ATOLL RESEARCH BULLETIN NO. 251 BIOGEOGRAPHY OF THE

Transcription

ATOLL RESEARCH BULLETIN
NO. 251
BIOGEOGRAPHY OF THE PUERTO RICAN BANK
by Harold Heatwole, Richard Levins and Michael D. Byer
Issued by
THE SMITHSONIAN INSTITUTION
Washington, D. C., U.S.A.
July 1981
VIRGIN ISLANDS
CULEBRA
PUERTO R l C O
F i g . 1. Map of t h e P u e r t o Rican I s l a n d S h e l f . R e c t a n g l e s A - E i n d i c a t e b o u n d a r i e s of maps
p r e s e n t e d i n more d e t a i l i n Appendix I . 1. Cayo S a n t i a g o , 2 . Cayo B a t a t a , 3. Cayo de
Afuera, 4. Cayo de T i e r r a , 5 . Cardona Key, 6 . P r o t e s t a n t Key, 7 . Green Key ( s t . ~ r o i x ) ,
8. Caiia Azul
ATOLL RESEARCH BULLETIN 251
ERRATUM
The following caption should be inserted for figure 7:
Fig. 7. Temperature in and near a small clump of vegetation
on Cayo Ahogado. Dots: 5 cm deep in soil under clump.
Circles: 1 cm deep in soil under clump. Triangles: Soil
surface under clump. Squares: Surface of vegetation.
X's: Air at center of clump. Broken line indicates
intervals of more than one hour between measurements.
BIOGEOGRAPHY OF THE PUERTO RICAN BANK
by Harold Heatwolel, Richard Levins2 and Michael D. Byer3
INTRODUCTION
There h a s been a r e c e n t s u r g e of i n t e r e s t i n t h e biogeography of
a r c h i p e l a g o e s owing t o a r e i n t e r p r e t a t i o n of c l a s s i c a l concepts of
e v o l u t i o n of i n s u l a r p o p u l a t i o n s , f a c t o r s c o n t r o l l i n g numbers of
s p e c i e s on i s l a n d s , and t h e dynamics o f i n t e r - i s l a n d d i s p e r s a l .
The
l i t e r a t u r e on t h e s e s u b j e c t s i s r a p i d l y accumulating; g e n e r a l reviews
a r e p r e s e n t e d by Mayr (1963) , and Baker and S t e b b i n s (1965)
C a r l q u i s t (1965, 1 9 7 4 ) , P r e s t o n (1962 a , b ) , ~ a c ~ r t h u
and
r Wilson
(1963, 1967) , MacArthur et al. (1973) , Hamilton and Rubinoff (1963,
1 9 6 7 ) , Hamilton et al. (1963) , Crowell (19641, Johnson (1975) ,
Whitehead and Jones ( 1 9 6 9 ) , Simberloff (1969, 19701, Simberloff and
Wilson ( 1 9 6 9 ) , Wilson and Taylor (19671, Carson ( 1 9 7 0 ) , Heatwole and
Levins (1973) , Abbott (1974) , Johnson and Raven (1973) and Lynch and
Johnson ( 1 9 7 4 ) , have provided major impetuses through t h e o r e t i c a l and/
o r g e n e r a l p a p e r s on numbers of s p e c i e s on i s l a n d s and t h e dynamics of
i n s u l a r biogeography and e v o l u t i o n .
Other work h a s d e a l t w i t h
s p e c i f i c problems such a s d i s p e r s a l ( G r e s s i t t and Yoshimoto 1963,
C a r l q u i s t 1966 a-c, 1967, Heatwole and Levins 1972b) , i n t e r s p e c i f i c
competition (Grant 1965, 1966, 1968, Crowell 1962, 1968, Sheppard et al.
1 9 6 8 ) , e f f e c t of d i s a s t e r s ( B r a t t s t r o m 1963, Sauer 19621, t r o p h i c
r e l a t i o n s (Heatwole 1971, Heatwole and Levins 1972a1, c o l o n i z a t i o n
(Bassot and B a l l 1971, Hunt and Hunt 1974, Becker 1975, Schoener 1975
and Williams 19691, and endemism ( ~ a y r1965, Heatwole and MacKenzie
.
Department of Zoology, U n i v e r s i t y of New England, Armidale, N.S.W.,
2351, A u s t r a l i a .
Department of P o p u l a t i o n S c i e n c e s , Harvard School of P u b l i c H e a l t h ,
665 Huntington Avenue, Boston, Massachusetts, 02115, U.S.A.
Departmento de ~ i o l o ~ i Universidad
a,
de P u e r t o Rico, Rio P i e d r a s ,
P u e r t o Rico, 00931.
1967, Thornton 1967).
I n a d d i t i o n t h e r e have been a l a r g e number of
s t u d i e s t r e a t i n g t h e d i s t r i b u t i o n and v a r i a t i o n of s p e c i f i c groups of
organisms on i s l a n d s and t h e r o l e of h i s t o r i c a l f a c t o r s upon such
patterns.
W i t h this broadening base of t h e o r e t i c a l background and
accumulation of information on s p e c i f i c a s p e c t s , t h e f i r s t two a u t h o r s
f e l t i t opportune t o c a r r y o u t a comprehensive study of an archipelago
conta.ining many i s l a n d s with a v a r i e t y of s i z e s , i n t e r - i s l a n d d i s t a n c e s ,
topography, v e g e t a t i o n , time of s e p a r a t i o n , use by humans, and o t h e r
c h a r a c t e r i s t i c s , and t o s t u d y a s many a s p e c t s of t h e biogeography of a s
wide a range of organisms a s f e a s i b l e .
Our aim was t o look a t the
s u b j e c t a s a whole and thereby c o n t r i b u t e toward a s y n t h e t i c
understanding o f t h e broad a s p e c t s of t h e problem.
The o r i g i n a l
design of t h e p r o j e c t was p r e s e n t e d i n an e a r l y paper (Levins and
Heatwole 1963) and some of t h e d a t a have been used i n g e n e r a l reviews
(Levins 1968, 1969, Heatwole 1976a).
This paper p r e s e n t s t h e
geography, v e g e t a t i o n , and e c o l o g i c a l c h a r a c t e r i s t i c s of t h e P u e r t o
Rican Bank and c o n s t i t u t e s a background f o r t h e r e s t of t h e s e r i e s
(some o f which have a l r e a d y been p u b l i s h e d , e . g. Heatwole and Levins
1972b, 1973, Levins and Heatwole 1973, Levins e t a l . 1 9 7 3 ) , and f o r
papers by a v a r i e t y of s p e c i a l i s t s using t h e m a t e r i a l we c o l l e c t e d
d u r i n g our study.
GENERAL DESCRIPTION OF THE REGION AND ITS CLIMATE
The a r e a s t u d i e d c o n s i s t s of a l l t h e i s l a n d s and cays i n t h e p a r t
of t h e A n t i l l e a n chain making up t h e p o l i t i c a l u n i t s of P u e r t o Rico,
t h e American Virgin I s l a n d s , and t h e B r i t i s h V i r g i n I s l a n d s , sometimes
referred t o as
"Greater P u e r t o Rico'' (Thomas and Schwartz 1966)
This i n c l u d e s a l l l a n d masses between l a t i t u d e s 17O 40' N and 18O 45' N
and s t r e t c h i n g from t h e i s l a n d of Monito ( l o n g i t u d e 67O 57' W ) t o
Anegada ( l o n g i t u d e 64O 16' W ) .
There a r e 243 i s l a n d s and cays i n t h i s
a r e a ( n o t i n c l u d i n g those emergent ones p e r i o d i c a l l y washed over by
t i d e s and l a c k i n g t e r r e s t r i a l v e g e t a t i o n ) .
Of t h e s e we surveyed 141,
i n c l u d i n g a l l of t h e l a r g e r ones.
Figure 1 shows a map of t h e general
region and appendix I provides d e t a i l e d ones of s e l e c t e d a r e a s
c o n t a i n i n g t h e small cays which were s t u d i e d .
Where p o s s i b l e we
followed t h e o f f i c i a l names f o r i s l a n d s a s l i s t e d by t h e U.S. Department
When no o f f i c i a l name
of Commerce, Coast and Geodetic Survey (1962).
could be found, we used names s u p p l i e d by l o c a l fishermen.
In c a s e s of
nomenclatural d i s p u t e s among l o c a l a u t h o r i t i e s , we simply chose one of
the alternatives.
Sometimes we were unable t o o b t a i n any name a t a l l
and have named i s l a n d s o u r s e l v e s .
.
The c l i m a t e of t h e a r e a i s t r o p i c a l and s l i g h t l y modified by
a l t i t u d e on t h e l a r g e r i s l a n d s .
R a i n f a l l on P u e r t o Rico i s
g e o g r a p h i c a l l y v a r i a b l e , g r e a t e r i n the c e n t r a l mountains than on t h e
coastal plain.
I n a d d i t i o n , t h e r e i s a g e n e r a l east-west d i f f e r e n c e .
On t h e Luquillo Mountains a t t h e n o r t h e a s t e r n t i p of t h e i s l a n d ,
r a i n f a l l may exceed 5,080 mm (200 i n c h e s ) annually; the southwestern
lowlands r e c e i v e l e s s than 750 mm (30 i n c h e s ) annually ( P i c 6 1954 and
s e e Fig. 2 ) .
The p r i n c i p a l V i r g i n I s l a n d s a r e s i m i l a r i n r a i n f a l l t o t h e
c o a s t a l a r e a s o f P u e r t o Rico; s m a l l e r , o r more i s o l a t e d i s l a n d s
r e p r e s e n t e d by Culebra, Vieques, and Mona i n f i g u r e 2, a r e more x e r i c .
Mona i s d r i e r t h a n i t s r a i n f a l l would s u g g e s t , a s i t i s honeycombed
w i t h limestone caverns which r a p i d l y d r a i n o f f s u r f a c e water.
Temperatures a r e remarkably uniform s e a s o n a l l y and from y e a r t o
However, on P u e r t o Rico,
y e a r i n a given l o c a l i t y ( F i g s . 3, 4)
temperature follows e a s t-wes t and a l t i t u d i n a l t r e n d s .
Guineo
R e s e r v o i r ( e l e v a t i o n 1,000 m) h a s lower t e m p e r a t u r e s than Rio P i e d r a s
Rio P i e d r a s i s i n t u r n
( e l e v a t i o n 15 m) on t h e n o r t h e a s t e r n c o a s t .
c o o l e r t h a n Magueyes I s l a n d (10 m) which i s r e p r e s e n t a t i v e o f t h e
The i s o t h e r m f o r mean annual
southwestern c o a s t (Fig. 4)
t e m p e r a t u r e s o f 24O C s e p a r a t e s t h e c o a s t a l p l a i n and some v a l l e y s
(below 150 m e l e v a t i o n and where mean annual v a l u e s exceeding 26O may
occur) from t h e mountains where mean annual v a l u e s below 20° C a r e
encountered i n t h e h i g h e s t p a r t s ; t h e h i g h e s t temperature e v e r
recorded f o r t h e i s l a n d was 39.5O C , t h e l o w e s t 5O C ( P i c 6 1 9 5 4 ) .
The h o t t e r c o a s t a l a r e a s o f southwest P u e r t o Rico have temperatures
s i m i l a r t o t h o s f o f t h e p r i n c i p a l Virgin I s l a n d s (Fig. 4 ) .
.
.
Very few meteorological d a t a a r e a v a i l a b l e f o r s m a l l c a y s , though
it can be s a i d t h a t most a r e warm and dry.
Figure 3 p r e s e n t s d a t a f o r
Magueyes I s l a n d .
However, t h i s i s l a n d i s s o c l o s e t o P u e r t o Rico t h a t
it i s r e p r e s e n t a t i v e of c o n d i t i o n s o f c o a s t a l southwest P u e r t o Rico
r a t h e r t h a n t h a t o f more i s o l a t e d i s l a n d s .
Table 1 g i v e s t h e average
c o n d i t i o n s on Cayo Ahogado, a s m a l l , n e a r l y b a r r e n sandy cay a b o u t 1 km
e a s t o f P u e r t o Rico, d u r i n g s e v e r a l days i n March, 1966.
F i g u r e s 5-7
show t h e d e t a i l s of t h e d a i l y regimen f o r t h e two most extreme days,
March 10 which was b r i g h t and sunny, and March 19 which was h e a v i l y
overcast with occasional f i n e r a i n .
The h o t t e s t environment on t h e
cay was t h e s o i l s u r f a c e where midday temperatures exceeded 38O C. A t
1 cm and 5 cm below t h e s o i l s u r f a c e , temperatures decreased somewhat
A i r temperatures
b u t even t h e r e reached 30° C o r above on h o t days.
n e a r t h e ground reached 32O C on warm days, temperature d e c r e a s e d w i t h
h e i g h t above ground.
However, except f o r a few f l y i n g forms which
e x p e r i e n c e above-ground t e m p e r a t u r e s , most i n h a b i t a n t s o f s m a l l cays
a r e exposed t o t h e thermal c o n d i t i o n s n e a r t h e ground.
The low
v e g e t a t i o n on Cayo Ahogado a m e l i o r a t e s t e m p e r a t u r e s somewhat (Fig. 71,
although on h o t days a i r temperatures n e a r t h e s u r f a c e o f s m a l l clumps
S u r f a c e t e m p e r a t u r e s o f l e a v e s were
of p l a n t s r e a c h 32 - 33O C.
s e v e r a l d e g r e e s lower, presumably because o f t r a n s p i r a t i o n .
Temperature 1 cm down i n t h e s o i l was a s much a s 2 O C c o o l e r under
A t 5 cm depth t h e d i f f e r e n c e was a s much
v e g e t a t i o n t h a n i n t h e open.
a s 4O C.
Although t h e p l a n t s were s m a l l and s p a r s e l y d i s t r i b u t e d , and
hence d i d n o t g r e a t l y a f f e c t environmental t e m p e r a t u r e s , t h e few d e g r e e s
o f d i f f e r e n c e t h e y caused were i n a range t h a t may reduce t h e
i n s o l a t i o n d i r e c t l y r e a c h i n g small animals.
The e f f e c t o f
u n o b s t r u c t e d r a d i a n t energy can be seen from t h e h i g h b l a c k b u l b (TBB)
t e m p e r a t u r e s (Fig. 5 ) .
The n e a r l y b a r e , rocky cays probably have even more extreme
c o n d i t i o n s than Cayo Ahogado, more h e a v i l y v e g e t a t e d ones a r e probably
more moderate.
The p r e v a i l i n g winds a r e from the n o r t h e a s t and apparently have
been so a t l e a s t back i n t o t h e Pleistocene (Kaye 1959).
Marine
c u r r e n t s a r e e a s t t o west, although a northward c u r r e n t e x i s t s between
Mona and Puerto Rico, and t h e r e i s a northward d e f l e c t i o n between
Vieques and Puerto Rico (Kaye 1959).
Hurricanes u s u a l l y t r a v e l i n a general e a s t t o west d i r e c t i o n .
VEGETATION
The Puerto Rican and Virgin I s l a n d f l o r a has been most r e c e n t l y
summarized by B r i t t o n and Wilson (1923-30) and by L i t t l e and Wadsworth
(1964).
Ecological treatments of a broad n a t u r e have been presented
by Gleason and Cook (1927) , Cook and Gleason (1928) and Dansereau
(1966).
Only a few a r t i c l e s have d e a l t with t h e p l a n t s on the small
i s l a n d s (e.g. B r i t t o n 1924, 1933 on Hicacos and Culebra, Heatwole e t a l .
1963 on Cayo Santiago and Cayo Batata, and D'Arcy 1971, 1975 on
Our records a r e presented i n Appendix 11.
Anegada)
.
Analysis of the v e g e t a t i o n of t h e i s l a n d s t r e a t e d here would be a
major p r o j e c t i n i t s e l f and was n o t p o s s i b l e within t h e scope of t h e
present investigation.
However, broad vegetation types could be
recognized and were grouped on a physiognomic b a s i s i n t o 12 c a t e g o r i e s ,
most of which include more than one a s s o c i a t i o n , o r a t l e a s t v a r i e d
somewhat i n f l o r i s t i c composition from p l a c e t o place.
The
c l a s s i f i c a t i o n i s n o t intended t o be phytosociological, b u t r a t h e r a
u s e f u l d e s c r i p t i o n of t h e type and d i v e r s i t y of a v a i l a b l e s t r u c t u r a l
h a b i t a t s f o r animals.
The s p a t i a l r e l a t i o n s h i p s of t h e c a t e g o r i e s t o
one another and t o s e v e r a l major environmental parameters a r e
c l a r i f i e d i n Figs. 8 and 9.
The c a t e g o r i e s a r e as follows:
1. Lack of Vegetation.
Extensive a r e a s of bare rock o r sand,
u s u a l l y c o a s t a l , though occurring l o c a l l y f u r t h e r inland.
2.
Succulent Beach Vegetation.
Often n e a r l y pure stands of low,
s u c c u l e n t , halophytic herbs.
Sesuvium portulacastrum and P h i l o x e r i s
vermicularis, u s u a l l y under 30 cm i n h e i g h t , occur alone o r i n
combination on r e l a t i v e l y undisturbed g r a v e l l y or sandy beaches1.
