SYNOPSIS OF BIOLOGICAL DATA

NOAA Technical M m andum NMFS

SYNOPSIS OF BIOLOGICAL DATA ON THE GREEN TURTLE

IN THE HAWAIIAN ISLANDS

George H. Balazs

fi

m

NOM-TNHVMFS-S WFC-7

October 1980

U.S. DEPARTMENT OF COMMERCE National Oceanic and Atmospheric Administration National Marine Fisheries Service

Southwest Fisheries Center

NOAA Technical Memorandum NMFS

The National Oceanic and Atmospheric Administration (NOAA) was organized in 1970. It has evolved into an agency which establishes national policies and manages and conserves our oceanic coastal, and atmospheric resources. I t provides managerial, research, and technical expertise to produce practical services and essential information for the programs concerned with such resources.

The National Marine Fisheries Service (NMFS) provides the United States with an integrated program of management, research, and services concerned about the protection and rational use of living marine resources for their aesthetic, economic, and recreational value. NMFS determines the consequences of the naturally varying environment and human activit ies on living marine resources. NMFS provides knowledge and services to foster the efficient and judicious use of those resources. NMFS provides for domestic and for international management and conservation of these living resources of the sea.

To carry out i t s mission, the organization also provides for communication of NMFS information. In addition to i t s formal publications, NMFS uses the NOAA Technical Memorandum series for informal scientific and technical publications. These documents are specialized reports that require multiple copies when complete formal review and editorial processing are not appropriate or feasible. The management and control of Technical Memorandums has been delegated to the Research Centers, Regional Offices, and corresponding staff offices within NMFS. Therefore, requests for copies of Technical Memorandums should be sent to the author or to the originating office for the material.

4

c

c

NOAA Technical Memorandum NMFS

SYNOPSIS OF BIOLOGICAL DATA ON THE GREEN TURTLE

IN THE HAWAIIAN ISLANDS

George H. Balazs

National Marine Fisheries Service Southwest Fisheries Center

Honolulu Laboratory Honolulu, Hawaii 9681 2

October 1980

U.S. DEPARTMENT OF COMMERCE Philip M. Klutznick, Secretary

National Oceanic and Atmospheric Administration Richard A. Frank, Administrator

National Marine Fisheries Service Terry L. Leitzell, Assistant Administrator for Fisheries

iii

Adult female Hawaiian green t u r t l e basking on T r i g I s l a n d a t French F r i g a t e Shoals . The s m a l l depth recorder a t t a c h e d t o t h e carapace w a s used t o measure maximum d i v i n g depth during t h e i n t e r n e s t i n g i n t e r v a l . (Photograph by G. H. Balazs , June 1979.)

V

TABLE OF CONTENTS

Page

TNTRODUCTION ................................................................ 1

1. IDENTITY

1.1 Nomenclature and Taxonomy. ........................................ 2

1 . 2 Common Names...,....... ........................................... 4

1 . 3 S i z e Categor ies ................................................... 4

1.4 Tagging and Measuring ............................................. 4

2 . DISTRIBUTION

2 . 1 T o t a l A r e a . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

2.2 D i f f e r e n t i a l D i s t r i b u t i o n . . . . . . . . . . . . . . . . . . . . . . . . . . . . , , . . . . . . . . . . . 7

2 . 2 1 Adults ...................................................... 7 2.22 Hatchl ings, j u v e n i l e s and subadul t s ......................... 8

FL

a

3. ECOLOGY AND LIFE HISTORY

3.1 Reproduction ...................................................... 9

3.11 Sexual i ty ................................................... 9 3.12 Maturi ty .................................................... ' 9 3.13 ~ t i n g ...................................................... 10 3.14 F e r t i l i z a t i o n . .............................................. 11 3.15 Nesting and h a b i t a t c h a r a c t e r i s t i c s . . . . . . . . . . . . . . . . . . . . . . . . . 11 3.16 Reproductive c y c l e s ......................................... 14 3.17 Eggs .......................................... "............. 15

3.2 Hatchl ing Phase................................................... 16

3 . 3 Juveni le , Subadult and Adult Phase. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1 7

3.31 Longevity ................................................... 1 7 3.32 Competitors ................................................. 18 3.33 Predators................................................... 18 3.34 Epizoics , d i s e a s e s , abnormal i t ies and i n j u r i e s . ............. 20

3.4 N u t r i t i o n and Growth.............................................. 23

23 25 27

3.41 Food s o u r c e s . . . . . . . . . . . . . . . . . . . . . . . . . , . . . . . . . . . . . . . . . . . . . . . . 3.42 Feeding behavior . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . * . . . . . . . . . . 3.43 Growth rates................................................

V i

Page L

3.5 Post-Hatchling Movements .......................................... 29

3.51 Dispersa l and developmental migrat ions ...................... 29

3.53 Navigation and o r i e n t a t i o n .................................. 32 3.52 Remigrations ................................................ 31

3.6 Basking ........................................................... 34

4 . POPULATION

4 .1 S t r u c t u r e ......................................................... 38

4.11 Sex r a t i o ................................................... 38 4.12 S i z e composition ............................................ 39

4.2 Abundance and Density ............................................. 40

4.3 Reproduction Rates ................................................ 41

a-

c

4.4 Mor ta l i ty ......................................................... 41

4.5 Recruitment ....................................................... 42 c

5 . EXPLOITATION

5 .1 H i s t o r i c a l Overview ............................................... 43

5.2 Commercial Catch Reports .......................................... 45 Y

5.3 Noncommercial Catch ............................................... 47

6 . PROTECTION AND MANAGEMENT

6.1 Regulatory Measures ................................................ 48

6.2 Management P r a c t i c e s and Options .................................. 49

6.21 A r t i f i c i a l s tocking ......................................... 49 6.22 P r e d a t o r c o n t r o l ............................................ 50 6.23 Enforcement and educat ion ................................... 51 6.24 Aquaculture ................................................. 51

BIBLIOGRAPHY ................................................................ 53

a

SUBJECT INDEX ............................................................... 98

v i i

LIST OF TABLES

Page

Number of CheZonk tagged throughout t h e Hawaiian Archipelago s i n c e 1973......................... .............. 100

Y

L

Table 1.

Table 2.

Table 3.

Table 4.

Table 5 .

Table 6 .

Table 7.

Table 8.

Table 9.

Table 10.

Table 11.

Table 12.

Table 13.

Table 14,

S t r a i g h t carapace l eng th and width of a d u l t female green t u r t l e s a t French F r i g a t e Shoals . . . . . . . . . . . . . . . . . . . . . . . 101

Curved carapace l eng th and width of a d u l t female green t u r t l e s a t French F r i g a t e Shoals . . . . . . . . . . . . . . . . . .oe . . . 102

Summary of growth rates and p r o j e c t e d y e a r s t o ma tu r i ty f o r green t u r t l e s sampled a t r e s i d e n t fo rag ing areas i n t h e Hawaiian Archipelago ..................................... 103

Number of egg c l u t c h e s l a i d by green t u r t l e s a t East I s l and , French F r i g a t e Shoals ................................ 104

I n t e r n e s t i n g i n t e r v a l s of green t u r t l e s a t East I s l a n d , French F r i g a t e 105

Resul t s of 40 precounted egg c l u t c h e s excavated and examined a t East I s l and fo l lowing the na tura l emergence of ha t ch l ings . . . . . . ................................ 105

Number of n e s t i n g green t u r t l e s tagged and subsequent ly recovered a t East I s l a n d , French F r i g a t e Shoals, 1973-1979... 106

Food sources recorded f o r green t u r t l e s in t h e Hawaiian Archipelago .................................................. 107

Major food sources of green t u r t l e s i n t h e Hawaiian Archipelago .................................................. 110

Growth rates of Hawaiian green t u r t l e s r e l eased from c a p t i v i t y and r a i s e d i n captivity... . . . . . . . . . . . . . . . . . . . . . . . . . 111

Growth rates of a d u l t green t u r t l e s recovered a t French F r i g a t e Shoals s i n c e June 1973............................... 112

Growth r a t e s of Hawaiian green t u r t l e s captured from t h e wi ld and h e l d f o r extended pe r iods i n c a p t i v i t y a t Sea L i f e Park.................,...........................~~.o~~~~ 113

Movements of Hawaiian green t u r t l e s i n t h e main i s l a n d s a f t e r being he ld i n c a p t i v i t y f o r extended periods. . . . . . . . . . . 114

v i i i c

Page

Table 15. Locat ions and numbers of t ag r ecove r i e s documenting long d i s t a n c e migra t ions by a d u l t green t u r t l e s i n t h e Hawaiian Archipelago ..................................... 115

Table 16.

Table 1 7 .

Table 18.

Long d i s t a n c e migra t ions of a d u l t Hawaiian green t u r t l e s recovered s i n c e June 1973.. . . . . ...................... 116

Mean weights of t u r t l e s from t h e Hawaiian I s l a n d s r epor t ed as being commercially captured , 1948-1973. .......... 118

Maximum numbers of green t u r t l e s seen basking a t one t i m e dur ing surveys i n t h e Northwestern Hawaiian I s l a n d s s i n c e 1950 ................................................... 1 2 1

Table 19. Tiger sha rks , GaZeocerdo cuvier , captured a t l o c a t i ons i n t h e Hawaiian Archipelago where t u r t l e s w e r e found t o be prey i t e m s . ............................................ 122

Table 20.

Table 21 .

Monthly number of t u r t l e s s o l d i n t h e Honolulu F ish Market, 1900, 1902-1904 ...................................... 123

Kilograms and percent of t u r t l e from the-Hawaiian I s l a n d s r epor t ed as be ing commercially captured by va r ious f i s h i n g methods, 1948-1973 ........................................... 124

Table 22. Kilograms and pe rcen t of t u r t l e from t h e Hawaiian I s l a n d s r epor t ed as being commercially captured by month, 1948-1973 .................................................... 128

L I S T OF FIGURES

Hawaiian Archipelago ......................................... 129 Figure 1.

Figure 2.

F igure 3.

French F r i g a t e Shoals ........................................ 130

Some of t h e important r e s i d e n t areas i n t h e main i s l a n d s where Hawaii-an green t u r t l e s feed and rest . . . . . . . . . . . . . . . . . . . 131

Figure 4. Surface d r i f t t r a j e c t o r i e s r ep resen t ing t h e t h e o r e t i c a l movement of h a t c h l i n g and pos t -ha tch l ing Chelonia l eav ing French F r i g a t e S h o a l s . .............................. 132

Sea s u r f a c e temperatures a t French F r i g a t e Shoals . . . . . . . . . . . . 133

c

Figure 5.

F igure 6 . Locat ions of 220 n e s t s recorded o n East I s l and from June t o August of 1974... . ................................... 134 P

i x

Figure 7 .

Figure 8.

F igure 9.

F igure 10.

F igure 11.

Figure 1 2 .

Figure 13.

Percent of green t u r t l e s s u c c e s s f u l l y n e s t i n g each n i g h t a t East I s l and , French F r i g a t e Shoals. . . . . . . . . . . .*

Tag r e c o v e r i e s of f i v e of the 4 1 green t u r t l e s r e tu rned t o t h e wi ld a f t e r extended pe r iods i n c a p t i v i t y .................................................... L i f e h i s t o r y and h a b i t a t model f o r Hawaiian Chelonia.........

Long d i s t a n c e movements of a d u l t green t u r t l e s In t h e Hawaiian Archipelago ......................................... Inc idence of d a i l y basking a t East I s l and , French F r i g a t e Shoals. .............................................. Number of females n e s t i n g annua l ly a t French F r i g a t e Shoals . . . . . . . . . . . . . . . ........................................ Percent t u r t l e r e p o r t e d as being commercially captured f o r each of t h e main i s l a n d s , 1948-1973.......,,.............

Page

135

136

137

138

139

140

141

INTRODUCTION

x

The green t u r t l e , CheZonia mydas, is t h e p r i n c i p a l marine t u r t l e s p e c i e s i n t h e Hawaiian I s l a n d s . It has been t h e r e c i p i e n t of n e a r l y a l l r e sea rch conducted t o d a t e w i t h i n t h i s reg ion of t h e P a c i f i c . Although s i g n i f i c a n t advances have been made dur ing r ecen t yea r s i n t h e knowledge of Hawaiian CheZonia, important a s p e c t s of t h e t u r t l e ' s n a t u r a l h i s t o r y remain t o be e luc ida ted . The purpose of t h i s synops is i s t o b r i n g toge the r a l l of t h e b i o l o g i c a l in format ion p r e s e n t l y known on t h e popula t ion (as of September 1979) and, i n doing s o , focus a t t e n t i o n on f u t u r e r e sea rch needs. c h a r a c t e r i s t i c s of Hawaiian Chelonia d e t a i l e d i n t h i s synops is w i l l a l s o serve t o emphasize t h e e x c e l l e n t r e sea rch o p p o r t u n i t i e s f o r ga in ing informat ion app l i - c a b l e t o a l l green t u r t l e popula t ions . The informat ion presented has been der ived from a comprehensive review of t h e l i t e r a t u r e and from unpublished o r i g i n a l r e sea rch conducted by t h e au thor .

The unique behav io ra l and popula t ion

The format used i n t h i s synopsis is pa t t e rned a f t e r t h e synops is series on s p e c i e s and s t o c k s i s sued by t h e Food and Agr i cu l tu re Organiza t ion (FAO) of t h e United Nat ions. A synops is of b i o l o g i c a l d a t a dea l ing w i t h t h e green t u r t l e on a g l o b a l b a s i s w a s publ ished i n 1971 as p a r t of t h i s series (Hi r th 265). The p resen t synops is complements and updates t h i s earlier document, which d e a l t only i n gene ra l terms w i t h Hawaiian Chelonia. synopsis is t h e inco rpora t ion of an ex tens ive l i s t of r e fe rences covering Hawaiian c u l t u r a l and legendary a s p e c t s of marine t u r t l e s .

An a d d i t i o n a l f e a t u r e of t h e p re sen t

The two o t h e r s p e c i e s of marine t u r t l e s found i n t h e Hawaiian I s l a n d s are t h e hawskbi l l , EretmocheZys imbricata, and the l ea the rback , Demochelys coriacea. The hawksbi l l is only known t o occur i n s m a l l numbers exc lus ive ly a t t h e south- e a s t e r n end of t h e Hawaiian Archipelago where a few n e s t i n g s have been recorded i n r ecen t y e a r s on t h e i s l a n d s of Molokai and H a w a i i . s i g h t e d i n o f f s h o r e waters of t h e Hawaiian I s l a n d s , b u t n e s t i n g is n o t known t o t ake p l ace . oliuacea, have a l s o been documented i n Hawaiian waters, b u t on ly as rare v i s i t o r s . A l l of t h e known l i t e r a t u r e r e l a t i n g t o t h e s e f o u r s p e c i e s i n t h e Hawaiian I s l a n d s as of September 1979 has been inc luded i n the .b ib l iog raphy of t h i s synops is .

The l ea the rback is r e g u l a r l y

The loggerhead, Caretta caret ta , and t h e o l i v e r i d l e y , LepidocheZys

This synops is w a s o r i g i n a l l y prepared by t h e au tho r under c o n t r a c t No. 79-ABA-02422 and c o n s t i t u t e d Working Paper No. 13 of t h e J o i n t South P a c i f i c Commission/National Marine F i s h e r i e s Se rv ice Workshop on Marine T u r t l e s i n t h e Trop ica l P a c i f i c I s l a n d s (11-14 December 1979, Noumea, New Caledonia) ,

The a u t h o r ' s r e sea rch of t h e Hawaiian green t u r t l e has been funded dur ing previous yea r s by t h e Un ive r s i ty of H a w a i i Sea Grant Col lege Program, t h e S t a t e of H a w a i i Of f i ce of t h e Marine A f f a i r s Coordinator , t h e Nat iona l Geographic Socie ty , t h e U.S. F i sh and W i l d l i f e Se rv ice (USFWS), and t h e New York Zoologica l Society. The au tho r is c u r r e n t l y a f f i l i a t e d w i t h t h e Southwest F i s h e r i e s Center Honolulu Laboratory under an Intergovernmental Personnel A c t c o n t r a c t w i t h t h e Un ive r s i ty of H a w a i i , H a w a i i I n s t i t u t e of Marine Biology.

This work is t h e r e s u l t of r e sea rch sponsored i n p a r t by t h e Un ive r s i ty of H a w a i i Sea Grant Col lege Program under I n s t i t u t i o n a l Grant Nos. 04-6-158-44114, 04-7-158-44129, and 04-8-MOI-178 from NOAA Of f i ce of Sea Grant , Department of Commerce. Sea Grant Cooperative Report UNIHI-SEAGRANT CR-81-02,

...

2

I IDENTITY

1.1 Nomenclature and Taxonomy P

The valid name for the green turtle is Chelonia mydas (Linnaeus) 1768, with the mydus complex consisting of a number of genetically isolated populations distributed worldwide in tropical and subtropical latitudes. populations have been assigned subspecific names on the basis of coloration, carapace morphology, and geographical location; however, for the most part detailed taxonomic studies have not been carried out.

Some of these

The Atlantic green turtle, Chelonia mydas mydas (Linnaeus) (type locality-Ascension Island) has a predominantly brown coloration to its dorsal surfaces, a white to light yellow plastron, and a carapace displaying a low pro- file, especially from the middle of the lateral laminae to the periphery. At the present time there is no clear distinction between Chelonia mydas mydas and green turtles of the major Caribbean breeding colony at Tortuguero, Costa Rica. However, Carr (151) has expressed confidence that small morphometric, behavioral, and physiological differences exist and, once demonstrated, the name Chelonia mydas v ir idis (Schneider) will apply.

In the eastern Pacific, Chelonia mydas agassizi Bodourt has been described from the coasts of Central and South America and the Galapagos Islands. This subspecies is characterized by a greenish-olive dorsal coloration with the carapace at times strongly marked with black. agassiz$ carapace is highly arched with indentations over the hind flippers. Chelonia mydas carmhegra Caldwell, the stock of turtles with heavy black pigmentation described from the Gulf of California and Pacific coast o f Baja California, is now considered by some authorities to be synonymous with Chelonia mydus agassizi.

In contrast with m y d u s , the

The Hawaiian Chelonia population has thus far not been given nomenclature recognition. Pearl and Hermes Reef in the Northwestern Hawaiian Islands (MWHI), Carr (150, 151) indicated that carrinegra or agassizi characteristics are present. Similar features have also been recorded by Amerson et a l . (32) and Balazs (57, 62). In contrast, Caldwell (147) and Pritchard (418) could not distinguish the small number of Hawaiian specimens they observed from mydus stock. possible predominance of mydas-like stock in the Hawaiian Islands have also been suggested by Hirth and Carr (267). unspecified data, Oliver and Shaw (379) thought that the Hawaiian population might represent the easternmost outpost of the western Pacific green turtle, Chelonia mydas japonica (Thunberg).

Based on turtles observed at French Frigate Shoals and

The presence and

Based on zoogeographic considerations and

Since 1972 the author has observed approximately 2,000 naturally occurring Hawaiian CheZonia of nearly all sizes, including large samples o f adult males and females at the breeding colony of French Frigate Shoals. A reference collection of color photographs including many tagged animals has been compiled along with these field observations, carapace coloration of adult females is heavy black with infusions of yellowish gold and olive. In most cases, the black component comprises an estimated 80% or more of the carapace area. The head and dorsal surfaces of the flippers are

Findings to date show that the predominant

P

R

A

3

"..

a l s o mainly b l ack w i t h l i g h t seams f r e q u e n t l y o u t l i n i n g t h e s c a l e s on each s i d e of t h e head. Along wi th t h e s e predominantly b l ack tur t les , a small number (<3%) of a d u l t females possess a b a s i c a l l y brown colored carapace wi th p a t t e r n s of yel low, gold, and r edd i sh brown. The head and d o r s a l s u r f a c e s of t h e f l i p p e r s of t h e s e i n d i v i d u a l s are a l s o l i g h t e r co lored .

Within t h e a d u l t females of t h e Hawaiian popula t ion , t h e r e is g e n e r a l l y a h igh ly arched appearance t o t h e carapace. The measurement of t h i s c h a r a c t e r i s - t i c has been undertaken by c a l c u l a t i n g t h e r a t i o of t h e curved and s t r a i g h t carapace widths as an estimate of carapace he igh t . Although t h i s r a t i o would no t be v a l i d f o r comparisons between r a d i c a l l y d i f f e r e n t carapace shapes of t h e same s t r a i g h t width ( i . e . domed vs . a f l a t t o p ) , i t n e v e r t h e l e s s r e p r e s e n t s a s i m p l i f i e d index of some va lue . I n a sample of 93 females from French F r i g a t e Shoals, t h i s r a t i o w a s found t o be 1.28 f 0.05 wi th a range of 1.17-1.44. l a r measurements w i l l be necessary i n o t h e r CheZonia popula t ions i n o r d e r t o determine i f s i g n i f i c a n t d i f f e r e n c e s do indeed e x i s t .

S i m i -

The carapace of t h e a d u l t male i n t h e Hawaiian popula t ion can a l s o con ta in cons ide rab le b l ack pigmentat ion, however, g r e a t e r v a r i a b i l i t y h a s been recorded. Ol ive and ye l lowish gold components c o n s i s t i n g of s e v e r a l d i f f e r e n t p a t t e r n s can a t t i m e s p r e v a i l . f l i p p e r s are similar i n pigmentat ion t o t h e black-phase female. 'The male's carapace seems t o p r e s e n t a lower p r o f i l e t han t h e female, however, f u r t h e r morphometric d a t a need t o be ga thered t o c l a r i f y t h i s apparent s exua l dimorphism t h a t was a l s o a p a r t of Caldwell ' s (147) d e s c r i p t i o n of carrinegra.

I n g e n e r a l , t h e head and d o r s a l s u r f a c e s of t h e

P l a s t r o n c o l a t a t i o n i n t h e a d u l t s of bo th sexes ranges from orange t o ye l lowish orange. margina ls and t h e in f r amarg ina l s , which are c l e a r l y v i s i b l e dur ing pe r iods of land basking and n e s t i n g . a long t h e a n t e r i o r edges of t h e second, t h i r d , and f o u r t h margina ls on bo th s i d e s of t h e carapace. carrhzegra by Caldwell (147) has only been found i n a few of t h e a d u l t Hawaiian CheZonia t hus f a r examined by t h e au thor . b lack pigmented p l a s t r o n of an a d u l t a t French F r i g a t e Shoals appears i n a 1954 ar t ic le by Eagle (204).

This c o l o r a l s o encompasses t h e v e n t r a l s u r f a c e s of t h e

A border of orange o r ye l lowish orange o f t e n ex tends

The b l ack o r gray pigmentat ion i n t h e p l a s t r o n descr ibed f o r

A photograph showing t h e d i s t i n c t l y

Inden ta t ions o r c o n s t r i c t i o n s of t h e marginals over t h e h ind f l i p p e r s have been found i n vary ing degrees i n Hawaiian CheZonia of both sexes . For t h e female, such a mod i f i ca t ion may s e r v e t o lessen t h e i n j u r y t o d b r s a l s u r f a c e s of t h e hind f l i p p e r s t h a t can r e s u l t from repea ted con tac t w i th t h e margina ls dur ing excavat ion of t h e egg chamber.

Immature green t u r t l e s observed throughout t h e Hawaiian Archipelago have been found t o show cons ide rab le v a r i a t i o n i n carapace c o l o r a t i o n and p a t t e r n , bo th between and w i t h i n d i f f e r e n t s i z e c a t e g o r i e s . combinations of yel lowish gold, brown, r edd i sh brown, b l ack , and b l i v e . Like t h e a d u l t s , such c o l o r a t i o n s can only be p rope r ly descr ibed i n conjunct ion wi th a d o s s i e r of c o l o r photographs. Pre l iminary evidence h a s been ga thered ind ica- t i n g t h a t some c o l o r phases , p a r t i c u l a r l y those wi th b l ack pigmentat ion, are more p reva len t i n c e r t a i n r e s i d e n t fo rag ing areas. environmental i n f l u e n c e , r e s u l t i n g p o s s i b l y from food sources o r o t h e r characteristics of t h e h a b i t a t . Changes i n t h e c o l o r a t i o n of bo th t h e carapace and

Co lo ra t ions inc lude

This would sugges t an

4

p l a s t r o n have been recorded i n Hawaiian Chelonia r a i s e d under cap t ive condi t ions . For t h e p l a s t r o n , ex tens ive b lack pigmentation develops i n j u v e n i l e s . b y t h e t i m e they reach 8 cm i n s t r a i g h t carapace l eng th , but fades and d isappears by a s i z e of approximately 20 cm. and i t s adap t ive s i g n i f i c a n c e is p r e s e n t l y unknown. I n a d d i t i o n t o gradual and s u b t l e t ransformat ions occurr ing dur ing normal growth, some co lo r changes recorded i n cap t ive specimens have been both d i s c r e t e and r a d i c a l w i th t h e causa- t i v e f a c t o r suspected t o be of an environmental na tu re .

This has not been repor ted i n o the r Chelonia popula t ions

P

With r e s p e c t t o carapace morphology, the h igh ly arched c h a r a c t e r i s t i c has no t been found t o manifest i t s e l f among immature Hawaiian Chelonia measuring less than approximately 65 cm i n s t r a i g h t carapace length .

The o v e r a l l ques t ion of taxonomic s t a t u s f o r Hawaiian Chelonia is c l e a r l y a complex matter t h a t may prove t o be d i f f i c u l t t o r e so lve s o l e l y on t h e b a s i s of c o l o r a t i o n and carapace morphology. exh ib i t ed wi th in t h e popula t ion could i n f a c t be t h e most v a l i d claim f o r even- t u a l des igna t ion as a s e p a r a t e subspec ies .

The unique land basking behavior

1 . 2 Common Names c

Common names used f o r Chelonia i n t h e Hawaiian Archipelago inc lude green t u r t l e , green sea t u r t l e , and honu (Hawaiian). Amerson e t al. (32) have used t h e name "black t u r t l e . I'

1.3 S ize Categor ies

The s i z e ca t egor i e s used f o r Hawaiian Chelonia are as fol lows: hatchl ing--umbil ical s c a r s t i l l p resen t ; juveni le--posthatchl ing t o 65 cm (25.5 i n . ) s t r a i g h t carapace l eng th ; subadult--65 t o 81 cm (31.5 i n . ) s t r a i g h t carapace l eng th ; adult-->81 cm and reproduct ive ly mature.

These d e f i n i t i o n s are modified from t h e ones o f f e red by Hi r th (265) i n t h a t t h e d i v i s i o n between j u v e n i l e and subadul t ca t egor i e s is placed at 65 cm i n s t e a d of 40 cm. This is j u s t i f i e d on t h e b a s i s t h a t 65 cm is t h e minimum s i z e a t which sexua l dimorphism ( lengthening of t h e male's t a i l ) has been observed t o start t ak ing p lace . n a t u r a l l y occur r ing pos tha tch l ing t u r t l e s <35 cm are extremely rare i n t h e Hawaiian Archipelago. Retent ion of H i r t h ' s (265) 40 c m d i v i s i o n would conse- quent ly l i m i t t h e j u v e n i l e category, f o r p r a c t i c a l purposes, t o a narrow 35-40 cm range.

Furthermore, human con tac t and research o p p o r t u n i t i e s wi th

1.4 Tagging and Measuring

The f i r s t recorded occurrence of CheZonia being "tagged" i n t h e Hawaiian Archipelago involved t h e e a r l y 1900's c u l t u r a l p r a c t i c e among some Japanese r e s i d e n t s of i n s c r i b i n g c h a r a c t e r s on a carapace and r e tu rn ing t h e t u r t l e t o t h e sea (Cobb 176). The f i r s t known t a g s a c t u a l l y a t t ached t o Hawaiian green t u r t l e s occurred i n June 1934 when b ras s t a g s in sc r ibed wi th "U.S.S. Itasca 1934" were placed on t h r e e a d u l t s p r i o r t o release a t sea nea r Laysan I s l a n d (Bayl is 104; Clapp and Wirtz 171). During s h o r t v i s i t s t o Laysan I s l a n d and P e a r l and Hemes Reef i n June 1950, Brock (133) tagged an unknown

x

c

5

number of green t u r t l e s using s m a l l s t a i n l e s s steel plates fas tened t o t h e carapace ( P a c i f i c Ocean F i s h e r i e s I n v e s t i g a t i o n s 387; R. Brock, personal communication; Anonymous 574).

The u s e of s i z e 49 c a t t l e ear t a g s manufactured from Monel' 400 a l l o y t h e Nat iona l Band and Tag Company of Newport, Kentucky,was i n i t i a t e d dur ing by

t h e e a r l y 1960's by the S t a t e of Hawaii Div is ion of F i sh and Game (HDFG) (Walker 532, 533; Woodside 554). This type of t a g had a l r eady been s u c c e s s f u l l y used on Chelonia i n both Sarawak and Costa R i c a . Personnel of t h e USFWS and t h e P a c i f i c Ocean B io log ica l Survey Program (Smithsonian I n s t i t u t i o n ) subsequent ly a l s o used t h i s t a g on green t u r t l e s , as w e l l as Hawaiian monk seals, Monuchus schuuinslandi, dur ing i n t e r m i t t e n t survey and in spec t ion v i s i t s t o t h e Hawaiian I s l a n d s NBtional W i l d l i f e Refuge. I n 1967 t h e USFWS made t h e s e t ags a v a i l a b l e t o the Koral Kings Diving Club a t t h e U.S. Navy s t a t i o n on Midway f o r use on j u v e n i l e and subadul t t u r t l e s captured during r e c r e a t i o n a l diving. 800 t u r t l e s had been tagged as a r e s u l t of t h e s e combined e f f o r t s . Nearly a l l of t h e taggings were i n t h e NWHI, and t h e ma jo r i ty cons i s t ed of a d u l t s of both sexes captured whi le basking ashore (Amerson 30; Amerson e t aZ. 32; Clapp and Wirtz 1 7 1 ; Clapp and K r i d l e r 172 ; Ely and Clapp 211; K r i d l e r 311-327; Olsen 380- 385; Woodward 556).

By 1972 an approximate t o t a l of

Systematic tagging and monitoring of n e s t i n g green t u r t l e s a t t h e French F r i g a t e Shoals breeding colony w a s i n i t i a t e d by t h e au thor i n June 1973. This r e sea rch program has continued dur ing subsequent yea r s and has been expanded t o inc lude j u v e n i l e s , subadu l t s , and a d u l t s basking ashore and r e s i d i n g i n forag ing areas throughout t h e Hawaiian Archipelago. Between 1973 and September 1976, s i z e 49, as w e l l as t h e smaller s i z e 681, Monel t a g s were used i n t h i s program. Since t h a t t i m e only s i z e 681 Inconel 625 a l l o y t a g s , s p e c i a l l y manufactured by t h e Nat iona l Band and Tag Company, have been used as t h e primary means of i n d i v i d u a l i d e n t i f i c a t i o n . 29 mm. The change t o Inconel w a s made fol lowing a de te rmina t ion by t h e au thor t h a t cons iderable co r ros ion and subsequent t a g l o s s had occurred i n many of the Monel t a g s placed on Hawaiian Chelonia (Balazs 66; Mrosovsky 363). The implica- t i o n s of t h i s cor ros ion wi th r e spec t t o t h e in t roduc t ion of heavy metals (copper and n i c k e l ) i n t o an an imal ' s system are p o t e n t i a l l y far-reaching. S imi l a r cor ros ion has been found i n Monel t a g s used i n a number of o t h e r marine t u r t l e populat ions. No s i g n s of cor ros ion have thus f a r been found i n t h e Inconel t a g s placed on Hawaiian Chelonia.

These t a g s weigh 3.5 g and measure 8 by

The tagging sites used on Hawaiian Chelonia have been t h e t r a i l i n g edges of t h e f r o n t f l i p p e r s , both i n t h e f o l d of f l e s h proximal t o t h e body and between s c a l e s a t a d i s t a l s i te . This lat ter l o c a t i o n is more conducive t o t h e v i s u a l recovery of t a g numbers from basking t u r t l e s without causing d is turbance . When p o s s i b l e and appropr i a t e , as many as t h r e e Inconel t a g s are placed a t t h e d i f f e r e n t tagging s i tes on each t u r t l e .

'Reference t o t r a d e names does no t imply endorsement by t h e Nat iona l Marine F i s h e r i e s Serv ice , N O M .

6

A number of o t h e r i d e n t i f i c a t i o n techniques have been t e s t e d f o r p o s s i b l e use i n conjunct ion wi th t h e b a s i c t a g . This has included t h e product ion of i d e n t i f i a b l e a n t i b o d i e s (Benedict 107, 108; Hendrickson 262), epoxy p a i n t app l i ed t o t h e carapace (Kr id l e r 324; Anonymous 579) , t a t t o o i n g (Balazs 77) , carapace notch ing of j u v e n i l e s , v i n y l s t r i p s a t t ached through t h e carapace , and p l a s t i c t a g s manufactured by va r ious companies. None of t hese techniques have proven t o be very s a t i s f a c t o r y f o r long-term i d e n t i f i c a t i o n .

Numbers formed on t h e carapace of n e s t i n g and basking t u r t l e s wi th a e r o s o l p a i n t (DuPont Luci te ) have been s u c c e s s f u l l y u t i l i z e d f o r short- term i d e n t i f i c a t i o n and monitor ing of d a i l y ac t iv i t ies (Balazs 49, 51).

Under t h e p re sen t r e sea rch program, a t o t a l of 1,102 Chelonia has thus f a r been tagged throughout t h e Hawaiian Archipelago (Table 1 ) . 444 immature t u r t l e s and 140 a d u l t m a l e s , two c a t e g o r i e s t h a t are not normally tagged i n most o t h e r marine t u r t l e popula t ions .

This i nc ludes

The measurements recorded f o r Hawaiian Chelonia have cons i s t ed o f : s t r a i g h t and curved carapace l e n g t h along t h e midl ine ; s t r a i g h t and curved carapace width a t t h e wides t po in t (u sua l ly t h e s i x t h margina l ) , head width; s t r a i g h t p l a s t r o n l e n g t h along t h e midl ine; t a i l l eng th from t h e p o s t e r i o r edge of t h e p l a s t r o n a l o n g t h e midl ine; and body weight. n e s t i n g t u r t l e s have no t been r e g u l a r l y turned over o r o therwise r e s t r a i n e d i n o rde r t o c a r r y ou t tagging and measuring. The suspension of t h i s prev ious p r a c t i c e has lessened t h e o p p o r t u n i t i e s t o record p l a s t r o n and t a i l measurements, body weight and, a t t i m e s , o t h e r measurements. Never the less , t h i s reduct ion of impact on t u r t l e s from resea rch a c t i v i t i e s w a s deemed necessary i n o rde r t o ensure t h e con t inua t ion of normal basking and n e s t i n g behavior ( see Wallace 535).

Since June 1973, basking and

e

c

2. DISTRIBUTION

2 . 1 To ta l Area

Green t u r t l e s are d i s t r i b u t e d a t select l o c a t i o n s throughout t h e 2,450-km long Hawaiian Archipelago. This nea r ly l i n e a r cha in c o n s i s t s of 132 i s l a n d s , islets, and r e e f s extending from l a t . 18'54'N, long. 154'40'W t o la t . 28'15'N, long. 178'20'W i n an i s o l a t e d reg ion of t h e North Central P a c i f i c Ocean (Figure 1). Eight main and inhab i t ed i s l a n d s (Hawaii, Maui, Kahoolawe, Lanai, Molokai, Oahu, Kauai, and Niihau) loca t ed i n t h e sou theas t e rn segment of t h e Hawaiian Archipelago comprise over 99% o r 16,650 km2 of t h e t o t a l l and area. remainder c o n s i s t s of o f f shore islets and t h e s m a l l i s l a n d s extending t o t h e northwest of Kauai and Niihau known as t h e Leeward I s l a n d s o r t h e NWHI. Except for Kure and Midway, t h e i s l a n d s and c e r t a i n ad jacen t waters i n t h i s segment of t h e cha in c o n s t i t u t e t h e Hawaiian I s l a n d s Nat iona l W i l d l i f e Refuge.

The

There are 1,210 km of c o a s t l i n e i n t h e Hawaiian Archipelago, w i th t h e main i s l a n d s account ing f o r 1 ,165 km o r 96% of t h e t o t a l . underwater c o a s t a l she l f where most green t u r t l e s r e s i d e i s gene ra l ly very narrow wi th t h e 20-fathom curve (37 m) o f t e n only a few k i lome te r s from shore . A number of l a r g e banks w i t h l i t t l e o r no a s soc ia t ed emergent land occur i n t h e northwestern segment of t h e Hawaiian Archipelago. Within t h e 100-fathom curve (182 m>

However, t h e ad jacent

t hese submerged areas encompass approximately 16,000 km2

c

L

7

The c l o s e s t i s l a n d a r e a t o the Hawaiian Archipelago is Johnston A t o l l l oca t ed 820 km south of French F r i g a t e Shoals a t l a t . 16'45'N, long. 169'31'W. A green t u r t l e aggregat ion of undetermined composition and s i z e occurs a t Johnston, however, i t s r e l a t i o n s h i p , i f any, t o Hawaiian CheZonia is no t known a t t h e p re sen t t i m e . An i s o l a t e d green t u r t l e aggregat ion a lso occurs a t Wake I s l and ( l a t . 19'18'N, long. 166'36'E), which is 1,900 km southwest of Midway and t h e c l o s e s t i s l a n d area loca ted t o the w e s t of t h e Hawaiian Archipelago. One of t h e j u v e n i l e s tagged a t Midway w a s subsequent ly recovered a t Wake, however, t h e weakened and apparent ly pathologic .a l condi t ion of t h e t u r t l e sugges ts t h a t i t may have pass ive ly d r i f t e d t h e r e wi th p r e v a i l i n g winds and c u r r e n t s (Balazs 69, 81; Anonymous 641). Nevertheless , t h e r e l a t i o n s h i p of t h e Wake aggrega t ion remains t 6 be determined.

The d i s t r i b u t i o n of green t u r t l e s i n t h e Hawaiian Archipelago has been A breeding colony that formerly occurred a t reduced wi th in h i s t o r i c a l times.

Pol ihua Beach on t h e i s l a n d of Lanai no longer exists (Balazs 55; Emory 213; Kahaulel io 281; Tabrah 473). of green t u r t l e s i n t h e i r r e s i d e n t forag ing areas ad jacen t t o the main i s l a n d s (Balazs 46, 48; Hendrickson 262; S t a t e of H a w a i i 459-463). I n the "HI, the l a r g e aggrega t ion of green t u r t l e s t h a t formerly occurred a t Laysan I s l and has how n e a r l y disappeared (Ely and Clapp 211). Reef appears t o be fol lowing a similar p a t t e r n and t h e s i t u a t i o n a t t h i s l o c a t i o n warran ts a t t e n t i o n .

Reductions have a l s o occurred i n t h e d i s t r i b u t i o n

The aggregat ion a t P e a r l and Hermes

2'2 D i f f e r e n t i a l D i s t r i b u t i o n

2.21 Adults

The d i s t r i b u t i o n of a d u l t Ch320n&2 i n t h e Hawaiian Archipelago is determined p r imar i ly by t h e l o c a t i o n s of accep tab le breeding, feeding , and resting h a b i t a t and, of course, t h e ex i s t ence of s u f f i c i e n t numbers of animals t o u t i l i z e such areasI

c

In excess of 90% of a l l breeding by Hawaiian Chelonia occurs a t French F r i g a t e Shoals (Figure 21, a 35-km long crescent-shaped a t o l l s i t u a t e d i n t h e middle of t h e Archipelago (Amerson 30; Balazs 51, 55, 57; Hendrickson 262; H i r t h 265; K r i d l e r 311-3271 Olsen 380-385). Small groups of t u r t l e s and s e p a r a t e l y n e s t i n g i n d i v i d u a l s us ing Laysan and L i s i a n s k i I s l a n d s and P e a r l and Hermes Reef account €or t h e remaining reproduct ive e f f o r t . Only a few n e s t i n g s have ever been recorded a t Kure and Midway (Balazs 76) .

Feeding and r e s t i n g areas where a d u l t Hawaiian Chelonia l ive t h e g r e a t e r p o r t i o n of t h e i r lives dur ing nonbreeding pe r iods are loca ted i n c o a s t a l waters of both t h e main i s l a n d s and t h e NWHI. The p r i n c i p a l food source , marine b e a t h i c a l g a e of several genera, is r e s t r i c t e d t o shal low depths where s u n l i g h t , s u b s t r a t e and n u t r i e n t s are conducive t o p l a n t growth. Feeding p a s t u r e s used by a d u l t s are usua l ly less than 10 m deep, and f r equen t ly no t more than 3 m deep. The underwater sites where a d u l t s r e g u l a r l y retreat f o r pe r iods o f quiesence inc lude c o r a l r eces ses , t h e undersides of ledges , and sand bottom areas (ca l l ed "nests") t h a t are r e l a t i v e l y f r e e of s t r o n g c u r r e n t s and d i s tu rbance from n a t u r a l

8

p r e d a t o r s and man. occur a t depths >20 m, bu t probably no t normally exceeding 50 m, Avai lab le informat ion i n d i c a t e s t h a t t h e r e s t i n g areas are i n proximity t o t h e feeding pas tu re . whi le f l o a t i n g a t t h e s u r f a c e dur ing l i g h t winds and calm seas. basking undoubtedly y i e l d s a thermal advantage due t o s o l a r r a d i a t i o n d i r e c t l y on t h e carapace and t h e warmer l a y e r of water p resen t a t t h e s u r f a c e . I n a d d i t i o n , less energy would be expended by not having t o p e r i o d i c a l l y s w i m t o t h e s u r f a c e f o r r e s p i r a t i o n .

These r e s t i n g areas f o r a d u l t s i n the main i s l a n d s u s u a l l y

Pe r iods of rest nea r t h e f eed ing p a s t u r e are a l s o known t o t a k e p l ace This s u r f a c e

Some of t h e important r e s i d e n t areas i n t h e main i s l a n d s where a d u l t CheZonia f eed and rest are shown i n F igure 3 and inc lude t h e fol lowing: Kau and North Kohala Districts; Maui--Hana D i s t r i c t and P a i a ; Lanai--northem and n o r t h e a s t e r n c o a s t a l areas border ing t h e Kalohi and Auau Channels; Molokai-- southern c o a s t a l areas from Kamalo t o Halena; Oahu--Kailua and Kaneohc Bays, and nor thwes tern c o a s t a l areas from Mokuleia t o Kawailoa Beach; Kauai--Princevi l le , northwestern c o a s t a l areas of N a P a l i , and southern c o a s t a l areas from Kukuiula t o Makahuena Po in t . Addi t iona l i n v e s t i g a t i o n s are needed t o adequately d e l i n e a t e t h e s e and o t h e r c o a s t a l areas.

Hawaii--

In t h e NWHI, r e s i d e n t aggrega t ions of a d u l t s a r e known t o occur a t Necker I s l a n d , French F r i g a t e Shoals , L i s i a n s k i I s l a n d , Pearl and Hermes Reef, and, t o a lesser e x t e n t , Laysan, Midway, and Kure I s l a n d s . Except f o r Midway, r e s t i n g h a b i t a t a t t h e s e l o c a t i o n s a l s o inc ludes s h o r e l i n e areas where land basking r e g u l a r l y t a k e s p l ace . i t is still unknown i f a d u l t s , o r any o t h e r s i z e c a t e g o r i e s , reside on o r i n some way u t i l i z e t h e suhmerged banks wi th no emergent land l o c a t e d i n t h e northwestern segment of t h e Hawaiian Archipelago. Gardner P innac les .

Although a few random s i g h t i n g s have been made,

Data are a l s o l ack ing f o r Nihoa and

While i n t r ans i t dur ing r ep roduc t ive mig ra t ions , a d u l t Hawaiian Chelonia are d i s t r i b u t e d a t unknown l o c a t i o n s i n t h e p e l a g i c environment between t h e i s l a n d s of t h e Hawaiian Archipelago.

2.22 Ha tch l ings , j u v e n i l e s , and subadu l t s

Hatchl ings emerge from t h e i r n e s t s and e n t e r t h e water a t French F r i g a t e Shoals and t h e o t h e r breeding si tes i n t h e MI between mid-July and e a r l y October. l o s t t o almost a l l human con tac t . been found t o occur i n o t h e r Chelonia popula t ions (Hi r th 265). t u r t l e h a t c h l i n g s are thought t o be subsequent ly d i spe r sed i n t h e p e l a g i c environment by c u r r e n t s and vigorous swimming. p l o t t e d f o r t h e t h e o r e t i c a l movement of h a t c h l i n g s l eav ing French F r i g a t e Shoals suggest a predominant wes te r ly d i s p e r s a l (Figure 4). However, CheZonia hatch- l i n g s are s t r o n g swimmers and t h e d i s t i n c t p o s s i b i l i t y e x i s t s t h a t cons ide rab le movement t a k e s p l a c e a t va r i ance wi th p r e v a i l i n g s u r f a c e c u r r e n t s .

Following a r a p i d depa r tu re from t h e ad jacen t waters, they are

Hawaiian green This disappearance is c o n s i s t e n t w i th what has

Surface d r i f t t r a j e c t o r i e s

The 28 March 1779 l o g of t h e Resolution r eco rds t h e p e l a g i c s i g h t i n g of a pos tha tch l ing t u r t l e of unknown s p e c i e s t o t h e southwest of t h e H a w a i i a n Archi- pelago a t l a t . 20°15'N, long. 179"20'E (Beaglehole 105).

c

4

c

9

The d i s t r i b u t i o n of j u v e n i l e s up t o 35 c m i n t h e Hawaiian Archipelago is a l s o unknown, due aga in t o a n e a r l y complete absence of human con tac t . T u r t l e s of t h i s s i z e are almost environment where t h e chances of s ee ing them are g r e a t l y reduced. p reda t ion by c e r t a i n p e l a g i c f i s h e s such as t h e oceanic w h i t e t i p sha rk , Carcharhinus longimanus, could be expected t o t ake p l ace . be ing made t o recover j u v e n i l e s of t h i s " l o s t " s i z e from t h e stomachs of p o t e n t i a l p r e d a t o r s (Balazs 70, 79; Hendricks 261; Anonymous 658).

c e r t a i n l y s t i l l r e s i d i n g i n t h e p e l a g i c Never the less ,

E f f o r t s are t h e r e f o r e

J u v e n i l e s l a r g e r than 35 cm as w e l l as subadu l t s can be found feeding and r e s t i n g i n c o a s t a l a r e a s throughout t h e Hawaiian Archipelago. These t u r t l e s f r e q u e n t l y r e s i d e i n t h e s a m e gene ra l area as t h e a d u l t s . There is , however, t h e tendency among j u v e n i l e s and subadu l t s t o u t i l i z e r e s t i n g h a b i t a t l o c a t e d a t a shal lower depth. p a s t u r e s due t o t h e ve ry sha l low depths involved.

I n a d d i t i o n , on ly j u v e n i l e s are a b l e t o use some feeding

3. ECOLOGY AND LIFE HISTORY

3 .1 Reproduction

3.11 Sexua l i ty

Adult males of t h e Hawaiian Chelonia popula t ion have a 35 t o 45 cm long

The a d u l t female ' s p r e h e n s i l e t a i l t h a t ex tends beyond t h e h ind f l i p p e r s wh i l e swimming and is l a r g e r i n o v e r a l l diameter t han t h e t a i l of t h e a d u l t female. t a i l ranges from 20 t o 25 cm i n l e n g t h and only ex tends t o about t h e middle of t h e h ind f l i p p e r s . f l i p p e r is a l s o longer i n t h e a d u l t male. However, i n bo th sexes t h i s s t r u c t u r e can be cons iderably worn down, probably due mostly t o ab ras ion wi th hard sub- strate encountered whi le feeding and r e s t i n g underwater. Furthermore, t h e n a i l s on t h e female r e c e i v e some wear dur ing n e s t i n g , whi le n a i l s on t h e male may be abraded dur ing copula t ion . A heav i ly k e r a t i n i z e d t i p is p resen t at t h e end of t h e male's t a i l , bu t t h i s may be miss ing i n many i n d i v i d u a l s due t o i n j u r y from o t h e r t u r t l e s o r p reda to r s .

The s i n g l e n a i l p re sen t on t h e foremargin of each f r o n t

It is p r e s e n t l y only p o s s i b l e t o determine sex on t h e b a s i s of e x t e r n a l c h a r a c t e r i s t i c s i n i n d i v i d u a l s l a r g e r t han 65 cm. Even then, cau t ion must be exe rc i sed i n t h a t l engthening of t h e t a i l i n some males does not start u n t i l a g r e a t e r s i z e is reached.

3.12 Matur i ty

The smallest female thus f a r found n e s t i n g a t French F r i g a t e Shoals measured 81 c m i n s t r a i g h t carapace l e n g t h , w i th 92 c m being t h e o v e r a l l m e a n (Table 2). t ive purposes. Data by K r i d l e r (311-327) and Olsen (380-385) show t h a t t h e mean weight of a d u l t females is 110 kg (range 68-148 kg N=69). (265), n e s t i n g aggrega t ions of Chelonia may c o n s i s t of some females t h a t grew t o a l a r g e s i z e a f t e r reaching ma tu r i ty , and some t h a t d i d not mature u n t i l reach- i n g a l a r g e s i z e . It is p r e s e n t l y no t p o s s i b l e t o i d e n t i f y newly matured i n d i v i d u a l s ( r e c r u i t s ) i n

Curved carapace measurements are p resen ted i n Table 3 f o r compara-

As noted by H i r t h

Such a phenomenon could a l s o be expected f o r a d u l t males.

t h e French F r i g a t e Shoals breeding assemblage, however, continued tagging and c o l l e c t i o n of s ize-frequency d a t a may even tua l ly provide t h e b a s i s f o r t h i s determinat ion.

The age a t which green t u r t l e s mature under n a t u r a l cond i t ions has rece ived very l i t t l e research a t t e n t i o n i n most popula t ions , a l though estimates based p r i n c i p a l l y on cap t ive growth rates have appeared i n t h e l i t e r a t u r e (Hir th 265). j u v e n i l e s have demonstrated s i g n i f i c a n t d i f f e r e n c e s i n t h e rates of growth, hence age a t ma tu r i ty , between r e s i d e n t areas throughout t h e Hawaiian Archipelago (Balms 81) . The p ro jec t ed l eng ths of t i m e f o r 35 c m j u v e n i l e s t o reach t h e mean s i z e of a d u l t females range from 11 t o 59 y r (Table 4 ) . T u r t l e s t h a t r e s i d e i n t h e NWHI seem t o r e q u i r e t h e longes t per iods of t i m e t o mature.

For Hawaiian Chelonia, t a g and r ecap tu re experiments w i th wi ld c

3.13 Mating

Court.ship and copula t ion t ake p l ace i n t h e shal low waters of French F r i g a t e Shoals , u s u a l l y wl th in 2 km of t h e 11 small i s l a n d s present a t t h i s l o c a t i o n . Some observa t ions have suggested t h a t t h e g r e a t e s t i n f l u x of males t o t h e area occurs p r i o r t o the a r r i v a l of most females (Balazs 4 9 ; Olsen 382). This i s complicated, however, by t h e presence of a d u l t males t h a t are known t o r e s i d e f o r extended pe r iods a t French F r i g a t e Shoals.

