarging salt looks as if it is "crying".Carr and Ogren (1960) suggested that -the "crying"of nesting females also serves to keep the eyesolean of sand.

Holmes and McBean (1964) have clearly shoanthat the "salt glanda in mydas is -the predoTsinattroute of sodium and potassium excretion, Theyfurther hypothesize that mea turtles drink unawater not only to obtain osmotically free waterbut also -Lo enable the excretion of. potassiumingested as food, Such information say beimportant for fu-tune turtle farmers in providing-the proper milieu for their capbives,

Ellis and. Abel (1964a, b) describe how thetun-bis' s salt gland tissue is in some waysdifferent from the salt-secreting epithelia offish and birds, Abel and Ellis (1966) statethat histologically the modified. lacrymal glandof marine turtles may be classified. as compoundbranched tubular glands and that the salt-secreting nasal glands of marine birds, therectal. glands of elasmobrancha and the lacrymalglands of marine turtles have many commoncytochemical and cytological features.

The marine plants that -the turtles grazeon probably have a salt concentration similarto that of sea water and no doubt the saltgland of the tun-tic operates in response to-this electrolytic load.

Thorson (1968) measured the body water insub-adult . riwdas and C,. agasøijL and

3:20 IIH14 S85 Green turtle

determined its apportionment among the major fluidcompartments. Both subspecies were similar inhaving total body water account for about 64% ofbody weight.

Pbr some aspects of diving physiolo'-, seesection 3.41.

3,5 Poa-t-hatchling movements and, baskingbehaviour

3.51 Developmental migrations

Developmental migrations may be a naturalpart of the life cycle of the green turtle,although little is known about the subject. Anon-breeding population ranging in weight from 5to 52 kg appears on the feeding pastures off theupper west coast of Florida in April and. disappearsin November (Carr and Ca],dwell, 1956). Thispopulation is made up chiefly of sub-adults and afew juveniles, The Florida pastures may be a stopin the developmental migration of turtles whichhatch on the rookeries of Mexico and. the YucatanPeninsula. Sub-adults are also taken betweenMaroh and November in the state of Sonora, Nexic.Carr (1961) draws a parallel between this Sonoremcolony and the fleet of young individuals whiöbappear off the west coast of Florida. Schmidt(1916) may have been describing a developmentalmigration when he said ,"Whether the green turtle,the species most commonly cau.t, breeds in theDanish. West Indios, or whether it first arrivesthere as specimen 20-30 cm in length is,strangely enough, unknown, and it is a matter ofgreat importance".

An itinerant population of sub-adult greenturtles is found near Bermuda and is probablycarried there by the Gulf Stream (Iwbray andCaldwell, 1958). A juvenile and sub-adult(lengths 22.5 and 42,5 cm) have been recoveredon the beaches of Long Island, N.Y. (Latham,1969), but apparently were waifs.

3.52 Dispersal and remigrations

1bst adult green turtles customarily makeseasonal trips between feeding pastures and breed-ing grounds, and some of these movements mayencompass thousands of km (see section 2 and 3.1).Fig. 12 shows some places where adult females havebeen recovered after having been tagged on theirnesting beaohes, All the arrows in the figure donot indicate location of feèding pastures, orroutes of travel, but they do indicate themagnitude of the spread from three major nestingbeaches and thus give an indication of the "totalrange" of at least some individuals in thepopulation. The two best studied reznigrationpatterns are those of the Tortuguero and AscensionIsland colonies,

From 4,200 mature green turtles that havebeen tagged at the Tortuguero rookery, there have

58 Green urtle

been 175 international recoveries (Cari', 1967o)dMst of the recaptures have been made on thefeeding pastures off the Nicaraguan coast. Oneindividual was recovered in Trinidad, some2,400 kai from the site of tagging. Twoindividuals marked on the Tortuguero beach nerolater taken about 800 kin away in Colombia, Itwas calculated that the minimum speed of travelfor one individual over a 34-day period was23.5 km per day and that the second turtletravelled.aminintum of 21.6 km per day over a37-day period (Cai'x' and Ogren, 1960). It nowappears that two main fleets converge uponTortuguero to nest - one comes from the south(i.e., from Panama and Colombia) and the otherfrom the Miskito Coya and Nicaraguan coast(Cari', 196m), One of the most wnazing factspieced together from tagging work is that theTortuguero turtles return to the same senentof beach to nest, after absences of from two tofour years (see section 3.16).

Some 556 turtles have been tagged whilenesting on Ascension Island and ten havesubsequently been recovered on the feedingpastures off Brazil, a dispersal of some 2,240km (Cari', 1967c). These turtleB also show apropensity to return to specific, small beacheson Ascension for'nesting (Cari', 1965),

ve females tagged on Musa and Sharmabeaches in Southern Yemen have been recapturedon the east coast of Somalia (Hirth and Cari',1970). Assuming that the turtles swam withthe prevailing surface currents, the longestrecapture would constitute a movement of about3,200 ion (i.e., 'rom Sharma to Chisimayo aroundthe Gulf of Aden), Minimum rate of travel fortwo Yemeni green turtles was computed at 22.4ion per day for 85 days.

Pritchard (1969) reported that one enturtle tagged in Surinaxn was recovered in Brasilafter hav'ingtravelled adistance of about 1,600ka in six weeks. Schulz (1969) also disclosedthe recovery of two Surinam turtles in Brasil.I- thus appears that the turtle grass pastureeoff the coast of Brazil are the feeding groundsfor two distinct nesting populations, viz., theAscension Island nesters and the Surinas nestere.

Harrisson (1960) stated that a femaletagged at Talang Besar, Sarawak in 1953 wasrecovered in 1959 in a net off Kimanis, NorthBorneo, This repreaerrted a dispersal of about600km.

Hendrickson (1969) reported that two greenetugged on French Frigate Shoals were later rèoop.--tured in the main Hawaiian Islands about 800 km

ESE; and that two individuals marked at Pearl andHermes Reef were res ighted on French FrigateShoals - again a movement of about 800 lun ESF.

Green turtles tagged on Heron Island,

:22 1IhN 53 Greon turtle

Fig. 12 Longdistance recoveries of green turtles. Arrows indicate spread from thenesting beach and are not intended to suggest routes. A = tagging site wasTortuguero, Costa Rica (adapted from Carr, 196Th); B tagging site wasMusa and Sharzna Beaches, Southern Yemen (adapted from Hirth aM Carr, 1970);C = tagging site was Ascension Island, South Atlantic (adapted from Carr,

1967b)

3Q0

20°-

loo-

00

loo-

2Q0

3Q0

B'I 58 Green turtle

Queensland have been recaptured in Papua and NewCaledonia b3ustard, 1969c).

Montoya (1969) found a general movement northalong the Mexican coast after oviposition.

It is postulated that the green turtlesnesting on the southern atolls of the SeychellesIslaids remigrate there from feeding pastures offthe East African coast and from the MozambiqueChannel, althou there have never been any tagrecoveries to substantiate th.is., Large numbersof green turtles, including 5uvenileo, are seenoff the beaches in Tongaland, but they do not breed.there (Hughes, 19691,).

In some of the earliest tagging experiments,Schmidt (1916) released 65 marked green turtlesof various sizes around St. Thomas and St. Jan 5.nthe West Indies and recaptured 14% from three toeleven months later. st recoveries were madejust, a few kg from sites of release. The longestmovement recorded was about 80 km and thisindividual was recaptured. -ten months after beingtagged. Schmidt concluded that young individualsare rather stationary.