These two s p e c i e s p l u s B a t i s maritima and S a l i c o r n i a p e r e n n i s form
v a r i o u s combinations a f t e r t h e elimination of mangrove (81, o r i n
otherwise i n t e r m i t t e n t l y - f l o o d e d , shallow, s a l i n e c o a s t a l depressions.
B a t i s may reach 70 cm i n height.
3. Mixed Beach Vegetation.
A v a r i a b l e mixture of herbs (Cakile
l a n c e o l a t a ) , small s u f f r u t e s c e n t s ( ~ u p h o r b i ab u x i f o l i a ) , rhizomaceous
Nomenclature follows B r i t t o n and Wilson (1923-30)
.
grasses ( S p o r o b d u s
v i r g i n i c u s , S p a r t i n a p a t e n s ) and c r e e p i n g v i n e s
(Ipomoea p e s - c a p r a e , C a n a v a l i a m a r i t i m a , V i g n a m a r i n a ) , sometimes w i t h
Sesuvium portulacastrum.
Vegetation u s u a l l y l e s s than 1 m t a l l , with
low c o v e r .
On s a n d , g r a v e l , o r c r e v i c e s i n c o a s t a l s h e l f l i m e s t o n e ,
f a i r l y r e s i s t a n t t o d i s t u r b a n c e , and r e c o v e r s r a p i d l y .
4.
Beach Shrubs.
Small, s c a t t e r e d shrubs intermixed with h e r b s
and g r a s s e s , o c c a s i o n a l l y c o n t a i n i n g a h i g h p r o p o r t i o n o f v i n e s which
form a n e a r l y c o n t i n u o u s mat.
H e i g h t u s u a l l y 1-2 m, found i n l a n d from
t h e p r e v i o u s type.
Common s p e c i e s a r e S u r i a n a mari t i m a , T o u r n e f o r t i a
g n a p h a l o d e s , and T h e s p e s i a p o p u l n e a , w i t h t h e s p e c i e s c h a r a c t e r i s t i c
Fairly resistant to
o f t h e p r e v i o u s t y p e ( 3 ) a s ground cover.
disturbance
.
5 . Graminoid.
S t a n d s havTng t h e graminoid l i f e form dominant,
whether composed o f t r u e g r a s s e s o r o f sedges.
Usually a l s o contains
a number o f f o r b s .
U s u a l l y 1 m o r l e s s h i g h , though t a l l e r s t a n d s a r e
sometimes found.
Found i n t h e i n t e r i o r o f i n t e r m e d i a t e - s i z e d and
Always a d i s c l i m a x o r s u c c e s s i o n a l s t a g e i n t h e
larger islands.
c l i m a t i c zone s t u d i e d ( R i c h a r d s 19571, m a i n t a i n e d by g r a z i n g , f i r e , o r
p e r i o d i c c u t t i n g o r mowing.
Various graminoid s p e c i e s c o v e r t h e f u l l
r a n g e o f m o i s t u r e and e d a p h i c c o n d i t i o n s found i n t h e s t u d y r e g i o n .
6 . Scrub.
Shrubs of v a r i o u s t y p e s , c a c t u s , and v i n e s , f r e q u e n t l y
i n t e r m i x e d w i t h h e r b s and g r a s s e s .
V a r i e s from l e s s t h a n 50% t o 100%
c o v e r , e a s i l y p e n e t r a b l e t o i m p e n e t r a b l e , and l e s s t h a n 1 m t o o v e r 2 m,
occasionally higher.
F r e q u e n t dominants a r e Croton a s t r o i t e s , C .
r i g i d u s ( b o t h r e p l a c e d by C . d i s c o l o r i n Anegada) , Lantana i n v o l u c r a t a ,
An t h a c a n t h u s s p i n o s u s , and P i t h e c e l l o b i u m u n g u i s c a t i
A disclimax
m a i n t a i n e d by o v e r g r a z i n g and overbrowsing, p r i m a r i l y by f e r a l g o a t s ,
i n t h e i n t e r i o r of i n t e r m e d i a t e - s i z e d , d r i e r i s l a n d s and on t h e leeward
(rain-shadow) s i d e s o f t h e l a r g e s t i s l a n d s .
Shrub s t a g e s i n t h e zone
where m o i s t f o r e s t (10) i s climax a r e s u c c e s s i o n a l and c o n s i s t c f more
mesomorphic s p e c i e s , o f t e n s e e d l i n g s and s a p l i n g s of l a r g e r , s e c o n d a r y
forest trees.
.
7.
C o a s t a l Groves.
S t a n d s o f s m a l l t r e e s o r s h r u b s , i n l a n d from
( 4 ) , r a n g i n g from n e a r l y p r o s t r a t e and l e s s t h a n 1 m i n h e i g h t on
windswept c o a s t s and d u n e s , t o 4 m o r more i n p r o t e c t e d s i t e s .
U s u a l l y p r o v i d e dense shade w i t h a heavy accumulation o f l e a f l i t t e r .
F r e q u e n t l y p u r e s t a n d s of Coccoloba u v i f e r a o r Hippomane m a n c i n e l l a , o r
m i x t u r e s of s e v e r a l t o many s p e c i e s ; E l a e o d e n d r o n x y l o c a r p u m ,
~ r i t h a l i sf r u t i c o s a , Conocarpus e r e c t a , and t h e t r e e - c a c t u s O p u n t i a
r u b e s c e n s a r e prominent.
8. Mangroves.
S t a n d s o f mangrove r a n g i n g from s c a t t e r e d
s a p l i n g s a b o u t 1 m i n h e i g h t t o f o r e s t s o v e r 10 m, p r o v i d i n g r a t h e r
deep s h a d e , i n permanently- o r i n t e r m i t t e n t l y - f l o o d e d d e p r e s s i o n s
behind c o a s t s , and a l o n g c o a s t s and on o l d r e e f s e t c . which a r e
R h i zophora m a n g l e , A v i c e n n i a n i t i d a ,
s h e l t e r e d from wave a c t i o n .
L a g u n c u l a r i a racemosa and Conocarpus e r e c t a , e i t h e r a s s i n g l e - s p e c i e s
s t a n d s o r i n v a r i o u s combinations.
9. Xeric Forest.
Deciduous, open f o r e s t , frequently with sparse
understory of cactus, thorny shrubs, vines, and annual grasses.
I n t e r i o r of intermediate-sized i s l a n d s and
Height usually over 8 m.
le~eward (rain-shadow) s i d e s of l a r g e s t i s l a n d s .
May be absent on the
windward (humid) s i d e s of t h e l a t t e r , where mesic f o r e s t (10) l i e s
adjacent t o c o a s t a l formations, except where l o c a l edaphic-topographic
conditions permit x e r i c f o r e s t .
Probably includes Dansereau' s (1966)
category of Semi-deciduous Forest and L i t t l e and Wadsworth's (1964) Dry
Coastal and Dry Limestone f o r e s t s .
Dominant t r e e s v a r i a b l e from
place t o p l a c e ; P i s o n i a s u b c o r d a t a , T o r r u b i a f r a g r a n s , B u r s e r a s i m a r o u b a ,
C i t h a r e x y l u m f r u t i c o s u m , T a b e b u i a h e t e r o p h y l l a , and the g i a n t cactus
C e p h a l o c e r e u s r o y e n i a r e representative.
10. Mesic Forest.
Includes a v a r i e t y of moister, a l t i t u d i n a l l y zoned types including those c a l l e d Moist-coastal, Moist Limestone,
Lower and Upper C o r d i l l e r a , and Lower and Upper Luquillo Forest by
L i t t l e and Wadsworth (19641, and Lowland Rain Forest, Lower Montane
Rain F o r e s t , Montane Forest and Montane Scrub by Dansereau (1966).
Physiognomy and f l o r i s t i c s are v a r i a b l e (see Beard 1944, Richards 1957).
Such a v a r i e t y i s included under one heading since i n the region
s t u d i e d , t h e s e f o r e s t s occur only on t h e l a r g e s t i s l a n d s , some types
only on Puerto Rico.
Finer d i s t i n c t i o n s would be unnecessary f o r our
purposes
.
11. Coconut Groves.
Usually planted by man i n f l a t , sandy
c o a s t a l areas.
Light shade, understory usually composed of species
of t h e mixed beach vegetation (3) and v a r i a b l e .
12.
Other.
Any vegetation not f i t t i n g any of t h e above
c a t e g o r i e s , e.g. crops, r u d e r a l s , dooryards, o r vegetation of o t h e r
highly modified areas.
During t h e survey of a p a r t i c u l a r i s l a n d , the vegetation types
were recorded, and where f e a s i b l e , t h e percentage of t h e i s l a n d
covered by each one estimated and a rough sketch-map drawn.
When
only a few individual p l a n t s were p r e s e n t on an i s l a n d , complete
censuses were made.
In some i n s t a n c e s , vegetational reconnaissances
were made.
By way of summary it can
The r e s u l t s a r e presented i n Table 2 .
be s t a t e d t h a t Puerto Rico contains a l l the vegetation types o u t l i n e d
(1 - 2
Culebra and Vieques have x e r i c t o mesic f o r e s t s (9, 1 0 ) ;
Mona, Caja de Muertos and Desecheo support x e r i c f o r e s t , a l l f i v e have
Monito has predominantly
a v a r i e t y of the simplest types (1 - 8 ) .
scrub ( 6 ) .
The small cays near t h e southern and southwestern coast
of Puerto Rico a r e mostly mangrove i s l a n d s ( 8 ) ; those f u r t h e r offshore
tend t o be sandy and nearly bare (1) o r w i t h succulents ( 2 ) mixed beach
vegetation (31, o r beach shrubs ( 4 ) .
The group of cays extending toward Culebra from northeastern
Puerto Rico a r e mostly steep-sided, limestone o r cemented sand dunes,
although a few low sandy i s l a n d s a l s o occur.
Most support scrub (61,
/
-
succulent o r mixed beach vegetation ( 2 , 3 ) , although the l a r g e r ones
have shrubs (4) and occasionally t r e e s .
The cays south of t h i s chain
( e a s t of Puerto Rico) and around Culebra I s l a n d a r e v a r i a b l e , most being
e i t h e r rocky with a beach, o r sandy.
The l a r g e r ones support x e r i c
f o r e s t (91, the s m a l l e r ones only succulent o r mixed beach vegetation
( 2 , 3 ) o r scrub ( 6 ) .
The Virgin I s l a n d s , S t . John, Tortola and S t . Croix have mesic
f o r e s t p l u s a l l o t h e r c a t e g o r i e s (1 - 1 2 ) .
The o t h e r p r i n c i p a l
i s l a n d s except Anegada, a s w e l l a s a few somewhat smaller ones such a s
Guana, Greater Caminoe, P e t e r , Norman, and Greater Thatch have x e r i c
forest.
The remaining cays and small i s l a n d s p l u s Anegada a r e
variable.
However, few support vegetation more lush than scrub (61,
o r a t most l i m i t e d a r e a s of x e r i c f o r e s t (9)
The a v a i l a b l e climate
and s o i l d a t a , corroborated by conversation with o l d e r i n h a b i t a n t s ,
i n d i c a t e t h a t a t one time f o r e s t was more extensive on both t h e l a r g e r
and t h e s m a l l e r i s l a n d s .
Apparently it has been decimated by
charcoal burners, c u l t i v a t o r s , and f e r a l animals, a widespread
phenomenon throughout t h e Caribbean a s indeed i n the t r o p i c s
generally.
.
GEOLOGY
Rosen (1975) has reviewed t h e previous h i s t o r y of the Caribbean.
Although t h e r e i s n o t unanimity among a u t h o r i t i e s on t h e t o p i c , he f e l t
t h a t t h e most reasonable s y n t h e s i s of t h e geologic and biogeographic
I n t h e l a t e Mesozoic when
d a t a was t h e following sequence of events.
South America was s e p a r a t i n g from A f r i c a , a subduction zone consuming
t h e e a s t e r n P a c i f i c s e a f l o o r bordered t h e western c o a s t of both North
and South America and t h e a r e a between them, the l a t t e r c o n s i s t i n g of a
v o l c a n i c archipelago, t h e Proto-Antilles.
Subsequently, s e p a r a t e
North and South American p l a t e s arose by t h e formation of two decoupling
f a u l t s on e i t h e r s i d e of t h e Proto-Antilles.
With the continued
r e l a t i v e westward d r i f t of t h e North American and South American
p l a t e s , t h e Proto-Antillean region became displaced eastward towards
i t s p r e s e n t r e l a t i v e p o s i t i o n ; a new f a u l t zone occurred which
s e p a r a t e d t h e Proto-Antilles i n t o two subregions which subsequently
became t h e Greater A n t i l l e s and Lesser A n t i l l e s .
With t h e continued
westward movement of t h e major American c o n t i n e n t s , f u r t h e r f a u l t i n g
occurred breaking up the Central American region i n t o an archipelago
which subsequently joined t o form the p r e s e n t day isthmus; the
Caribbean i s l a n d s aligned themselves i n t o t h e i r p r e s e n t r e l a t i v e
positions.
I f t h i s model i s c o r r e c t , the Greater A n t i l l e s have been an
archipelago f o r a long time, contrary t o previous opinion t h a t t h e
major A n t i l l e a n i s l a n d s were broadly connected (Schuchert 1935).
Connections between a d j a c e n t banks were r a r e , t h e Puerto Rican one
never having been connected t o the n e a r e s t one t o t h e e a s t ( S t . Maarten)
I f it was ever connected t o Hispaniola, it l o s t t h e
( B u t t e r l i n 1956).
connection i n the Pliocene o r e a r l i e r itchel ell 1954)
.
The P u e r t o Rican Bank ( P u e r t o Rico., i t s o u t l y i n g i s l a n d s , t h e
American and B r i t i s h Virgin I s l a n d s ) r e s u l t e d from v u l c a n i z a t i o n i n t h e
Cretaceous (Meyerhoff 1933) and became emergent d u r i n g o r o g e n i c
movements i n t h e l a t e Eocene ( B u t t e r l i n 1956).
In c o n t r a s t t o t h e
p a u c i t y of connections between banks, connections of i s l a n d s w i t h i n t h e
P u e r t o Rican Bank have been e x t e n s i v e and r e c e n t .
Heatwole and
MacKenzie (1967) have shown t h a t t h e p r i n c i p a l i s l a n d s of t h e Virgin
group ( e x c e p t S t . Croix) l o s t t h e i r connection with each o t h e r and w i t h
P u e r t o Rico only about 8,000-10,000 y e a r s ago, due t o e u s t a t i c r i s e i n
sea level.
Culebra l o s t i t s connection w i t h P u e r t o Rico and t h e
V i r g i n s a t t h e same time, although Vieques, Caja de Muertos, and many
s m a l l c a y s remained connected t o P u e r t o Rico u n t i l about 6,000 y e a r s
ago.
The s m a l l e r i s l a n d s around t h e p r i n c i p a l V i r g i n s were
s e p a r a t e d from t h e l a t t e r a t v a r i o u s times b e f o r e 8,000-10,000 B.P.
By c o n t r a s t , Mona, Monito, Desecheo and S t . Croix h a w been
i s o l a t e d much l o n g e r , n o t having been connected t o any o t h e r i s l a n d s
s i n c e a t l e a s t t h e P l i o c e n e , i f then.
A number of i s l a n d s were p r e s e n t a t v a r i o u s times which have s i n c e
submerged.
ISLAND AND COASTLINE FORMS
The l a r g e r i s l a n d s a r e o f v o l c a n i c o r i g i n and c o n s i s t mainly of
igneous and metamorphic rock w i t h q u i t e s t e e p topography.
P u e r t o Rico
h a s , i n a d d i t i o n , an e x t e n s i v e n o r t h e r n and a s m a l l e r s o u t h e r n s t r i p of
cenozoic limestone which h a s been e x t e n s i v e l y eroded, g i v i n g r i s e t o
c h a r a c t e r i s t i c low, blocky, v e r t i c a l - s i d e d h i l l s w i t h many caves.
Mona, Monito and t h e h i g h e r i s l a n d s to t h e n o r t h e a s t of P u e r t o Rico
a r e of s i m i l a r o r i g i n , w h i l e Anegada i s a low, f l a t limestone s h e l f ,
c o n t r a s t i n g markedly w i t h a l l t h e o t h e r i s l a n d s of t h e P u e r t o Rican
bank save f o r a few s m a l l cays.
Form of t h e s m a l l e r i s l a n d s v a r i e s c o n s i d e r a b l y , and ranges from
low, sandy cays w i t h g e n t l y s l o p i n g beaches t o mushroom-shaped ones
surrounded by v e r t i c a l c l i f f s u n d e r c u t a t t h e base by wave a c t i o n .
Many i s l a n d s have a g r e a t p r o p o r t i o n of t h e c o a s t l i n e made up o f rock
which i s n e i t h e r v e r t i c a l c l i f f , nor beach.
We termed t h i s
A f i n a l c o a s t l i n e t y p e c o n s i s t e d of mangrove
"intermediate type".
swamp.
Most l a r g e i s l a n d s have s e v e r a l c o a s t l i n e t y p e s , i n c l u d i n g
beach, u n d e r c u t c l i f f s , and i n t e r m e d i a t e type.