Copulat ion has been observed t ak ing p l ace a t t h e s u r f a c e , on t h e bottom, and a t va r ious depths w i t h i n t h e water column. Seve ra l males are almost always i n a t tendance wi th a copu la t ing p a i r , c i r c l i n g , smel l ing and b i t i n g t h e t a i l and f l i p p e r s of t h e s u c c e s s f u l l y mounted m a l e . The i n j u r i e s i n f l i c t e d by t h i s behavior are f r equen t ly ex tens ive wi th scales and f l e s h be ing t o r n away and t h e under ly ing limb bones exposed. dur ing previous seasons can o f t e n be recognized by t h e missing areas along t h e t r a i l i n g edges of t h e i r f r o n t f l i p p e r s and t h e presence of t h i c k , nonpigmented scar t i s s u e . neophyte males a t t h e breeding colony.

Males t h a t have undergone these a s s a u l t s

These scars may even tua l ly be u s e f u l i n t h e i d e n t i f i c a t i o n of

The male main ta ins h i s p o s i t i o n dur ing copula t ion by grasp ing t h e margins of t h e female 's carapace wi th t h e d i s t a l areas of h i s f o u r f l i p p e r s . The n a i l on t h e f r o n t as w e l l as t h e hind f l i p p e r s s e e m t o provide some assis- tance t o t h i s e f f o r t . The muscular p r e h e n s i l e t a i l a l s o serves as a hold ing device by extending under t h e female t o a p o s i t i o n where t h e pen i s can be i n s e r t e d i n t o t h e c loaca . of a l l movement s i n c e he r four f l i p p e r s are not r e s t r a i n e d . Copulating p a i r s s t i l l locked toge the r have been observed both d i u r n a l l y and noc tu rna l ly i n water only s e v e r a l cen t ime te r s deep ad jacen t t o t h e i s l a n d s a t French F r i g a t e Shoals. This beaching maneuver is presumably c a r r i e d ou t by t h e female i n an at tempt t o de tach t h e male.

During copula t ion t h e female i s i n complete c o n t r o l

Most copula t ion a t French F r i g a t e Shoals occurs during t h e e a r l y p o r t i o n of t h e breeding season , between mid-April and e a r l y June. i d e n t i f i c a t i o n numbers pa in ted on t h e carapace of n e s t i n g t u r t l e s has i n d i c a t e d t h a t females w i l l not copula te a f t e r l a y i n g t h e i r f i r s t c l u t c h of eggs. Copulations recorded l a t e r i n t h e breeding season appear t o involve females t h a t have only r e c e n t l y a r r i v e d a t French F r i g a t e Shoals.

The use of

c

11

Copulat ing t u r t l e s have been seen a t L i s i a n s k i I s l and dur ing A p r i l 1976 (C. F i scus , personal communication) and a t P e a r l and Hemes Reef dur ing March 1961, l a t e May 1967 (Amerson e t aZ. 32), and A p r i l 1976 (DeLong e t aZ. 195). No records e x i s t f o r Laysan, Kure, and Midway.

Two a d u l t males locked toge the r i n a copula tory p o s i t i o n have been observed on s e v e r a l occasions a t French F r i g a t e Shoals , both dur ing t h e breeding season and i n the month of December. p o s i t i o n was maintained f o r a t least 20 min. S imi l a r behavior among males has also been documented a t Necker, a small rock i s l a n d wi th no sand beaches loca ted 155 km east of French F r i g a t e Shoals (Balazs 72).

During one of t h e s e encounters , t h e

3.14 F e r t i l i z a t i o n

It is unce r t a in i f t h e eggs of Chelonia (and o t h e r marine t u r t l e s ) are f e r t i l i z e d as a r e s u l t of copula t ion t h a t occurs wi th in t h e same breeding season, o r from spermatozoa s t o r e d i n t h e reproduct ive t rac t r e s u l t i n g from copu la t ion dur ing a previous season. I f s t o r a g e is a p r e r e q u i s i t e f o r t h e f e r t i l i z a t i o n , and copula t ion only occurs nea r t h e n e s t i n g beaches, then t h e neophyte female would have t o make her f i r s t v i s i t t o French F r i g a t e Shoals s o l e l y f o r mating purposes , Such a t r i p could, however, a l s o involve n e s t i n g excurs ions a shore , bu t wi th only i n f e r t i l e eggs o r no eggs a t a l l being l a i d . occurrences have been documented a t French F r i g a t e Shoals , o t h e r f a c t o r s could very w e l l be r e spons ib l e .

Although both of t h e s e

I n an e f f o r t t o determine i f f e r t i l i t y decreases at French F r i g a t e Shoals as t h e breeding season p rogres ses , 40 precounted egg c l u t c h e s l a i d over a 72-day per iod (June-August 1974) were excavated and examined fo l lowing t h e n a t u r a l emergence of h a t c h l i n g s . Regression a n a l y s i s of t h e s e d a t a showed a tendency toward decreased f e r t i l i t y w i t h t i m e , however, t h i s w a s no t found t o be s i g n i f i c a n t (Pro. 08).

3.15 Nest ing and h a b i t a t c h a r a c t e r s i t i c s

c

Nest ing t akes p l a c e a t French F r i g a t e Shoals on ly on t h e i s l a n d s of East, Whale-Skate, T r ig , Tern, Gin, and L i t t l e Gin (F igure 2 ) . . Of t h e females p re sen t f o r each breeding season, approximately 55% n e s t on E a s t and 35% nest on Whale-Skate. East I s l a n d has consequently been t h e s i t e of i n t e n s i v e r e sea rch dur ing each breeding season s i n c e 1973 w i t h surveys being made a t r e g u l a r i n t e r - vals on t h e o t h e r i s l a n d s . i t s peak dur ing l a t e June, and d e c l i n e s t o a very low level by e a r l y August. Some sporad ic n e s t i n g may occur u n t i l mid-September. depos i t i on t a k e s p l ace a t n i g h t , o r a t least by s u n r i s e , females have been recorded ashore excavat ing a n e s t s i t e as e a r l y as 2 h p r i o r t o sunse t .

Nest ing commences dur ing t h e middle of May, reaches

Although a c t u a l egg

East and Whale-Skate are the two l a r g e s t n a t u r a l l y occur r ing i s l a n d s a t French F r i g a t e Shoals , c o n s i s t i n g of 4.0 and 6.8 ha, r e s p e c t i v e l y , wi th e l e v a t i o n s no t exceeding 2 m. Both i s l a n d s have a ca lcareous s u b s t r a t e composed of f i n e t o coa r se fragments of c o r a l s , c o r a l l i n e a l g a e , molluscs , and ba rnac le s . The i n t e r i o r s of both i s l a n d s have low v e g e t a t i o n (Tribulus, Chenopodium, Portulaca, Boerhauia) and are r i c h w i t h humus o r i g i n a t i n g from n e s t i n g seab i rd

1 2

co lonies . From 1944 t o 1952 E a s t I s l and w a s t h e s i t e of a 26-man U.S. Coast Guard naviga t ion (LORAN) s t a t i o n . Although some clean-up a c t i v i t i e s took p l ace i n 1965 and 1973, remains of t h i s abandoned f a c i l i t y are s t i l l prominent on t h e i s l a n d . Tern I s l a n d , l oca t ed 11 km t o t h e northwest of East I s l a n d , is a r econs t ruc t ed i s l a n d t h a t w a s en la rged by dredging and l a n d f i l l from 4.5 t o 23 ha f o r m i l i t a r y purposes i n 1942 (Amerson 30; Berger 125; Olsen 383). From 1952 t o 1979, a U.S. Coast Guard LORAN s t a t i o n w a s a l s o p re sen t on T e r n I s l and .

Mean monthly sea-surface temperatures recorded a t French F r i g a t e Shoals between 1974 and 1976 were found t o range from 23.8' t o 28.3OCY with t h e warmest water gene ra l ly occur r ing from June through October (F igure 5 ) . This is similar t o sea-sur face temperatures found i n t h e main Hawaiian I s l a n d s . The mean annual r a i n f a l l a t French F r i g a t e Shoals i s 115 cm wi th t h e d r i e s t months be ing A p r i l through November. Addi t iona l c l i m a t i c d a t a f o r t h e area are presented by Amerson (30)

Green t u r t l e s n e s t i n g a t East I s l a n d have been recorded l a y i n g as many as six egg c l u t c h e s w i t h i n a season, however, t h e mean number is only 1.8 (Table 5). 10% make n e s t i n g a t t empt s on s e v e r a l consecut ive n i g h t s , bu t do n o t l a y eggs and o f t e n are no t seen aga in . Only a few t u r t l e s (-3%) have been recorded each season changing i s l a n d s once n e s t i n g has s t a r t e d . This f a c t o r a lone would, t h e r e f o r e , n o t f u l l y e x p l a i n t h e absence of o v i p o s i t i o n by some ind iv idua l s . It is p l a u s i b l e t h a t some of t hese t u r t l e s are r e l e a s i n g t h e i r eggs i n t h e water due t o a n e s t i n g dysfunct ion , whi le o t h e r s could b e m o r t a l i t i e s due t o p reda t ion by t i g e r sha rks , Caleocerdo cuuier.

Approximately 40% of t h e t u r t l e s l a y only once i n a season. Almost

The l eng th of t i m e between o v i p o s i t i o n i n t u r t l e s t h a t l a y more than once i n a season ranges from 11 t o 18 days, wi th t h e mean being 13 days (Table 6 ) . During t h i s i n t e r n e s t i n g i n t e r v a l , t u r t l e s i d e n t i f i e d by pa in t ed numbers are r e g u l a r l y seen basking a t E a s t I s l a n d and swimming i n t h e ad jacen t waters. The unders ides of r e e f s t h a t extend both t o t h e sou theas t and northwest of t h e i s l a n d are a l s o known t o be u t i l i z e d dur ing t h e i n t e r n e s t i n g per iod. a pre l iminary experiment conducted dur ing June 1979, t h e m a x i m u m d i v i n g depth recorded w i t h small depth gauges a t t ached t o two females a t T r i g and Whale-Skate I s l a n d s was 1 2 . 8 m. This sugges t s t h a t movement t o deeper waters o u t s i d e the lagoon may n o t b e t ak ing p l ace dur ing t h e breeding season.

In

The degree of s i t e f i d e l i t y d isp layed by green t u r t l e s n e s t i n g at East I s l and w i t h i n each season has been examined by d iv id ing t h e s h o r e l i n e i n t o seventeen 50-m long r e fe rence areas. Observat ions made over t h e 1973, 1974, and 1975 seasons revea led t h a t 32% of t h e egg c lu t ches l a i d dur ing r e n e s t i n g s w e r e i n t h e same 50-m area as earlier nes t ings . Approximately 60% w e r e i n t he same o r an ad jacen t 50-m area, whi le 83% of t h e t u r t l e s emerged on t h e same s i d e of t h e i s l a n d as t h e earlier n e s t i n g s . Some i n d i v i d u a l s showed cons iderable s i te f i x i t y , as demonstrated by two t u r t l e s i n 1974 t h a t each nes t ed f o u r t i m e s i n t h e same 50-m area, and a t u r t l e t h a t nes t ed s i x t i m e s , a l t e r n a t i n g r e g u l a r l y between t h e same two areas (1 and 6 ) . Also dur ing 1974, r e n e s t i n g s by f i v e t u r t l e s involved egg c l u t c h e s t h a t w e r e l a i d only 1.5 t o 3.5 m from an earlier c lu tch .

P

P

a

1 3

c-

The l o c a t i o n s of 220 egg c lu t ches l a i d a t East I s l a n d during t h e 1974 season are presented i n Figure 6. areas 2-3 and 14-16 con ta in several low concre te foundat ions from t h e abandoned Coast Guard f a c i l i t y . Although s u i t a b l e s u b s t r a t e f o r n e s t i n g is a l s o p r e s e n t , very few t u r t l e s have ever been recorded e n t e r i n g t h i s reg ion of t h e i s l a n d .

The u n u t i l i z e d p o r t i o n s of t h e i n t e r i o r of

Usual ly less than h a l f of t h e t u r t l e s t h a t emerge f o r n e s t i n g on any one n i g h t s u c c e s s f u l l y l a y eggs (Figure 7 ) . The remaining t u r t l e s cont inue t o emerge on subsequent n i g h t s u n t i l o v i p o s i t i o n i s achieved o r , as previous ly descr ibed , t h e t u r t l e is no longer p re sen t . Many of t h e s e n e s t i n g a t t empt s involve t h e n e a r l y complete excavat ion of an egg chamber be fo re abandonment takes p l a c e and another s i t e is s e l e c t e d on t h e i s l a n d . mentary body p i t s are dug be fo re a s i te is abandoned. c o n t r i b u t e t o t h e incomplete excavat ion of a n e s t i nc lude i n j u r i e s o r amputations of a t u r t l e ' s h ind f l i p p e r s , i n s u f f i c i e n t moisture i n t h e s u b s t r a t e due t o low r a i n f a l l , and con tac t w i t h bu r i ed antenna w i r e and o t h e r d e b r i s p re sen t on East I s l and . because t h e complement of eggs has n o t y e t developed t o a s t a g e f o r o v i p o s i t i o n t o occur . emergences iti which t h e t r a c k s form a p a r a b o l i c curve (Carr e t aZ. 153; H i r t h 265) has only r a r e l y been recorded i n Hawaiian Chelonia. The "sand-smelling" behavior exh ib i t ed by newly emerged females i n some green t u r t l e popula t ions has never been observed a t French F r i g a t e Shoals.

I n o t h e r cases, only rudi - Three f a c t o r s t h a t

It is a l s o p o s s i b l e t h a t some of t h e s e n e s t i n g a t tempts are unsuccessfu l

The tendency of some green t u r t l e s t o make s h o r t , nonegg-laying

h r i n g t h e n e s t i n g process , Hawaiian ChgZonia are least s u s c e p t i b l e t o d i s tu rbance immediately fo l lowing o v i p o s i t i o n when n e s t covering has j u s t s t a r t e d t o t ake p lace . Carapace measurements and tagging are t h e r e f o r e usua l ly c a r r i e d out dur ing t h i s per iod .

A sample of 50 egg c l u t c h e s counted dur ing o v i p o s i t i o n over a 75-day per iod (June-August 1974) a t East I s l a n d revea led a mean of 104 eggs p e r c l u t c h w i t h a range of 38 t o 145 eggs. Mul t ip l e r eg res s ion a n a l y s i s of t h e s e d a t a was conducted t o determine i f s i g n i f i c a n t r e l a t i o n s h i p s exis t between t h e number of eggs i n a c l u t c h and t h e independent v a r i a b l e s of : t i m e of o v i p o s i t i o n w i t h i n t h e season ( X I ) , r a t i o of t h e curved and s t r a i g h t carapace widths of t h e female (x2), and s t r a i g h t carapace l eng th of t h e female (x3). Larger females were found t o l a y s i g n i f i c a n t l y (PC0.05) more eggs p e r c lu t ch . dency f o r fewer eggs p e r c l u t c h t o b e l a i d as t h e season progressed , t h i s w a s no t s i g n i f i c a n t (P=O.lO). The r e l a t i o n s h i p between t h e curved-s t ra ight width r a t i o and number of eggs p e r c l u t c h w a s a l s o no t found t o b e s i g n i f i c a n t (p10.30). This sugges t s t h a t body th i ckness a l o n e is not an adap ta t ion f o r t h e s t o r a g e of g r e a t e r numbers of eggs. eggs i n a c l u t c h (y) i s y = -268.704 4- (-0.271)xl + 93 .768~2 + 2 . 8 1 9 ~ 3 .

Although t h e r e w a s a ten-

The r e s u l t i n g formula f o r p r e d i c t i n g t h e number of

The mean incubat ion pe r iod (days t o h a t c h l i n g emergence a t t h e su r face ) of 38 n e s t s l a i d a t East I s l a n d from June t o August 1974 w a s found t o be 64.5 days w i t h a range of 54 t o 88 days. Mul t ip l e r eg res s ion a n a l y s i s of t h e s e d a t a found no s i g n i f i c a n t r e l a t i o n s h i p s between incubat ion pe r iod and t i m e of ovi- p o s i t i o n w i t h i n t h e season, coarseness o f t h e n e s t s u b s t r a t e , o r depth of t h e egg chamber. c a n t r e l a t i o n s h i p coarseness of s u b s t r a t e , w i t h sha l lower chambers .being excavated i n c o a r s e r

The mean egg chamber depth w a s 60 c m (range 48-74 c m ) . A s i g n i f i - (P<0.05) w a s found between depth of t h e egg chamber and

14

s u b s t r a t e . s i z e of t h e n e s t i n g female.

N o s i g n i f i c a n t r e l a t i o n s h i p w a s found between egg chamber depth and

The mean width and l eng th of t h e s u r f a c e area excavated by 19 t u r t l e s t h a t s u c c e s s f u l l y nes t ed on E a s t I s l a n d w a s 2.0 m ( range 1.7-2.7 m) and 3.8 m (range 2.3-5.0 m), r e s p e c t i v e l y . No s i g n i f i c a n t r e l a t i o n s h i p was found between t h e s e measurements and t h e s i z e of t h e n e s t i n g female.

T u r t l e s on both East and Whale-Skate I s l a n d s r e g u l a r l y come i n t o con tac t w i th 10 s p e c i e s of n e s t i n g s e a b i r d s , t h e most abundant of which are sooty t e r n s , Sterna fuscata, and wedge-tailed shearwaters , Puffinus pacif icus. The crushing of eggs and young ch icks by n e s t i n g t u r t l e s consequently t a k e s p l a c e t o varying degrees depending on t h e t i m e of t he season. I n a d d i t i o n , a d u l t shearwaters occas iona l ly become bur i ed i n t h e i r underground burrows as a r e s u l t of t u r t l e a c t i v i t y . are a l s o encountered a t t i m e s by n e s t i n g t u r t l e s , bu t wi th t h e except ion of mothers w i t h pups, u s u a l l y l i t t l e i f any i n t e r a c t i o n t akes p lace .

Monk seals hauled o u t a long t h e s h o r e l i n e and i n t h e v e g e t a t i o n zone

Table 7 p r e s e n t s t h e r e s u l t s of 40 precounted egg c lu t ches l a i d a t East I s l a n d from June t o August 1974 t h a t were excavated and examined fol lowing t h e n a t u r a l emergence of ha t ch l ings . Mul t ip l e r eg res s ion ana lyses of t h e s e d a t a w e r e conducted t o determine i f s i g n i f i c a n t r e l a t i o n s h i p s exist between cent of eggs ha tched , 2) percent of h a t c h l i n g emergence, 3) pe rcen t of dead ha tch l ings , 4 ) percen t of p a r t i a l l y developed bu t dead embryos, 5) percent of eggs wi th no apparent development, and the independent v a r i a b l e s of 1 ) t i m e of o v i p o s i t i o n w i t h i n t h e season, 2) coarseness of t h e n e s t s u b s t r a t e , 3) depth of n e s t , and 4 ) s t r a i g h t carapace l eng th of t h e female. The only s i g n i f i c a n t r e l a t i o n s h i p (P<0.05) found w a s t h a t t h e percent of h a t c h l i n g s emerging a t t h e s u r f a c e decreases i n egg c lu t ches t h a t are l a i d as t h e season progresses .

1 ) per-

c

3.16 Reproductive cyc le s c

Reproductive cyc le s , as measured by remigra t ion i n t e r v a l s , have been documented i n 2 1 n e s t i n g females a t E a s t I s l a n d s i n c e June 1973 (Table 8). Fourteen of t h e s e t u r t l e s (66.7%) d isp layed a 2-yr cyc le , s i x (28.6%) a 3-yr cyc le , and one t u r t l e (4 .7%) w a s no t seen aga in u n t i l 6 y r a f t e r be ing tagged. Thus f a r , no n e s t i n g t u r t l e s have been recovered a t French F r i g a t e Shoals a f t e r on ly a 1 y r absence. The p resen t predominance i n record ings of 2-yr n e s t i n g c y c l e s is due t o t h e s i g n i f i c a n t i n c r e a s e i n r ecove r i e s made dur ing t h e 1979 breeding season of t u r t l e s tagged 2 y r earlier i n t h e 1977 season (Table 8). Because 1977 w a s t h e f i r s t season i n which t h e more durable Inconel t a g s w e r e used a t French F r i g a t e Shoals , cont inued monitor ing and tagging w i l l be neces- s a r y dur ing f u t u r e breeding seasons ( p a r t i c u l a r l y 1980 and 1981) i n o r d e r t o accu ra t e ly determine t h e most common n e s t i n g cyc le . per iods appa ren t ly r equ i r ed f o r Hawaiian Chelonia t o reach ma tu r i ty a t c e r t a i n r e s i d e n t fo rag ing areas ( s e c t i o n 3.12) , i t is reasonable t o assume t h a t t h e same c o n t r o l l i n g e c o l o g i c a l f a c t o r s would a l s o cause p r o t r a c t e d pe r iods f o r repro- duc t ive r ead iness t o be reached between remigra t ions . The occurrence of some n e s t i n g c y c l e s f a r g r e a t e r than 3 o r even 6 y r should t h e r e f o r e be considered as a d i s t i n c t p o s s i b i l i t y . Furthermore, t h e low recovery rates of tagged a d u l t females i n both t h e Hawaiian and Caribbean (Carr e t a l . 153) Chelonia popula t ions

I n v i e w of t h e long t i m e

c

15

... ._ ..

u

suggest t h e p o s s i b i l i t y t h a t many t u r t l e s may only be involved i n a s i n g l e breeding season dur ing t h e course of t h e i r e n t i r e l i f e t i m e .

The reproduct ive cyc le s of males a t French F r i g a t e Shoals have been documented i n 16 cases. Nine (56.2%) of t h e s e represented a l -yr cyc le , f ive (31.3%) a 2-yr cyc le , and two (12.5%) a 3-yr cycle . r ecove r i e s i n d i c a t e s t h a t t h e l -y r cyc le involves remigra t ions r a t h e r t han simply a res idency a t French F r i g a t e Shoals. The apparent predominance of a l -yr cyc le i n males may be due t o a lower energy requirement f o r reaching reproduct ive r ead iness between remigra t ions i n comparison t o females. Although t h e occurrence of 2- and 3-yr cyc le s d i f f i c u l t i e s are encountered i n r e l i a b l y determining male reproduct ive c y c l e s t h a t are longer than 1 y r . t h a t a l l males w i l l come ashore t o bask during a given season and, t h e r e f o r e , be a v a i l a b l e f o r tagging and t a g recovery.

The n a t u r e of t h e s e t a g

i s be l i eved t o be v a l i d , i t should be noted t h a t g r e a t e r

This i s due t o t h e f a c t t h a t t h e r e i s no assurance

The modulation of r ep roduc t ive c y c l e s (see Carr e t aZ. 153) h a s thus f a r been recorded i n two t u r t l e s a t French F r i g a t e Shoals. change from a 3- t o a 2-yr cyc le i n a female, and a change from a 2- to a l-yr cyc le i n a male. are p r e s e n t l y unknown. p e r i o d i c changes i n p r o d u c t i v i t y o r o t h e r e c o l o g i c a l cond i t ions of t h e t u r t l e ' s r e s i d e n t fo rag ing area.

This involved a phase

The e x t e n t and s i g n i f i c a n c e of such s h i f t s i n Hawaiian Chelonia Modulation of reproduct ive cyc le s may b e a r e f l e c t i o n of

3.17 Eggs

The mean weight of eggs l a i d a t East I s l and by a 96 c m s t r a i g h t carapace l eng th female w a s 50 g (range 45-54 g , N-99). t hese eggs w a s 44 mm w i t h a range of 43 t o 46 mm. observed dur ing o v i p o s i t i o n a t East I s l a n d were found t o con ta in a few of t h e abnormally small o r nonspher ica l eggs t h a t have been r epor t ed i n o t h e r Chelonia popula t ions . f u l l y excavated egg chamber ranges from approximately 20 t o 30 min. cons iderably longer than t h e 10 min r epor t ed by H i r t h (265) f o r o t h e r Chelonia popula t ions .

The mean diameter of Three of 50 egg c lu t ches

The t i m e r equ i r ed f o r an egg c l u t c h t o b e depos i ted i n t o a success- This is

Preda t ion on eggs does not normally t a k e p l a c e a t French F r i g a t e Although two s p e c i e s of ghost c r abs (Ocypode eeratophthuZmus and 0. Shoals.

Zaevis) are p resen t i n r e l a t i v e l y small numbers, n e i t h e r of t h e s e c rus t aceans have been found burrowing i n t o a n e s t . Although i t is a rare occurrence a t French F r i g a t e Shoals , n e s t i n g t u r t l e s w i l l a t t i m e s i n a d v e r t e n t l y d i g int.0 an e x i s t i n g c l u t c h of eggs. Broken and exposed eggs from n e s t s t h a t have been d i s tu rbed i n t h i s manner have been observed be ing e a t e n by 0. Zaevis, as w e l l as migratory sho re b i r d s (ruddy tu rns tones , Arenaria interpres and golden p love r s , PZuviaZis dominiea). On a s i n g l e occasion, a wedge-tailed shearwater w a s recorded as having acc iden t ly burrowed into a c l u t c h of eggs a t E a s t I s l and . In a few i n s t a n c e s , egg c lu t ches have been l a i d too c l o s e t o t h e s h o r e l i n e , thereby being uncovered and washed away by subsequent wave a c t i o n . Also a few t u r t l e s have been observed breaking l a r g e numbers of t h e i r own eggs as a r e s u l t of harsh con tac t by a p a r t i a l l y amputated hind f l i p p e r during t h e n e s t covering process .

Prom 1973 t o 1979, an e s t ima ted 500 kg of unhatched green t u r t l e eggs and dead h a t c h l i n g s have remained bur ied i n t h e ground each season a t French

16 c

F r i g a t e Shoals and undergone decomposition. Although probably small i n comparison t o d e p o s i t s from n e s t i n g s e a b i r d s , t h i s chelonian organic matter neve r the l e s s r ep resen t s a va luab le con t r ibu t ion t o t h e s o i l f e r t i l i t y of t h e i s l a n d s .

3.2 Hatchl ing Phase

Newly emerged h a t c h l i n g s a t French F r i g a t e Shoals measure 53 mm i n s t r a i g h t carapace l eng th (range 48-59 m, N=556) and weigh 31 g (range 25-35 mm, N=120). The d o r s a l s u r f a c e s of t h e ha t ch l ings are b lack wi th whi te edges 2- t o 3-nun wide on t h e f l i p p e r s and a 1-mm wide whi te edge a long t h e per iphery of t h e marginal laminae. The v e n t r a l s u r f a c e s of t h e ha t ch l ings are whi t e w i t h areas of b lack loca ted on t h e f l i p p e r s ( see Balazs 55, 62 f o r c o l o r photographs).

P r i o r t o emergence from t h e n e s t , Hawaiian green t u r t l e h a t c h l i n g s have been found t o e x h i b i t sudden b u r s t s of a c t i v i t y when subjec ted t o loud sound, such as produced by j e t a i r c r a f t (Balazs and R o s s 89; Barr 101). e f f e c t s of such n o i s e on h a t c h l i n g s , as w e l l as developing embryos, are p r e s e n t l y unknown.

The b i o l o g i c a l

The emergence of h a t c h l i n g s a t French F r i g a t e Shoals i s only known t o occur at n i g h t , u s u a l l y w i t h i n a few hours a f t e r sunse t . A f i r s t h a n d observa t ion of n a t u r a l depa r tu re from t h e underground nest w a s made by t h e au thor dur ing August 1974. Once emergence w a s i n i t i a t e d , t h e h a t c h l i n g s crawled out s i n g u l a r l y o r i n p a i r s i n a moderately slow b u t continuous stream over a 4- t o 5-min per iod . This w a s followed by very slow c i r c u l a r movements i n t h e immediate area of t h e n e s t s i te . During t h i s pe r iod , which l a s t e d f o r about 2 t o 3 min, t h e ha t ch l ings ' heads were extended upward g iv ing t h e appearance of i n t e n t l y observing some f e a t u r e of t h e sky o r horizon. Following t h e s e d i s t i n c t movements, t h e h a t c h l i n g s movedrap id ly to t h e water i n a number of d i f f e r e n t d i r e c t i o n s .

Although most h a t c h l i n g s s e e m t o e n t e r t h e water a t t h e s h o r e l i n e c l o s e s t t o t h e i r n e s t s i t e , a l l d i r e c t i o n s (except on Tern I s l a n d ) w i l l u l t i m a t e l y r e s u l t i n success due t o t h e s m a l l l and areas involved. excurs ions would, however, r e s u l t i n an increased energy expendi ture t h a t could conceivably l e s s e n t h e ha t ch l ings ' s u r v i v a l c a p a b i l i t i e s dur ing t h e f i r s t few days i n t h e p e l a g i c environment. Groups of ha t ch l ings on East I s l a n d have been recorded t r a v e l i n g i n t h e d i r e c t i o n of a b r i g h t moon, thereby cons iderably inc reas ing t h e d i s t a n c e covered t o reach t h e water.

Unnecessary terrestrial

Preda t ion on ha tch l ings a t French F r i g a t e Shoals t akes p l a c e by bo th s p e c i e s of ghost c r abs , bu t 0. ceratophthuhus is c o n s i s t e n t l y more success fu l due t o i t s l a r g e r s i z e . i n t e r t i d a l zone where both c rabs p e r i o d i c a l l y d i g t h e i r burrows. s e i zed by t h e head, neck, o r f r o n t f l i p p e r s and ea t en e i t h e r on t h e . s u r f a c e o r i n s i d e a burrow. The i n t e r n a l organs and po r t ions of t h e head, neck, and p e c t o r a l muscles are usua l ly t h e only p a r t s consumed. from t h e popula t ion by ghost c r a b s probably does no t exceed 5% o r an es t imated seasonal average of 1,200 ind iv idua l s . t h e course of tagging and o t h e r research act ivi t ies o f f e r s some p o t e n t i a l as a management p r a c t i c e .

Preda t ion only occurs i n o r immediately above t h e narrow Hatchl ings are

The number of h a t c h l i n g s e l imina ted

Reducing t h e number of ghost crabs during

c

c

L

1 7

Although f r i g a t e b i r d s , Fregtzlo m%ncjr, are among t h e s e a b i r d s p re sen t a t French F r i g a t e Shoals , they are no t known t o prey on ha tch l ings on land o r i n t h e in shore waters such as repor ted i n c e r t a i n o t h e r Chelonia popula t ions (Hir th 2 6 5 ) . Nocturnal emergence undoubtedly se rves as a p r o t e c t i v e mechanism aga ins t such preda t ion . Examinations of f e c a l matter a t t h e n e s t i n g and roos t ing sites of f r i g a t e b i r d s have no t r e s u l t e d i n t h e recovery of laminae o r o t h e r i n d i g e s t i b l e p a r t i c l e s of ha t ch l ings . This sugges ts t h a t p reda t ion i s a l s o n o t occur r ing i n the p e l a g i c waters surrounding French F r i g a t e Shoals where f r i g a t e b i r d s per iodi - c a l l y forage .

During t h e U.S. Coast Guard t enure a t East and T e r n I s l a n d s , newly emerged ha tch l ings were r e g u l a r l y d i s o r i e n t e d and a t t r a c t e d t o a r t i f i c i a l l i g h t s and poss ib ly v i b r a t i o n s from d i e s e l gene ra to r s a s soc ia t ed wi th s t a t i o n f a c i l i t i e s . M o r t a l i t i e s from thermal exposure subsequent ly r e s u l t e d i n those ha t ch l ings t h a t were n o t discovered by Coast Guard personnel and t r anspor t ed t o t h e water.

Preda t ion on ha tch l ings by carnivorous f i s h e s a t French F r i g a t e Shoals does not appear t o be s i g n i f i c a n t . It is d i f f i c u l t , however, t o a s s e s s such impacts w i th in t h e marine environment. From J u l y t o October 1974, 101 u lua , Caram ignobi l i s , C. melampygus; 16 wrasses, Thalassoma purpuxewn, Bodianus bilunulatus; and 1 3 gray r ee f sharks, Carcharhinus mblyrhynchos; were captured i n t h e v i c i n i t y of East I s l and t o examine stomach and i n t e s t i n a l con ten t s f o r t h e presence of ha t ch l ings . Major food items recovered included f i l e f i s h , Permagor spiZosoma, and p ieces of u n i d e n t i f i e d c rabs and cephalopods, bu t no evidence of ha t ch l ing preda t ion w a s found.

Hatchl ings t h a t emerge from n e s t s i n t h e c e n t e r of East I s l and are coated wi th p a r t i c l e s of humus, whi le those o r i g i n a t i n g from p u r e ca lcareous sand areas along t h e per iphery and a t t h e w e s t end emerge p e r f e c t l y c lean . For an unknown per iod of t i m e a f t e r e n t e r i n g t h e water, ha t ch l ings coated wi th humus e m i t a t r a i l of organic material. o l f a c t o r y cues t o p o t e n t i a l f i s h p reda to r s , o r somehow poss ib ly serves as a d e t e r r e n t t o preda t ion .

It would be of va lue t o know i f th i s provides

The types and l e v e l s of p reda t ion on ha tch l ings t h a t emerge from n e s t s a t o t h e r i s l a n d s i n t h e northwestern segment of t h e Hawaiian Archipelago are p r e s e n t l y unknown. Both s p e c i e s of ghost c rabs as w e l l as many of t h e same ca rn i - vorous f i s h e s are a l s o p re sen t a t t h e s e loca t ions .

Hawaiian Chelonia ha tch l ings t h a t emerged from a captive n e s t a t Sea L i f e Park on Oahu i n 1976 w e r e ex t ens ive ly preyed upon by wi ld mongoose, Herpestes awopunctatus, (Bourke et a l . 131) . H a w a i i i n 1883 and is now p resen t on a l l of t h e main i s l a n d s except Kahoolawe, Lanai, and Niihau.

This mammal w a s f i r s t in t roduced t o

3.3 J u v e n i l e s , Subadul ts , and Adults

3.31 Longevity

There is v i r t u a l l y no information on t h e l i f e spans of Hawaiian CheZonia o r marine t u r t l e s of any o t h e r populat ion. could,however, be cons iderable i n view of t h e f ind ings t h a t t u r t l e s a t c e r t a i n r e s i d e n t forag ing areas may r e q u i r e more than 59 yr t o reach matur i ty (Table 4 ) .

The maximum age obta ined

18 n

Tag recove r i e s of f o u r females and e i g h t males have been made i n the Hawaiian Archipelago a f t e r per iods exceeding 9 y r . A l l of t h e s e t u r t l e s were recorded as being a d u l t s when o r i g i n a l l y tagged. The longes t t i m e between i n i t i a l t agging and las t recovery f o r a female i s 12.1 y r , whi le t h e longes t f o r a male i s 14.3 y r . t h e longes t e lapsed t i m e r epor t ed f o r t h e recovery of a tagged n e s t i n g green t u r t l e w a s 19 y r (Carr e t aZ. 153) .

I n a 22-yr o l d r e sea rch program a t Tortuguero, Costa R i c a ,

I n v e s t i g a t i o n s by t h e au thor are c u r r e n t l y underway t o develop a method of e s t ima t ing t h e age of Hawaiian CheZonia based on a n n u l i discovered i n t h e p l a s t r a l andl imb bones. Thesebands are most prominent i n t h e d o r s a l segments of t h e hyoplas t rons . Prel iminary r e s u l t s have ind ica t ed a p o s i t i v e c o r r e l a t i o n between t h e number of bands and t h e s i z e of t h e t u r t l e . To supplement t h i s r e sea rch , au toradiographs are be ing at tempted wi th s e c t i o n s of bones and laminae obta ined from Chelonia a t Enewetak A t o l l (Marshall I s l a n d s ) where nuc lea r devices were detonated dur ing t h e 1950's .

3.32 Competitors

Monk seals wi th pups a t t i m e s compete wi th green t u r t l e s f o r p re fe r r ed basking space a long t h e s h o r e l i n e s a t French F r i g a t e Shoals. Competition f o r basking space wi th t h i s s p e c i e s has a l s o been documented a t t h e small s lop ing rock ledge a t Necker I s l a n d (Balazs 72) . competed wi th CheZonia f o r terrestrial space. form of m i l i t a r y e x e r c i s e s , cons t ruc t ion of f a c i l i t i e s , commercial f i s h i n g , w i l d l i f e r e sea rch , and p u r s u i t of ornamental g l a s s f i s h i n g f l o a t s (Amerson 30; Amerson e t a l . 32; Clapp and W i r t z 1 7 1 ; Clapp and K r i d l e r 1 7 2 ; Ely and Clapp 211; Woodward 556).

Humans i n t h e NWHI have p e r i o d i c a l l y This has p r i n c i p a l l y been i n t h e

A t some of t h e r e s i d e n t fo rag ing areas i n t h e main i s l a n d s , humans compete w i t h green t u r t l e s f o r space by s e t t i n g g i l l n e t s and b a i t e d hooks, and by r epea ted ly i n t r u d i n g i n t o h a b i t a t w i th boa t s and scuba. a s i g n i f i c a n t f a c t o r a t t h e p re sen t t i m e , compet i t ion a l s o exists f o r c e r t a i n k inds of ben th ic a l g a e (Codiwn, Ulva, Gracilaria) t h a t are ea t en by both green t u r t l e s and humans (Abbott and Williamson 1; For tne r 228). Herbivorous f i s h e s and some i n v e r t e b r a t e s i n t h e Hawaiian Archipelago a l s o u t i l i z e ben th ic a lgae as food sources , bu t t h e e x t e n t of t h i s compet i t ion is n o t known.

Although probably not

3.33 P reda to r s

Tiger sha rks are v i r t u a l l y t h e only known n a t u r a l p reda to r s of j u v e n i l e , subadul t and a d u l t Hawaiian Chelonia. Considering j u s t t h e marine environment, t h i s is e s s e n t i a l l y t h e case f o r a l l popula t ions of Cheloniidae. I n t e n s i v e shark r e sea rch and e r a d i c a t i o n programs p e r i o d i c a l l y conducted around t h e main Hawaiian I s l a n d s found t h a t 18% (Ikehara 275), 10.8% (Fujimoto and Sakuda 230>, and 12.7% (Tes t e r 487) of t h e t i g e r sharks examined wi th food i n t h e i r stomachs had been feeding on t u r t l e s . During 1974 t h e au thor captured four t i g e r sha rks a t French F r i g a t e Shoals ranging from 1.6 t o 3.4 m i n length . Two of t h e s e sharks contained p i eces of t u r t l e s r ep resen t ing t h r e e i n d i v i d u a l s , one of which w a s an a d u l t . A t French F r i g a t e Shoals and P e a r l and Hemes Reef, green t u r t l e s were recorded by Taylor and N a f t e l (481, 482) i n 31% and 36%, r e s p e c t i v e l y , of

c

c

c

19

P

u

the t i g e r sharks captured t h a t contained food. bones, laminae and beaks revea led t h e presence of n ine d i f f e r e n t t u r t l e s i n t h e f i v e t i g e r sharks from French F r i g a t e Shoals , and e i g h t d i f f e r e n t t u r t l e s i n t h e fou r t i g e r sharks from P e a r l and Hermes Reef (Balazs 70). and Hermes Reef accounted f o r f i v e t u r t l e s t h a t ranged from an es t imated 53 t o 64 cm i n s t r a i g h t carapace length . Four of t h e t u r t l e s recovered from sha rks a t French F r i g a t e Shoals were found t o be a d u l t s of unknown s e x ranging from an es t imated 8 1 t o 94 c m i n s t r a i g h t carapace length . The s i z e s of t h e t i g e r sharks captured ranged from 2.8 t o 4.3 m, bu t s i g n i f i c a n t r e l a t i o n s h i p s (P<0.05) were not found between s i z e of t h e t u r t l e s e a t e n and t h e sha rk ' s l eng th o r mouth width, which ranged from 26 t o 51 cm. r e v i s e t h e preva len t b e l i e f t h a t n a t u r a l p reda t ion on a d u l t green t u r t l e s i s minimal (Hir th 265). are unknown, t h e r e f o r e i t i s not p o s s i b l e t o determine how long t h i s material may have been r e t a i n e d i n each stomach. laminae and beaks p re sen t among t h e recovered items, i t seems l i k e l y t h a t ke ra t in - i z e d s t r u c t u r e s are t h e most d i f f i c u l t t o d i g e s t , if i n f a c t they can be d iges t ed a t a l l . be p e r i o d i c a l l y e l imina ted by r e g u r g i t a t i o n .

An a n a l y s i s of t h e recovered

A s i n g l e shark a t P e a r l

Based on t h e s e f i n d i n g s , i t may be necessary t o

The d i g e s t i o n rates of green t u r t l e p a r t s i n t i g e r sha rks

Due t o t h e comparatively l a r g e q u a n t i t y of

It has been suggested t h a t i n d i g e s t i b l e i t e m s e a t e n by t i g e r sha rks may

The feeding mechanism of t h e t i g e r shark inc ludes heavy sawing a c t i o n s by both j a w s which can e f f e c t i v e l y cu t up l a r g e green t u r t l e s i n t o p i eces s u i t a b l e t o swallow. However, f o r smaller s i z e t u r t l e s t h i s i s appa ren t ly not necessary. On numerous occasions j u v e n i l e Hawaiian CheZonia ranging from 35 t o 50 cm i n s t r a i g h t carapace l eng th have been recovered whole from t i g e r shark stomachs (Taylor and N a f t e l 481, 482; Tester 487; C. Chiswick and D. Her tz , pe r sona l cornu- n i c a t i o n s ) . Tinker (490, 492) repor ted t h a t a 23 kg t u r t l e w a s found i n a l a r g e t i g e r shark caught o f f Oahu i n 1935.

Tiger sharks have been r e g u l a r l y recorded a t French F r i g a t e Shoals i n water only a few meters deep ad jacen t t o t h e va r ious i s l a n d s (Balazs and Whittow 90). I n A p r i l 1956, a 3-m shark w a s observed a t t a c k i n g a l a r g e t u r t l e o f f Whale- Skate I s l a n d ( P a c i f i c Ocean F i s h e r i e s I n v e s t i g a t i o n s 392). I n June 1974, t h e au thor watched a 2-m t i g e r shark beach i t s e l f momentarily on E a s t I s l and dur ing t h e unsuccessfu l pursuit : of a j u v e n i l e green t u r t l e t h a t had f l e d t o t h e sho re l ine . Two t i g e r sharks w e r e a l s o recorded swimming c l o s e t o a copula t ing p a i r of t u r t l e s and t h r e e male e s c o r t s , bu t no a t t a c k r e s u l t e d . I n August 1978, a l a r g e t i g e r shark w a s observed f o r 2 h repea ted ly t r y i n g t o reach a dead a d u l t t u r t l e w i th a missing head and f l i p p e r s t h a t had become beached i n t h e wavewash a t Tr ig I s l a n d (G. C. Whittow, personal communication). I n view of t h e s e s i g h t i n g s , i t is s i g n i f i c a n t t o no te t h a t dur ing an 11-mo per iod on E a s t I s l and i n 1946, former Coast Guardsman H . E. Finch (personal communication) never s a w a t i g e r shark,even though h e r e g u l a r l y set b a i t e d hooks and caught numerous Carcharhinus ambly- rhynchos.

Approximately 10% of t h e t u r t l e s n e s t i n g on East I s l and have healed o r f r e s h i n j u r i e s o r amputations of t h e i r hind f l i p p e r s t h a t are sugges t ive of shark a t t a c k . (Anonymous 611, 613, 616) w a s observed 25 mo Later basking on Whale-Skate I s l and wi th a f r o n t f l i p p e r f r e s h l y amputated a t t h e humerus. a f r e s h l y amputated f r o n t f l i p p e r w a s seen basking a t E a s t I s l a n d during J u l y 1976. Two o t h e r a d u l t males wi th a completely missing bu t healed f r o n t f l i p p e r

A tagged female r e l eased from c a p t i v i t y o f f Oahu i n December 1973

An a d u l t m a l e w i th

20 L

have a l s o been recorded wh i l e basking. A tagged a d u l t female wi th one f r o n t and both hind f l i p p e r s missing, but p a r t i a l l y hea led , w a s seen basking a t T r i g I s l and i n June 1973 (Olsen 384). An a d u l t female tagged a t E a s t I s l and i n 1973 w a s recovered 1 3 mo l a t e r o f f Maui and found t o have a r e c e n t l y amputated f r o n t f l i p p e r (Balazs 57) . A t u r t l e wi th a missing f r o n t f l i p p e r w a s seen a t Gin I s l and i n 1923 (Wetmore 541). A t L i s i a n s k i I s l a n d , Taylor (475) recorded an a d u l t male i n a moribund condi t ion wi th i t s t a i l f r e s h l y amputated.

A

I n a t r i a l c a l c u l a t i o n of t h e dep le t ion ra te i n t h e Hawaiian Chelonia popula t ion , Hendrickson (262) specu la t ed t h a t no less than 100 and poss ib ly as many as 1,000 t u r t l e s pe r year of 23 kg and above w e r e l o s t t o t i g e r sha rk preda t ion .

The only o t h e r n a t u r a l p reda to r of pos tha tch l ing green t u r t l e s documented i n t h e Hawaiian Archipelago i s t h e l a r g e grouper , Epinephelus tauvina. In October 1974, a 205 kg specimen caught o f f Molokai w a s found t o con ta in a p a r t i a l l y d iges t ed bu t whole j u v e n i l e t u r t l e measuring approximately 52 cm i n s t r a i g h t carapace l eng th (K. Mench, pe r sona l communication). S igh t ings and cap tu res of t h e s e groupers are rare i n t h e Hawaiian Archipelago, bu t specimens as l a r g e as 365 kg have been documented.

3.34 Epizoics , d i s e a s e s , abnorma l i t i e s , and i n j u r i e s

Barnacles commonly found on Hawaiian CheZonia inc lude CheZonibia tes tu- dinaria (Bryan 143; Edmondson 205; Gordon 237) and Platylepas hexasty 20s ( i d e n t i - f i e d by W . J. Cooke). CheZonibia i s u s u a l l y confined t o t h e carapace, p l a s t r o n and head, whi le PlatyZepas a t t a c h e s i t s e l f p r i n c i p a l l y t o t h e s k i n r eg ions , occas iona l ly i n l a r g e numbers. Large specimens of CheZonibia can measure 50 mm i n diameter a t t h e base , whi le PlatyZepas usua l ly does no t exceed 6 mm. t u r t l e s c o n s i s t e n t l y h o s t l a r g e r specimens of t h e s e two ba rnac le s than t h e j u v e n i l e o r subadul t t u r t l e s , ( i d e n t i f i e d by W. J. Cooke) has been recovered from t h e f r o n t f l i p p e r of a j u v e n i l e i n Kaneohe Bay, Oahu, b u t is apparent ly a rare spec ie s i n Hawaiian Che 2 onia .

Adult

The burrowing ba rnac le , Stephanolepas mw?icata,

Two specimens of t h e p i s c i c o l i d leech,Ozobranchus branchiatus, ( i d e n t i - f i e d by M. D. Dai ley) have been recovered from a j u v e n i l e green t u r t l e a t Necker I s l and . During 1978 green, hawksbi l l and loggerhead t u r t l e s on d i s p l a y at Sea L i f e Park (Oahu) became heav i ly p a r a s i t i z e d wi th 0. rnargoi ( i d e n t i f i e d by M. D. Dai ley) . loggerhead t h a t had been found of f Molokai and brought i n t o c a p t i v i t y (Altonn 20; McKinney 351). A t p r e s e n t , no o t h e r records e x i s t f o r Ozobranchus i n t h e Hawaiian Archipelago.

This i n f e s t a t i o n i s thought t o have o r i g i n a t e d from a s t r a y

c

c

The bucca l c a v i t y of Hawaiian Chelonia occas iona l ly con ta ins one o r two

Th i s c rus t acean has a l s o been found on t h e neck and hind f l i p p e r s l i v i n g specimens of t h e t a l i t r o i d e a n amphipod, Hyachelia tortugae ( i d e n t i f i e d by W. J. Cooke). i n a s s o c i a t i o n w i t h s u p e r f i c i a l s k i n l e s i o n s .

In June of 1974, t h e au thor observed a remora, Echenesis s p . , a t t ached t o t h e p l a s t r o n of an a d u l t female o f f East I s l a n d .

c

2 1

Q

The red a l g a , Polysiphonia tsudana, ( i d e n t i f i e d by D. J. R u s s e l l ) , commonly occurs on t h e s k i n of Hawaiian Chelonia and occas iona l ly on t h e p l a s t r o n and Inconel t ags ( see Newbert e t aZ. 375 f o r co lo r photograph). This s p e c i e s w a s f i r s t descr ibed from the neck of a green t u r t l e a t Laysan I s l a n d (Hollenberg 270). Other a lgae t h a t have been p e r i o d i c a l l y found by t h e au thor on t h e s k i n , carapace and p l a s t r o n of Hawaiian green t u r t l e s inc lude t h e fol lowing ( i d e n t i f i e d by D. J . Russe l l ) : Rhodophyta ( red) - Polysiphonia dotyi , Acrochaetium graeile, Falken- bergia rufolanosa, Melobesia s p . ( enc rus t ing ) ; Phaeophyta (brown) - Sphacelmia furcigeria, S. novae-hoZZandiae, Ectocarpus indicus; chlorophyta (green) - Enteromorpha eZathrata; Cyanophyta (blue-green) - Lyngbya majuscula, L. cinerescens, Oseillatoria sp . A t Necker I s l and , i n t e r t i d a l c r abs , Grapsus grapsus, have been observed on t h e carapace and f l i p p e r s of basking t u r t l e s feeding on a l g a l mats . When t h e t u r t l e s r e t u r n t o t h e water, surgeonf i sh , Acanthurus sandvicensis, are a l s o a t t r a c t e d t o these e p i z o i c s f o r graz ing purposes (Balazs 72). where green t u r t l e s r e g u l a r l y p o s i t i o n themselves while wrasses and surgeonf i sh feed on ep izo ic s . n o r t h shore of Tern I s l a n d appears i n an ar t ic le by E l i o t (208).

