NO. 397

(1)

NO. ₃₉₇

(2)

Tab1 e o f C o n t e n t s

Page ...

L i s t o f F i g u r e s

...

¹¹¹

_{. .}

L i s t of P l a t e s

...

vii

L i s t o f T a b l e s

... ^x

A b s t r a c t

...

¹

A

.

I n t r o d u c t i o n

...

³

...

B

.

H i s t o r y o f C a r o l i n e A t o l l 6

...

Pre-European H i s t o r y : Tuamotuan P e r i o d 6 Post-European H i s t o r y : 1 7 t h t o 1 9 t h C e n t u r i e s

...

6

...

The Guano Era 9

...

S o l a r E c l i p s e E x p e d i t i o n 10

...

The L a t e 1 9 t h and 20th C e n t u r i e s 10

...

2 0 t h Century S c i e n t i f i c S t u d i e s 12 C

.

^Methods

. . .

¹²

. . .

F i e l d Techniques 1 2 Naming C a r o l i n e ' s Motus ^. ^. ^. ^. ^. ^. ^. ^. ^. ^. ^. ^. ^. ^. ^. ^. ^. ^. ^. ^~ ^~ ^o ^o ^~ ^~ ^~ ^~ ^~ ^~ ^. ^~ ^. ^. ^. ^. ^.

. . . .

17

. . .

D o P r e l i m i n a r y S t r u c t u r e and Topography 17 Background Geological S e t t i n g

. . .

^{1 8} Sea Level Changes

. . .

19

General Account

. . .

19

. . .

Reef F l a t s

...

²⁰

Beaches

. . .

²³

Lagoon

. . .

²⁴

Lagoon Hydrology

. . .

²⁴

Patch Reefs

...

²⁵

. . .

T r i d a c n a - A c r o p o r a Reefs 25 Lagoon Reef F l a t s

...

²⁷

Lagoon Reef Fauna: A B r i e f Summary

. . .

27

S u b s t r a t a

. . .

28

. . .

Hydrology 30

. . .

C l i m a t e 30

. . .

Wind and R a i n f a l l 31 C y c l o n i c Storms

. . .

31

. . .

Sea C o n d i t i o n s 33 T i d e s

. . .

³³

E

.

F l o r a : V a s c u l a r P l a n t s and F l o r i s t i c s

. . .

³⁴

B o t a n i c a l H i s t o r y . . . e a s * . . .

. . . .

34

. . .

V a s c u l a r P l a n t s of C a r o l i n e A t o l l 34 P l a n t C o l l e c t i o n s

. . .

34

S p e c i e s L i s t s , Annotated C h e c k l i s t and Maps o f T e r r e s t r i a l V a s c u l a r P l a n t s

. . .

3 5 F l o r i s t i c s and Ecology o f t h e Motus

. . .

64

S i z e o f t h e F l o r a

. . .

⁶⁴

. . .

Numbers o f I n d i g e n o u s P l a n t s 65

. . .

Composition o f t h e F l o r a 68 F

.

E c o l o g i c a l S u c c e s s i o n

. . .

69

. . .

B a s i c Sera1 S t a g e s 69

. . .

E c o l o g i c a l S u c c e s s i o n on Motus of D i f f e r e n t S i z e C l a s s e s 73 S p e c i e s - A r e a R e l a t i o n s h i p s

. . .

Comparisons o f S p e c i e s - A r e a R e l a t i o n s h i p s w i t h O t h e r A t o l l s

. . .

87

(3)

The Q u e s t i o n o f Fresh Water

...

8 7 Motu S i z e i n R e l a t i o n t o t h e D i s t r i b u t i o n o f T r e e s .

...

Shrubs and Herbs 89

...

G

.

P l a n t Communities 90

...

General Account 90

...

Natural Herb Mat 90

...

S e a b i r d Use 93

Beach Scrub w i t h S u r i a n a

. . .

⁹³

Pandanus F o r e s t

...

93

T o u r n e f o r t i a Scrub and F o r e s t

. . .

94

General D i s t r i b u t i o n

. . .

94

. . .

S p e c i e s D i v e r s i t y i n T o u r n e f o r t i a Woodlands 96

...

S t a t u r e and Area Coverage 97 Ecology

...

98

S e a b i r d Use

. . .

99

Cordia F o r e s t

. . .

99

General D i s t r i b u t i o n

...

99

H i s t o r y

...*

^{* . 100}

. . .

Abundance and D i s t r i b u t i o n * - l o 0

. . .

S e a b i r d Use e s n a * l O O

. . .

P i s o n i a F o r e s t 101

. . .

General D i s t r i b u t i o n 101 An H i s t o r i c a l P e r s p e c t i v e

. . .

101

. . .

Annual Growth Rates 108 S p e c i e s D i v e r s i t y i n P i s o n i a F o r e s t s

. . .

109

. . .

Ecology 110 R e l a t i o n s h i p s Between P i s o n i a F o r e s t Height and Motu

. . .

Dimensions 110

. . .

P i s o n i a - S e a b i r d R e l a t i o n s h i p s 112 Remnant P i s o n i a F o r e s t s i n t h e P a c i f i c

. . .

113

. . .

Coconut Woodlands 113

. . .

General D i s t r i b u t i o n 113 D i s t r i b u t i o n and Abundance

...

-117

. . .

S e a b i r d Use 118

. . .

Absent P l a n t Communities H

.

D e s c r i p t i o n and Ecology of t h e Motus

. . . .

I Conclusion . . . o . . a e e . . . . o . . . o . . . 171

J

.

L i t e r a t u r e C i t e d . . . q . . 9 . . o . 9 . a . . . e . . 172

Appendix I : Reef I n f o r m a t i o n f o r N a v i g a t o r s

...

A 8 3 Appendix 11: Weather D a t a , 1989-1990

. . .

¹⁸⁴

(4)

L i s t of Fiqures

F r o n t i s p i e c e Air-mosaic of Caroline A t o l l , RNZAF 6569.

Reproduced by permission of t h e Department of Lands and Survey Information, Wellington, New Zeal and.

Page 1 Line I s l a n d s : geographic l o c a t i o n in t h e P a c i f i c Ocean

...

4 2 Carol i ne At01 1

,

Republ i c of K i r i b a t i

,

with newly-named

i s l e t s

...

⁵

3 Main marae on Nake I s l a n d , Caroline A t o l l , based on a plan pub1 ished by t h e S o l a r Ecl i p s e Expedition (Holden &

Qualtrough 1884)

. . .

7 4 Caroline A t o l l , as surveyed by John Arundel, 1883, and

s t i l l used as t h e standard hydrological c h a r t

. . .

7 5 C a r o l i n e A t o l l , as c h a r t e d by t h e S o l a r E c l i p s e P a r t y ,

a l s o in 1883 (Holden & Qualtrough 1884) ,...eee.+. 14 6 C a r o l i n e A t o l l , by Bryan (1942), a modified version of

t h e Solar E c l i p s e P a r t y ' s map (1883)

. . .

¹⁴

7 A map by Clapp & S i b l e y (1971a), based on Bryan's 1942

map ( F i g . 6)

. . .

¹⁵

8 C a r o l i n e A t o l l : survey t r a n s e c t s . The d i s t a n c e covered

was 13.3 km

. . .

¹⁵

9 Caroline A t o l l : perimeter s u r v e y s , The d i s t a n c e covered

was 19.3 km

. . .

¹⁶

10 Re1 a t i o n s h i p between mean annual r a i n f a l l and

d i s t r i b u t i o n of phosphate i s l a n d s in t h e t r o p i c a l P a c i f i c

(reproduced from Stoddart & S c o f f i n 1983)

...

²⁶

11 E n t i r e d i s t r i b u t i o n map of r a r e and/or l o c a l i z e d p l a n t s on Caroline A t o l l : H i b i s c u s t i l i a c e u s , Lepidium

b i d e n t a t u m , Pandanus t e c t o r i u s , P h y l l a n t h u s amarus, P s i l o t u m nudum, Scaevola taccada, S i d a f a l l a x , Tacca

l e o n t o p e t a 7 0 i d e s , Thespesia populnea, T r i b u l u s c i s t o i d e s ,

Ximenia americana, and Species A . . . 41 12 Transect d i s t r i b u t i o n map of t h e f e r n Phymatosorus

s c o l o p e n d r i a on C a r o l i n e Atoll

. . .