Carr and Sweat (1969) report that a yearlinggreen turtle tagged and released in the lowerFlorida Keys on 17 October 1967 was retaken on 15November 1968 off Cape Hatteras, North Carolina.

Oliver (1955) made 23 observations on anadult green turtle as it cruised leisurely in alarge salt water stockade and he reported acruising speed of 1.4 to 2.2 km per hour, Variousswimming patterns of captive turtles are furtherdescribed by Parrish (1958).

3.53 Navigation and. orientation

The ability of green turtles to navigate (=ability to initiate and. maintain directed movementindependently of learned landmarks) across vastexpanses of featureless ocean is one of the ,noøtintriguing aspects of their biology.

Working with six loggerheads displaced off theFlorida coast, CarD (1962a) was able to show thatat least the initial movements of some individualswere non-random, In the same paper, Carr discussesthe value of telemetry in unravel1in the mys-teriesof open-sea navi -tion.

The evolution anc3 ramifications of theAscension-Brazil pattern are discussed by Carr andHir-bh (1962) and Carr (1962b, 1964, 1965, 1967a),It is hypothesized that the hatchlings are carriedto Brasil by the South Equatorial Current and that.for adults returning to the island. the ehorteatroute would be directly eastward from the hump ofBrazil against the same current. Evidently food.would be available for the surface-swimminghatchlings, since according to Walford (1958), thesurface waters of at least some branches of the

South Equatorial Current are quite fertile, Atplaces where the current diverges, there wouldalso be more nutrient cycling through upwelling.According to Carr (1967b) it would. take abatchling about -three weeks to t to Brasilfrom Ascension.

The Ascension-Brazil pattern may not veoriginated under present physiographicalconditions, but nay be an artifact of' continentaldrift. Th± idea is discussed by Carr (1967a).Ii this connection, Myers (1967) has found thatthe fish evidnce siipports -the concept that thefinal parting of Africa and South ,nerica, and.hence the origin of' the South Atlantic Ocean,took place at some time during the earlier halfof' the Mesozoic.

Carr (1965) considered celestial navìtionas a possiblo guidance mechanism in long-rangemovements of the green turtle, but at least bi-coordinate star navigation no longer seems o bca realistic hypothesis in view of the fact thatEhrenfeld and Koch (1967) found the Atlanticgreen turtle to be extremely myopic when itseyes are ou-t of water. However, other types ofnavigation involving the sun or moon have notyet bean critically evaluated or -tested.

Several -types of orientation and navigation,with special reference to the Ascension-Brazilremigration, are discussed by Carr (1967b).These include orientation based upon olfactorycues, light-compass sense, maetic sense,perception of Coriolis force and inertialguidance. He also discusses some of the problemsa green -turtle would have to overcome in order tonavigate using celestial information.

It would be informative to be able to tracke. "fleet" of traen turtles in their movement froethe coast of Brazil to Ascension - assuming thatthey do travel in concert. Future research mayshow that swimming together improves accuracy oforientatìou, especially when the goal is anisland. Hamilton (1967) has already discussedthis idea in relationship to avisa remigra-tioneand has postulated that orientation of a flockof birds is more accurato than individualorientation; and. that flocking may be helpfulto juveniles making the -trip for the firat timo.

Now that some older theories of non-visualnavigstion have boon resurrected (for review,seo Griffin, 1969), it uld be worthwhile toexamine a green turtle' a sensitivity to subtlegsOphya&Ql and oceanographic cues, i.e.,forces whiih separately or synergistically couldprovide a bi-coordinate grid. A step in thisdirection has been taken by Koch, CarE' antiEhreuf old (1969). They postulate that olfactorycues emanating frote Ascension Island could berecouized by turtles off the Brazilian coarid thus play a part in the Ascension-Brazilromigration. Homing studies aro now underw5y on

3:24 FIRM/S85 Green tu

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

Ascenajon Island. Preliminary results indicatenon-random movements by a turtle out of sight offixed landmarks (Carr, pers. comm.).

Ilicigvay et al, (1969) studied sensitivity tomouths in three sub-adult green turtles and. theyfound maximum sensitivity in the region of 300 to400 Hz, They conclude that the oar is clearly alow-frequency receptor with a useful span ofprobably 60 to 1,000 Hz, and they further statethat "both on land and in the sea, this ear nodoubt enables the animal to perceive many importentsignals". Is it possible that green turtles possesssome kind of sonar sense?

It is most likely that turtle navigation isa composite process employing different senses andbased upon a multiplicity of cues,

Preliminary attempts at tracking turtles forshort distances have been made with floats,balloons end radio-telemetry (see Fig. 13). It

mag well be that tracking turtles with earth-orbiting satellites will be the only way to followmovements with any great precision.

3,54 Basking

Some Pacific green turtles are known to baskor rest on shore (see Fig. 14). The basking habithas been reported from such widely scatteredPacific localities as Islas Revillagigedo, HawaiianLeeworda and. the Galpagos (Mellen, 1925; Wetmore,1925; Grant, 1927; Carr, 1952; Kenyon and Rice,1959; Parsons, 1962),

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

3 2

Several hypotheses have heen offered toexplain the banking habit anoag all kiods ofturtles, Ongle (1950), Neill and Alien (1954)and Boyer (1965) suggest that basking might be aform of thermo-regulation, aid ecdypis, free theturtle of ectoparasites or elimlnate epizoioalgae0 Pritchard and Greenhood (1968) suggestthat basking aids in the synthesis of Vitamin D.The habit may also aid digestion or it may be aprelude to oviposition (but in the HawaiianLeewards male green turtles bask too), In the

Gulf of Carpentaria female green turtles bankduring the mating season (Buntard, MS), It is

also possible that banking serves a socialfunction,

In view of the ever-expanding human popula-tion (and the quest for food) banking aggregationsof green turtles are mow at a disadvantage intheir struggle for survival, Banking has notbeen observed in Atlantic green. turtle populations.If the habit wan not more widespread. in earlieryears, its occurrence today in sosie populationsand mot others may be just behavioural polymor-phism.

Green turtles, like all reptiles, are cota-therme, Hjrth (1962) found that the cloacaltemperatures cf adult, female, green turtles atTortugilero, Costa Rica, upon emerging from thesea for oviposition, were very similar to thesea water temperature (average cf turtles 29,9 C;

range of sea water temperature 27,5-28,5°C).

1_Oreen turtle

4 POPULATION

4.1 Structure

4.11 Sex ratio

Natural sex ratios in green turtle populationsare virtually unknown. No studies have been madewhich elucidate the sex ratios in clutches ofhatchlings and virtually nothing is known aboutsex ratios in the juvenile and sub-adult popula-.-tiens. It is only when larger individuals arecaught on the feeding and breeding grounds thatbiologists are able to glean something of the sexratios and these data are not entirely accurate.

Hirth and Cs.rr (1970) provide evidence thatfemales outnumber males on some of the feedingpastures off Southern Yemen. An overall sex ratioof 56% females and 44% males is based on a totalof 4,376 individuals collected over a period ofseven years. The figures include sub-adults andadults.