A t French F r i g a t e Shoals, d i s c r e t e underwater sites have been i d e n t i f i e d

A co lo r photograph showing t h i s symbiot ic behavior o f f t h e

Neoplas t ic growths ranging from s m a l l w a r t s t o huge masses 25 c m i n diameter o c c u r o n 5 % t o 10% of t h e green t u r t l e s observed dur ing each breeding season a t French F r i g a t e Shoals. Common sites of t h e s e neoplasms inc lude the neck (Kr id l e r 322), f l i p p e r s , t a i l , and f o l d of f l e s h where t h e t ags are a t tached . I n many of t h e a f f e c t e d t u r t l e s small growths a l s o occur i n a s s o c i a t i o n wi th t h e eyes, apparent ly o r i g i n a t i n g from t h e conjunct iva and s c l e r a . Two a d u l t males seen basking a t East I s l a n d had eyegrowths s o ex tens ive t h a t most v i s i o n w a s e l imina ted . However, except f o r t h e presence of t hese growths, t h e t u r t l e s appeared hea l thy . subadul t , and a d u l t t u r t l e s found o f f Oahu s i n c e 1977, as w e l l as from an ind iv idua l he ld i n c a p t i v i t y f o r more than 5 y r . t o t he Regis t ry of Tumors i n Lower Animals (Smithsonian I n s t i t u t i o n ) where they w e r e analyzed and subsequent ly i d e n t i f i e d as benign f ibropapi l lomas t h a t have been occas iona l ly repor ted i n o t h e r marine t u r t l e populat ions. The formation of t hese growths i n Hawaiian Chelonia i s descr ibed as fol lows. The epidermis pro- l i f e r a t e s and forms pegs extending i n t o the dermal connect ive t i s s u e . A t t h e same t i m e , dermal p a p i l l a e conta in ing connect ive t i s s u e , vascu la r channels , and a s soc ia t ed melanophores grow between t h e epidermal pegs t o provide nourishment. The s u r f a c e epidermis produces abundant l a y e r s of k e r a t i n and occas iona l ly t rapped c e l l s w i th in t h e dermis form p e a r l s o r epidermal c y s t s conta in ing concen t r i c r i n g s of k e r a t i n . Subsequent enlargement r e s u l t s t o an inc reas ing e x t e n t w i th expansion of t h e f i b r o u s component. Larger tumors are t h e r e f o r e mostly a monotonous f i e l d of f i b r o c y t e s wi th some i n t e r s p e r s e d melanophores and a few epidermal pegs near t h e su r face . (J. C. Harshbarger, personal communication and 255, 256, 257). The e t i o l o g y of f ibropapi l lomas is unknown. I n Hawaiian Chelonia, they are n o t a s soc ia t ed wi th t h e f o r e i g n organic materials ( leeches , ba rnac le s , a lgae ) t h a t have been suggested as p o s s i b l e causes i n o t h e r marine t u r t l e popula t ions . The p o s s i b i l i t y of a v i r a l o r i g i n i s p r e s e n t l y being i n v e s t i g a t e d by e l e c t r o n microscopy of t i s s u e b i o p s i e s from t h e p r i c k l e c e l l l a y e r of t h e epidermis of e a r l y growths on Hawaiian Chelonia (J. C. Harshbarger, personal communication). Other p o s s i b l e causes inc lude repeated con tac t wi th carc inogenic subs tances , such as might occur a t sewage d ischarges o r from corroding Monel t ags and excess ive exposure t o s o l a r r a d i a t i o n

A n a r r a y of growths has been obtained from moribund j u v e n i l e ,

These specimens were submitted

Invasion of t h e underlying muscle does no t u s u a l l y t ake p l ace

22

whi le f l o a t i n g a t t h e surface o r basking. Fibropapi l lomas on green t u r t l e s may n o t , however, be a new occurrence i n t h e Hawaiian popula t ion . A photograph taken i n 1923 a t French F r i g a t e Shoals shows a basking t u r t l e wi th what appears t o be a growth on t h e d o r s a l s u r f a c e of i t s l e f t f r o n t f l i p p e r (Tinker 492; Anonymous 568). It is l i k e l y t h a t some t u r t l e s wi th f ibropapi l lomas are more s u s c e p t i b l e t o t i g e r shark p reda t ion due t o decreased v i s i o n and maneuverabi l i ty whi le swimming.

A sa l t c r y s t a l depos i t weighing 8 g and measuring 20 mm i n diameter w a s e x t r a c t e d from t h e corner of t he l e f t eye of a j u v e n i l e green t u r t l e a t L i s i a n s k i I s land . This cond i t ion probably represented some dysfunct ion of t h e lacrimal gland or blockage of t h e duct t h a t t r a n s p o r t s t h e hype r sa l ine s e c r e t i o n .

During J u l y 1978, a t u r t l e t h a t nes t ed on Laysan I s l a n d approximately 285 m in l and became d i s o r i e n t e d dur ing h e r r e t u r n t o t h e oc"ean and d i ed t h e fo l lowing morning from thermal exposure (P. and B. Johnson, personal communica- t i o n ) . exp la in t h i s abnormal behavior . have a l s o been recorded by t h e au thor a t Canton, a l a r g e 2O5O'S, long. 171'43'W ( see P r i t c h a r d 418).

There were no growths on t h e t u r t l e ' s eyes o r o t h e r e x t e r n a l s i g n s t o S imi l a r o r i e n t a t i o n problems by n e s t i n g Chelonia

low c o r a l a t o l l a t la t .

Capt ive growth s t u d i e s conducted wi th 120 green t u r t l e s c o l l e c t e d as ha tch l ings a t French F r i g a t e Shoals r e s u l t e d i n 31 m o r t a l i t i e s from va r ious causes by 7 mo of age (Bardach 98; Bardach and He l f r i ch 99, 100) . Postmortem examina- t i o n s revea led t h a t s i x of t h e s e t u r t l e s had l e s i o n s c o n s i s t i n g of f o c a l granu- lomas c h a r a c t e r i s t i c of t u b e r c u l o s i s . Serotype 8 w e r e i s o l a t e d from one of t h e two specimens subsequent ly c u l t u r e d (Brock e t aZ. 134) . The M. aviwn complex is usua l ly a s s o c i a t e d wi th p o u l t r y , ca t t le , and swine, and t h i s i s o l a t i o n from Chelonia appa ren t ly represented t h e f i r s t record f o r a poiki lotherm. The source of t h e i n f e c t i o n w a s no t determined, however neona ta l con tac t i n t h e underground n e s t wi th s u b s t r a t e contaminated by s e a b i r d s would appear t o be a p l a u s i b l e rou te . t h e occurrence of Mycobacteriwn i n n a t u r a l l y occurr ing s e a b i r d s o r green t u r t l e s i n t h e Hawaiian Archipelago.

Bac i l l i i d e n t i f i e d as Mycobacterium av&m

No information p r e s e n t l y exists on

SalmoneZla weZteureden ( i d e n t i f i e d by R. M. Nakamura) and S. san-diego Type B ( i d e n t i f i e d by M. H. Crumrine) have been i s o l a t e d from t h e f e c e s of cap t ive r ea red Hawaiian CheZonia, bu t aga in noth ing i s known of occurrence i n t h e wi ld .

P a r a l y s i s of a f r o n t f l i p p e r due t o a s e p t i c n e c r o s i s of t h e head of t h e humerus has been observed i n two c a p t i v e r ea red Hawaiian Chelonia. The e t i o l o g y of t h i s a f f l i c t i o n could n o t be determined (M. H. Crumrine, M. W . Balk, N . E. Palumbo, and G. Liese, personal communications). Necrosis of unknown o r i g i n has a l s o been found i n t h e t e rmina l scales of t h e f r o n t f l i p p e r s of j u v e n i l e s a t Kure (Balazs 73) .

I n j u r i e s t o t h e carapace and p l a s t r o n of Hawaiian Chelonia are known t o r e s u l t from v i o l e n t con tac t w i th rocks and o t h e r underwater s u b s t r a t e acci- d e n t a l l y encountered wh i l e fo rag ing i n rough s u r f c l o s e t o shore (Wright e t a l . 557). A t c e r t a i n c o a s t a l areas on t h e i s l a n d of H a w a i i , tsunamis have c a r r i e d j u v e n i l e and subadul t green t u r t l e s over 100 m in l and where they became s t randed and unable t o r e t u r n t o t h e water without human a s s i s t a n c e (A. L. Howard,

c

c

c

2 3

personal communication). along remote coastal areas of the Hawaiian Archipelago following a large tsunami.

It is unknown how extensive these strandings may be

Both injuries and mortalities of Hawaiian green turtles are known to occur incidental to fishing activities directed at other species (Anonymous 570, 583). This can result from contact with fishhooks and from entanglement in gill nets, rope, and monofilament fishing line. In addition, discarded items can at times be hazardous to turtles, as evidenced by a juvenile captured off Oahu that had a part from a plastic container stuck around its neck.

3.4 Nutrition and Growth

3.41 Food sources L

4

Green turtles that reside in coastal areas of the Hawaiian Archipelago have been documented by the author feeding on 56 species of algae, 1 marine angiosperm and 9 types of invertebrates (Table 9). However, 9 species of algae out of the approximately 400 species present in the Hawaiian Archipelago account for the major food sources utilized (Table 10). components of juveniles, subadults, and adults in both segments of the Hawaiian Archipelago, joined by Pterocladia and Amansia in the main islands, and Caulerpa, Turbinaria and Spyridia in the NWHI (Ralazs 71, 72, 75, 81, 82). In 1933 Galtsoff (231), quoted by Amerson e t a l . ( 3 2 ) , noted that the stomachs of green turtles examined at Pearl and Hermes Reef contained large amounts of Codiwn. Abbott and Williamson (1) indicated that Hawaiian green turtles are known to eat Sargassum, thereby probably accounting for its native name of limu honu. listed "turtle limu" as an alternate common name for UZva. Hirth (265) stated that "green algae" was the food of adult turtles off Maui. Wetmore (542), quoted by Mellen ( 3 5 3 ) , stated that green turtles in the NWHI "browse in submarine fields of algae." the only published information on the food sources of Hawaiian CheZonia.

Codiwn and UZva are major dietary

Fortner (228)

Prior to the author's research program, these references constituted

The methods used to determine the food sources of Hawaiian Chetonia listed in Tables 9 and 10 have included: 1) analysis of gastrointestinal contents salvaged from mortalities due to tiger shark predation, accidental and intentional human capture, and strandings of unknown causes; 2) direct observations of foraging made from shore and underwater with scuba; 3) recovery of incompletely digested food from fecal pellets found at resident foraging areas; 4) recovery of food directly from the mouths of turtles captured uninjured while actively feeding; and 5) sampling of stomach contents using a flexible plastic tube (Balazs 86) and a small diameter flexible grasping tool (Kam and Balazs 286) inserted through the esophagus.

The distribution and abundance of benthic algae in the Hawaiian Archi- pelago are not well known, however standing crop densities of the species preferred by green turtles appear to be far greater in the main islands. For example, certain foraging areas around the islands of Hawaii, Maui, Oahu, and Kauai have dense growths of the red alga, Pterocladia cap i l keea ; while in the NWHI, this is a rare species only known to occur in small quantities at Lisianski Island. island foraging areas, but relatively scarce in the northwestern segment of the

Concomitantly, Amansia glomerata is abundant at many main

2 4

Hawaiian Archipelago. at certain sites in the NWHI, has never been found as a dietary component in the main islands even though it occurs at a number of locations. note that the genus Caulerpa, and particularly the species racemosa, contain the toxic constituent "caulerpicin" which can produce symptoms in humans similar to ciguatera fish poisoning (Doty and Aguilar-Santos 203). It is unknown what effects, if any, caulerpicin may have on green turtles. Tuxbinaria o m t a and Spyridia filamentosa, dietary components in the "HI, are also not eaten in the main islands where they at times grow in the same general area as the favored PterocZadia capillacea and h a n s i a glomerata.

Caulerpa racemosa, a green alga regularly eaten by Chelonia

It is significant to

Marine angiosperms, such as turtle grass, Thalassia testudinwn, constitute the principal food source for a number of Chelonia populations (Hirth 265), however hawaiiana. meadows of low densities around the main islands and at Midway.

the only species represented in the Hawaiian Archipelago is HaZophiZa This seagrass has oval blades 1 to 3 cm long and only occurs in small

It is of interest to note that two species of red algae, Aeanthophora spieifera and Hynea musei fomis , recorded in the diets of green turtles in the main islands are introductions to Hawaiian waters from Guam and Florida, respec- tively. In addition, the recovery of Polysiphonia tsuduna from the stomach of a juvenile at Kure suggests that this turtle was grazing on either its own or another turtle's epizoic algal mat.

Juvenile and subadult turtles in the M I , particularly at Midway and

The eyes Kure, have been observed voraciously feeding on the invertebrates PhysaZia, VeZeZZa, and Janthina that periodically drift into the coastal areas. and gastrointestinal tracts of these turtles are apparently not adversely affected by the nematocysts present in both PhysaZia and Velella. islands the small black sponge, Chondrosz"a chuealla, is also periodically eaten by green turtles, usually in association with Codiwn edule. Examinations of fecal pellets have shown that these sponges, as well as pieces of Codiwn and other algal species, are able to pass through the gastrointestinal tract in a relatively undigested state. Although small crustaceans can sometimes be found living on algal filaments, these animals have not been present in the stomach samples examined.

In the main

Synthetic items such as plastic fragments, pieces of plastic bags, cloth, string, and small diameter polypropylene line have occasionally been found in Hawaiian Chelonia. An adult female captured by a fisherman in November 1978 off the north shore of Lanai contained large pieces of both black and clear plastic bags throughout its intestines. This material was also present in fecal matter in the rectum, thereby indicating that blockage of the tract had not taken place. Plastic items of this nature found floating at the surface appear to be mistaken by some turtles for coelenterates or other edible invertebrates. Fragments of terrestrial vegetation in the stomach as well as tar stains in the mouth have also been occasionally found in Hawaiian Chelonia.

The food sources of Hawaiian Chelonia <35 cm living in the pelagic environment are completely unknown due to the absence of human contact (section 2.22). It is logical to assume that during this period the turtles are carnivores

P

R

a

c

c

2 7

.”.

Differences have a t t i m e s been found i n t h e spec ie s of a lgae p re sen t a t va r ious d i s t a n c e s a long the i n t e s t i n e s , and i n t h e two compartments t h a t comprise t h e stomach. Pe r iod ic d i e t a r y s h i f t s of t h i s na tu re probably h e l p Hawaiian Chelonia m e e t t h e i r requirements f o r e s s e n t i a l n u t r i e n t s (i.e. amino a c i d s , f a t t y a c i d s , v i tamins , minera ls ) t h a t may be more concentrated i n some s p e c i e s of a lgae . Ce r t a in of t h e s e n u t r i e n t s may a l s o be synthes ized by mic rob ia l a c t i o n w i t h i n t h e i n t e s t i n e . t u r t l e s i n t h e wi ld would vary wi th age, a c t i v i t y , and reproduct ive condi t ion .

A s w i th o t h e r v e r t e b r a t e s , t h e n u t r i t i o n a l requirements of green

It i s no t unusual f o r j u v e n i l e and subadul t green t u r t l e s i n t h e Hawaiian Archipelago t o b i t e on hooks b a i t e d wi th squid , shrimp, and f i s h f l e s h (Carter 158; Anonymous 583). r eg res s ions t o t h e feeding h a b i t s e x h i b i t e d wh i l e l i v i n g i n t h e p e l a g i c environ- men t .

These carnivorous i n t e r l u d e s probably r ep resen t

3.43 Growth rates

The mean rates of growth of immature green t u r t l e s (37-59 cm) occur r ing n a t u r a l l y a t seven r e s i d e n t areas i n t h e Hawaiian Archipelago have been found t o range from 0.08 t o 0.44 cm/mo i n s t r a i g h t carapace l eng th (Table 4) . r ap id growth t akes p l a c e a long t h e Kau c o a s t l i n e of H a w a i i (0.38-0.52 cm/mo), whi le t h e s lowes t occurs a t French F r i g a t e Shoals (0.02-0.13 cm/mo) and Kure (0.04-0.12 cm/mo). I n a d d i t i o n t o t h e growth d a t a presented i n Table 4 , 34 hea l thy appearing immature t u r t l e s have been recovered a f t e r pe r iods ranging from 2 t o 20 mo i n which no measurable growth could be de t ec t ed . This has inc luded 6 t u r t l e s a t Midway, 3 a t L i s i a n s k i , 24 a t French F r i g a t e Shoals , and 1 a t Necker. One of t h e t u r t l e s a t French F r i g a t e Shoals w a s a 68 c m subadul t t h a t showed no i n c r e a s e i n s t r a i g h t carapace l eng th a f t e r an i n t e r v a l of 20 mo. The t u r t l e a t Necker w a s a 42.5 c m j u v e n i l e t h a t w a s recaptured a f t e r a 17-mo i n t e r v a l . Although t h e apparent absence of growth could, i n some cases, poss ib ly be a t t r i b - u ted t o measuring e r r o r s , t h i s i s n o t be l i eved t o be a s i g n i f i c a n t f a c t o r i n t h a t t h e au tho r has pe r sona l ly taken most of t h e i n i t a l and recovery measurements. The causes and impl i ca t ions of c e s s a t i o n of growth among some t u r t l e s are unknown and f u r t h e r i n v e s t i g a t i o n s are warranted.

The most

I n a d d i t i o n t o u t i l i z i n g large-mesh t ang le n e t s , t h e . c a p t u r e of immature t u r t l e s f o r growth s t u d i e s i n t h e Hawaiian Archipelago has been accomplished wi th long-handled scoop n e t s I n t h e NWHI, p a r t i c u l a r l y at L i s i a n s k i , p e r i o d i c basking a l s o provides access t o t h e s e t u r t l e s . The use of curved carapace l eng th f o r d e t e c t i n g growth has been found t o be gene ra l ly u n r e l i a b l e due t o v a r i a t i o n s i n p o s i t i o n i n g t h e f l e x i b l e measuring t ape (see KrPdler 322), and changes i n t h e curva ture of t h e carapace t h a t appear t o be independent of an inc rease i n s i z e . t o be u n r e l i a b l e . This is probably due t o d i f f e r e n c e s i n t h e amount of food material i n t h e g a s t r o i n t e s t i n a l t r a c t , a component t h a t can comprise up t o 18% of t h e weight of j u v e n i l e Hawaiian Chelonia (Balazs 81).

and by hand whi le d iv ing wi th scuba.

The use of body weight has a l s o been found

The d i f f e r e n t rates of growth exh ib i t ed by immature t u r t l e s a t v a r i o u s l o c a t i o n s i n t h e Hawaiian Archipelago are most l i k e l y a func t ion of t h e sources and abundance of food a t t h e r e s i d e n t areas ( s e c t i o n 3.41). Seawater temperature would be expected t o have some in f luence , bu t t h i s i s no t ev ident based on t h e a v a i l a b l e da t a . A t Kure and Midway, and probably extending t o the

28

sou theas t as f a r as L i s i a n s k i I s l a n d , mean monthly sea-surface temperatures range from a low of 20.5"C dur ing February, t o a h igh of 26.2"C dur ing August and September (Clapp and W i r t z 172; Ga l t so f f 231). French F r i g a t e Shoals (F igure 5) where e s s e n t i a l l y t h e s a m e growth rates have been recorded. where cons iderably f a s t e r growth occurs . accounts f o r t h e d i f f e r e n c e s i n growth cons ider ing t h e polygamous behavior d i sp layed by a d u l t s dur ing each breeding season a t French F r i g a t e Shoals (Balazs 57) .

This i s 2" t o 3 O C coo le r than

Temperatures a t French F r i g a t e Shoals are similar t o t h e main i s l a n d s It is u n l i k e l y t h a t a g e n e t i c b a s i s

A growth rate of 0.18 cm/mo w a s recorded a t French F r i g a t e Shoals f o r a subadul t female r e l e a s e d o f f Oahu 2 8 mo earlier a f t e r an extended pe r iod iti c a p t i v i t y (Table 11). found t o be t h e same s i z e when recovered by t h e au tho r 11 mo la ter wh i l e d iv ing a long t h e Kau c o a s t l i n e . r e l e a s e d a t French F r i g a t e Shoals (Altonn 15) e s t a b l i s h e d res idency around an i r o n seawall o f f t h e northwestern co rne r of Tern I s l and . U . S . Coast Guard personnel r e g u l a r l y fed t h e s e t u r t l e s f r e s h f i s h s c r a p s and t h e au thor w a s subsequent ly a b l e t o cap tu re them f o r growth measurements (Table 11). 0.71 cm/mo over an 8-mo pe r iod , thereby c o n s t i t u t i n g t h e most r a p i d growth rate thus f a r documented f o r a Hawaiian green t u r t l e l i v i n g i n t h e wi ld . This t u r t l e subsequent ly d isappeared , wh i l e t h e o t h e r one w a s found washed up on t h e beach dead of unknown causes .

A 42.5 c m j u v e n i l e r a i s e d i n c a p t i v i t y from a h a t c h l i n g was

Two of 26 o t h e r j u v e n i l e s r a i s e d from h a t c h l i n g s and

One of t h e t u r t l e s grew

For comparative purposes , growth d a t a are a l s o presented i n Table 11 f o r f o u r Hawaiian Chelonia r a i s e d i n c a p t i v i t y from h a t c h l i n g s f o r pe r iods ranging from 57 t o 81 mo. The d i e t s f ed t o t h e s e t u r t l e s cons i s t ed of va r ious combinations of p e l l e t e d dry feed i n g r e d i e n t s ( i . e . meal, soybean meal, and co rn ) , as w e l l as f r e s h f rozen squid and f i s h . These d a t a show t h a t a t least 19 mo were requ i r ed t o reach a s i z e of 35 cm, a f t e r which t h e rate of growth dec l ined cons iderably . Bourke e t a l . (131) p re sen t growth d a t a i n t h e form of weight i n c r e a s e s f o r 46 Hawaiian CheZonia hatched i n c a p t i v i t y and r a i s e d t o 6 mo of age.

f i s h meal, meat and bone

The growth rates of Hawaiian Chelonia a f t e r matura t ion have been documented through t h e remeasurement of a d u l t s dur ing each breeding season a t French F r i g a t e Shoals . Inc reases f o r 1 7 females and 1 male ranged from 0.01 t o 0.12 cm/mo (24-75 mo), w i th a mean of 0.04 cm/mo (Table 12 ) . t h e rates of growth f o r a t least some Hawaiian Chelonia d e c l i n e s i g n i f i c a n t l y once ma tu r i ty has been reached. A cons ide rab le l e n g t h of t ime would t h e r e f o r e be requi red f o r a t u r t l e t h a t matured a t a s m a l l s i z e (81 cm) t o grow t o a l a r g e s i z e (see s e c t i o n s 3.12, 3.31 and H i r t h 265). The d i f f e r e n c e s i n growth rates recorded f o r m a t u r e t u r t l e s (0.01-0.12 cm/mo) may a l s o be a func t ion of t h e sources and abundance of food a t t h e r e s i d e n t area ( s e c t i o n 3.16). I f t h i s is indeed t h e case, mature t u r t l e s t h a t show t h e most r ap id growth could be i d e n t i - f i e d as having remigrated from areas i n t h e main i s l a n d s . Table 12 provide some suppor t f o r t h i s p ropos i t i on i n t h a t female 5016 exh ib i t ed one of t h e most r a p i d growth rates (0.09 cm/mo) and is known t o have o r i g i n a t e d from t h e Kau Di s t r i c t on t h e i s l a n d of H a w a i i ( s e c t i o n 3.52).

This sugges t s t h a t

Data presented i n

a

c

c

P

c

29

Table 1 3 p resen t s growth rates f o r both immature and a d u l t Hawaiian Chelonia captured from the wild and he ld f o r extended per iods i n c a p t i v i t y a t Sea L i f e Park on Oahu. .

and f i s h . The d i e t s of t hese t u r t l e s cons i s t ed of f r e s h f rozen squid

3.5 Pos tha tch l ing Movements

3 .51 Dispersal and developmental migra t ions

The d i s p e r s a l of green t u r t l e ha t ch l ings from French F r i g a t e Shoals i n t o t h e p e l a g i c environment t akes p l ace by s u r f a c e c u r r e n t s and vigorous swimming ( s e c t i o n 2.22). Af t e r an undetermined per iod of t i m e i n t h e open ocean, dur ing which unknown r o u t e s are followed, j u v e n i l e s of approximately 35 cm a r r i v e a t c o a s t a l areas throughout t h e Hawaiian Archipelago. The recru i tment of t h e s e t u r t l e s a t i s l a n d s t o t h e northwest of French F r i g a t e Shoals could be a d i r e c t r e s u l t of nor thwes ter ly c u r r e n t s t h a t p r e v a i l f o r ha t ch l ings e n t e r i n g t h e water during t h e month of J u l y (Figure 4) . I n a d d i t i o n , t h e low l e v e l of breeding t h a t occurs p r i n c i p a l l y a t Laysan and L i s i a n s k i I s l a n d s and P e a r l and Hemes Reef may be a f u r t h e r source of j u v e n i l e s i n t h i s segment of t h e Archipelago. The recru i tment of j u v e n i l e s i n t h e main i s l a n d s is more d i f f i c u l t t o t heo r i ze . One p o s s i b i l i t y i s t h a t dur ing t h e peak ha tch ing month of August, p r e v a i l i n g s u r f a c e c u r r e n t s t r a n s p o r t ha t ch l ings i n a n o r t h e r l y d i r e c t i o n f o r a 2-mo per iod t o approximately l a t . 28'40" (Figure 4). southwest a long wi th t h e same c u r r e n t system, t h e t u r t l e s s w i m t o t h e v i c i n i t y of la t . 30' t o 31'N where win te r s u r f a c e c u r r e n t s of 19* t o 20'C t rave1 ,eas tward . Over an ensuing per iod of 6 mo o r longer , t h e t u r t l e s could be c a r r i e d by a gyre t h a t u l t i m a t e l y d e l i v e r s them t o t h e main i s l a n d s . a f a r l a r g e r c i r c u l a r t r a n s p o r t system is involved i n which ha tch l ings are c a r r i e d w e l l t o t h e w e s t of t h e Hawaiian Archipelago and around a vast area of t h e North P a c i f i c back t o t h e main i s l a n d s . Of course i t is a l s o conceivable t h a t some of t h e t u r t l e s l eav ing French F r i g a t e Shoals are no t c a r r i e d by t h e c u r r e n t s , bu t r a t h e r s w i m a g a i n s t t h e c u r r e n t s on a course d i r e c t l y toward t h e main i s l ands . Never the less , whatever oceanic rou te s of d i s p e r s a l are involved, t h e r e is l i t t l e . doubt t h a t j u v e n i l e s <35cm are r e s i d i n g somewhere o u t s i d e of t h e c o a s t a l areas where larger t u r t l e s feed and rest. of d i r e c t s i g h t i n g s , as w e l l as by t h e absence of j u v e n i l e s <35cm i n t h e stomachs of t i g e r sharks . French F r i g a t e Shoals (Balazs 57) w a s undoubtedly a rare occurrence involv ing a s t r a y ind iv idua l .

A t t h i s p o i n t i n s t e a d of t u rn ing t o t h e

Another p o s s i b i l i t y is t h a t

This is supported by t h e dea r th

The s i n g l e observa t ion of a 20 t o 25 cm j u v e n i l e repor ted a t

Juven i l e s measuring 35 t o 40 cm t h a t are be l ieved t o be r ecen t arrivals t o c o a s t a l areas have, on a few occasions, been observed by t h e au thor a t Kure, Midway, Oahu, and along t h e Kau D i s t r i c t (Wright e t a l . 557). These new r e c r u i t s were d i s c e r n i b l e by an absence of e p i z o i c s and s u p e r f i c i a l s c r a t c h e s , and by t h e presence of t h i n t r a n s l u c e n t edges t o t h e per iphery of t h e marginal laminae and te rmina l s c a l e s on t h e f l i p p e r s . I n a d d i t i o n , t he c u t t i n g edges of t h e lower beak had more pronounced s e r r a t i o n s than o t h e r j u v e n i l e s of t h e same o r a s l i g h t l y l a r g e r s i z e . a f t e r e s t a b l i s h i n g r e s t i n g and herbivorous forag ing h a b i t s i n a c o a s t a l area.

A l l of t hese c h a r a c t e r i s t i c s could be expected t o disappear r a p i d l y

Most evidence accumulated t o d a t e i n d i c a t e s t h a t a f t e r l eav ing the p e l a g i c environment, Hawaiian Chelonia r e s i d e i n t h e same gene ra l c o a s t a l area for

30

exttwded pe r iods , poss ib ly throughout t h e i r e n t i r e l i f e t i m e except f o r remigra- t i o n s f o r reproduct ion . This extended res idency concept is at least p a r t i a l l y supportcd by t h e f a c t t h a t a l l s i z e s of t u r t l e s from 35 c m j u v e n i l e s t o mature a d u l t s Some s p e c i f i c forag ing sites have been i d e n t i f i e d t h a t can only be used by j u v e n i l e s ( s e c t i o n 3.42), however h a b i t a t employed by l a r g e r t u r t l e s i s usua l ly only a s h o r t d i s t a n c e away. The recovery of tagged immature t u r t l e s a f t e r pe r iods ranging up t o 37 mo has a l s o provided evidence f o r extended residency. With t h e except ion of two t u r t l e s , a l l A-

r ecap tu res (146 out of 524--Table 4) have been made i n t h e same fo rag ing and r e s t i n g areas where i n i t i a l t agging occurred . A t French F r i g a t e Shoals , r ecove r i e s have shown t h a t no movement t akes p l a c e between sites sepa ra t ed by as s h o r t a d i s t a n c e as 8 km. A t Kure a tagged t u r t l e w a s found r e s t i n g under t h e same c o r a l l edge where it had been captured 1 3 mo earlier (Balazs 81). r ecove r i e s t h a t i n d i c a t e d movement of any d i s t a n c e involved t h e weak 38 c m juve- n i l e tagged a t Midway and found downwind a t Wake I s l and ( s e c t i o n 2 .1>, and a 40 c m j u v e n i l e a l s o tagged a t Midway t h a t w a s repor ted t o t h e au thor 7 mo la ter as having been recovered and r e l eased a l i v e i n Hi lo Bay on t h e i s l a n d of H a w a i i . This l a t t e r case involves an ocean d i s t a n c e of approximately 2,300 km a g a i n s t t h e p r e v a i l i n g winds and c u r r e n t s i n t h e l a t i t u d e s of t h e Hawaiian Archipelago. Monel t a g s w e r e o r i g i n a l l y p laced on t h i s t u r t l e , only one t a g w a s found and recorded a t t h e t i m e of recovery. The p o s s i b i l i t y must t h e r e f o r e be considered t h a t t h e t a g number w a s misread due t o co r ros ion o r o t h e r causes . I f such movements can, i n f a c t , be s u b s t a n t i a t e d through a d d i t i o n a l r ecove r i e s , immature t u r t l e s r e s i d i n g i n t h e northwestern segment of t h e Hawaiian Archipelago would c o n s t i t u t e a s i g n i f i c a n t f a c t o r i n recru i tment t o t h e main i s l a n d s . Such a recru i tment system w a s t heo r i zed by t h e au thor (Balazs 57) p r i o r t o t h e accumula- t i o n of e x i s t i n g d a t a which suppor t s t h e concept of an extended res idency a t t h e c o a s t a l area en te red from t h e p e l a g i c environment. p a r t , on obse rva t ions of groups of j u v e n i l e s 35 t o 60 c m p e r i o d i c a l l y occur r ing a t French F r i g a t e Shoals . These s i g h t i n g s suggested t h a t developmental migra t ions of some n a t u r e w e r e t ak ing p l ace (Balazs 57). t h a t t h e same t u r t l e s were involved, and n o t new aggrega t ions as o r i g i n a l l y specula ted . The r e g u l a r disappearance of t hese t u r t l e s f o r several weeks and poss ib ly even months a t a t i m e has s t i l l no t been resolved. be t h a t a form of dormancy i s p e r i o d i c a l l y being undertaken a t s h e l t e r e d under- w a t e r l o c a t i o n s , C a l i f o r n i a , bu t i s p r e s e n t l y unknown f o r Hawaiian Chelonia.

are f r e q u e n t l y p re sen t a long a given c o a s t a l area.

The only two

Although two

This theory w a s based, i n

However, subsequent tagging revea led

One explana t ion would

Such behavior has been documented f o r carrinegra i n t h e Gulf of

c

I f s i g n i f i c a n t l e v e l s of recru i tment t o t h e main i s l a n d s occur through developmental migra t ions from t h e northwestern segment of t h e Hawaiian Archipelago, then r e v i s i o n s would be necessary i n t h e p ro jec t ed number of y e a r s needed f o r Hawaiian green t u r t l e s t o reach ma tu r i ty ( s e c t i o n 3.12 and Table 4 ) .

Since January 1973, 10 a d u l t and 31 subadul t green t u r t l e s have been re turned t o t h e wi ld a f t e r extended pe r iods i n c a p t i v i t y . of t h e f i v e t u r t l e s t h a t have been recaptured i n the main i s l a n d s are presented i n Table 14 and Figure 8.

The d i s p e r s a l p a t t e r n s

An experimental model f o r t h e l i f e h i s t o r y and h a b i t a t s of Hawaiian Chelonia i s presented i n F igure 9.

c

31

3.52 Remigrations

Adult Hawaiian CheZonia periodically travel between their resident foraging areas and French Frigate Shoals for reproduction (sections 2.21, 3.16). These remigrations have been documented for both males and females through 52 long distance tag receoveries, 31 of which involved French Frigate Shoals and the main islands, and 21 that involved French Frigate Shoals and the northwestern locations of Laysan and Lisianski Islands and Pearl and Hermes Reef (Table 15, Figure 10). Details of the 16 female and 4 male tag recoveries that have been made since June 1973 are presented in Table 16.

The recoveries in the main islands, of turtles that were tagged while at French Frigate Shoals, mostly represent mortalities resulting from direct capture by fishermen. However, recoveries at Laysan and Lisianski Islands and Pearl and Hemes Reef were of basking turtles that remained viable members of the population and, in several cases, were recovered again at later dates. Records of remigrations in the Hawaiian Archipelago are unique among marine turtle populations due to the two-way tagging opportunities afforded by the basking behavior, and by the research emphasis placed on turtles in their resident foraging areas. These factors have made it possible to document movements from the resident areas back to the breeding grounds, a missing segment in all one-way tagging programs where it is only feasible to tag nesting turtles. movements by adult males is, of course, in itself a rare research occurrence. The results of these various opportunities with Hawaiian CheZonia are dramatically illustrated by male 1060 in Table 16 which was tagged at Pearl and Hermes Reef in 1964, recovered at East Island, French Frigate Shoals in 1976, and recorded back at Pearl Hermes Reef 1 yr later in 1977.

The documentation of long-distance

Tag recoveries demonstrate that the breeding assemblage at French Frigate Shoals is comprised of adults that remigrate from widely separated resident areas. The longest voyages thus far recorded are from French Frigate Shoals to Hilo Bay (tag 2229, Table 16), and from the Kau District to French Frigate Shoals (tag 5016, Table 16), both of which represent one-way ocean distances of approximately 1,100 km. Fourteen other recoveries (4 males, 10 females) have been made between Pearl and Hemes Reef and French Frigate Shoals, a distance of 1,050 km. Mating is therefore taking place between some males and females that live in areas separated by as many as 2,150 km. A minimum rate of travel of 23 km/day during these remigrations has been computed from a relatively short-term recovery between French Frigate Shoals and Kauai (tag 853, Table 16). arched (less streamline) in Hawaiian CheZonia as suspected (section l.l), then a greater expenditure of energy, and perhaps even slower swimming speeds, would be expected to occur in comparison to Atlantic and Caribbean Chelonia populations.

If the contour of the adult female's carapace is indeed more highly

It is significant to note that adults from the main islands, as well as from Laysan and Lisianski Islands and Pearl and Hermes Reef have only been recorded traveling from their respective segments of the Hawaiian Archipelago as far as French Frigate Shoals, and not beyond. that Hawaiian CheZonia return to their same resident foraging area at the end of each breeding season. widely recognized for CheZonia populations, far less attention has been given to

This further supports the belief

Although a homing instinct for the breeding grounds is

32

t h e pronounced f i x a t i o n which a l s o appears t o e x i s t f o r t he r e s i d e n t area (Balazs 57; H i r t h 266). weak j u v e n i l e found a t Wake I s l and , none of t h e t u r t l e s tagged i n t h e Hawaiian Archipelago have been recovered a t any o t h e r l o c a t i o n i n t h e P a c i f i c .

It i s a l s o s i g n i f i c a n t t o no te t h a t , except f o r t h e c

It i s unknown i f t h e low l e v e l of breeding t h a t occurs a t i s l a n d s t o t h e northwest of French F r i g a t e Shoals involves nonmigratory r e s i d e n t s of t hese areas, o r a d u l t s t h a t have remigrated from o the r l o c a t i o n s i n t h e Hawaiian Archipelago. g e n e t i c remnants of what a t one t i m e may have been f a r l a r g e r breeding colonies . Such reproduct ive voyages could conceivably be t ak ing p l ace from r e s i d e n t areas anywhere i n t h e Hawaiian Archipelago. Shoals , where tagged a d u l t females known t o l i v e i n t h e area have never been recorded n e s t i n g a t t h a t l o c a t i o n . Although migra t ions from French F r i g a t e Shoals t o o t h e r areas f o r breeding purposes would seem t o be t o t a l l y unnecessary, t h i s form of cyc le i s t ak ing p l ace when a d u l t s t r a v e l from t h e breeding areas of Laysan and L i s i a n s k i I s l a n d s and P e a r l and Hermes Reef t o reproduce a t French F r i g a t e Shoals. Another p o s s i b l e explana t ion t o account f o r n e s t i n g a t i s l a n d s t o t h e northwest of French F r i g a t e Shoals i s t h a t some of t h e females are re tu rn ing t o their r e s i d e n t area from French F r i g a t e Shoals w i t h an un la id c l u t c h o r p a r t i a l c l u t c h of eggs. Kauai has been repor ted by a fisherman ( tag 853, Table 1 6 ) , bu t d e t a i l s could not be obta ined and no r ecen t n e s t i n g s have been recorded f o r t h i s i s l a n d .

The occurrence of remigra t ions would suggest t h a t t h e s e t u r t l e s are

This could even inc lude French F r i g a t e

0

The presence of eggs i n a female s h o r t l y a f t e r h e r r e t u r n t o

It i s unknown what mot iva t ing f a c t o r s may have been involved i n t h e 440 km movement of an a d u l t female tagged while basking a t P e a r l and H e r m e s Reef i n March 1965, and recovered whi le basking a t Laysan I s l a n d i n December 1967 (Kr id le r 3 2 4 , Figure 10 ) .

3.53 Navigation and o r i e n t a t i o n

The p e r i o d i c migra t ions c a r r i e d out i n CheZonia popula t ions between r e s i d e n t forag ing areas and d i s t a n t breeding grounds r e q u i r e some form of nav iga t iona l a b i l i t y . be i n use f o r long d i s t a n c e movements involv ing small oceanic i s l a n d s , such as from t h e coas t of B r a z i l 2,000 km t o Ascension I s l and (Carr 150, 151). H i r t h (265) has suggested t h a t naviga t ion by green t u r t l e s i s l i k e l y t o be a composite process employing d i f f e r e n t senses t h a t are based on a m u l t i p l i c i t y of cues. The var ious mechanisms thus f a r suggested t o exp la in t h i s a b i l i t y have a l s o been summarized by H i r t h (265) and inc lude b i coord ina te c e l e s t i a l naviga t ion , l i g h t - compass sense , o l f a c t o r y cues, percept ion of C o r i o l i s f o r c e , magnetic sense , i n e r t i a l guidance, sonar sense , and s u b t l e oceanographic cues such as c u r r e n t s and wave p a t t e r n s . unknown. long d i s t a n c e s of open ocean us ing boa t s , a i r p l a n e s , o r land-based monitoring techniques. taken, thereby a i d i n g g r e a t l y i n determining what cues are being employed. r ecen t a v a i l a b i l i t y of ea r th -o rb i t i ng sa te l l i t es f o r w i l d l i f e te lemet ry o f f e r s cons iderable p o t e n t i a l f o r answering t h i s nav iga t iona l ques t ion f o r a green t u r t l e populat ion.

Furthermore, a h ighly r e f i n e d guidance system would have t o

However, t h e a c t u a l system o r systems u t i l i z e d remain t o t a l l y This is mostly due t o t h e unresolved problems of t r ack ing a t u r t l e over

The s u c c e s s f u l t r ack ing of a t u r t l e would r e v e a l t h e a c t u a l rou te s The

ri

c

c

33

A b a s i c assumption i n t h e migra t ions and naviga t ion of Chelonia i s that: the a d u l t s r e t u r n t o breed a t t h e same l o c a t i o n where they o r i g i n a t e d as hatch- l i n g s . i n d i r e c t evidence, is thought t o r e s u l t from some form of "imprinting" t h a t t akes p l ace when t h e ha t ch l ings emerge from t h e n e s t o r depa r t through ad jacen t waters. However, no th ing is known about t h i s process , i f indeed one e x i s t s . I f a d u l t s remigra te i n groups t o improve t h e i r accuracy of o r i e n t a t i o n , as has been suggested (Hir th 265), t h i s could a l s o s e r v e t h e purpose of provid ing a l e a r n i n g mechanism f o r newly matured t u r t l e s t o f i n d t h e i r way back t o t h e breeding grounds.

This f i x a t i o n on t h e n a t a l beach, which is supported by cons ' iderable

For Hawaiian CheZonia, t h e least complex system f o r f i n d i n g French F r i g a t e Shoals when remigra t ing from r e s i d e n t forag ing areas would appear t o be one i n which t h e va r ious i s l a n d s and submerged areas wi th no emergent land are followed and used as n a v i g a t i o n a l gu idepos ts . i s l a n d s could fo l low t h e c o a s t l i n e s i n a nor thwes ter ly d i r e c t i o n , c r o s s i n g t h e r e l a t i v e l y narrow i n t e r i s l a n d channels of 15 t o 120 km u n t i l reaching Kauai o r Niihau, The next i s l a n d g u i d e p o s t could then be Nihoa ( a r e a 63 ha , e l e v a t i o n 273 m) l oca t ed 225 km t o the northwest , w i t h a 19 fathom (35 m) shoa l occu r r ing a t 125 km a long t h i s course. The d i s t a n c e from Nihoa t o Necker (area 1 7 h a , e leva- t i o n 85 m) is 295 km, bu t t h r e e s m a l l s h o a l s are loca ted a t about t h e midpoint between t h e two i s l a n d s . I n a d d i t i o n , Necker i t s e l f i s s i t u a t e d on a l a r g e 28 by 74 km bank w i t h depths ranging from 8 t o 23 fathoms (15-42 m). One of t h e few repor t ed s i g h t i n g s i n t h e Hawaiian Archipelago of a green t u r t l e away from land w a s made i n March 1979 a long t h e southern edge of t h e Necker Bank. This involved an a d u l t , b u t i t could no t be determined i f t h e t u r t l e w a s i n t r a n s i t o r a r e s i d e n t of t h e area (G. L. N a f t e l , pe r sona l communication). From Necker, t h e f i n a l goa l of French F r i g a t e Shoals f o r a remigra t ing t u r t l e i s 155 km d i r e c t l y t o t h e w e s t .

T u r t l e s r e s i d i n g i n t h e main

T u r t l e s a t r e s i d e n t forag ing areas t o t h e northwest of French F r i g a t e Shoals would have t o f o l l o w the Hawatian Archipelago i n a s o u t h e a s t e r l y d i r e c t i o n dur ing remigra t ions , submerged area, o r two s e p a r a t e submerged areas, would be 155 km, t h e d i s t a n c e between Raita Bank and Gardner P innac les (area 1 h a , e l e v a t i o n 45 m) . Between P e a r l and Hermes Reef and French F r i g a t e Shoals , t h e r e are 1 4 submerged areas and t h r e e i s l a n d s wi th a s s o c i a t e d banks. The l a r g e s t of t h e s e is Gardner P innac les bank which is 37 by 93 km, with depths of 10 t o 25 fathoms (18-46 m ) .

Along t h i s r o u t e , t h e maximum span between an i s l a n d and a

T u r t l e s t r a v e l i n g from r e s i d e n t forag ing areas i n t h e main i s l a n d s t o French F r i g a t e Shoals would be moving gene ra l ly wi th t h e p r e v a i l i n g c u r r e n t s and t h e r e f o r e would n o t be a b l e t o d i r e c t l y u t i l i z e chemical cues o r plumes t h a t may emanate from t h e va r ious i s l a n d s and submerged areas. I n d i r e c t u se could r e s u l t , however, by sens ing t h e p o s i t i o n of t h e s e areas once they have been passed and are upstream. T u r t l e s t r a v e l i n g from t h e northwestern r e s i d e n t fo rag ing areas t o French F r i g a t e Shoals would be moving gene ra l ly a g a i n s t t h e p r e v a i l i n g c u r r e n t s and t h e r e f o r e could conceivably make d i r e c t use of chemical cues . During t h e r e t u r n t r i p from French F r i g a t e Shoals back t o t h e r e s i d e n t areas, t u r t l e s from each segment of t h e Hawaiian Archipelago would be confronted wi th an oppos i t e set of c u r r e n t cond i t ions from t h e i r ear l ier voyage.

I n t h e l a t i t u d e s of t h e main i s l a n d s , t h e sun i s a t t h e z e n i t h dur ing la te May as i t t r a v e l s t o i t s f a r t h e s t po in t no r th , and aga in i n l a t e J u l y as i t

34

returns south. Turtles departing from French Frigate Shoals to return to the main islands during July could set a course each morning directly into the rising sun.

therefore

Another navigational aid that could possibly be used by Hawaiian Chelonia is the subtle color differences that are present in clouds due to reflec- tions from shallower ocean depths where banks occur. However, the use of visual cues may not be essential considering that two males found with extensive fibro- papillomas on their eyes were believed to have been recent migrants to French Frigate Shoals (section 3.34) .

The evolution of reproductive migrations by Chelonia to midocean' Ascension Island is thought to be an adaptation to the gradual separation of South America and West Africa resulting from seafloor spreading (Carr 150, 151; Hirth 265). Similar geophysical mechanisms may also help to account for evolution of the Hawaiian Chelonia pattern with respect to voyages occurring between the geologically older northwestern areas of the Hawaiian Archipelago and the more recently formed French Frigate Shoals. This migratory component appears to be equivalent to the Ascension Island pattern. and disappeared, ancestors of Hawaiian Chelonia may also have gradually extended their traveling distances between resident foraging areas and aggregate breeding grounds. This theory does not, however, provide an explanation for reproductive migrations occurring from the main islands, which are known to be geologically younger than French Frigate Shoals.

As new islands progressively appeared

While at French Frigate Shoals, nesting turtles demonstrate a distinct preference for a certain island and, to a lesser extent, show some favoritism for certain areas on that island (section 3.15). The retention of this island fixity between breeding seasons in the 21 remigrations of females thus far documented (section 3.16) suggests the presence of a refined memory system for short-range orientation. Direct vision as well as olfactory cues are undoubtedly major components of this discrimination process. The orientation of turtles underwater at night prior to emergence for nesting could be expected to involve some of the same sensory cues used by turtles foraging at night in the resident areas (section 3.42).

c

c

3.6 Basking

Hawaiian green turtles exhibit the behavioral trait of coming ashore to bask or rest at certain undisturbed sites in the M I . This includes adults of both sexes and, to a lesser extent, immature turtles of all sizes larger than 35 cm. The land basking habit is rare among marine turtles, with Hawaiian and Galapagos CheZonia being the only two populations in which it has been well documented in the historical literature. However, basking in the Galapagos Archipelago has not been observed for a number of years and longer be a characteristic of the population. There is no historical evidence of Hawaiian Chelonia basking on any of the main islands. made at Johnston Atoll suggests that on rare occasions green turtles may bask at this location (Amerson and Shelton 31; Balazs 78). At the Wellesley Isles in Australia's Gulf of Carpentaria, female green turtles alone are known to come ashore during the breeding season to avoid males.

therefore may no

However, an observation

The 27 December 1777 log of

a

35

I U

L

a.

t h e ResoZution states t h a t a t Christmas I s l a n d green t u r t l e s were turned on t h e i r backs wh i l e "asleep," and t h a t t h i s w a s a common method of cap tu re (Beaglehole 105) . t u r t l e s has been observed a t Lizard I s l a n d on t h e Great Barrier Reef (W. Anderson and J. L e i s , personal communications) and a t North West Cape, Western A u s t r a l i a (J. Bradley, personal communication).

I n a d d i t i o n , p rev ious ly unreported basking behavior by green

The presence of basking t u r t l e s i n t h e NWHI w a s noted by many of t h e e a r l y v i s i t o r s dur ing t h e 1800's (Brooks, 135, 136; F a r r e l l 220; Horne l l 271; Lis iansky 335; Morre l l 360; Munro 365-370; Pa ty 402; Read 429; Rothschi ld 435; Walker 531; Anonymous 564, 566). However, widespread knowledge of t h i s behavior d i d n o t occur u n t i l p u b l i c a t i o n of an account of t h e 1923 Tanager Expedi t ion i n a 1925 i s s u e of National Geographic Magazine (Wetmore 542). photograph from t h i s a r t i c l e of a basking t u r t l e a t L i s i a n s k i I s l a n d w a s republ i shed s h o r t l y t h e r e a f t e r i n t h e New York Zoological Society Bul le t in , a long wi th f u r t h e r d e s c r i p t i v e informat ion by D r . Alexander Wetmore (Mellen 353). by Hawaiian Chelonia as a r a r i t y among marine t u r t l e s (Carr 150, 152; Grant 238; Loveridge 336; Pope 415). During aerial censuses of Hawaiian monk seals i n 1957 and 1958, Kenyon and Rice (288) r e g u l a r l y observed t u r t l e s basking along t h e s h o r e l i n e s of t h e NWHI (see a l s o Parsons 400). From 1950 t o 1973, bask ing t u r t l e s were a t va r ious t i m e s recorded and counted i n conjunct ion wi th b i o l o g i c a l surveys by t h e P a c i f i c Ocean F i s h e r i e s I n v e s t i g a t i o n (386-398), t h e HDFG (Brock 133; K r a m e r 308, 309; Kramer and Beardley 310; Walker 532, 533; Woodside 554; Woodside and K r a m e r 555) , t h e P a c i f i c Ocean B io log ica l Survey Program ( see Subjec t Index) , and t h e USFWS (Kr id l e r 311-327; Olsen 380-385). Summaries of t h e s e obse rva t ions and censuses have been publ ished by Amerson (30); Amerson e t a l . (32); Clapp and Wirtz (171); Clapp and K r i d l e r (172); Clapp e t a l . (173); Ely and Clapp (211); and Woodward (556). During r ecen t y e a r s , photographs of basking Hawaiian Chelonia have appeared along wi th art icles and r e p o r t s by Altonn ( 6 ) ; Balazs (55 wi th Monachus, 57, 6 1 w i t h ocypode, 62, 78 ) ; E l i o t (208) ; H i r t h (265) ; Lipman (333) ; Moake (358 wi th Chelonibia) ; and Anonymous (642, 657, 667).

A

Severa l o t h e r au tho r s subsequent ly mentioned t h e occurrence of basking

Basking i n t h e NWHI t akes p l a c e on ca lcareous sand beaches of vary ing p a r t i c l e s i z e and composition loca ted a t French F r i g a t e Shoals , Laysan and L i s i a n s k i I s l a n d s , P e a r l and H e r m e s Reef , Kure and, on a s i n g l e repor ted occasion, Midway (Balazs 78). A t Laysan (P, and B. Johnson, pe r sona l communica- t i o n ) and L i s i a n s k i I s l a n d s , some t u r t l e s a l s o emerge on ca lcareous beachrock s l a b s . A t Nihoa, two t u r t l e s have been seen on a s i n g l e occasion basking on a rock a t t h e base of t h e i s l a n d ' s northwest c l i f f (Clapp e t a l . 173 ; K r i d l e r 326). A t Necke? I s l a n d , basking t u r t l e s r e g u l a r l y use a small s lop ing rock ledge measuring approximately 4 by 5 m. wes t e r ly ocean s w e l l s , some t u r t l e s bask a t an a l t e r n a t e p ro tec t ed s h o r e l i n e area a s h o r t d i s t a n c e away t h a t is comprised of smooth waterworn boulders . The r e s i d e n t aggrega t ion of green t u r t l e s t h a t basks and f eeds a t Necker I s l a n d is undoubtedly one of t h e most i n t e r e s t i n g , unusual, and f r a g i l e components of t h e Hawaiian Chelonia popula t ion (Balazs 72).