41 13 Transect d i s t r i b u t i o n map of t h e g r a s s Lepturus repens on

C a r o l i n e Atoll

...

(5)

14 E n t i r e d i s t r i b u t i o n map o f the coconut Cocos n u c i f e r a on

C a r o l i n e A t o l l

...

45

15 T r a n s e c t d i s t r i b u t i o n map o f Laportea r u d e r a 7 i s on C a r o l i n e A t o l l

...

47

16 T r a n s e c t d i s t r i b u t i o n map o f Achyranthes c a n e s c e n s . . . 47

17 T r a n s e c t d i s t r i b u t i o n map of Boerhavia repens, C a r o l i n e

...

A t o l l 49 18 E n t i r e d i s t r i b u t i o n map o f P i s o n i a g r a n d i s , C a r o l i n e A t o l l

. . .

50

19 T r a n s e c t d i s t r i b u t i o n map of Portu7aca 7utea, C a r o l i n e

. . .

A t o l l 53 20 T r a n s e c t and p e r i m e t e r s u r v e y d i s t r i b u t i o n map o f S u r i a n a m a r i t i m a ^. ^. ^. ^. ^. ^. ^. ^. ^. ^. ^, ^. ^. ^. ^. ^. ^. ^. ^~ ^, ê ê ^. ^~ ^. ^, ^~ ê ^. ^~ ⁹ ^. ^.53 ^. ^. ^.

21 T r a n s e c t d i s t r i b u t i o n map of lpomoea macrantha. E n t i r e d i s t r i b u t i o n i s shown f o r South I s l a n d

. . .

⁵⁵

2 2 E n t i r e d i s t r i b u t i o n map o f Cordia subcordata

. . .

⁵⁷

23 E n t i r e d i s t r i b u t i o n map of He7iotropium anoma7um

...

59

2 4 E n t i r e d i s t r i b u t i o n map of T o u r n e f o r t i a argentea

...

60

25 T r a n s e c t d i s t r i b u t i o n map o f Morinda c i t r i f o l i a . The o u t l i n e d a r e a on T r i d a c n a I s l a n d ( n o r t h e a s t of South I s 1 and) e n c l o s e s T o u r n e f o r t ia-Morinda f o r e s t .

. . .

^.^,^,^,

.

⁶²

26 Evidence f o r t h e i n d i g e n o u s s t a t u s of Morinda c i t r i f o 7 i a on C a r o l i n e A t o l l : p e r c e n t a g e c o v e r on t r a n s e c t s w i t h i n

...

n a t u r a l and a n t h r o p o g e n i c f o r e s t s

...,...-**.

⁶²

27- P l a n t communities and amount of i n d i g e n o u s v e g e t a t i o n on 30 t h e d i f f e r e n t s i z e c l a s s e s of motus, a s i l l u s t r a t e d by " p i e " diagrams

. . .

74

31 T o t a l numbers o f p l a n t subcommunities ( u p p e r g r a p h ) and s p e c i e s ( l o w e r g r a p h ) i n r e l a t i o n t o motu a r e a , d e m o n s t r a t i n g p l a n t s u c c e s s i o n on t h e d i f f e r e n t s i z e d motus e n c i r c l i n g C a r o l i n e ' s lagoon. Fine d o t s i n d i c a t e t r e n d l i n e s

. . .

84

32 Schemati c p r o f i 1 e through Arundel Is1 e t , r e c o v e r i n g from d i s t u r b a n c e o v e r 60 y e a r s ago, showing n a t u r a l h e r b m a t s , T o u r n e f o r t i a s c r u b and f o r e s t , and 5 s p e c i e s o f b r e e d i n g

...

95

(6)

Maximum h e i g h t s o f P i s o n i a f o r e s t s i n r e l a t i o n t o w i d t h

o f t h e motus

...

¹⁰²

Maximum canopy h e i g h t s o f P i s o n i a f o r e s t s i n r e l a t i o n t o

motu a r e a

...

¹⁰²

Schematic p r o f i l e t h r o u g h Long I s l a n d , T r a n s e c t 0.

A l t h o u g h Long I s l a n d has been formed i n t h e r e c e n t p a s t b y a merger o f 5 s m a l l e r i s l e t s , t h i s s e c t i o n o f t h e i s l e t i s v e r y mature, c o n t a i n i n g n a t u r a l h e r b mats,

T o u r n e f o r t i a s c r u b and f o r e s t , and t a l l P i s o n i a f o r e s t .

...

^.lo3

Schematic p r o f i 1 e t h r o u g h S o u t h I s 1 and, where >90% o f t h e l a n d s u r f a c e i s c o v e r e d w i t h Cocos f o r e s t s , p r i m a r i l y i n

a d y i n g s t a t e

...

¹¹⁴

Nake I s l a n d : v e g e t a t i o n and p h y s i o g r a p h y . . . ~ ~ ~ . ~ ~ . 0 . . e ~ ~ - 0 1 2 1 bong I s l a n d : v e g e t a t i o n and p h y s i o g r a p h y ...s-a...----...A24

Long I s l a n d : n o r t h - s o u t h t r a n s e c t showing d i v i s i o n i n t o f o r m e r i s l e t s , f l o r i s t i c c o m p o s i t i o n , r e l a t i v e abundance o f p l a n t s p e c i e s , d e g r e e o f s p e c i e s o v e r l a p , and canopy

h e i g h t s

. . .

125 Long I s l a n d : e a s t - w e s t c r o s s - s e c t i o n t h r o u g h T r a n s e c t C,

a f o r m e r i n t e r i s l e t c h a n n e l , showing f l o r i s t i c

c o m p o s i t i o n , r e l a t i v e abundance o f p l a n t s p e c i e s , d e g r e e

o f s p e c i e s o v e r 1 ap, and canopy h e i g h t s . .

. . . .. . . .

^.I26

bong I s l a n d : e a s t - w e s t c r o s s - s e c t i o n t h r o u g h T r a n s e c t 8, w h i c h passes t h r o u g h m a t u r e i n t e r i o r P i s o n i a f o r e s t o f l a r g e s t o f Long's c o a l e s c e d motus. D a t a i n c l ude f l o r i s t i c c o m p o s i t i o n , r e l a t i v e abundance o f p l a n t s p e c i e s , d e g r e e

o f s p e c i e s o v e r l a p , and canopy h e i g h t s

...

....i26 V e g e t a t i o n and p h y s i o g r a p h y o f Windward I s l e t no. 1:

Eo'sun B i r d I s l e t

. . .

1 3 1 V e g e t a t i o n and p h y s i o g r a p h y o f Windward I s l e t s nos. 2, 3

and 4 : Windward and C r e s c e n t I s l e t s , and Motu A t i b u

( " C o r a l Rubble I s l e t " )

. . .

¹³³

V e g e t a t i o n and p h y s i o g r a p h y o f Windward I s l e t s n o s . 5 t h r o u g h 9: N o r t h P i g , P i g , S k u l l , N o r t h B r o t h e r s , and

B r o t h e r s I s l e t s

. . .

135 P i g I s l e t : e a s t - w e s t c r o s s - s e c t i o n t h r o u g h c e n t e r o f

i s l e t . D a t a i n c l u d e s f l o r i s t i c c o m p o s i t i o n , r e l a t i v e abundance o f p l a n t s p e c i e s , d e g r e e o f s p e c i e s o v e r l a p and

. . .

canopy h e i g h t s

(7)

Brothers of i s l e t . abundance canopy he

I s l e t : east-west c r o s s - s e c t i o n through c e n t e r Data includes f l o r i s t i c composition, r e l a t i v e of p l a n t species, degree of species overlap and

i g h t s

...

140 Vegetation and physiography of Windward I s l e t s nos. 10

through 1 2 : Noddy Rock, N o r t h Arundel, and Arundel

I s l e t s

...

¹⁴¹

Vegetation and physiography of Windward I s l e t no. 13:

Tridacna I s l e t

. . .

144 Tridacna I s l e t : east-west c r o s s - s e c t i o n through lower

c e n t e r of motu. Data includes f l o r i s t i c composition, r e l a t i v e abundance of plant s p e c i e s , degree of s p e c i e s

overlap and canopy heights

. . .

145 South I s l a n d : vegetation and physiography

. . .