Calduell (1962b) found that in Baja Californiawaters, over four sampling periods in which thenumber of landed turtles varied from 104 to 465,females comprised between 68 and 92% of the catch.Presumably these were taken on the feeding pasturesand were sub-adults and adults,

Cax'r axai Giovannoli (1957) examined twooargoes of green turtles caught on the feedingpastures off Nicaragua between February and April,end in one case there were 27 males and 66 females;in the second cargo were 105 males and 271 females,The sizes were not given, but all were probablysub-adults and adults.

Fxpert turtle fishermen in the Kingdom ofTonga have advised Hirth (NS) that females out-number males in the feeding pastures off Tongatapu.

11 is common to see more males than females,at least for part of the breeding seanon, off someof the major nesting beaches, Reasons for thisaz's discussed in section 3.13.

4.12 Size composition

According to Galdwell and Caidwell (1969),the Iscarcity of records of small turtles seen inthe ocean as compared with adult or sub-adultindividuals is a most intriguing and puzzlingcircumstance. While the apparent rarity maar resultfrom not looking in the proper places, it may bethat the lack of numbers is real. Considering thehigh mortality of the very young, the rapid growthof those that survive, and long life after reach-ing maturity, it is quite possible for thepopulation -to consist mostly of larger turtles".

Data relative to the sizes of nesting femalesare given in Fig. 6. Sizes of grazing males andfemales in the Gulf of Men are given in Table IX.

4:1

4.2 Abundance end deiiity (of popultioj

4.21 Average abundance

It is estimated that about 10,000 femalesmake up the entire population nesting on theSarawak Turtle Islands (Banks, 1937; Parsons,1962; Honegger, 1968). Schulz (1969) estimate-that there aro about 2,500 females in the entireSurinais population. Carr (1969b) calculatesthat about 6,000 females nest annually atTortu,guero. Hir-th and Carr (1970) estimate thatless than 1,000 females nest annually on Aldabr%.Dr. Bustard estimates the total breeding femalepopulation of the three common species (green,flatback and loggerhead) in Queensland waters tobe not less than 75,000 (Anon, 1970g).

Relative sizes of other nesting popula-tiónWcan be ascertained by referring to referencesgiven in Table II,

Cromie (1966) states that green turtlesrepresent about 20% of all turtles foundthroughout the warm seas of the world.

4.22 Changes in abundance (see section5.4 for a discussion of thesignificant decreases in sizes ofturtle populations)

4.3 Natality and recruitment

4.31 Reproduction rates

Details of egg production are provided inseotion8 3,16, 3.17 and 3.2. Some of theimportant aspects of reproduction pertaining topopulation studies are (i) a typical, adultfemale produces about 550 eggs each reproductiveseason and. this usually is every two or threeyears, (2) some femaleamay not retain reproduc-tive capacity -throughout life, (3) at some majorrookeries about half the eggs laid do no-t hatch,and. (4) on high density nesting beaches someincubating eggs are destroyed by turtles nestingla-t er,

4.32 Factors affecting reproduction:see sections 3 and 4.5

4.33 Recruitment (Some factorsdetermining recruitment such asgrowth and seasonal movementsare discussed In sections 3.4and 3.5)

4.4 ISrtalIty and morbidity

4.41 Nortality rates: see section3.22 and Fig.. 9

4.42 Factors causing or affectingmortality (Predation on eggs,young and adults has been

asoil in sections 3.11, 3.22and 3,34, The direct and indiresteffects of f jailing are discussedin section 5)

4.43 Factors affecting morbidity (Someparasites are listed in section3.35 and chelonitoxication isdiscussed in section 3.44)

4.5 Dynamics of populations (as a who]e)

Bustard and. Tognetti (1969) have produced amodel to show how nest destruction is dependent Onpopulation density and how this provides a mecha-ui.nm to regulate population size (see 'ig. 15).The mechanism tends to keep a population withincertain limits, They state that "at a low popu-lation density its effects are negligible; however,if a population increased greatly, it would notstay ai the new levo1 due o hia mec1iansm, whichtends to restoro the population to its originallevel." They also postulate that the BarrierReef region of Australia may be the last placowhore natural regulation of population size by themechanism of density-dependent neat destructioncan still be observed today. This is becaUse onmany other nesting beaches the species w beenseriously ovorexploited and the population densityis below the level at which nest destruction byturtles would operate as a regulatory mechanism,

Bustard and. Tognetti's model would also seemto be operational for such sites as the SarawakTurtle Islands where many eg, if not dug by man,would be destroyed by turtles nesting later.

4.6 The poLatiOn in the communitem

Green turtles are large, egg-laying, marine

F11U1JS85 Green turtle

reptiles wefl adapted to their aquatic habitat,but who periodically crawl onto dry land to laytheir eggs. At such times, when on their nest-ing beach, they may be the dominant nocturnalvertebrate of the strand association. Theircrawling and digging activities at least tempo-rarily affect the beach topography and theireggs and hatohlinga are a source of food for avariety of predators (see sections 3.21 and3.22).

The distribution of adult groen turtles inthe littoral and sublittoral zones of tropicalseas is directly re].aied to their nestingbeaches, feeding pastures and perhaps oceancurrents. Long-ren: biennial or triennialnesting remigrations across deep water aretypical for at leási some members of some popu-lations, i.e., the population of green turtlesin any given region may include two components,a migratory group, and a group that remains inthe area year-round and nests on the nearestoptimal beach (mee section 3.5).

The relationship of green turtles Withcompetitors in the "turtle grass community" isdicused in section 3.42. most adults occupythe second trophic level in a simple predatorfood chain, viz., plant-turtle. Hatchiinga andmaall juveniles are chiefly carnivorous andhence occupy at least the third trophic levelin a predator food chain, viz., green plant-herbivore-hatchling.

The green turtle occupies a broad ecolo-gical niche. This is partly due to the lackof' strong interspecific competition.

lOO 200 300TURTLE POPULATION

400

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

500

FIRM/335 Green turtle 4:3

FI- SB Green turtle

5 EXPLOITATION

5.1 Fishing eQuipment

5.11 Gears and methods

Indigenous methods of turtle fishing includediving for them and using harpoons, spearguns,traps, seines, auckerfish and decoys.

Green turtles are usually caught on theirfeeding pastures by- using nets and seines. Car-rand Caidwell (1956) describe how turtle fishermenseeking juvenile and sub-adult individuals on thegrass pastures off the Gulf coast of Florida usetangle nets from about 97 to 194 m long, 2.4 to3 n deep, and with a 20.3 to 30.5 cm bar mesh.

Duncan (1944) describes how Carman Islanderscapture turtles w-ith nets on the feeding pasturesoff Nicaragua, and Parsons (1962) further describeshow turtlers on the Nicaraguan pastures "set theirnets over marked rocks to which the turtlescustomarily retire at night and wait for them tobecome entangled in the nets when they surface toblow". Huxley (1962) gives a very brief accountof turtling by the Cayman Islanders.

The beach seines used by the Yexneni in theGulf of Men to catch sub-adults and adults varyfrom about 97 to 388 m in length and are up to12 m deep with a 10 to 46 cm mesh (Hirth and Carr,1970). According to Tressler and Lemon (1951) themost popular form of gear for catching turtles isthe gill net,

Vasa and Straatmans (1954) describe turtlenetting on the pastures in the Tonga Islands asfollows: "A fisherman, having located an area ofseaweed between the small, uninhabited islands,drops his net in a straight line at right angle tothe flowing current and returns home. After twodays he goes back to retrieve his net and usuallyfinds turtles of all sizes caught in the not. Ina 50 by 12 ft net, as many as ten turtles can beexpected and often more, depending upon the lengthand width of the net and the area where it isdropped".