During pe r iods of l a r g e wes te r ly and north-

Although basking occurs p r i n c i p a l l y i n t h e daytime between approximately 1000 and 1830 h ( v a r i a b l e wi th season) , t u r t l e s a t Necker I s l a n d have

36

been found to also commonly emerge at night. At times this nocturnal emergence appears to be correlated with the setting of a bright moon (Balazs 72). The author has also observed small numbers of turtles resting along the shoreline at night at French Frigate Shoals and Laysan Island, but not at Lisianski Island. Amerman (24), Kridler (319), and Sibley (444, 445) frequently found turtles resting at night on Southeast Island at Pearl and Hemes Reef. (32) attributed nighttime emergence to the disturbing activities of research personnel during the daytime. However, this seems unlikely in view of the fact that, for the September 1967 visit to Southeast Island by USFWS personnel, Kridler (319) stated "When the island was first landed upon, there were no turtles present on the beach. At dusk the first day, however, one after another hauled up on the beach on the lagoon side near camp until 19 were present."

Amerson e t al.

With the exception of La Perouse Pinnacle, basking occurs on all of the islands at French Frigate Shoals (Figure 2 ) , as well as on several unnamed seasonally occurring sandbars. northeastern shore of Whale-Skate Island are the most heavily utilized during all months of the year by the resident aggregation. At East Island, basking tends to coincide more with the breeding season (April-September). Turtles use the entire coastline, but higher concentrations consistently occur at the southeastern end (Figure 6, areas 8-9). A s with nesting, turtles display a fixity for basking on a particular island and, to a lesser extent, for regularly basking at the same site on that island. turtles are found throughout French Frigate Shoals during late May and June due to the presence of the migratory breeding assemblage. The incidence of basking then declines as the breeding season progresses (Figure 11). The greatest number of basking turtles seen by the author at any one time was 52 (18 males, 18 females, and 16 unknown) along the northeastern shore of Whale-Skate on 12 June 1978 at 1630 h. It is of interest to note that turtles have never been observed entering into nesting activity directly from a basking position.

The northern shore of Trig Island and the

The greatest numbers of basking

Some adult Hawaiian Chelonia apparently do not bask. During each breeding season at French Frigate Shoals since 1973, approximately 46% (range 32.7%-56.2%) of the turtles identified with painted numbers on their carapace while nesting were subsequently never seen basking. Some of these females are undoubtedly among the 40% that only lay a single clutch of eggs in a breeding season (section 3.15). present to bask because they have either departed on the return trip to their resident area, or have been eliminated by tiger sharks. Females that attempt to nest but no not lay eggs and are subsequently not seen again (section 3.15) also constitute some of the turtles that are not seen basking. It is possible, however, that turtles in both of these categories do bask, but only during the time prior to first coming ashore to nest or make nesting attempts.

Consequently, after nesting the turtles may not be

c

c

No differences were found between the lengths of the internesting intervals of basking and nonbasking turtles observed during the 1974 and 1975 breeding seasons. the in v i v o development of eggs for oviposition as might be expected.

This indicates that basking does not significantly hasten

a

37

9

It i s important t o no te t h a t a t least some members of t h e seasonal breeding aggrega t ion t h a t are from r e s i d e n t areas i n t h e main i s l a n d s do, i n f a c t , bask whi le they are a t French F r i g a t e Shoals. i n t h a t t h e 54% recorded basking each season could conceivably c o n s i s t of only those females from Laysan basking is a r e g u l a r occurrence. Both males and females observed basking a t French F r i g a t e Shoals have been among t h e tagged t u r t l e s recovered i n t h e main i s l a n d s (Table 15) . I n a d d i t i o n , basking has been d isp layed a t French F r i g a t e Shoals by t h r e e t u r t l e s t h a t were re turned t o t h e wild a f t e r extended pe r iods i n c a p t i v e f a c i l i t i e s where emergence f o r basking w a s no t p o s s i b l e (Table 16 ) .

This i s a s ign i ' f i can t po in t

and L i s i a n s k i I s l a n d s and Pearl and Hermes Reef where

I n v e s t i g a t i o n s of t h e thermal ecology of basking Hawaiian Chelonia are c u r r e n t l y being conducted by Whittow and Balazs (549). t o d a t e have shown t h a t t h e s u r f a c e temperature of t h e carapace measured with a radiometer can a t t a i n va lues as g r e a t as 42°C. t a t i o n p resen t i n most a d u l t Hawaiian CheZonia undoubtedly c o n t r i b u t e t o h igh s u r f a c e temperatures during per iods of i n t e n s e s o l a r r a d i a t i o n . The g r e a t e s t i n t e r n a l body temperature recorded through t h e c loaca w a s 31.3"C a t a t i m e when the ambient seawater temperature w a s 26.3"C. some t u r t l e s open t h e i r mouth t o a wide p o s i t i o n and r e g u r g i t a t e small amounts of l i q u i d ( see Anonymous 670 f o r photograph). This is probably a r e s u l t of compres- s i o n of t h e stomach due t o t h e body weight p re s s ing on t h e p l a s t r o n whi le on land. T u r t l e s e x h i b i t very l i t t l e a c t i v i t y whi le basking except f o r occas iona l ly f l i p p i n g sand on t h e i r carapace f o r thermoregulat ion ( see Tinker 492 f o r photograph) . They do n o t , however, seem t o o r i e n t t h e i r p o s i t i o n i n r e l a t i o n t o t h e sun. I n males, t h e t a i l w i l l f r equen t ly be cu r l ed c l o s e t o t h e body r a t h e r than f u l l y extended. The l eng th of time spent basking appears t o be i n v e r s e l y r e l a t e d t o t h e black-globe temperature. found t o c o n s i s t of breath-holds averaging 3 . 6 min, followed by a s i n g l e shal low brea th .

Resu l t s of t h i s work

The l a r g e areas of b l ack pigmen-

Shor t ly a f t e r coming ashore t o bask,

Respi ra t ion p a t t e r n s whi le basking have been

Basking i n f reshwater t u r t l e s (and o t h e r poiki lotherms) i s p r i n c i p a l l y a behav io ra l s t r a t e g y t o raise body temperature and a c c e l e r a t e metabol ic processes such as d i g e s t i o n and growth. Other mot iva t ing f a c t o r s o r b e n e f i t s t h a t have been suggested inc lude s y n t h e s i s of Vitamin D from s k i n s t e r o l s (P r i t cha rd and Green- hood 419), removal of e p i z o i c s through dry ing , and s o c i a l i n t e r a c t i o n (Boyer 132). These f a c t o r s could a l s o be a p p l i c a b l e t o basking by Hawaiian green t u r t l e s .

With r e spec t t o s o c i a l i z a t i o n (Hir th 265) , cons iderable gregar ious behavior is d isp layed while basking a t French F r i g a t e Shoals , wi th t u r t l e s o f t e n i n d i r e c t con tac t o r even p a r t i a l l y on top of one another (Balazs 57). Thie is, of course, c o n s i s t e n t w i th t h e o t h e r l i f e a c t i v i t i e s of Chelonia such as feeding toge the r , breeding toge the r i n l a r g e aggrega t ions , and probably even t r a v e l i n g toge the r a c r o s s t h e open ocean. Hawaiian monk seals sometimes show gregar ious tendencies toward basking t u r t l e s , both a t French F r i g a t e Shoals and a t Necker. This has involved seals of a l l s i z e s from young pups t o a d u l t s . Mothers w i th newborn pups may, however, make aggres s ive ges tu re s and v o c a l i z a t i o n s a t t u r t l e s emerging t o bask i n t h e immediate s h o r e l i n e area.

One of t h e advantages t o Hawaiian Chelonia obta ined from basking is t h e reduct ion i n exposure t o p reda t ion by t i g e r sharks . even be t h e mot iva t ing f a c t o r f o r basking ( s e c t i o n 3 . 3 3 ) .

I n some cases, t h i s could Emergence t o land a t

38

n i g h t may be e s p e c i a l l y advantageous, cons ider ing t h a t t i g e r sha rks are be l i eved t o be p r i n c i p a l l y n o c t u r n a l p reda to r s . could, however, a l s o be due t o a s c a r c i t y of accep tab le r e s t i n g site's loca ted underwater. A f u r t h e r advantage t o r e s t i n g on land would be t h e conserva t ion of energy by not having t o p e r i o d i c a l l y s w i m t o t h e s u r f a c e f o r r e s p i r a t i o n .

Res t ing along t h e s h o r e l i n e a t n i g h t

Hawaiian Chelonia w i l l bask under c a p t i v e cond i t ions i f they are provided w i t h a s l o p i n g area s u i t a b l e f o r emergence (Balazs and Ross 88) . Sea L i f e Park and t h e Kahala H i l t o n Hote l on Oahu have d i s p l a y f a c i l i t i e s i n which Hawaiian green t u r t l e s r e g u l a r l y bask. However, hawksbi l l s and loggerheads i n t h e same pool a t Sea L i f e Park never e x h i b i t t h i s behavior . A subadul t green t u r t l e of Hawaiian o r i g i n a t t h e Kewalo Marine Laboratory, Un ive r s i ty of H a w a i i , a l s o r e g u l a r l y basks on a wooden ramp i n s t a l l e d i n i t s concre te tank (whittow and Balazs 549). There are no r e p o r t s i n t h e l i t e r a t u r e of green t u r t l e s from o t h e r popula t ions basking i n c a p t i v i t y . However, most f a c i l i t i e s where marine t u r t l e s are maintained do no t have s lop ing areas where emergence can occur .

Both

The basking behavior of Hawaiian Chslonia could very w e l l be an i n h e r i t e d c h a r a c t e r i s t i c t h a t evolved as a p r o t e c t i v e mechanism a g a i n s t t i g e r sha rksand as a method f o r ga in ing body h e a t a t l o c a t i o n s where seawater tempera- t u r e s were marginal . The mean temperature of 20.5"C i n February a t t h e northwestern end of t h e Hawaiian Archipelago ( s e c t i o n 3.43) could c o n s i t u t e such an environment a t t h e p r e s e n t t i m e , and perhaps c o o l e r ocean cond i t ions p reva i l ed dur ing t h e e a r l y development of t h e popula t ion . The s u r v i v a l va lue of basking, however, dec l ined s i g n i f i c a n t l y w i t h t h e f i r s t arrival of humans i n t h e NWHI i n t h e la te 1700's. p l a c e can be a t t r i b u t e d most ly t o t h e basking behavior which provides a c c e s s i b i l - i t y f o r easy h a r v e s t i n g i n t h e terrestr ia l environment.

The ex tens ive e x p l o i t a t i o n of t u r t l e s t h a t subsequent ly took

I n a d d i t i o n t o t h e unique r e sea rch o p p o r t u n i t i e s f o r tagging ( s e c t i o n 3.52) and depth record ings ( s e c t i o n 3.15), t h e basking behavior of Hawaiian Chelonia o f f e r s cons ide rab le p o t e n t i a l f o r i n v e s t i g a t i o n s of hea r ing and v i s i o n on land under n a t u r a l condi t ions . a spec t of v i s i o n , due t o t h e f a c t t h a t green t u r t l e s are be l i eved t o be myoptic when o u t of t h e water (see H i r t h 265). evo lu t iona ry development cons ider ing t h a t t h e t u r t l e ' s eyes leave t h e water each

The au tho r i s p a r t i c u l a r l y i n t e r e s t e d i n t h e

This would appear t o be an unusual

4. POPULATION

4 .1 S t r u c t u r e

4.11 Sex r a t i o

The i n a b i l i t y t o d i s t i n g u i s h immature m

Chelonia d i s p l a y cons ide rab le v i s u a l s e n s i t i v i t y wh i l e basking.

1

t i m e a b r e a t h is taken. Observat ions t o d a t e have i n d i c a t e d t h a t Hawaiian

b a s i 1 s from fern on t h of e x t e r n a l c h a r a c t e r i s t i c s is a l i m i t i n g f a c t o r i n determining n a t u r a l s e x r a t i o s i n popula t ions of Chelonia and o t h e r marine t u r t l e s . Shoals dur ing June of 1973, 1978, and 1979 on days when r e l a t i v e l y l a r g e numbers of a d u l t bask ing t u r t l e s were ashore revea led a sex r a t i o of 66% females and 34% males (range 50%-81% females , 23%-50% males), t h e pe rcen t of basking males has been found t o d e c l i n e cons iderably .

Observat ions a t French F r i g a t e

As t h e breeding season p rogres ses , Of t h e 33

P

39

basking adults tagged by Amerson (28) at French Frigate Shoals during May and June of 1967, 73% were females and 27% were males. During May of 1972, Olsen (382) tagged 52 basking adults at French Frigate Shoals, with the resulting sex ratio of 54% females and 46% males. These various counts at the breeding colony would, however, be biased if a differential exists in the levels of basking undertaken by each sex. sexually receptive females are in the surrounding waters. At the same time, females that have already laid their first clutch of eggs for the season may be more inclined to bask in order to avoid the attention of males. An additional complicating factor is the occurrence of shorter reproductive cycles in males (section 3.16). This would cause the breeding aggregation to be an unrepresenta- tive sample of the adults present in the total population, with the bias in favor of males.

For example, males might be less inclined to bask when

Observations of adults basking at other locations in the "HI have resulted in the following sex ratios: Necker Island - 71% females, 29% males; Lisianski Island - 62% females, 38% males; Pearl and Hermes Reef - 60% females, 40% males.

In the main islands, 48 adult green turtles were reported to the HDFG as being captured by fishermen for noncommercial purposes from 1974 to 1977. Approximately 52% were listed as females and 48% as males. However, it is unknown if these determinations were made on the basis of external characteristics, or an internal examination of the reproductive tract. The reports for each island are listed as follows: Hawaii - 4 females, 0 male, Maui - 3 females, 1 male; Lanai - 0 female, 1 male; Molokai - 0 female, 3 males; Kauai - 10 females, 7 males; Oahu - 8 females, 11 males. The reports on file with the HDFG for turtles commercially captured prior to the prohibition of 1974 do not contain information on sex.

The underwater sampling of green turtles with scuba along the Kau coastline from July to September of 1976 resulted in the capture of one adult female, 8 subadults, and 29 juveniles. None of the subadults showed evidence of tail enlargement (Wright e t aZ. 557). Subsequent samplings along this resident foraging area using scuba, as well as large-mesh tangle nets, have resulted in the capture of 5 more adult females, 9 subadults, and 26 juveniles, One of the subadults showed initial signs of tail enlargement (Balazs 64, Kam and Balazs 286).

The greater number of adult females that appear to exist in the Hawaiian green turtle population is consistent with the limited data on sex ratios summarized by Hirth (265) for several other CheZonia populations. However, further investigations are necessary to determine if tail enlargement among some males may be occurring after an "adult" size has been reached. mortality rates may be higher in males than in females (section 4.4).

It is also plausible that

4.12 Size composition

The size composition of Hawaiian CheZonia, based on a sample of 77 turtles captured along the Kau coastline, has been found to be 71.4% juveniles, 22.1% subadults, and 6.5% adults. These values are generally in agreement with sightings of turtles made during underwater surveys at Kau and other areas in the main islands.

Est imat ions of t h e weight composition of t u r t l e s repor ted as being commercially captured i n t h e main i s l a n d s from 1948 t o 1973 a r e presented i n Table 1 7 . These d a t a were c a l c u l a t e d from r e p o r t s submit ted each yea r t h a t l i s t e d t h e number of t u r t l e s captured a long wi th t h e t o t a l weight. The r e s u l t i n g y e a r l y average weights range from 4.5 t o 115.7 kg, wi th t h e o v e r a l l averages f o r each i s l a n d ranging from 38.8 kg f o r H a w a i i t o 50.0 kg f o r both Molokai and Oahu. The s t anda rd d e v i a t i o n s f o r t h e y e a r l y weights i n d i c a t e t h a t most of t h e t u r t l e s captured were immature, s i n c e t h e minimum weight f o r an a d u l t female is 68 kg ( s e c t i o n 3.12).

Caldwell and Caldwell (148), quoted by H i r t h (265), t heo r i zed t h a t green t u r t l e popula t ions may c o n s i s t of most ly l a r g e r t u r t l e s due t o h igh m o r t a l i t y of t h e very young, r a p i d growth of those t h a t su rv ive , and long l i f e a f t e r reaching ma tu r i ty . This i s n o t i n agreement wi th t h e f i n d i n g s t o d a t e f o r Hawaiian Chdonia, where t h e r e i s a preponderance of j u v e n i l e s t h a t grow a t a r e l a t i v e l y slow rate, and many of t h e a d u l t females appear t o be involved i n only a s i n g l e breeding season du r ing t h e course of t h e i r l i f e t i m e .

4.2 Abundance and Densi ty

Based on sys t ema t i c tagging and monitor ing dur ing each breeding season s i n c e 1973, t h e approximate number of females n e s t i n g annual ly a t French F r i g a t e Shoals has been found t o range from 94 i n 1976 t o 248 i n 1978 (F igure 12 ) . Mean annual number of females f o r t h i s 7-yr per iod i s 180. Not more than an es t imated t o t a l of 20 females are thought t o n e s t annual ly a t Laysan and L i s i a n s k i I s l a n d s and P e a r l and H e r m e s Reef.

P r i o r t o 1973, t h e breeding colony a t French F r i g a t e Shoals w a s thought t o be cons iderably l a r g e r due t o inadequate da t a . by personnel of t h e P a c i f i c Ocean B io log ica l Survey Program ranged from 650 t o 1,300 a d u l t females and males (Amerson 30) . However, t h i s w a s based on t h e sho r t - term tagging of bask ing t u r t l e s , and t h e erroneous assumption t h a t a l a r g e d a i l y exchange of new t u r t l e s cont inuously t akes p l a c e a t each i s l a n d throughout t h e breeding season. Using t h e s e va lues , Hendrickson (262), quoted by Amerson (30) , i n c o r r e c t l y specu la t ed t h a t 650 t o 1,300 d i f f e r e n t t u r t l e s would a l s o be p re sen t i n June, and twice as many would be p re sen t i n Ju ly . The t o t a l b reeding assem- b lage w a s t h e r e f o r e p laced a t between 2,600 and 5,200 t u r t l e s . Hendrickson (262) f u r t h e r q u a l i f i e d t h e s e va lues by i n d i c a t i n g , "While i t is very important t o s ta te f l a t l y t h a t t h i s estimate has l i t t l e b a s i s and is not t o b e t r u s t e d , one can a t least s a y t h a t i t does n o t appear t o c o n f l i c t v i o l e n t l y wi th any o t h e r a v a i l a b l e q u a n t i t a t i v e information."

Estimates f o r August of 1965

An es t ima t ion of t h e former s i z e of t h e breeding colony can be obta ined from an account i n June of 1891 which states t h a t a t E a s t I s l a n d , formerly known as T u r t l e I s l a n d , hundreds of t u r t l e s w e r e found basking on t h e beach, and t e n t i m e s as many were s e e n i n t h e w a t e r (Walker 531).

The d e n s i t y of t u r t l e s n e s t i n g on each of t h e i s l a n d s a t French F r i g a t e Shoals i s p r e s e n t l y a t a level where t h e d e s t r u c t i o n of p rev ious ly l a i d egg c l u t c h e s i s n e g l i g i b l e ( s e c t i o n 3.17). However, i f l a r g e r breeding aggrega t ions formerly e x i s t e d under t h e same terrestrial cond i t ions , density-dependent n e s t

c

L

rp

IC

4 1

**

d e s t r u c t i o n may have n a t u r a l l y regula ted t h e popula t ion s i z e ( see H i r t h 265). This could be a s i g n i f i c a n t f a c t o r i n t h e f u t u r e , i n t h a t density-dependent n e s t d e s t r u c t i o n can a l s o r e s u l t from reduct ions i n t h e amount of a v a i l a b l e n e s t i n g h a b i t a t a t a t i m e when the popula t ion s i z e i s cons tan t . a v a i l a b l e h a b i t a t w a s reduced i n 1942 wi th t h e ex tens ive modi f ica t ion of Tern I s l a n d and es tab l i shment of permanent f a c i l i t i e s . h a b i t a t due appa ren t ly t o n a t u r a l causes has a l s o been documented by t h e au thor a t E a s t I s l a n d . A t t h i s l o c a t i o n , t h e no r theas t e rn c o a s t l i n e has eroded a t least 20 m i n t o t h e vege ta t ion zone s i n c e 1948, wi th a l o s s of 13% of the land area,

A t French F r i g a t e Shoals ,

The reduct ion of n e s t i n g

Counts of basking t u r t l e s i n t h e NWHI made s i n c e 1950 ( s e c t i o n 3.6) provide some i n d i c a t i o n of r e l a t i v e abundance a t t h e s e r e s i d e n t forag ing areas (Table 18) . However, cons iderable v a r i a t i o n can e x i s t i n both t h e numbers ashore a t any one t i m e , and t h e d a i l y exchange of d i f f e r e n t t u r t l e s . Systematic tagging and observa t ions over a def ined per iod are L i s i a n s k i I s l a n d and P e a r l and H e r m e s Ree f , i n o rde r t o adequately assess t h e p re sen t s t a t u s of t h e s e aggrega t ions . have occurred w i t h i n h i s t o r i c a l t i m e s a t most of t h e NWHI, bu t p a r t i c u l a r l y a t Laysan (Ely and Clapp 211) and L i s i a n s k i (Clapp and Wirtz 171) I s l a n d s .

t h e r e f o r e needed, p a r t i c u l a r l y a t

Reductions i n t h e number of basking t u r t l e s

Underwater surveys wi th scuba along t h e Kau c o a s t l i n e r e s u l t e d i n t h e s i g h t i n g of approximately t h r e e t u r t l e s dur ing each hour of d iv ing t i m e . of a l a r g e mesh t a n g l e n e t measuring 3.5 by 80 m r e s u l t e d i n t h e average cap tu re of four t u r t l e s f o r each 1 4 h sample per iod (Balazs 64; Kam and B a l a z s 286).

The use

4 .3 Reproduction Rates

The b a s i c reproduct ive d a t a f o r Hawaiian Chelonia t h a t have been determined a t French F r i g a t e Shoals ( s e c t i o n s 3.15, 4.2) i nc lude 1 ) 104 eggs i n each c l u t c h , 2) 1 .8 egg c lu t ches l a i d wi th in a season by each female, 3) 70.8% emergence of ha t ch l ings from each n e s t , and 4) 180 females n e s t i n g annual ly . Using t h e s e va lues , a mean annual product ion of 23,857 ha tch l ings would occur a t French F r i g a t e Shoals . Since t h e number of n e s t i n g females over t h e 7-yr s tudy per iod ranged from 94 t o 248, t h e range i n product ion would be 12,459 t o 32,869 ha tch l ings . emergence of ha t ch l ings are assumed f o r t he 20 females us ing Laysan and L i s i a n s k i I s l a n d s and P e a r l and Hermes Reef, then 2,651 h a t c h l i n g s would r e s u l t from t h e s e l o c a t i o n s . of t h e Hawaiian Archipelago would be 26,508.

I f t h e same egg c l u t c h s i z e , number of egg c l u t c h e s , and percent

The mean annual product ton of green t u r t l e ha t ch l ings f o r a l l areas

4.4 Mor ta l i t y

The known f a c t o r s a f f e c t i n g t h e n a t u r a l m o r t a l i t y of Hawaiian CheZonia have been presented i n s e c t i o n s 2 .22 , 3 ,2 , 3.33, and 3.34. Tiger sharks are almost c e r t a i n l y t h e p r i n c i p a l cause of m o r t a l i t y fol lowing t h e t r a n s i t i o n from t h e p e l a g i c environment t o res idency i n a c o a s t a l area. Although t h e a c t u a l m o r t a l i t y rates r e s u l t i n g from t h i s p reda to r are unknown, d i f f e r e n c e s i n t h e l e v e l s of p reda t ion almost c e r t a i n l y exis t between r e s i d e n t forag ing areas. The precent of t i g e r sharks conta in ing t u r t l e s has been found t o vary cons iderably between some l o c a t i o n s i n t h e Hawaiian Archipelago, wi th t h e range extending from 6 . 7 % t o 75% (Table 19) . Furthermore, a t r e s i d e n t forag ing areas i n t h e NWHI where

s Iowcs i 1;i-owt-Ii o c * ( . r i r H , c.onip;ir;il I v c . 1 ~ 1 1 Il:hcr mort;il Lty r a t e s would be expected f o r 1 r tv (X i1 I I vti tlrw I o t i i o y,t-cI;i t vr 1 c1nf;tli o f L l m c Lticy a r t cxpoHed t o t i g e r sha rks as sii i: iI I t i i r t I P S . ‘I’IiIs 1.q ; i H s i i m L n I : , o f courHc!, t h a t a t l e a s t Rome i n c r e a s e i n p r o t e c t t o n rroin prc%tl;itLoti I H a f f o r t l c . d ;IH 3 t r i r t l e grows l a r g e r . Of t h e t u r t l e s rc .covcrcd from t if;cnr s1i;irks ;it French Frlgatc Shoi i l s , 4 4 % were a d u l t s and 592 wc’rc Jiivcnllc-s ( B ; i l n z ~ 70; Taylor and N n f t f l 4 8 2 ) . This may i n d i c a t e t h a t l a r g e r t r i r t l r s arc’ 1cs.s s t w c e p t i h l c t o a t t a c k , o r I t cotild slmply r e f l e c t a g r e a t e r ahrindrince of .)uvcnl_l CH.

‘l’hc s h o r t e r reprodirc.1 fvc c y c l cs of males could s e r v e t o i n c r e a s e t h e mor tn l i t y r a t e f o r t h i s sex. Ily undertaking morc f r equen t remigra t ions t o French F r i g a t e Shoals , males would bc spending more t i m e than females i n an area where t h e r i s k of shark a t t a c k appears t o be g r e a t e r . I n a d d i t i o n , t h e males t h a t u s u a l l y surround a copu la t ing p a i r a r e undoubtedly more vu lne rab le t o a t t a c k due t o t h e i r errat ic behavior and preoccupat ion wi th c o u r t s h i p a c t i v i t i e s . These accompanying males may a l s o be i n a d v e r t e n t l y se rv ing as a b u f f e r a g a i n s t shark a t t a c k f o r t h e c o p u l a t j n g p a i r . i nc lude c e r t a i n hazards t h a t i n c r e a s e m o r t a l i t y , such as d i s o r i e n t a t i o n i n t h e open ocean and exposure t o l a r g e p reda to r s no t p r e s e n t l y impl ica ted wi th Hawaiian Cholonia (i .e. k i l l e r whales, Oreinus orca, and g r e a t whi te s h a r k s , Carcharodon carcharias).

The long-distance remigra t ion i n i t s e l f may

The minimum s u r v i v a l ra te necessary t o main ta in t h e Hawaiian green t u r t l e popula t ion a t a s t a b l e l e v e l i s one i n which each female r e p l a c e s h e r s e l f w i th a female o f f s p r i n g t h a t s u r v i v e s t o l a y f e r t i l e eggs dur ing a t least one b reed ing season. Assuming an equa l sex r a t i o a t t h e t i m e of ha t ch ing , each a d u l t female a t French F r i g a t e Shoals produces an average of 66 female ha t ch l ings . I f a female i s on ly involved i n one breeding season du r ing h e r l i f e t i m e , t hen a t least 1.5% of h e r female h a t c h l i n g s would have t o s u r v i v e f o r t h e popula t ion to b e s t a b l e . Some Hawaiian Chelonia are , however, involved i n more than one breed- i n g season. This would i n c r e a s e t h e number of female h a t c h l i n g s produced, thereby lowering t h e percentage t h a t must reach ma tu r i ty . The s u r v i v a l rates and repro- d u c t i v e c y c l e s of a d u l t females a f t e r t h e i r f i r s t b reeding season are only p a r t i a l l y known a t t h e p re sen t t i m e ( s e c t i o n s 3.16, 3.3). Due t o t h e tagging e f f o r t t hus f a r expended a t French F r i g a t e Shoals and o t h e r l o c a t i o n s i n the Hawaiian Archipelago, t h e r e i s an e x c e l l e n t p o t e n t i a l f o r a c q u t r i n g t h i s informa- t ion w i t h i n a 5-yr pe r iod .

It should be emphasized t h a t i t is unknown i f t h e Hawaiian green t u r t l e popula t ion i s s t a b l e a t t h e p r e s e n t t i m e . The number of females n e s t i n g annual ly s i n c e 1973 has f l u c t u a t e d s u b s t a n t i a l l y , and no t r e n d s can p r e s e n t l y be de t ec t ed (F igure 12 ) .

4.5 Recruitment

The known f a c t o r s a f f e c t i n g recru i tment i n t h e Hawaiian green t u r t l e popula t ion are presented i n s e c t i o n s 2.22, 3.12, 3.33, 3.34, 3.43, 3.51, 4.3, 4.4, and 5. The t h r e e p r i n c i p a l l e v e l s of t h i s recru i tment are: 1) recru i tment of h a t c h l i n g s t o t h e p e l a g i c environment, 2) r ec ru i tmen t of j u v e n i l e s t o coastal fo rag ing areas, and 3) recru i tment of a d u l t males and females t o t h e breeding colony. The informat ion acqui red t o d a t e i n d i c a t e s t h a t t h e s e p rocesses are

c

43

*-

occurr ing i n a c losed system w i t h i n t h e Hawaiian Archipelago and i t s surrounding p e l a g i c waters. t h e Hawaiian Archipelago is occur r ing by j u v e n i l e s t h a t o r i g i n a t e d from o t h e r CheZonia popula t ions i n t h e P a c i f i c . With t h e p o s s i b l e except ion of Johnston A t o l l and Wake, t h e r e is a l s o no i n d i c a t i o n t h a t green t u r t l e s of Hawaiian o r i g i n are u t i l i z i n g any r e s i d e n t fo rag ing area o u t s i d e of t h e Hawaiian Archipelago. However, t h e p o s s i b i l i t y t h a t such i n t e r a c t i o n s may e x i s t should no t be e n t i r e l y excluded a t t h e p re sen t t i m e . A s i n a l l marine t u r t l e popu la t ions , b a s i c d a t a need t o be acqui red on t h e p e l a g i c segment of j u v e n i l e development.

There i s no i n d i c a t i o n t h a t recru i tment t o r e s i d e n t areas w i t h i n

5. EXPLOITATION

5 .1 H i s t o r i c a l Overview

The e x p l o i t a t i o n of green t u r t l e s f o r food w a s found t o be p a r t of the n a t i v e c u l t u r e when Captain James Cook red iscovered t h e Hawaiian I s l a n d s i n 1778 (Beaglehole 105; Anonymous 659). This t r a d i t i o n a l usage undoubtedly s t a r t e d as e a r l y as A.D. 600 w i t h t h e i n i t i a l occupat ion of H a w a i i by Polynesians from o t h e r P a c i f i c areas. i n e f f e c t u n t i l 1819, t u r t l e s could only be e a t e n by men who were n o b i l i t y o r p r i e s t s (Kalakaua 285). T u r t l e s were captured p r i n c i p a l l y by hand whi l e d iv ing underwater, w i th s p e a r s o r harpoons from shore , and wi th n e t s made of 'cord from t h e bark of t h e olona p l a n t , Touchardia ZatifoZia (Cobb 176) and ahu'awa p l a n t , Gypems javanicus (Buck 145) . The beveled edge of a p l e u r a l bone from a t u r t l e ' s carapace w a s o f t e n used t o sc rape t h i s bark and extract t h e durable f i b e r s (Malo 341; Stokes 471; TenBruggencate 484). Another method of cap tu re involved t h e use of two 7-cm hooks lashed t o a f l a t s t o n e t h a t w a s a t t ached t o a long l i n e (Cobb 176 wi th i l l u s t r a t i o n ) . This w a s appa ren t ly used t o hook t u r t l e s bo th from shore and wh i l e d iv ing i n areas where r e s t i n g t akes p l ace .

Under t h e s t r i c t l y enforced Hawaiian "kapu" system t h a t remained

Except f o r r e fe rences t o Nihoa i n chants and legends , n a t i v e Hawaiians w e r e appa ren t ly no t aware of t h e northwestern segment of t h e Hawaiian Archipelago a t t h e t i m e of Captain Cook's a r r i v a l (Emory 212). t h e r e f o r e confined t o the main i s l a n d s , a l though a t a much earlier d a t e some t u r t l e s were probably taken by t h e s m a l l groups of Hawaiians (or o t h e r Polynesians) t h a t occupied Nihoa and Necker. t u r t l e s by Hawaiians i s t h e i s l a n d which they c a l l e d K a Moku Papapa. This w a s descr ibed t o Captain Cook and h i s o f f i c e r s on s e v e r a l occas ions i n both 1778 and 1779 as a low sandy i s l a n d t h a t w a s sometimes v i s i t e d t o ca t ch t u r t l e s and sea- b i r d s . K a Moku Papapa w a s s a i d t o be loca ted t o t h e west-southwest of Kaula (lat. 21°39'N, long. 16Oo33'W), which i s a s m a l l rock i s l a n d 35 km t o t h e south- w e s t of Niihau (Figure 1 ) . From Kaula, where an overn ight v i s i t was u s u a l l y made, t h e n a t i v e s r epor t ed t h a t they could "very e a s i l y paddle t h e r e i n t h e course of t h e fol lowing day" (Beaglehole 105) . On 16 and 17 March 1779, t h e Xseovsry searched f o r K a Moku Papapa without success wh i l e depa r t ing from Hawaiian waters. Although no i s l a n d has eve r been found i n t h i s v i c i n i t y , a s m a l l s h o a l w i t h a depth of 9 m i s s i t u a t e d 6.5 k m t o t h e northwest of Kaula. I n a d d i t i o n , an area of d i sco lo red w a t e r poss ib ly r ep resen t ing a shoa l w a s r epor t ed i n 1955 a t a s i t e 35 km t o t h e southwest of Kaula a t l a t . 21"28'N, long. 160'45'W. Nevertheless , f o r t h e s h o r t per iod of t i m e involved and the absence of geo log ica l a c t i v i t y with- i n t h i s r eg ion of t h e Hawaiian Archipelago, i t would no t have been p o s s i b l e f o r an i s l a n d t o have s e t t l e d i n t o t h e ocean (G. A , MacDonald, pe r sona l communication).

E x p l o i t a t i o n of t u r t l e s w a s

An unresolved a spec t of t h e e x p l o i t a t i o n of

44

The t r a d i t i o n a l c o n t r o l l e d e x p l o i t a t i o n of t u r t l e s by Hawaiians g radua l ly disappeared w i t h t h e a b o l i t i o n of t he kapu system, t h e i n f l u x of Caucasians and o t h e r rac ia l groups, and t h e discovery of t h e unexplo' i ted and uninhabi ted northwestern segment of t h e Hawaiian Archipelago. Numerous commercial exped i t ions t o t h e NWHI took p l a c e dur ing t h e 1800's and e a r l y 1900's t o e x p l o i t green t u r t l e s , s e a b i r d s , monk seals, sha rks , and beche-de-mer (Holothuria spp.) . T u r t l e s were taken p r i n c i p a l l y f o r meat, o i l and, a long wi th monk seals, f o r use as a s u p e r i o r sha rk b a i t . v i s i t e d these i s l a n d s f o r 5 m o i n 1882, a t least 410 t u r t l e s were taken o f f t h e beaches and from t h e ad jacen t waters (Hornell 271). t o t h e NWHI t h a t involved t h e e x p l o i t a t i o n of t u r t l e s dur ing t h i s e a r l y pe r iod are descr ibed by Amerson ( 3 0 ) , Amerson e t a l . (32) , Clapp and Ely (170) , Ely and Clapp (211), F a r r e l l (220) , Gal t sof f (231), K e m b l e (287) , Munro (365-370), Walker (531), Woodward (556) , and Anonymous (569). I n June of 1923, t h e Tanager Expedi- t i o n found evidence t h a t t u r t l e s had been r e c e n t l y k i l l e d a t French F r i g a t e Shoals (Wetmore 541). of 1867 desc r ibed t h e rare presence of t u r t l e eggs i n t h e Honolulu market t h a t had been imported from t h e NWHI (Anonymous 565).

When t h e Japanese-chartered f i s h i n g v e s s e l A d a

Other commercial exped i t ions

6,

It is a l s o of i n t e r e s t t o n o t e t h a t a newspaper a r t ic le i n June

Another f a c t o r i n t h e e x p l o i t a t i o n of Hawaiian CheZonia w a s t h e occurrence of shipwrecks. o f t e n had t o depend on t u r t l e s and o t h e r marine and terrestrial animals f o r food sources (Clapp and Wirtz 1 7 1 ; L is iansky 335; Read 429; Ward 538; Anonymous 566). The 30 s t r anded c r e w members of a whaling vessel wrecked a t French F r i g a t e Shoals i n March of 1859 k i l l e d i n excess of 100 t u r t l e s be fo re being rescued (Amerson 30; Walker 531).

The s u r v i v o r s of v e s s e l s t h a t s t r u c k r e e f s i n t h e "HI

I n 1909 a l l of t h e NWHI except Midway were dec lared a United S t a t e s of I n America p rese rve f o r n a t i v e b i r d s known as t h e Hawaiian I s l a n d s Reservat ion.

1936 Kure w a s removed from t h i s Reservat ion f o r m i l i t a r y purposes and la ter , i n 1952, t r a n s f e r r e d t o t h e T e r r i t o r y of H a w a i i . Hawaiian I s l a n d s Reservat ion w e r e redes igna ted as t h e Hawaiian I s l a n d s Nat iona l W i l d l i f e Refuge. However, u n t i l r e c e n t yea r s t h i s p ro tec t ed re fuge s t a t u s has no t served as a s i g n i f i c a n t d e t e r r e n t t o t h e e x p l o i t a t i o n of t u r t l e s . I n 1946 a commercial f i s h i n g base w a s e s t a b l i s h e d by t h e T e r r i t o r y of H a w a i i a t French F r i g a t e Shoals . Both t u r t l e s and f i s h w e r e cap tured i n t h e area and t r anspor t ed t o Honolulu by s m a l l a i r c r a f t us ing t h e abandoned m i l i t a r y landing s t r i p on Tern I s l and . T u r t l e meat a l s o became a main food source i n t h e d i e t s of t h e r e s i d e n t f ishermen. about 200 t u r t l e s from 1946 u n t i l they te rmina ted ope ra t ions i n 1948 (L. K. Agard i n litt. t o Amerson 30) . Over t h i s pe r iod , t h e fishermen noted a d e c l i n e i n t h e numbers of t u r t l e s . Amerson (30) specu la t ed t h a t t h i s w a s more t h e r e s u l t of human d i s tu rbance than a c t u a l k i l l i n g , however such an explana t ion seem un l ike ly . During t h e summer of 1959, t u r t l e s were aga in exp lo i t ed on t h e i s l a n d s a t French F r i g a t e Shoals by a commercial f i s h i n g company based i n Honolulu ( P a c i f i c Ocean F i s h e r i e s I n v e s t i g a t i o n s 395; D. W. S t r a sburg , personal communication; J. J. Di r sche l , personal c o m u n i c a t i o n ) .

I n 1940 t h e remaining areas of t h e

One of t h e two f i s h i n g companies us ing t h i s base es t imated t ak ing

R.

L

C'

c

Both the commercial and noncommercial e x p l o i t a t i o n of green t u r t l e s i n

a b o l i t i o n of t h e kapu system, u n t i l t h e adopt ion of a p r o t e c t i v e r e g u l a t i o n by t h e t h e main i s l a n d s proceeded w i t h v i r t u a l l y no c o n t r o l s from t h e t i m e of t h e 1819 P

x

45

u

HDFG i n May of 1974. During t h i s pe r iod , t h e hunt ing techniques increased i n both e f f i c i e n c y and s o p h i s t i c a t i o n , and t h e hunt ing range expanded t o n e a r l y a l l c o a s t a l areas. This w a s due p r i n c i p a l l y t o t h e a v a i l a b i l i t y of outboard motors, motor v e h i c l e s , f ire;irms, spearguns, inexpensive machine-made n e t s and, a t a l a te r d a t e , scuba equipment. I n a d d i t i o n , t h e commercial demand f o r t u r t l e meat and o t h e r Hawaiian seafoods increased cons iderably w i t h t h e advent of l a rge - sca l e tourism fo l lowing s ta tehood i n 1959 (Balazs 48; Benson 115; Hendrickson 262; Smyser 448). e n t l y no t been commercially exported s i n c e t h e t u r n of t h e century. However, such commerce wi th Japan w a s proposed a s r e c e n t l y as 1973 (Borreca 128, Linder 332).

Based on t h e a v a i l a b l e r eco rds , Hawaiian green t u r t l e s have appar-

A s a gene ra l index of p o t e n t i a l e x p l o i t a t i v e p r e s s u r e s , i t is of va lue t o n o t e t h a t t h e r e s i d e n t human popula t ion of t h e main Hawaiian I s l a n d s w a s approximately 130,000 i n 1831, 54,000 i n 1876, and 770,000 i n 1970. Since 1831 t h e d i s t r i b u t i o n of t h e human popula t ion has changed from 23% on Oahu and 77% on t h e o t h e r main i s l a n d s , t o 82% on Oahu and 18% on t h e o t h e r main i s l a n d s . I n 1778 t h e r e were an es t imated 200,000 t o 300,000 Hawaiian i n h a b i t a n t s . I n t h e NWHI, Midway w a s f i r s t colonized i n 1902 by t h e Commercial P a c i f i c Cable Company. U.S. t e r r i t o r y , t h e a t o l l has been occupied cont inuously s i n c e t h a t t i m e under t h e j u r i s d i c t i o n of t h e Navy. I n 1978 t h e r e s i d e n t popula t ion w a s reduced t o approximately 400 from t h e former 2,200 where i t had been maintained f o r a number of years . Kure has been cont inuously inhab i t ed by approximately 20 personnel s i n c e t h e cons t ruc t ion of a U.S. Coast Guard LORAN s t a t i o n i n 1960. I s l a n d , a s m a l l r e s i d e n t popula t ion e x i s t e d from 1891 t o 1910, p r i m a r i l y i n conjunct ion wi th t h e mining of guano (Bryan 139-141; Ely and Clapp 211) .

As a

A t Laysan

5.2 Commercial Catch Reports

T u r t l e s captured i n t h e Hawaiian I s l a n d s f o r commercial purposes have been p e r i o d i c a l l y recorded under t h e ca tegory of "honu ( t u r t l e ) " i n conjunct ion w i t h o t h e r f i s h e r i e s s t a t i s t i c s ( see Sub jec t Index) . cases where hawksbi l l s and l ea the rbacks have been taken , t h e s e r eco rds are only known t o invo lve green t u r t l e s .

Except f o r t h e few rare

Cobb (176) p r e s e n t s d a t a showing t h a t over a 4-yr pe r iod a t t h e t u r n of t h e century , from 168 t o 258 t u r t l e s were inspec ted and s o l d annual ly i n t h e Honolulu f i s h market (Table 20). The t o t a l weight recorded f o r t h e 184 t u r t l e s s o l d i n 1900 w a s 1 ,248 kg, which y i e l d s an average of 6.8 kg p e r t u r t l e . q u a n t i t y t h e r e f o r e probably only r e p r e s e n t s t h e m e a t and o t h e r e d i b l e p o r t i o n s a f t e r bu tcher ing . Based on t h i s r epor t ed weight , approximately 96% of t h e t u r t l e s were captured wi th s p e a r s and 4% w i t h n e t s . Ice w a s no t used i n t h e Honolulu f i s h market o r a t markets on t h e o t h e r i s l a n d s du r ing t h i s pe r iod due t o t h e p r o h i b i t i v e c o s t . Spearing must t h e r e f o r e have been c a r r i e d o u t i n a manner t h a t would a l low t h e t u r t l e s t o b e kep t a l ive u n t i l butchered and s o l d . For 1900, Cobb (176) a l s o r eco rds t h e sale of 443 kg of t u r t l e on Maui, 364 kg on H a w a i i , and 68 kg on Molokai, a l l of which w e r e taken w i t h spea r s . It is a l s o of i n t e r e s t t o note t h a t 192 l - l b cans of green t u r t l e w e r e among t h e l a r g e q u a n t i t i e s of f i s h e r i e s prod- u c t s imported t o t h e Hawaiian I s l a n d s dur ing t h e yea r 1900.

This

Since a t least 1944, commercial f ishermen i n t h e Hawaiian I s l a n d s have been r equ i r ed t o be l i c e n s e d and t o f i l e c a t c h r e p o r t s on a l l s p e c i e s s o l d . From

46

1944 t o 1947, t h e amount of t u r t l e r epor t ed annual ly by l ive weight under t h i s system was as fo l lows: 1944 - 18,007 kg (no r e p o r t s f o r January and February, r e p o r t missing f o r A p r i l ) ; 1945 - 8,954 kg ( r e p o r t missing f o r A p r i l ) ; 1946 - 3,717 kg (no r e p o r t s f o r June, J u l y , and August); 1947 - 4,317 kg. Table 21 p r e s e n t s t h e amount of t u r t l e r epor t ed annual ly by v a r i o u s methods of cap tu re f o r t h e y e a r s 1948 t o 1973. These d a t a f l u c t u a t e cons iderably , w i t h annual amounts ranging from 173 kg i n 1963 t o 11,628 kg i n 1972. A t o t a l of 90,803 kg w a s r epor t ed du r ing t h e 26-yr per iod . Due t o t h e absence of a v e r i f i c a t i o n system, unde r repor t ing and nonrepor t ing undoubtedly had a s i g n i f i c a n t i n f luence on t h e s e va lues (Hendrickson 262). On Kauai, no r e p o r t s f o r t u r t l e s were f i l e d from 1957 t o 1969 (Table 1 7 ) . However, in format ion communicated t o t h e au tho r by a knowledgeable source i n d i c a t e d t h a t approximately 4,500 kg of t u r t l e were purchased from fishermen by a s i n g l e t o u r i s t - o r i e n t e d r e s t a u r a n t on Kauai dur ing t h a t yea r . an absence of any t u r t l e r e p o r t s f o r Oahu dur ing t h e yea r s 1957, 1960, 1961, and 1968, as w e l l as i n 1 4 of t h e 26 y r f o r t h e i s l a n d of H a w a i i (Table 17 ) .

I n 1973 only 182 kg w e r e r epor t ed f o r t h i s i s l a n d .

There was a l s o

The l i s t i n g of t h e methods used i n t h e cap tu re ope ra t ions w a s no t w e l l def ined i n t h e commercial r e p o r t s . This r e s u l t e d i n a number of unc lea r cate- g o r i e s t h a t are s u b j e c t t o some i n t e r p r e t a t i o n (Table 21) . 1961 most of t h e r e p o r t s d i d n o t s p e c i f y t h e method of capture .

In a d d i t i o n , p r i o r t o

x

The most f r equen t ly r epor t ed cap tu re method f o r a l l y e a r s was n e t s

and t h e r e f o r e should be combined w i t h t h e (49.5%). The c a t e g o r i e s of scuba (5.6%) and d i v e (0.5%) probably involved t u r t l e s t h a t were captured s o l e l y by hand hand (2.3%) ca tegory t o g ive a t o t a l of 8.4%. The f i r s t r epor t ed use of scuba f o r li

cap tu r ing t u r t l e s commercially w a s i n 1958. The noose ( 0 . 3 % ) and handl ine (1.5%) methods are a l s o probably synonymous. This Involves p l ac ing a l i n e around t h e f l i p p e r o r neck of a t u r t l e s l e e p i n g underwater. The t u r t l e is then p u l l e d t o t h e s u r f a c e by o t h e r f ishermen wa i t ing i n a boa t . The spea r (3.4%) ca tegory , which was repor ted most ly af ter 1964, could inc lude spearguns used whi le d iv ing , as w e l l

r epo r t ed e x c l u s i v e l y a f t e r 1968, a long wi th t h e hook and l i n e (0.3%) ca tegory , are be l i eved t o invo lve v a r i a t i o n s of t h e earlier Hawaiian p r a c t i c e of ca t ch ing t u r t l e s w i t h a l i n e a t t a c h e d t o a s t o n e and two hooks (Cobb 1 7 6 ) . i nco rpora t e s a bu tche r ' s hook a t t a c h e d t o a l i n e t h a t l e a d s e i t h e r t o 8 boat where o t h e r f ishermen are wa i t ing , o r t o a f r e e - f l o a t i n g buoy a t t h e su r face . The hook

p u l l e d t o t h e s u r f a c e o r allowed t o s w i m u n t i l exhausted from towing t h e buoy. Another g a f t o r hook and l i n e technique is c a r r i e d o u t from shore wi th barbed hooks cast i n t o a l e a d weight t h a t is a t t ached t o a po le and l i n e . c o a s t a l c l i f f s a long foraging areas, t u r t l e s are then hooked i n t h e f l i p p e r s o r neck when they s w i m w i t h i n range, The s i n g l e r e p o r t l i s t e d under t h e ca tegory of

No in format ion is c u r r e n t l y a v a i l a b l e on t h e ca tegory des igna ted as t r a p (0.4%) which appeared i n t h e records dur ing 1962, 1963, and 1969. Poss ib ly t h i s involved the e n t r y of t u r t l e s i n t o l a r g e w i r e cages t h a t are used by some fishermen t o t r a p f i s h .

as s p e a r s o r harpoons thrown from a b o a t o r from shore . The g a f t (10.2%) category d=,

One procedure

is t hen implanted i n t o a l a r g e t u r t l e found s l eep ing and subsequent ly e i t h e r c

Working from

t r o l l i n g (0.1%) probably involved an a c c i d e n t a l snagging whi le t r o l l i n g f o r f i s h . 6

I n a d d i t i o n t o t h e methods presented i n Table 21 , t h e commercial cap tu re of t u r t l e s i s known t o have inc luded t h e i l l e g a l use of powerheads o r

47

underwater shark guns, as well as rifles used from shore in conjunction with a boat to retrieve the dead or wounded turtles.

Y

a

Over the 26-yr period, the greatest amount of turtle was reported for March (11.6%), and the least amount reported for November (4.8%) (Table 22). Figure 13 shows that the greatest amount of turtle was reported from Maui (37.1%), followed by Oahu (34.1%), and Molokai (17.6%). However, the absence of reports from Kauai and Hawaii during a number of years undoubtedly biased these percent- ages.

Along with catch data, commercial fishermen occasionally included informative observational notes in their reports to the HDFG. A report filed for July 1968 recording the capture of 15 turtles in the area between Maui, Molokai, and Lana.i stated "This area in 1948-1950 I used to catch at least 100 in 4 to 5 days fishing - for some reason there are no turtles there now."

Some of the turtles commercially captured in the "HI were reported to the HDFG, These data are listed as follows. Pearl and Hermes Reef: 1953 - 161 kg, 1958 - 817 kg; Gardner Pinnacles: 1948 - 627 kg; Necker: 1951 - 690 kg; French Frigate Shoals: 1948 - 201 kg, 1950 - 531 kg, 1951 - 520 kg, 1953 - 730 kg, 1957 - 155 kg. cially at French Frigate Shoals from 1946 to 1948, or the captures made in 1959 (section 5.1).

Apparently no reports were filed for the 200 turtles taken commer-

5.3 Noncommercial Catch

The noncommercial capture of green turtles in the Hawaiian Archipelago has resulted principally from sport and trophy hunting, and an esteem for turtles as food. The capture of turtles for true subsistence purposes has not been documented, but nevertheless may have occurred (and may still occur) in small numbers at some rural locations.

No records exist for the noncommercial capture of turtles prior to May of 1974. The annual amounts taken are therefore problematic but may have been substantial, particularly with the advent of scuba equipment, of the HDFG, quoted in 1969 by Hendrickson (262), stated that '.'I have little doubt that the sport fishery take plus possible unlicensed commercial take far exceeds the legitimate commercial take." From May of 1974 to June of 1977, 48 adults and 1 juvenile (an illegal capture) were reported in compliance with the HDFG regulation that permitted only noncommercial exploitation.