146 South Island: d i s t r i b u t i o n and abundance of p l a n t

s p e c i e s along Transect 2 , which runs a t an angle of 60"

from the lagoon t o the south shore through t h e western c e n t e r of t h e i s l e t . Data includes f l o r i s t i c

composition, r e l a t i v e abundance of p l a n t s p e c i e s , degree

of species overlap and canopy heights

. . .

149 Vegetation and physiography of t h e 7 South Nake I s l e t s :

Pandanus, Danger, Booby, Coral, and Lone Palm I s l e t s , Motu Kota ("Red-footed Booby I s l e t " ) , and Motu Mouakena

("Masked Booby I s l e t " )

. . .

152 Vegetation and physiography of Central Leeward Is1 e t

no. 1 : Motu Mannikiba ("Seabird I s l e t " ) . . . e . e s a 0 s a . e + 1 5 7 Vegetation and physiography of t h e Central Leeward I s l e t s

nos. 2 t h r o u g h 4 : Blackfin I s l e t , Motu Matawa ( " F a i r y

Tern I s l e t " ) , and Emerald I s l e

. . .

160 Vegetation and physiography o f t h e Central Leeward I s l e t s

nos. 5 through 11: Shark and S c a r l e t C r a b I s l e t s , Motu Nautonga ("Sea Cucumber I s l e t " ) , Azure I s l e , R e e f - f l a t ,

B i r d and Fishball I s l e t s

. . .

162 Fi shball Is1 e t ( n o . 11, Central Leewards) : east-west

c r o s s - s e c t i o n through t h e c e n t e r of t h i s young motu, which e x h i b i t s e a r l y s t a g e s of geological and biological e v o l u t i o n . Data includes f l o r i s t i c composition, r e l a t i v e abundance of p l a n t s p e c i e s , degree of species overlap a n d

. . .

canopy heights

(8)

vii 57 Vegetation and physiography of t h e 5 Southern Leeward

I s l e t s : Motus Raurau ("Blue-gray Noddy I s l e t n ) . E i t e i ( " F r i g a t e b i r d Is1 e t " ) . Pisonia I s l e t . Kimoa ("Rat

I s l e t M ) . and Ana-Ana ("Anne's I s l e t " )

...

¹⁶⁸

L i s t of P l a t e s Page 1 A dawn view of Caroline. 1988

...

¹⁸⁶

2 The main c l e a r i n g on South Is1 and. 1883

. . .

¹⁸⁶

. . .

3 One of 3 European-style houses b u i l t on Caroline. 1883 187 4 An a r t i s t ' s rendering of Caroline. 1883

. . .

¹⁸⁷

5 Map of t h e . s e t t l e m e n t .

.

South I s l a n d . 1883

. . .

188

6a. b Two lagoon views a century ago

. . .

a e e " s 0 + 1 8 8 7a Motu Ana.Ana. in 1883 e s s e n t i a l l y i d e n t i c a l t o today

. . .

¹⁸⁹

7b A T o u r n e f o r t i a t r e e along South I s l a n d ' s lagoon edge

. . .

189

8 The j u n i o r author in I p o m o e a m a c r a n t h a t h i c k e t . South Island

. . .

¹⁸⁹

9 "Rat City" base camp. Long Island

. . .

¹⁹⁰

10 Black-tipped r e e f shark. C a r o l i n e ' s lagoon

. . .

¹⁹⁰

1 1 L a t e r a l l y continuous upraised o l d e r r e e f s and wide r e e f f l a t s . southwest Nake Island . . . a . e . . . e e e e a a s a 1 9 1 12 Floating a srnall boat across t h e r e e f f l a t s from " e n t r a n c e " t o " l a n d i n g . " ^. ^. ^~ ^. ^. ^~ ^. ^~ ^s â ^~ ^~ ^. ^~ ê ô ê â â ^. â s a e a s . e . e * . . 1 9 1 â ^~ ^~ ^. â ^. 13 Beach c r e s t . r u b b l e . seaward "lagoon" and r e e f f l a t s o f f s o u t h e a s t Nake Island

. . .

¹⁹²

14 Extensive lagoon r e e f f l a t s south of Arundel I s l e t

. . .

192

1% A n i n c i p i e n t motu south of Motu Mouakena

. . .

193

16 Ridges of coral rubble. Nake Island

. . .

193

17 Channel between t h e 2 northern i s l e t s . Long and Nake

. . .

194

18 Noddy Rock. windward r e e f f l a t s

. . .

¹⁹⁴

19 Windward beach with flotsam and jetsam. Long I s l a n d

. . .

195

(9)

(10)

4 1 A c h y r a n t h e s c a n e s c e n s i n a c l e a r i n g a d j a c e n t t o P i s o n i a

f o r e s t . P i g I s l e t

...

²⁰⁷

42 B o e r h a v i a f r u i t s on f e a t h e r s and b i l l o f a G r e a t

F r i g a t e b i r d

...

^{2 0 8}

43 I n s i d e a m a t u r e P i s o n i a g r a n d i s f o r e s t . Nake I s l a n d

. . .

^{2 0 8}

44 F r i n g e o f S u r i a n a . n o r t h e a s t p o i n t . S o u t h I s l a n d

...

²⁰⁹

45 W e l l - d e v e l o p e d n a t u r a l h e r b mat

. . .

²¹⁰

46 H e l i o t r o p i u m anomalum w i t h t r o p i c b i r d s k u l l ( f i r s t

e v i d e n c e o f t h i s s p e c i e s ) . S k u l l I s l e t

. . .

²¹⁰

47 T o u r n e f o r t i a s c r u b . f r i n g e d by a n a t u r a l h e r b mat. and

o c c u p i e d by a c o l o n y o f S o o t y T e r n s

. . .

²¹¹

48 S e c o n d a r y T o u r n e f o r t i a - M o r i n d a f o r e s t w i t h n e s t i n g Brown

N o d d i e s . T r i d a c n a I s l e t

. . .

212 49 S k u l l I s l e t l o o k i n g e a s t t o t h e windward r e e f

...

²¹³

50 M a t u r e P i s o n i a g r a n d i s c a n o p y w i t h i n c u b a t i n g B l a c k

N o d d i e s and a White T e r n . P i g I s l e t

. . .

²¹³

51 A z u r e I s l e . an e x a m p l e o f a motu c o n t a i n i n g a s i n g l e

P i s o n i a t r e e

. . .

214 52 C a r o l i n e ' s s i n g l e c l e a r i n g w i t h T a h i t i a n - s t y l e h u t s . Motu

Ana-Ana ^. ^. ^. ^. ^. ^. ^. ^. ^. ^. ^. ^. ê ^~ ê ^. ê ê ^~ ^. ⁶ ê ^~ ^. ^. ^. ^. ^. ^. ^. ^. ^. ^. ^. ^. ^. ^. ^. ^. ^. ^. ^. ^. ^. ^. ^. ^. ^. ^. ^. ^. 214 53 P i l e s o f f i b r o u s s h a v i n g s . . c o c o n u t c r a b s i g n

. . .

215 54 B e a c h r o c k . Long I s l a n d

. . .

215 55 Windward I s l e t s nos

.

5 - 9 : N o r t h P i g . P i g . S k u l l . North

B r o t h e r s and B r o t h e r s

. . .

216 56 Arundel I s l e t ( f o r e g r o u n d ) . l o o k i n g s o u t h - s o u t h w e s t

a c r o s s T r i d a c n a I s l e t t o S o u t h I s l a n d

. . .

216 57 D e t a i l . T r i d a c n a maxima r e e f s . l a g o o n s i d e o f T r i d a c n a

I s l e t

. . .

217 5 8 Pandanus I s l e t ( c e n t e r )

. . .

²¹⁷

59 Danger I s l e t

. . .

2 1 8 60 S o u t h Nake I s l e t s n o s

.

3 - 6 : Booby. C o r a l . Lone Palm. and

Kota

. ...

²¹

6 1 Lone Palm I s l e t

. . .

219

(11)

(12)

Widths of pioneer herb mats on seaward- and lagoon-facing

shores, Caroline Atoll

. . .

^{9 2}

Stature and extent of Tournefortia in the major habitats

of Caroline Atoll

...

9 8 Distribution of well-developed (r10 m height) Pisonia

forests on the motus of Caroline Atoll

. . .