METHOD OF SURVEY
Each i s l a n d surveyed was d e s c r i b e d a s t o i t s v e g e t a t i o n ( s e e above),
g e o l o g i c a l t y p e , s o i l , p r e s e n c e o r absence o f s t a n d i n g w a t e r , maximum
h e i g h t , a r e a , n a t u r e of s h o r e l i n e , and p r o p o r t i o n of each type of c o a s t ,
p r o p o r t i o n surrounded by b a r r i e r r e e f , d i s t a n c e t o n e a r e s t o t h e r landmass on t h e P u e r t o Rican s h e l f i n each of 4 q u a d r a n t s , degree of
d i s t u r b a n c e by humans and time of i s o l a t i o n from o t h e r l a n d masses.
P r o p o r t i o n s of v a r i o u s s h o r e l i n e t y p e s were o b t a i n e d by measuring
t h e p e r i m e t e r on a map o r a e r i a l photograph w i t h a Radix map measurer,
keeping v a l u e s f o r each c o a s t l i n e type s e p a r a t e i n accordance w i t h
o b s e r v a t i o n s made while on t h e i s l a n d .
P e r c e n t a g e s of each type
were then c a l c u l a t e d .
I s l a n d a r e a s were o b t a i n e d e i t h e r from t h e
l i t e r a t u r e , from maps u s i n g a p l a n i m e t e r , o r by d i r e c t measurement.
Maximum h e i g h t was o b t a i n e d from topographic maps, o r i n t h e case of
s m a l l cays by d i r e c t e s t i m a t i o n o r measurement.
The o t h e r information
was o b t a i n e d by d i r e c t o b s e r v a t i o n supplemented by maps, a e r i a l
photographs, n a u t i c a l c h a r t s , c o n v e r s a t i c n s with i n h a b i t a n t s o f v a r i o u s
i s l a n d s , and t h e l i t e r a t u r e .
On e v e r y i s l a n d v i s i t e d , each of t h e above-defined v e g e t a t i o n
c a t e g o r i e s which was p r e s e n t was searched f o r t h e f o l l o w i n g group of
animals; t e r r e s t r i a l i s o p o d s , c e n t i p e d e s , m i l l i p e d e s , pseudoscorpions,
amblypigids (= p h r y n i c h i d s o r t a i l l e s s whip s c o r p i o n s ) , s c o r p i o n s ,
s p i d e r s , a n t s , Drosophila, l a n d s n a i l s , amphibians and r e p t i l e s ; o t h e r
groups provided supplementary information on c e r t a i n a s p e c t s .
We were
o p p o r t u n i s t i c i n t h a t e a s e of sampling and t h e p r o b a b i l i t y of o b t a i n i n g
adequate samples i n a r e l a t i v e l y s h o r t time was a c o n s i d e r a t i o n i n o u r
choice of groups.
However, t h e animal groups we used were chosen t o
i n c l u d e b o t h e u r y t o p i c and s t e n o t o p i c forms, t h o s e with weak and t h o s e
w i t h s t r o n g d i s p e r s a l powers, and groups w i t h a wide range of s u s p e c t e d
evolutionary r a t e s .
S p e c i a l i s t s i n v a r i o u s groups i d e n t i f i e d ( o r a r e
i n t h e p r o c e s s of i d e n t i f y i n g ) o u r m a t e r i a l and advised on t e c h n i q u e s
which would provide a s complete a r e p r e s e n t a t i o n of an i s l a n d ' s fauna
as possible.
These methods i n c l u d e d use of (1) T u l l g r e n f u n n e l
e x t r a c t i o n o f l i t t e r samples t r a n s p o r t e d t o t h e l a b o r a t o r y i n p l a s t i c
b a g s , ( 2 ) sweeping v e g e t a t i o n with an i n s e c t n e t , ( 3 ) b e a t i n g
v e g e t a t i o n over a white s h e e t , ( 4 ) examining l i t t e r on a white s h e e t ,
( 5 ) s e t t i n g o u t b a i t (mixed f r u i t s f o r Drosophila, s u g a r and t u n a f o r
a n t s , e t c . ) , ( 6 ) s h o o t i n g o r noosing ( r e p t i l e s ) , ( 7 ) c a r e f u l s e a r c h of
t h e s u r f a c e of t h e ground and v e g e t a t i o n and under rocks and o t h e r
o b j e c t s , and ( 8 ) b r e a k i n g open t w i g s , o l d t e r m i t e n e s t s , l o g s , and
other debris.
A l l d a t a were s t o r e d on computer c a r d s .
S t a t i s t i c a l treatment is
d i s c u s s e d l a t e r i n a p p r o p r i a t e p a p e r s , a s a r e o t h e r s p e c i a l methods used
f o r i n d i v i d u a l p a r t s of t h e p r o j e c t .
Biogeographic and taxonomic
t r e a t m e n t s w i l l be p u b l i s h e d by t h e v a r i o u s s p e c i a l i s t s t h a t have
received material.
The appendices l i s t t h e r e s u l t s f o r some groups
f o r which a number o f i d e n t i f i c a t i o n s a r e now a v a i l a b l e .
SPECIAL NOTES
The i s l a n d
I t i s a mass of
I t i s about 100
nearly v e r t i c a l
o f S a i l Rock (Appendix I ) r e q u i r e s s p e c i a l mention.
rock r i s i n g 38 m (125 f t ) p r e c i p i t o u s l y o u t of t h e s e a .
m i n diameter.
The c l i f f s around it a r e v e r t i c a l o r
I t i s unusual i n t h a t
and a r e undercut a t t h e base.
it completely lacks t e r r e s t r i a l vascular p l a n t s and hence no l o c a l
autotrophic energy base f o r t h e s u r p r i s i n g l y l a r g e number of
t e r r e s t r i a l animals.
Indeed, the only t e r r e s t r i a l p l a n t s observed
a t a l l was a green a l g a i n one crevice on the north s i d e .
A few
i n d i v i d u a l s of a marine p l a n t s , Sargassum, c o l l e c t e d by sea b i r d s o r
m a t e r i a l was p r e s e n t b u t d i d not appear t o have any t e r r e s t r i a l animals
a s s o c i a t e d with it o r feeding upon it.
W
e v i s i t e d t h e i s l a n d i n May 1966, s c a l e d t h e c l i f f s and made an
i n t e n s i v e search f o r organisms on t h e top.
Numerous cracks and c r e v i c e s occur i n t h e rock on t o p , and on the
few f l a t , o r n e a r l y f l a t a r e a s , t h e r e a r e some loose boulders and
stones.
From these p l a c e s we c o l l e c t e d s p e c i e s of t e r r e s t r i a l
v e r t e b r a t e s and one v e r t e b r a t e ( a gekkonid l i z a r d , Sphaerodactylus
macrolepis macrolepis); many of t h e species were q u i t e abundant.
By
It
f a r t h e most abundant arthropod p r e s e n t was a t e r r e s t r i a l isopod.
occurred i n s u r p r i s i n g l y dense populations and probably served a s the
b a s i s f o r much of t h e r e s t of t h e t e r r e s t r i a l food chain which
consisted l a r g e l y of p r e d a t o r s , t h e r e was 1 s p e c i e s each of centipede,
t a i l l e s s whip scorpion (amblypigid = phrynichid) , scorpion and s p i d e r .
There was i n addition one s p e c i e s of a n t , s e v e r a l o t h e r s p e c i e s of
i n s e c t s , and a land crab.
Unfortunately none of t h e i n v e r t e b r a t e s
were i d e n t i f i e d t o species as our c o l l e c t i o n was l o s t i n t h e mail when
s e n t t o s p e c i a l i s t s f o r i d e n t i f i c a t i o n and we have n o t had opportunity
t o v i s i t t h e i s l a n d again.
However, s u f f i c i e n t i s known about the
genuine feeding h a b i t s of some of t h e groups t o comment on t h e
probable t r o p h i c r e l a t i o n s .
The t o p of the i s l a n d was used e x t e n s i v e l y by s e a b i r d s f o r
n e s t i n g and t h e i r excrement l i t e r a l l y covers t h e i s l a n d , giving it a
white-washed appearance.
These b i r d s feed on f i s h and hence it i s
probable t h a t they serve a s t r a n s f e r organisms (Heatwole 1975)
obtaining energy from t h e marine community and depositing a food
source f o r t h e t e r r e s t r i a l community on t h e i s l a n d i n the form of
guano and on occasions a s c a r r i o n (dead a d u l t s o r chicks) o r broken
eggs.
The guano i s believed t o be t h e most important source a s i s
c a r r i o n was observed and t h e only species l i k e l y t o be scavengers t h a t
were p r e s e n t was the land crab and possibly t h e a n t .
The undercut
nature of t h e i s l a n d and i t s h e i g h t would make work up of marine
carrion unlikely.
Isopods generally feed on d e t r i t u s o r o t h e r f i n e l y
divided organic matter and we suspect t h a t it fed on guano and i n t u r n
was preyed upon by the predatory i n v e r t e b r a t e s .
The l a r g e r p r e d a t o r s
such a s scorpion, whip scorpion, and l i z a r d s probably e a t isopods and
some of t h e i n v e r t e b r a t e s t h a t prey on isopods.
None of the l i z a r d s
we c o l l e c t e d had prey i n t h e i r stomachs.
However, we d i d obtain t h r e e
recognizable items from the s e c t i o n of one animal.
There were
remains of an a n t , the head of an i n s e c t l a r v a , and s u r p r i s i n g l y , p a r t s
of a species not otherwise known from t h e i s l a n d , a homopteran.
The
l a s t i s a p l a n t feeder and must r e p r e s e n t a waif a r r i v i n g a e r i a l l y
( t h e n e a r e s t vegetated land mass, Culebra I s l a n d i s 14 km away).
In summary, t h e probable t r o p h i c r e l a t i o n s a r e (1) a t r a n s f e r of
marine-derived energy v i a s e a b i r d s i n the form of guano and c a r r i o n ,
( 2 ) i t s u t i l i z a t i o n by isopods which ( 3 ) serve a s prey f o r a v a r i e t y
of p r e d a t o r s , ( 4 ) leading t o secondary p r e d a t o r s .
The endogenous
prey i s probably supplemented by ( 5 ) a e r i a l waifs.
Complete
dependence of an i n s u l a r t e r r e s t r i a l community upon exogenous, marinederived energy, although unusual, i s n o t unique. Heatwole (1975) r e p o r t s
communities of s e v e r a l t e r r e s t r i a l s p e c i e s ( i n c l u d i n g p r e d a t o r s ) on
v e g e t a t i o n - f r e e sand cays i n t h e Coral Sea.
The source of t h e t e r r e s t r i a l fauna on S a i l Rock i s problematic.
S a i l Rock became i s o l a t e d from t h e once continuous land mass of
Greater Puerto Rico (about 8,000 years B.P.) (Heatwole and MacKenzie
1 9 6 7 ) , and it seems u n l i k e l y t h a t t h e r e s i d e n t s p e c i e s could have
maintained themselves on such a small barren i s l a n d .
On the whole
t h e undercut nature of t h e i s l a n d ' s base would suggest t h a t a t p r e s e n t
flotsam t r a n s p o r t would be u n l i k e l y and t h e f l i g h t l e s s n a t u r e of many
o f t h e r e s i d e n t species would m i l i t a t e a g a i n s t a e r i a l t r a n s p o r t .
The
d i f f i c u l t y of access t o the i s l a n d would make human t r a n s p o r t u n l i k e l y ,
although t h e r e i s an automatic l i g h t house on top which i s p e r i o d i c a l l y
serviced.
I n t r o d u c t i o n of s o many s p e c i e s by t h i s means, however,
seems u n l i k e l y .
From t h e above, i t i s c l e a r t h a t S a i l Rock i s e s p e c i a l l y
i n t e r e s t i n g from t h e s t a n d p o i n t s of i s l a n d biogeography and community
ecology.
I t would warrant q u a n t i t a t i v e study by someone i n a p o s i t i o n
t o c a r r y o u t a long-term observation t h e r e .
DISCUSSION
The b r i e f d e s c r i p t i o n of t h e general ecology of t h e archipelago
presented above provides a summary of l o c a l conditions during t h e
8-year period i n which d a t a were c o l l e c t e d .
Many e c o l o g i c a l
c h a r a c t e r i s t i c s a r e , however, i n t h e process of r a p i d change, l a r g e l y
through human i n f l u e n c e .
An assessment of t h e n a t u r e of such changes
i s r e q u i s i t e f o r p l a c i n g the p r e s e n t study i n p e r s p e c t i v e and f o r
e v a l u a t i o n of f u t u r e d a t a .
I n pre-Columbian times a succession of c u l t u r e s occurred i n the
a r e a , e s p e c i a l l y on t h e l a r g e r i s l a n d s .
However, even those a s small
a s Culebra (about 1 4 x 6 kms) had a t l e a s t semi-resident populations,
a midden containing v a r i o u s a r t i f a c t s was found on t h i s i s l a n d during
one of our f i e l d t r i p s .
Some s p e c i e s , such a s u s e f u l p l a n t s and t h e
e d i b l e iguana, may p o s s i b l y owe t h e i r d i s t r i b u t i o n i n p a r t t o
t r a n s p o r t by these e a r l y r e s i d e n t s .
I n post-Columbian times, man's
e f f e c t was a c c e l e r a t e d through a d d i t i o n a l c l e a r i n g of land and
establishment of an European-type c u l t u r e .
Domestic animals and a
v a r i e t y of ornamental and e d i b l e p l a n t s were purposefully introduced
a s well a s various weeds, rodents, i n s e c t s and o t h e r i n v o l u n t a r i l y t r a n s p o r t e d species.
F e r a l c a t s and e s p e c i a l l y r a t s have p o s s i b l y
had an adverse e f f e c t on the now r a r e and endangered Puerto Rican
p a r r o t (Amazona v. v i t t a t a ) through depredation of n e s t s (RodriguezVidal 1 9 5 9 ) .
The mongoose was introduced i n o r d e r t o c o n t r o l r o d e n t s
(unsuccessfully).
The r o l e of t h e mongoose i n the d e c l i n e and
e x t i n c t i o n of West Indian n a t i v e fauna may not be a s g r e a t a s i s
sometimes a t t r i b u t e d t o it, b u t it i s almost c e r t a i n l y r e s p o n s i b l e f o r
l o c a l e x t i n c t i o n of some l i z a r d s , snakes and ground-nesting b i r d s (see
Barbour 1930, Heatwole and Torres 1967, P h i l i b o s i a n and Ruibal 1 9 7 1 ) .
The cane t o a d (Bufo marinus) was brought t o Puerto Rico t o c o n t r o l
i n s e c t p e s t s of sugar cane i n 1920 ( B a r t l e t t 1949) and i s now abundant
on v a r i o u s i s l a n d s .
Cayo Santiago h a s had a f e r a l band of rhesus
monkeys f o r a number of y e a r s .
Recently, i n t r o d u c t i o n of some of them
i n t o s e v e r a l a d d i t i o n a l i s l a n d s was made.
These p o p u l a t i o n s w i l l
undoubtedly have an e c o l o g i c a l e f f e c t .
Goats and sheep a r e perhaps t h e most d e s t r u c t i v e of any introduced
s p e c i e s , and t h e r e a r e probably few i s l a n d s i n t h e a r c h i p e l a g o which
have n o t been d i s t u r b e d t o some e x t e n t by t h e s e animals a t some time o r
other.
The e a r l y custom of r e l e a s i n g g o a t s and o t h e r domestic
ungulates on small i s l a n d s and cays f o r l a t e r use by mariners was
a p p a r e n t l y common (Davis 1 9 7 1 ) .
Mona and Desecheo I s l a n d s s t i l l
maintain l o c a l p o p u l a t i o n s o r i g i n a t i n g i n t h i s way; the former i s l a n d
s e r v e s a s a r e s e r v e f o r sportsmen t o hunt g o a t s and f e r a l p i g s .
Many
of t h e cays i n t h e V i r g i n I s l a n d s have g o a t h e r d s which belong t o
r e s i d e n t s from l a r g e r i s l a n d s and from which i n d i v i d u a l s a r e
p e r i o d i c a l l y k i l l e d f o r meat.
Perhaps t h e i s l a n d most a f f e c t e d by
f e r a l animals i s S a l t I s l a n d , B . V . I . ,
i n which t h e g r e a t e r p a r t of t h e
v e g e t a t i o n has been reduced t o a low, open s t r a n d of Croton r i g i d u s and
C. a s t r o i t e s , s p e c i e s u n p a l a t a b l e even t o goats.
Except f o r creeping
Opuntia r e p e n s , t h e spaces between Croton bushes a r e n e a r l y devoid of
o t h e r v e g e t a t i o n , and g o a t s have been observed gleaning the t i n y
g r a s s e s and annuals which do appear.
On S a l t I s l a n d a t l e a s t , free-roaming sheep and c a t t l e a l s o played
a r o l e i n decimating once-considerable g r a s s y a r e a s ( M r s . B e a t r i c e
Smith, p e r s o n a l communication).
These animals d i e d o u t , s i n c e they
cannot s u b s i s t on a d i e t of woody d i c o t s a s can g o a t s .