Yonge (1930) has described how natives usedto dive for sleeping turtles off the Great BarrierReef. Some of the ways of fishing for turtles inthe Trust Territory are described in the Anthropo-logical Working Papers (1957).

Suckerfish or remera (Echeneis sp.) have beenused to capture turtles in such widely scatteredplaces as the South China Sea, northern Australia.,east coast of Africa and the Caribbean, Parsons(1962) gives an excellent account of this extrae-ordinary custom.

Scuba divers use Bpeargims to get greenturtles of various sizes in such places as theSeychelles Islands, Caribbean Islands, and some

South Pacific Islands.

Harpoons are widely usad to capture bothsexes when they conregate off the nestinggrounds. McCarthy (1955) describes, in a popu-lar way, how aboriginale harpoon turtles in theGulf of Carpentaria.

Schmidt (1916) described how the fishermenin the Dutch West Indies used wooden decoys inthe shape of a turtle, along with nets, tocapture green turtles. Wooden decoys are stillused in several areas of Central America. Theyare especially useful in attracting males.

On the nesting grounds adult females arosimply "turned on their backs", This can bedone by one man, No special equipment is neededto locate and dig up the eggs on the beaches,although a straight stick to probe for the eis useful.

Mechanized turtle hunting is becoming morewidespread. Two companies operate in the feed-ing pastures off western Australia and thefollowing account describes their methods(Anon, 1969e): "Each licensed freezer boat hasseveral anali 16-ft scooter catcher boats,powered by 40 hp outboard motors, These scooterboats operate within a one-mile radius of themother freezer boat, in the relatively shallowwater inside the offabare reefs, where theturtles graze on the brown and green algae ofthe rocky sea bed. When a turtle is located,it is harpooned from the scooter boats as itraces through water from 3 to 8 ft deep. Onattaining a full load of about 10 turtles, thescooter boats unload their catch on board thefreezer boat fo: processing, Turtles are gutted,beheadeit, washed, drained and blast frozen,Each carcass weighs about 120 lbs dressed. Whenthe freezer boal; attains a full load, usuallyabout 300 turtLes taken in about three days'fishing, it returns t.o port to unload itscatch".

5,12 Boats: see preceding section

5.2 Fishing areas

5.21 General geographic distribution

Host green turtles are caught on theirfeeding pastures, in shallow water just off thecasting beaches, and on the nesting beaches(see section 2).

5,3 Fishing seasons

5.31 General pattern of seasons: seesections 2 and 6

5.32 Dates of beginning, peak andend of seasons: see section 6and Table XI

Hawaiian Leewards

1iidwar IslandTrust Territory of

the PacificKingdom of Tonga

Fiji IslandaRose Atoll,

Lu ericen Samoa

Malarsia

Queensland,Australia

Mexico

Costa Rica

Panama

Surinam

renoh Guiana

Trinidad andTobago

kscension Island

British IndianOcean Territoryand SeyoheiiesIslands

Pacific Ocean and adjacent areas

complete protection for ail marine turtles and eggs(Anon, 1968b; Hendrickson, 1969; Laycock, 1970).protection for turtles less than 60 cm in carapace length.full protection for turtles and, eggs on shore; and in the water for turtleswith carapace length lese than 61 orn (Wilson, 1969).following regulations now wait ing approval by Government complet e year-aroundprotection for eg and for turtles with carapace length more than 87.5 cm;protection for all turtles of all sizes from I Novmaber 'to 28 February; ban onthe sale or export of anr turtle shell greater than 87.5 cm; complete protec-tion for the leatherback of all sizes at all tines,same regulations as for Tonga, awaiting Government approval.complete pLu(ection for turtles and eggs,

complete protection for nesting turtles; Government controls harvesting ofeggs (about one million ennuall7); (arch is closed season for eggs in Sabah(Harrisson 1950_1969; Hendrickson, 1958; de Silva, 1968, 1969b).full protection for all marine turtles end eggs (Bustard, 1969d),

Atlantic Ocean and adjacent areas

full protection for egga permit 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,adults and eggs fully protected on nesting beaches; and protection for turtleswithin 4.8 km of nesting beaches (but harpooners can operate beyond this limit),full protection for the green turtle (Myers, 1970),complete protection for nenti,ng 'turtles mid eggs on some of the major nestingbeaches (see text),protection for adults and eggs during May, June, July (Anon 19691;Pritchard, i969.protection for turtles and eggs from 1 June through 30 September,

full protection for turtles and eggs,

Ipdian Ocean and adjacent areas

complete protection for turtles end eggs (but see text).

Location Regulation

52 FIR1 E38 Green turtle

TABLE Xl

Current regulations regarding the harvesting of green turtles and their eggs

FIRM SB Green turtle

5.33 Variation in date or duration ofseason: see Tables II, III andsection 6

5.4 Fahin operations and. ret],.

Of the seven species of marine turtles, thegreen -turtle is of the greatest economic value toman. Its flesh, eg:, and "calipee" (= cartilaused to maki soup) have been eaten for centuries.Now the growing market for ibs skin and oil, coup:Led:ribh he oven greater need for, animal proteinespecially in the tropics, places -this reptile inen especially important but precarious position.,Carr Ç1952) called the green turtle "the moslvaluable reptile in the world."

The numbers of turtles caught on -the feeding.pas-bures as well as the numbers of females takenfrom the nesting beaches have declined rapidly inthe current oen-bury. Parsons (1962) provides acomprehensive cultural and historical survey ofman's attitudes toward the flesh arid eggs of thegreen turtle.

In some countries bordering lhe South ChinaSea, eggs rather than turtles are harvested. Banks(1937) reported that the Sarawak Turtle Islandsyielded. 2 119 912 eggs in 1927 and these h513 a valueof U.S. $3,O(0. Of these, 70% were consumed orsold locally, Ir: 1935, 98% of the 924,000 eggscollected (value U.S. $7, 769) were consumed locally.

Harriason (1962b, 192d, 1965a, 196Th) hasdocumented the decline of green turtles in Malaysiathrough an analysis of' egg-harvest records. Hopoints out that in the thirties a year with overtwo million eggs was commonplace, while in theearly and middle sixties a year with less thanthree-quarters of a million (only 99,307 in 1966)was commonplace, He speculates that the reasonsfor the decline to be both local (i.e., disturbanceof nesting females; increased small boatingactivity off nesting beaches) and. international(i.e., large steamers and. mechanized fishing fleetsin area; catching adults at sea).

During the peak months of nesting on Langawan,one of the Philippine "Turtle Islands", about45,000 eggs are collected and sold each month(Domantay, 1952-53). The same number of eggs, ormore, are harvested on at least two other islandsin the group. Risher, Simon arid Vincent (1969)estimate that one million eggs are collectedannually in the Philippines,

In recent years, two western Australia process-.in companies have undertaken commercial exploita-ion of the green turtle off the coast of westernaralia (Anon, 1969c). The areas fished comprise

the territorial waters extending along the coast-line for approximately 160 km north from southlatitude 23°10'. In 1968 and 1967, respectively,the export of processed turtle meat from westernAustralia was about 164,558 and. 135,922 kg. Thefishing season generally commences in June or Julyand terminates in September or October.