104 Area and dimens

and 5 islets of Number of trees islets during th showing remnant

ons of Pisonia grandis on Vostok, Flint,

Caroline Atoll

. . .

105 and areas planted in Cocos on Caroline's

e major planting era (l9l6-l92O), a Cocos data for 1988

. . .

Species diversity in Pisonia forests of decreasing maturity, Caroline Atoll

. . .

(13)

PART L EIISTORY, PHYSIOGRAP ISLET DESCRIPTIONS

ANGELA K. KEPLER' AND CAMERON B. KEPLER~

ABSTRACT

C a r o l i n e A t o l l ( F r o n t i s p i e c e ) i s s i t u a t e d a t 1O000'S l a t i t u d e and 150°13'W l o n g i t u d e i n t h e s o u t h - c e n t r a l P a c i f i c Ocean. C a r o l i n e i s t h e s o u t h e a s t e r n m o s t o f t h e S o u t h e r n Line I s l a n d s , a g r o u p o f 3 i s l a n d s which a l s o i n c l u d e s Vostok and F l i n t , l y i n g 230 km t o i t s west and

s o u t h w e s t , r e s p e c t i v e l y . Although a r c h a e o l o g i c a l l y and g e o g r a p h i c a l l y w i t h i n P o l y n e s i a , C a r o l i n e i s owned by t h e R e p u b l i c o f K i r i b a t i

( f o r m e r l y Gi 1 b e r t I s 1 a n d s )

.

C a r o l i n e , 9 . 7 km l o n g , 2 . 3 km wide a t i t s w i d e s t p o i n t , and 2 6 . 9 km i n c i r c u m f e r e n c e , i s a c r e s c e n t i c c o r a l r i n g w i t h 39 i s l e t s (motus) c e n t e r e d on a c o n t i n u o u s r e e f e n c l o s i n g a r e l a t i v e l y s h a l l o w l a g o o n . I t s t o t a l l a n d a r e a above h i g h w a t e r i s 399 h a . , w i t h motus r a n g i n g i n s i z e from 0 . 0 2 t o 1 0 7 . 5 ha. Motus e x t e n d a l o n g 55% o f t h e r e e f p e r i m e t e r . The c l o s e d l a g o o n , r i c h i n marine l i f e , c o n t a i n s a maze o f p a t c h r e e f s and impeccably c l e a r w a t e r .

The a t o l l , u n i n h a b i t e d , was " d i s c o v e r e d " by d e Q u i r o s i n 1606.

Although t r a c e s o f an a n c i e n t Tuamotuan c u l t u r e s t i l l e x i s t , t h e a t o l l a p p a r e n t l y n e v e r s u p p o r t e d a l o n g - t e r m permanent p o p u l a t i o n and h a s been l e s s a f f e c t e d by man t h a n most P a c i f i c i s l a n d s . I t s European h i s t o r y i n c l u d e s guano e x p o r t , a m u l t i n a t i o n a l e x p e d i t i o n t o o b s e r v e a s o l a r e c l i p s e , and c o p r a p r o d u c t i o n . I t has been u n i n h a b i t e d s i n c e t h e e a r l y 1930s ( a f a c t o r c o n t r i b u t i n g t o i t s r e l a t i v e l y u n d i s t u r b e d e c o l o g y ) , e x c e p t f o r t h e p r e s e n c e o f one f a m i l y from 1987 t o 1991. The p r i m a r y f a c t o r s r e s p o n s i b l e f o r i t s l a c k o f permanent s e t t l e m e n t a r e r e m o t e n e s s , a p p a r e n t a b s e n c e o f u s a b l e ground w a t e r , r e p e a t e d f a i l u r e o f i t s c o c o n u t p l a n t a t i o n s ( d i s e a s e s , d e s t r u c t i o n by c o c o n u t c r a b s , r a t s and s e a b i r d s , s m o t h e r i n g by v i n e s ) , a b s e n c e o f a p a s s a g e i n t o t h e l a g o o n , and a

p a u c i t y of s a f e b o a t a n c h o r a g e s .

U n t i l t h e 1988 USSR/USA e x p e d i t i o n , o n l y an 1883 c h a r t was

a v a i l a b l e , which named 7 i s l e t s . We d r a f t e d an a c c u r a t e map based on f i e l d work and r e c e n t a e r i a l p h o t o g r a p h s , naming 32 p r e v i o u s l y unnamed i s l e t s , 4 i s l e t g r o u p s and an i n l e t . During 8 d a y s ' i n t e n s i v e f i e l d work, we s u r v e y e d 38 i s l e t s , w a l k i n g 33 km i n s y s t e m a t i c c r o s s - i s l e t t r a n s e c t s and around i s l e t p e r i m e t e r s . T h i s p a p e r p r e s e n t s much new d a t a on C a r o l i n e from 3 v i s i t s i n 1988 and 1990, and a t t e m p t s t o summarize, expand and s y n t h e s i z e p r e v i o u s i n f o r m a t i o n i n t h e l i g h t o f new f i n d i n g s .

1 400 Snapfinger Drive, Athens, Georgia, 30605, USA 2 National Bioiogical Survey, School of Forest Resources

The University of Georgia, Athens, Georgia, 30602, USA

manuscript received 8 April 1991; revised 26 October 1993

(14)

S o i l s , p r i n c i p a l l y of c o r a l , mollusc, and a l g a l o r i g i n , a r e

c a t e g o r i z e d i n t o 5 t y p e s , from barren coral rubble t o rubble mixed with humus and guano. Caroline provides an e x c e l l e n t example of s o i l

development through d i f f e r e n t age and s i z e c l a s s e s of motus.

Preliminary d e s c r i p t i o n s of t h e a t o l l ' s r e e f , motu and lagoon

morphology a r e given, including t h e background geological s e t t i n g of t h e Line I s l a n d s . Physiographic f e a t u r e s include inland upraised r e e f s ( f e o ) and deep sand d e p o s i t s , coalesced i s l e t s , exposed o l d e r r e e f s , 1 i t h i f i ed beachrock, a congl omerate pl a t form, a "perched 1 agoon, " a nonfunctional hoa, and changes in motu s i z e and shape during t h e p a s t c e n t u r y .

C a r o l i n e ' s lush v e g e t a t i o n supports 26 s p e c i e s of p l a n t s organized i n t o 7 p l a n t communities, 6 n a t u r a l and one anthropogenic. The a t o l l - wide d i s t r i b u t i o n of each p l a n t s p e c i e s i s mapped. Plant s p e c i e s

( i n c l uding 5 new is1 and r e c o r d s ) and communities a r e d e t a i l e d , emphasizing t h e a t o l l ' s past h i s t o r y . The a t o l l ' s i n s u l a r f l o r a , although impoverished due t o i t s geographical l o c a t i o n , i s 89%

indigenous ( p o s s i b l y 92%), an extremely high f i g u r e f o r anywhere in t h e world. Although C a r o l i n e ' s motus a r e covered with e x t e n s i v e t r a c t s of

indigenous p l a n t communities, t h e P i s o n i a g r a n d i s f o r e s t s , u p t o 21 m high and covering 22% of t h e woodlands, a r e p a r t i c u l a r l y notable as they c o n s t i t u t e some of t h e b e s t groves l e f t in t h e P a c i f i c . P i s o n i a i s t r e a t e d in d e t a i l , including d a t a on r a p i d recovery and growth r a t e s during t h e p a s t 70-odd y e a r s . Tournefortia argentea (43% of t h e woodlands) i s abundant, and Cordia subcordata, becoming q u i t e r a r e elsewhere, occurs on 21 motus (54%). Cocos i s p r e s e n t , but only dominates one i s l e t ; 2 2 i s l e t s harbor wholly indigenous v e g e t a t i o n .