The Director

The methods used for the noncommercial capture of turtles are the same as those employed for commercial purposes (section 5.2 and Subject Index), Some recreational fishermen have also successfully taken turtles using a surf rod and reel with a baited hook (Carter 158). of a hunting bow with a line attached to an arrow (Anonymous 583).

Another novel technique involved the use

The capture of Hawaiian Chelonia for noncommercial purposes was stimulated in 1975 with the publication of a popular book that described how to prepare sunburn lotion from green turtles (McBride 3 4 8 ) . attack by tiger sharks resulting from use of such ointments before entering the ocean have been pointed out (Balazs 65).

The potential risks of

48

P r i o r t o 1977, green t u r t l e s a t Midway were a t times e x p l o i t e d by r e s i d e n t personnel f o r t r o p h i e s and food. S imi l a r cap tu res were s p o r a d i c a l l y made by m i l i t a r y personnel s t a t i o n e d a t Kure and French F r i g a t e Shoals .

'

6. PROTECTION AND MANAGEMIINT

6 .1 Regulatory Measures

Regulatory c o n t r o l s on t h e cap tu re of Hawaiian Chelonia p r i o r t o May of 1974 inc luded a ban on t h e use of f i r ea rms and, s i n c e 1949, a p r o h i b i t i o n on t h e sale of t u r t l e s taken w i t h s p e a r s ( T e r r i t o r y of H a w a i i 486; S t a t e of H a w a i i 457, 458). Both of t h e s e measures a l s o covered c rus t aceans , mollusks, a q u a t i c mammals, and f i s h (o the r t han s h a r k s ) . The p r o h i b i t i o n on t h e sale of t u r t l e s taken wi th s p e a r s appa ren t ly w a s never enforced (Table 21).

I n May of 1974, t h e adopt ion of Regulat ion 36 of t h e HDFG banned t h e commercial e x p l o i t a t i o n of green t u r t l e s and t h e use of t a n g l e n e t s . Adults measuring 36 i n . (91 cm) and l a r g e r i n s t r a i g h t carapace l eng th could be taken f o r "home consumption," b u t on ly from waters around t h e e i g h t main i s l a n d s . The eggs and progeny of c a p t i v e Hawaiian Chelonia could be s o l d under permi t , a long wi th green t u r t l e products imported from any area o u t s i d e of t h e Hawaiian I s l a n d s ( S t a t e of H a w a i i 465). state level t o t h e hawksbi l l and l ea the rback , which were a l r eady l i s t e d i n t h e Endangered ca tegory under t h e U.S. Endangered Species Act. surrounding t h e adopt ion of Regulat ion 36 are descr ibed i n ar t ic les by Altonn (8, 9 , 11, 12) , Balazs (46-48) Benson (115, 116) , Chiav ie l lo (160), Ching and McCabe (161), Kido (289-298), Moake (358, 359) , Smyser (448, 449) , Standbury and F l a t t a u (453) , S t a t e of H a w a i i (459-463) , S t o f f e l (470) , and Anonymous (594, 595,

This r e g u l a t o r y measure a l s o gave f u l l p r o t e c t i o n a t t h e

The series of events

597, 599, 600, 602-608, 610, 612, 618, 621, 622).

I n September of 1978, Regulat ion 36 w a s superseded wi th t h e l i s t i n g of Hawaiian Chelonia i n t h e Threatened ca tegory under t h e U.S. Endangered Species A c t (U.S. Department of Commerce 503, 504; Anonymous 668, 669). The popula t ion c u r r e n t l y receives f u l l p r o t e c t i o n through Federa l r e g u l a t i o n s i s s u e d a long wi th t h i s l i s t i n g . Under an agreement formalized i n J u l y of 1977, the j u r i s d i c t i o n over a l l marine t u r t l e s s u b j e c t t o Federa l l a w i s shared j o i n t l y by t h e Nat iona l Marine F i s h e r i e s Service (NMFS) and t h e USFWS,

I n thr? NWHI, f u l l p r o t e c t i o n f o r green t u r t l e s i n t h e Fede ra l ly c o n t r o l l e d Hawaiian I s l a n d s Nat iona l W i l d l i f e Refuge has a t least t h e o r e t i c a l l y been a f fo rded s i n c e 1940. Although t h e exac t re fuge boundaries of each i s l and- area have n o t always been c l e a r l y def ined , as a minimum they encompassed a l l emergent l and and t h e s h o r e l i n e waters. It is r e l e v a n t t o n o t e t h a t i n A p r i l of 1952 Refuge as a w i l d l i f e re fuge under t e r r i t o r i a l l a w . However, according t o t h i s d e c l a r a t i o n (Resolu t ion 7 of t h e Board of Commissioners of Agr i cu l tu re and F o r e s t r y ) , t h i s w a s only " . . . f o r management as a re fuge f o r t h e mammal and b i r d w i l d l i f e found thereon. ..." Aspects of t h e disagreement between t h e State and Federa l governments over t h e re fuge boundaries and t h e involvement of Hawaiian Chelonia appear i n ar t ic les by Benson (113, 121-123), Conant (180, l S l ) , Kakesako (284), Miller (356) , U.S. Department of t h e I n t e r i o r (524), Zalburg (563), and Anonymous (650).

t h e T e r r i t o r y of H a w a i i des igna ted t h e Hawaiian I s l a n d s Nat iona l W i l d l i f e

49

u

The f i r s t known regu la to ry measure on t h e cap tu re of Hawaiian Chelonia a t Midway w a s i s sued by t h e U.S. Navy s t a t i o n i n 1969, and la te r formal ly incorpora ted i n t o t h e area's w i l d l i f e management p l a n (U.S. N a v y 528). Under t h i s r e g u l a t i o n , green t u r t l e s measuring 24 in (61 cm) and l a r g e r i n s t r a i g h t carapace l eng th could be taken , but only by hand and n o t more than one p e r person each day. I n 1975 t h i s s i z e r e s t r i c t i o n w a s increased t o 36 i n . , and i n 1977 t h e t u r t l e s were a f fo rded f u l l p r o t e c t i o n .

Between 1973 and 1976, a number of b i l l s and r e s o l u t i o n s r e l a t i n g t o t h e p r o t e c t i o n and r e sea rch of Hawaiian Chelonia were in t roduced i n t o t h e H a w a i i S t a t e L e g i s l a t u r e ( see Subjec t Index). l e g i s l a t i v e approval w a s t h e appropr i a t ion of funds t h a t have been used t o ' p a r t i a l l y suppor t t h e au tho r ' s p re sen t r e sea rch program (King 305; Anonymous 642).

The only measure t o even tua l ly receive

6.2 Management P r a c t i c e s and Options

6 .21 A r t i f i c i a l s tock ing

There are c u r r e n t l y no proven techniques f o r r ep len i sh ing marine t u r t l e popula t ions through s tock ing o r o t h e r a r t i f i c i a l manipulat ions. However, i n a number of popula t ions t h e h igh level of p reda t ion on eggs and h a t c h l i n g s warran ts t h e use of experimental measures. These inc lude t h e t r a n s f e r of eggs t o p ro tec t ed areas f o r incubat ion and ha tch ing , and t h e subsequent release of h a t c h l i n g s under guarded cond i t ions on t h e beach o r a f e w k i lome te r s o f f s h o r e (Hi r th 265; Parsons 400). Another experimental procedure is t h e "heads ta r t ing" o r r a i s i n g of h a t c h l i n g s i n c a p t i v i t y t o a j u v e n i l e s i z e where, presumably, less p reda t ion w i l l occur when they are r e l e a s e d i n t o t h e wi ld .

The experimental s tock ing of Hawaiian Chelonia has thus f a r on ly been I n 1976, 295 of t h e 398 green t u r t l e h a t c h l i n g s undertaken on a l i m i t e d b a s i s .

t h a t emerged from n e s t s l a i d i n c a p t i v i t y a t Sea L i f e Park were r e l e a s e d o f f Oahu as a s tock ing e f f o r t . cons t ruc t ion i n 1974 of a n 8 by 15 m pool and ad jacen t sand beach t o s e r v e as a d i s p l a y and experimental breeding f a c i l i t y (Bourke et a l . 131; Anonymous 636, 643). A l l of t h e green t u r t l e s i n t h i s pool (9 females , 3 males, 6 subadu l t s ) had been captured i n t h e main i s l a n d s a t va r ious unrecorded times s i n c e 1964. On f o u r occas ions , eggs w e r e recovered from t h e bottom of an earlier d i s p l a y pool i n use a t Sea L i f e Park, however, incubat ion a t t empt s w e r e no t success fu l . Mating a c t i v i t y w a s occas iona l ly observed a f t e r t h e t u r t l e s were moved t o t h e new f a c i l i t y i n 1974, bu t n e s t i n g d i d no t commence u n t i l June of 1976. Only f o u r of t h e n i n e females w e r e be l ieved t o have been involved i n t h e nes t ings . Although some mating a c t i v i t y has occurred s i n c e 1976, a d d i t i o n a l egg l a y i n g has no t y e t r e s u l t e d . Mating a c t i v i t y , as w e l l a s t h e depos i t i on of eggs i n d i s p l a y tanks l ack ing a sand beach, have a l s o been p e r i o d i c a l l y recorded i n Hawaiian Chelonia a t t h e Waikiki Aquarium ( D e Luca 193; Mowbray 362) and t h e Kahala H i l t o n Ho te l (Daacon 192) .

The propagat ion of t h e s e t u r t l e s w a s made p o s s i b l e by t h e

Twenty-four of t h e h a t c h l i n g s t h a t o r i g i n a t e d from Sea L i f e Park were subsequent ly r a i s e d i n c a p t i v i t y f o r 5 mo be fo re be ing tagged and r e l eased o f f t h e western c o a s t of Lanai (Bourke et a l . 131) . None of t h e s e t u r t l e s have thus f a r been recovered.

50

Between A p r i l of 1974 and August of 1977 a t o t a l of 54 j u v e n i l e s , 20 t o 60 mo of age , were r e l eased o f f Oahu a f t e r be ing r a i s e d i n c a p t i v i t y from h a t c h l i n g s t h a t were obta ined a t French F r i g a t e Shoals (Altonn 5) . Pourteen of t h e s e t u r t l e s have been recovered under c o n t r a s t i n g circumstances. One t u r t l e w a s found r e s t i n g under a c o r a l l edge a long t h e Kau c o a s t l i n e , a minimum movement of 380 km from t h e release s i t e ( s e c t i o n 3.43). Another t u r t l e t r a v e l e d 110 km and w a s speared by a fisherman o f f t h e no r th sho re of Lanai 11 m o a f t e r being r e l eased . several o t h e r Chelonia. of t h e n a t u r a l l y occur r ing t u r t l e s r a p i d l y l e f t t h e area. Other r ecove r i e s included two t u r t l e s found swimming c l o s e t o shore i n an e r r a t i c manner 2 days a f t e r release; one t u r t l e tangled i n a c rab n e t 2 days a f t e r release; one t u r t l e found on shore i n s i d e an enc losure used f o r porpoise 6 wk a f t e r release; one t u r t l e speared 2 wk a f t e r release; and e i g h t t u r t l e s found w i t h i n 200 m of t h e release s i te over pe r iods ranging up t o 4 mo a f t e r release.

This w a s an i n t e r e s t i n g recovery s i n c e t h e t u r t l e w a s observed wi th However, when t h e f isherman's presence w a s de t ec t ed , a l l

I n May of 1975, 26 o t h e r j u v e n i l e s t h a t had been r a i s e d i n c a p t i v i t y from h a t c h l i n g s w e r e r e tu rned t o French F r i g a t e Shoals (Altonn 15 ) . t u r t l e s have been recovered, w i th two i n d i v i d u a l s e s t a b l i s h i n g a per iod of res idency n e a r Tern I s l a n d ( s e c t i o n 3.43, Table 11). The o t h e r t h r e e r ecove r i e s were made 4 days a f t e r release, and included a t u r t l e found crawling a long t h e beach on Tern I s l a n d , a t u r t l e found basking nea r an a d u l t on T r i g I s l a n d , and a t u r t l e found swimming i n shal low water ad jacen t t o Whale-Skate I s l and .

Five of t h e s e

A p o t e n t i a l e x i s t s f o r i n c r e a s i n g t h e number of l ive h a t c h l i n g s a t French F r i g a t e Shoals by excavat ing n e s t s a f t e r n a t u r a l emergence has occurred i n o rde r t o retrieve those i n d i v i d u a l s t h a t d id no t reach t h e su r face . Although 76.7% of t h e eggs l a i d i n each n e s t a t French F r i g a t e Shoals produce l i v e hatch- l i n g s , only 70.8% r e s u l t i n l i ve h a t c h l i n g s reaching t h e s u r f a c e ( s e c t i o n 3.15, Table 7) . i n m o r t a l i t y . cond i t ion i f they are excavated w i t h i n a 3- t o 5-day per iod .

The remaining 5 .9%,or an annual average of 1 ,988 h a t c h l i n g s , r e s u l t Many of t h e s e h a t c h l i n g s can be salvaged i n an appa ren t ly hea l thy

6.22 P reda to r con t ro l

No measures have thus f a r been taken t o reduce t h e nupber of ghost c r abs a t French F r i g a t e Shoals i n o r d e r t o enhance t h e s u r v i v a l of h a t c h l i n g s when they leave t h e n e s t and c r o s s through t h e i n t e r t i d a l zone. no t excess ive compared t o many o t h e r green t u r t l e breeding areas, up t o 1,200 h a t c h l i n g s (5%) may be e l imina ted annual ly a t French F r i g a t e Shoals by t h e s e c rus t aceans ( s e c t i o n 3 .2) . duc t ion of a l i m i t e d experimental c o n t r o l program.

Although p reda t ion is

The s m a l l s i z e of the i s l a n d s would s impl i fy t h e con-

Shark f i s h i n g programs have been c a r r i e d ou t on several occas ions i n t h e main i s l a n d s f o r t he purpose of decreas ing t h e r i s k of a t t a c k on humans, promoting shark meat a s a food source , and f o r b i o l o g i c a l research . These in ten- sive f i s h i n g programs w e r e undoubtedly b e n e f i c i a l t o Hawaiian CheZonia i n view of t h e l a r g e number of t i g e r sha rks e l imina ted . t i g e r sha rks has taken p l a c e a t a reduced level i n conjunct ion wi th predator-prey r e sea rch of t h e monk seal (Taylor and N a f t e l 482) and c u r r e n t l y as p a r t of t h e t r o p h i c a n a l y s i s of f i s h communities ( P a r r i s h and Taylor 399). c o n t r o l program a t French F r i g a t e Shoals would r e s u l t i n immediate s u r v i v a l

I n t h e NWHI, t h e cap tu re of

While an i n t e n s i v e

P

c

51

P

Y

enhancement for green turtles, the possible long-term implications of such action would have to be carefully assessed. One possible adverse impact might be an increase in the number of small sharks after the large predators have been removed (Taylor and Naftel 482; Tester 487). This, in turn, could conceivably result in greater predation on hatchlings.

6.23 Enforcement and education

The enforcement of Federal regulations protecting green turtles in the Hawaiian Archipelago is currently the responsibility of two agents with the USFWS and two agents with the NMFS. are responsible for enforcing numerous other Federal wildlife and fisheries regulations (Benson 110; Vernon 530). Officers of the Department of Land and Natural Resources, State of Hawaii, continue to enforce the applicable provisions of Regulation 3 6 . The Commanding Officer of the U.S. Navy station at Midway also has enforcement responsibilities for the protection of Hawaiian Chelonia under his jurisdiction (U.S. Navy 528). tions, enforcement efforts with Hawaiian Chelonia are not complicated by international migrations. However, illegal exploitation is known to occur within the Hawaiian Archipelago and the magnitude of this catch needs to be determined.

These personnel are based in Honolulu and

Unlike nearly all other green turtle popula-

Since 1973 a number of magazine and newspaper articles,, as well as lectures, have served to provide information to the general public about Hawaiian Chelonia (Baldwin 94; Balazs 44, 48, 55, 60, 62; Chong 162, 165; Eliot 208; Hendricks 261; Kim 299; Lipman 333; Sekora 439; Whitten 547; Anonymous 580, 608, 649, 660, 661, 663). In addition, the display facilities and educational programs at Sea Life Park and the Wakiki Aquarium have helped to generate interest in the biology and conservation of this population (Engle 215; Taylor $77-479; Anonymous 636).

6.24 Aquaculture

The intensive culture and breeding of Hawaiian Chelonia for conservation purposes, such as artificial stocking of hatchlings and headstarting experimenta- tion, offer some potential as a management technique (section 6.21). However, this is likely to be an expensive undertaking unless carried out in conjunction with existing display facilities.

Periodic interest has been expressed in the commercial aquaculture of Hawaiian Chelonia as a source of marketable meat, curios, and soup stock (Altonn 13; Benson 118; Barreca 129; Finnerty 221; Haugen 258; Mahikoa 340; Pryor 420-422; Anonymous 615). However, no operations were ever actually started. Investigations conducted by the author suggested that the commercial aquaculture of Hawaiian green turtles was neither biologically nor economically feasible under existing circumstances. Furthermore, such facilities were not considered to.be in the conservation interests of naturally occurring marine turtle populations (Altonn 15; Balazs 52, 58-60; Benson 119; Ehrenfeld 206; Pfund 407). Under the present Federal regulations, commerce in green turtle products is not permitted, and turtles cannot be acquired from the wild for use in commercial aquaculture facilities.

52

The only aquaculture o f Chelonia known to have been carried out by the, early Hawaiians involved a 15 by 150 m coastal pond on Oahu named Pahonu that was used to maintain turtles until they were ready to be eaten (McAllister 345; Whitten 544).

P

L

c

c

53

BIBLIOGRAPHY OF MAZTNE TURTLES I N THE HAWAIIAN ISLANDS

a.

1, Abbott, I. A. and E . i l . d i I tiamson (1974) L I P ~ I : An e thnobotanica l s tudy of some e d i b l e i idwuian seawwds. Gardens, L a w a i , Kauai, 2 1 p p .

P a r i Lic Trop ica l Botanica l

2 . Akaj i , S. (1961) Hook, l i n e anti :;.i.Eft(,i : HGE,\ f i s h i n g c o n t e s t . The Carden Island (Lihue, Kauai ) , 25 Octoht$r, 6 .

3. A l t i e r y , M . , Senator (1974) Re la t ing t o t h e p re se rva t ion of green sea t u r t l e s . S t a t e of H a w a i i , The Senate , Seventh Leg i s l a tu re : A . B i 1 1 f o r an Act, Senate B i l l No. 1530.

4. Altonn, H. (1970) Ecologis t seeks t u r t l e farming sites. Honolulu Star-Bulletin, 1 7 September, C:2.

5 . Altonn, H. (1972) He's incubat ing a new indus t ry : S c i e n t i s t seeks t o t r a n s p l a n t green t u r t l e t o Oahu. 21 June, C:12.

Honolulu Star-Bulletin,

.- 6. Altonn, H. (1973) Secr.et base le ts t u r t l e o u t of bag. , HonoZulu

Star-Bulletin, 24 May, B:4. (1973) 34, 1:9.

Reprinted i n p a r t i n The Elepaio

x

e.

7. Altonn, H. (1973) Here i n H a w a i i : L a s t o u t p o s t f o r t h e green t u r t l e . Honolulu Star-Bulletin, 3 October, A:18 .

8.

9.

10.

11.

12.

13

14.

15.

16.

17.

Altonn, H. (1973) Sea t u r t l e i s s u e t o g e t f u r t h e r a i r i n g . HQ~OZUZU Star-Bulletin, 2 7 October, A:2.

Altonn, H. (1974) Sea t u r t l e p r o t e c t i o n code ready, Honolulu Star- Bulletin, 8 January, B:6.

Altonn, H. (1974) T u r t l e s t e a k n o t h i s d i s h . Honolulu Star-Bulletin,

Altonn, H. (1974) T u r t l e r u l e crawls forward. HonoZUlu Star-Bulletin,

28 February, D:6. I

9 March, A:3.

Altonn, H. (1974) Panel agrees on t u r t l e r u l e : HonoZuZu Sk-BulZetin, 6 A p r i l , A:3.

Altonn, H, (1974) Pryor has hopes f o r sea farm. Honolulu Star-Bulletin, 10 Apr i l , F : l .

Altonn, H. (1974) Land board a c t i o n may be chal lenged: S t a t e f a c e s l awsu i t on U-drive concessions. FlonoZuZu Star-Bulletin. 1 2 Apr i l , B:8.

Altonn, H. (1975) Transplanted t u r t l e s go home t o French F r i g a t e Shoals. Honolulu Star-Bulletin, 16 May, C:8.

Altonn, H. (1976) Grounded s h i p t h r e a t t o Kure A t o l l w i l d l i f e . Honolulu Star-Bulletin, 20 February, B : 2.

Altonn, H. (1976) Aquarium releases 7 t u r t l e s . Homluzu Star-Bullet+, 2 June, B:8. Reprinted i n 'Elepaio (1976) 37, 1:9-10.

54

18.

19.

20.

21 L

Altonn, H. (1976) Oahu f i s h farm g e t s S t a t e go-ahead. H o n i i Z U Z U Star- Bulletin, 29 December, C:5.

P

Altonn, €1. (1977) T u r t l e s and seals f i n d a home. Honolulu Star- Bulletin, 1 2 October, G : l .

Altonn, H. (1978). Bat te red vagrant g e t s a l i f t and f i n d s a home i n Hawaii. Honolulu Star-RulZetin, 10 March, A : l .

Altonn, H. (1978) Elephant s ea l , eag le come from f a r away: Unique v i s i t o r s a r r i v e by a i r , by sea. HonoZuZu Star-Bulletin, 15 March, C : l . Reprinted i n fie Islander, U. S. Naval S t a t i o n Midway (1978) 28 Apr i l , 1.

c

Altonn, H. (1978) H e vows f i g h t f o r w i l d l i f e . Honolulu Star-Bulletin, 6 A p r i l , A : 1 2 .

22.

23.

24.

Altonn, H. (1978) Aquarium green t u r t l e now l i v i n g f r e e i n n a t u r a l h a b i t a t . Honoluh Star-Bulletin, 8 J u l y , A:2.

Amerman, K. E. (1964) Leeward I s l a n d s survey: P e a r l and Hermes Reef, L i s i a n s k i I s l and , August 1964. Leeward survey no. 4.' P a c i f i c Ocean B io log ica l Survey Program, Smithsonian I n s t i t u t i o n , Washington, D. C. , unpublished r e p o r t , 11 pp.

25. Amerson, A. B., Jr. (1964) Northwest Hawaiian I s l a n d s t r i p r e p o r t , March 1964. Leeward survey no. 3. P a c i f i c Ocean B io log ica l Survey Program, Smithsonian I n s t i t u t i o n , Washington, D. C . , unpublished r e p o r t , 20 pp.

26. Amerson, A. B . , Jr. (1965) French F r i g a t e Shoals t r i p r e p o r t , August 1965. Leeward survey no. 10. P a c i f i c Ocean B io log ica l Survey Program, Smithsonian I n s t i t u t i o n , Washington, D. C. , unpublished r e p o r t , 48 pp.

27.

28.

Amerson, A . B . , Jr. (1966) French F r i g a t e Shoals t r i p r e p o r t , June 1966. Leeward survey no. 12. P a c i f i c Ocean B io log ica l Survey Program, Smithsonian I n s t i t u t i o n , Washington, D. C. , unpublished r e p o r t , 32 pp.

Amerson, A. B . , Jr , (1967) Leeward I s l a n d survey no. 20, French F r i g a t e Shoals , May-June 1967. P a c i f i c Ocean B io log ica l Survey Program, Smithsonian I n s t i t u t i o n , Washington, D. C. , unpublished r e p o r t , 26 pp.

29. Amerson, A. B . , Jr. (1968) Leeward I s l a n d s survey no. 23, French F r i g a t e Shoals , May-June 1968. P a c i f i c Ocean B io log ica l Survey Program, Smithsonian I n s t i t u t i o n , Washington, D. C., unpublished r e p o r t , 1 9 pp.

30. Amerson, A. B . , Jr. (1971) The n a t u r a l h i s t o r y of French F r i g a t e Shoals , Northwestern Hawaiian I s l ands . 15O:l-303.

A t o l l Reseamh Bulletin,

55

31.

'..

32.

33.

34.

35.

36.

37.

38.

39.

40.

41.

42.

43.

44.

45.

Amerson, A. B . , Jr. and P. C. Shel ton (1976) The n a t u r a l h i s t o r y of Johnston A t o l l , Cen t r a l P a c i f i c Ocean. A t o z z Research Bulletin, 192~1-479.

Amerson, A. B . , Jr . , Clapp, R. B. and W. 0. Wirtz 11. (1974) The n a t u r a l Atoll h i s t o r y of Pear l and H e r m e s Reef, Northwestern Hawaiian I s l ands .

Research Bulletin, 174 :1-306.

Apple, R. and P. Apple (1974) Lonely L i s i a n s k i i s le t . Honozuzu Star-Bulletin, 30 November, A:13.

Armstrong, D. (1973) E n t i r e community becomes a classroom. Honoiulu Star-Bulletin, 28 December, A : 1 2 .

Ashdown, I. (1976) Kahoolawe p l a c e names. P a c i f i c S c i e n t i f i c Information Center , Bernice P . Bishop Museum, Honolulu, unpublished r e p o r t , 3 pp.

Atkinson, A. L. C . and W . A. Bryan (1913) A rare seal. New York Zoological Soc ie ty B u l l e t i n , 16:1050-1051.

Bac l ig , A. e d i t o r (1975) Second s t r a i g h t year : Clutch of t u r t l e s ha t ch a t Punaluu Beach. Ka'u News (Hawaii), 24 September, 3.

Bai ley, A. M. (1918) The monk seal of t he Southern P a c i f i c . NaturaZ History, 18 : 396-399.

Bai ley, A. M. (1956) Bi rds of Midway and Laysan I s l ands . Museum P i c t o r i a l , Denver Museum of Natura l His tory , 130 pp.

Bai ley , A. M. and R. J . Niedrach (1951) Stepping s t o n e s ac ross t h e P a c i f i c . Museum P i c t o r i a l , Denver Museum of Natura l His tory , 63 pp.

Baker, A. S . (1922) Petroglyphs of Ka'u. In T. G. Thrum ( e d i t o r ) Hawaiian almanac and annual f o r 1922, Honolulu, 49-58.

Baker, A. S. (1967) More petroglyphs: Puuanahulu and Honokohau. In J. H. Cox ( e d i t o r ) Hawaiian petroglyph sources , 1825-1967. Univers i ty of H a w a i i , Hamilton Library , Hawaiian Co l l ec t ion .

Bakus, G . J. (1978) W i l d l i f e re fuges and endangered s p e c i e s of t h e Hawaiian I s l a n d s and t h e Trus t T e r r i t o r y of t h e P a c i f i c I s l a n d s . In L i t e r a t u r e review and s y n t h e s i s of information on P a c i f i c i s l a n d ecosystems. Documentation Assoc ia tes Information Se rv ices Incorpora ted , Los Angeles, Ca l i fo rn ia . U. S. F ish and W i l d l i f e Service c o n t r a c t number 14-16-009-77-075, Por t land , 1:l-37.

Balazs , G. H. (1973) The sea t u r t l e : Gent le c r e a t u r e f ac ing e x t i n c t i o n . Waikiki Aquarium p u b l i c a t i o n , l e a f l e t , 4 pp.

Balazs , G. H. (1973) Testimony concerning House B i l l No. 1635 which relates t o r e sea rch and management f o r t h e conserva t ion of sea t u r t l e s i n Hawaiian waters. Univers i ty of H a w a i i , H a w a i i I n s t i t u t e of Marine Biology, Kaneohe, 28 March 1973, unpublished r e p o r t , 3 pp.

56

46.

4 7 .

48.

49.

50.

51.

52.

53.

54.

55.

56.

57.

58.

59.

60.

Balazs, G . H . (1973) I t . ;timony presented on Silptember 21, 1973 concerning Regulation 36 which re la tes t o ~ \ i e p r o t e c t i o n of marine t u r t l e s i n H a w a i i . U n i v e r s i t y of H a w a i i , H t w a i i I n s t i t u t e of Marine Biology, Kaneohe, unpublished r e p o r t , 9 pp.

Balazs, G. H. (1973) Statti., of marine t u r t l e s in t h e Hawaiian I s l a n d s . The GZepa,?:o, 33, 12:l-5.

Balazs, G. H. (1973) A p a r t i c u l a r p o i n t of view: P r o t e c t i n g Hawaiian sea t u r t l e s . Hono Zulu Staz--RuZZetin, 18 December, A: 25.

Balazs, G. H . (1973) Summary r e p o r t on the 1973 French F r i g a t e Shoals ' g reen t u r t l e i n v e s t i g a t i o n s . Un ive r s i ty of H a w a i i , H a w a i i I n s t i t u t e of Marine Biology, Kaneohe, unpublished r e p o r t , December, 35 pp.

Balazs, G . H . (1974) Testimony concerning House B i l l No. 1635 which relates t o management s t u d i e s of t he H a w a i i a n green t u r t l e . Un ive r s i ty of H a w a i i , H a w a i i I n s t i t u t e of Marine Biology, Kaneohe, 26 February 1974, unpublished r e p o r t , 4 pp.

Balazs, G. H . (1974) Surv iva l s t a t u s of t h e green t u r t l e (Chelonia mydas) n e s t i n g and basking colony a t French F r i g a t e Shoals, Northwestern Hawaiian I s l a n d s . P r e s e n t a t i o n t o 104th Annual Meeting of t h e American F i s h e r i e s Soc ie ty , 1 0 September, Honolulu, unpublished r e p o r t , 8 pp.

Balazs, G. H. (1974) Green t u r t l e ' loophole ' c l a r i f i e d . The Honolulu Advertiser, 23 September, A: 7 .

Balazs, G. H. (1975) Letters t o t he e d i t o r : Good example. HornZulu Star-Bulletin, 1 2 February.

Balazs, G. H. (1975) The e d i t o r ' s mailbag: Green sea t u r t l e s . Pacifie IsZands MonthZy, March, 16.

Balazs, G. H . (1975) Green t u r t l e ' s u n c e r t a i n f u t u r e . Defenders, 50, 6:521-523.

Balazs , G . H . (1975) Testimony concerning House B i l l No. 191, and Senate B i l l No. 548 which relate t o a green sea t u r t l e r e source management study. Univers i ty of Hawaii, H a w a i i I n s t i t u t e of Marine Biology, unpublished r e p o r t , 2 pp.

Balazs, G. H. (1976) Green t u r t l e migra t ions i n t h e Hawaiian Archipelago. Biological Conservations, 9:125-140.

Balazs, G. H . (1976) Ex t inc t ion threa tened . The Sunday Star-BulZstin (f; Advertiser (Honolulu), 1 February, B:5.

Balazs, G. H. (1976) A p a r t i c u l a r p o i n t of view: The f a c t s on t u r t l e farms. HonoZuZu Star-Bulletin, 5 June, A: 11.

Balazs, G. H. (1976) Sea t u r t l e conserva t ion . 'EZepaio, 36, 7:79-85.

A

57

6 11.

62.

63.

64.

65.

66.

67.

68.

69.

"m.

70.

71.

72.

Bal .am, G. H. (1976) Hawaiian green ' t u r t l e (Chelonia mydas agass iz i ) and ghos t c rab (Ocypode ceratophthalmus) basking on t h e beach a t French F r i g a t e Shoals (photograph and c a p t i o n ) . HerpetoZogical Review, 7 , 1 : 36.

Balazs , G. H . (1976) H a w a i i ' s s e a b i r d s , t u r t l e s and seals. World Wide D i s t r i b u t o r s , Honolulu, 32 pp.

Balazs, G. H. (1977) Survey and assessment of the green sea t u r t l e resource of t h e Northwestern Hawaiian Archipelago. Sea Grant I n s t i t u t i o n a l Program, B ienn ia l Proposal Years 10-11 (1977-19791, Univers i ty of Hawaii , .Sea Grant College Program, B14-B19.

Balazs , G. H. (1977) Report on t h e experimental use of l a r g e mesh n e t s as a l i ve -cap tu re technique i n t h e r e sea rch of Hawaiian green t u r t l e s . Univers i ty of H a w a i i , H a w a i i I n s t i t u t e of Marine Biology, Kaneohe. Report prepared f o r t h e S t a t e of H a w a i i , Department of Land and Na tu ra l Resources, Div is ion of F ish and Game, 11 pp.

Balazs , G. H. (1977) Letters t o the e d i t o r : Live b a i t ? Surfer, 18, 2:29.

Balazs, G. H. (1977) Comments on Inconel t ags . IUCN/SSC Marine Turtle Newsletter, 2 : 7-8.

Balazs, G. H. (1977) S a l e of t u r t l e products promoted i n H a w a i i . IUCN/SSC Mzrine Turtle Newsletter, 4:4. Reprinted i n 'Elepaio (1977) 38, 1:11.

Balazs , G. H. (1977) South P a c i f i c Commission T u r t l e P r o j e c t : A c o n s t r u c t i v e review and eva lua t ion wi th recommendations f o r f u t u r e a c t i o n . Univers i ty of H a w a i i , H a w a i i I n s t i t u t e of Marine Biology, Kaneohe. Report prepared f o r t h e South P a c i f i c Commission (SPC), Noumea, New Caledonia, Apr i l , 54 pp.

Balazs , G. H. (1977) An i n v e s t i g a t i o n of t he growth and migra t ions of immature green t u r t l e s under n a t u r a l cond i t ions . H a w a i i , H a w a i i I n s t i t u t e of Marine Biology, Kaneohe. Research proposa l submit ted t o t h e World W i l d l i f e Fund, Washington, D. C.,

Univers i ty of

May, 1 3 PP.

Balazs , G. H . (1977) Analyses of t u r t l e p a r t s recovered from sha rk stomachs sampled a t P e a r l and H e r m e s Reef and French' F r i g a t e Shoals , Northwestern Hawaiian I s l a n d s . Univers i ty of H a w a i i , H a w a i i I n s t i t u t e of Marine Biology, Kaneohe, unpublished r e p o r t , June, 8 pp.

Balazs , G. H. (1977) MAC green t u r t l e p r o j e c t a c t iv i t i e s r epor t : J u l y 1976 t o Ju ly 1977. Univers i ty of H a w a i i , H a w a i i I n s t i t u t e of Marine Biology, Kaneohe. Report prepared f o r t h e S t a t e of H a w a i i , Of f i ce of t h e Marine A f f a i r s Coordinator , August, 75 pp.

Balazs , G. H. (1977) Ecologica l a s p e c t s of green t u r t l e s a t Necker I s l and . Univers i ty of H a w a i i , H a w a i i I n s t i t u t e of Marine Biology, Kaneohe, unpublished r e p o r t , October, 27 pp.

c

58

73. Balazs, G. H . (1978) Northwestern Hawaiian I s l a n d s f i s h e r i e s i n v e s t i g a t i o n s : Survey and assessment of t he green sea t u r t l e resource of t he Northwestern Hawaiian I s l ands . Sea Grant I n s t i t u t i o n a l Program Proposa l , Univers i ty of H a w a i i , Sea Grant Col lege Program, Year 11 (1978-1979), A43-A50.

74. Balazs , G. H. (1978) Northwestern Hawaiian I s l a n d s f i s h e r i e s i n v e s t i g a t i o n s : Survey and assessment of t h e green sea t u r t l e resource of t he Northwestern Hawaiian I s l a n d s . Sea Grant I n s t i t u t i o n a l Program Proposal , Univers i ty of H a w a i i , Sea Grant College Program, Years 12-13 (1979-1980), A50-A55.

75. Balazs, G. H . (1978) P r o j e c t p rog res s r e p o r t : Hawaiian green sea t u r t l e management s tudy , August 1977-April 1978. Un ive r s i ty of H a w a i i , H a w a i i I n s t i t u t e of Marine Biology, Kaneohe. Report prepared f o r t h e S t a t e of H a w a i i , Of f i ce of t h e Marine A f f a i r s Coordinator , A p r i l , 4 pp.

Balazs , G. H. (1978) A hawksbi l l t u r t l e i n Kaneohe Bay, Oahu. ‘EZepaio, 38, 11~128-129.

Balazs , G. H. (1978) Tat tooing green t u r t l e s . IUCN/SSC Murine Turt le Newsletter, 8: 3 .

76.

77.

78. Balazs , G. H. (1978) Terrestrial c r i t i c a l h a b i t a t f o r sea t u r t l e s under United S t a t e s j u r i s d i c t i o n i n t h e P a c i f i c reg ion . ’EZepuio, 39, 4~37-41 .

79.

80.

Balazs, G. H. (1978) A ca l l f o r observa t ions : T u r t l e s i n f i s h stomachs. ‘EZepaio, 39, 5:52.

c Balazs, G. H. (1978) Sea t u r t l e s of Kahoolawe I s l and : A pre l iminary survey. Univers i ty of H a w a i i , H a w a i i I n s t i t u t e of Marine Biology, Kaneohe, Report prepared f o r t h e U. S. Naval F a c i l i t i e s Engineer ing Command, P e a r l Harbor, November, 21 pp.

81.

82.

Balazs , G. H. (1979) Growth, food sources and migra t ions of immature Hawaiian Chelonia. IUCfl/SSC M&ne !l’urtZe News Zetter, 10 : 1-3.

Balazs , G. H. (1979) Marine b e n t h i c a l g a e c o l l e c t e d from Kure A t o l l , Mar0 Reef and Necker Bank, Northwestern Hawaiian I s l ands . ‘EZepuio, 39, 9 : 110-111,

83.

8 4 .

85.

86.

Balazs, G. H. (1979) Forum, t h e r e a d e r ’ s page: T u r t l e s t e a k s . HonoZuZu Star-BuZZetin, 5 March, A:13.

Y

Balazs , G. H. (1979) Letters t o the e d i t o r : P r o t e c t i n g t u r t l e s . The HonoZuZu Advertiser, 1 3 March, A : l l .

Balazs, G. H . (1979) Loggerhead t u r t l e recovered from a t i g e r shark a t Kure Ato l l . ‘Elepaio, 39, 12:145-147.

Balazs , G. H. (1980) F i e l d methods f o r sampling the d i e t a r y componen*s of green t u r t l e s , Chelonk mydas. Herpeto ZogicaZ Review, 17., 1:5-6.

59

87.

88.

89.

90.

91.

92.

93.

94.

95.

96.

Balazs , G. H. and E. Ross (1973) Reared i n c a p t i v i t y . InternatiomZ Turtle and Tortoise Soeiety Journal, 7 , 1:6-9, 33.

Balazs , G. H. and E. Ross (1974) Observat ions on t h e basking h a b i t i n t h e cap t ive j u v e n i l e P a c i f i c green t u r t l e . Copeia, 2:542-544. -

Balazs , G. H. and E . Ross (1974) Observat ions on t h e preemergence behavior of t h e green t u r t l e . Copeia, 4:986-988.

Balazs , G. H. and G. C . Whittow (1979) F i r s t record of a t i g e r sha rk observed feeding on a Hawaiian monk seal. 'Elepaio, 39, 9:107-109.

Balazs , G. H., Higa, L. H. and A . Baird (1979) Tide ca lendar : Sea- b i r d s of the Hawaiian I s l a n d s . Dill ingham Corporat ion, Honolulu.

Balcomb, K. C. (1966) Pre l iminary r e p o r t on L i s i a n s k i I s l and . P a c i f i c Ocean B io log ica l Survey Program, Smithsonian I n s t i t u t i o n , Washington, D. C . , unpublished r e p o r t , 26 pp.

Balcomb, K. C. (1966) Pre l iminary r e p o r t , Laysan I s l and , October 1966. P a c i f i c Ocean B io log ica l Survey Program, Smithsonian I n s t i t u t i o n , Washington, D. C . , unpublished r e p o r t , 1 3 pp.

Baldwin, H. S . (1978) Afoot & a f i e l d i n H a w a i i : T u r t l e man and a peek i n t o our fu tu re . Hawaii Tribune-HeraZd, 9 J u l y , 6.

Ball, S. C. (1923) F i e l d no te book taken dur ing the v i s i t of t h e Tanager Expedi t ion t o t h e Northwestern Hawaiian I s l a n d s i n A p r i l and May 1923. Bernice P. Bishop Museum, Honolulu.

Banko, W . E. (1965) T r i p r e p o r t , Hawaiian Leeward I s l a n d s survey, March 11-23, 1965. Leeward survey no. 8. P a c i f i c Ocean B io log ica l Survey Program, Smithsonian I n s t i t u t i o n , Washington, D. C., unpublished r e p o r t , 11 pp.

97. Banko, W. E. (1967) Endangered w i l d l i f e i n H a w a i i . The EZepaio, 27, 11:98-100.

98. Bardach, J. E. (1975) Trop ica l animal aquacul ture . Sea Grant I n s t i t u t i o n a l Program Proposal , Un ive r s i ty of H a w a i i , Sea Grant College Program, Year 08 (1975-1976), A3-A21.

99. Bardach, J. E. and P. He l f r i ch (1973) Tropica l animal aquacul ture . Sea Grant I n s t i t u t i o n a l Program Proposa l , Un ive r s i ty of H a w a i i , Sea Grant College Program, Year 06 (1973-1974), A3-Al5 .

100. Bardach, J. E. and P. He l f r i ch (1974) Tropica l animal aquacul ture . Sea Grant I n s t i t u t i o n a l Program Proposa l , Univers i ty of H a w a i i , Sea Grant College Program, Year 07 (1974-1975), A3-Al7.

101. Barr, R. (1972) Noise could hamper sea t u r t l e ha tch ing . H O n O z u z U Stm-Bulletin, 11 September, A: 22.

102. Bauer, E . A. (1972) Adventure o f f H a w a i i : Voyage of t h e Buttonwood. Outdoor Li fe , May, 57-58, 118-122.

60

103.

104.

105.

106.

107.

108.

109.

110.

111.

112.

113.

114.

115.

116.

117.

118.

119.

Bauer, E . A. (1972) Hawaiian I s l a n d s Nat iona l W i l d l i f e Refuge. Sea Frontiers, 18, 6:346-356.

Bayl i s , J. S . (1934) Cruise r e p o r t f o r t h e Itasca f o r t h e month of June 1934. Record Group 26, Nat iona l Archives, Washington, D. C., 6 PP*

Beaglehole, J. C . , e d i t o r (1967) The j o u r n a l s of Captain James Cook: The voyage of t h e Resolution and Discovery, 1776-1780. Socie ty , Cambridge, 1647 pp.

Hakluyt

Beckwith, M. W. (1970) Hawaiian mythology. Un ive r s i ty of H a w a i i . P re s s , Honolulu; 575 pp .

Benedict , A. A. and L. W. P o l l a r d (1972) Three classes of immuno- g lobu l ins found i n t h e sea t u r t l e , Chelonia mydas. Folia klierobwZogiea, 1 7 :75-78.

Benedict , A. A. and L. P o l l a r d (1977) t u r t l e immunoglobulins. In J. B. Solomon and J. D. Hortan ( e d i t o r s ) Developmental Immunobiology, Elsevier P r e s s , Amsterdam, 315-322.

Bennet t , W. C. (1931) Archaeology of Kauai Bernice P. Bishop Museum

The ontogeny and s t r u c t u r e a f sea

Bullet in 80, 156 pp.

Benson, B. (1973) No gunsmoke f o r Marshal l D i l lon - j u s t kangaroo. The HonoZulu Advertiser, 6 A p r i l .

Benson, B. (1973) Layson [ s ic ] I s l a n d t h e l a s t refuge. !The Honozuzu Advertiser, 8 Apr i l , H:4.

Benson, B. (1973) Ci ty l i m i t s a l s o would inc lude French F r i g a t e Shoals. The HonoZuZu Advertiser, 9 A p r i l , D:3.

Benson, B. (1973) Is le wi lderness proposa l t o g e t p u b l i c hear ing . The HonoZuZu Advertiser, 11 A p r i l , A : 1 7 .

Benson, B. (1973) Hearing ton igh t on r e g u l a t i o n s t o save dwindling t u r t l e s . The HonoZuZu Advertiser, 2 1 September , A: 5.

Benson, B. (1973) Rise i n tourism seen as p e r i l t o i s l e t u r t l e s . The Honolulu Advertiser, 22 September, B : l .

Benson, B. (1974) Tur t le -s teak ban begins tomorrow. The H O Y W ~ ~ U Advertiser, 29 May, A : 5 .

Benson, B. (1974) Leeward I s l a n d s debate: F ish ing r i g h t s snag sea wi lderness p lan . The Honolulu Advertiser, 14 June, A: 15.

Benson, B. (1974) Loophole p u t s sea t u r t l e i n t o t h e f r y i n g pan. The HonoZuZu Advertiser, 18 September, A: 3.

Benson, B. (1974) Words ' 'out of context" : T u r t l e farming n o t favored. The Honolulu Advertiser, 26 October, C:14.

c

c

c

c

61

120. Benson, B. (1976) Old law a new t h r e a t t o t u r t l e s . The HonoZuZu Advertiser, 16 January , A: 15.

121 . Benson, B. (1976) Commercial f i s h i n g urged i n w i l d l i f e haven. The Sunday Star-Bulletin & Advertiser (Honolulu), 9 May, B:9.

122. Benson, B. (1976) Sea refuge - a l l want t o g e t i n t o the a c t . The How Zulu Advertiser, 26 June, A: 8 .

123. Benson, B. (1976) Audubon: Don't cu t s i z e of w i l d l i f e refuge. The HotooZuZu Advertiser, 30 Ju ly , B:6.

124. Berger, A. J. (1970) The Leeward I s l ands . Oceans, 3:22-33.

125. Berger, A. J. (1972) Hawaiian b i r d l i f e . The Un ive r s i ty P r e s s of H a w a i i , Honolulu, 270 pp.

126. Blair, J, (1973) Sea t u r t l e s f a c e e x t i n c t i o n . lTke Leeward Cormnutity Co I lege NeGspaper, 9 : 3.

127. Boom, R. and C. Chris tensen (1978) Important Hawaiian p l a c e names. Bob Boom Books, Hilo, Hawaii., 47 pp.

128. Borreca, R. (1973) Buy goods, not land, McClung urges Japan. H o w ~ u L U Skrr-BtiZZetin, 31 J u l y , D:4.

Borreca, R. (1974) Isle seafood farm t o start. Honotulu Stcrr-BuZ&?t&z, 15 November.

129.

Bostwick, J. (1977) Dining cues: The Second Floor. fiono~u~uJ 12, 5:159-160.

130 ,

131.

132.

Bourke, R. E., Balazs, G . and E. W. Sha l l enbe rge r (1977) Breeding of t he green sea t u r t l e ( C h ? h & z mydas) a t Sea Life Park, Hawaii. Drwtr and Cmaker, October, 4-9.

Iloyer, D. R. (1965) Ecology of t h e bask ing h a b i t i n t u r t l e s . &obgy, 46:99-118.

Brock, V. E. (1951) Some obse rva t ions on the k y s a n duck, Anas uyo<Zl<ana laysavzensis. A d J 68, 3:371-372.

133.

134. Brock, J. A , , Nakamura, R. M . , Miyahara, A. Y. and E. N. L, Chang. (1976) T'rovzsacthns of the A m ~ b KsherCes Smtkf;y, 105, 6:564-566.

Tuberculosis i n P a c f f i c green sea t u r t l e s , G!h&w&z mgdas.

135. Brooks, M. C. ((1859) Cru i se of t h e G m b k k c i f i c ? Lbmmemekt ddt?ert;&er, 11 August .

136.

137-

Brooks, W. C. (1860) I s l ands and reefs West-Worth-West of the Sandwrch Islands, P a c i f i c , #uut&euu! J&gaaa&te, 29:499-504.

Bryan, E. H,, Jr. (1938) Andent Hawaiian llfe, AdvertLsec Pwkolishing conpay, Honolu1u, 113 pp.

138.

139.

140.

141 .

Bryan, E. H . , . J r . (1938) L i s i a n s k i , an i s l a n d of H a w a i i . Parachke of thc Paci f ic?, 50, 2:31-34.

Bryan, E . H . , Jr. (1938) Laysan, an i s l a n d of H a w a i i . Paradise of the Pacific, 5 0 , 5:21, 28-30.

Bryan, E. H . , J r . (1942) American Polynes ia and t h e Hawaiian cha in . Tongg Pub l i sh ing Company, Honolulu, 253 pp.

Bryan, E. H . , Jr. (1978) The Northwestern Hawaiian I s l a n d s . An annota ted b ib l iography. U . S . Fish and W i l d l i f e Se rv ice , Honolulu, 167 pp.

Bryan, W. A . (1911) Laysan I s l a n d : A v i s i t t o H a w a i i ’ s b i r d r e s e r v a t i o n . The M<d-Puc?i fi.c Magazine, 2 , 4 : 303-315.

c 1.42.

143.

144.

145.

146.

147.

Bryan, W. A . (1915) N a t u r a l h i s t o r y of H a w a i i . The Hawaiian Gazette Company, Ltd. , Honolulu, 596 pp.

Buck, P. H. , S i r (1964) A r t s and c r a f t s of H a w a i i : Food. Bernice 2’. Bishop Museum Special. PubZication 4 5 , 1-73.

c

Buck, E’. H . , S i r (1964) A r t s and c r a f t s of H a w a i i : F i sh ing . Rernice P. Bishop Musewn Special Publication 4 5 , 285-585.

Buck, P. H . , S i r (1964) A r t s and c r a f t s of H a w a i i : Death and b u r i a l . Bernice P. B.l:shop Museum Spcc?:aZ Publication 45 , 565-585.

Caldwell, D. K. (1962) Sea t u r t l e s i n Baja C a l i f o r n i a waters (with s p e c i a l r e f e rence t o those of t h e Gulf of C a l i f o r n i a ) , and t h e d e s c r i p t i o n of a new subspec ies of Nor theas te rn P a c i f i c green t u r t l e . ContAbutions i n Science, Los Angeles County Museum, 61:l-31. P

148. Caldwell, D. K. and M. C . Caldwell (1969) Sea t u r t l e s . In F. E. F i r t h ( e d i t o r ) The encyclopedia of marine resources . Van Nostrand Reinhold Company, New York, 740 pp.

149.

150.

151.

152.

153.

C a r l q u i s t , S. (1970) H a w a i i : A n a t u r a l h i s t o r y . The Na tu ra l His tory P res s , Garden C i ty , New York, 463 pp.

h

Carr, A . (1964) Transoceanic migra t ions of t h e green t u r t l e . Bioscience, 14 49-52.

Carr, A . (1975) The Ascension I s l a n d green t u r t l e colony. Copeia, 3:547-555.

P

Carr, A. (1977) The r e p t i l e s . Time-Life Books, Alexandria, V i rg in i a , 192 pp.

Carr, A. , Carr, M. H. and A. B. Meylan (1978) The ecology and migra t ions of sea t u r t l e s , 7 . The West Caribbean green t u r t l e colony. BuZZeth of the American Musewn of NaturaZ History, 162, 1:l-46.

63

154.

155.

156.

157.

158.

159.

160 i

161.

162.

163.

164.

165.

C a r r o l l , J. S., Representa t ive (1973) Requesting the people of Hawaii t o a i d i n t h e p re se rva t ion of t h e green sea t u r t l e which i s considered an endangered s p e c i e s throughout t he world. S t a t e of H a w a i i , House of Representa t ives , Seventh Leg i s l a tu re : House Concurrent Resolut ion No. 9 ; House Resolut ion No. 41.