Observations
on e x c l o s u r e s over a three-year p e r i o d (Byer, i n p r e p a r a t i o n ) s u g g e s t
t h a t once ground cover i s e l i m i n a t e d , re-establishment of woody p l a n t s
i s r e t a r d e d even i n t h e absence of goats.
P o s s i b l y i n c r e a s e d run-off
e i t h e r washes seeds away d i r e c t l y o r removes o r g a n i c matter necessary
f o r t h e i r germination.
The degree of d i s t u r b a n c e t o a small i s l a n d seems t o be r e l a t e d t o
i t s geomorphologic type.
Monito, which h a s s t e e p s i d e s undercut a t
the b a s e , i s r e l a t i v e l y i n a c c e s s i b l e and h a s probably been v i s i t e d by
humans only a few times (Rolle e t a l . 1964).
The same i s t r u e of a
number of t h e small i s l a n d s i n t h e chain j u s t n o r t h e a s t of P u e r t o Rico.
The l a t t e r , however, were used f o r s t r a f i n g p r a c t i c e d u r i n g World War
I1 and remnants of s h e l l s can s t i l l be found embedded i n rock on t h e
t o p of some.
Others of t h e r e l a t i v e l y i n a c c e s s i b l e s t e e p - s i d e d
i s l a n d s ( e . g. S a i l Rock) c o n s t i t u t e shipping dangers and have l i g h t -
houses o r o t h e r n a v i g a t i o n a l a i d s which must be p e r i o d i c a l l y s e r v i c e d .
Many of t h e small a c c e s s i b l e i s l a n d s a r e f r e q u e n t l y v i s i t e d f o r
s h o r t p e r i o d s by fishermen who camp on t h e beach b u t o t h e r w i s e have a
minimal e f f e c t .
Some i s l a n d s c o n t a i n i n g l a r g e c o l o n i e s of n e s t i n g s e a
b i r d s a r e p e r i o d i c a l l y v i s i t e d by egg g a t h e r e r s (e.g. Frenchcap Cay).
A number of t h e more a c c e s s i b l e i s l a n d s have people l i v i n g on t h e m ,
even some a s small a s Ramos I s l a n d ( < % k m i n g r e a t e s t d i a m e t e r ) .
Lobos I s l a n d ( s l i g h t l y more t h a n % km i n g r e a t e s t d i a m e t e r ) h a s a
t o u r i s t h o t e l , and I s l e t a Marine s e r v e s a s a y a c h t b a s i n .
Most
i s l a n d s with some beach and l a r g e r than 1 km i n g r e a t e s t d i a m e t e r a r e
i n h a b i t e d e i t h e r permanently o r s e a s o n a l l y .
Some i n h a b i t e d i s l a n d s
show a l m o s t no d i s t u r b a n c e by their r e s i d e n t s ( e . 9 . Guana I s l a n d ) ;
most however, a r e c l e a r l y modified by a g r i c u l t u r a l o r o t h e r p r a c t i c e s .
Cayo Luis Peiia and p a r t of Culebra I s l a n d a r e used f o r bombing and
n a v a l gunnery p r a c t i c e .
During World War 11, Desecheo was used f o r
bombing p r a c t i c e and bomb fragments can s t i l l be found on v a r i o u s p a r t s
of t h e island.
Human e f f e c t s have a c c e l e r a t e d g r e a t l y d u r i n g o u r study.
On many
s m a l l u n i n h a b i t e d i s l a n d s i n t h e beginning of t h e s t u d y one was reminded
o f t h e p r o x i m i t y of human p o p u l a t i o n s o n l y by the l a r g e numbers of
b o t t l e s , l i g h t b u l b s , s h o e s , e t c . encountered i n t h e beach d r i f t .
Most, however, e s p e c i a l l y i f c u r r e n t l y g o a t - f r e e , appeared i n a
relatively natural state.
Various such i s l a n d s c l o s e t o P u e r t o Rico,
were s e l e c t e d f o r d e t a i l e d , long-term s t u d y (e.9. Cayo Ahogado and
Palominitos).
I t was f o r t u n a t e t h a t o u r s t u d y was i n i t i a t e d a s e a r l y
a s it was because today one would have t o s e l e c t l e s s a c c e s s i b l e
i s l a n d s i n o r d e r t o c a r r y o u t such s t u d i e s .
Those n e a r P u e r t o Rico
with good anchorage f o r small c r a f t a r e now v i s i t e d by l a r g e numbers of
weekend e x c u r s i o n i s t s , who have t u r n e d t h e cays i n t o v e r i t a b l e garbage
and t r a s h dumps and have c u t woody v e g e t a t i o n f o r campfires and t e n t
supports.
I t a p p e a r s t h a t t h e s m a l l , i n h a b i t e d i s l a n d s whose owners
d i s c o u r a g e t r e s p a s s i n g a r e l i k e l y t o r e t a i n some s i m i l a r i t y t o t h e i r
n a t u r a l s t a t e , whereas u n i n h a b i t e d , p u b l i c ones a r e a l r e a d y b e i n g
r a p i d l y destroyed.
The l a r g e r i s l a n d s a l s o show an alarming degree of human
d i s t u r b a n c e s t o n a t u r a l communities.
For example, i n a s t u d y
c o n t r a c t e d by t h e P u e r t o Rican government, Heatwole (1970) found t h a t
t h a t i s l a n d had l o s t a mean of 1.7% of i t s coverage of mangrove p e r
decade between 1930 and t h e mid-1960's.
Between 1965 and 1970, l a r g e
t r a c t s were l o s t , some swamps dying completely.
Industrial pollution
and i n t e r f e r e n c e with d r a i n a q e p a t t e r n s by c o n s t r u c t i o n a c t i v i t i e s ,
d r e d g i n g , and f i l l i n g c o n t r i b u t e d t o t h e i r demise.
Proposed t o u r i s t i c
development i n t h e Vacia Talega a r e a w i l l probably e l i m i n a t e much o f
P u e r t o R i c o ' s b e s t remaining mangrove, and i s a l r e a d y f l o o d i n g t h e
i s l a n d ' s l a r g e s t s t a n d of A v i c e n n i a , n e a r Boguer&, w i t h f r e s h w a t e r
i n o r d e r t o c o n v e r t it i n t o a Typha marsh t o be used f o r waterfowl
hunting.
The f l o o d i n g was undertaken because a l l s i z e a b l e n a t u r a l
f r e s h - w a t e r marshes, once t h o u g h t v a l u e l e s s , had been e l i m i n a t e d some
t i m e i n t h e p a s t by a g r i c u l t u r a l development.
The d e s t r u c t i o n of h a b i t a t w i t h i n c r e a s i n g l e v e l s o f human
p o p u l a t i o n i s o f c o u r s e , n o t a problem unique t o i s l a n d s .
However,
t h e r e may be unique a s p e c t s a r i s i n g from i n s u l a r i t y .
I t would a p p e a r
t h a t t h r e e major t a s k s f a c e i s l a n d e c o l o g i s t s , (1) d e t a i l e d s t u d y o f
t h e f u n c t i o n i n g o f r e l a t i v e l y u n d i s t u r b e d i n s u l a r ecosystems w h i l e such
a r e s t i l l a v a i l a b l e , ( 2 ) t h e a s s e s s m e n t o f t h e e f f e c t o f human p r e s s u r e
on s u c h ecosystems, and ( 3 ) recommending p r a c t i c e s f o r u s e o f s m a l l
i s l a n d s i n ways t h a t w i l l n o t r e s u l t i n t h e i r d e s t r u c t i o n and l o s s t o
t h e human p o p u l a t i o n .
The s e r i e s , t o which t h i s p a p e r r e p r e s e n t s t h e
i n t r o d u c t i o n , c o n c e n t r a t e s l a r g e l y on t h e f i r s t of t h e s e t a s k s .
ACKNOWLEDGEMENTS
We a r e g r a t e f u l t o t h e c i t i z e n s of t h e U n i t e d S t a t e s o f America
who f i n a n c e d t h i s p r o j e c t through a N a t i o n a l S c i e n c e Foundation g r a n t
D r . Maxim Cerame-Vivas a i d e d by p u t t i n g t h e Marine
(GB-2906).
Research V e s s e l , Medusa, a t o u r d i s p o s a l .
The s u c c e s s o f t h e p r o j e c t
was g r e a t l y enhanced by t h e d e d i c a t i o n o f t h e r e s e a r c h a s s i s t a n t s ,
F a u s t i n o McKenzie and Frank T o r r e s , whose e f f o r t s f r e q u e n t l y went
beyond t h e c a l l o f d u t y .
Many s t u d e n t s and s t a f f o f t h e U n i v e r s i t y
o f P u e r t o R i c o a i d e d i n t h e f i e l d work.
V a r i o u s p e o p l e and
i n s t i t u t i o n s p r o v i d e d f a c i l i t i e s , l o d g i n g , t r a n s p o r t a t i o n and o t h e r
e s s e n t i a l a i d during f i e l d expeditions.
These were Eunice Boardman,
R e g i n a l d Penn, t h e P o l i c e Department o f Cabo Rojo, P.R., t h e U.S.
C o a s t Guard, t h e U.S. Navy, t h e I n s t i t u t e o f Marine Biology a t La
P a r g u e r a , P.R., t h e Department o f A g r i c u l t u r e o f t h e B r i t i s h V i r g i n
I s l a n d s , t h e C a t h o l i c U n i v e r s i t y o f P u e r t o R i c o , and t h e l a t e T i t o
Wirshing.
John Cooper and Steven D. Garber s u p p l i e d t h e i r
d i s t r i b u t i o n a l r e c o r d s f o r o u r use.
We a r e g r a t e f u l t o Frank T o r r e s ,
A l i c e Matta and A n g e l i c a M d i z f o r summarizing t h e a r a c h n i d d a t e .
Roy Woodbury a i d e d i n i d e n t i f i c a t i o n o f some o f t h e p l a n t s .
Terence
Done, E l i z a b e t h Cameron and Barbara S a y l o r Done c r i t i c i z e d t h e
m a n u s c r i p t and V i o l a Watt, Neva Walden, R u s s e l l Hobbs, Sue Dixon and
Audry Heatwole a i d e d i n i t s p r e p a r a t i o n .
LITERATURE: CITED
Abbott, I . 1974 Numbers o f p l a n t , i n s e c t and l a n d b i r d s p e c i e s on
n i n e t e e n remote i s l a n d s i n t h e S o u t h e r n Hemisphere.
B i o l . J.
L i n n . S o c . 6: 143-152.
Baker, H. G. and G.L. S t e b b i n s
Academic P r e s s , N e w York.
1965
The genetics o f colonizing species.
and E.E. B a l l 1972 B i o l o g i c a l c o l o n i z a t i o n o f r e c e n t l y
B a s s o t , J.M.
c r e a t e d i s l a n d s i n Lake Wisdom, Long I s l a n d , Papua New Guinea, w i t h
o b s e r v a t i o n s on t h e f a u n a o f t h e l a k e . Papua New Guinea S c i . S o c .
Proc. 23: 26-35.
Beard, J.S.
1944
127-158.
Becker, P.
1975
Coleoptera.
Climax v e g e t a t i o n i n t r o p i c a l America.
E c o l o g y 25:
I s l a n d c o l o n i z a t i o n by c a r n i v o r o u s and herbivorous
E c o l . 44: 893-906.
J . Anim.
B r a t t s t r o m , B.H.
1963 Barcena Volcano, 1952: I t s e f f e c t on t h e fauna
and f l o r a of San Benedicto I s l a n d , Mexico. Symposium 10th
P a c i f i c S c i . C o n g r e s s , pp. 499-524.
1924
B r i t t o n , N.L.
La vegetaci6n de Cayo I c a c o s .
Revista Agric.
P t o R i c o 1 2 : 91-96.
1933 An undescribed c a c t u s of Culebra I s l a n d , P u e r t o
B r i t t o n , N.L.
Rico.
C a c t u s and S u c c u l e n t J o u r n a l 5: 469.
B r i t t o n , N.L. and P. Wilson 1923-1930 D e s c r i p t i v e f l o r a Spermatophyta. Botany of P o r t o Rico and t h e V i r g i n I s l a n d s .
N . Y . A c a d . S c i . S u r v . P.R. and V i r g i n I s l s .
Vols. 5 and 6.
1956 La c o n s t i t u t i o n g 6 o l o g i q u e e t l a s t r u c t u r e d e s
A n t i l l e s : Centre National de l a Recherche S c i e n t i f i q u e , 453 pp.
B u t t e r l i n , J.
1965 I s l a n d L i f e .
C a r l q u i s t , S.
New York, 451 pp.
N a t u r a l H i s t o r y P r e s s , Garden C i t y ,
I.
C a r l q u i s t , S. 1966a The b i o t a of long-distance d i s p e r s a l .
P r i n c i p l e s of d i s p e r s a l and e v o l u t i o n . Q u a r t . R e v . B i o l . 41:
247-270.
1966b The b i o t a of long-distance d i s p e r s a l .
11. Loss
E v o l u t i o n 20: 30-48.
of d i s p e r s i b i l i t y i n P a c i f i c Compositae.
1966c The b i o t a of long-distance d i s p e r s a l .
IV.
Genetic systems i n t h e f l o r a s of o c e a n i c i s l a n d s . E v o l u t i o n 20:
433-455.
Plant
C a r l q u i s t , S. 1967 The b i o t a of long-distance d i s p e r s a l . V .
d i s p e r s a l t o P a c i f i c I s l a n d s . B u l l . T o r r e y B o t . C l u b 94: 129-161.
1974
York, 660 pp.
Carson, H.L.
2 : 3-6.
1970
Island Biology.
Columbia U n i v e r s i t y P r e s s , New
Chromosomal t r a c e r s of founder e v e n t s .
1960-1967
U.S. Dept. of Commerce Weather Bureau, A s h e v i l l e .
C l i m a t o l o g i c a l Data P u e r t o R i c o and V i r g i n I s l a n d s .
6-12.
Biotropica
vols.
Cook, M.T. and H.A. Gleason 1928 Ecological survey of t h e f l o r a of
P o r t o R i c o . P t o . R i c o D e p t . o f A g r i c . J o u r . 1 2 : 1-139.
C r o w e l l , K.L.
1962 Reduced i n t e r s p e c i f i c c o m p e t i t i o n among t h e b i r d s
E c o l o g y 43: 75-88.
of Bermuda.
1964
194-196.
On d e t e r m i n a n t s of i n s u l a r f a u n a s .
Amer. N a t . 47:
1968 R a t e s o f c o m p e t i t i v e e x c l u s i o n by t h e A r g e n t i n e
a n t i n Bermuda.
E c o l o g y 49: 551-555.
Dansereau, P. 1966 S t u d i e s on t h e v e g e t a t i o n o f P u e r t o Rico.
I.
D e s c r i p t i o n and i n t e g r a t i o n o f t h e p l a n t communities. Univ. P t o
R i c o I n s t i t . C a r i b . S t u d . S p e c . P u b l . 1: 1-45, 56-287.
D'Arcy, W.G.
1971 The i s l a n d o f Anegada and i t s f l o r a .
B u l l . No. 139: 1-21.
D'Arcy, W.G.
Bull.
1975 Anegada I s l a n d : v e g e t a t i o n and f l o r a .
188: 1-40.
Gleason, H.A.
A t o l l Res.
A t o l l Res.
and M.T. Cook 1927 P l a n t e c o l o g y o f P o r t o Rico.
7: 1-173.
A c a d . S c i . S u r v . P.R. and V i r g i n I s l s .
N.Y.
G r a n t , P.R.
1965 The a d a p t i v e s i g n i f i c a n c e o f some s i z e t r e n d s i n
E v o l u t i o n 19 : 355-367.
island birds.
G r a n t , P.R.
1966
Amer. N a t .
E c o l o g i c a l c o m p a t i b i l i t y o f b i r d s p e c i e s on i s l a n d s .
100: 451-462.
G r a n t , P.R.
1968 B i l l s i z e , body s i z e , and t h e e c o l o g i c a l a d a p t a t i o n s
o f b i r d s p e c i e s t o c o m p e t i t i v e s i t u a t i o n s on i s l a n d s . S y s t e m a t i c
Z o o l o g y 319-333.
G r e s s i t t , J . L . and C.M. Yoshimoto 1963 D i s p e r s a l o f a n i m a l s i n t h e
P a c i f i c . pp. 283-292 i n : p a c i f i c B a s i n B i o g e o g r a p h y , a S y m p o s i u m .
Bishop Mus. P r e s s , Honolulu.
Hamilton, T . H . and I . Rubinoff 1963 I s o l a t i o n , endemism, and
m u l t i p l i c a t i o n o f s p e c i e s i n t h e Darwin F i n c h e s .
E v o l u t i o n 17:
388-403.
Hamilton, T . H . , I . Rubinoff and R.H. B a r t h Jr. 1963 S p e c i e s
abundance: N a t u r a l r e g u l a t i o n of i n s u l a r v a r i a t i o n . S c i e n c e 142:
1575-1577.
Hamilton, T . H . and I . Rubinoff 1967 On p r e d i c t i n g i n s u l a r v a r i a t i o n
i n endemism and sympathy f o r t h e Darwin F i n c h e s i n the Galapagos
A r c h i p e l a g o . Amer. N a t . 101: 161-171.