Motus of varied age and s i z e c l a s s e s provide e x c e l l e n t examples of s u b s t r a t e and vegetation development, accompanied by an i n c r e a s i n g d i v e r s i t y of bird l i f e . On account of i t s r e l a t i v e l y low human d i s t u r b a n c e and r a p i d f o r e s t recovery t o a more n a t u r a l s t a t e ,

e s p e c i a l l y s i n c e 1920, Caroline i s one of t h e few P a c i f i c i s l a n d s t h a t i s t r u l y an "outdoor ecological l a b o r a t o r y " : many motus have recovered so remarkably they a r e almost i n d i s t i n g u i s h a b l e from t h o s e t h a t have remained p r i s t i n e , while o t h e r s a r e i n d i f f e r e n t s t a g e s of recovery r e s u l t i n g from v a r i e d management ( o r non-management) p r a c t i c e s . Exotic p l a n t s p e c i e s a r e very few: a n c i e n t Polynesian-introduced (Cocos, p o s s i b l y Pandanus), r e c e n t Polynesian-introduced ( H i b i s c u s t i l i a c e u s , Thespesia populnea, Tacca 7eontopeta7oides, Ximenia americana), 20th century e x o t i c s ( P h y l l a n t h u s amarus, in one small a r e a ) . Some garden s p e c i e s , c u l t i v a t e d from 1987-1990 have an u n c e r t a i n f u t u r e and a r e not t r e a t e d as p a r t of t h e a t o l l ' s v i a b l e f l o r a .

An a n a l y s i s of ecological succession on motus of i n c r e a s i n g s i z e r e v e a l s t h a t by t h e time a motu reaches 0 . 8 ha in s i z e , a l l t h e n a t u r a l p l a n t communities, most p l a n t s p e c i e s , and most s p e c i e s of s e a b i r d s a r e p r e s e n t . This i s in s t r i k i n g c o n t r a s t t o s p e c i e s - a r e a r e l a t i o n s h i p s on i n h a b i t e d a t o l l s with more introduced

(15)

f e a t u r e 1 a t

Each motu i s i n d i v i d u a l l y mapped and t h e main p h y s i o g r a p h i c r e s , known h i s t o r y , v e g e t a t i o n p a t t e r n s ( i n c l u d i n g s p e c i e s - a r e a i o n s h i p s ) , s e a b i r d s , and m i s c e l l a n e o u s b i o t a ( c o c o n u t c r a b s , r a t s , l i z a r d s ) a r e d e t a i l e d . Appendices p r o v i d e w e a t h e r d a t a and d e s c r i b e t h e p r a c t i c a l i t i e s o f a n c h o r i n g b o a t s , l a n d i n g , and moving small b o a t s

around t h e r e e f s and l a g o o n .

Permanent p r o t e c t i o n of C a r o l i n e i s c u r r e n t l y underway a s The Nature Conservancy of Hawaii n e g o t i a t e s w i t h t h e government of K i r i b a t i f o r a S o u t h e r n Line I s l a n d s W i l d l i f e P r e s e r v e , which i n c l u d e s C a r o l i n e , Vostok and F l i n t .

A. INTRODUCTION

Carol i n e toll

'

( F r o n t i s p i e c e ; F i g s . 1 , 2 ) i s a s m a l l , 1 ow c o r a l i s l a n d s i t u a t e d a t 10°0O'S l a t i t u d e , 15O013'W l o n g i t u d e i n t h e s o u t h - c e n t r a l P a c i f i c Ocean. I t s maximum h e i g h t i s l e s s t h a n 3 m above MSL.

I t l i e s 2,800 km s o u t h o f Hawaii, 830 km n o r t h o f T a h i t i , and 1 , 0 3 0 km west o f t h e Marquesas I s l a n d s . I t s n e a r e s t n e i g h b o r s a r e Vostok and F l i n t , 230 km t o t h e west and s o u t h w e s t , r e s p e c t i v e l y .

Recent measurements by t h e ICBP 1990 Line and Phoenix I s l a n d s

E x p e d i t i o n , u s i n g a compact s a t e l l i t e n a v i g a t i o n computer "Magellan" NAV 1000, i n d i c a t e t h a t t h e a t o l l l i e s one n a u t i c a l m i l e e a s t o f i t s

p r e v i o u s l y c h a r t e d l o n g i t u d e p o s i t i o n , 150°14'W. I t s r a n g e o f c o o r d i n a t e s a r e : 0g054' t o 1 0 ° 0 1 ' S l a t i t u d e , 150'12' t o 150'14' W l o n g i t u d e . The g i v e n c o o r d i n a t e s , 10°0O'S and 150°13'W, i n t e r s e c t i n t h e l o w e r lagoon j u s t west o f t h e " b l i n d p a s s a g e . "

Although a r c h a e o l o g i c a l l y and g e o g r a p h i c a l l y w i t h i n P o l y n e s i a , t h e Line Group was u n i n h a b i t e d when d i s c o v e r e d by Europeans; i t s i s l a n d s were v a r i o u s l y c l a i m e d by t h e United S t a t e s and England. With t h e e x c e p t i o n of U.S.-owned J a r v i s , Palmyra, J o h n s t o n and Kingman Reef, a l l a r e now governed by t h e R e p u b l i c o f K i r i b a t i ( f o r m e r l y G i l b e r t I s l a n d s ) ,

C a r o i i n e , 9 . 7 km l o n g by 2 . 3 km wide a t i t s w i d e s t p o i n t , i s a c r e s c e n t i c c o r a l r i n g 2 6 . 9 km i n c i r c u m f e r e n c e . Composed o f 39 i s l e t s

( F i g . 2 , T a b l e 9 ) and t h r e e i n c i p i e n t i s l e t s , i t i s c e n t e r e d on a

c o n t i n u o u s r e e f f l a t , submerged a t high t i d e , t h a t e n c l o s e s a r e l a t i v e l y s h a l l o w l a g o o n . Most a r e well-wooded, b u t 4 t i n y o n e s , l e s s t h a n 0 . 1 ha i n s i z e , a r e s c a r c e l y more t h a n c o r a l r u b b l e p i l e d on t h e r e e f ,

s u p p o r t i n g s p a r s e p a t c h e s o f T o u r n e f o r t i a and H e l i o t r o p i u m . One i s l e t , Noddy Rock ( P I . 1 8 ) , i s a v e s t i g e of a f o r m e r r e e f segment. The t o t a l l a n d a r e a above high w a t e r i s 399 ha. One o f t h e o c e a n i c i s l a n d s c o n t r i b u t i n g t o Darwin's t h e o r y o f a t o l l f o r m a t i o n (Darwin 1 8 4 2 ) , i t s

' c a r o l i n e At01 1 i s n e i t h e r p h y s i c a l l y , g e o g r a p h i c a l l y , n o r pol i t i c a l 1 a s s o c i a t e d w i t h t h e C a r o l i n e I s l a n d s , now p a r t o f t h e F e d e r a t e d S t a t e s of M i c r o n e s i a , more t h a n 6,000 km t o t h e n o r t h w e s t . Because o f t h i s c o n f u s i o n , we use t h e name " C a r o l i n e A t o l l " i n s t e a d o f " C a r o l i n e I s l a n d . "

(16)

H A W A I I

.

S C A L E A T E Q U A T O R

K I N G M A N R E E F

. J A R V I S I S . \ .

Q

M A L D E N . 9

I S . I

0 . F I L I P P O REEF^

S T A R B U C K I S . I-- -

C I

\

\ I

\ -a I

+\

I V O S T O K 1s. , I

I

1 F L I N T I S .

L - - - - ---,--,,- J

I R A R O T O N G A " I

I ^I_I^{I .}

tropic o

--- -- ---

^I

I I I B

Figure 1. Line Islands: geographic location in the Pacific Ocean.

After map by the Hawaii Geographic Society (1981).

(17)

r

P A N O A N U S I S L E T 1

S O U T H N I S L E T

CENTRAL L E E W A R D

I S L E T S

D A N G E R I S L E T

-____

AKE

1

8 0 0 8 ~ I S L E T

-4

C C O R A L I S L E T

/

L O N E P A L M I S L E T \

M O T U K O T A L M O T U

I S L A N D

B O ' S U N B I R D

i

B L A C K F I N I S M O T U U A T A W

E M E R A L D I S L W I N D W A R D I S 1

C R E S C E N T I S L

N O R T H P I G I S L E T

WINDWARD I S L E T S

B R O T H E R S I S L E T M O D D Y R O C K N O R T H A R U N D E L I S L E T

P I S O N I A I S L E T / a R U M D E L I S L E T

S C A L E 1 : 3 4 . 0 0 0

5 0 0

-

0 m e t e r s I 0 0 1000 2 0 0 0

Figure 2 . C a r o l i n e A t o l l , Republic of K i r i b a t i , with newly-named is1 e t s . Based on photos by t h e Royal New Zealand Air Force (RNZAF 1986).