C a r r o l l , J. S. , Representat ive (1973) Making an appropr i a t ion f o r a r e sea rch management s tudy of the popula t ion of t he green sea t u r t l e i n Hawaiian waters. State of H a w a i i , House of Representat ives , Seventh Leg i s l a tu re : A B i l l f o r an Act, House B i l l No. 245.

Ca r ro l l , J. S . , Representat ive (1975) Requesting the people of H a w a i i t o a i d i n the p re se rva t ion of t h e green sea t u r t l e which is considered an endangered spec ie s throughout t he world. S t a t e of H a w a i i , House of Representa t ives , Eighth Leg i s l a tu re , House Concurrent Resolut ion No. 16.

C a r r o l l , J. S . , Representa t ive (1975) Requesting a moratorium on the ca tch ing of green sea t u r t l e s . S t a t e of H a w a i i , House of Representa t ives , Eighth Leg i s l a tu re : House Resolut ion No. 538.

Carter, B. (1969) T a l l s t o r i e s froin fishermen. The Sunday Star - Bulletin & Advertiser (Honolulu), 2 November, D:6.

Chase, R. G. (1968) The growth rate and feeding e f f i c i e n c y of t he green sea t u r t l e (CheZonia m y d a s ) . Research proposal , Univers i ty of H a w a i i , Department of Oceanography, 9 pp.

Ch iav ie l lo , T. (1973) Fishermen d i s p u t e proposed r u l e s l i m i t i n g t u r t l e ca tch . The Garden IsZand. (Lihue, Kauai), 19 December, F:4.

Ching, H. and R. McCabe (1973) Divided viewpoints on marine t u r t l e con t ro l s . HonoZuZu Star-BuZZetin, 6 December, D:13 .

Chong, A., Representa t ive (1973) Green t u r t l e & r e l a t e d conserva t ion news. S t a t e of H a w a i i , House of Representa t ives , Newsletter No. 5 , 23 October, 6 pp.

Chong, A . , Representa t ive (1973) Making appropr i a t ions f o r r e sea rch management s t u d i e s of t he populat ion of t he green sea t u r t l e i n H a w a i i a n waters and s t e p s which may l ead t o the p re se rva t ion and i n c r e a s e of t h a t populat ion. State of H a w a i i , House of Representa- t i v e s , Seventh Leg i s l a tu re : A B i l l f o r an Act, House B i l l No. 1635.

Chong, A . , Representa t ive (1973) Requesting t h a t t he people of H a w a i i a i d i n h a l t i n g the d e c l i n e o f , and i f p o s s i b l e i n t h e r e s t o r a t i o n of , t h e green sea t u r t l e popula t ions which f r equen t t h e waters ad jacen t t o these Hawaiian I s l ands , and r eques t ing the Department of Land and Natura l Resources t o s tudy means of p r o t e c t i n g sea t u r t l e s and the f e a s i b i l i t y of r a i s i n g t u r t l e s commercially and t o r e p o r t i t s f ind ings t o the House of Representat ives . State of H a w a i i , House of Representa t ives , Seventh Leg i s l a tu re , House Resolut ion 303.

Chong, A., Representa t ive (1974) Green t u r t l e news le t t e r , Stare of H a w a i i , House of Representat ives , Newsletter No. 6, 22 January, 2 pp.

64

166. Clapp, R. B . (1968) Prel iminary report , Tern Island, French F r i g a t e Shoals , LS 22, 11-15 March 1968. P a c i f i c Ocean B io log ica l Survey Program, Smithsonian I n s t i t u t i o n , Washington, D. C . , unpublished r e p o r t , 9 pp.

167. Clapp, R. B. (1968) Leeward survey no. 22, prel iminary r e p o r t , P e a r l and Mermes Reef, 22-24 March 1968. P a c i f i c Ocean Bio logica l Survey Program, Smithsonian I n s t i t u t i o n , Washington, D. C . , unpublished r e p o r t , 20 pp.

168. Clapp, R. B. (1968) Leeward survey no. 22, p re l iminary r e p o r t , Nihoa I s l a n d . P a c i f i c Ocean B io log ica l Survey Program, Smithsonian I n s t i t u t i o n , unpublished r e p o r t , 9 pp.

c

169.

170.

.Clapp, R. B. (1972) The n a t u r a l h i s t o r y of Gardner P innac les , Northwestern Hawaiian I s l ands . AtoZZ Research BuZZetin, 163:l-29.

Clapp, R. B . and C. A. E l y (1967) Leeward survey no. 20, p re l iminary r e p o r t , P e a r l and H e r m e s Reef , 28-30 August 1967. P a c i f i c Ocean B io log ica l Survey Program, Smithsonian I n s t i t u t i o n , Washington, D. C . , unpublished rep.or t , 16 pp.

171. Clapp, R. B . and W. 0. Wirtz I1 (1975) The n a t u r a l h i s t o r y of L i s i a n s k i I s l and , Northwestern Hawaiian I s l ands . Atoll Research Bulletin, 186~1-196.

c

Clapp, R. B. and E. K r i d l e r (1977) The n a t u r a l h i s t o r y of Necker I s l and , Northwestern Hawaiian I s l ands . AtoZZ Research Bulletin, 206:l-102.

172.

173. Clapp, R. E . , K r i d l e r , E. and R. R. F l e e t (1977) The n a t u r a l h i s t o r y of Nihoa I s l and , Northwestern Hawaiian I s l ands . AtoZZ Research BuZletin, 207: 1-147.

174.

175.

176.

177.

Clark, H. (1975) Pryor r aps t a s k f o r c e on sea farm se tup . The Honozuzu Advertiser, 21 March, A:5.

Clark, J. R. K. (1977) The beaches of O'ahu. The Univers i ty P r e s s of H a w a i i , Honolulu, 193 pp.

Cobb, J. N. (1905) Commercial f i s h e r i e s of t he Hawaiian I s l a n d s i n 1903. U. S. Bureau of F i s h e r i e s , document no. 590:433-512.

Coleman, R. (1977) Nar ra t ive r e p o r t , NOAA vessel Townsend CromeZZ, 7/4/77-8/17/77. U. S. Fish and Wi ld l i f e Serv ice , Hawaiian I s l a n d s Nat iona l W i l d l i f e Refuge, Honolulu, unpublished r e p o r t , 10 pp.

178. Colum, P. (1924) A t t he gateways of the day: The S to ry of Kona, t he youth who could s t r e t c h himself upward; The daughter of t h e King of Ku-ai-he-lani. Oxford Univers i ty P r e s s , London, 217 pp.

Colum, P. (1966) Legends of H a w a i i : The daughter of t h e King of Ku-ai-he-lani. Yale Univers i ty P res s , London, 220 pp.

179.

180. Conant, S. (1977) Letter t o Di rec to r Lynn A. Greenwalt, U. S . F i sh and W i l d l i f e Service. 'EZepaio, 37. 11:134-135.

65

P

181.

182.

183.

184.

185.

186.

187.

188.

189.

190.

191.

192.

193.

194.

Conant, S. (1977) Letter t o Di rec to r Lynn Greenwalt, U. S. F ish and WiLdlife Serv ice . 'EZepaio, 37, 12:145-146.

Cooper, A . (1978) Pub l i c hear ing testimony, March 1 7 , 1978 - House C0nferenc.e Room 11330. Hawaii Fishing News, 2 , 4:19.

Corbin, J. S . (1976) Aquaculture i n H a w a i i 1976: Sea Grant College Program, H a w a i i I n s t i t u t e of Marine Biology. S t a t e of H a w a i i , Department of Planning and Economic Development, January, 107 pp.

Coul tas , W. F. (1936) Notes taken on Laysan I s l a n d , December 1936. U. S. F i sh and W i l d l i f e Serv ice , Bureau of Spor t F i s h e r i e s and Wi ld l i f e , Honolulu, unpublished, 1 p.

Cox, J. H. (1971) Resul t s of a pre l iminary i n v e s t i g a t i o n of t h e Anaehoomalu petroglyphs. In Barrera, W. P re l iminary r e p o r t o f s a lvage r e sea rch i n South Kohala, H a w a i i . Bernice Pauahi Bishop Museum, Department of Anthropology, Honolulu, 179 pp.

Cox, , J . H. and E. S tasack (1971) Hawaiian pe t roglyphs . Bernice P. Bishop Museum Spacial Publication 60, Honolulu, 100 pp.

Crossin, R. S. (1965) I s l a n d r e p o r t - J u l y 1965. P a c i f i c Ocean B io log ica l Survey Program, Smithsonian I n s t i t u t i o n , Washington, D. C. , unpublished r e p o r t , 14 pp.

Crossin, R. S. (1965) I s l a n d r e p o r t , August 1965. P a c i f i c Ocean B io log ica l Survey Program, Smithsonian I n s t i t u t i o n , unpublished r e p o r t , 16 pp.

Crossin, R. S. (1966) Leeward I s l a n d survey no. 13, June 1966. P a c i f i c Ocean B io log ica l Survey Program, Smithsonian I n s t i t u t i o n , unpublished r e p o r t , 18 pp.

C u r t i s , C. (1970) L i f e i n o l d H a w a i i : N a w a i t h e n e t maker. Kamehameha Schools P res s , Honolulu, 273 pp.

Daacon, G. (1977) S t a r spangled ban te r . The HonoZuZu Advertiser, 1 2 January, A : 3 .

Daacon, G. (1978) Monday melange. The HonoZulu Advertiser, 1 2 June.

D e Luca, C. , e d i t o r (1974) T u r t l e s seek new home. Lue Ahi (Waikiki Aquarium), 1, 1:l.

DeLong, R. L. and R. L. Brownell (1977) Hawaiian monk seal (&nachus schauinslandi) h a b i t a t and popula t ion survey i n the Northwestern (Leeward) Hawaiian I s l a n d s , A p r i l 1977. Nat iona l Marine F i s h e r i e s Serv ice , Northwest and Alaska F i s h e r i e s Center, S e a t t l e , Washington, processed r e p o r t , August, 1-30.

66

195. DeLong, R. L. , F i scus , C . H. and K. W. Kenyon (1976) Survey of monk seal (Monuchs s c h & z s l a n d i ) popula t ions of t h e Northwestern (Leeward) Hawaiian I s l ands : Prel iminary r epor t . Nat iona l Marine F i s h e r i e s Serv ice , Northwest F i s h e r i e s Center, Marine Mammal Divis ion, S e a t t l e , Washington, processed r e p o r t , May, 36 pp.

P

Dennon, G. B. , e d i t o r (1974) Two s i d e s t o the t m r t l e ques t ion . The Aquaculture Newsletter ( S e a t t l e ) , November, 2 .

196.

197.

198.

Dickey, D. R. (1923) Notes taken during t h e Tanager Expedition. Smithsonian I n s t i t u t i o n , Washington, D. C.

Diggs, J. T. (1918) Report of t h e c r u i s e of t h e U. S. S . Hemes among t h e i s l a n d s of t he Hawaiian group, 1918. Nat iona l Archives, 24 pp.

Record Group 45, c

199. D i l l , H , R , and W. A. Bryan (1912) Report on an exped i t ion t o Laysan I s l a n d i n 1911. U. S. Department of Agriculture Survey Bulletin, 42 : 1-30.

200.

201.

202.

203.

204.

205.

206.

Dodd, E. H. (1967) The r i n g of f i r e : Polynesian ar t . Dodd-Mead, New York, 368 pp.

Y

Dodd, C. K. (1978) Terrestrial c r i t i ca l h a b i t a t and marine t u r t l e s . Bullet in of the Maryland Herpetological SocCety, 1 4 , 4:233-240.

Donnelly, D. (1975) Dave Donnelly 's H a w a i i . Honolulu Star-Bulletin, 16 October, A t 4 .

Doty, M. S . and G. Aguilar-Santos (1966) Caulerp ic in , a t o x i c con- s t i t u e n t of Caulerpa. Nature, 211:990.

Eagle, L. (1954) Underwater exp lo re r s . The Honoluh Advertiser (Hawaiian L i f e Weekend Magazine), 31 October, 16-17.

Edmondson, C. H. (1946) Reef and shore fauna of H a w a i i . Bernice P. Bishop Musewn Special Publication 22.

Ehrenfeld, D. W. (1978) Commercial breeding i n c a p t i v e sea t u r t l e s : S t a t u s and prospec ts . 21st Annual Meeting of t h e Socie ty f o r t h e Study of Amphibians and R e p t i l e s , Arizona S t a t e Un ive r s i ty , Tempe, 11 pp.

207 E l b e r t , S. H., e d i t o r (1959) Se lec t ions from Fornander 's Hawaiian a n t i q u i t i e s and fo lk- lore : T rad i t ion of Kamapuaa. Univers i ty of H a w a i i P re s s , Honolulu, 297 pp.

E l i o t , J. L. (1978) H a w a i i ' s f a r - f lung w i l d l i f e paradise . Na~wnul Geographic, 153, 5: 670-691.

208.

209.

210.

E l l i o t , W. (1947) Kane i n t h e k i t chen . H a w i i f a r m CWLd home, 10, 9:51.

Elachner, C. (1915) The leeward i s l a n d s of t h e Hawaiian group. Reprinted from The Sunday Advertiser, Honolulu, 68 pp.

6

67

211.

P

212.

213.

214.

215.

216.

217.

218.

219.

220.

221.

222.

223. e

224.

225.

Ely, C , A. and R. B. Clapp (1973) The n a t u r a l h i s t o r y of Laysan I s l and , Northwestern Hawaiian I s l a n d s . Atoll Ressarch BuJZetin, 1 7 1 : 1-361.

Emory, K. P. (1928) Archaeology of Nihoa and Necker I s l ands . Bernice P. Bishop Musewn Bulletin, 53: 1-124.

Emory, K. P. (1969) The I s l and of Lanai: A survey of n a t i v e c u l t u r e . Bernice P. Bishop Musewn Bulletin 12, Honolulu, 129 pp.

Emory, K. P., Bonk, W. J. and Y . H. S inoto (1968) Fishhooks. Bernice P. Bishop Musewn Speciul Publication 47, 62 pp.

Engle, M. (1974) Looking through ... our 'window on t h e sea'. !The Sunday Star-Bulletin & Advertiser (Honolulu), 13 October, C:4.

Engle, M. (1978) Coming up: Today. The Sunday Star-Bulletin & Advertiser (Honolulu), 23 Apr i l , C : l l .

E rns t , C. H. and R. W. Barbour (1972) T u r t l e s of t h e United S t a t e s . The Univers i ty Press of Kentucky, Lexington, 347 pp.

Evans, L. (1973) Proposals t o save our t u r t l e s . Kapiolani Colrprmnity College Newspaper, 2 October.

Evans, L. R. (1975) Kaula: Forgot ten b i r d i s l a n d a l s o bomb t a r g e t . The Sunday Star-Bulletin & Advertiser (Honolulu), 16 November, F : l , 6. Reprinted i n 'Elepaio (1976) 27, 6:61-63.

F a r r e l l , A . , e d i t o r (1928) John Cameron's odyssey. The Macmillan Company, New York, 461 pp.

F inner ty , A. F. J. (1974) Food f o r thought: Business i d e a - raise eel o r t u r t l e s f o r expor t? The Garden Island (Lihue, Kauai), 1 7 Apr i l , B:7.

F iscus , C. H. (1977) Voyage t o Laysan I s l and , February-March 1977, a p a r t of t he j o i n t NMFS-FWS 1977 monk seal (Monuchus schauinstandi) r e s e a r c h program. Appendix I t o DeLong and Brownell (1977). Nat iona l Marine F i s h e r i e s Serv ice , Northwest and A l a s k a F i s h e r i e s Center , S e a t t l e , Washington, processed r e p o r t , 31-43.

F l e e t , R. (1964) Leeward I s l a n d s survey no. 5, September 1964. P a c i f i c Ocean B io log ica l Survey Program, Smithsonian I n s t i t u t i o n , Washington, D. C. , unpublished r e p o r t , 1 5 pp.

Fornander, A. (1917) Fornander c o l l e c t i o n of Hawaiian a n t i q u i t i e s and fo lk- lore : Memoirs of the Bernice P. Bishop Museum, 4, 436-449.

Legend of Kona and Niheu (Rela t ing t o Haupu H i l l ) .

Fornander, A. (1917) Fornander c o l l e c t i o n of Hawaiian a n t i q u i t i e s and fo lk - lo re : Legend of Laukiamanuikahiki. Memoirs of the Bernice P. Bishop Museum, 4, 3:596-608.

68

226. Farnnnder, A . (1919) Fornander c o l l e c t i o n of Hawaiian a n t i q u i t i e s and f o l k-lore: Myth concerning Molokini. Memoirs of the Bernice P , 87:Rh/)r> M~.q(<2071, 5, 3 : 514-521 .

229 ‘

228.

Famander , A . (1920) Fornander c o l l e c t t o n of Hawaiian a n t i q u i t i e s and fo lk- lore : Prayer of Malaehaakoa. Memoirs of the Bernice P. R7:shoy M1A.’:n7Am, 6, 3 : 492-498.

Fo r tne r , H. .I. (1978) The l i m u e a t e r : A cookbook of Hawaiian seaweed. S r a Grant MJ scel laneoris Report UNIHI-SEAGRANT-MR-79-01, November, 1 I) 7 p p .

c

F r a z i e r , T, (1355) T u r t l e s o u p . HonoluZu Stnr-Bulletin (Hawaiian L i f e Weekend Magazine), 16 December, 7.

229.

230.

A

Ftijimoto, M. M. and H. M. Sakrida (1972) The 1971 s h a r k c o n t r o l and r e s e a r c h program f i n a l r e p o r t . S t a t e of H a w a i i , Department of Land and N a t i i r n l Resources, Div is ion of F ish and Game , 37 pp.

231. G a l t s o f f , P. S . (1.933) P e a r l and Hermes Reef, H a w a i i hydrographical and b i o l o g i c a l observa t ions . Bernice P. Bishop Musewn Bulletin, 107 : 1-49.

c

232.

233.

234.

235.

236.

237,

Cauthey, J . R , (1968) F i e l d t r i p t o Ulupau Head, A p r i l 21, 1968. The EZepaia, 28, 1 2 : 109.

Gee, H . (1974) S t a t e se t t les t u r t l e s h e l l f raud case. Honolulu Star- Bu%Zd&, 2 1 November, A : I , 9.

P

Given, . l . (1976) 4 g i a n t t u r t l e s sea-soned. The Honoluh Advertiser, 10 November, A : 3 .

Goodwin, H. A . (1972) Endangered and e x t i n c t w i l d l i f e of H a w a i i . ?%he Elepab, 3 3 , 2 : 14-15.

c

Goodwin, H. A. (1972) The endangered s p e c i e s conservat ion program f o r Hawaii. The Elepaio, 33, 3:23-27.

Gordon, J. A. (1970) An annotated c h e c k l i s t of Hawaiian barnac les ( c l a s s Crustacea; s u b c l a s s C i r r i p e d i a ) wi th n o t e s on t h e i r nomen- c l a t u r e , h a b i t s and Hawaiian l o c a l i t i e s , Hawa-ti t n s t l t u t e o f FIar-Fne Biology, Technical Report No. 19 , 130 pp.

Univers i ty of H a w a i i ,

238.

239.

Grant, C. (1927) Note on sea t u r t l e s . Copeia, 164:69.

Green, L. C. S. (1926) Folk- ta les from H a w a i i : The shark-gods of Kau. Vassar College, Poughkeepsie, New York, 126 pp.

240. Grigg, R. W. (1978) Northwestern Hawaiian I s l a n d s F i s h e r i e s Inves t iga- t i o n s : Reef and prec ious c o r a l resources - comparative ecology w i t h i n the Hawaiian Archipelago. UniverRity of Hawaii, Sea Grant College Program, Year 11 (1978-1979),

Sea Grant I n s t i t u t i o n a l Program Proposal ,

A.32-A37.

c

69

241.

242

243.

244.

245.

246.

247.

248.

249.

250.

251.

Grigg, R. W. (1978) Northwestern Hawaiian I s l a n d s F i s h e r i e s Inves t iga t ions : Archipelago. of H a w a i i , Sea Grant Col lege Program, Years 12-13 A30-A37.

Reef and s h e l f ben th ic ecology of t h e Hawaiian

(1979-1981), Sea Grant I n s t i t u t i o n a l Program Proposal , Univers i ty

Guest, S . (1974) Excerpts : Minutes of H a w a i i Audubon Socie ty genera l meeting, 17 September 1973. The EZepaio, 34, 7:82.

Guest, S. (1974) Excerpts from minutes of H a w a i i Audubon Socie ty gene ra l meeting, 15 October 1973. n e Elepaio, 34, 9:107.

Gutmanis, J. (1977) Kahuna l a ' a u lapa 'au : The p r a c t i c e of Hawaiian h e r b a l medicine. Trans la ted by Theodore Kelsey. I s l a n d Her i tage , Norfolk I s l a n d , A u s t r a l i a , 267 pp.

Hackman, C. D, (1967) Pre l iminary r e p o r t of Leeward I s l a n d survey no. 18, March 6 t o March 27, 1967. P a c i f i c Ocean B io log ica l Survey Program, Smithsonian I n s t i t u t i o n , Washington, D. C., unpublished r e p o r t , 12 pp.

Hammel, F. and S. Levey (1975-1976) H a w a i i on $15 a day: Molokai. Arthur Frommer Pub l i ca t ion , New York.

Handy, E. S. and M. Pukui (1972) The Polynesian family system i n Ka'u H a w a i i . Char les E. T u t t l e Company, Rutland, Vermont, 259 pp.

Handy, E. S. C. , Pukui, M. K. and K. Livermore (1934) Ou t l ine of Hawaiian phys ica l t he rapeu t i c s . 126, 51 pp.

Bernice P. Bishop Musewn Bulletin

Handy, E. S. C., Handy, E. G. and M. K. Pukui (1972) Native p l a n t e r s i n o l d H a w a i i : The i r l i f e , l o r e and environment. Bernice P. B i s h o p Musewn BuZZetin 233, 641 pp.

Harr ington, B. A. (1966) French F r i g a t e Shoals t r i p r e p o r t , August- September 1966. Leeward survey no. 15. P a c i f i c Ocean B io log ica l Survey Program, Smithsonian I n s t i t u t i o n , Washington, D. C., unpublished r e p o r t , 30 pp.

Harris J. (1977) I n harmony wi th t h e sea: The p r e s e n t day kapu. f i e Garden IsZand (Lihue, Kauai), 16 December, A:8 .

252. Harris, J. (1977) I n harmony w i t h t h e sea: Today's f i s h i n g r e g s r e f l e c t o l d Haw'n kapu system. 23 December, B:3.

The Garden Island (Lihue, Kauai),

253. Harris, J. (1977) H a w a i i i n harmony wi th t h e sea: I l l e g a l f i s h i n g techniques. Univers i ty of H a w a i i , Sea Grant Advisory Pamphlet, UNIHI-SEAGRANT-AB-77-05. (Lihue, Kauai), (1978) 1 3 January, 9 .

Reprinted i n p a r t i n The Garden Ishnd

70

254.

255.

256.

257.

258.

259.

260.

261.

262.

263.

264.

265.

266.

267.

Harris, J. (1979) H a w a i i i n harmony wi th t h e sea: I ' a of H a w a i i , present-day kapu. Univers i ty of H a w a i i , Sea Grant Advisory Brochure , UNIHI-SEAGRANT-AB-79-04.

Harshbarger, J. C . (no da te ) A c t i v i t i e s r epor t : Regis t ry of tumors i n lower animals , 1965-1973. Nat ional Museum of Natura l His tory , Smithsonian I n s t i t u t i o n , Washington, D. C . , 141 pp.

Harshbarger, J. C. (no da te ) A c t i v i t i e s r epor t : Regis t ry of tumors i n lower animals, 1975 supplement. Nat iona l Museum of Natura l His tory , Smithsonian I n s t i t u t i o n , Washington, D. C . , c o n t r a c t no. N01-CB-33874, 42 pp.

Harshbarger, J. C. (no da t e ) A c t i v i t i e s r e p o r t : Regis t ry of tumors i n lower animals , 1977 supplement. Nat iona l Museum of Nat iona l His tory , Smithsonian I n s t i t u t i o n , Washington, D. C. , N I H c o n t r a c t no. N01-CB-33874, 44 pp.

c

x

Haugen, K. (1974) Af t e r l e a r n i n g i t ' s i l l e g a l : S to re p u l l s t u r t l e jewelry. Honolulu Star-Bulletin, 18 September, A c 6 .

L

Haugen, K. (1974) Saves wi ld animals , he says: T u r t l e ' 'farmer" exp la ins h i s s i d e . Honolulu Star-Bulletin, 24 September , B: 2.

Hendricks, P. (1978) H a w a i i ' s Leeward I s l ands : Leeward I s l and r e sea rch p l an approved. West Hawaii Today, 30 June, 32.

Hendricks, P. (1978) T u r t l e s i g h t i n g s , f i s h ca tch r e p o r t s h e l p f u l t o s t u d i e s . West Hawaii Today, 7 J u l y , 14.

Hendrickson, J. R. (1969) Report on Hawaiian marine t u r t l e populat ions. Proceedings of t he Working Meeting of Marine T u r t l e S p e c i a l i s t s , IUCN Publicatwns New Series, 20:89-95.

H i l l , B. H. (1973) F i r s t b i e n n i a l r e p o r t : Marine Option Program. Univers i ty of H a w a i i , Sea Grant Miscellaneous Report UNIHI- SEAGRANT-MS-73-02, June, 64 pp.

H i r t h , H. F. (1971) South P a c i f i c i s l a n d s - marine t u r t l e resources . A r e p o r t prepared f o r t h e F i s h e r i e s Development Agency P r o j e c t . Food and Agr i cu l tu re Organizat ion of t h e United Nat ions, Rome, 34 pp.

H i r th , HI. F. (1971) Synopsis of b i o l o g i c a l d a t a on the green t u r t l e Chelonia mydas (Linnaeus) 1758. FA0 Fisheries Synopsis No. 85, 1:1-8:19.

H i r th , H. F. (1978) A model of t he evo lu t ion of green t u r t l e (CheZonia m y d a s ) remigra t ions . Herpetologica, 34 , 2: 141-147.

H i r t h , H. F. and A. Carr (1970) The green t u r t l e i n t h e Gulf of Aden and t h e Seychel les I s l ands . NederZandse Akademie van Wetenschappen, A f d . Natuurkunde Tweede Reeks, 58:l-44.

verjiande Zingen der KoninkZyke c

71

268.

269.

270.

c

271.

272.

273,

274.

275.

276.

277.

278.

279.

280.

281.

282.

283.

284.

Hochachka, P. W., Owens, T. G. , Al len, J. F. and G. C. Whittow (1975) Mul t ip l e end products of anae rob ios i s i n d i v i n g v e r t e b r a t e s . Comparative Biochemistry and Physiology, 50B:17-22.

Hodge, P. (1976) Green sea t u r t l e . 'Elepaio, 36, 8:101,

Hollenberg, G. J. (1968) An account of t he s p e c i e s of t h e r e d a l g a Polysiphonia of t h e Cen t ra l and Western P a c i f i c Ocean. Polysiphonia. Pacific Science, 22, 2~198-207.

11.

Hornel l , J. (1934) Log of t h e schooner A d a on a f i s h i n g c r u i s e i n t h e North P a c i f i c , 1882. The Mariner's Mirror, 20:426-437.

Hosaka, E. Y. (1944) Spor t f i s h i n g i n H a w a i i . Bond's, Honolulu, 198 pp.

Hosaka, E. Y. (1973) Shore f i s h i n g i n H a w a i i . Petroglyph P r e s s , Ltd. , Hi lo , H a w a i i , 176 pp.

Hylton, R. D., e d i t o r (1976) Keep t u r t l e s ou t of soup. Kind, November, 12-13.

Ikehara , I. I. (1961) B i l l y Weaver sha rk r e sea rch and c o n t r o l program: F i n a l r e p o r t , S t a t e of H a w a i i , Department of Agr i cu l tu re and Conservation, Div is ion of F i sh and G a m e , 20 Apr i l , 25 pp.

Imbleau, L. L. (1979) The Northwestern Hawaiian I s l ands : Economic ga in o r environmental l o s s ? Student Symposium on Marine A f f a i r s , UNIHI-SEAGRANT-MR-79-02, 207-213.

Proceedings of t he Fourth Annual

Inouye, D. K., Senator (1978) Northwest Hawaiian I s l ands . Honolulu Star-Bulletin, 31 J u l y , A: 15.

Johnson, W. A. (1976) Letters t o t h e e d i t o r : T u r t l e s . Honolulu StaP- BulZetin, 18 August

Judd, H. P . , Pukui, M. K. and J. F. G . Stokes (1977) I n t r o d u c t i o n t o t h e Hawaiian language. Tongg Publ i sh ing Company, Honolulu, 314 pp.

Kadooka, L. (1975) Returning t o t h e n e s t . Hawaii !7!dmne-Hemld, 9 Ju ly , 8.

Kahaulel io , A. D. (1902) F ish ing Lore. Trans la ted by M. K. Pukui from K a Nu epa Kuokoa. Marine Bioyogy , l i b r a r y manuscript , 89 pp . Elepaio, 29, 11:102-103.

Elepaio, 30, 12:116-117.

Un ive r s i ty of H a w a i i , Hawaii I n s t i t u t e of

Kaigler , H. (1968) F i e l d t r i p 13 A p r i l 1969 t o Ulupau Head. 2%

Kaigler , C. G. (1970) F i e l d t r i p t o Ulupau Head, 1 2 A p r i l 1970. The

Kakesako, G. K. (1978) S t a t e f i g h t s takeover of shoa l s , isles. The Honolulu Star-Bulletin, 6 January, A: 2.

72

285.

286.

287.

288.

289.

290.

291.

292.

293,

294 a

295

296.

Kalakaua, D. , King (1888) The legends and myths of H a w a i i . C. L . Webster and Company, New York, 530 pp.

Kam, A . K. and G . H. Balazs. (1978) A survey of green t u r t l e s a t select l o c a t i o n s i n the Hawaiian Archipelago. Report prepared f o r the Marine Option Program, Univers i ty of H a w a i i , January, 21 pp.

Kemble, J. H . , e d i t o r (1966) To C a l i f o r n i a and t h e South Seas: The d i a r y of Alber t G. Osbun, 1849-1851. The Huntington Library , San Marirao, C a l i f o r n i a , 233 pp.

Kenyon, K. W . and D. W. Rice (1959) L i f e h i s t o r y of t h e Hawaiian monk seal. Pacif ic Science, 1 3 , 3:215-252.

Kido, S . , Chairman (1973) Notice of pub l i c hear ing: Regulat ion 36 r e l a t i n g t o t h e p r o t e c t i o n of marine t u r t l e s . S t a t e of H a w a i i , Board of Land and Natura l Resources, Th.e Maui News, 18 August, C : 4 .

Kido, S. Chairman (1973) Notice of p u b l i c hear ing: Regulat ion 36 r e l a t i n g t o the p r o t e c t i o n of marine t u r t l e s . S t a t e of H a w a i i , Board of Land and Na tu ra l Resources, The Sunday Star-Bulletin 13 Adver-i;iser (Honolulu), 19 August, A : 1 8 .

Kido, S., Chairman (1973) Notice of p u b l i c hear ing: Regulat ion 36 r e l a t i n g t o the p r o t e c t i o n of marine t u r t l e s . State of H a w a i i , Board of Land and Natura l Resources, The Garden Island (Lihue, Kauai) , 20 August, 11.

Kido, S . , Chairman (1973) Notice of pub l i c hear ing: Regulat ion 36 r e l a t i n g t o the p r o t e c t i o n of marine t u r t l e s . S t a t e of H a w a i i , Board of Land and Na tu ra l Resources, H a w a i i Tribune-Herald, 20 August, 7.

Kido, S., Chairman (1973) Notice of pub l i c hear ing: Regulat ion 36 r e l a t i n g t o the p r o t e c t i o n of marine t u r t l e s . S t a t e of H a w a i i , Board of Land and Na tu ra l Resources, Hawaii Tribune-Herald, 9 November, 3.

kid^, S . , Chairman (1973) Notice of p u b l i c hear ing: Regulat ion 36 r e l a t i n g t o the p r o t e c t i o n of marine t u r t l e s . S t a t e of H a w a i i , Board of Land and Natura l Resources, HonoZuZu Stm-Bulletin, 9 November, D : 8 .

Kido, S . , Chairman (1973) Notice of p u b l i c hear ing: Regulat ion 36 r e l a t i n g t o the p r o t e c t i o n of marine t u r t l e s . S t a t e of H a w a i i , Board of Land and Natura l Resources, The Maui News, 10 November, B:9.

Kido, S . , Chairman (1973) Notice of p u b l i c hear ing: Regulat ion 36 r e l a t i n g t o t h e p r o t e c t i o n of marine t u r t l e s . Board of Land and Natura l Resources, The Garden Island (Lihue, Kauai), 1 2 November, 10.

S t a t e of H a w a i i ,

c

c

c

e-

a

73

297. Y

298.

299.

300.

301.

302.

303.

304.

305 .,

306.

307.

308.

309.

Kido, S., Chairman (1974) Notice of pub l i c hearing: Regulation 36 r e l a t i n g t o t h e p r o t e c t i o n of marine t u r t l e s . State of H a w a i i , Board of Land and Natural Resources, The &ui News, 5 February.

Kido, S., Chairman (1974) Notice of pub l i c hear ing: Regulation 36 r e l a t i n g t o the p r o t e c t i o n of marine t u r t l e s . S t a t e of H a w a i i , Board of Land and Natura l Resources, Honolulu Star-Bulletin, 5 February, D:6.

K i m , M. (1977) Sea t u r t l e s . Marine Aquarim Observer (The Sa l twater Aquarium Society of Hawaii), 8:lO-13.

KimBrer, W. J. and W. W. Durbin, Jr. (1975) The p o t e n t i a l f o r a d d i t i o n a l marine conservat ion d i s t r i c t s on Oahu and H a w a i i . Univers i ty of Hawaii, Sea Grant Technical Report, UNIHI-SEAGWT- TR-76-03, December, 108 pp.

King, F, (1975) More of Francee 's d in ing out : I l i k a i Hotel . The Sunday Star-Bulletin & Advertiser (Honolulu), Dining Out ( adve r t i s ing supplement) 28 December.

King, F. (1976) A min i - ed i to r i a l : On t h e s u b j e c t df t u r t l e s . The Swzday Star-Bulletin & Advertiser (Honolulu), Dining,Out ( a d v e r t i s i n g supplement), 18 January, 10.

King, F. (1979) L i t t l e George's is bigger and b e t t e r . The Sunday StQr-BuZZet5n & Advertiser (Honolulu), Dining Out ( adve r t i s ing supplement), 18 February, 1, 3.

King, F, W. (1975) Research r e p o r t s . Animal King&m, February- March, 29.

King, J. , Senator (1975) Making appropr i a t ions f o r a management s tudy of t he green sea t u r t l e resource i n the Hawaiian Archipelago. State of H a w a i i , The Senate , Eighth Leg i s l a tu re : A B i l l f o r an A c t , Senate B i l l No. 548.

King, W. B. (1973) Conservation status of b i r d s of Cent ra l P a c i f i c i s l ands . The Wilson Bulletin, 85, 1:89-103. Reprinted i n t h r e e i n s t a l l m e n t s i n The Elepaio (1973) 34, 6:58-62; (1974) 34,7:70-74; (1974) 34, 8:92-94.

Kitney, C. (1976) A memorable day i n H a w a i i . 'Elepaio, 36, 11:140-141.

Kramer, R. J. (1962) A r e p o r t on a survey t r i p t o t h e Hawaiian I s l ands Nat iona l W i l d l i f e Refuge, June 1962. S t a t e of H a w a i i , Department of Land and Natura l Resources, Divis ion of F ish and Game, unpublished r e p o r t , 18 pp.

Kramer, R. J. (1963) A r e p o r t on a survey t r i p t o the Hawaiian I s l ands Nat ional W i l d l i f e Refuge, February 1963. S t a t e of H a w a i i , Department of Land and Natura l Resources, Divis ion of F ish and Game , unpublished r e p o r t , 10 pp.

74

310.

311.

312.

313.

314.

315.

316.

317.

318.

319.

320.

321.

322.

Kramer, R. J . and J. W . Beardsley (1962) A r e p o r t on a survey t c i p , and an entomological r e p o r t t o Hawaiian I s l a n d s Nat iona l W i l d l i f e Refuge, June 1962. S t a t e of H a w a i i , Department of Land and Natural Resources, Divis ion of Fish and G a m e , unpublished r e p o r t , 21 pp.

K r i d l e r , E. (1964) Hawaiian I s l a n d s Nat iona l W i l d l i f e Refuge, J u l y 25-27, 1964. U. S . F ish and W i l d l i f e Service, Bureau of Spor t F i s h e r i e s and W i l d l i f e , unpublished r e p o r t , 8 pp.

I”’

K r i d l e r , E . (1964) Hawaiian I s l a n d s Nat iona l W i l d l i f e Refuge, September 16-27, 1964. U. S. F i s h and W i l d l i f e Serv ice , Bureau of Spor t F i s h e r i e s and W i l d l i f e , unpublished r e p o r t , 8 pp.

K r i d l e r , E . (1965) Hawaiian Islands Nat iona l W i l d l i f e Refuge, March 11-23, 1965. U. S. F i s h and W i l d l i f e Serv ice , Bureau of Spor t F i s h e r i e s and W i l d l i f e , unpublished r e p o r t , 20 pp.

K r i d l e r , E. (1966) Hawaiian I s l a n d s Nat iona l W i l d l i f e Refuge t r i p r e p o r t , March 17-April 16 , 1966. U. S . F i s h and W i l d l i f e Se rv ice , Bureau of Spor t F i s h e r i e s and W i l d l i f e , unpublished r e p o r t , 24 pp. c

K r i d l e r , E. (1966) Pre l iminary r e p o r t - French F r i g a t e Shoals, Hawaiian I s l a n d s Nat iona l W i l d l i f e Refuge, H a w a i i , March 21-24, 1966. U. S. F i s h and W i l d l i f e Service, Bureau of Sport F i s h e r i e s and W i l d l i f e , unpublished r e p o r t , 8 pp.

Kridler, E . (1966) Hawaiian I s l a n d s Nat iona l W i l d l i f e Refuge t r i p , September 8-28, 1966. U. S . F i sh and W i l d l i f e Serv ice , Bureau of Spor t F i s h e r i e s and W i l d l i f e , unpublished r e p o r t , 34 pp.

K r i d l e r , E. (1967) French F r i g a t e Shoals t r i p r e p o r t , March 1967. U . S . F i s h and W i l d l i f e Serv ice , Bureau of Sport F i s h e r i e s and W i l d l i f e , a

unpublished r e p o r t .

Kridler, E. (1967) Refuge l o g book f o r 6 March-1 A p r i l 1967. U.S. F i s h and W i l d l i f e Se rv ice , Bureau of S p o r t F i s h e r i e s and W i l d l i f e , unpublished r e p o r t , 39 pp.

K r i d l e r , E . (1967) Refuge log book f o r 19-29 September 1967. U. S . F i s h and W i l d l i f e Serv ice , Bureau of Spor t F i s h e r i e s and Wildlife, unpublished r e p o r t , 12 pp.

K r i d l e r , E . (1967) Hawaiian I s l a n d s Nat iona l W i l d l i f e Refuge, pre l iminary t r i p r e p o r t , December 7-18, 1967. U. S. F i s h and W i l d l i f e Service, Bureau of S p o r t F i s h e r i e s and W i l d l i f e , unpublished r e p o r t , 13 PP.

K r i d l e r , E. (1968) HawaiianIslands Nat iona l W i l d l i f e Refuge survey, March 1968. U. S. F i s h and W i l d l i f e Se rv ice , Bureau of S p o r t F i s h e r i e s and W i l d l i f e , unpublished f i e l d n o t e s .

K r i d l e r , E. (1969) Hawaiian I s l a n d s Nat ional g i l d l i f e Refuge s p r i n g t r i p , March 19-April 6 , 1969. U. S. F i s h and W i l d l i f e Service, Bureau of Sport F i s h e r i e s and W i l d l i f e , unpubldshed r e p o r t , 32 pp.

75

Y

323. K r i d l e r , E. (1969) Hawaiian I s l a n d s Nat iona l W i l d l i f e Refuge f i e l d t r i p , August 19-SeptembeK 23, 1969. U. S. F ish and W i l d l i f e Se rv ice , Bureau of Spor t F i s h e r i e s and W i l d l i f e , unpublished r e p o r t , 32 pp.

324. K r i d l e r , E. (1969) Report of w i l d l i f e management study. Popula t ions and movement of t h e green s e a t u r t l e on t h e Hawaiian I s l a n d s Na t iona l W i l d l i f e Refuge, August 31, 1969. U . S . F i s h and W i l d l i f e Se rv ice , Bureau of Spor t F i s h e r i e s and W i l d l i f e , unpublished r e p o r t . 6 pp.

325. K r i d l e r , E ? (1969) Hawaiian I s l ands Nat iona l W i l d l i f e Refuge exped i t ion repor t , November 9-13, 1969. U . S. F i sh and W i l d l i f e Serv ice , Bureau of Spor t F i s h e r i e s and W i l d l i f e , unpublished *

r e p o r t

,'-

e

326. K r i d l e r , E. (1971) Hawaiian I s l a n d s Na t iona l W i l d l i f e Refuge t r i p r e p o r t , September 2-17, 1971. U. S . F i s h and W i l d l i f e Se rv ice , Bureau of Spor t F i s h e r i e s and W i l d l i f e , unpublished r e p o r t , 30 pp.

327. Kr id l e r , E. (1972) Expedition r e p o r t , Hawaiian I s l a n d s Nat iona l W i l d l i f e Refuge, September 3-17, 1972. U. S. F i s h and W i l d l i f e Se rv ice , Bureau of Spor t F i s h e r i e s and W i l d l i f e , unpublished r e p o r t , 28 pp.

328. Kuroda, J . , Senator (1977) Re la t ing t o f i s h i n g . S t a t e of H a w a i i , Ninth L e g i s l a t u r e . A B i l l f o r a n A c t , Sena te B i l l No. 855.

329. Kuroda, J., Senator (1979) Re la t ing t o f i s h i n g . S t a t e of H a w a i i , Tenth Leg i s l a tu re . A B i l l f o r an A c t , Sena te B i l l No. 1430.

330. Laycock, G. (1970) The Hawaiian I s l a n d s of b i r d s . Audubon, 72, 1:44-61.

331. Leech, M. (1975) How a Br i ton views t h e H a w a i i e x h i b i t . !The Hono.hZu Advertiser, 23 December, B:2.

332. Linder, S . (1973) F i sh ing f l e e t s and sea t u r t l e s . Honozuzu Star- Bulletin, 1 6 August, A:23.

333. Lipman, V. (1978) H a w a i i ' s endangered w i l d l i f e : What chances f o r s u r v i v a l ? HonoZuZu, 13, 5:47-62.

334. Lipske, M. C. (1977) Sea t u r t l e s s u f f e r as bu reauc ra t s b i cke r . Defenders, August, 227-229.

335. Lis iansky , U. (1814) A voyage round t h e world i n t h e y e a r s 1803-1806 i n t h e s h i p Neva. John Booth, London, 388 pp.

336. Loveridge, A. (1945) R e p t i l e s o f t h e P a c i f i c world. The MacMillan Company, New York, 259 pp.

337. Lunasco, O. , Represen ta t ive (1977) Re la t ing t o speared f i s h . S t a t e o f H a w a i i , Ninth L e g i s l a t u r e . A B i l l f o r an A c t , House B i l l No. 515.

76

338.

339.

340.

341.

342.

343.

344.

345.

346.

347.

348.

349.

350.

351.

352.

353.

Lyons, A. E . (1890) I n b i r d land: P a r t of t h e j o u r n a l of a v i s i t t o Laysan I s l a n d . The Friend, 48, 12:90-91.

c

MacKelland, J , S . (1968) H a w a i i goes f i s h i n g : T u r t l e hunt ing . Charles E. T u t t l e Company, Rutland, Vermont, 160 pp.

Mahikoa, K. (1977) Meat from Caymans be ing s o l d now: Impor te rs hope to s t a r t t u r t l e farm. Pacific Business News (Honolulu) 14 May.

9

Malo, D. (1951) Hawaiian a n t i q u i t i e s . T rans l a t ed from the Hawaiian by D r . N. B. Emerson, Bishop Museum P r e s s , Honolulu, 278 pp.

Marquez, R. , Vi l lanueva , A. and C. Pena f lo re s (1976) S inops i s de d a t o s b i o l o g i c o s sob re l a t o r t u g a g o l f i n a : Lepidochelys olivacea (Eschscholtz, 1829). I n s t i t u t o Nacional de Pesca, Mexico, D. F., 61 PP.

Matsuura, P. (1975) There's something f i s h y going on! The HonoZuZu Advertiser, 26 August, B : l .

May, R., e d i t o r (1973) P a c i f i c I s l a n d Mar i cu l tu re Conference, February 6-8, 1973. Un ive r s i ty of H a w a i i , H a w a i i I n s t i t u t e of Marine Biology, Kaneohe, unpublished r e p o r t , 88 pp.

McAllister, J. G. (1933) Archaeology of Oahu. Bernice P. Bishop Musewn Bullet in 104, 201 pp.

McAllister, J. G. (1933) Archaeology of Kahoolawe. Bernice P. Bishop Museum Bulletin 1 1 5 , 61 pp.

McBride, L. R. (1969) Pe t roglyphs of H a w a i i . Petroglyph P r e s s , Hi lo , H a w a i i , 48 pp.

McBride, L. R. (1975) P r a c t i c a l f o l k medicine of H a w a i i . Pe t roglyph P r e s s , Hi lo , H a w a i i , 104 pp.

McCabe, R. (1975) Man f a c e s f i n e f o r t ak ing t u r t l e . HonoZuZu Star- BuZZetin, 5 March, A:4.

McKeom, S. (1978) Hawaiian r e p t i l e s and amphibians. The Oriental Pub l i sh ing Company, Honolulu, 80 pp.

McKinney, A. (1978) Barn ic l e [ s i c ] B i l l t h e d r i f t e r . Hawaii E ' L d z i ~ g News, A p r i l , 23.

McMahon, J . J. (1977) Un ive r s i ty of H a w a i i Marine Option Program: Annual r e p o r t 1976-77. Un ive r s i ty of H a w a i i , Sea Grant Miscellaneous Report UNIHI-SEAGWT-MR-78-02, 34 pp.

Mellen, I. M. (1925) Marine t u r t l e s s l e e p on Hawaiian sands. New York Zoological Society Bul le t in , 28:160-161.

x

P

5.

354.

355.

356,

357.

358.

359.

360.

361.

362.

363.

364.

365.

366,

367.

368.

369.

370.

Menard, W. (1967) 20., 1: 10-15.

Michioku, S. (J97

77

The las t paradise . Pacific Discovery,

) But programs can ' p l e a ' : UH t o absorb "udget cutbacks. kh Nuhou Kau Wela (Univers i ty of Hawaii); 3 Ju ly .

Miller, R. C , (1975) Lindbergh f i s h , b i r d sanc tuary proposed. The +Sunday Star-Bulletin -& Advertiser (Honolulu) ? 13 A p r i l , A:8.

Mi t che l l , W. (1967) T u r t l e s t odd le a long road t o s u r v i v a l . The Hono Zulu Advertiser, 3 May , B : 1.

Moake, B. (1973) Hearing Dec. 5: T u r t l e r e g s proposed. The Maui News, 27 November, B:8, 10.

Moake, B. (1973) P r o t e c t i o n of sea t u r t l e s favored by speakers . The Maui News, 11 December? B : l .

Morre l l , B. (1832) A n a r r a t i v e of f o u r voyages, t o t h e South Sea, North and South P a c i f i c Ocean, Chinese Sea E th iop ic and Southern A t l a n t i c Ocean, Ind ian and A n t a r c t i c Ocean. J . SI .J . Harper, New York, 492 pp.

Morse, L. A . (1944) Going up (photograph). Paradise of the Pacific, 58, 12:96.

Mowbray, L. S . (1965) Hawaiian monk seals Monachus schauinslandi and green t u r t l e s Chelonia mydas a t Waikiki Aquarium. Zoo Yearbook, 5 : 146-147.

International

Mrosovsky, N. , e d i t o r (1976) The t a g l o s s problem. IVCN/SSC W i n e Turtle Newsletter, 1 : 3-4.

Mull, M. E . (1973) Excerpts : Minutes of t h e H a w a i i Audubon Socie ty gene ra l meeting, 15 January 1973. The Elepaio, 33, 12:134-136.

Munro, G. C. (1940) Bi rds of H a w a i i . Tongg Pub l i sh ing Company, Honolulu, 189 pp.

Munro, G. C. (1941) Bi rds of H a w a i i and adventures i n b i r d s tudy: An ocean c r u i s e , No. 4. The Elepaio, 2, 8, 56-58.

Munro, G. C. (1942) Bi rds of Hawai.L and adventures i n b i r d s tudy: An ocean c r u i s e , No. 5. The Eleptzio, 2, 9:63-64.

Munro, G. C. (1942) Bi rds of H a w a i i and adventures i n b i r d s tudy: An ocean c r u i s e , N o . 6. The Elepuio, 2, 10:67-68.

Munro, G. C. (1942) Bi rds of H a w a i i and adventures i n b i r d s tudy: An ocean c r u i s e , No. 7. The EZepaio, 2, 11;71-72.

Munro, G. C. (1942) B i rds of H a w a i i and adventures i n b i r d s tudy: An ocean c r u i s e , N o . 8. The Elepaio, 2, 12:2-6.

78

371. Munter, W . H . (1916) Report dur ing c r u i s e , i s l a n d s t o the westward (USCGC Thet i s ) 24 January-13 February 1916. Records of t h e U. S. Coast Guard, Record Group 26, Nat iona l Archives, Washington, D. C . , 19 pp.

372.

373.

374.

375.

Murphy, R. (1968) A certain i s l a n d . The Hearst Corporation, New York, 239 pp.

Murphy, R. W. (1968) Wild s a n c t u a r i e s : Hawaiian I s l ands . Dutton Company, New York, 288 pp.

Nelson, L. (1976) I s o l a t i o n r u l e s on Kure A t o l l . HonoluZu Star - Bulletin, 24 March, A:12 .

P

Newbert, C . , Guard, R. T . , Yamato, B. and A. Baird (1977) Tide Calendar: I n t h e depths of t h e sea. Dillingham Corporation, Honolulu .

Newman, T. S. (1968) Hawaiian f i s h i n g and farming on t h e I s l a n d of H a w a i i i n A. D. 1 7 7 8 . S t a t e of H a w a i i , Department of Land and N a t u r a l Resources, Div is ion of S t a t e Parks , 305 pp.

376 A

377. Newman, T. S. (1972) Man i n t h e p r e h i s t o r i c Hawaiian ecosystem. In E. A. Kay ( e d i t o r ) A n a t u r a l h i s t o r y of t h e Hawaiian I s l a n d s : Se lec t ed readings . The Un ive r s i ty P r e s s of H a w a i i , 653 pp.

378.

379.

Nor thsh ie ld , R. (1969) The Leeward I s l a n d s . NatumZ History, 78: 60-66.

Oliver, J. A. and C. E . Shaw (1953) The amphibians and r e p t i l e s of t h e Hawaiian I s l a n d s . Zoologica: New York Zoological Society, 38: 65-95.