1971 Marine-dependent t e r r e s t r i a l b i o t i c communities on
Heatwole, H.
some c a y s i n t h e c o r a l s e a . E c o l o g y 52: 363-366.
Heatwole, H.
1976a The e c o l o g y and b i o g e o g r a p h y o f c o r a l c a y s .
In:
B i o l o g y and G e o l o g y o f C o r a l R e e f s . Vol. 3 ( B i o l o g y . 1 1 ) ( e d s .
O.A. J o n e s a n d R . E n d e a n ) , Academic P r e s s , New York.
H e a t w o l e , H.
1976b H e r p e t o g e o g r a p h y o f P u e r t o R i c o . V I I .
Geographic
v a r i a t i o n i n t h e A n o l i s c r i s t a t e l l u s complex i n P u e r t o R i c o and
the Virgin Islands.
Occ. P a p e r s Mus. N a t . H i s t . Univ. K a n s a s 46:
1-18.
H e a t w o l e , H. and R . L e v i n s 1972a T r o p h i c s t r u c t u r e s t a b i l i t y and
E c o l o g y 53: 531-534.
f a u n a l change d u r i n g r e c o l o n i z a t i o n .
Heatwole, H. a n d R. L e v i n s 1972b Biogeography o f t h e P u e r t o R i c a n
Bank: f l o t s a m t r a n s p o r t o f t e r r e s t r i a l a n i m a l s .
E c o l o g y 53:
112-117.
H e a t w o l e , H . and R . L e v i n s 1973 Biogeography o f t h e P u e r t o R i c a n
Bank: s p e c i e s - t u r n o v e r on a s m a l l c a y , Cayo Ahogado. E c o l o g y 54:
1042-1055.
H e a t w o l e , H. a n d F. MacKenzie 1967 H e r p e t o g e o g r a p h y o f P u e r t o R i c o .
IV.
P a l e o g e o g r a p h y , f a u n a l s i m i l a r i t y and endemism.
Evolution
21: 429-438.
H e a t w o l e , H . , D.S. Sade a n d R. H i l d r e t h 1 9 6 3 H e r p e t o g e o g r a p h y o f
P u e r t o Rico.
I.
H e r p e t o f a u n a o f Cayo S a n t i a g o and Cayo B a t a t a .
C a r i b . J. S c i . 3: 1-5.
Hunt, G.L. a n d M.W. Hunt 1974 T r o p h i c l e v e l s a n d t u r n o v e r r a t e s : t h e
avifauna of Santa Barbara I s l a n d , C a l i f o r n i a .
Condor 76: 363-369.
J o h n s o n , M.P. a n d P.H. Raven 1973 S p e c i e s numbers and endemism: t h e
Galapagos Archipelago r e v i s i t e d .
S c i e n c e 179: 893-897.
1975 C o n t r o l s o f number o f b i r d s p e c i e s on montane
Johnson, N.K.
i s l a n d s i n the G r e a t B a s i n .
E v o l u t i o n 29: 545-563.
1959 S h o r e l i n e f e a t u r e s a n d q u a t e r n a r y s h o r e l i n e c h a n g e s
Kaye , C. A.
P u e r t o Rico.
U.S. G e o l . S u r v . P r o f e s s . Pap. 317-B: 49-140.
L e v i n s , R.
1969 Thermal a c c l i m a t i o n and h e a t r e s i s t a n c e i n D r o s o p h i l a
s p e c i e s . Amer. N a t . 103: 483-499.
L e v i n s , R. 1 9 6 8 E v o l u t i o n i n C h a n g i n g E n v i r o n m e n t s .
Press. Princeton, N.J.
P r i n c e t o n Univ.
L e v i n s , R . and H. Heatwole 1 9 6 3 On t h e d i s t r i b u t i o n o f o r g a n i s m s on
islands.
C a r i b . J. S c i . 3: 173-177.
L e v i n s , R . , M.L. P r e s s i c k a n d H. Heatwole
o f a n t s . Amer. S c i e n t . 61: 463-472.
1973
Coexistence p a t t e r n s
L e v i n s , R. and H. Heatwole 1 9 7 3 Biogeography o f t h e P u e r t o Rican Bank:
I n t r o d u c t i o n o f s p e c i e s on t o Palominitos I s l a n d .
Ecology 54:
1056-1064.
L i t t l e , E.L. Jr. and F.H. Wadsworth 1964 Common T r e e s o f P u e r t o R i c o
and t h e V i r g i n I s l a n d s . A g r i c . Handbook 249: 548 p p . U.S.D.A.
F o r e s t r y S e r v i c e , Washington D .C.
Lynch, J . F . a n d N.K. Johnson 1974 T u r n o v e r a n d e q u i l i b r i a i n i n s u l a r
a v i f a u n a s , w i t h s p e c i a l r e f e r e n c e t o t h e C a l i f o r n i a Channel
I s l a n d s . Condor 76: 370-384.
MacArthur, R.H. and E.O. Wilson 1963 An e q u i l i b r i u m t h e o r y o f i n s u l a r
zoogeography. E v o l u t i o n 1 7 : 373-387.
MacArthur, R.H. and E.O. Wilson 1967 T h e T h e o r y o f I s l a n d ~ i o g e o g r a p h y .
P r i n c e t o n Univ. P r e s s , P r i n c e t o n , N . J . ,
203 pp.
MacArthur, R . , J. MacArthur, D. MacArthur and A. MacArthur 1973 The
e f f e c t o f i s l a n d a r e a on p o p u l a t i o n d e n s i t i e s . E c o l o g y 5 4 : 657658.
MacLean, W.P., R . K e l l n e r and H . Dennis 1977 I s l a n d l i s t s o f W e s t
I n d i a n amphibian and r e p t i l e s . S m i t h s o n i a n H e r p e t o l o g i c a l
I n f o r m a t i o n S e r v i c e 40: 1-47.
1963 Animal S p e c i e s a n d E v o l u t i o n .
Mayr, E.
Cambridge, Mass. , 797 pp.
Mayr, E.
1965
Avifauna:
Turnover on i s l a n d s .
1933 Geology o f P u e r t o R i c o .
Meyerhoff, H.A.
S e r . B . , 1: 1-306.
H a r v a r d Univ. P r e s s ,
S c i e n c e 150: 1587-1588.
Univ. P t o R i c o Monogr.
M i t c h e l l , R.C.
1954 A s u r v e y o f t h e g e o l o g y o f P u e r t o Rico.
P t o R i c o A g r i c . E x p . S t a . T e c h . Pap. 1 3 : 1-167.
Univ.
P h i l i b o s i a n , R . and J . A . Yntema 1976 R e c o r d s and s t a t u s o f some
I . 1968-1975.
r e p t i l e s and a m p h i b i a n s i n t h e V i r g i n I s l a n d s .
H e r p e t o l o g i c a 32 : 81-85.
P h i l i b o s i a n , R . and Yntema, J . A .
Records and s t a t u s o f some r e p t i l e s
and a m p h i b i a n s i n t h e V i r g i n I s l a n d s .
11. 1975-1976.
Herpetologica, i n press.
P i c 6 , R.
1954 ~ e o g r a f y ad e P u e r t o R i c o .
P.R., 243 p p .
P r e s t o n , F. W.
1962a
r a r i t y : P a r t I.
E d i t o r i a l Univ.,
Rio P i e d r a s ,
The c a n o n i c a l d i s t r i b u t i o n o f commonness a n d
E c o l o g y 43: 185-215.
P r e s t o n , F.W.
1962b The c a n o n i c a l d i s t r i b u t i o n o f
r a r i t y : P a r t 11. E c o l o g y 43: 410-432.
R i c h a r d s , P.W.
1957 T h e T r o p i c a l R a i n Forest.
Cambridge, 450 pp.
commonness and
Cambridge Univ. P r e s s ,
Rosen, D.E.
1975 A v i c a r i a n c e model o f Caribbean biogeography.
S y s t e m a t i c Z o o l o g y 24: 431-464.
1962 E f f e c t s o f r e c e n t t r o p i c a l c y c l o n e s on the c o a s t a l
Sauer, J.D.
vegetation of Mauritius.
J o u r . E c o l . 50: 275-290.
Schoener, A.
1975
mini-islands.
E x p e r i m e n t a l zoogeography: c o l o n i z a t i o n o f marine
Amer. N a t . 108: 715-738.
1935 H i s t o r i c a l G e o l o g y o f t h e An t i l l e a n - C a r i b b e a n
S c h u c h e r t , C.
John Wiley and Sons, N e w York, 811 pp.
Region.
Schwartz, A. and R. Thomas 1975 A c h e c k - l i s t o f West I n d i a n
amphibians a n d r e p t i l e s . C a r n e g i e Mus. N a t H i s t . S p e c i a l Pub1
1: 1-216.
.
.
Sheppard, D . H . , P.H. K l o p f e r and H. Oelke 1968 H a b i t a t s e l e c t i o n :
D i f f e r e n c e s i n s t e r e o t y p y between i n s u l a r and c o n t i n e n t a l b i r d s .
Wilson B u l l . 80: 452-457.
S i m b e r l o f f , D.S.
1969 E x p e r i m e n t a l zoogeography o f i s l a n d s .
f o r i n s u l a r c o l o n i z a t i o n . E c o l o g y 50: 296-314.
A model
S i m b e r l o f f , D.S.
1970 Taxonomic d i v e r s i t y of i s l a n d b i o t a s .
~ v o l u t i o n24: 23-47.
S i m b e r l o f f , D.S. and E.O. Wilson 1969 E x p e r i m e n t a l zoogeography o f
E c o l o g y 50: 278-296.
i s l a n d s . The c o l o n i z a t i o n o f empty i s l a n d s .
Thomas, R. and A. S c h w a r t z 1966
Greater Puerto Rico region.
S p h a e r o d a c t y l u s (Gekkonidae) i n t h e
B u l l . F l a . S t a t e Mus. 10: 193-260.
Thornton, I.W.B.
1967 The measurement o f i s o l a t i o n on a r c h i p e l a g o s ,
and i t s r e l a t i o n t o i n s u l a r f a u n a l s i z e and endemism. E v o l u t i o n
21: 842-849.
U.S.
Dept. Commerce C o a s t and G e o d e t i c Survey.
1962 U n i t e d S t a t e s
C o a s t P i l o t 5 . A t l a n t i c C o a s t G u l f o f Mexico, P u e r t o R i c o , and
5 t h e d i t i o n . U.S. G o v ' t P r i n t i n g O f f i c e ,
Virgin Islands.
Washinqton, D. C . , 264 pp. p l u s 1965 supplement, 1 7 pp.
Whitehead, D.R. and C.E. J o n e s 1969
t h e o r y o f i n s u l a r biogeography.
S m a l l i s l a n d s and the e q u i l i b r i u m
E v o l u t i o n 23: 171-179.
W i l l i a m s , E.E.
1969 The e c o l o g y o f c o l o n i z a t i o n a s s e e n i n t h e
Quart. Rev.
zoogeography o f a n o l i n e l i z a r d s on s m a l l i s l a n d s .
B i o l . 44: 345-389.
Wilson, E.O. and R.W. Taylor 1967 An e s t i m a t e o f t h e p o t e n t i a l
e v o l u t i o n a r y i n c r e a s e i n s p e c i e s d e n s i t y i n t h e Polynesian a n t
fauna.
Evolution 21: 1-10.
TABLE 1.
TEMPERATURES (MEANS AND RANGES) I N DIFFERENT MICRO-HABITATS ON CAY0 AHOGADO, MARCH 16-21,
AIR TEMPERATURES
DATE
Rh %
T~~
1 cm
L a t e e v e n i n g and n i g h t o n l y .
**
E a r l y morning o n l y .
1 5 cm
SOIL TEMPERATURES
1m
5 cm d e e p
1 cm d e e p
1966.
TEMPERATURES AT A CLUMP OF VEGETATION
Surface
5 cm i n
s o i1
1 cm i n
soil
S o i l S u r f a c e S u r f a c e of
u n d e r Clump V e g e t a t i o n
Center of
Clump
Table 2.
Vegetation t y p e s p r e s e n t on t h e i s l a n d s o f t h e P u e r t o Rican
Bank. X i n d i c a t e s presence b u t cover e i t h e r n o t e s t i m a t e d
o r e s t i m a t e d t o have been l e s s than 1%. Where numbers a r e
g i v e n , they approximate t h e percentage o f t h e i s l a n d ' s a r e a
covered by a given v e g e t a t i o n t y p e .
I s l a n d s which do n o t
a p p e a r , e i t h e r were n o t s t u d i e d from t h e v e g e t a t i o n s t a n d p o i n t , o r w i l l be t r e a t e d i n more d e t a i l i n f u t u r e p a p e r s .
For names of v e g e t a t i o n types s e e t e x t .
VEGETATION TYPE
-
ISLAND NAME
Abel Key
A l i n a Key
Anegada
Bare Cactus
Beef I s l a n d
Big Cockroach
Big Hans L o l l i c k
Big Tobago
Blanquilla
Booby Hatch
Botella
Byer Bache
Caballo Blanco
Cabez de P e r r o
CajadeMuertos
Caiia Azul
Cayo Ahogado
Cayo B a t a t a
Cayo Enmedio
Cayo Enrique
Cayo G i l b e r t o
Cayo I s a b e l
Cayo Majimo
Cayo Norte
Cayo Olga
Cayo S a n t i a g o
Cayo Turramotecito
Chicken I s l a n d
Cooper Is l a n d
Cucaracha
(Las Cucarachas)
Culebra
Culebrit a
Dead Man's Chest
De s e cheo
Diablo
Double I s l a n d
East ~ a r a l l 6 n
~~-
1
2
3
x
4
5
6
x
x
x
x
x
10
x
x
x
x
x
8
9
100
100
5
85
x
x
x
x
x
x
80
10
x
60
35
10
85
x
x
65
x
7
1
0
1
1
1
2
10
X
X
x
x
x
x
x
40
x
5
100
100
x
x
x
90
75
5
25
x
x
x
5
x
x
5
5
x
20
x
x
x
x
5
75
x
x
100
100
90
5
x
x
x
x
x
x
100
x
90
x
x
x
x
x
x
x
x
x
x
x
10
x
x
4
1
50
x
x
x
x
x
40
5
x
x
x
x
x
x
x
45
75
50
25
x
x
x
x
3
x
x
x
5
48
x
x
1
x
x
x
x
48
x
x
x
x
Table 2
cont'd.
East ~ e n i q u i
E a s t S e a l Dog
Eustatia
F a l l e n Jerusalem
Frank Key
Frenchman' s Cap
(French Cap Key)
Frenchman ' s Cay
George Dog
Ginger I s l a n d
G r a n t Rock
Green Cay
(Near S t . C r o i x )
Guana
Gusano
Herb Key
Hicacos
Island 0
Island Q
Island R
I s l e t a Marina
J o s 6 Key
J u l i e Key
Key R e i s t
Konyoki
La G a t a
( I s l a La G a t a )
Laurel
Levin ' s Rock
L i t t l e Hans L o l l i c k
L i t t l e Saba I s l a n d
L i t t l e Tobago
Long Key
Magueyes
Mangrove Key
Marina Cay
Mata S e c a
Media Luna
Minnesota Mangrove
Mona
Mon it o
Mosquito I s l a n d
Necker I s l a n d
Norman I s l a n d
Outer Caracol
(Cavo C a r a c o l e s )
Palominitos
Palominos
Table 2
cont'd.
Pelican Key
Peter Island
Platillo
P r i c k l y Pear
P r o t e s t a n t Key
Puerto Rico
Ramos
Ratones
(Cayo Ratones)
Romero I1
Romero I11
Round Key
S a i l Rock
S a l t Island
San C r i s t o b a l
S h e i l a Rock
Solito
(Cayo S o l i t o )
Spaghetti Key
Spiny Butte
S t . Croix
S t . John
S t . Thomas
S u r p r i s e Key
Tortola
Turramot e
Vieques
V i l l a d e l Mar
Virgin Gorda
Watson Rock
West Dog
West ~ a r a l l 6 n
West Seal Dog
90
X
X
5
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
70
X
X
X
5
X
30
X
X
X
X
X
x
x
X
X
80
x
6
X
4
100
5
65
X
10
X
X
X
X
X
X
x
30
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
20
50
15
15
X
X
X
X
25
50
X
X
25
X
X
X
X
X
90
X
X
X
90
95
x
X
X
x
X
X
X
X
X
X
X
X
25
60
15
5
5
5
X
5
5
X
SPINY BUTTE
ROUND KEY
S A N CRlSTOBALo
CAYO ENMEDlOo
oLAUREL
LITTLE
M E D I A L U N A O'MEDIA
TURRAMOTECITO
LUNA
D~~~~~
TU RRAMOTE
C A Y O TERREMOTOO
0
km
2
Q+
PALOMINOS
0PALOMlNlTOS
.CAY0 A H O G A D O
PUERTO
c31SLA CABRAS
QCAYO A L G O D O N E S
0
km
2
w
LAS CUCARACHAS
EAST FARALLON
W E S T FARAILON-..