(18)

g e o l o g y , s o i l s , c l i m a t e , and v e g e t a t i o n a r e t y p i c a l o f low l a t i t u d e a t 0 1 1 s and a r e r e 1 a t i v e l y unmodified by man.

T h i s p a p e r p r e s e n t s much new d a t a on C a r o l i n e from 3 v i s i t s i n 1988 (September) and 1990 (March, May) and a t t e m p t s t o summarize, expand and s y n t h e s i z e w i d e l y s c a t t e r e d i n f o r m a t i o n i n t h e l i g h t o f new

f i n d i n g s . The o n l y s i g n i f i c a n t p r e v i o u s b i o l o g i c a l i n f o r m a t i o n on C a r o l i n e was from t h e S m i t h s o n i a n ' s P a c i f i c I s l a n d B i o l o g i c a l Survey Program's 2-day v i s i t i n 1965 (Clapp & S i b l e y 1 9 7 1 ) .

TOLL

C e n t u r i e s b e f o r e Europeans e n c o u n t e r e d C a r o l i n e , t h e a t o l l was i n h a b i t e d by P o l y n e s i a n s . No o r a l t r a d i t i o n s o f t h i s o c c u p a t i o n a r e known, but e v i d e n c e o f former h a b i t a t i o n was e v i d e n t when de Q u i r o s found t h e a t o l l i n 1606. He n o t e d "an o l d c a n o e , l y i n g on h e r s i d e , "

and a small g r o v e of c o c o n u t s p l a n t e d on South I s l a n d ( B e n n e t t 1840, Markham 1 9 0 4 ) .

No f u r t h e r c l u e s were u n e a r t h e d u n t i l Messrs. Brown, B r o t h e r s , and Arundel exposed about 50 a n c i e n t P o l y n e s i a n s i t e s i n t h e 1870s w h i l e d i g g i n g f o r guano (Holden 1884, Arundel 1 8 9 0 ) . P o l y n e s i a n i n h a b i t a n t s a t t h a t t i m e c a l l e d them " m a r a i " ( m a r a e ) , Arundel photographed and drew p l a n s of them ( F i g . 3 ) : d e p i c t e d a r e a p l a t f o r m o f c o r a l and

c o n g l o m e r a t e r o c k , s u r r o u n d e d by 10 s m a l l e r s l a b s r e s e m b l i n g

g r a v e s t o n e s , a l l a r r a n g e d i n a r e c t a n g u l a r p l a n , Although t h e l a r g e s t 2 were marked a s " g r a v e s " on A r u n d e l ' s 1883 map ( F i g . 4 ) , no b o n e s , a s h e s o r human remains were f o u n d . T h e i r f i n d i n g s were l a t e r i d e n t i f i e d a s Tuamotuan marae (Emory 19471, Marae, a c c o r d i n g t o a n c i e n t b e l i e f ,

"bound t h e a n c e s t r a l s p i r i t s and gods o f t h e k i n d r e d t o t h e l a n d ,

p u t t i n g i t under t h e i r e t e r n a l g u a r d i a n s h i p " (Emory 1 9 4 7 ) . The l a r g e s t marae was on n o r t h w e s t Nake I s l a n d , and a s m a l l e r one was found n e a r t h e

s o u t h e r n t i p o f Long I s l a n d , Both l o c a t i o n s conform t o such

p r e r e q u i s i t e s f o r b u i l d i n g marae a s nearby s h o r e l i n e s and b i r d s ( s e e S e c t . M.1), which Tuamotuans b e l i e v e d housed d i v i n e s p i r i t s (Emory 1947, p. 1 2 3 ) . Although AKK, G . Wragg and R , F a l c o n e r l o c a t e d , photographed

( P I . 3 6 ) , and measured t h e s e i n 1990 (A. K e p l e r 1 9 9 0 d ) , no f i e l d work by a r c h a e o l o g i s t s h a s been c o n d u c t e d , Our d a t a and numerous p h o t o g r a p h s have been s e n t t o Dr. J e f f I r w i n , Auckland U n i v e r s i t y , New Z e a l a n d . P o s t - E u r o ~ e a n H i s t o r y : 1 7 t h t o 1 9 t h C e n t u r i e s

On 21 February 1606, t h e P o r t u g u e s e e x p l o r e r Pedro Fernandez

d e Q u i r o s , employed by S p a i n , " d i s c o v e r e d " C a r o l i n e A t o l l (Markham 1904, S t e v e n s & Barwick 1 9 3 0 ) , naming i t San Bernardo. D e s p i t e i t s

r e m o t e n e s s , C a r o l i n e was e n c o u n t e r e d e a r l y i n P a c i f i c h i s t o r y , l o n g b e f o r e T a h i t i , Rarotonga, and Hawaii.

l o c a t i o n , f o r e a r l y n a v i g a t o r s t e n d e d t o s a i l due w e s t from South

(19)

(20)

America a l o n g l i n e s o f l a t i t u d e , and 1 0 " s was an o b v i o u s c h o i c e . De Q u i r o s , t h e l a s t a d v e n t u r e r i n t h e S p a n i s h age o f d i s c o v e r y , was l e a d i n g h i s second major t r a n s - P a c i f i c e x p e d i t i o n w i t h 3 s h i p s and 150 men o b s e s s e d w i t h f i n d i n g t h e f a b l e d " T e r r a A u s t r a l i s I n c o g n i t o . " The d e s c r i p t i o n s o f C a r o l i n e by h i s crew, a l t h o u g h a t v a r i a n c e w i t h one a n o t h e r , s t i l l a p p l y t o d a y ( P I . 1 ) . T h e i r f i r s t a t - s e a i m p r e s s i o n was t h a t i t was " d i v i d e d i n t o f o u r o r f i v e hummocks, and a l l t h e r e s t submerged. I t s c i r c u m f e r e n c e appeared t o be t e n l e a g u e s " (Markham 1 9 0 4 ) . A f t e r l a n d i n g , t h e y found t h a t

"There was a g r e a t number of f i s h i n s h o r e , and, owing t o t h e w a t e r b e i n g v e r y s h a l l o w , t h e y were k i l l e d w i t h swords and p o l e s . T h e r e were g r e a t numbers of l o b s t e r and c r a w f i s h , and o t h e r k i n d s of marine a n i m a l s . They found a g r e a t q u a n t i t y o f c o c o a - n u t s i n a heap a t t h e f o o t of t h e palm t r e e s , many

l a r g e , and o f d i f f e r e n t s i z e s . There were a g r e a t q u a n t i t y o f s e a b i r d s o f s e v e r a l k i n d s , and s o i m p o r t u n a t e t h a t t h e y

seemed t o want t o a t t a c k t h e men. We t o o k p l e n t y o f a l l t h e s e t h i n g s

. . .

I t seemed t o t h e C a p t a i n t h a t on an i s l a n d where t h e r e a r e s o many t r e e s t h e r e c o u l d n o t f a i l t o be w a t e r "

(Markham 1 9 0 4 ) .

Fresh w a t e r was c r u c i a l t o de Quiros and h i s crew, who were

s u f f e r i n g from l a c k o f food and w a t e r . D e s p i t e t h e i r e f f o r t s , however, t h e y f a i l e d t o o b t a i n w a t e r . D i s a p p o i n t e d and l a c k i n g e n e r g y , t h e y c o n t i n u e d t h e i r voyage t h e f o l l o w i n g morning. T h e i r d e m o r a l i z e d s t a t e may e x p l a i n one s t a t e m e n t t h a t C a r o l i n e " c o n s i s t e d o f t w e n t y - t w o i s l e t s , u n i n h a b i t e d and w i t h o u t w a t e r , t r e e s o r s c r u b f o r wood."

In 1795, C a p t . W . R . Broughton, on t h e B r i t i s h s l o o p P r o v i d e n c e , r e d i s c o v e r e d and named t h e a t o l l w h i l e voyaging from T a h i t i t o Hawaii

(Broughton 1804):

"The s o u t h e r n e x t r e m i t y was t h e h i g h e s t p a r t , c o v e r e d w i t h t r e e s , most p r o b a b l y c o c o a - n u t from t h e i r a p p e a r a n c e , a s t h e y s t o o d i n d e t a c h e d clumps along t h e s h o r e . The i s l a n d

. . .

appeared t o be low, and covered w i t h t r e e s , and i f I am r i g h t i n i t s e s t i m a t e d d i s t a n c e , i t s l e n g t h w i l l be about f i v e m i l e s i n a n o r t h and s o u t h d i r e c t i o n . I named i t C a r o l i n a I s l a n d i n compliment t o t h e d a u g h t e r o f S i r P. S t e p h e n s of t h e

Admiral t y

.