L

380. Olsen, D. L. (1969) Hawaiian I s l a n d s Nat iona l W i l d l i f e Refuge f i e l d t r i p , May 28-June 11, 1969. U. S. F ish and W i l d l i f e Se rv ice , Bureau of Spor t F i s h e r i e s and W i l d l i f e , unpublished r e p o r t , 18 pp.

Olsen, D. L. (1971) Expedi t ion r e p o r t , Hawaiian I s l a n d s Nat iona l W i l d l i f e Refuge, Nihoa, Necker, French F r i g a t e Shoals, August 16-26, 1971. U. S . F i s h and W i l d l i f e Se rv ice , Bureau of Spor t F i s h e r i e s and W i l d l i f e , unpublished r e p o r t , 1 6 pp.

381.

382. Olsen, D. L. (1972) Hawaiian I s l a n d s Nat iona l Wi ld l i f e Refuge exped i t ion r e p o r t , French F r i g a t e Shoals, May 4-11, 1972. U. S. F ish and W i l d l i f e Se rv ice , Bureau of Spor t F i s h e r i e s and Wi ld l i f e , unpublished r e p o r t , 1 3 pp.

c

383. Olsen, D. L. (1973) Expedition r e p o r t , Hawaiian I s l a n d s Nat iona l W i l d l i f e Refuge, A p r i l 24-27, 1973. U. S. F i sh and W i l d l i f e Serv ice , Bureau of Spor t F i s h e r i e s and W i l d l i f e , unpublished r e p o r t , 6 pp.

384. Olsen, D. L. (1973) Hawaiian I s l a n d s Nat iona l W i l d l i f e Refuge e x p e d i t i o n r e p o r t French F r i g a t e Shoals, June 1-5, 1973. U. S . F i sh and W i l d l i f e Se rv ice , Bureau of Spor t F i s h e r i e s and W i l d l i f e , unpublished r e p o r t , 5 pp.

79

385.

"L

c

,-

386.

387.

388.

389.

390.

391.

392.

393.

394.

'395.

396.

397.

Olsen, D. L . (1973) Expedition r e p o r t , Hawaiian I s l a n d s Nat iona l W i l d l i f e Refuge, J u l y 22-August 1, 1973. U. S . F i sh and W i l d l i f e Service, Bureau of Spor t F i s h e r i e s and W i l d l i f e , unpublished r e p o r t , 1 7 pp.

P a c i f i c Ocean F i s h e r i e s I n v e s t i g a t i o n s (1950) FWS M/V Hugh M. Snrith, r e p o r t on c r u i s e 2 , 16 January-9 February. Bureau of Commercial F i s h e r i e s , Honolulu, unpublished r e p o r t .

P a c i f i c Ocean F i s h e r i e s I n v e s t i g a t i o n s (1950) N a r r a t i v e r e p o r t : Hugh M. Smith, c r u i s e 5 , 16 June-6 August. Bureau of Commercial F i s h e r i e s , Honolulu, unpublished r e p o r t .

P a c i f i c Ocean f i s h e r i e s I n v e s t i g a t i o n s (1950) S c i e n t i s t s ' l o g f o r t h e June-August 1950 c r u i s e of t h e H . M. Smi th ,c ru ise 5. Bureau of Commercial F i s h e r i e s , Honolulu, unpublished r e p o r t .

P a c i f i c Ocean F i s h e r i e s I n v e s t i g a t i o n s (1951) N a r r a t i v e r e p o r t Hugh M. Smith, c r u i s e 9, 5 May-2 Ju ly . Bureau of Commercial F i s h e r i e s , Honolulu, unpublished r epor t .

P a c i f i c Ocean F i s h e r i e s I n v e s t i g a t i o n (1954) N a r r a t i v e ' r e p o r t of t h e September-November 1954 c r u i s e of t he C. H. GiZber t ,c ru ise 1 7 . Bureau of Commercial F i s h e r i e s , Honolulu, unpublished r e p o r t .

P a c i f i c Ocean F i s h e r i e s I n v e s t i g a t i o n (1955) Cruise r e p o r t FWS M/V John R. Manning, c r u i s e 25, 2 May-7 June. Bureau of Commercial F i s h e r i e s , Honolulu, unpublished r e p o r t .

P a c i f i c Ocean F i s h e r i e s I n v e s t i g a t i o n s (1956) Cruise r e p o r t : EWS M/V John R. Manning, c r u i s e 30, 15 March-20 Apr i l . Bureau of Commercial F i s h e r i e s , Honolulu, unpublished r e p o r t .

P a c i f i c Ocean F i s h e r i e s I n v e s t i g a t i o n s (1956) Cruise r e p o r t : FWS M/V J o h R. Manning, c r u i s e 31, 19 May-22 June. Bureau of Commercial F i s h e r i e s , Honolulu, unpublished r e p o r t .

P a c i f i c Ocean F i s h e r i e s I n v e s t i g a t i o n s (1957) Narrative r e p o r t Hugh M. Smith, c r u i s e 39, 19 April-30 May. Bureau of Commercial F i s h e r i e s , Honolulu, unpublished r e p o r t .

P a c i f i c Ocean F i s h e r i e s I n v e s t i g a t i o n s (1959) N a r r a t i v e r e p o r t Charles H. Gi lber t , c r u i s e 45, 6 July-3 September. Bureau of Commercial F i s h e r i e s , Honolulu, unpublished r epor t .

P a c i f i c Ocean F i s h e r i e s I n v e s t i g a t i o n s (1961) Notes from EWS M/V ChWles H. G i l b e r t , c r u i s e 51, 9-19 February. Bureau of Commercial F i s h e r i e s , Honolulu, unpublished r e p o r t .

P a c i f i c Ocean F i s h e r i e s I n v e s t i g a t i o n s (1961) Notes from FWS M/V Charles H . Gilber t , c r u i s e 53, 13-14 J u l y - Bureau of Commercial F i s h e r i e s , Honolulu, unpublished r epor t .

80

398.

399.

400.

40.L.

402.

403.

404.

4 0 5 ,

406 ..

407,

408.

409.

410.

411.

412.

413.

P a c i f i c Ocean F i s h e r i e s I n v e s t i g a t i o n s (1966) Notes from FWS M / V Charles H . Gilbert, c r u i s e 91, 12-15 May. Bureau of Commercial F i s h e r i e s , Honolulu, unpublished r e p o r t .

P a r r i s h , J . D. and L . R . Taylor Jr. (1978) Northwestern Hawaiian I s l a n d s F i s h e r i e s I n v e s t i g a t i o n s : Trophic a n a l y s i s of shallow- water f i s h communities i n the Northwestern Hawaiian I s l a n d s : E f f e c t s of n a t u r a l and human preda t ion . Sea Grant I n s t i t u t i o n a l Program Proposa l , Univers i ty of H a w a i i , Sea Grant College Program, Years 12-13 (1979-1981), A38-A49.

c

Parsons, J. J. (1962) The green t u r t l e and man. G a i n e s v i l l e , Un ive r s i ty of F lo r ida Press. 126 pp.

Parsons, J . J . (1972) The hawksbill t u r t l e and t h e t o r t o i s e s h e l l Etudes de geo9raphi.e tropicale o f f e r t e s a Pierre Gourou, t r a d e .

45-60.

Pa ty , J. (1857) Account of t h e Idanukawai - i n t e r e s t i n g account of h e r e x p l o r a t i o n s . The Polynesian, 6 June, 40 : 2-3,

Peters, H. H . , Representa t ive (1977) Re la t ing t o f i s h i n g . S t a t e of H a w a i i , Ninth L e g i s l a t u r e : A B i l l f o r an Act, House B i l l No. 1163.

Pfund, R . , edi . tor (1972) Baby t u r t l e s are HIMB lunch g u e s t s . Sea Grant Ne7~sZetter (Univers i ty of Hawaii), 2 , 11 : 4.

Pfund, R . , e d i t o r (1973) "Tur t l e , t u r t l e , come up t o brea the ." Sea Grant Newsletter (Un ive r s i ty of H a w a i i ) , 3 , 9:1, 4 .

Pfund, R . , e d i t o r (1974) Haw'n sea t u r t l e . Sea Grant Newsletter (Univers i ty o f Hawaii), 4 , 4:3-4.

Pfund, R . , e d i t o r (1974) Turtle mar i cu l tu re : Boon o r bane. Sea Grant Newsletter (Univers i ty of H a w a i i ) , 4 , 4:5.

Pfund, R . , e d i t o r (1974) Green sea t u r t l e . Sea Grant Newsletter (Univers i ty of Hawaii), 4 , 5:5.

Pfund, R . , e d i t o r (1977) Living toge the r i n peace (photograph and c a p t i o n ) . Sea Grant Newsletter (Univers i ty of Hawaii), 7, 7.

Pfund, R. , e d i t o r (1978) MAP marine miscellany. Sea Grant Newsletter (Univers i ty of Hawaii), 8, 12:4.

Pfund, R . , e d i t o r (1978) F ishery c r u i s e t r a p s 79 sha rks . Sea Grant Neusletter (Un ive r s i ty of Hawaii), 8, 4:8.

Pfund, R . , e d i t o r (1.978) An idea whose t i m e has come: Northwestern Hawaiian I s l a n d s provide p o t e n t i a l f o r f i s h e r y development. Sea Grant Newsletter (Univers i ty of Hawaii), 8, 1 : l -3 .

P h i l l i p s , M. (1973) A day on Coconut I s l and . Sun Jose Mercury, 12 September.

F

c

c

8 1

414. Polunin, N . V. C. (1975) Sea t u r t l e s : Reports on Thailand, West Malaysia and Indonesia , w i th a synops is of d a t a on t h e ' conserva t ion s t a t u s ' of sea t u r t l e s i n the Indo-West P a c i f i c reg ion . Report t o t he World Wi ld l i f e Fund, March, 113 pp.

415. Pope, C. H. (1939) T u r t l e s of t he United S t a t e s and Canada. Al f red A . Knope, New York, 343 pp.

416, P o t t e r , N . W. (1947) Wanna wrestle w i t h a t u r t l e ? Hawziian Digest, 1, 1:13-14. Reprinted from Hauaii Sporting News.

417., P r i t c h a r d , P. C. H. (1967) Living t u r t l e s of t h e world. T. F. H.' Pub l i ca t ions , J e r sey Ci ty , New Je r sey , 288 pp.

418. P r i t c h a r d , P. C. H. (1977) Marine t u r t l e s of Micronesia. Chelonia P res s , San Francisco, 83 pp.

419, P r i t c h a r d , P . C. H. and W . F. Greenhood (1968) The sun and t h e t u r t l e . International Turtle and Tortoise Society Journal, 2 : 20-25 , 34- 35.

420. Pryor , T . (1974) Systemculture: A marine p l a n t a t i o n f o r North Kohala. Kohala Task Force Grant Submi t ta l , Scope of Work Statement , 29 pp.

r )

4..

421. Pryor , T. A . (1976) Habi ta t : The b lue r evo lu t ion . NationaZ ' Development, May, 64-67.

422. Pryor , T. A . (1978) 'Systemculture ' works on Oahu: Cul ture of phytoplankton provides f eed f o r o y s t e r farm. Farmer and Aquaculture News, 4, 3: 24-26.

The ComercCaZ Fish

423. Pukui, M. K. and C. C u r t i s (1951) The water of Kane: The t u r t l e i n the sweet-potato pa tch . The Kamehameha Schools P res s , Honolulu, 249 pp.

424. Pukui, M. K. and C . C u r t i s (1971) P i k o i and o t h e r legends of t h e I s l and of H a w a i i : The r e scue of Hina. The Kamehameha Schools P res s , Honolulu, 282 pp.

425. Pukui, M. K. and S . H. E l b e r t (1971) Hawaiian d i c t i o n a r y . The Univers i ty P r e s s of H a w a i i , Honolulu, 402 pp + 188 pp.

426. Pukui, M. K . , E l b e r t , S. H . and E. T . Mookini (1976) P l a c e names of H a w a i i . The Un ive r s i ty P r e s s of H a w a i i , Honolulu, 28 pp.

427. Pukui, M. K. , E l b e r t , S. H. and E. T. Mookini (1978) The pocket Hawaiian d i c t i o n a r y . The Un ive r s i ty P r e s s of H a w a i i , Honolulu, 276pp.

428. Rauzon, M., Kenyon, K . and A . M. Johnson (1977) Observat ions of monk seals, French F r i g a t e Shoals , 1 7 February t o 27 May 1977. United S t a t e s Department of t he I n t e r i o r , F ish and Wdldl i fe Service, Nat iona l F ish and W i l d l i f e Laboratory, Anchorage F ie ld S t a t i o n , unpublished r e p o r t , 34 pp.

82

429.

430.

431.

432.

433.

434.

435.

436.

437.

438,

439,

440.

441.

442,

443.

444.

Read, G. H. (1912) The l a s t c r u i s e of t h e Saginaw. Houghton M i f f l i n Company, Boston, 1 2 8 pp.

4

Ressencourt , E . (1962) The t u r t l e rock and the l o t u s s tone : Molokai's myster ious f e r t i l i t y s h r i n e . Hamilton Library , Hawaiian Co l l ec t ion , unpublished r e p o r t , 32 pp.

Univers i ty of H a w a i i ,

Rice, W. H. (1923) Hawaiian legends. The Museum, Honolulu, 137 pp. c

Richardson, F. (1954) Notes on the b i r d s of French F r i g a t e Shoal. P a r t I. General account of v i s i t of October 1953. The EZepaio, 1 4 , 9:61-63.

Ridgeway, S . H., Idever, E . G . , McCormick, J . G . , P a l i n , J . and J. H. Anderson (1969) Hearing i n t h e g i a n t sea t u r t l e , CheZonia mydas. Proceedings of t h e Nat iona l Academy of Science, 1 5 November, 64, 3: 885-890.

Ri ley , R. (1966) H a w a i i ' s petroglyph puzzles: What were they t r y i n g t o t e l l us? f i e Sunday Star-Bulletin & Advertiser (Honolulu), 11 December, D:6.

Rothschi ld , W. (1893-1900) The av i fauna of Laysan and t h e neighboring i s l a n d s . R. H. P o r t e r , London.

Sa l i sbu ry , G. R. (1913) Report on Laysan exped i t ion , December 5 , 1912 t o A p r i l 11, 1913. Record Group 22, Nat iona l Archives, Washington, D. C . , 8 pp.

x

Schauinsland, H. (1899) Drei monate auf e i n e r Koral len-Inseln. Bremen, Noss le r , 104 pp.

n

Schlemmer, M. and E. L. Schlemmer (1915) Copy of t he l o g of t he yacht "Helene" and d i a r y w r i t t e n on Laysan I s l a n d from June 25 t o December 2, 1915. Record Group 22, Nat iona l Archives, Washington, D. C. , 32 pp.

Sekora, P. (1975) Refuge f o r rare s p e c i e s , Defenders 50, 6:506-511.

Se l land , J. (1974) Clutch of t u r t l e s ha tch i n n e s t a t Punalu 'u Beach. H a u a i i Tribune-HeraZd: Orchid I s l e (Hawaii), 27 October- 2 November, 5-6.

Sha l lenberger , E. W. (1979) Kaula I s l a n d . Hawaii Fishing,News, January, 15.

Shor t , P. (1967) Tern I s land- lone ly outpos t . HonoZuZu Star-Bulletin, 19 February, B:5.

Shura, E. (1923) French F r i g a t e Shoal explored by lea thernecks of the Rainbow. The Hono Zu Zu Aduertiser, 8 Ju ly , A: 1.

Sib ley , F. C . (1963) P a c i f i c p r o j e c t f i e l d t r i p , June 1963. Leeward survey no. 2. P a c i f i c Ocean B io log ica l Survey Program, Smithsonian I n s t i t u t i o n , Washington, D. C . , unpublished r e p o r t , 9 pp.

a3

445.

446.

447.

448.

449.

450.

451.

452.

453.

454.

455.

456. P

457.

458.

Sib ley , F. C. (1964) Pre l iminary r e p o r t on ATF t r i p no. 1, February-March 1963. Leeward survey no. 1. P a c i f i c Ocean Bio log ica l Survey Program, Smithsonian I n s t i t u t i o n , Washington, D. C. , unpublished r e p o r t , 1 3 pp.

Smith, C. H. (1966) Hi lo legends. Pe t roglyph P r e s s , Hi lo , H a w a i i , 29 pp.

Smith, T. and J . Stimson (1978) S t a t u s of ha rves t ed r e sources i n t h e Hawaiian I s l a n d s and t h e P a c i f i c T rus t T e r r i t o r y . review and s y n t h e s i s of in format ion on P a c i f i c i s l a n d ecosystems. Documentation Assoc ia tes Informat ion Se rv ices Incorpora ted , Los Angeles, C a l i f o r n i a . U . S. F i sh and W i l d l i f e Se rv ice c o n t r a c t number 14-16-009-77-075, Po r t l and , 9:l-13.

In L i t e r a t u r e

Smyser, A . A . , e d i t o r (1974) H a w a i i ' s green t u r t l e . HonoZuZu Star- Bulletin, 4 A p r i l , A:20.

Smyser, A . A., e d i t o r (1974) Endangered t u r t l e s . Honok~Zu S tar - Bulletin, 31 May, A: 20.

Snow, C. E. (1974) Ea r ly Hawaiians: An i n i t i a l s tudy of s k e l e t a l remains from Mokapu, Oahu. Lexington, 179 pp.

The Unive r s i ty Press of Kentucky,

S t a d e l , D. L. (1967) Pre l iminary r e p o r t , Laysan I s l a n d , Leeward I s l a n d survey 19 , June 7-12, 1967. P a c i f i c Ocean B i o l o g i c a l Survey Program, Smithsonian I n s t i t u t i o n , unpublished r e p o r t , 18 pp.

S t a f f o r d , J. (1973) Opera t ion t u r t l e o r a l l you ever wanted t o know about CheZoniu mydas bu t were a f r a i d t o ask. (Fourteenth Coast Guard District) , March, 8-10.

Pac i f i c Shield

Stansbury, J . and E. F l a t t a u (1974) T u r t l e l e s s soup. Los Angeles T i m e s Syndicate, 1 3 March;

S t a t e of H a w a i i (1948-1974) Commercial f i s h ca t ch b y , s p e c i e s , ca l enda r y e a r s 1948-1974: Honu ( t u r t l e ) . Department of Land and Na tu ra l Resources, Div is ion of F i s h and G a m e .

S t a t e of H a w a i i (1959-1974) Commercial f i s h c a t c h by s p e c i e s , f i s c a l y e a r s 1959-1974: Honu ( t u r t l e ) . Department of Land and Na tu ra l Resources, Div is ion of F ish and Game .

S t a t e of H a w a i i (1960-1974) Monthly commercial f i s h ca t ch . by s p e c i e s , January 1960-March 1974: Honu ( t u r t l e ) . Department of Land and Na tu ra l Resources, Div is ion o f F i s h and Game .

S t a t e o f H a w a i i (1971) Digest of c e r t a i n s a l t w a t e r f i s h i n g l a w s and r e g u l a t i o n s . Department of Land and Na tu ra l Resources, Div is ion of F i sh and Game, 9 pp.

S t a t e of H a w a i i (1971) F i sh ing w i t h f i r ea rms , s p e a r s . H a w a i i Revised S t a t u t e s , Sec t ion 188-25.

84 c

459. S t a t e of H a w a i i (1973) Minutes of t h e pub l i c hea r ing of t h e Board of Land and Na tu ra l Resources on Regulat ion 36 r e l a t i n g t o t h e p r o t e c t i o n of marine t u r t l e s : 2 1 September 1973 Honolulu. Department of Land and Natural Resources, Div is ion of F i s h and G a m e , 8 pp.

m

460. S t a t e of H a w a i i (1973) Minutes of p u b l i c hea r ing of Board of Land and Natura l Resources, Div is ion of F ish and Game: 6 December 1973 Hi lo , H a w a i i . Department of Land and Natura l Resources, Div is ion of F i sh and G a m e , 4 pp.

461. S t a t e of H a w a i i (1973) Minutes of t h e p u b l i c hea r ing of t h e Board of Land and Natura l Resources on Regulat ion 36 r e l a t i n g t o t h e p r o t e c t i o n of marine t u r t l e s : 5 December 1973 Wailuku, Maui. Department of Land and Natura l Resources, D iv i s ion of F i sh and Game, 5 pp.

462. S t a t e of H a w a i i (1973) Minutes of t h e pub l i c hear ing of t h e Board of Land and Natura l Resources on Regulat ion 36 r e l a t i n g t o the p r o t e c t i o n of marine t u r t l e s : Department of Land and Natura l Resources, Div is ion of F ish and Game, 5 pp.

5 December 1973 Lihue, Kauai.

4 6 3 , S t a t e of H a w a i i (1974) Minutes of t h e pub l i c hea r ing of t h e Board of Land and Natural Resources on Regulation 36 r e l a t i n g t o the p r o t e c t i o n of marine t u r t l e s : 2 8 February 1974 Kaurrakakai, Molokai. Department of Land and Natura l Resources, Div is ion of F i s h and Game, 5 PP*

S t a t e of H a w a i i (1974) Rare and endangered spec ie s . Report t o t he Governor 19 72-19 73. January.

Department of Land and Natura l Resources, 464.

465. S t a t e of H a w a i i (1974) Regulat ion No. 36: Regulat ion r e l a t i n g t o t h e p r o t e c t i o n of marine t u r t l e s . Department of Land and Natura l Resources, Div is ion of F ish and G a m e .

466.

467.

S t a t e of H a w a i i (1977) H a w a i i aquacul ture program i n t e r i m repor t . Department of Planning and Economic Development, January, 21 pp.

S t a t e of H a w a i i (1978) Aquaculture development f o r H a w a i i : Assessments Department of Planning and Economic Development, and recommendations.

Aquaculture Planning Program, 222 pp.

468. S t a t e of H a w a i i (no d a t e ) Marine t u r t l e s found i n Hawaiian waters. Department of Land and Natura l Resources, Div is ion of F ish and Game, l e a f l e t .

469,

470.

S t e jnege r , L. (1899) The land r e p t i l e s of t h e Hawaiian I s l a n d s . Proceedings U. S. Nationa2 Museum 2 1 , 1174:783-813.

S t o f f e l , P. (1974) Rules should p r o t e c t both t u r t l e s and fishermen, s a y s Yamagata. The Garden. Is2an.d (Lihue, Kauai), 23 January, A:9.

85

P

471,

472

473.

474.

475,

476.

477.

478.

479.

480.

I-

481.

482,

483.

484.

485.

Stokes, J. F. G . (1906) Hawaiian n e t s and n e t t i n g . Memoirs of the Bernice P. Bishop @usem, 2:105-162.

Svih la , A. (1957) Observat ions on French F r i g a t e Shoals , February 1956. Atol l Research Bulletin, 51:l-2.

Tabrah, R . (1976) Lanai. I s l a n d Heri tage, Norfolk I s l and , A u s t r a l i a , 120 pp.

Takemoto, R. , Tabata, R. and M. Doughtie (1978) H a w a i i f i s h i n g l a w s and r e g u l a t i o n s f o r sa l t water s p o r t s f i s h i n g . Univers i ty of H a w a i i Sea Grant College Program, Marine Advisory Program, UNIHI-SEAGRANT-AB-78-07.

Taylor , L t R. Jr. (1973) . Prel€minary observa t ions of t h e in shore 'mar ine ecosystem i n the Leeward Hawaiian I s l ands . Univers i ty of

H a w a i i , Cooperative F ishery Unit , unpublished r e p o r t , August, 1 0 pp.

Taylor, L. (1975) The aquarium g e t s a f a c e l i f t . HonoZuZu Star- Bullst<n, 11 November C : l .

Taylor, L. (1978) T u r t l e u n g i r t . Kilo i ' a (Waikiki: Aquarium), 3:l-2.

Taylor, L. (1978) A f u l l year of Friend-ship. Kilo i ' a (Waikiki Aquarium) , 4 : 1-2.

Taylor , L. (1978) T u r t l e homecoming. Kilo i 'a (Waikiki Aquarium) , 5:2.

Taylor, L. R. , Jr. (1978) Northwestern Hawaiian I s l a n d s F i s h e r i e s I n v e s t i g a t i o n s : Ecology of top-level ca rn ivo res i n r ee f communities of t h e Northwestern Hawaiian I s l ands . Sea Grant I n s t i t u t i o n a l Program Proposal , Un ive r s i ty of H a w a i i , Sea Grant Col lege Program, Year 11 (1978-1979) , A38-A42.

Taylor, L. R. and G. N a f t e l (1977) How t o avoid sha rk a t t a c k ( I f you happen t o be a Hawaiian monk s e a l ) . Oceans, 10, 6:21-23.

Taylor, L. R. and G. N a f t e l (1978) Pre l iminary i n v e s t i g a t i o n s of sha rk p reda t ion on the Hawaiian monk seal a t P e a r l and H e r m e s Reef and French F r i g a t e Shoals. F i n a l Report , U. S. Marine Mammal Commission c o n t r a c t N o . 7ACOl1, NTIS PB285-626, 34 pp.

T e l f e r , T. C. (1973) A r e p o r t on a t r i p t o Kure W i l d l i f e Kefuge, May 25 t o 30, 1973. S t a t e of H a w a i i , Department of Land and Natura l Resources, Divis ion of F ish and G a m e , unpublished r e p o r t , June, 1 7 pp.

TenBruggencate, J. (1974) Kauai t a l k i n g about p r o j e c t ' s bad luck . The Honolulu Advertiser, 2 October, A : l .

TenBruggencate, J. (1979) I s le c u l t u r e is entwined wi th olona. The Sunday Star-Bulletin & Advertiser (Honolulu), 11 February, A:32.

86

486. T e r r i t o r y of H a w a i i (1955) F ish ing w i t h f i rearms . Board of Commissioners of Agr i cu l tu re and Fores t ry , Div is ion of F i s h and Game, F i sh and Game Laws, Sec t ion 1229, 95.

487 .. 488.

489,

Tester, A. L. (1969) Cooperative sha rk r e sea rch and c o n t r o l program: F i n a l r e p o r t 1967-69. Un ive r s i ty of H a w a i i , 47 pp.

Thrum, T. C. , e d i t o r (1921) Hawaiian f o l k tales: A c o l l e c t i o n of native legends , A. C. McClung and Company, Chicago, 284 pp.

Tinker, S . W. (1935) Wild animals of H a w a i i : A series of b r i e f n a t u r e ar t ic les . No. 4 - The t runk t u r t l e . HonoZuZu Star-BdZetin, 24 August, 7 .

490 Tinker, S. W. (1935) Wild animals of H a w a i i : A series of b r i e f n a t u r e art icles. No. 5 - The green t u r t l e . Honolulu Star-Bulletin, 31 August, 16.

491. Tinker, S. W. (1935) Wild animals of H a w a i i : A series of b r i e f n a t u r e ar t ic les . No. 6 - The hawk's-bil l o r t o r t o i s e - s h e l l t u r t l e . IlonoZulu Star-Bulletin, 7 September, 6. r

492. Tinker, S. (1941) Animals of H a w a i i . Tongg Pub l i sh ing Company, Honolulu, 190 pp.

493, Titcomb, M. (1969) Excerpts from t h e minutes, H a w a i i Audubon Socie ty General Meeting, 21 J u l y 1969. The Elepaio, 30, 3:27.

494.

495.

Tomich, P. Q. (1975) Tr ip r e p o r t . The Elepaio, 36, 1:6-7.

Trempe, A. D. (1936) Report on t h e b i r d l i f e observed on t h e c r u i s e of t h e ReZiance i n March 1936. Washington, D. C., 5 pp.

Record Group 26, Nat iona l Archives,

496,

497.

498.

499.

Tus le r , S. (1974) Keep t u r t l e s a l ive, h igh rises down. The Garden Island (Lihue, Kauai), 7 January, 2.

U. S. Coast Guard (1975) Loran S t a t i o n French F r i g a t e Shoals: General in format ion book. Fourteenth Coast Guard Dis t r ic t , 13 pp. 5

U. S. Coast Guard (1976) Loran S t a t i o n Kure: General informat ion book. Four teenth Coast Guard District, 14 pp.

U. S. Department of Commerce (1968) H a w a i i : Outer I s l a n d s . United S t a t e s Coast P i l o t 7: P a c i f i c Coast, C a l i f o r n i a , Oregon, Washington and H a w a i i . Tenth Ed i t ion , 380 pp.

500. U. S. Department of Commerce (1971) F i she ry statistics of t h e United S t a t e s , 1968: H a w a i i f i s h e r i e s . Nat iona l Oceanic and Atmospheric Adminis t ra t ion , Nat iona l Marine F i s h e r i e s Serv ice , Washington, D. C., S t a t i s t i c a l Diges t No. 62, 578 pp.

87

501.

502.

503..

504.

505.

506.

507.

I / -

508.

509.

510.

U. S. Department of Commerce (1972) F ishery s t a t i s t i c s of t h e United S t a t e s , 1969: H a w a i i f i s h e r i e s . Nat iona l Oceanic and Atmospheric Adminis t ra t ion, Nat iona l Marine F i s h e r i e s Service, Washington, D. C., S ta t is t ical Digest No. 63, 474 pp.

U. S. Department of Commerce (1973) F ishery s ta t is t ics of t h e United States, 1970: H a w a i i f i s h e r i e s . Nat iona l Oceanic and Atmospheric Adminis t ra t ion , Nat ional Marine F i s h e r i e s Serv ice , Washington, D. C., S t a t i s t i c a l Digest No. 64, 489 pp.

U. S. Department of Commerce (1976) Dra f t Environmental Impact Statement: Proposed l i s t i n g of the green sea t u r t l e (Chetonia mydas), loggerhead sea t u r t l e (Caretta care t ta) , and P a c i f i c r i d l e y sea t u r t l e (LepidocheZys otivacea) as Threatened s p e c i e s under t h e Endangered Species Act of 1973. Adminis t ra t ion , Nat iona l Mar ine F i s h e r i e s Serv ice , Washington, D. C. , January, 167 pp.

Nat iona l Oceanic and Atmospheric

U. S . Department of Commerce (1978) F i n a l Environmental Impact Statement: mydas), loggerhead sea t u r t l e (Caretta care t tu) , and P a c i f i c r i d l e y sea t u r t l e (LepidocheZys olivacea) under the Endangered Spec ies A c t of 1973. Nat ional Oceanic and Atmospheric Adminis t ra t ion, Nat iona l Marine F i s h e r i e s Serv ice , Washington, D. C. Ju ly , 144 pp.

L i s t i n g and p r o t e c t i n g the green sea t u r t l e (C?lebnia

U. S. Department of Commerce (no 'da t e ) Fishery statist ics of t h e United States, 1971: H a w a i i f i s h e r i e s . Nat iona l Oceanic and Atmospheric Adminis t ra t ion, Nat iona l Marine F i s h e r i e s Se rv ice , Washington, D. C., S t a t i s t i c a l Digest No. 65, 424 pp.

U. S, Department of Commerce (no d a t e ) F ishery statist ics of t h e United S t a t e s , 1972: Hawaii f i s h e r i e s . Nat iona l Oceanic and Atmospheric Adminis t ra t ion, Nat iona l Marine F i s h e r i e s S e r v i c e , Washington, D. C. , S t a t i s t i ca l Digest No. 66, 517 pp.

U. S . Department of Commerce (no da te ) Fishery s ta t is t ics of t h e United States, 1973: H a w a i i f i s h e r i e s . Nat iona l Oceanic and Atmospheric Adminis t ra t ion, Nat iona l Marine F i s h e r i e s Se rv ice , Washington, D. C . , S t a t i s t i c a l Digest No. 67, 458 pp.

U. S. Department of Commerce (no d a t e ) Fishery statistics of the United S t a t e s , 1974: H a w a i i f i s h e r i e s . Na t iona l Oceanic and Atmospheric Adminis t ra t ion, National Marine F i s h e r i e s Service, Washington, D. C., S t a t i s t i c a l Diges t No. 68, 424 pp.

U. S . Department of t h e I n t e r i o r (1950) F ishery statist ics of t he United S t a t e s , 1946: Hawaiian f i s h e r i e s . F ish and W i l d l i f e Serv ice , Washington, D. C., S t a t i s t i c a l Digest 19, 265 pp.

U. S . Department of the I n t e r i o r (1650) Fishery statistics of the United States, 1947: Hawaiian f i s h e r i e s , F i sh and W i l d l i f e Se rv ice , Washington, D. C., S t a t i s t i c a l Digest 21, 285 pp.

88

511. U. S . Department of t h e I n t e r i o r (1960) Fishery s ta t i s t ics of t h e United S t a t e s , 1958: H a w a i i f i s h e r i e s . F ish and W i l d l i f e Se rv ice , Bureau of Commercial F i s h e r i e s , Washington, D. C. , S t a t i s t i c a l Digest N o . 49, 424 pp.

P

512. U. S . Department of t h e I n t e r i o r (1961) Fishery s t a t i s t i c s o f t he United S t a t e s , 1959: Hawaii f i s h e r i e s . F ish and W i l d l i f e Service, Bureau of Commercial F i s h e r i e s Washington, D. C . , S t a t i s t i c a l Digest No. 51, 457 pp.

w

513.

514.

U. S. Department of t h e I n t e r i o r (1962) Fishery s ta t i s t ics o f t h e United S t a t e s , 1960: H a w a i i f i s h e r i e s . F ish and W i l d l i f e Serv ice , Bureau of Commercial F i s h e r i e s , Washington, D. C. , S t a t i s t i c a l Diges t No. 53, 529 pp.

U. S. Department of t h e I n t e r i o r (1963) Fishery s t a t i s t i c s o f t h e United S t a t e s , 1961: H a w a i i f i s h e r i e s . Fish and W i l d l i f e Se rv ice , Bureau of Commercial F i s h e r i e s , Washington, D. C . , S t a t i s t i c a l Digest No. 54, 460 pp.

c 515. U. S . Department of t h e I n t e r i o r (1964) Fishery s ta t i s t ics o f t h e

United S t a t e s , 1962: H a w a i i f i s h e r i e s . F i s h and W i l d l i f e Serv ice , Bureau of Commercial F i s h e r i e s , Washington, D. C . , S t a t i s t i c a l Diges t N o . 56, 466 pp.

516.

517.

U. S . Department o f t h e I n t e r i o r (1964) Hawaiian I s l a n d s Nat iona l W i l d l i f e Refuge. F i s h and W i l d l i f e Service, Bureau o f Spor t F i s h e r i e s and W i l d l i f e , L-508, P o r t l a n d , 8 pp.

U. S. Department of t h e I n t e r i o r (1965) Fishery s t a t i s t i c s o f t h e United S t a t e s , 1963: H a w a i i f i s h e r i e s . F i s h and W i l d l i f e Se rv ice , Bureau o f Commercial F i s h e r i e s , Washington, D. C . , S t a t i s t i c a l Diges t No. 57, 522 pp.

518. U. S . Department of t h e I n t e r i o r (1966) Fishery s ta t i s t ics of t he United S t a t e s , 1964: H a w a i i f i s h e r i e s . F i s h and W i l d l i f e Serv ice , Bureau o f Commercial F i s h e r i e s , Washington, D. C. , S t a t i s t i c a l Digest No. 58, 541 pp.

519. U. S . Department of t h e I n t e r i o r (1967) Fishery s ta t i s t ics o f t h e United S t a t e s , 1965: H a w a i i f i s h e r i e s . F i s h and W i l d l i f e Serv ice , Bureau o f Commercial F i s h e r i e s , Washington, D. C . , S t a t i s t i c a l Digest N o . 59, 756 pp.

520.

521.

U. S . Department o f t h e I n t e r i o r (1968) Fishery s ta t i s t ics of t h e United S t a t e s , 1966: H a w a i i f i s h e r i e s . F i s h and W i l d l i f e Serv ice , Bureau of Commercial F i s h e r i e s , Washington, D. C . , S t a t i s t i c a l Diges t No. 60, 679 pp.

U. S . Department o f the I n t e r i o r (1969) Fishery s t a t i s t i c s of t h e United S t a t e s , 1967: H a w a i i f i s h e r i e s . F ish and W i l d l i f e Serv ice , Bureau o f Commercial F i s h e r i e s , Washington, D. C . , S t a t i s t i c a l Diges t No. 61, 490 pp.

c

89

522. U , S . Department of t he I n t e r i o r (1969) Hawaiian I s l a n d s Nat iona l W i l d l i f e Refuge. F i sh and W i l d l i f e Serv ice , Bureau of Spor t F i s h e r i e s and Wi ld l i f e , l e a f l e t 508-R2, May.

U. S . Department of t he I n t e r i o r (1970) Hawaiian I s l a n d s Nat iona l W i l d l i f e Refuge. F ish and W i l d l i f e Serv ice , Bureau of Spor t F i s h e r i e s and Wi ld l i f e , A p r i l , 28 pp.

523.

,-

U. S. Department of t h e I n t e r i o r (1973) Hawaiian I s l a n d s wi lderness proposal : Hawaiian I s l ands Nat iona l W i l d l i f e Refuge, Ci ty and County of Honolulu, H a w a i i . F i sh and W i l d l i f e Serv ice , Bureau of Spor t F i s h e r i e s and Wi ld l i f e , 26 pp.

524.

U. S. Department of t h e I n t e r i o r (1974) H a w a i i I s l a n d s Nat iona l W i l d l i f e Refuge. Fish and Wi ld l i f e Serv ice , l e a f l e t 1352700-1, March.

525.

526. U. S . Department of t he I n t e r i o r and U. S . Department of Commerce (1974) Sea Tur t l e s : Review of s t a t u s . Federa2 Register, 16 August, 39, 160:29605-29606,

U. S . Department of t he I n t e r i o r (1975) Sport f i s h e r y and w i l d l i f e research 1973-74: A c t i v i t i e s i n t h e d i v i s i o n s of r e sea rch f o r the ca lendar yea r 1973 and the f i r s t h a l f of 1974. F i sh and W i l d l i f e Serv ice , Div is ions of Research, Washington, D. C . , 178 pp.

527.

U. S . Navy (1973) Midway I s l a n d Navy W i l d l i f e Refuge management p lan . SOPAMIDWAY Ins t r u c t i o n 11015.1 , Apr i l , 9 pp . 528.

Udvardy, M. D. F. (1961) The Harold J. Coolidge expedi t ion t o Laysan I s l and , 1961. The EZepaio, 22, 6~43-47 .

529.

Vernon, S . (1975) Crossroads f o r contraband. Defenders, 50, 6:512-515. 530.

531. Walker, F. D. (1909) Log of t h e KaaZokai. The Hawaiian Gazet te Company, Ltd . , Honolulu, 64 pp.

Walker, R. L. (1963) Notes on a v i s i t t o Laysan Isl.and, December 3-10, 1963. S t a t e of H a w a i i , Department of Land and Natura l Resources, Div is ion of F i sh and G a m e , unpublished r e p o r t , 25 pp.

532.

Walker, R. L. (1964) Report on a t r i p t o t h e Hawaiian I s l a n d s Nat iona l W i l d l i f e Refuge, March 1964: F i e l d no te s . S t a t e of H a w a i i , Department of Land and Natura l Resources, Divis ion of F ish and Game, unpublished r e p o r t , 35 p p .

Walker, R. L. (1972) The w i l d l i f e of Keehi Lagoon, Oahu. The EZepaio, 32 , 11: 106-108.

533.

534.

Wallace, R. (1972) H a w a i i : The American Wilderness. Refuge on a r e e f . Time-Life Books, New York, 184 pp.

535.

Walsh, J. (1977) New York c i t y boy v i s i t s K a u a i . 'Ezepaio, 37, 7:76-77. 536.

537. Walters, J. F. (1977) H a w a i i Audubon Socie ty f i e l d t r i p t o Kauai, 1 9 March 1977. 'EZepaio, 38, 1:9.

Y

90

538.

539.

540.

541.

542.

543.

544.

545.

546.

547.

548.

549.

550.

551.

552,

553,

554.

Ward, R. G . , e d i t o r (1967) American ac t iv i t ies i n t h e Central P a c i f i c 1790-1870. Volume 4. The Gregg Press, Ridgewood. New .Jersey, 695 pp.

Westerve l t , W . D. (1964) Hawaiian legends of ghos ts and ghost-gods: Ke-au-nini. Charles E. T u t t l e Company, Rutland, Vermont, 262 pp.

Wes te rve l t , W . D. (1968) Ghosts of H i lo h i l l s . The Pe t roglyph P r e s s , L td . , H i lo , H a w a i i , 16 pp.

Wetmore, A . (1923) F i e l d n o t e s taken on t h e 1923 Tanager Expedition. Smithsonian I n s t i t u t i o n , Washington, D. C.

Wetmore, A . (1925) Bird l i f e among l a v a rock and c o r a l sand. !&.e NatwnaZ Gkographic Magazine, 48: 77-108.

Wheeler, P. (1953) Hawaiian wonder tales: Flash-of-the paddle and listener-to-the-heavenly-voice. The Beechhurst P res s , New York, 232 pp.

Whitten, H. (1972) Sunday ou t ing : Waimanalo pond recalls c h i e f ' s s p e c i a l d e l i c a c y , lionolulu Star-Bulletin, 29 January, B:14.

Whitten, H . (1976) Sea t u r t l e s . Honolulu Star-Bulletin. 27 September, A: 14.

Whitten, H. (1977) Leeward I s l a n d s s tudy . HonoluZu Star-Bulletin, 25 A p r i l , A:17.

Whitten, H. (1978) Necker I s l and . HornZulu Star-Bulletin, 6 February, A: 13.

Whitten, H. (1978) Notes. Honolulu Star-Bulletin, 24 Apr i l , A : 1 3 .

whittow, G. C. and G. H. Balazs (1977) Thermal ecology of Hawaiian basking green sea t u r t l e s . Research proposa l accepred by t h e Na t iona l Geographic Soc ie ty , Washington, D. C., 15 pp.

Whittow, G. C. (1978) Tern I s l a n d . Sea Frontiers, 24, 3:150-158.

c

6

Wilder, G. P. (1905) A s h o r t t r i p t o Midway wi th Captain A. P. Niblack i n t h e U . S . S . Iroquois. The Hamiian Forester and AgricuZturist, 2, 12:390-396.

u-

Wirtz, W . O . , I1 (1965) Leeward I s l a n d s survey no. 7, March 1965. P a c i f i c Ocean B io log ica l Survey Program, Smithsonian I n s t i t u t i o n , Washington, D. C . , unpublished r e p o r t , 26 pp.

Wong, F., Sena tor (1977) Re la t ing t o f i s h i n g . S t a t e of H a w a i i , Ninth Leg i s l a tu re . A B i l l f o r an A c t , Sena te B i l l No. 706.

Woodside, D. H. (1961) Report on a survey of Laysan I s l a n d , September 1961. S t a t e of H a w a i i , Department of Land and Natura l Resources, Div is ion of F ish and G a m e , unpublished r e p o r t , 32 pp.

91

555.

556.

557,

558.

559.

560.

561.

562. N

563.

564,

_P

565.

566.

567.

568.

569.

570.

Woodside, D. H. and R. J. Kramer (1961) A r e p o r t on a survey t r i p t o t h e Hawaiian I s l a n d s Nat ional W i l d l i f e Refuge, March 1961. State of H a w a i i , Department of Land and Natura l Resources, Divis ion of F ish and G a m e , unpublished r e p o r t , 30 pp.

Woodward, P. W. (1972) The n a t u r a l h i s t o r y of Kure A t o l l , Northwestern Hawaiian I s l ands . A t o l l Research Bulletin, 164:l-318.

Wright, R. J . , Lay, M. and G. H. Balazs (1977) An i n v e s t i g a t i o n of the green t u r t l e i n the feeding pas tu re s of Ka'u, H a w a i i . Report prepared f o r t he Marine Option Program, Universi ty of H a w a i i a t Hilo, 23 pp.

Yamada, S. (1977) Ceramics i n H a w a i i . Honolulu, 78 pp .

Yamamoto, H. and F. D. Bicoy (1972) Skin d iv ing h i n t s . CAP Newezetter (Molokai), 1 Ju ly , 66th i s s u e .

Yamashita, S. (1968) 'Forgot ten H a w a i i ' : The Leeward I s l ands . The Sunday Star-Bulletin & Advertiser (Honolulu), 1 3 October, A:8.

Y i m , S. (1977) The sunshine, sea.. .and l o n e l i n e s s . HonoZuZu Star- Bulletin, 6 A p r i l , F : l .

Yoshida, J. (1973) Rep. Anson Chong's r eques t f o r r e p r i n t s of green sea t u r t l e paper . The Elepaio, 34, 4:44.

Zalburg, S . (1978) Fisherman seeks test case a t French F r i g a t e Shoals. The Honolulu Advertiser, 16 June, B:4.

Anonymous (1859) Memoranda: The Hawaiian bark Cambia. The PoZynes%cz, 13 August.

Anonymous (1867) T u r t l e ' s eggs. Pacific Comercia2 Advertiser, 29 June, 3:5.

Anonymous (1887) Hawaiian almanac and d i r e c t o r y : Wreck of t h e Dunnottar Castle - Annexation of Ocean I s l and . 99-101.

Anonymous (1918) Cruise of t he U. S. S. Hemes among i s l a n d s of t h e Hawaiian group, 1918. Record Group 45, Nat ional Archives, Washington, D. C. 24 pp.

Anonymous (1923) On H a w a i i ' s nor thwest wonder i s l a n d s . HomluZu Sta.r-BulZetin, 2 1 J u l y , 3 : l .

Anonymous (1927) F ish ing and s h e l l hunt ing exped i t ion r e t u r n s from P e a r l and Hermes Reef. The HonoZuZu Advertiser, 13 May, A: l .

Anonymous (1930) Large ca t ch of haha le lu made a t Anahola Beach. !The Carden IsZand (Lihue, Kauai), 9 September, 1.

92

571.

5 7 2 .

Anonymous (1935) 445 pound t u r t l e caught by sampan. H O Y W Z U ~ Star-Bulletin, 8 A p r i l , 3.

Anonymous (1936) Aircraft advanced base r e p o r t of the U. S. S. Chi lds : Report of t he survey of P e a r l and Hermes Reef and Ocean I s l and . Record Group 37, Nat iona l Archives, Washington, D. C.

573.

574 .

575.

576.

577.

578.

579.

580.

Anonymous (1951) The Laysan I s l and cyclorama and la te news from Laysan. The EZepaio, 1 2 , 1:l-2.

Anonymous (1956) T e r r i t o r y hopes t o expand marine t u r t l e popula t ion . Honolulu Star-Bulletin, 9 March, 14.

Anonymous (1956) 3 b i g t u r t l e s flown t o Vancouver zoo: Flip-flopped t u r t l e s ready f o r journey. HonoZuZu S-tur-BuZletin, 1 2 June, A:12 .

c

Anonymous (1958) Aquarium g e t s rare t u r t l e , s h e ' s moo ted by o l d t enan t s . Honolulu Star-Bulletin, 30 May, B:17.

Anonymous (1960) Aloha t u r t l e s go t o Canada. HonoZuZu Star-Bulletin, 29 August.

Anonymous (1962) 1 sea t u r t l e found dead,3 dying i n scummy water a t Marineland. Honolulu Star-Bulletin, 23 February, 1 7 .

Anonymous (1966) B i o l o g i s t s p u t 'X ' on t u r t l e s . The HonOZuzu Advertiser, 30 Apr i l , C : 4 .

c

Anonymous (1966) H a w a i i ' s o t h e r i s l a n d s . Photographs by W. Roll, Honolulu Star-Bulletin, 30 November, B : 3 ; 12 December, C:2; 22 December, D:5; 24 December, A:9.

581.

582.

Anonymous (1967) 764-pound t u r t l e caught a The Honolulu Advertiser, 1 7 November, A : l .

Anonymous (1969) Alexander 's Nursery ... a weal th of cur ious and e x o t i c c o l o r f u l p l a n t s . The Garden Island (Lihue, Kauai), 1 December, B:2.

563. Anonymous (1970) Ou t - tu r t l i ng a t u r t l e . Honolulu Star-BuZZetin, 3 March, B:12.

584. Anonymous (1970) Laysan w i l d l i f e t h r i v i n g - t r e s p a s s e r ' s j ou rna l . The HonoluZu Advertiser, 14 October, C : 4 .

585. Anonymous (1971) Growing wi th Kauai. The Garden Island, Annual Edi t ion (Lihue, Kauai) , 1 5 March, A: 3.

586.

587.

Anonymous (1971) Kahuku: August 28, 1971. The Elepaio, 32, 4:39.

Anonymous (1971) 18 October 1971. The EZepaio, 33, 1:8-9.

93

588.

589.

590.

591.

592.

593.

594.

595.

596.

597.

598,

599,

600.

601.

602,

603 ,

604.

Anonymous (1972) B i o l o g i s t s tudying sea t u r t l e p o t e n t i a l on gourmet market. Co Zmbia Record, 16 September .

Anonymous (1972) B i o l o g i s t i n s t e w over t a s t y t u r t l e . Dallas Times Hera td , 18 September.

Anonymous (1972) Hawaiian crow becoming scarce. The HonoluZu Advertiser, 29 November, B : 7.

Anonymous (1972) Crowned wi th a comb. HonoZuZu Star-BuZZetin, 6 December, C:2;

Anonymous (1972) Rare and endangered spec ie s of w i l d l i f e of H a w a i i . lnrle EZepaw, 33, 2 : 15-16.

Anonymous (1973) The Big I s l a n d . Destination Hawaii, East/West Network, 1, 4 .

Anonymous (1973) T i m e t o s t o p t u r t l e t r a d e i n H a w a i i . Oryx, October, 160-161.

Anonymous (1973) F i r s t s t e p taken t o p r o t e c t sea t u r t l e s . HonoZuzu Star-Bulletin, 1 3 January, A: 2.

Anonymous (1973) Crowding problem solved: T u r t l e s r e tu rned t o the sea. The HonoZuZu Advertiser, 31 January, A:14.

Anonymous (1973) T u r t l e s , sheep and deer i n advisory pane l ' s vo te . The HonoZuZu Advertiser, 1 0 February, A:15.

Anonymous (1973) 11 t u r t l e s t o g e t r i d e o u t t o sea. The HonoZuZu Advertiser, 7 March, C:3.

Anonymous (1973) T u r t l e ' s t he i s s u e . m e HonoZuZu Advertiser, 2 1 March, B:2.

Anonymous (1973) Turt le-hunt ing r u l e s o f f e red . The.Honolulu Advertiser, 11 August, A: 1 2 .

Anonymous (1973) P r o t e i n from the sea. Los AngeZes Hera ld -Edner , 22 August, C:9.

Anonymous (1973) Hearing set on sea t u r t l e r egu la t ions . HonoZuZu Star-BuZZetin, 13 September, A:8.

Anonymous (1973) Marine b i o l o g i s t George H. Balazs wi th a hawksbi l l sea t u r t l e , now an endangered spec ie s . MiZwaukee SentineZ, 1 November.

Anonymous (1973) T o u r i s t s ' a p p e t i t e s t h r e a t t o t u r t l e s . HaZifm ChronicZe, 5 November.

94 c

605.

606.

607.

608.

609.

Anonymous (1973) Notice of pub l i c hear ing. Honolulu Star-Bulletin, 9 November, D:8 .

Anonymous (1973) T u r t l e hear ings. The Honohlu Advertiser, 10 November, D:3.

Anonymous (1973) Green t u r t l e colony faces e x t i n c t i o n . The Detroit News, 22 November, G:2 .

