STUDIES ON
THE FAUNA
OF CURAÇAO
CARIBBEAN ISLANDS:
The tree-climbing
No.
AND
OTHER
175
crabs of
Trinidad
by
Heinrich-Otto
von
Hagen
(Philipps-Universitàt, Marburg, D.B.R.)
26
Introduction
Localities,
habitats
and
systematics
28
Grapsidae
Goniopsis
Aratus
33
cruentata
34
pisonii
Metasesarma
Sesarma
27
36
rubripes
roberti
36
i
Sesarma ricordi,
37
Sesarma
38
angustipes
Pachygrapsus gracilis
39
40
Ocypodidae
Uca
Uca
vocator
40
rapax
41
Ucides
cordatus
Gecarcinidae,
Cardisoma
Xanthidae
Discussion
and Coenobitidae
guanhumi
Hexapanopeus
Coenobita
41
caribbaeus
clypeatus
and Conclusions
42
42
42
42
43
Summary
50
References
50
26
INTRODUCTION
The remoteness of crab and tree in the scheme of nature
to
1
by DARWIN,
is
839)
as
much taken for
the lack of relations between fish and
relations nevertheless exist
therefore
can't
mentioned
probably
1939:
Since the
on
climb
mangrove roots
little
"Terrestrial
crabs
into the
The
comparative
Adaptations
9th
cf.
in
Crustacea"
normally
was
HOLTHUIS,
has been
at
tree-climbing
general affinity
ground
or
plants
general
eral
up
remarks
species
on
on
groups
GUINOT,
HILL,
1947)
thus
to
and
1935,
were
of
are
rather
On
observed
to be Sesarma
Milne Edwards
1961), Geograpsus
tree-climbing
a
on
up
terres-
grayi
present
in
tree-climbing
always imply
close
(H.
within
the other
brachyuran
far
scaling
to the
or
upon
1943, for
hand,
inland
trees.
or
sev-
high
Typical
foxi Kemp (TWEEDIE,
(DE MAN,
Milne
paper to
need
constantly sub-
WESENBERG-LUND,
and Coenobita rubescens Greeff
provide
climbing
"more
as
crabs to
progressing very
never
seem
H.
It is the purpose of the
the habit of
that
phytotelmic animals).
1940), S. trapezoideum
un-
symposium
species
regarded
phytotelmata, i.e. micro-pools
of terrestrial crabs
mountains
(Mer-
ascending
1959a). However,
climbing species keep
mangrove roots
inland forms of this kind
&
see
(Amer. Zoologist 9, 3, 1969)
should be
Some
THIENEMANN,
into the
the
was
forms.
adaptations.
stay
shrimp
a
observed
done. The
all, though
of terrestrial
Others live in
(see
be
trees
(SOLEYMAN,
Even
published.
habits does not mean, however, that these habits
merged.
must
("Who
may
coconut
century
Hippolytidae)
work
ground-living
of terrestrial
This
1972).
climbing
tropics)
peculiar
merchant SOLEYMAN many accounts
1970;
(ABELE,
atmosphere
trial" than
Arab
have been
Natantia:
mentioned
hardly
the
as
as
284-285).
tree-climbing
now
early
as
in the
something
POLUNIN,
(Birgus)
(referred
opinion
our
Observations that such
(mostly
reporting
crab
days of the
guia rhizophorae,
til
of
granted
tree.
somewhere
trees?",
the robber
why
REYNE,
the
create
fish
says,
reason
idea
in
1902; FOREST
Edwards)
(GIBSON-
(GREEFF, 1882, 1884).
present
and
a
first review
anomuran
certain basis for future research.
on
crabs and
27
This review
tree-climbing
I
greatly
am
nidad)
for his
lands)
for
tention
indebted to
many-sided
several
menclature used
observations in Trinidad
by
(West Indies),
where several
occur.
identifying
to
The
induced
was
crabs
Prof.
that
2
in Table
visit to Trinidad
mention
and for
(1965/66)
Stiftung Volkswagenwerk,
KENNY
and to
of the West
(University
Prof.
L. B.
material collected
the crab
papers
S.