,
S H E I L A ROCK
'f,c
0-..
.
.
H ICAQU I
H l CACOS
C A Y 0 RATONES
PRESSICK ROCK
KON YOKl
DOUBLE I S L A N D
A U D R Y ROCK-*
R
,
S U R P R I S E KEY
C A Y 0 LOBOS
BOOBY ROCK
O
BLANQUILLA 2
G R A N T ROCK
0 0
P
M A C K E N Z I E KEY
&<-
6
<
'.$
0.
/
.
0.
C A Y O SOLITO-0
CAY0
NORTE
oROCA B A L L E N A
OROCA T I BURON
owEST GENlQUl
-EAST G E N l Q U l
0
LOS
HERMANOSO
QBARE CACTUS
Appendix I .
Maps o f s m a l l i s l a n d s on t h e P u e r t o Rican Bank.
I n c o n s i s t e n c y i n u s e o f " c a y " and "key" r e f l e c t s
d i f f e r e n c e s between t h e American and B r i t i s h V i r g i n
We p r e f e r
I s l a n d s i n t h e way t h e word i s s p e l l e d .
t h e former and u s e i t u n l e s s "key" i s p a r t of t h e
p r o p e r name o f an i s l a n d .
Cays i n t h e La P a r g u e r a r e g i o n of P u e r t o Rico ( e n l a r g e m e n t o f
r e c t a n g l e A i n Figure 1).
Cays immediately e a s t o f P u e r t o Rico ( e n l a r g e m e n t o f r e c t a n g l e B
i n F i g u r e 1)
1. Levin' s Rock
.
Cays between P u e r t o Rico and C u l e b r a I s l a n d ( e n l a r g e m e n t of
rectangle C i n Figure 1 ) .
C u l e b r a I s l a n d and i t s s u r r o u n d i n g c a y s ( e n l a r g e m e n t of r e c t a n g l e
D i n F i g u r e 1)
1. Chicken I s l a n d , 2 .
Mangrove Key.
.
The i s l a n d s and c a y s o f t h e American and B r i t i s h V i r g i n I s l a n d s ,
e x c e p t f o r S t . C r o i x and i t s n e a r b y i s l e t s ( e n l a r g e m e n t o f
rectangle E i n Figure 1 ) .
S a i l Rock
Savanna I s l a n d
C r i c k e t Rock
Cockroach I s l a n d ( n e a r S t . Thomas)
Dutchman' s Cap
S a l t Cay
West Cay
L i t t l e Saba I s l a n d
Outer Brass I s l a n d
Inner Brass I s l a n d
Hassel I s l a n d
Water I s l a n d
Buck I s l a n d ( n e a r S t . Thomas)
Cape1l a I s l a n d
Frenchman's Cap (French Cap ~ e y )
B i g Hans L o l l i c k
Thatch Key
G r e a t e r S t . James
L i t t l e S t . James
Big Tobago
L i t t l e Tobago
Mingo Key
Congo Key
Lovango Key
J o s t van Dyke
L i t t l e J o s t van Dyke
Sandy Key
G r e a t Thatch I s l a n d
L i t t l e Thatch I s l a n d
Frenchman's Key
Leduck I s l a n d
Flannigan I s l a n d
Norman I s l a n d
Peter Island
Dead Man's C h e s t
S a l t Island
Cooper I s l a n d
Ginger I s l a n d
Round Rock
Guana I s l a n d
Buck I s l a n d ( n e a r T o r t o l a )
Beef Is l a n d
Marina Cay
Scrub I s l a n d
G r e a t e r Camanoe
The Dog I s l a n d s : George
Dog, West Dog, G r e a t Dog
and B i g Cockroach ( n e a r
V i r g i n Gorda)
S e a l Dogs: E a s t S e a l Dog
and West Se-.?l Dog
P r i c k l y Pear
Necker I s l a n d
Watson Rock
P e l i c a n Key
55.
56.
57.
58.
59.
60.
61.
A s e r i e s of t i n y cays, R , 0 ,
Q and Byer Bache
Green Cay ( J o s t van Dyke)
Sandy S p i t
Eustatia
Saba Rock
L i t t l e Camanoe
Broken Jerusalem
Appendix 11.
F l o r a l l i s t s from some small i s l a n d s and cays i n t h e
Puerto Rican and Virgin I s l a n d a r e a . These should be
considered a s p l a n t records r a t h e r than complete f l o r a s
except f o r those i s l a n d s marked with an ( * ) f o r which
attempts were made t o c o l l e c t a l l species.
Major
i s l a n d s n o t included; f o r treatment of t h e i r f l o r a and
v e g e t a t i o n see B r i t t o n and Wilson (1923-30) , Gleason
and Cook (192 7) , L i t t l e and Wadsworth (1964) and
Byer h a s a long-term, comprehensive
Dansereau (1966)
study of t h e f l o r a of small i s l a n d s of t h i s region i n
progress.
.
ANEGADA
CAYO AHOGADO
See D ' Arcy (1971, 1975)
B I G COCKROACH
Cephalocereus royeni
Opuntia d i l l e n i i
B I G HANS LOLLICK
Thrinax argen t e a
Piscidia piscipul a
C itharexylum fruticosum
T o r r u b i a f r a g r a n s (?)
P i thecellobi um unguis-ca ti
Croton a s t r o i t e s
Agave mi s s i o n u m
Vigna marina
Coccol oba u v i f e r a
Hippomane m a n c i n e l l a
Bursera simarouba
C a p p a r i s i n d i c a (?)
See Heatwole and ~ e v i n s(1973)
CANA
AZUL
*
Avicennia n i t i d a
R h i zophora mangle
CAYO BATATA
See Heatwole, Sade and
H i l d r e t h (1963)
*
CAYO ENMEDIO
Sesuvium p o r t u l a c a s t r u m
*
CAYO ENRIQUE
CAYO MAJIMO
*
S e s u v i um p o r t u l a c a s t r u m
CAYO SANTIAGO
B I G TOBAGO
Conocarpus e r e c t a
Bulbostylis vestita
Agave m i s s i o n u m
C e p h a l o c e r e u s r oy e n i
Tabebuia h e t e r o p h y l l a
P i thecellobium unguis-cati
Torrubia fragrans
Bursera simarouba
Opuntia r e p e n s
See Heatwole, Sade and
H i l d r e t h (1963)
CAYO TURREMOTECITO
*
Sesuvium portulacastrum
COOPER ISLAND
GINGER ISLAND
B u r s e r a simarouba
Cakile lanceolata
C a r i c a papaya
Coccoloba u v i f e r a
Cocos n u c i f e r a
Croton a s t r o i tes
Euphorbi a b u x i f 01i a
Musa s p .
Sporobol u s v i r g i n i c u s
DOUBLE KEY
Avi cennia n i t i d a
C r ot o n a s troi tes
Croton r i g i d u s
Pisoni a subcordata
HICACOS
See B r i t t o n (1924)
ISLAND 0
*
*
ISLAND R
DEAD MAN'S CHEST
Croton s p .
Sporobolus v i r g i n i c u s
E A S T FARALLON
*
*
ISLAND Q
*
Borri chi a a r b o r e s c e n s
L a g u n c u l a r i a racemosa
S a l i c o x n i a ambigua
L E V I N S ROCK
EUSTATIA
Cakile lanceolat a
Croton a st r o i tes
S p o r o b o l u s vi r g i n i c u s
FALLEN JERUSALEM
Tabebuia h e t e r o p h y l l a
FRENCHCAP CAY
C a n a v a l i a mari t i m a
Clerodendron aculeatum
Cyperus p l a n i f o l i u s
Ficus laevigata
Mollugo v e r t i c i l l a t a
Portulaca g r a n d i f l o r a
GEORGE DOG ISLAND
*
C a n a v a l i a mari t i m a
E r i thralis fruticosa
Paspal um g l abrum
L I T T L E HANS LOLLICK
Cephalocereus r o y e n i
Croton a s troi tes
C r o t o n discolor
Jacquinia barbasco
Opuntia repens
Opuntia r u b e s c e n s
P i thecel l o b i u m u n g u i s - c a t i
Rivina humilis
Sporobolus v i r g i n i c u s
Thrinax argent e a
Torrubia fragrans
L I T T L E SABA
Cephal o c e r e u s r o y e n i
L I T T L E TOBAGO
Agave mi s s i o n u m
Bursera simarouba
Jacquemon t i a jamai censis
Lantana i n v o l u c r a t a
P i thecellobium unguis-cati
Sporobolus v i r g i n ic u s
Torrubia fragrans
Vigna marina
MOSQUITO ISLAND
C a r i c a papaya
Coccol oba u vi f e r a
T a b e b u i a he t e r o p h y l l a
T o r r u b i a f r a g r a n s ( ?)
NECKER
Bursera simarouba
Cactus i n t o r t u s
Guilandia c r i s t a
PALOMINITOS *
B o r r i chi a a r b o r e s c e n s
Cakile lanceolata
Canavalia mari tima
Cassytha f i l i f o r m i s
Cenchrus e c h i n a t a
Cenchrus p a u c i f l o r u s
Cyperus pl a n if o l i u s
Ere thrali s fruticosa
Ermodia 1i t t o r a l i s
Euphorbia b u x i f o l i a
Melanthera n i v e a
Scaevola plumeri
Suriana mari tima
Vigna marina
3 unidentified
PELICAN KEY
Capparis flexuosa
( d i s a p p e a r e d again before
June 1968)
Opuntia repens
Portulaca oleracea
PLATILLO
NORMAN ISLAND
Amaranthus d u b i u s
A t r i p l e x sp.
Bursera simarouba
Cappari s f l e x u o s a
Euphorbi a s p .
Laguncul a r i a racemosa
P i soni a a 1 bi da
Tamarindus i n d i c a
PRICKLY PEAR
Agave m i s s i o n u m
RATONES
S A I L ROCK *
Barren
SALT ISLAND
SAN CRISTOBAL
*
Philoxerus vermicularis
R h i zophora mangl e
SHEILA ROCK
*
TURREMOTO (CORRAL)
WEST DOG ISLAND
WEST FARAWLON
SOLITO
*
*
WEST SEAL DOG
SPINY BUTTE
*
Acuan v i r g a tum
Ayenia p u s i l l a
Boerhaavi a e r e c t a
C a p p a r i s fl e x u o s a
Commel i n a e l egans
Cordia a n g u s t i f o l i a
C r o t o n d i s c o 1 or
Croton h u m i l i s
Euphorbi a b e r t e r i a n a
Hibiscus b r a s i l i e n s i s
Leptocereus quadricost at u s
M o l l uchia tomentosa
Panicum maximum
Pectis linifolia
P i thecell obi um u n g u i s - c a t i
Portulaca pi1 osa
Pteramnus l a b i a l i s
Rauwolfia lamarckii
Rhizophora mangle
Ruellia tuberosa
S i d a glabra
S i da procumbens
Stigmaphyllon lingulatum
3 unidentified
Coccol oba u v i fe r a
Appendix 111.
R e p t i l e s and Amphibians o f t h e P u e r t o Rico V i r g i n I s l a n d Archipelago
The i s l a n d s from which each s p e c i e s was c o l l e c t e d a r e l i s t e d
f i r s t , ,followed by a l i s t i n g of a l l of t h e herpetofauna f o r each
An a t t e m p t was made t o do a complete survey on each i s l a n d .
island.
These l i s t s a r e based on our own c o l l e c t i o n s , t h o s e o f v a r i o u s
museums, l i t e r a t u r e r e c o r d s up t o 1970 which we have n o t v e r i f i e d (*I ,
and our own s i g h t r e c o r d s (**I
They do n o t i n c l u d e t h e r e c o r d s
o b t a i n e d by o t h e r i n v e s t i g a t o r s a f t e r 1972, and whose c o l l e c t i o n s we
have n o t seen.
Some of t h e s e l a t e r c o l l e c t i o n s add s p e c i e s f o r
i s l a n d s we have surveyed and p r e s e n t r e c o r d s from a number of i s l a n d s
A complete l i s t i n g of t h e known i n s u l a r
we d i d n o t v i s i t .
d i s t r i b u t i o n s of t h e P u e r t o Rican - V i r g i n I s l a n d s p e c i e s can be
o b t a i n e d from a combination of t h e p r e s e n t p a p e r , and t h e p u b l i c a t i o n s
of Schwartz and Thomas (1975) , P h i l i b o s i a n and Yntema (1976, i n p r e s s )
and Maclean e t a l . (1977).
I n o u r l i s t , i s l a n d names i n p a r e n t h e s e s
i n d i c a t e t h a t t h e s p e c i e s i n q u e s t i o n was recorded from t h a t i s l a n d b u t
probably does n o t o c c u r t h e r e a s a permanently e s t a b l i s h e d b r e e d i n g
population.
T h i s l i s t c o r r e c t s t h a t of Schwartz and Thomas (1975) f o r
Anolis c r i s t a t e l l u s c r i s t a t e l l u s and Anolis c r i s t a t e l l u s w i l e y a e i n
l i n e with t h e taxonomic assessment of Heatwole (1976b).
.
Schwartz and Thomas i n d i c a t e d A. c. c r i s t a t e l l u s t o i n h a b i t many
i s l a n d s e a s t o f P u e r t o Rico, which a r e i n f a c t i n h a b i t e d by A . c .
wileyae instead.
D i s t r i b u t i o n of S ~ e c i e s
LIZARDS
I uuan i d a e
Anolis acutus
Buck I s l a n d ( n e a r S t . Croix)
Green Key ( n e a r S t . Croix)
S t . Croix
Anolis cooki
Caja de Muertos
P u e r t o Rico
Anolis c r i s t a t e l l u s c r i s t a t e l l u s
Algodones
Caja de Muertos
Cardona Key
(Cayo Ahogado )
Cayo B a t a t a
Isleta ~ a r i n a '
Long I s l a n d
Magueyes I s l a n d
Platillo
P u e r t o Rico
Ramo s
Also h y b r i d s between A . c. c r i s t a t e l l u s and A . c . w i l e y a e .
Anol i s cris tatellus wi 1eyae
*
*
*
Algodones
Anegada
Bare C a c t u s I s l a n d
Beef I s l a n d ( n e a r T o r t o l a )
Blanquilla
Booby Hatch
Botella Island
Buck I s l a n d
( n e a r S t . Thomas)
Cabeza de P e r r o
Cayo Lobos
Cayo Norte
Cayo Ratones ( n e a r Hicacos)
Chicken I s l a n d
Cockroach I s l a n d
( n e a r S t . Thomas)
Cooper I s l a n d
Congo Cay
Culebra
Culebrita
Dead Man ' s Chest
I s l e t a Marina
Konyokf
L i t t l e Camanoe
L i t t l e Saba ( n e a r S t . Thomas)
L i t t l e Tobago
Lovango Key
Marina Cay
Norman I s l a n d
Necker I s l a n d
Palominitos
Palominos
Peter Island
Piiieros
Pifieritos
P r i c k l y Pear I s l a n d
Puerto ~ i c o l
*
*
*
*
*
*
*
*
*
**
Diablo
Dog I s l a n d
Dutchman's Cap
E a s t S e a l Dog
Eustatia
George Dog I s l a n d
Ginger I s l a n d
Grant Rock
G r e a t Dog I s l a n d
G r e a t e r Thatch I s l a n d
Green Cay ( n e a r T o r t o l a )
Guana I s l a n d
Hans L o l l i c k
Hassel I s l a n d
Hicacos
I s l a Cabras ( n e a r Roosevelt
Rds P.R.)
Ramos
S t . James
S t . John
S t . Thomas
S a l t Island
S a l t Key
Sandy Cay ( n e a r T o r t o l a )
Sandy S p i t
Savanna I s l a n d
Scrub I s l a n d
S u r p r i s e Key
Tobago
Tortola
Vieques
V i l l a d e l Mar
V i r g i n Gorda
Water I s l a n d
West Dog I s l a n d
West S e a l Dog I s l a n d
Anol is cuvi eri
P u e r t o Rico
*
Vieques
Also h y b r i d s between A. c. cristatellus and A. c. wileyae.
An01 i s desechensi s
De s e cheo
Anoli s evermanni
Puerto Rico
Anoli s gundlachi
Puerto Rico
Anolis krugi
Puerto Rico
An01 is monensi s
Monito
Mona
Anolis occul tus
Puerto Rico
Anolis poncensis
Puerto Rico
m o l i s pulchellus
*
*
*
Algodones
Anegada
Cabeza de P e r r o
Ca j a de Muertos
(Cayo hoga ado)
Cayo Batata
Guana I s l a n d
Hicacos
I s l e t a Marina
J o s t van Dyke
Lovango Key
Palominos
Peter Island
*
*
*
Cayo Norte
Cayo Ratones (near Hicacos)
Cayo Santiago
Culebra
Greater Camanoe
Piiieros
Puerto Rico
S t . James
S t . John
S t . Thomas
Tortola
Vieque s
Virgin Gorda
Water I s l a n d
An01 i s roosevel ti
*
Culebra
Anolis stratulus
*
*
*
Beef I s l a n d ( n e a r T o r t o l a )
Cayo Santiago
Congo Key
Culebra
Ginger I s l a n d
G r e a t e r Thatch I s l a n d
Guana I s l a n d
J o s t van Dyke
Marina Cay
Necker I s l a n d
*
Norman I s l a n d
Peter Island
Piiieros
P r i c k l y Pear
P u e r t o Rico
S t John
S t . Thomas
Savanna I s l a n d
Scrub I s l a n d
Tortola
Vieques
V i r g i n Gorda
Water I s l a n d
.