Anonymous (1973) I n T-sh i r t campaign: F l a t ou t f o r t u r t l e s . Honolulu Star-Bulletin, 3 December, A:13.

Anonymous (1973) I t ' s up t o you: A weekly l i s t i n g of pub l i c hear ings and open meetings. Honolulu Star-Bulletin, 3 December, c:20.

610. Anonymous (1973) Kauaians urged t o he lp save sea t u r t l e s from e x t i n c t i o n . The Garden Island (Lihue, Kauai), 3 December, A:8.

611. Anonymous (1973) Over she goes. Honolulu Star-Bulletin, 1 3 December, A:23.

612 - Anonymous (1973) Tour i s t t r ade p e r i l s green sea t u r t l e s . Washington Stur-News, 14 December, A:22 .

613. Anonymous (1973) Sea t u r t l e tagged, r e l eased o f f WW shore. Windward Sun Press (Oahu), 19-25 December.

614.

615.

Anonymous (1973) 11 December 1972. The Elepaio, 33, 11:125-126.

Anonymous (1973) Countdown a t Kohala. Hawaii Business, 18, 6: 23-24, 27-30, 33-34, 37.

616. Anonymous (1974) Hawaii 's concern f o r i t s t u r t l e s . Sea Secrets, 18, 1:12-13.

617. Anonymous (1974) The Anchorage. Neighbor Islands: This Week, 9-15 January, 96.

618. Anonymous (1974) T u r t l e p r o t e c t i o n i s delayed. HornZulu Star-Bulletin, 5 February, C:5.

Anonymous (1974) P u b l i c hear ing. Ka Molokai, 2 1 February. 619.

620. Anonymous (1974) Buzz's Wharf. Neighbor Islands: This Week, Maui, 24-30 Apr i l , 38.

621. Anonymous (1974) New t u r t l e r u l e s e f f e c t i v e May 30. f i e Carden Island (Lihue, Kauai) , 27 May, A : 6.

622. Anonymous (1974) Marine t u r t l e r egu la t ions begin soon. Honohlu Star-Bulletin, 28 May, C:5.

95

x

623.

624,

625.

626.

627.

628

629.

630 .

631.

632.

633.

634.

635.

636.

637.

638.

639.

Anonymous (1974) To mom wi th love: Genuine t u r t l e s k i n handbag now $160.00. HonoluZu Star-Bulletin, 5 August.

Anonymous (1974) Green Garden Res t a u r a n t . Neighbor Is~ands: This Week, Kauai, 21-27 August, 112.

Anonymous (1974) The Ri tz : Exot ic c ros s c u l t u r e jewel ry . f i e Sundag Star-Bulletin 8 Advertiser (Honolulu), 20 October, 5.

Anonymous (1974) Woman i s f i n e d i n f raud case. !The HornZulu Advertiser, 22 November, A: 1 7 .

Anonymous (1974) Consul Genera l ' s w i f e f i ned - bus iness f raud . Sun Francisco Chronicle, 23 November.

Anonymous (1975) F i r s t case i n H a w a i i : Kauai cour t f i n e s poacher and r e s t a u r a n t . f i e Garden Island (Lihue, Kauai), 20 January, 1.

Anonymous (1975) Rare hawksbi l l t u r t l e found i n t i d e pool . Honolulu Star-Bulletin, 1 7 January, A:14.

Anonymous (1975) I n s a f e hands. The Hono l u h Advertiser, January, A : l .

Anonymous (1975) More t o green s e a t u r t l e than s h e l l . HornZulu Star-Bulletin, 27 February, E : 2 .

Anonymous (1975) Seafood d inner menu: A t H a w a i i ' s most unique garden-res taurant , Pagoda F loa t ing Restaurant . Honolulu Star- Bulletin, 15 Apr i l .

Anonymous (1975) Bird pa rad i se i n memory of Lindbergh. The &lay Mail, 2 May, 16.

Anonymous (1975) U. S. F ish & W i l d l i f e Serv ice , Por t land , Oregon, news r e l e a s e , 23 May 1975: Sea t u r t l e s to be a d d e d . t o Threatened list. The EZepaio, 36, 2:24-25.

Anonymous (1975) Real e s t a t e f o r sale: Sea t u r t l e s . The Sunday Star-BulZet&z & Advertiser (Honolulu), 1 3 J u l y

Anonymous (1975) Sea L i f e Park. Sea L i f e Park, Makapuu P o i n t , Oahu, H a w a i i , 26 pp.

Anonymous (1976) Backdoor. Swzbwns (Honolulu), 8, 7:15.

Anonymous (1976) Aquaculture d i s p l a y set. HonoluZu Star-Bulletin, 1 January, E:22.

Anonymous (1976) F ish ing boa t breaking up. HomluZu Star-BuZZetin, 25 February, A:4.

96

646 I

641.

642.

643.

644.

645.

646.

647.

648.

649.

650.

651.

682.

653.

654.

655.

Anonymous (1976) Hawksbill is no slowpoke a t t h e t a b l e . Honolulu Star-Bulletin, 14 A p r i l , A: 6 .

Anonymous (1976) Midway t u r t l e mystery. The Islander (U. S. Naval S t a t i o n Midway), 6 August, 1.

Anonymous (1976) Green sea t u r t l e s tudy w i l l begin t h i s month. HomluZu Star-Bulletin, 11 September, B: 16.

Anonymous (1976) Popula t ion explos ion . Honolulu Stur-Bulletin, 18 October, C:16.

Anonymous (19 76) 715-Misc. f o r sale : Col l ec to r s , deco ra to r s : Endangered s p e c i e s t u r t l e s h e l l s . Honolulu Stur-Bulletin, 9 November

Anonymous (1976) Hawaiian ep icu re : A menu guide t o Oahu, Kauai, H a w a i i and Maui. Peanut Bu t t e r P a r t n e r s , S e a t t l e , 240 pp.

Anonymous (1977) Rescue of crewmen under way: O i l t anker breaks up o f f Midway. Honolulu Star-Bulletin, 18 January, A : l .

Anonymous (1977) Tanker endangers i s l a n d . Desert Sun (Palm Springs, C a l i f o r n i a ) , 19 January.

Anonymous (1977) Dead f i s h wash ashore a t Kualoa. Honoluzu Star- Bulletin, 31 January, A : 1 4 .

Anonymous (1977) The Midway grapevine: T u r t l e l e c t u r e . The Islander (U. S . Naval S t a t i o n Midway), 27 May, 10.

Anonymous (1977) W i l d l i f e assessment t o cover Northwestern Hawaiian I s l a n d s . The Sunday Star-Bulletin & Advertiser (Honolulu), 14 August

Anonymous (1977) Survey and assessment of green sea t u r t l e r e source of t he Northwestern Hawaiian Archipelago. Act ivi t ies i n r e sea rch , educa t ion f o r f i s c a l y e a r 1977-1978. Univers i ty o f H a w a i i , Sea Grant College Program, UNIHI-SEAGRANT-MB-78-01.

Anonymous (1977) The o u t e r o u t e r i s l a n d s . Honozulu, 12, 2:34-37.

Anonymous (1978) Treasures of t he P a c i f i c : A r t i f i c i a l c u r i o s i t i e s of t he 18 th century . Bernice P . Bishop Museum, 21 pp.

Anonymous (1978) S k i e r s complete San Diego- to-Waikiki c ros s ing . Los Angeles E m i n e r , 1 January.

Anonymous (1978) Environmental s e t t i n g and h i s t o r y of Midway I s l a n d a lbacore tuna development p r o j e c t . S t a t e of H a w a i i , O f f i c e of t h e Marine A f f a i r s Coordinator, unpublished r e p o r t , 32 pp.

c

97

656.

657.

658.

659.

660.

661.

662 ..

663.

664.

665.

666.

667.

668.

669.

670,

Anonymous (1978) The seasons of Mahina: A Hawaiian f i s h i n g and farming moon ca lendar 1979. Honolulu.

Graphic A r t s and Letters Unlimited,

Anonymous (1978) Rare behaviora l t raits , Sea Secrets, 22, 3:8-9.

Anonymous (1978) T u r t l e in format ion needed. Hawaii Coastal Zone News, 3, 1:8.

Anonymous (1978) A meticulous i n v e s t i g a t i o n i n t o the v i c t u a l s of t h e s e peoples . The Honolulu Advertiser, 14 February, 1:28.

Anonymous (1978) W i l d l i f e and geology of t h e Hawaiian Leeward I s l a n d Chain, Hawaii *ibune-Herald, 28 June, 4 .

Anonymous (1978) Marine l i f e l e c t u r e : B i o l o g i s t speaking tomorrow on Leeward I s l ands . West Hamii Today, 30 June, 7 .

Anonymous (1978) TASS he lps w i t h s e a l i f e research . The Hawaiian Falcon (Hickam A i r Force Base), 18 August, 4 .

Anonymous (1978) W i l d l i f e and geology of the Leeward I s l a n d s . The &vden Island (Lihue, Kauai), 11 October, B:8.

Anonymous (1978) Isle tuna f l e e t g e t s OK t o use Midway. Hon~ZUzu Star-Bulletin, 17 October, A : 1 8 .

Anonymous (1977) RCUH annual r e p o r t f i s c a l 1977. The Research Corporat ion of t h e Univers i ty of H a w a i i , Honolulu, 36 pp.

Anonymous (1978) RCUH annual r e p o r t f i s c a l 1978. The Research Corporat ion of t he Univers i ty of H a w a i i , Honolulu, 39 pp.

Anonymous (1979) The odd couple. National WiZdZife, 17 , 2:16.

Anonymous (1979) H a w a i i warned on t u r t l e f i s h i n g . flonolulu Star- Bulletin, 24 January, C:14.

Anonymous (1979) Sea t u r t l e s off l i m i t s . The Sunday Star-Bulletin dl Advertiser (Honolulu), 4 March, A:5.

Anonymous (1979) T u r t l e imports t o U. S. ha l t ed . Focus (World W i l d l i f e Fund-U.S.), 1, 2:6.

c

98

SUBJECT INDEX

c

HAWKSBILL (ERE!lYiOCHELYS IEBRICATA): 30, 31, 37, 46, 47, 55, 62, 76, 78, 80, 105,110, 176, 201, 202, 217, 258, 262, 264, 280, 299, 333, 348, 350, 360, 379, 401, 440,

447, 457, 460, 461, 462, 465, 468, 491, 492, 530, 556, 557, 591, 594, 603, 629,

630, 636, 640, 653, 661. s

LEATHERBACK (DEIWOCHELYS CORIACEA): 46, 47, 55, 80, 262, 264, 299, 350, 357, 379, 457, 460, 461, 465, 468, 489, 492, 571, 576, 581, 594, 661

LOGGERHEAD (CARETTA CARETTA): 26, 38, 85, 262, 351, 379.

OLIVE RIDLEY (LEPIDOCHELYS OLIVACEA) : 262, 379, 459 I*

' CAPTIVE MAINTENANCE AND AQUACULTURE: 4, 5, 13, 15, 17, 18, 22, 34, 52, 59, 60, 67, 63, 83, 87, 88, 89, 98, 99, 100, 101, 118, 119, 120, 129, 130, 131, 134, 159,

164, 174, 183, 192, 193, 196, 206, 215, 221, 234, 259, 262, 264, 274, 278,

303, 340, 344, 407, 413, 420, 421, 422, 452, 459, 461, 462, 466, 467, 476,

477, 479, 494, 503, 504, 535, 544, 575, 576, 577, 578, 588, 596, 598, 601,

611, 613, 615, 616, 636, 638, 643

c

COMMERCIAL CATCH STATISTICS: 47, 72, 176, 262, 264, 447, 454, 455, 456, 460, 462, m

500, 501, 502, 505, 506, 507, 508, 509, 510, 511, 512, 513, 514, 515, 517,

518, 519, 520, 521, 594

OAPTURE TECHNIQUES: 47, 64, 85, 176, 204, 209, 229, 249, 264, 281, 339, 354, 416, *

457, 458, 459, 460, 486, 559, 583, 593

HAWAII STATE LEGISLATION (BILLS AND RESOLUTIONS): 3 , 154, 155, 156, 157, 163, 164, 305, 328, 329, 337, 403, 553

99

HAWAIIAN CULTURE : 35, 41, 42, 72, 105, 109, 127, L37, 144, 145, 14h,

176, 185, 186, 190, 200, 212, 213, 214, 244, 247, 248, 249, 25i, 252, 254,

262, 279, 281, 331, 341, 345, 346, 347, 348, 376, 377, 425, 426, 427, 430,

434, 450, 471, 484, 485, 489, 544, 656, 659

HAWAIIAN LEGENDS: 106, 178, 179, 207, 224, 225, 226, 227, 239, 285, 423, 424,

431,446, 488, 539, 540, 543

P A C I F I C OCEAN BIOLOGICAL SURVEY PROGRAM (POBSP) : 25, 26, 27, 28, 29, 92, 93,

96, 166, 167, 168, 170, 187, 188, 189, 223, 245, 250, 405, 444, 445, 451,

552

E P I Z O I C S AND DISEASES: 20, 71, 76, 85, 134, 205, 237, 255,'256, 257, 270,

351

NATURAL PREDATION: 20, 47, 49, 51, 62, 70, 71, 72, 73, 79, 86, 13'1, 230, 262,

265, 275, 350, 351, 399, 480, 481, 482, 487, 490, 492, 557, 658

100

Table 1. Number of Chelonia tagged throughout t he Hawaiian Archipelago s i n c e 1973.

Locat ion Immature Adult T o t a l male female

Kure A t o l l 2 1 21 1,

65 1 66 Midway I s l a n d s

Pearl and Hermes Reef 5 4 9

Lisians k i 23 3 7 33

Laysan 1 1 2

Mar0 Reef

Gardner P innac les

French F r i g a t e Shoals

Necker

Nihoa

Kaula

Niihau/Lehua

Kauai

Oahu

Mo 1 oka i

Lanai

Kahoolawe

Maui

H a w a i i

R

c

130

7

122 483 735

4 6 17

4 2 5 11

2 2

75 2 6 83

28 1 1 30

1 1

8 7 0 5 92

T o t a l 444 140 5 18 1102

x

F

101

. rn d (d 0 % al U (d M .rl k k

2 8 k k U (d

rn al d U k 3 U

d al a, k M al

1 a,

W

U rl 3 a (d

ICI 0

5

3 5

a *rl 3 a

M E al rl

a 0 (d a (d 0

U

*rl (d Fc U cn

2

5

N

a rl P (d I3

4 VI

* u3 03

I N

N \o

rl

N h

N VI

* m 0 4

I m 0 00

.

"! N m

m h m rl

m m

W

VI 03

I VI m W

rl

N h

rl m

N

VI rl rl

h

00 h

I m * W

m 0 h

m rl rl

e

GI I 0

y\ 00

h

N m .

e h m rl

\ o c u 0 - 0

\o

4

e h

I W

00 W

m 0 h

h

00

m m I

\o

h 00

0 m m

ro h m rl

W m

m VI h

I m m VI

.

U 4 h

.

\o m

m 0 0 4

I 00

0 00

m m 00

h h m rl

u) U

VI

rl 00 I 4

0 h

.

.

m 0 h

ro e

\o

d 0 rl I 0 N 00

.

.

N

N m

00 h m rl

N m

N

00 h

I VI m ro

\o

rl h

.

3

m N 0 4

I e 00 00

h

m m .

m h m rl

m h cr)

4

\o 00 I m m VI

e rl h

m h m

N

u) 0 d

I 00

Q 00

.

N

N m

rl rl td ld E

102

a

E 1 V

m aJ rl P

8

-5 U

d N 0 rl I 0

0 03

h

0 m

U U

m h 0 rl I rl

co co

m h m

m 3 -I

h

co 0 rl

U m 0 rl

I \D

N m

rl

d m

03 0 I4

0

m d - 4 I

N

0 m

co h m

U h m rl

cn m d

rl

N 0 rl I 0

0 m

m 0 o\

vr m rl

0

al 0 d I rl

cn co

.

m \o m

rn h m rl

m 4

rl

N 0 rl

I m

m

m m 00

00 rl

h

m 0 d

I N

m m

m h m

\o h m rl

h m

co I- o\ I

W

rl W

.

m 0 m

h m

h

\o 0 rl I

N

rl m

vr I- m

h h m rl

<

I-

\o 0 rl I m rl 00

N

rl m

\o U

N

I- O rl I h

m co

m I- m

co h m rl

\o N

0

rl 0 rl I m m m

N

N m

h hl

vr co 0 rl I h

N m .

d

m m

m h m rl

N rl U

h

\o 0 rl

I 0

0 co

N

rl m

vr rl U

0 e3 rl rl I d li7 00

m h m

rl rl cd k

6

c

c

c

L

103

L

Table 4. Summary of growth rates and p ro jec t ed y e a r s t o ma tu r i ty for green t u r t l e s sampled a t r e s i d e n t fo rag ing areas i n t h e Hawaiian Archipelago.

~ - ____I .-. .

Growth rates Years t o N o , of Range of- cm p e r month,' ma tu r i ty (35-92 cm) growth i n t e r v a l s

Location Range Mean Range Mean measurements i n months - -^_ .._ - -__...__- --

Main I s l a n d s Kau, H a w a i i .38-. 52 .44 9-12 10.8 4

No. cap tures--79 No. turtles---72

7-17

Bellows, Oahu .19-. 21 .20 22-25 23.8 2 13-22 No. cap tures--24 No. turt les---21

Nor thwes t e r n Is l ands

No . captures---9 N o . turt1,es---- 7

Necker . 1 4 33.9

French F r i g a t e .02-. 1 3 '.OS 36-237 59.4 Shoals

No. captures-214 No. tu r t les - -130

L i s i a n s k i No. cap tures--30 N o . turt les---23

.13 36.5

Midway .03-. 2 1 .09 23-158 52.8 No. captures-293 No. tu r t les - -250

1

'19

20

3-36

2

6-37

Kure .04-. 12 .08 40-119 59.4 2 13-24 No. captures--23 No. turt les---21

.__- r_ . . ... .. .. . . - . . . ._.... ,.,,. ~~

* s t r a i g h t carapace l e n g t h

104

Table 5. Number of egg c l u t c h e s l a i d by green t u r t l e s a t East I s l a n d , French F r i g a t e Shoals.

c’

- 1975 1974-197s 1974 No. of egg No. of No. of No. of

c l u t c h e s t u r t l e s % turtles % _I_____ t u r t l e s %

0 1 3 12 .1 7 7.0

1 31 2 8 . 7 53 53.0

20 9.6

84 40.4 c

2 20 18.5 34 34.0 54 26.0

3 26 24.1 6 6.0

4 12 11.1 0 0

I

32 15.4

12 5.7

5 5 4.6 0 0 5 2.4

6 1 0.9 0 0 1 0.5 c.

1.08 100.0 100 100.0 208 100.1 To t a l

Mean 2 . 1 1 . 4 1.8

c

c

c

105

Table 6. I n t e r n e s t i n g i n t e r v a l s of green t u r t l e s a t East I s l a n d , French F r i g a t e Shoals.

1974 1975 1974-1975 I n t e r v a l N o . of No. of No. of

i n day i n t e r v a l s % i n t e r v a l s % i n t e r v a l s %

11 6 8.0 1 6.7 7 7.9

1 2 1 7 23.0 4 26.7 2 1 23.6

1 3 26 35.1 2 13 .3 28 31.5

1 4 13 17.6 5 33.3 18 20.2

15 8 10.8 2 13 .3 10 11 .2

16 2 2.7 1 6.7 3 3.4

ii 1 1 . 4 0 0 1 1.1

18 1 1.4 0 0 1 1.1

T o t a l 74 100.0 15 100.0 89 100.0

Mean 13 .2 days 13.4 days .13.2 days

Table 7. R e s u l t s of 40 precounted egg c l u t c h e s excavated and examined a t East I s l a n d fol lowing t h e n a t u r a l emergence of h a t c h l i n g s .

Y

Standard Mean d e v i a t i o n Range

% of eggs hatched 76.7 24.2 0-100

% of h a t c h l i n g s emerging at t h e s u r f a c e

% dead h a t c h l i n g s i n t h e n e s t

W of eggs wi th p a r t i a l l y developed b u t dead embryos

% of eggs wi th no apparent development (presumably i n f e r t i l e )

70.8 24.0 0-97.6

5.9 9.4 0-52.1

10.8 9.9 0-50.0

12.5 22.4 0-100

106

a al 3

w m a l M k

aCd d U & ( d ( d rww

8

' 0 2 3

5

al U a M .r(

k n 0 m N Y rl d

.

m N .

n m (v v m

.

0 rl .

n h

h

N

. U

h

h

n n 4 0 h N . .

m

I

c,

3 l-4 m H U m (d w U (d

a a, &

0 U al k

R rl U c al 7

s s v)

a d (d

a al M M (d U

rl

m . 0 3

l-4 rl c

r l r l m r l

r l r l d r l 4

0 0 rl

m 0 rl

\o N

U U

m h

(v \o

. 03

al rl

w % m h m rl

m h m rl

h h o\ rl

03 h m PI

m h o\ rl

h m rl

c

10 7

Table 9 . Food sources recorded for green t u r t l e s i n the Hawaiian Archipelago.

Location Principal constituents Other food sources

Hawaii R* - Pterocladia capillacea Amansia glomerata

Yaui R - Pt,nrocludiu capillacea

Kahoolawe ? R - Gelidiwn s p .

Lanai

Oahu

Kauai

Necker

R - Amansia glomerata R - Hypnea cemicornis

B - Sargassum polyphyllum LeveilZea jungermannioides Acant hop b r a spici f era Champia parvula

Gracilaria sp . G - Codiwn phasmaticum B - Dictyotu acuteloba

Padina japonica A - Halophila hawaiiana

G - Codim Bdule Codiwn arabicwn Codiwn phasmatimm Ulva fasciatu Ulva r e t i a l a t a

R - PterocZadia capillacea Armnsia glomerata AhnfeZtia concinna

R - Pterocladia capillacea

G - Caulerpa racemsa

R - GeZidiella acerosa Gelidiwn pusiZlwn GraciZaria coronopi fo t i a Gracilaria bursapastoris Hypnea musciformis Hypnea chorducea Hypnea cernicornis Acanthophora spicifera Grateloupia. f izicinu

G - Caulerpa serrutata CauZerpa ssrtu lurioide s Dictyosphaeria versZuysii Enteromorpha s p . Cladophora fasicu Zaris

B - Dictyota acuteloba Padina japonica

A - Halophila hamiiana T - Chondrosia chucalla

Spongia Oceania Brae hidon t e s ere br i s tr ia tu s

BG - Lyngbya majuscuZa

G - Halime& discoidea R - Ammtsia glomerata

Hypnea s p .

108

Table 9 (cont inued) ~.

Locat ion P r i n c i p a l c o n s t i t u e n t s

French F r i g a t e G - Codiwn arabicwn Sho a 1s Codiwn phasmaticwn

Codiwn eduZe CauZerpa racemosa UZva fasciata

B - Turbimria ornata R - Spyr id ia fiZamentosa

Lays an

L i s i a n s k i

?

G - CauZerpa racemosa B - Turbinmia o m t a

Other food sources

B - Lobophora variegata Rosenvingea orientaZis Sphacelmia tribuZoides Roschera s p .

R - GraciZaria new spec ie s Porolithon gmdineri Ceramiwn s p . Po Zysiphonia s p . Liagora s p .

Ch Zorodesmis h i Zdebmndtii Ha lime& discoidea

BG - MicrocoZeus Zyngbyaceus

G - Microdictyon setcheZZianwn P

I - Physalia physalis C e r i t h i i d a e An t h r o zo a

D - AsterioneZZa notata

R - Spyridia fiZarnentosa Amansia gZomerata

G - Microdictyon japonicwn Pssudabryo s i s oahuensis Hazimeda 3 zscoidea

B - Turbinaria ornata Padina crassa Zonmia variegata

R - Ge Zidiopsis s p . PterocZadia capiZlacea Jania capi Z laced: Po Zysiphonia s p .

G - Halimeda discoidea ChZoreZZa sp .

BG - Lyngbya majuscula OsciZZatoria sp .

I - VeZeZZa veZeZZa Janthina exigua

D - Melosira sp.

*-

a*

Y 109

Table 9 (continued)

Location P r i n c i p a l c o n s t i t u e n t s ^__- Other food sources

Midway

m

Kure

G - Codiwn eduZe G - Codiwn coneatwn R - Spyridia filamentosa Caulerpa serrulata

CauZerpa sertuarioides Dictyosphaeria vers luys i i

R - Amansia glomerata Ceramiwn sp . FaZkenbergia s p . Padina s p .

Oscillatoriu sp . A - Halophila hawaiiana I - Physalia physalis

VelelZa ve le t la Janthina exQua Echinoidea

B - Sphacelaria tribuloides

BG - Lyngbya majuscula

G - Codiwn edule R - Polysiphonia tsudana I - Physalia physalis

Janthina exigua Echinoidea

* R - r e d a l g a e , Rhodophyta B - brown a l g a e , Phaeophyta G - green a lgae , Chlorophyta BG - blue-green a lgae , Cyanophyta D - diatoms, Bac i l l a rophy ta A - angiosperm ( seag ras s ) I - i n v e r t e b r a t e s

Algae i d e n t i f i e d by D. J. Russe l l and M. S. Doty; i n v e r t e b r a t e s i d e n t i f i e d by W. J. Cooke; s e a g r a s s i d e n t i f i e d by K. Bridges.

P

110

Table 10. Major food sources of green turtles in the Hawaiian Archipelago.

_ _ _ _ I _ _ _ - _ . _ _ _ ~ _ I Benthic algae Location

Main Islands (sample size N=85)

Main Islands and Northwestern Hawaiian Islands

Northwestern Hawaiian Islands (sample size N=56)

~ f i - Pterocladia capillacea Aman s i a glomera ta

G - Codiwn edule L'odiuni arabicwn Codiwn phasmticwn Ulva fasciata

G - Caulepa racemsa R - s p y r i d i u filamentosa B - Turbinaria ornata

* R - red algae, Rhodophyta G - green algae, Chlorophyta B - brown algae, Phaeophyta

P

Y

L

c

111

*

h u *rl 3 *rl U

U

C *rl

a 0) m .rl (d k

a c (d

h U .rl 3

*I4 u a (d U

%

k w a aJ

0) rl aJ k

m a, rl U k 1 U

3

3

9 2 ti

aJ k bo

*rl .rl

X w 0

m Q) u (d k

5 50 k c3

rl rl

aJ rl P (d H

0 3 0 o r l c a m rl b N m N 4 . .

oca O U O h m m w hlr l r l co co F m U O U rl U

. . . . .

0 m m

Q) M (d

w o h

"8 f i U

ai

m a G U O m

I-- rl

U CrJ

n N I- \ co W

4 rl 0 N

rl N O m

N

h In

0 m m

0) M (d

w o h

"8 m a v G U o m

I-- rl U (d

n m I- \ 0 rl

m rl 0 hl

W

a 0 (d rl m H

G k aJ

E-c

G 0

rl aJ c C 0 v) k aJ a a k (d

3 U m (d 0 u

m

=; h P m

i? U m

% .rl W

a aJ w h rl k (d rl 1 bo aJ k U

11.2

Table 1 2 . Growth rates of a d u l t green t u r t l e s recovcred a t French F r i g a t e Shoals s i n c e June 1973.

___----__ S t r a i g h t -carapace l eng th , c m - I n t e r v a l Growth rate

i n months i n c m p e r month __--- --I_

Tag no. I n i t i a l Recovery

Females

699Fw, 642

362Fw, 648 391FW, 779 T60FW, 723 976FW, 763 655FW, 828 959Fw 650, 1075 1029, 2605 1047, 2729 958FW, 2787 2235, 3145 2209, 2210 842, 2270 2253, 3228 2268, 3257 2255, 3266 5016, 5018 651, 3264

372na, 641 94.9 89.5 92.1 94.6 88.3 94.6 90.2 95.3 95.3 94.0

100.6 97.0 90.8 89.8 94.6 92.7 90.8 90.2 90.5 87.0

95.3 90.2 95.3 97.8 91.4 96.5 92.7 94.0 94.0 95.6

101.6 98.1 91.8 90.8 95.6 92.7 91.8 90.5 92.7 87.9

60 35 35 47 25 37 75 49 24 36 36 73 24 24 24 2 4 24 24 25 72

. 0 1

.02

.09

.07.

.12

.05

.03

- .04 .03 .02 .04 .04 .04

0 .04 . O l .09 . 0 1

747FW, 2616 81.3 82.6 64 .02

Mean 91.8 range 81.3-100.6

.04

N=18 24-75 .01- .12

C

a

c

113 x

Table 1.3. Growth rates o f Hawaiian green turtles captured from the w i l d and h e l d for extended periods i n capt iv i ty a t Sea Life Park (diet--fresh frozen

rr squid and f i s h ) .

Interval Growth rates Straight carapace lengths, cm i n months i n c m per month

Tag no. 1 2 3 1 2 1 2

Immn Lure

2057

2067

2014

2065

2071

2018

Adul t Females

2021

2013

684

2017

2062

Adul t Males

2015

2019

2069

n

-

4.

x

-.

".

69.2

62.2

63.5

64.8

76.8

73.3

85.7

86 .4

94.6

92.7

85 .1

78.1

82.6

78.7

70.8

69.2

68.6

73.6

84.8

80.3

87.6

86.7

95.6

93.7

86.0

78.7

84.5

79.7

80.0 9

79.4 9

79 .1 9

1 3

89.2 1 3

84.8 1 3

59

59

35

46

46

59

35

46

1 3 .18 .71

1 3 .78 .78

37 .57 .28

.68

24 .62 .18

24 .54 .19

.03

.01

.03

.02

.02

. 0 1

.05

.02

114 c

c

m 0 0 m h \o W r-. l-i m m

rl

L

n cd 3

I 4 3

n

2 43 c d f d 5 0

c

0 m m rl rl hl a3

c Pl

a aJ k

a J a J U ? ( d o n u aJ

k

m h * I h rl I rl rt

m In h m

b I U

h I I N

\o rl

co m m

\o m In

n m m h h I I rl rl

m rr) h h I I rl m

m h I rl

9 H c

B b U \o rl \o m

0

m ul

8 a aJ m

c \o 03 rl 0 m m

0 03 rl rl m hl

P 115

Table 15. Locations and numbers of t a g r e c o v e r i e s documenting long d i s t a n c e migra t ions by a d u l t green t u r t l e s i n t h e Hawaiian Archipelago (sources - Balazs 57 and unpublished d a t a , Kr id l e r 31.1-327, Olsen 380-385) -

~

Males Femo 1 e s Tota l Tagging and recovery l o c a t i o n s

French F r i g a t e Shoals t o : Kauai 1 Oahu 2 Mo l o ka i 1 Ma u i 0 H a w a i i 1

9 11

4 2 2

Lis i a n s k i Pearl & H e r m e s Reef

0 1.

2 3

2 4

H a w a i i t o : French F r i g a t e Shoals 0

Oahu t o : French F r i g a t e Shoals 1

Niihau t o : French F r i g a t e Shoals 0

L i s i a n s k i to : French F r i g a t e Shoals 0 4

Laysan t o : French F r i g a t e Shoals 0 1

1 0 P e a r l & H e r m e s Reef t o :

French F r i g a t e Shoals 3

Pearl & H e r m e s Reef t o : Laysan 0 1 1

French F r i g a t e Slroals t o main i s l a n d s 5 Main i s l a n d s t a French F r i g a t e Shoals 1

Tota l 6

23 2

25

28 3

31

French F r i g a t e Shoals t o nor thwes tern i s l a n d s 1 Northwestern i s l a n d s t o French F r i g a t e Shoals 3

T o t a l 4

5 12 17

6 15 21

A

11 6

a 8 $ 8 a, &

a, u (d cl

e h m rl

3 c,

3

m I h

Y rn Fr k

u

W

*

3

a I (II c a, X a rl & (d a, PI

w w m rl

&

4 a, u

VI

W N

e

e m h

$ h m -01 rlh

w h m rl

h rl 1 c, 0 rl

w I- m rl

h rl 1 c, 0 rl

m 1

m

a I rn r4 Fr

bo *I4 &

I

E - c .

rn I

i a,

X 4

rl $4

% !

w w tn rl

& a, .o

u a a, VI

0 N

8

m 0 m rl

2 cn rl

3 c, N

m I 0 rl

T 5=:

3

rn

* u

W

a I v)

E 2l a rl & (d a, PI

4 w h rl

5 3 m rl

W h m rl

z @-J

h

m I h

7 E

8

r4

u

W

*

a 1 v) c a , . X 4

rl & fa 4

m w m rl

rl rl &

2 rl cn N rl

$ m oo co

m h m rl

a,

I-, 4

B .I4 (II

a a,

* 0 L) a, &

v) a, rl u & 1 u d a, a, & bo

3

8 4 4

B X u rl 3 a (d

W 0

UI El 0 rl U (d &

m 04 0 0 N N r l w rl 4 . rl

I I 1 1 I I I I I I m m N O rl 0 m w m w

0 I

1 'd 2

a 0 I rn k Fr

a, u (d

I a, rl

I+

i!

z I

VI h Fr

T E E

5 a a

a,

(d a, (d rl

4 * u

ii .A J

a I v)

8 a, X

a I

a rl rl

$4 (d a, c4

$4 a a, PI

v) .rl cl

oo w m rl

Q) W m rl

0 h m l-i

m I- m 4

4 h m rl

9 c,

m ?I

5 8 &

0 N

h l-i 1 c,

a a, VI 9

c, W N 0

?I

a oo VI PI

$W c u o m m rlrl

i: In E

m w m N woo

O m W N w r l

N W W w o o

or m

117

00 h m d

h rl 5 r, cv

0 1 rl

0 I

.I4 4 (d

X

h (d

0 d .I4 X

5 n

m

5;; m Erc Erc

m U m (d W

I- - m rl

al

b PI N

Ei

m cv cv N

0 d o I I rlrl rl

a I m

I a l m x k E r c 4

L X 4

rl

al PC

?d

e \o m rl

a 91 m \o l-i

N I- N d

00 h m d

al

b

rl rl

2

00 I 0

m I

E *

U

W 3

U p4

!I

3

2 i s"

0

rl *

h I- m rl

Fc aJ P 0 U u 0 m l-i

* rl 4. N

5

91 J U (d

I 1 1 1 1 m Erc ~ r c m z o p : u

m h

m f- r- e Fc 0 rl h d m

m hl

0 U rl 4 9) 9) m * *

P

118

h rl rl (d -rl CJ k

i 0 V

M C

.rl 01 e m cd a aJ U Ir 0 a aJ k

m a

rl VI H

d (d *rl

9) c U

I- rl

9) ?-I

c r. 0 o ! m m 0 m r - J . . a 3 w a ~ ~ o ~ o o o - t r - ~ o m o o m o m o c o o m m m 1 4 c l r l r - I 4 4 c l 4 1 1 r l r l 4 4 4 1

' * ' o o o o o c o o t - 0 ~ o o u o o ; o r ; o

A

a 4 0 o o o * m - C n rl* u O r n h \ o O - r t b c o m m b N h N m m r l m O U m r l r l N m -

N I l m N l l r l l N N m m - f N u 3

. . . . . . . . . . . . . . . .N

c o m m a J N . . . e . m r l m o w r l m u c a

c

A

M . I

0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0

1 1 r l r l 1 r - i r l 1 1 1 1 r l 1 1 1 1 r l 1 r l 1 r l r l r - i m l l

c

m m r-ia In d u m a r l m U I I N I 1 ' 1 I I 1 1 I 1 I 1 m r l N I I c o rl

a 3 m 0 . . 0 m m m o b . * - -

NO\ -3 o m rl I l r l r l l I I I I I I I I I I I r l l r l N I I

0 a c

119

*

c

m c o m r l m d 0 - a m 0 0 m 0 . 0 . . . 0

1 1 a 1 r l d h 0 0 ~ 1 1 1 1 1 1 1 1 r l 1 1 l l l l m l

b b N m a m rl rl 0 0 a m m I l r l l N r l m m l I 1 I 1 I I l r l l I I I m m m

N

o u m m . . . . 0 0 u * O N 0 m 0

I I I I I r l w m l l l l l l l l r l l l l I I I N

0 0 0 0 0 0 0 0 0 0 0 0000 0 r l 1 1 1 1 1 1 1 1 1 1 1 r l r l 1 1 1 1 r l r l r l r l l r l l l

rl m r l m o r l m N 0 1 1 1 1 1 1 1 1 1 1 1 r l I I I I N I I l r n

u m o o u h O

rl m m u m . . 0

I I I I I I I I I I I r l I I I l r l I m l I

I h a, a, a) a, a, a l a , a, a, a, a o m a, a, a, a l u m m h a J N a, a, m b * c c c c c c c c c c c * c c c c * c ‘ c e ’ N 0 0 0 0 0 0 0 0 0 0 o o m 0 0 0 O m 0 0 u N o m 0 0 m N N c c c c c c c c c c c u e c c c c 0 0 u m N c u c c ucw

120

i i a E-c

0 . ' 0 o m 40 l l l l l l l 4 h l l l l l l l l l l l l l l ~ r i l

m m

?: h m

I I I I I I I 1 1 1 1 1 1 1 1 1 1 1 1 1 1 4 1 1

0 m

N r i m m r i c o m \ o o v m o m l d l I I I I I 1 v m m m c n U N m v l - l 4 1 I

O c n h h c o o r i m c n U 0 'Do4

m c o c o m p . m N c n a h l m 4 N l l r i l l l l l l l 4 V I 1 1 m r i m

. . . . . . . .

N

k a a h a 5 h 0 IU

a a U U

D 7 rn rn U k 0 a aJ bl

2

2 *

P

c

1 2 1

Table 18. Maximum numbers of green t u r t l e s seen basking a t one time dur ing surveys i n t h e Northwestern Hawaiian I s l a n d s s i n c e 1950.

Location Number of

basking t u r t l e s Mon th-Year Reference

Nihoa 2 (23"06'N, 161"58'W)

September 1971 K r i d l e r (326)

Necker 7 November 1977 Balazs, un- (23"35'N, 164"42'W) publ ished d a t a

French F r i g a t e Shoals 22 (23O45'NY 166"lO'W) (nonbreeding months)

Laysan 6 (25"43'N, 171"44'W)

February 1975 Balazs , un- publ i shed d a t a

September 1966 K r i d l e r (316)

L i s i a n s k i 1 3 March 1964 Clapp & Wirtz (26"02'N, 174"OO'W) (171)

e

P e a r l and H e r m e s Reef (27,'55'N, 175'45'W)

Southeast I s l a n d North I s l and

50 10-20

L i t t l e North I s l and 10-20 Bird and Sand I s l ands 8

Midway (28"13'N, 177"21'W)

Kure (28"25'N, 178"lO'W)

September 1'966 Kr id l e r (316) 195 7- 58 Kenyon & Rice

(288) personal communication t o Parsons (400)

11 1957-58 January 1962 A. F. Carr,

personal communication t o Amerson e t a t . (32)

May 1975 J. Bradley, (only record) pe r sona l com-

mun i c a t i o n

January 1979 J. , S t a r k , personal com- mun ica t i o n

122 c

Tab1 .e 19. Tiger sharks , RzZeocerdo mvier, captured a t l o c a t i o n s i n the Hawaiian Archipelago where tu r t les were found t o be prey items. A. Tester 487 and unpublished d a t a ; Taylor and Naf t e l 482).

(Sources -

No. No. Percent t i g e r sharks t i g e r sharks t i g e r sharks

Location conta in ing con t a i n i n g conta in ing I s l and Coas t l i n e food tu r t l e s t u r t l e s

Main I8 lands * Hawaii southwest

nor thwes t 1 2

9 1 4

11.1 14 .3

Maui southwes t west

1 1

9 5

11.1 20.0

Oahu nor theas t sou theas t south

24 6

32

3 1 5

12.5 16.7 15.6

c

Kauai nor theas t southwest

4 18

3 2

75.0 11.1

Niihau ea8 t west

13 1 5

2 1

15.4 6.7

Northweetern Hawiian Is Zanda

French F r i g a t e Shoals 16 5 31.2

P e a r l and Hermes Reef 11 4 36.3

* 25 t i g e r sharks t h a t contained food bu t n o t t u r t l e s were captured by Tester (487) a t o t h e r l o c a t i o n s i n the main i s l a n d s

I’

c

123

a

I

3 $ k 3

v

e 0 m rl I

N 0 m rl

ti

3

o\ rl

U .)

4 c, d k

3 rl 1 rl 0.

31 a

g

5 E d a d 0 m

z rl U k 7 U

w 0 k a, P

E - 9

rlrl c tz g 8

0 N

a, rl P (d El

U

3 :: a PI e

0 m rl

k a,

3 z

U

CJ k a, PI

G m .o rl

k a

Ei z

(\1 0 m rl

U d a, CJ k a pc

k a

fi 2;

5 g c

m m .

m

9

m

9

rl

m

9

I

I

h k (d 1 E (d c,

I

I

e m

o\

9

rr)

h

N

cr)

9

b td 1 k P a,

Frc

m m

m

0 9

0 rl

a3

o\

cn rl

N

N .

e

5 5

U m

U rl

h

0 rl

do rl

U cv l-l

U N

rn 9

N 4

rl d k 2

m cn

U cv

m m rl

9 N

h

9

m I4

0 \o

rl rl

9

m U .

rl rl

m b rl

m cv

h

b

m rl

m N rl

cr) N

B ?

rl m

a3

m r( rl

cn 4

N

h

e rl

9

do m

2

h rl 1 c,

m rl

U

rn 9

rl rl

m h l-l

e m

h

00

9 rl

U

bo 3 3

m

;;i

N a3

Q: m

0 li

o\

rl rl

rr) N

9

r-

e rl

k a P fi a, U a a, v)

m o\

e N

b

h

m rl

m

=1

N N

rl

rl

N

k a 0 U CJ 0

n

9

m rl

rn m

N

e

r-

b

In

rl rl

m 0 rl

.

m rl

bl a, n 8 3 0 2

h

e rl

m m

Q: m

0 rl

rl

m

9

m m

9

k a,

5 i

0 0 rl

co m N

8 rl

do 9 rl

0 0 rl

U m rl

0 0 d

U Q) rl

d (d U

EEi

124 A'

a J 1 Pcd

cd

aJ

W E a&

rl N

aJ

n w rl W

U

n &O rl

0 N

rl N 00

rl

W

h

co U cn rl

n w U W

b N

n w N m W

cn 0 m

W

4-

cn m rl

n M m m

rl a0 rl

W

n M m N co

N \D U U

W

- 0 m m 4

n W rD

m N

cn cn m

W

rl m m rl

n $s m m m

m co W

W

N m m rl

n M N

N U

U rl co rl

W

*

m m cn 4

n $s N

hl m

m U

W

U m m rl

n M hl

m .

W

rD cn N m - m m cn rl

n M rl

a) m

0 N rl

. W

b m m rl

n M

U rl

00 0 N

? W

a-

*

0 c n N m m N rl m

n

N

rl rl W

x

n ae 0 m

b cn

W

125

0 \o 0

cn *

n M

N \o

cn cn \o

m

0'. W

m

n M In rl U

N \o m rl

h

n M cn rl

In h 4

W

m e a\ rl

h m rl

h A

n w 4

u3 m

rl 0 \o

N

W

h

4 co m In

8,

n

co m rl

rl e

b\O

W

h

O? Pl m N

n

n M 4

4 h

0 e h

rl

W

m

m 0 m 4

A

h M 00 h In

cn co e N

W

*

N a3 m l-l

m

n d o W

h \o W

h m m

W m 0

m A

n

co b\'

;fm W

0 \D h

rl h

\o rl m

n M cn rl \o

\o cn rl

W

co In e rl .,

n M m \o In

rl N co

. W

4 N m

n M 0 h W

h N N

OI cn u3

rl A

n M In

\o cn

rl e \o

rl

W

n

N N m

c

n

N b y

In

\o h

W

0 rD cn rl

rl \D cn rl

cn e cn rl

0 In cn

rl m OI rl

P4 In cn rl

m m cn rl

U In m rl

m m cn rl

\o m cn rl

h m cn 4

cn In m rl rl

126

n w w W

N ln

n

W

rl U U

n OS? U

n

m bw n

h e h

m

m m m

W

n O Y m N

h h 0

N

W

L

n

m b\o

O H * m

0 m rl

W

n w d

v

m * W

U rl W W

h N

N m m

cu w \o

@ d

n w m m b W

m .j ln

n w * r( rl W

n n O N w F 7 - n

n w U m

m m 0 m

W

L

n O N w m

* m 0 m

W

m

n

cu 0 rl

N

b y

W

s m

m

n

In m U

m VI m U U

b y

v

L

1 com U h m m rlrl

U W r - i w w w

a3 a cv

m o m * N d N - 4

c n w m rl d W

n w W

W d W

m m n

rl

n OW U h N

rl co rl

m m

W

h w rl

m U

m 4 N

m

W

m

N h m rl

n A0 oc, m

d U m

W

n

m 0 m

In 0 w U

bS?

W

L

n

h

0 m

w h h

cu

b-

W

L

rl fi m rl

n

U N

m 00 0 N

b-

W

I,

n w 0 d

\D w W

W

n O N J

m W

rD r-i m

m m

m h m rl

0 In v)

n O N m U

b cu N

W

n w co \o W

m co 0 m

W

I,

m \o m d

n w U h W

m N U

c

n 0- m oc, In

N 0 rl

W

n O N U co U

* 0 r i

d

W

m

h \o m rl

n 0- m W W

m m m rl

W

m

03 \o m rl

n O N m 0 co v

n w m U W

N m

e rD m rl

n A m m

w m

W

m W m rl

m m m

N m W W m m r l r l

x m rl

12 7

b rl N

n O N 0 0 rl

h 4 N

v

N UJ m 4

rl h m

n M rl

4. 4

N N rn

W

n OW N

UJ

Vl 4.

W

U \d - m

rl vr m N

A

n M co

s 4 rl rl

n M 4.

W

m

\o \o m rl

c n N h N

n M Ln

b

4

W

=1

n w 4

b N hl

W

n M 4

N W

b m

I-- PI 0

m A

n M UJ

rl

v) 4 rl

W

n M co d

rl \o rl

W

rl h m rl

m N \o

rl rl

A

n iM UJ

m

4.

. W

A r-4

N h m rl

In m cn 0 \o co co 0 m

A n

n b?

n o\ iM N v)

N W W

A

m N

n O M vr rl

N UJ m rl

W

A

n O N 4

W

UJ h

n M m W

m N N

n N 4 W

cn 00 m

n b\O h A &-

m

I m corn b 4.- m m m d rld

120 Y

Table 22. Kilograms and percent of t u r t l e from t h e Hawaiian I s l ands repor ted as being commercially captured by month, 1948-19730

c ---- -- --I_-_-------

Month Kilograms Percent of t o t a l

January 8,950 9.9

February 7,729 8.5

&

March 10,524 11.6

A p r i l 8,977 9.9

May 6,900 7.6

June 8,477 9.3

J u l y 7,680 a. 5

August 7,682 8 .5

Sep t emb e r 7,298 8.0

October 6,073 6.7

November 4 ,312 4.8

Decembe 1: 6,201 6.8

To t a l 90,803 100.0 c

129

_-

e

c

c

. a a E

0

w z

1 I 0 0 In 0 cv cv

130

c

4

P

131

3 0

I-

*

c

7 N N

0

3 a z)

0 z

x

3 0 I 7

0 I

a 0

E

! $ 2 Urn

al d k d

'd

$ 9 k a

u w 8 t n aal drl rnu a l k

U

8 8 ual k k ob0

3 M

132

KURE ' * MIDWAY 'PEARL AND HERMES

LlSlANSKl *LAYSAN

/-

e

.

JULY

9 NECKER

r O

QB.

.P

0 AUGUST

SEPTEMBER

OCTOBER

I 1 I

180" 170" W I 6 0 " W I7O0 E

Figure 4. hatchling and posthatchling Chelonia leaving French Frigate Shoals. of paths represent estimated distances covered during subsequent months. hatchlings emerge during August. Southwest Fisheries Center Honolulu Laboratory, National Marine Fisheries Service, NOM, Honolulu, Hawaii 96812.)

Surface drift trajectories representing the theoretical movement of

Most Segments

(From Balazs 57, courtesy of R. H. Barkley,

30"N

2 5"

20"

30'"

2 5"

2 0"

30"N

2 5 O

2 0"

5O"N

!5"

O0

c

R

1.33

n e

d

W

f !i c

30

29

26

27

maximum

m o a n

mlnlmum

2 2 1 1 1 1 1 1 1 1 ~ 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 (

J F M A M J J A S O N D J F M A M J J A S O N D J F M A M J J A S O N D

I974 I975 1976

Figure 5 . for each month repreeent an average of seven recordings. Balaze and U.S. Coaet Guard personnel.)

Sea aurface temperatures a t French Frigate Shoals. Data presented (Source: G . H .

a

134

6

EAST ISLAND

FRENCH FRIGATE SHOALS

N

Figure 6. of 1974. nesting areas that have been established for reference purposes.

Locations of 220 nests recorded on East Island from June to August The numbers on the island’s perimeter identify the 17, 50-m long

P

L

135

70’

60

50

40

30

20

IO

0 e

-

-

-

1973: Y- 25.442 + .1522X - I974 : Y-43*208 + .3039X

1975 : Y- 43. 055 + ,1451 X - 1976 : Y m 16.976 + ,9357 X

I I I I I I I

0 IO 20 30 40 50 60 70 81

NUMBER OF DAYS IN NESTING SEASON

Figure 7 . Island, French Frigate Shoals (day 1 = 1 June).

Percent of green tur t l e s successful ly nesting each night at East

136

7

c\l c\l

0

a 7

tu

rl Q O tutu

E m 3aJ u u a J 3 k 0

ucd

a u- l c d o v

M

c

A

137

INSHORE HABITAT

c

INSHORE HABITAT

FEEDING - RESTING

(JUVENILES 1

BEACH HABITAT

BASKING

JUVENILES

PELAGIC HABITAT

FEEDING - RESTING

-b

FEEDING - RESTING

(JUVENILES - SUBADULTS - ADULTS)

BASK1 NG ADULTS

I INSHORE HABITAT

COPULATION - FEEDING - RESTING

f I ADULTS

I 4

- HATCHLINGS BEACH HABITAT -l NESTING -BASKING

Figure 9 , L i f e h is tory and habitat model for Hawaiian CheZonia (adapted from Carr e t aZ. 153).

138

0 0 N

I I

c

L

c

n

P

139

20

v) W J I- 3 I-

(3 z Y v) U

L 0

a w m I 3 z

a

- m

15

IO

5

1973 : Y s 10.009 - .1523X

1974 : Y * 19.189 - .3199X

1975: Y 8 21.215 - .2483X

0 , I I I 1 I

'. 0 IO 20 30 40 50 60 70 00

NUMBER OF DAYS IN NESTING SEASON

Figure 11. Incidence of dai ly basking a t East Island, French Frigate Shoals (day 1 = 1 June).

x

140

1 I I I 1 4

FRENCH F RlGATE SHOALS

lL 0

250

roo

-

-

I973 L

I974 1975 L

I976 I977 I978 I979

NESTING SEASON

Figure 12. Number of females nesting annually at French Frigate Shoals.

141

a.

50

40

30

20

IO

I I I I 1

0 HAWAI I MAU I LANAI MOLOKAI OAHU KAUAI

Figure 13. Percent tur t l e reported as being commercially captured for each of the main is lands, 1948-1973.

Documents

SYNOPSIS OF BIOLOGICAL DATA