5:3

Parsons (1962) has documented the over-exploitation and demise of green turtles invarious parts of the world, specifically inBermuda, Dry Tortugas, the Caribbean Sea and inthe Indian Ocean, The decline of the greenturtle population in the Seychelles has beencommented on more recently by Honegger (1967),Gayiner (1968) and. Hirth and. Carr (1970).According to Acharji (1950), green turtles offCeylon are netted between November and March,-and the annual catoh is around 1,000,

An estimate of the economica of the 'centurtle fishery in some areas of the westernhemisphere is provided by Ingle and Smi-.h (1949).Statistics show that about 2,000 turtles wereoaugh-b annually in Costa Rica from about 1930bo 1948. Twenty-five men were en&ted in thefishery in 1948. These men caught turtles onthe nesting beaches between June and. September.The value of sea turtles exported from CostaRica over a 15-year period. varied. from $9,161in 1937 to $900 in 1942.

Murphy (1925) reported that a turtle fish-ery was opera-ted from time to time out of Pisco,Peru. Its existence today is unknown,

Cald.well (1963) gives a historical summaryof the turtle fishery in Baja California.Harpooning is the most common method of takingturtles in that area and. the fishery operatesyear-round.. Hubba (1960) also mentions theturtle fishery operating in that area. Townsend(1916) reported a catch of 162 green turtles ina single haul of a seine 90 m long in SanBartolome Bay in the Gulf of California on 11April 1889. rfley were reported to nest inApril and. Nay.

The average annual harvest of C.in. m,ydasfrom the east coast of Mexico between 1963 and1967 was approximately 225 t (R, Marquez, pers.comm.), The mean annual harvest of C.rn,carrinegra off western Mexico between 1963 and

was about 5,300 t (R. Marquez, pers. come,),

The development and decline of the greenturtle fishery in the Gull' of Mexico, Caribbeanarid British Honduras has been discussed by Inglearid Smith (1949), Westermann (1953), Carr (1954),Smith (1954), Caldwelland. Cari' (1957), Cari' andIngle (1959), arid Neill and Allen (1959). Cari'(1954) stated that "more than any other dietaryfactor, the green turtle supported the openingup of' the Caribbean".

An indication of the former abundance of'green turtles around the Bahamas end Cuba isgiven by Cateshy (1743).

The commercial catch of turtlea (almost allgreen) In Hawaii in 1969 was 4,625 kg (value U.S.$2 320). These figures should be consideredminimal because many more are taken but notreport ed..

Between 1953 and. 1967 up to 262 green turtleshave been sold annually in Papeete, Tahiti,

The Eijian's fondness for green turtles lawe lidocument ed in archaeological excava-t ions(Clifford, 1951).

Parsons (1962) estimated that between 15,000and 20,000 green turtles are taken annually tosupply coninercial markets. Reese (1917) mentionedthat in one year 15,000 were imported into England.

The catches of marine turtles have inoreasedeigsificantly in recent years (FAO. 1970). The

iThM/85 Green turtle

reported catches of sea turtlos of all speciesincreased from 20,000 metric tone in 1964 to 36and 33 thousand metric tons in 1967 and 1968,respectively. Whether these increases are realor due to better coverage in more recent yearsis unknown. Japan and !4exioo together accountedfor 95% of the landings (of ali species) in 1968.The breakdown by species is not known, but greensridleje and hawksbillø probably account for mostof the landings.

All the available evidence indicates thatgreen turtle populations are slow to recover fromovarexploitation.

FIPJV'S85 Green turtle

6 PROTECTION AND MANAGEMENT

6,1 Regulatory (legislative) measures

6.11 Limitation or reduction of totalcatch: see Table XI and followingsections

6,12 Protection of portions of popu.-lation

There are striking differences among variouscountries regarding protection of green turtles.There are no regulations at all pertaining to theircapture on the high seas. Current regulations,that were available to the author, are set out inTable XI, Various other recommendations and thephilosophy behind some of the ordinances arediscussed in the following regional sub-sections.

Pacific Ocean and adjacent areas

It should be noted. that the new Tonga and Fijiresolutions (see Table JOE) will provide protectionfor all eggs and for animals above a certain sizerather than below a certain size. The philosophybehind these ordinances is geared to the beliefthan an adult turtle remains reproductively activefor a considerable number of years and protectionof adults and their eggs should in time produce anincrease in the population. Turtle populations inFiji and Tonga should be monitored after these newlaws go into effect since the ordinances may serveas models for other areas in the South Pacific.(in Tonga and Fiji the carapace is measured alongits arch; straight line measurement iould placethe legal size a few centimetres below 87.5 cm).

As far as is known, the eggs are generallyeaten in Thailand, indonesia, Burma, Borneo andthe southern Philippine Islands; adults arepreferred in New Guinea and Melanesia (Penyapol,1958; Parsons, 1962; Harrisson, 1962d; Somadikarta,1968; and Anon, 1969d).

Atlantic Ocean and adjacent areas

In September and October 1969, representativesfrom Costa Rica, Nicaragua and Panama met in SanJose and agreed to suspend all turtling for aperiod of three years after which reappraisal ofthe status of the resource would be made andappropriate international controls could beestablished (Anon, 1970a; Carr, 197Oa). Ifsuccessful, this will be the first sigaificantinternational agreement on sea turtles,

About half of the big rookery at Tor-tuguero,Costa Rica will be enclosed in the new Costa RicaNational Park (Anon, 1968o; Anon, 1970h).

Complete protection is provided for adults aswell as for eggs on some of the major nestingbeaches in Surinam (Pritchard, 1969). On otherbeaches in the country, adults and their eggs areprotected only during certain seasons. The Galib:L

6:1

nesting grounds in Surinani were declared anature reserve in 1969 (includes Ellenti,Baboensanti, Pruimenboom and Galibi beaches),Lecal Caribs, however, are allowed. to take acertain quota of eggs each year.

Indian Ocean and adjacent areas

Legislation recently passed. by theSeychelles and. British Governments (August,1968) provides complete protection for bothsexes and. eggs -throuì.out the Seychelles Islandsand the British Indian Ocean Territory. Todaythere are no big, concentrated nesting placesin the Seychelles Islands, although in the lastcentury A.ldabra was one of the biggest greenturtle nesting grounds in the world. Thegeneral ecolor of Aldabra and its potentialrole as a site for a biological research station(in contrast to its proposed role as a militarylanding strip), have been discussed. by Stod.dart(1967, 1968a, 1968b) and Beamish (1970). Briefdescriptions of the Aldabran rookery areprovided by Abbott (1893), Keynes (1959) andHortmaz (1967). As in other places where thegreen turtle exhibits pronounced. migratorypatterns, the Seychellois are concerned. overthe fate of their turtles which now receive fullprotection on Seychelles beaches but whichmigrate elsewhere to feed and apparently get noprotection (Anon, 1970d.). The same situationled recently -to the Tripartite conference amongPanama, Conta Rica and Nicaragua (see precedingsection).