"

Because e a r l y n a v i g a t i o n t e c h n i q u e s and communication were f a r l e s s s o p h i s t i c a t e d t h a n t o d a y , e s p e c i a l l y w i t h r e g a r d s t o l o n g i t u d e , C a r o l i n e was s i g h t e d o r " d i s c o v e r e d " by s e v e r a l more e x p l o r e r s who were u n s u r e o f i t s i d e n t i t y . By 1821 t h e a t o l l had amassed a c o l l e c t i o n o f c o o r d i n a t e s and names: San Bernardo, I s l a n d o f F i s h , T h o r n t o n , H u r s t ' s , C l a r k ' s , Independence, and C a r o l i n a (which l a t e r became C a r o l i n e ) . Some n a v i g a t o r s e q u a t e d C a r o l i n e w i t h an i s l a n d named "San Bernardo" by t h e S p a n i s h e x p l o r e r Mendatia i n 1595. "San Bernardo" has r e c e n t l y been v e r i f i e d a s Puka-Puka, n o r t h e r n Cook I s l a n d s (Maude 1 9 6 8 ) .

(21)

The b e s t e a r l y d e s c r i p t i o n o f t h e a t o l l comes from an 1835 v i s i t by F . D . B e n n e t t , who was r e a s o n a b l y well v e r s e d i n n a t u r a l h i s t o r y ( B e n n e t t 1 8 4 0 ) . He n o t e d t h a t t h e i s l e t s t h e n , a s now, were " c o v e r e d w i t h v e r d u r e

...

s u r p r i s i n g l y l u x u r i a n t , when compared t o t h e a r i d s o i l i t c o v e r s . " A1 though B e n n e t t had v i s i t e d many a t 0 1 1 s , he was p a r t i c u l a r l y impressed w i t h t h e q u a l i t y o f C a r o l i n e ' s c o r a l r e e f s . His p a r t y

o b s e r v e d " r a t s o f a red-brown c o l o r " and v a r i o u s b i r d s b u t no r e p t i l e s ( P t . 1 1 ) Although he d i s c u s s e s " l a n d l o b s t e r s ( C o e n o b i t a s p e c i e s ) , " no mention i s made o f coconut c r a b s ( B i r g u s l a t r o ) : p e r h a p s t h e l a t t e r were lumped w i t h t h e f o r m e r .

First O c c u ~ a t i o n : The e x i s t e n c e o f 2 small coconut g r o v e s on C a r o l i n e prompted 2 B r i t i s h e n t r e p r e n e u r s , r e p r e s e n t i n g t h e T a h i t i a n f i r m C o l l i e and L u c e t t , t o e s t a b l i s h a s t o c k r a i s i n g v e n t u r e t h e r e i n 1846. T h i s f i r s t known s e t t l e m e n t was l o c a t e d a d j a c e n t t o t h e main c o c o n u t g r o v e on t h e n o r t h w e s t p e n i n s u l a o f South I s l a n d ; a s m a l l e r g r o v e e v i d e n t l y

e x i s t e d "on t h e s o u t h - s o u t h - w e s t s i d e " o f t h e same i s l a n d ( L u c e t t 1 8 5 1 ) . T a h i t i a n l a b o r e r s tended p i g s , hens, t u r k e y s , and grew many food p l a n t s , i n c l uding pumpkins and me1 o n s . They d r i e d and s a l t e d f i s h , pl a n t e d c o c o n u t s , and e x t r a c t e d coconut o i l (Maude 1942, G a r n e t t 1 9 8 3 ) , and were e v i d e n t l y s t i l l t h e r e i n May 1852 ( E l l s w o r t h 1 9 9 0 ) ,

n n e x a t i o n : Though i n h a b i t e d i n p r e h i s t o r y by Tuamotuans, o f f i c i a l l y " d i s c o v e r e d " by t h e S p a n i s h , and v i s i t e d by B r i t i s h , F r e n c h , and American s h i p s , i t t o o k c e n t u r i e s f o r C a r o l i n e t o a c q u i r e a

p o l i t i c a l i d e n t i t y . I t was f o r m a l l y annexed t o B r i t a i n by C a p t a i n Nares, R . N., who a r r i v e d i n t h e H.M.S. R e i n d e e r i n 1868, f i n d i n g 27 r e s i d e n t s .

C a r o l i n e was under t h e c o n t r o l of v a r i o u s merchants i n t h e l a t e 1 9 t h c e n t u r y : Lionel Brown, C a p t a i n B r o t h e r s , and l a t e r John A r u n d e l , a well-known businessman, t r a d e r , and guano merchant i n t h e P a c i f i c .

A r u n d e l ' s 1883 map ( F i g . 4 ) of C a r o l i n e i s t h e o n l y r e a s o n a b l y c o r r e c t c h a r t p u b l i s h e d p r i o r t o t h i s p a p e r ,

Though bonded under t h e American Guano Act i n 1860, no p h o s p h a t e was dug on C a r o l i n e u n t i l Arundel was g r a n t e d a 7 - y e a r l i c e n s e i n 1874, A few months e a r l i e r , a s e t o f moorings were l a i d o f f t h e l e e s i d e of South I s l a n d , a l l o w i n g s h i p s o f up t o 1 , 0 0 0 t o n s t o l i e s a f e l y d u r i n g t r a d e wind w e a t h e r . Guano was t h e o n l y s u c c e s s f u l b u s i n e s s v e n t u r e a t C a r o l i n e : a p p r o x i m a t e l y 10,000 t o n s were s h i p p e d t o C a l i f o r n i a and A u s t r a l i a between 1873 and 1895, when s u p p l i e s became e x h a u s t e d (Young c a . 1 9 2 2 ) . We have no d i r e c t i n f o r m a t i o n on which i s l a n d s were mined e x c e p t South and Nake. We s t r o n g l y s u s p e c t t h a t T r i d a c n a , A r u n d e l , Mannikiba, and p e r h a p s o t h e r s a l s o y i e l d e d guano. The t o n n a g e e x t r a c t e d from C a r o l i n e was small compared t o t h a t from d r y , b a r r e n , more

n o r t h e r l y i s l a n d s w i t h l i t t l e v e g e t a t i o n , where p o p u l a t i o n s o f Sooty

(22)

Sol ar Ecl i p s e E x p e d i t i o n

In 1883 C a r o l i n e r e c e i v e d i n t e r n a t i o n a l p u b l i c i t y when a s t r o n o m e r s c a l c u l a t e d t h a t i t l a y d i r e c t l y under t h e p a t h o f a pending s o l a r

e c l i p s e . T h r e e p a r t i e s o f a s t r o n o m e r s (American, B r i t i s h , French) a r r i v e d i n t h e U.S.S. H a r t f o r d and s e t up camp on South I s l a n d , making d e t a i l e d o b s e r v a t i o n s o f t h i s c e l e s t i a l e v e n t ( P I . 2 ) . A t t h a t t i m e C a r o l i n e was more famous, and housed more p e o p l e , t h a n b e f o r e o r s i n c e : 7 " n a t i v e s , " s c i e n t i s t s and crewmen t o t a l l e d 51 o c c u p a n t s . L e g a c i e s from f o r m e r i n h a b i t a n t s i n c l u d e d 3 houses ( P I . 3 ) , 2 s h e d s , 3 h u t s on s m a l l e r motus, n a u t i c a l f l o t s a m and j e t s a m , and 2 s h a l l o w w e l l s . To t h i s t h e y added t e n t s , o b s e r v a t o r y f r a m e s , a m a r b l e s l a b , f l a g p o l e , and b r i c k " p i e r s " f o r t h e i r t e l e s c o p e s , most o f which remained t e m p o r a r i l y a s t e c h n o l o g i c a l 1 i t t e r .