*
*
Cyclura cornuta
Mona
Cyclura p i n g u i s
Ane gada
Iguana i g u a n a
*
*
Guana I s l a n d
Hassel I s l a n d
Hicacos ( p e r s . com. S.D. Garber)
Peter Island
P u e r t o Rico
*
*
*
*
*
S t . Croix
S t . John
S t . Thomas
Tortola
Water I s l a n d
Te i idae
Ameiva a l b o g u t t a t a
Mona
Amei va d e s e c h e n s i s
Desecheo
Ameiva e x s u l
*
Algodones
Anegada
Beef I s l a n d
Blanquilla
Cabeza de P e r r o
Caja de Muertos
(Levin' s Rock)
L i t t l e Camanoe
Lovango Key
*
Cardona Key
(Cayo Ahogado)
Cayo B a t a t a
Cayo Lobos
Cayo Norte
Cayo Ratones
Cooper I s l a n d
Culebra
Culebrita
Dead Man's Chest
Diablo
Dutchman's Cap
Eustatia
Ginger I s l a n d
Guana I s l a n d
~ a nL
s ollick
Hassel I s l a n d
Hicacos
I n n e r Brass
I s l e t a Marina
~onyoki
*
Marina Cay
Necker I s l a n d
Norman I s l a n d
Palominitos
Palominos
Peter Island
Piiieros
Platillo
P r i c k l y Pear
P u e r t o Rico
Ramos
S t . James
S t John
S t . Thomas
S a l t Island
S a l t Key
Savanna I s l a n d
Scrub I s l a n d
Tobago
Tortola
Vieques
V i r g i n Gorda
Water I s l a n d
.
*
Amei va p o l ops
Green Key ( n e a r S t . Croix)
* S t . Croix
Ameiva wetmorei
Puerto Rico
Caja de Muertos
Scincidae
Mabu ya mabou ya sl oanei
*
*
Anegada
Cayo Norte
Culebra
Culebrita
Dead Man's Chest
Ginger I s l a n d
Hicacos
J o s t van Dyke
*
*
*
**
**
*
*
*
Mona
Monito
Necker I s l a n d
Peter Island
P u e r t o Rico
S t . John
S t . Thomas
S a l t Island
Vieques (Perhaps now l o c a l l y
extinct)
V i r g i n Gorda
Diploglossus pleii
P u e r t o Rico
Gekkonidae
Hemidact yl us brooki
Puerto Rico
Hemi dactyl us mabouia
*
*
Culebra
J o s t van Dyke
Mona
Peter Island
Puerto Rico
S t . Croix
S t . John
*
*
S t . Thomas
S a l t Island
Tobago
Tortola
Vieques
Water I s l a n d
Phyllodactylus wirshingi
*
*
C a j a de Muertos
P u e r t o Rico
Sphaerodactylus beattyi beattyi
Buck I s l a n d ( n e a r S t . C r o i x )
Green Key ( n e a r S t . C r o i x )
S t . Croix
Sphaerodact yl us beat tyi seamani
*
S t . Croix
Sphaerodactylus gai geae
Puerto Rico
Piiieros
Sphaerodactylus klauberi
P u e r t o Rico
Sphaerodactylus levinsi
Desecheo I s l a n d
Sphaerodactylus macrolepis ateles
P u e r t o Rico
Sphaerodact y l u s m a c r o l e p i s grandisquamis
Algodone s
Cabeza de Perro
Cayo Batata
Cayo Santiago
Puerto Rico
Sphaerodactylus macrolepis guarionex
Puerto Rico
Sphaerodactylus macrolepis i n i g o i
*
*
Cayo de Afuera
Cayo de T i e r r a
Vieques
Sphaerodactylus macrolepis macrolepis
*
*
*
Ane gada
Bare Cactus I s l a n d
Beef I s l a n d
Big Cockroach
Botella
Cooper I s l a n d
Congo Key
Culebra
Culebrita
Dead Man's Chest
E a s t Geniquf
East Seal Dog
Eustatia
Fa1 l e n Jerusalem
Ginger I s l a n d
Great Dog I s l a n d
Greater Camanoe
Greater Thatch
Green Cay (near T o r t o l a )
Guana I s l a n d
Hans L o l l i c k
J o s t van Dyke
L i t t l e Camanoe
L i t t l e Saba
*
*
Buck I s l a n d (near S t . Croix)
Cayo Luis Pefia
Cayo Norte
Chicken I s l a n d
Cockroach ( n e a r S t . Thomas)
L i t t l e Tobago
Marina Cay
Mosquito
Necker I s l a n d
Norman I s l a n d
Peter Island
P r i c k l y Pear
S a i l Rock
S t . Croix
S t John
S t . Thomas
S a l t Island
Sandy Cay (near T o r t o l a )
Sandy S p i t
Savanna I s l a n d
Scrub I s l a n d
Tobago
Tortola
V i l l a d e l Mar
Virgin Gorda
Water I s l a n d
Watson Rock
West Seal Dog I s l a n d
.
*
*
Sphaerodactyl u s macrolepis mimetes
Puerto Rico
Sphaerodactylus macrolepis phoberus
Puerto Rico
Sphaerodact yl us macro1 epis spani us
Puerto Rico
Sphaerodactylus macrolepis stibarus
Piiieros
Sphaerodactylus monensis
Mona
Sphaerodactylus nicholsi nicholsi
Spiny Butte
(Turramote Key)
Puerto Rico
Sphaerodactylus ni cholsi townsendi
Hicacos
I s l a Cabras (near Roosevelt
Rds., P.R.)
Konyokf
MacKenzie Key
Palominos
Piiieros
Platillo
Puerto Rico
Ramos
S u r p r i s e Key
Vieques
Blanquilla
Booby Hatch
*
*
*
Caja de Muertos
Cayo de Afuera ( n e a r ~ i e q u e s )
Cayo de T i e r r a ( n e a r ~ i e q u e s )
Cayo Lobos
Cayo Ratones ( n e a r ~ i c a c o s )
Culebra
Diablo
French Cap Key
Grant Rock
Sphaerodact yl us parthenopi on
Virgin Gorda
Sphaerodactylus roosevel ti
Caja de Muertos
*
Puerto Rico
Vieques
Thecadactylus rapicaudus
Necker I s l a n d
*
St.
AMPHISBAENIANS
Amphisbaenidae
Croix
Amphisbaena bakeri
Puerto Rico
Amphisbaena caeca
P u e r t o Rico
Amphisbaena fenes trata
*
*
*
*
Greater Camanoe
S t . James
S t . John
*
*
S t . Thomas
Tortola
Virgin Gorda
Amphisbaena schmi d ti
Puerto Rico
Amphisbaena xera
Caja de Muertos ( c a l l e d A. caeca
by Heatwole et al. 1965)
Puerto Rico
SNAKES
Boidae
Epi cra tes inorna tus
P u e r t o Rico
picr rates monensis
*
*
Mona
Tortola
Colubridae
Alsophis portori censis
Anegada
Buck I s l a n d (near S t . Thomas)
Cabeza de P e r r o
Ca j a de Muertos
Cayo Santiago
Culebra
Ginger I s l a n d
Green Cay ( n e a r T o r t o l a )
Guana I s l a n d
Mona
Mosquito
Necker I s l a n d
Norman I s l a n d
*
*
*
*
Peter Island
Platillo
Puerto Rico
S t . John
S t . Thomas
S a l t Island
*
*
*
*
J o s t van Dyke
L i t t l e Saba
L i t t l e Tobago
Savanna I s l a n d
Vieques
Virgin Gorda
Alsophis sancticrucis
*
S t . Croix
~ r ryht o n e x i guum
*
*
Anegada
Cayo Santiago
Culebra
Greater Camanoe
Hassel I s l a n d
Peter Island
*
Puerto Rico
S t . John
S t . Thomas
Tortola
Virgin Gorda
Typhlopidae
~ y p h l o p sg r a n ti
Puerto Rico
~ y p hol p s m o n e n s i s
Mona
~ y p h l o p sr i c h a r d i
*
Cayo Norte
Culebra
Diablo
Guana I s l a n d
Palominitos
*
*
*
Puerto Rico
S t . Croix
S t . John
S t . Thomas
Surprise Key
Tortola
Virgin Gorda
~ y p h l o p sr o s t e l l a t a
Puerto Rico
TURTLES (excluding marine ones )
Emydidae
C h r ysemys d e c u s s a t a
Puerto Rico
ste j n e g e r i
Tes t u d i n i d a e
Geochelone carbonaria
*
*
*
*
S t . John
S t . Thomas
Tortola
Water I s l a n d
ANU RANS
Bu £oni d a e
B u f o lemur
P u e r t o Rico
B u f o marinus (introduced)
P u e r t o Rico
S t . Croix
V i eques
A 1 godones
Culebra
Bufo t u r p i s
*
V i r g i n Gorda
Leptodactylidae
Eleutherodactylus a n t i l l e n s i s
Culebra
Piiieros
P u e r t o Rico
S t . Croix ( i n t r o d u c e d ? )
S t . John
S t . Thomas
Tortola
Vieques
V i r g i n Gorda
E l e u t h e r o d a c t y l u s bri t toni
P u e r t o Rico
E l e u t h e r o d a c t y l u s cochranae
*
Hassel I s l a n d
P u e r t o Rico
S t . John
S t . Thomas
Tortola
E l e u t h e r o d a c t y l u s cooki
P u e r t o Rico
Eleutherodactylus coqui
P u e r t o Rico
Eleutherodactylus eneidae
P u e r t o Rico
Eleutherodactylus gryllus
P u e r t o Rico
~le u t h e r o d a c t y l u s h e d r e c k i
P u e r t o Rico
Eleutherodactylus karlschmidti
P u e r t o Rico
Eleutherodactylus l e n t u s
S t . Thomas
S t . Croix
E l e u t h e r o d a c t y l u s 1o c u s t u s
P u e r t o Rico
Eleutherodactylus monensis
Mona
Eleutherodactylus portoricensis
P u e r t o Rico
Eleutherodact y l u s richmondi
P u e r t o Rico
Eleutherodactyl u s schwartzi
G r e a t Dog
S t . Croix (introduced?)
S t . John
Tortola
V i r g i n Gorda
E l eutherodactyl u s unicolor
P u e r t o Rico
E l e u t h e r o d a c t y l u s wightmanae
P u e r t o Rico
L e p t o d a c t y l u s a1 i b i l a b r i s
P u e r t o Rico
S t . Croix
S t . John
S t . Thomas
Tortola
Vieques
Algodones
Ane gada
Culebra
Hicacos
Piiieros
Rani dae
Rana ca t e s b e i a n a ( i n t r o d u c e d )
Vieques (J.E
com.
P u e r t o Rico
. Cooper,
pers
.
Fauna of I s l a n d s
Algodone s
Ameiva
Anolis
Anolis
Anolis
exsul
c r i s t at e l l u s c r i s t at e l l u s
c r i s t a t e l l u s wileyae
pulchell u s
Bufo marinus (introduced)
Leptodactylus a l b i l a b r i s
Sphaerodact yl u s m a c r o l e p i s g r a n d i s quami s
Anegada
*
A1 s o p h i s p o r t o r i c e n s i s
Ameiva e x s u l
Anolis c r i s t a t e l l u s wileyae
Anolis pulchellus
Arrhyton exigum
* Cyclura
pinguis
* Mabuya mabouya s l o a n e i
Bare Cactus I s l a n d
S p h a e r o d a c t y l u s m a c r o l e p i s macro1 e p i s
B e e f Island (near Tortola)
Ameiva e x s u l
Anolis stratulus
Big Cockroach
Ameiva e x s u l
S p h a e r o d a c t y l u s n i c h o l s i townsendi
Booby Hatch
S p h a e r o d a c t y l u s n i c h o l s i townsendi
Botella Island
*
*
*
*
Mabuya mabouya s l o a n e i
Buck I s l a n d ( n e a r S t . Croix)
Anolis acutus
Sphaerodact yl u s m a c r o l e p i s macro1 e p i s
Sphaerodact yl u s b e a t t y i bea t t y i
*
Ameiva e x s u l
Cabeza de P e r r o
Anolis pulchellus
Sphaerodactylus macrolepis
grandisquamis
d l sophis portoricensis
Ameiva e x s u l
Caja de Muertos
Alsophis portoricensis
Ameiva e x s u l
Amei va we tmorei
Amphi sbaena x e r a
A n o l i s cooki
*
An01 i s pul chell u s
Phyllodactylus wirshingi
S p h a e r o d a c t y l u s n i c h o l si townsendi
Sphaerodactylus r o o s e v e l t i
Cardona Key
Ameiva e x s u l
*
A n o l i s cris t a t e l l u s c r i s t a t e l l u s
Cayo Ahogado
(Ameiva e x s u l )
(Anolis pulchell us)
(Anolis cristatellus cristatel l u s )
Cayo B a t a t a
Ameiva e x s u l
Anolis c r i s t a t e l l u s cristatellus
Anolis pulchellus
Sphaerodactylus macrolepis grandisquami s
Cayo de Afuera
*
S p h a e r o d a c t y l u s macrol e p i s i n i g o i " S p h a e r o d a c t y l u s n i c h o l si townsendi
Cayo de T i e r r a
*
S p h a e r o d a c t y l u s macrol e p i s i n i g o i * S p h a e r o d a c ty l u s nicholsi townsendi
Cayo Lobos
Ameiva e x s u l
S p h a e r o d a c t y l u s n i chol si t o w n s e n d i
Cayo Luis Peiia
*
Cayo Norte
Ameiva e x s u l
Anolis pulchellus
Typhl o p s ri c h a r d i
Cayo Ratones
Ameiva e x s u l
Sphaerodact y l u s n i c h o l si t o w n s e n d i
Ameiva e x s u l
Anolis c r i s t a t e l l u s c r i s t a t e l l us
Anolis s t r a t u l u s
A r r h y ton
B u f o marinus (introduced)
Sphaerodactyl u s gaigeae
S p h a e r o d a c t y l u s macro1 e p i s g r a n d i s quami s
Chicken I s l a n d
*
Anolis c r i s t a t e l l u s w i l e y a e
S p h a e r o d a c t yl u s macro1 e p i s macro1 e p i s
Cooper I s l a n d
Ameiva e x s u l
Sphaerodactyl u s macrolepis
Congo Key
*
*
A n o l i s cris t a t e l l u s w i l e yae
*Sphaerodactyl us
macrol e p i s m a c r o l e p i s
Culebra
*
A1 s o p h i s p o r t o r i censis
Ameiva e x s u l
Anolis pulchel l u s
An01 i s roosevel ti
Arrhyton exiguum
B u f o m a r i n u s (introduced)
*
*
*
H e m i d a c t y l u s mabouia
Leptodact y l u s a l b i l a b r i s
S p h a e r o d a c t y l u s nicholsi townsendi
Typhlops r i c h a r d i
Culebrita
Ameiva e x s u l
*
Dead Man's Chest
Ameiva e x s u l
S p h a e r o d a c t y l u s m a c r o l e ~si m a c r o l e p i s
Dese cheo
Amei v a d e s e c h e n s i s
Anolis desechensis
S p h a e r o d a c t y l u s 1evinsi
Ameiva e x s u l
S p h a e r o d a c t y l u s n i cholsi t o w n s e n d i
Dog I s l a n d
*
Dutchman ' s Cap
Ameiva e x s u l
*
E a s t Geniqui
Sphaerodactyl u s macrol e p i s macrol e p i s
E a s t S e a l Dog
S p h a e r o d a c t y l u s macro1e p i s m a c r o l e p i s
Eustatia
~ m e i v ae x s u l
Anolis pulchellus
French Cap K e y
Sphaerodactyl u s n i c h o l si townsendi
Ameiva e x s u l
Anolis c r i s t a t e l lus wileyae
Ginger I s l a n d
Ameiva e x s u l
An01 i s c r i s t a t e l l u s w i l e yae
mabou ya sl o a n e i
* *Mabuya
Grant Rock
Sphaerodactylu s n i c h o l s i townsendi
Great Dog I s l a n d
*
Eleutherodactyl us schwartzi
*
Ameiva e x s u l
Amphi sbaena f e n e s t r a t a
Anolis p u l c h e l l u s
Anolis stratulus
Arrh yton exiguum
Greater Thatch I s l a n d
Anolis c r i s t atell u s wileyae
Anolis stratulus
Green Key (near S t . Croix)
Amei v a p o l o p s
Anolis acutus
Sphaerodactyl u s b e a t ty i b e a t t y i
Green Cay (near T o r t o l a )
Sphaerodactylus macrolepis inacrolepis
Guana I s l a n d
Ameiva e x s u l
A n o l i s p u l chell u s
*
Anolis stratulus
Iguana iguana
S p h a e r o d a c t y l u s macro1 e p i s m a c r o l e p i s
Hans Lollick
Ameiva e x s u l
Hassel I s l a n d
*
Ameiva e x s u l
A r r hyton exiguum
*
*
E l e u t h e r o d a c ty l u s cochranae
Iguana iguana
Hicacos
Ameiva
Anolis
Anolis
Iguana
exsul
pul c h e l l u s
i g u a n a ( p e r s . corn.