In and. near the Malagasy Republic,instituted. turtla reserves are located at NosyMambo, Nosy Iraxtja, Nosy Trozona, Nosy Va,Chesterfield. IsleZLd and. Europa Island. On someother beaches in. the Nalagasy- Republic it isforbidd.en to take sea turtles wheù they arelaying,

In Southern Yemen, green turtle flesh ismainly a commodity for export. In view of agrowing export trade and possibility of Over-exploitation, the Government of Southern Yemenwas advised. to limit the annual catch to 1 000,of which not more than half were to be females;and., to provide completo protection for nestingfemales and. eggs on the major rookeries in theeastern provinces (FAO, 1968). Whether or notthese recommendations have been adopted by theGovernment remains unknown,

Although there are no marine turtle con-.servation laws in West Pakistan, the greenturtle rookery at Hawke' s Bay seems in noimminent danger because local religious customsforbid. eating of turtle flesh and very fewindigents eat turtle eggs.

InSeptember, 1968,-the new African Conven-tion for Conservation of Nature and NaturalResources was sigeed by 38 heads of state ortheir representatives. All marino turtles are

6:2

placed in "Class A" which is defined as follows:"Class A species shall be totally protected through-out the entire territory of the contracting states,end the hunting, killing, capture or collection ofspecimens shall be permitted only on the authoriza.-tion in each case of the highest competent authorityand only if required in the national interest orfor scientific purposes", Ourry-Lindahi (1969)describes the events leading up to this Africanconservation convention.

6.13 Current status of the resource andaction programmes

The green turtle is currently listed as a"depleted species" in the I.U,C,N, Red Data Book(Honegger, 1968). Consequently, at least for thepresent time, the keeping of green turtles as petsshould be discouraged (cf. Murphy, 1968). Chelonianzydas is listed as a "peripheral" species in the1968 edition of Rare and Endangered Fish and Wild-life of the Thi-ted States (Anon, 1968a), and isincluded on Ziswiler's (1967) list of the mostgravely threatened animals.

The green turtle (and all other marine turtles)was mentioned repeatedly in conservation resolutionsadopted at the Second International Congress of theWorld Wildlife F\ind which met in London in November1970.

Many pleas for the wise management and conser-vation of green turtles have been made. Some ofthe more recent are those of Anon (1959), Hillaby(1965), Pritchard (1966), Carr (1967b, 1969d, 1970b),Bertram and Bertram (1968), Duniont (1969), Harrisson(1969b), Honegger (1969) and Crowo (1970),

In 1933 Moorhouse emphasized that much hasstill -to be learned about green turtles, especiallythe sex ratios in the nest, survival rates, whereyoung spend their lives, and growth rates. Fortyyears later biologists still know very little aboutthese four facets in the life of the green turtle,all of which are needed in order to establishrational management programmes.

Major action programmes recommended by a groupof sea turtle conservationists meeting in M,rges,Switzerland (10-13 March 1969) included (i) increasedincubation and hatchiing programmes, using proventechniques, (2) study and analysis of worldexploitation patterns, (3) a broad informationprogramme, (4) beach surveys where data are lacking,followed up by expert advice as required, and (5)establishment of special sanctuaries underscientific management. The recommendations receivedmuch publicity (for example, Anon, 1969k; Anon,1970b; and Anon, 1970c),

It is this author's opinion that in additionto the recommendations given above, major researchthrusts should be directed -toward populationdynamica, productivity and studies on the ontogenyof orientation,

PIFLM/S85 Green turtle

The establishment of breeding sanctuariesis certainly one way to perpetuate the stocks.Ascension in the South Atlantic, Europa andAldabra in the Southwest Indian Ocean, RoseAtoll in the South Pacific and French FrigateShoal in the Hawaiian Archipelago are goodexamples of such breeding reserves.

A series of marine national parks couldalso be established in key places where marineturtles are especially abundant. Kenya, forexample, has two marine parks ai-id is consideringa third (see Randall, 1969, for discussion ofmarine parks).

The problems manifest in green turtlemanagement and conservation are indeed complex.Factors that must be considered include turtlebiolo-; local customs and religions; demo-graphic trends; economics; federal regulations;and, perhaps most important of all, interna-.tiorial cooperation. The fact that fishingjurisdiction claimed by nations varies fromabout 5 km to 321 km complicates any rationalplan for the management of any marine turtle.

6.14 Public education

Because education plays a large part inany conservation and management programme,there should be more pictorial accounts likeJacques Cousteau's three film loops on thereproductive biolor and conservation of thegreen turtle (available from OceanographyUnlimited, Lodi, New Jersey) and more filmslike -the exoellent turtle conservation filniproduced in Sabak and shown on televisionaround the world (Bill Burred Television Co.,Los Angeles, California).

"Solomon, tue Sea Turtle" is a semi-.documentary film which portrays the homingprowess of a green turtle (see Anon, 1969e),

6.2 Control or alteration of physìcalfeatures of the environment

The encroachment of vegetation and theaccumulation of debris on some nesting beacheshas been mentioned in section 3,16.

A study of the physical parameters of good.turtle pasture is needed (see sections 2.22 and3.42).

Bustard and Greenham (1968) found that onthe Heron Island beaches no-t all egg chambersdug are successful. Failures are caused byobstruction of the digging action of the turtle1)7 large or thickly matted tree roots and bycollapse of the egg chamber sides due toinsufficient support of the sand.

FIRJ'/S85 Green turtle

6. Control or alteration of chemicalfeatures of the environment

Aocord.in to laboratory experiments of Bustariland. Groenhaxn (1968), green turtle eggs are able tohatch in chloride concentrations very much greater.than those occurring in the natural egg chamber.Chloride concentration is not, therefore, alimiting factor in incubation success except forthose few nests which aro laid below the springhit tide mark. Bustard. and Greenham also declarethat they have successfully incubated many turtleeg in coral sand moistened only with distilledwat er.

6.4 Control or alteration of biologicalf satures of the environment

A variety of species management programmesare in existence (see section 6.12).

Hendrickson (19589 1961) believes thatexploitation of eggs has a much less adverse effecton turtle populations then slaughter of adultfemales and that the exploitation for eggs is moreprofitable in terms of human nutrition than isexploitation for flesh and fat, Hendrickson'sstrategr of harvesting turtle eggs rather thanmeat can be briefly summarized as follows: theadult female lias a high reproductive potentialand the species is adapted to sustain enormouslosses a-t the very early stages of it-s life. Theharvesting of eggs constitutes exploitation at astage when the species is adapted to withstandsuch losses. The individuals which do survive toadulthood presumably remain reproductively activefor a considerable number of' years, If the averagefesale experiences more than three breeding seasonsin a lifetime, then the waight of protein availablein the form of eggs would. exceed the weightavailable in the form of flesh and. fat. Theelegance of Hendrickson's schomeat least inMalaysia, revolves around the fact that it is alsoin harmony with local customs and religion.

Hendrickson and Alfred (1961) estimated thatabout two million eggs constituting something over50 t of high grade protein were consumed each yearin Malaysia and -they also stated "preliminarystudies have indicated that as little as 2 or 3%of the nests, handled properly, could probablyprovide sufficient recruitment to the wild popu-lations to keep them at their present level,provided other circumstances remain unchanged".

Since in some arena of the tropics about halfof the eggs which are laid do not hatch, it wouldbe of' immense value to devise a method ofdistingaishing fertile from nonfertile eggs,These nonfertile eggs could then be utilized forhuman consumption.

Caz'r (196Th) stated that after over a decadeof research at Tortuguero, Costa Rica, more greenturtles are nesting now than in previous years,

6:1

chiefly because of the increased hatohling yield.At Toriuguero adults are taken 'for localconsumption and eggs are seldom harvested.