T h i s e x p e d i t i o n (Dixon 1884, Holden 1884, Holden & Q u a l t r o u g h 1884, T r e l e a s e 1884, Young 1884) a l s o marked t h e f i r s t a t t e m p t t o d e s c r i b e t h e t o p o g r a p h y , c l i m a t e , f l o r a , and f a u n a o f t h e a t o l l . Drawings i n c l u d e d an a r t i s t ' s r e n d e r i n g of C a r o l i n e , " s e t t l e m e n t " map

( P I . 5 ) and views a l o n g South Is1 a n d ' s 1 agoon s h o r e ( P l s o 6 , 7 ) , An a t o l l map was d r a f t e d b u t i s h i g h l y i n a c c u r a t e . T h e i r b i o l o g i c a l o b s e r v a t i o n s were s k e t c h y (Dixon 1884, B u t l e r & S t r e c k e r 1 8 8 4 ) . For example, Dixon, t h e z o o l o g i s t , l i s t e d such organisms a s " s h r i m p , "

" h e r m i t c r a b s , " " g n a t . " As w i t h B e n n e t t , t h e r e was no mention o f

c o c o n u t c r a b s , even though t h e y were e v i d e n t l y abundant on South I s l a n d i n 1910 (Young c a . 1 9 2 2 ) .

The L a t e 1 9 t h and 2 0 t h C e n t u r i e s

In 1875, C . D . Voy, a n a t u r a l i s t from C a l i f o r n i a , v i s i t e d Carol i n e , c o l l e c t i n g moll u s c s ( P i 1 s b r y & V a n a t t a 1 9 0 5 a , b) and f i s h

( F o w l e r 1901) ,

In 1885, Arunde! began t o c l e a r 1 and and p l a n t c o c o n u t s , b u t h i s p l a n n e d c o p r a i n d u s t r y was u n s u c c e s s f u l , In 1897 he s o l d h i s b u s i n e s s t o t h e P a c i f i c I s l a n d s Company, L t d . , which a l s o f a i l e d . The

p l a n t a t i o n s s u f f e r e d from d i s e a s e and poor v i t a l i t y , c o c o n u t c r a b s chewed on s e e d l i n g s , s e a b i r d s d e s t r o y e d t h e d e v e l o p i n g n u t s , Iponroea v i n e s s t r a n g l e d young t r e e s , and p o p u l a t i o n s o f P o l y n e s i a n r a t s a p p a r e n t l y e x p l o d e d , c a u s i n g f u r t h e r damage t o both i n t a c t n u t s and d r y i n g c o p r a (Young c a . 1922, Maude c a . 1938, 1 9 4 2 ) . By 1904, when t h e H.M.S. I c a r u s v i s i t e d C a r o l i n e , o n l y 6 P o l y n e s i a n s l i v e d t h e r e . A few months l a t e r t h e y were r e p a t r i a t e d t o Niue, and C a r o l i n e remained u n i n h a b i t e d u n t i l 1916, when a new e f f o r t was made t o d e v e l o p t h e c o c o n u t pl a n t a t i on by Messrs. S . R . Maxwell and Co., L t d .

During t h e u n i n h a b i t e d y e a r s , South I s l a n d ' s v e g e t a t i o n and w i l d l i f e began t o r e c o v e r from t h e e a r l i e r f o r e s t f e l l i n g ( P l s . 2 - 6 ) . When Mr. J . L . Young, t h e n managing d i r e c t o r f o r S . R . Maxwell and C o . , L t d . (Young c a . 1 9 2 2 ) , v i s i t e d t h e a t o l l i n J u l y 1910, he d e s c r i b e d i t a s a w i l d e r n e s s , teeming w i t h S o o t y T e r n s , f i s h and c o c o n u t c r a b s :

(23)

"The ground was covered with n e s t s of s e a b i r d s which l a t t e r r o s e l i k e a cloud when d i s t u r b e d : t h e noise of t h e i r s h r i e k i n g was so g r e a t t h a t one had t o shout t o enable oneself t o be heard by h i s companions. Hundreds of g r e a t Coconut Crabs were seen: 40 l a r g e ones were caught by t h e crew of t h e schooner in an hour. The r e e f and t h e lagoon swarmed with f i s h and small s h a r k s . "

From 1916 t o 1929, Caroline was a l t e r e d more than before o r s i n c e . All t h e a v a i l a b l e land on South was d e f o r e s t e d t o make room f o r

thousands of palms, and l a b o r e r s demolished huge numbers of coconut c r a b s and s e a b i r d s (Young c a . 1922). In a d d i t i o n , coconuts were planted on a l l of t h e main windward i s l e t s , southern Nake, and on Mannikiba.

(The windward i s l e t s recovered t h e i r f o r e s t s remarkably q u i c k l y , s e e S e c t . G). P l a n t a t i o n workers in g r e a t p a r t l i v e d o f f t h e l a n d , f e a s t i n g on f r e s h f i s h , s e a b i r d s , s e a b i r d eggs, t u r t l e s , and coconut c r a b s . Most of t h e leeward i s l e t s escaped a l t e r a t i o n f o r p l a n t a t i o n s .

Copra e x p o r t s averaged around 14 t o n s per y e a r from 1929 t o 1934, a f t e r which t h e company ran i n t o d e b t . Concurrently, t h e French

government forbade f u r t h e r recruitment from T a h i t i ; by 1936 only a few f a m i l i e s were l e f t (N.I.D. 1943). In 1941 t h e a t o l l c a r r i e d a p r i c e t a g of 600 English pounds (Maude, p e r s . comm.), but was never purchased.

Occupation l e a s e s f o r Caroline were c a n c e l l e d in 1943, a f t e r which t h e B r i t i s h Western P a c i f i c High Commission repossessed i t (Maude 1953).

However, new "queen's l e a s e s " were granted t o M . P . A . Bainbridge of Papeete, T a h i t i , 1951-1964 (Nicholson & Douglas 1969), then t o

Capt. Omer Darr of Moorea, French Polynesia, from 1964 t o 1989. When t h e B r i t i s h granted independence t o t h e G i l b e r t and E l l i c e I s l a n d s in 1979, a new country, t h e Republic of K i r i b a t i , assumed ownership of C a r o l i n e , along with most of t h e Line and Phoenix I s l a n d s .

Apart from occasional p a r t i e s of T a h i t i a n s c u t t i n g copra and a shipwrecked s a i l o r in t h e e a r l y 1980s, t h e a t o l l remained uninhabited f o r over 50 y e a r s . During t h i s time C a r o l i n e ' s v e g e t a t i o n and w i l d l i f e recovered t o such an e x t e n t t h a t , were i t not f o r unpublished

manuscripts (Maude c a . 1938, c a . 1942, and no d a t e , Young ca. 1922) and comparisons with F l i n t and VosLok ( S t . John S Fosberg 1937, A. Kepler 1990b-d, and Kepler, in p r e p . ) , we would have been unaware of t h e e x t e n t of previous human i n t e r f e r e n c e o r of t h e r a p i d i t y of f o r e s t recovery

( t h e f a c t t h a t 60% of C a r o l i n e ' s motus harbored wholly indigenous v e g e t a t i o n , and t h e presence of one small patch of one e x o t i c p l a n t a s i d e from a few standard Polynesian-introduced p l a n t s , seemed t o i n d i c a t e a r e l a t i v e l y p r i s t i n e a t o l l )

.

In 1987, t h e O f f i c e de l a Recherche S c i e n t i f i q u e e t Technique

Outre-Mer (ORSTOM, a French s c i e n t i f i c research agency) was requested by t h e K i r i b a t i government t o conduct a s h o r t study a t Caroline on t h e f e a s i b i l i t y of p e a r l - s h e l l c u l t u r e ( G . Monet, p e r s . comm.). Their

r e s u l t s concluded t h a t t h e a t o l l would be i n a p p r o p r i a t e f o r t h i s type of

NO. 397

...

. .

...

... x

...

.

...

...

.

...

...

...

...

...

...

.

. . .

. . .

. . . .

. . .

. . .

. . .

. . .

. . .

...

. . .

. . .

. . .

...

. . .

...

. . .

. . .

. . .

. . .

. . .

. . .

. . .

. . .

.

. . .

. . . .

. . .

. . .

. . .

. . .

. . .

. . .

. . .

.

. . .

. . .

. . .

. . .

. . .

...

...

...

.

...

...

...

. . .

...

. . .

. . .

. . .

...

...

. . .

. . .

...

...*

. . .

. . .

. . .

. . .

. . .

. . .

_{. .}

... ^x