Mabu ya mabou ya s l o a n e i
S p h a e r o d a c t y l u s nicholsi t o w n s e n d i
S.D. Garber)
Ameiva e x s u l
I s l a Cabras (near Roosevelt Rds ., P.R. )
Anolis c r i s t a t e l l us wileyae
I s l e t a Marina
Ameiva e x s u l
Anolis c r i s t a t e l l u s c r i s t a t e 1 l u s A . c . c r i s t a t e l l u s X A. c. w i l e y a e
J o s t van Dyke
*
*
*
*
*
Anolis stratulus
Sphaerodacty l u s m c r o l e p i s macrolepis
Konyokf
Ameiva e x s u l
Levin' s Rock
(Ameiva e x s u l )
L i t t l e Camanoe
Ameiva e x s u l
A1 s o p h i s r i c h a r d i
Ameiva e x s u l
Anolis cristatellus wileyae
Sphaerodactyl u s macrole p is macrole p i s
Amei v a e x s u l
Amphi s b a e n a f enes t r a t a
*
*
Ameiva e x s u l
*
*
*
Sphaerodactyl u s macrolepis macrolepis
*
Ameiva e x s u l
*
*
L i t t l e Tobago
Long I s l a n d
Anolis cristatellus cristatellus
Lovango Key
*
Ameiva e x s u l
*
MacKenzie Key
Amei v a w e t m o r e i
S p h a e r o d a c t y l u s n i cholsi n i chol si
Anolis c r i s t a t e l l u s c r i s t a t e l l u s S p h a e r o d a c t y l u s roosevel ti
Marina Cay
Ameiva e x s u l
Anolis stratulus
Mona
*
Ameiva a l b o y u t t a t a
An01 i s m o n e n s i s
Cyclura cornuta
Eleutherodactyl u s monensis
Epi c r a t e s m o n e n s i s
Hemi d a c t y l u s mabouia
Mabu ya mabou ya sl o a n e i
Sphaerodactylus monensis
T y p h l o p s monensi s
Monito
An01 i s m o n e n s i s
**Mabu ya mabouya s l o a n e i
A1 s o p h i s p o r t o r i censi s
Ameiva e x s u l
Sphaerodactyl u s parthenopion
Necker I s l a n d
Ameiva e x s u l
Anolis stratulus
Mabu ya mabou ya s l o a n e i
Thecadactylus r a p icaudus
Norman I s l a n d
d l sophi s portori c e n s is
Ameiva e x s u l
S p h a e r o d a c t y l u s macro1 e p i s m a c r o l e p i s
Palominitos
dineiva e x s u l
Typhl o p s r i c h a r d i
Palominos
dmeiva e x s u l
Anolis pulchellus
Sphaerodact y l u s n i c h o l si t o w n s e n d i
Peter Island
*
*
d lsophis portoricensis
dineiva e x s u l
Anolis pulchellus
d r r h y t o n exiguum
Hemidactyl u s mabouia
Iguana i g u a n a
dineiva e x s u l
Anolis stratulus
E l e u t h e r o d a c t yl u s a n t i l l e n s i s
Sphaerodactylus gaigeae
Sphaerodactylus macrolepis s t i b a r u s
Sphaerodact y l u s n i c h o l s i t o w n s e n d i
Piiieritos
d n o l i s c r i s t a t e l l u s wileyae
S p h a e r o d a c t y l u s n i c h o l si t o w n s e n d i
d l sophis portoricensis
**dineiva e x s u l
Prickly Pear Island
Ameiva e x s u l
Amei v a p o l o p s
Anolis acutus
P u e r t o Rico
Ameiva e x s u l
Amei v a we t m o r e i
Arrhyton exiguum
Amphisbaena b a k e r i
Bufo lemur
Amphisbaena c a e c a
~ u f m
o arinus (introduced)
Amphisbaena s c h m i d t i
Chrysemys d e c u s s a t a s t e j n e g e r i
Amphisbaena x e r a
Diploglossus p l e i i
Anolis cooki
A n o l i s c r i s t a t e l l u s c r i s t a t e l l u s E l e u t h e r o d a c t y l u s brittoni
Eleu therodactylu s cochranae
Anolis c. c r i s t a t e l l u s X A. c . E l e u t h e r o d a c t y l u s cooki
wileyae
Eleu t h e r o d a c t y l u s coqui
Anolis cuvieri
Anolis e v e r m a n n i
Eleu t h e r o d a c t y l u s eneidae
E l e u t h e r o d a c ty l u s g r y l l u s
Anolis g u n d l a c k i
E l eutherodacty l u s hedrecki
Anolis k r u g i
Eleutherodactylus karlschmidti
An01 i s o c c u l t u s
Anolis p o n c e n s i s
Eleutherodactylus locustus
Eleutherodactyl u s portoricensis
E l e u t h e r o d a c t y l u s r i chmondi
Sphaerodactylus macrolepis a t e l e s
Sphaerodacty l u s macrolepis grandisEleutherodactylus unicolor
quamis
E l e u t h e r o d a c t y l u s wightmanae
Sphaerodactylus macrolepis guarionex
Epicrates inornatus
Sphaerodactylus macrolepis mimetes
H e m i d a c t y l u s brooki
Sphaerodactylus m a c r o l e p i s phoberus
Sphaerodactylus macrolepis spanius
S p h a e r o d a c t y l u s nicholsi nicholsi
Iguana i guana
Leptodactyl u s a l b i l a b r i a
S p h a e r o d a c t y l u s roosevel ti
* Phyllodactylus wirshingi
Typhlops g r a n t i
Rana c a t e s b e i a n a ( i n t r o d u c e d )
Typhlo p s r ic h a r d i
Sphaerodactylus gaigei
Typhlops r o s t e l l a t a
Sphaerodactylus klauberi
Ramo s
Ameiva e x s u l
Anolis c r i s t a t e l l u s c r i s t a t e l l u s S p h a e r o d a c t y l u s nicholsi t o w n s e n d i
S a i l Rock
S t . Croix
*
*
Alsophis sancticrucis
Ameiva p o l o p s
Anolis acutus
Bufo marinus (introduced)
E l eutherodactylus antillensis
*
Iguana iguana
Sphaerodactylus b e a t t y i b e a t t y i
S p h a e r o d a c t y l u s b e a t t y i seamani
*
*
Thecadactylus rapicaudus
Typhlops richardi
*
( i n t r o d u c e d ?)
E l e u t h e r o d a c t y l u s 1en t u s
Eleutherodactylus schwart z i
(introduced?
S t . James
*
*
*
Ameiva e x s u l
Amphisbaena f e n e s t r a t a
S t . John
*
*
*
Ameiva e x s u l
Anolis stratulus
A r r h y t o n exiguum
*
*
*
*
Eleu t h e r o d a c t y l u s cochran ae
Eleutherodactylus schwart z i
Geochelone c a r b o n a r i a
Iguana iguana
Leptodactylus a l b i la h r i s
Typhlops richardi
S t . Thomas
*
*
*
Ameiva e x s u l
E l e u t h e r o d a c ty l u s cochranae
Eleutherodactylus l e n t u s
Geochelone c a r b o n a r i a
Iguana iguana
*
*
Arrhyton exiguum
E l e u t h e r o d a c t y l u s antillensis
Sphaerodactyl u s macrole p i s m a c r o l e p i s
S a l t Island
*
Ameiva e x s u l
*
Salt Key
*
Ameiva e x s u l
*
Sandy Cay ( n e a r T o r t o l a )
Sandy S p i t
Anolis cris t a t e l l u s w i l e y a e
Savanna I s l a n d
*
*
*
*
Ameiva e x s u l
Scrub I s l a n d
An01 i s s t r a t u l u s
Ameiva e x s u l
Spiny B u t t e
S p h a e r o d a c t y l u s nicholsi n i c h o l s i
S u r p r i s e Key
Sphaerodactyl u s n i c h o l s i townsendi
Tobago
Ameiva e x s u l
Hemi d a c t y l u s mabouia
Tortola
*
*
Ameiva e x s u l
Anolis s p . ( g i a n t form)
A r r h y t o n e x i guum
Eleutherodactylus a n t i l l e n s is
E l e ut h e r o d a c t y lu s cochranae
*
*
*
E l e u t h e r o d a c t y l u s schwart z i
Epicrates monensis
Geochelone carbonaria
Iguana iguana
Typhlops richardi
Turramote Key
( S p h a e r o d a c t y l u s nicholsi n i c h o l s i )
Vieques
*
*
Ameiva e x s u l
Anolis cuvieri
A n o l i s p u l chell u s
Anolis stratulus
~ uo fm a r i n u s ( i n t r o d u c e d )
E l e u t h e r o d a c t y l u s an t i l l e n s i s
Hemidact y l u s mabouia
*
*
Mabuya mabouya s l o a n e i (now probably
locally extinct)
Rana c a t e s b e i a n a ( i n t r o d u c e d ) (J.E .
Cooper, p e r s . comrn.
Sphaerodactylus macrolepis i n i g o i
S p h a e r o d a c t y l u s nicholsi t o m s e n d i
S p h a e r o d a c t y l u s roosevel ti
V i l l a d e l Mar
Virgin Gorda
*
Ameiva e x s u l
Anolis pulchell us
Anolis stratul us
A r r h y ton e x i g u u m
*
Bufo t u r p i s
E l e u t h e r o d a c t ydus s c h w a r t z i
Mabu ya mabouia s l o a n e i
Sphaerodactylu s parthenopion
Typhlo p s r ic h a r d i
Water I s l a n d
*
*
*
Ameiva
Anolis
Anolis
Anolis
*
*
*
*
exsul
pulchel l u s
stratulus
Geochel one c a r b o n a r i a
Hemidact y l u s mabouia
Iguana iguana
Watson Rock
**Anolis c r i s t a t e l l u s wileyae
West S e a l Dog I s l a n d
S p h a e r o d a c t y l u s macro1 e p i s macro1 e p i s
Appendix I V .
D i s t r i b u t i o n of t h e scorpion C e n t r u r o i d e s n i t i d u s and
t h e s p i d e r Gasteracantha t e t r a c a n t h a on the Puerto
Rican Bank
Cen t r u r o i d e s n i t i d u s
Anegada
Beef I s l a n d
Big H a n s L o l l i c k
Big Tobago
Broken Jerusalem
Caja de Muertos
Cayo Diablo
Cayo Norte
Cooper I s l a n d
Desecheo
Eustatia
George Dog
Great Dog
Greater Camanoe
Great Thatch I s l a n d
Guana I s l a n d
Hicacos
J o s t van Dyke
L i t t l e Tobago
Magueyes
Marina Cay
Necker I s l a n d
Norman I s l a n d
Palominos
Peter Island
Piiieros
P r i c k l y Pear
Puerto Rico
Ramos
S t . Croix
S t . John
S t . Thomas
S a l t Island
Scrub I s l a n d
Tortola
Virgin Gorda
Gasteracan t h a t e t r a c a n t h a
Anegada
Beef I s l a n d
Big Hans L o l l i c k
Big Tobago
Cayo Diablo
Culebrita
Desecheo
Eustatia
George Dog
Ginger I s l a n d
Great Dog
Greater Camanoe
Great Thatch I s l a n d
J o s t van Dyke
L i t t l e Camanoe
Necker I s l a n d
Norman I s l a n d
Pifieros
Puerto Rico
S a l t Island
Tortola
Virgin Gorda
CATHERINEBURG, ST. JOHN
r
-
20
74
54
53
49
-
- I .=
0
. 60
45
46
CRUZ BAY, ST. JOHN
42
20.
42
33
L
0
m
42
35
46
I
CHARLOTTE AMALIE, ST. THOMAS
52
36
CULEBRA
28
28
L
-
_-',
LC
.-C
m
0
20.
6')
77
61
28
24
I
I
I
I
-
RIO BLANCO UPPER. P R .
174
RIO P I E D R A S . P R .
67
59
132
74
58
0
MONA
48
30
F i g . 2. R a i n f a l l a t v a r i o u s l o c a l i t i e s on t h e P u e r t o Rican Bank, 1960-1967.
Data from
C l i m a t o l o g i c a l Data P u e r t o Rico and Virgin I s l a n d s , Vols. 6-12 (1960-1967). Numbers above
histograms i n d i c a t e y e a r l y t o t a l s .
F i g . 3. T e m p e r a t u r e s a t Magueyes I s l a n d , P u e r t o R i c o , 1960-1966.
Upper l i n e r e p r e s e n t s mean d a i l y
maxima, m i d d l e l i n e t h e monthly means, and l o w e r l i n e t h e mean d a i l y minima. Data f r o m same s o u r c e
as t h a t o f F i g u r e 2.
-
5
+
CRUZ BAY
--
-
CHARLOTTE
AMALIE
-
-
ALEX
HAMILTON
F l ELD
MAGUEYES
RIO PIEDRAS
GUINEO
RESERVATION
F i g . 4. Temperature a t v a r i o u s l o c a l i t i e s on t h e P u e r t o Rican s h e l f d u r i n g 1964. Crux Bay, S t . J o h n ;
C h a r l o t t e Arnalie, S t . Thomas; Alexander Hamilton F i e l d , S t . C r o i x ; Magueyes I s l a n d , Southwest P. R . ;
Rio P i e d r a s , c o a s t a l P. R . ; Guineo R e s e r v a t i o n , upland P. R . D o t t e d l i n e s i n d i c a t e m i s s i n g d a t a ,
d o t t h e August mean a t Guineo R e s e r v o i r . The upper and lower l i n e s f o r Rio P i e d r a s r e p r e s e n t
monthly maxima and minima r e s p e c t i v e l y . Other symbols a s i n F i g u r e 3. Data from C l i m a t o l o g i c a l
Data P u e r t o Rico and V i r g i n I s l a n d , Vol. 1 0 ( 1 9 6 4 ) .
M A R C H
18
M A R C H
19
I966
F i g . 5 . A i r and b l a c k b u l b t e m p e r a t u r e s on Cayo Ahogado, P . R .
Broken l i n e s i n d i c a t e i n t e r v a l s o f more t h a n 1 hour between
measurements. Dots: b l a c k b u l b t e m p e r a t u r e s 1 5 cm above t h e
ground. T r i a n g l e s : a i r , 1 cm above ground. C i r c l e s : a i r ,
1 5 cm above ground. Squares: a i r , 1 m above ground
F i g . 6. Unshaded soil temperatures on Cayo Ahogado. Broken lines
indicate i n t e r v a l s of more than one hour between measurements.
Triangles: a t soil surface. Circles: 1 c m deep. Dots: 5 cm deep
I
.
.
. . . . . . . . . . . . . . . . . .
12
24
12
MARCH
MARCH I 8
1966
IS
I
SHELTERED I OPEN
I
SHELTERED I OPEN
I
F i g . 8. Diagram-map o f h y p o t h e t i c a l i s l a n d showing s p a t i a l r e l a t i o n s h i p s o f numbered v e g e t a t i o n c a t e g o r i e s ( d e s c r i b e d i n t e x t ) t o o n e
a n o t h e r and t o t h r e e environmental parameters.
Three imaginary
s t r a i g h t l i n e s , whose t e r m i n i a r e show a n d l a b e l l e d , d i v i d e t h e
i s l a n d i n t o two c o n d i t i o n s e a c h o f wind e x p o s u r e , wave e x p o s u r e ,
and d i s t u r b a n c e . E l e v a t i o n i n c r e a s e s t o w a r d s t h e c e n t e r . A s
i s l a n d s i n c r e a s e i n a r e a a n d e l e v a t i o n , c o n c e n t r i c r i n g s a r e added
f r o m t h e c e n t e r , b e g i n n i n g w i t h t h e most m a r g i n a l r i n g n o t y e t
p r e s e n t , t h u s p r o g r e s s i v e l y expanding t h e r i n g s a l r e a d y p r e s e n t
t o w a r d t h e o u t s i d e . R e l a t i v e a r e a s a r e n o t t o s c a l e ; e . g . , most
o f t h e a r e a o f a l a r g e , l i t t l e - d i s t u r b e d i s l a n d might b e covered
by m o i s t f o r e s t ( 1 0 ) .
CONDENSATION
PRECIPITATION
WINDWARD
LEEWARD
EXPOSED
SHELTERED
2 4 1 1
2
F i g . 9 . V e r t i c a l s e c t i o n a c r o s s a h y p o t h e t i c a l i s l a n d showing
s p a t i a l r e l a t i o n s h i p s of numbered v e g e t a t i o n c a t e g o r i e s ( d e s c r i b e d
i n t e x t ) t o one a n o t h e r , d i s t a n c e from t h e s e a , topography,
e l e v a t i o n , wind d i r e c t i o n , and wave exposure. Not t o s c a l e .

ATOLL RESEARCH BULLETIN NO. 251 BIOGEOGRAPHY OF THE

Transcription

Similar documents

jacob feitel

Catalogo Eterea 2013