Although little is known regarding thenumbers o males in a natural population, it maybe -that some of the hunting pressure on femaloccould be relieved by cropping surplus maleo,

6.5 Artificial etockin

Mere than 100,000 ha.tchling green turtlesfrom Tor-tuguero have been distributed to 28different localities in Central and South Morion,Mexico, Nest Indies, Bahamas, Florida end Texasby the Caribbean Conserva-tien Corporation (Dr,Archie Carr, Technical Director) in hopes ofreplenishing turtle stocks. Mast of -thereleases are on sites where green turtlesformerly nested in abundance. Batches of'hatchiinga are released with the hope that theygo back to the place released. .-drawn there bysome kind of sensory imprinting to 'baste, smellor feel. To date there are no indications ofdefinite success; however, at some release sitesit appears -that some of -the hatchuings are stay-.ing on and. establishing resident populations(Carr, 1967b). A pictorial account of therestocking programme in the Caribbean is givenby Parsons (1962) and. Marquez (1966), Restock-ing programmes in Florida h-ave been discussedin a popular fashion by Stephens (1968).

The "Brotherhood of -the Green Turtle" is aloosely knit group whose efforts to s-ave thegreen turtle led to the founding of the CaribbeanConservation Corporation.

Turtle hatcheries also operate in Malaysiaand Australia. Harrisson (1950_1969) describesthe long established hatcheries on the Ssrawakturtle islands, In 1968 more than 100,000 eggswere dug up and reburied in the Sabeh hatchery.An egg hatchery has been in opera-tion forseveral years in Australia (&zs'tard and Greriham,1968),

The current practices are to releaseha-bchlings either on the beach or in the waterboth in the day-time and at night, Some arereared for several days or weeks and thenreleased in the sea just off the rookeries,This latter procedure reduces mortality fromlittoral predators. However, what effectsholding batchlings in captivity and thenreleasing them at sea have upon their naturalbehavioùr patterns are unknown.

At the present time the best (and simplest)method of natural stocking is to provideprotection for mes-bing females, eggs andhatchlins on the rookeries. If predation oneggs and/or hatchuings is high and cannot becontrolled, and in those few places ehero nestingdensity is so high that lato nesters destroy the

6:4

nests of their predecessors, then tile establishmentof centralized egg hatcheries on the nesting beachis justified Hatchlinge produced in thesehatcheries should be released as soon as possibleafter emerging on the surface. Although, aspreviously mentioned, the methods of releasinghatchllngs are debatable, i.e., whether to releasethem on the beach and let them crawl to the sea,

FIRN/585 Green turtle

or to release them from boats offshore, therebyeliminating littoral predation, this authorfavours the former because it most nearlyapproximates the natural behaviour of the animal.

The transplanting of eggs or hatelilingsoutside their natural nesting areas should bediscouraged until sorne evidence of success isobtained in the Caribbean restocking scheme.

FIRM/S85 Green -turtle

7 TURTLE MARICULIPURE

Oreen turtle ranching and farming are stillin the pioneer stages of development. True turtlefarming implies that the unit is completelyindependent of wilcistock (turtles live and mateunder controlled conditions and isy eggs on man-medo beaches), In turtle ranching the unit isdependent upon ,ild popu:'.ationc for eggs end theturtios arc kept in varying d.oeeo of oaptiv-lby(i.e.,, from freellving lndivduolo in feuoc1-.offpcittu'oo to completo containment in oonoiotc poo:Lu) o!Iost turtle biolo rite believe the;t turtle ronchin,ii' aHiomptcd t' all, ohould be only o tempcu'nvycohortio, the reaco» being that nalitu-al pop tiationoiil1 not withritceid ooatiaucil puaeitisatio&' cl'Iiiü oge. Oar (1969a) pointu out anotherclololieriouc effect of the p-cnature oprued ofturtle n oul'e and that io the rico of nowmerkotu und hih aioeu before the volume prodno..tien n000euuy to oateíy demando and relievoprouutu'o on natuael popula-tionri io achieved0 Necd.loso to coy, lilie ultimatu lu turtle inarisulturo io

riiug proven too}iuiqueri of agvi culturecina animal huobenthy. iwlliial o.bbonlptc at turtleequacalliuro uhould be on a emuli coule and. shouldbe conducted only if iioientifio expertisa ioavailable0

Some of the foctoru whioli uuot be oonoidercdbeforo initiating a turtle farei (or ranch) are thefollowings lon-torin financing, elimination ofpredators end compotibore in onclomtu'ss and onnesting beaches, protection from pollutants and.poachers, availability of veterinarians to adviseon disease problems, year-round availability ofnatural food (in nature, green turtle ha-bchlingsare chiefly carnivorous and adults are herbivorous).,growth rates obtainable with commeroia:L feed, waterand sanitary requirements, optimum density,pollution abatement, transportation to processingplants, expertise on how to utilize all turtleproducts (flesh, oil, skin and shell) and facilitiesto conduct scientific experiment s concomitantlywith the commercial scheme.

Sholbourne (1970) describes some prioritiesam far as marine fish cultivation is concerned amithe same would seem to apply to turtle culture.In addition to some of the factors mentioned above,he describes the provisions of "high quality" seedstock and physi co-chemical requirement s.

Iversen (1968) describes some of the advantagesof mariculture including the control of predationand artificial selection for rapid growth, highfecundity and resistance to disease. He alsodiscusses current techniques of milkfiah and. mulletculture and mentions the green turtle as a candidatefor sea farming.

Some general principles of aquaculture alongwith some of the problems and a few selective fishmodels have been discussed by Ba.rdaeh and ETther(1968).

The feasibility of turtle ranching has beenmentioned in a general way by Schroeder (1966).Ehrenfeld (1970) postulates that turtle ranchingmay have sorno promise if the sub-adults andadults can be induced to stay on natural pasturesnear a farm without being penned.

?'wbray (1965) reports that green turtlesmats regularly in the Waikiki Aquarium and thateg'i are dropped in the water. He believes thatthey would deposit their egi on a suitableartificial beach if one w provided.

Paruouri (i 9ci2) ropox-te how ycn'ng Southluiiericuu 'ivor ttu'tlou, Podoeuumiri ocrriiq, havebeen lirunuplanbccl nio L'-ko Valenoiu in the!lndoau higulaiithi end aro vowi1ig endthere, They uro »hic ubilicing artificialrLesting bochocj that have been oonritruotc onuomo 0± the lobado of thu lake.

7:1

)ioiiiuniI. bueli: toapino have buon riurcdcn'0000riL'uhly in tue IloibId tatoc miii methodeuced haro eiuy UOVO au addou for cou turtles,A brief modal l'or tur'apiu rouchuzig io Given bynglo cud mltui (1949).

borlcmr curjto. ulouu hava boon LLMCdfor 17 yoaro in the Unitccl States (Han, 1970)and. corno of the methode developed in l'arìing thrispecies might be adapted to oca tux't].oc,

Pinchot (1970) disousoec the pocalbility ofenriohing the lagoono of atolls and raisingvarious animal species therein, His ideas meritconsideration and would be especially applioabloin the South Pacifie reg-ion.

It may even prove possible as well aseconomical to grow certain plants via hlroponicsfor captive turtles, Several large European andAmerican zoos have employed these methods toproduce green grasc supplement for zoo animals.The zoo keepers believe that high nutritiongrasses may also stimulate reproductive processesand thus increase fecundity,

Preliminary experiments In Sarawak indicatethat the first six months are the most difficultfor rearing hatchiinge in captivity (Harrisson,1956a).

Significant pilot attempts at rearingloggerhead turtles in captivity (and resultinggrowth rates) have been described by Uehida(1967),

A pilot turtle ranch has been establishedon Great Inagua Islzid in the Bahamas, under theausp:Lces of the Caribbean Conservation Corporation.Here about 1,000 ha of turtle grass pasture havebeen walled in and hatchlings have grown up to3.6 kg in two years on a diet of conch (Gai-r,196Th),

Five Email green tur-ble ranches ara noti inoperation in Parres Strait Australia under thedirection of Dr, H.R, Bustard (Anon 1970fDuatard, MS).

A pilot turUe ranch (or farm) is alsooperating in Cuba. Details are lackin'.

At the present time a large turtle farn(iculture, Ltd., with about 40,000 turtles ofvarious aizea) is in operation on Grand Cayman,Dritish West Indies. Initia], reports indicate ahi.. degree of success in some aspeobs of the

operation. Popular acoowita of Maricmlturo areprovided by Gabier (1969), Anon (1969g) andAnon (1969j).

According to scott (1970) there are severalmajor economic obstacles io the development ofmarine aquaculture, including (i) the absanoc ofa strong demand for the high cost producta ofaquaculture (except for luxuries or freshwatercarp) in spite of the rising world population,and (2) the absence of property rights in naturalwaters and problems of international jurisdictionon the high seas.

*2 S8'5 Groen turtle

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Brattstrom, B.H,, Notes on tue herpetolo of the Revillagiged.o Islar3i, 1Iexioo. Aw,Eidl,Nat,,1955 54:219--29

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This is one of a series of documents issued by FAO, CSIRO and USFWS concerning species and stocks ofaquatic organisms of present or potential economic interest. The primary purpose of this series is to makeexisting information readily available to fishery scientists according to a standard pattern, and by so doing also todraw attention to gaps in knowledge. lt is hoped that synopses in this series will be useful to other scientistsinitiating investigations of the species concerned or of related ones, as a means of exchange of knowledge amongthose already working on the species, and as the basis for comparative study of fisheries resources. They will bebrought up to date from time to time as further information becomes available either as revisions of the entiredocument or their specific chapters.

series of documents are:

Fisheries Synopsis No.replacing, as from 1.1.63, FAO FisheriesBiology Synopsis No.

CSIRO Fisheries Synopsis No.and

USFWS FAO Fisheries Synopsis No.

SYNOPSIS OF FISHERIES BIOLOGICAL DATA

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 in this task.Additions and corrections to synopses already issued will also be most welcome. Comments including suggestionsfor the expansion of the outline and requests for information should be addressed to the coordinators and editorsof the issuing organizations.

FAO:Fishery Resources DivisionMarine Biology and Environment BranchFood and Agriculture Organizationof the United NationsVia delle Terme di Caracalia00100 Rome, Italy

USFWS: CSIRO:Chief, Scientific Publications UnitNational Marine Fisheries ServiceBuilding 67, U.S. Naval Support ActivitySeattle, Washington 98115, U.S.A.

Scientific EditorCSIRO Division of Fisheries and OceanographyBox 21Cronulla, N.S.W.2230 Australia

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

synopses in this series have been issued since January 1970:

Synopsis of biological data on the rainbow prawn, Parapenacopsis sculptiis(Heller, 1862)

Synopsis of biological data on the school prawn, Metapenaeus mac/say!(Haswell, 1879)

Synopsis of biological data on chum salmon Oncorhynchus keta (Walbaum) 1972Synopsis of biological data on the eel Anguilla anguilla (Linnaeus) 1758

Synopsis of biological data on the common shrimp Crangon crangon (Linnaeus, 1758)

Synopsis of biological data on the shrimp Panda/us montagui (Leach, 1814)Synopsis of biological data on the Jumbo tiger prawn Penaeus monodon Fabricius,1798

Synopsis of biological data on the Indian prawn Penaeus indicus H. MimeEdwards, 1837

Synopsis of biological data on the prawn Panda/us platyceros Brandt, 1851

Synopsis of biological data on the penaeid prawn Solenocera indica Nataraj, 1945

Synopsis of biological data on the penaeid prawn Metapenaeus dobsoni (Miers,1878)

Synopsis of biological data on the penaeid prawn Metapenaeus affinis (H. MimeEdwards, 1837)

FR/S

FB/S

D FO/S

NMFS/S

1970

1970

July 1970October 1970

October 1970

October 1970

October 1970

October 1970

October 1970

October 1970

October 1970

October 1970

The relevant

FAO

The following

DFO/S4

DFO/S5

BCF/S41

FIRI/S80Rev. 1

* FIRM/S91

* FIRM/S92* FIRM/S93

* FIRM/S94

* FIRM/S95* FIRM/S96

* FIRM/S97

* FIRM/S98

* FIRM/S99 Synopsis of biological data on the ocean shrimp Panda/us jordani Rathbun, 1902* FIRM/S100 Sinopsis de datos bìológicaos sobre el camarón blanco Penaeus schmitti Bur-

kenroad, 1936

* FIRM/Si 01 Synopsis of biological data on the white shrimp Penaeus setiferus (Linnaeus,1767)

'' FIRM/S102 Synopsis of biological data on the brown shrimp Penaeus aztecas aztecus ives,1891

* F1RM/S103 Synopsis of biological data on the pink shrimp Penaeus duorarum duorarumBurkenroad, 1939

* FIRM/S104 Synopsis of biological data on the penaeid prawn Metapenaeus tnonoceros(Fabricius, 1798)

* FIRM/Si05 Synopsis of biological data on the penaeid prawn Metapenaeus brevicornis(H. Milne Edwards, 1837)

* FIRM/5106 Synopsis of biological data on the penaeid prawn Parapenaeopsis stylifera(H. Milne Edwards, 1837)

* FIRM/S107 Sinopsis del camarón nailon Heterocarpus reediBahamonde, 1955

D FO/SS Synopsis of biological data on the greentail prawn Metapenaeus bennettae Racekand Dall, 1965

DFO/S7 Synopsis of biological data on the eastern king prawn Penaeusplebejus Hess, 1865

DFO/S8 Synopsis of biological data on the banana prawn Penaeus merguiensis de Man,1888

FIRM/S83 Synopsis of biological data on Saccorhiza polyschides (Lightf.) Batt.FIRM/S82 Synopsis of biological data on North Atlantic sandeels of the genus Ammodytes

(A. tobianas, A. dub/us, A. americanas and A. marinus)NMFS/S79 Synopsis of biological data on Pacific Ocean Perch, Sebastodes alutus (Gilbert),

Cramer (1895)

FIRM/S38 Synopsis of biological data on knobbed wrack Ascop/iyllum nodosum (Linnaeus)Rev. 1 Le Jolis

FIRM/S84 Synopsis of biological data on haddock Melanogrammus aeg/efinus (Linnaeus,1758)

FRm/587 Synopsis of biological data on Laminaria hyperborea (Gunnerus) FoslieF1RM/S85 Synopsis of biological data on the green turtle Chelonia mydas

(Linnaeus) 1758

October 1970

October 1970

October 1970

October 1970

October 1970

October 1970

October 1970

October 1970

October 1970

1970

1970

1970

October 1970

November 1970

December 1970

December 1970

December 1971

December 1971

December 1971

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

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