J.
assistance
HOLTHUIS
(except Uca),
reading
and
made
was
the Deutsche
the
criticizing
possible
for
for
tree-climbing crabs,
Tri-
Nether-
drawing
my
the
correcting
at-
no-
manuscript.
financial
by
Indies,
(Leiden,
of the
support
Ibero-Amerika-Stiftung and
the Deutsche
Forschungsgemeinschaft.
LOCALITIES,
General remarks
HABITATS AND
tions
1975).
of
a
given by KENNY,
are
The data
year
A
given
angustipes,
bita
devoted to the
in Table
the author,
from
give
The numeration
coasts
Tr
S~Tr
adjacent
(1970)
17
of Balandra
Tr
Mouth
of L'Ebranche
5
g
10
13
1
seen
Coeno-
climbing by
carapace
or
at
least
phytotelmic
width listed in
each
case
to
the east
coast, the
Details
VON
HAGEN( 1970b).
following
on
ones
to
pertinent regions
the
are
(1975).
of localities :
of
Tr
gracilis,
not
Trini-
Sesarma
caribbaeus,
of observation is taken from
refer
and STONLEY
Mouth
Tr
were
of
species
(Pachygrapsus
to the Gulf of Paria.
Mouth
Tr
crab
course
(genus Uca).
certain idea of the crabs' sizes.
Tr 6
Tr
collected in the
tree-climbing
tree-climbing
as
of the localities
north-western
BACON
were
of fiddler crabs
in Trinidad but
occur
a
figures
given by
study
(short
1970b and STONLEY,
guanhumi, Hexapanopeus
known
are
Identification
List
the
literature. The maximum
is meant to
HAGEN,
tidal
introduc-
nevertheless included in the table and in the text,
were
they
omitted here
VON
Five species
1.
Cardisoma
that
clypeatus)
.
because
Tr
of
systematic synopsis
dad is
are
in this paper
reported
mainly
1969,
vegetation and
climate,
topography,
on
pattern of the island of Trinidad
SYSTEMATICS
j
23
Salybia
L'Ebranche
Nariva
River
River
Highway
(east
Canal
of
Toco).
River.
River.
Bridge
from Nariva
Mouth of Lawai
Entrance
River
of Eastern Main
Bridge to
Road
near
Lower
Manzanilla.
the mouth.
River, Guayaguayare Bay.
(East-West
and Blue
River,
Trench
Caroni
no.
Swamp.
9)
between
Princess
Margaret
28
Tr
Blue
24
River, Caroni
Tr 26
El
Tr 27
Main
Tr
Cocorite
31
Socorro
waste-water
Tr 32
Mouth
Figure
8
Diego
outlines the
main habitats.
(indicating
Swamp,
of
Swamp.
Caroni
Entrance,
east
main
Diego
Swamp.
Martin
River.
River.
upstream
Species occurring
their
of
Martin
Swamp.
Laventille
ditch,
distribution of
in different
the
zones and
crabs
mentioned
habitats
are
listed
and
only
their
once
occurrence).
GRAPSIDAE
The
gories
a)
seven
of
species
known
Species
Species
known
frequently
seen
(Aratus pisonii,
c)
Species
climbing
S.
to be dealt with fall into three cate-
not
in
tree-climbing
as
only ( Pachygrapsus
b)
grapsids
:
gracilis,
tree-climbing
as
on
trees at many
or
known
as
Trinidad at
in
from literature and
Trinidad
Metasesarma
tree-climbing
restricted
literature
angustipes).
tree-living
places
Goniopsis cruentata,
from
phytotelmic
or
Sesarma
from
localities
by
the author
rubripes).
literature and
only (
Sesarma
found
roberti,
ricordi).
Fig.
8.
Upstream
distribution
and
habitats
text
A
B
C
D
Mangrove
swamp
Supralittoral zone
Laguncularia
Bamboo
with
of
the crab
species
mentioned
Rhizophora mangle,
sandy beach,
racemosa
wood.
of
(schematically).
mangroves
and
grassy
outskirts,
in the
29
The
cies
following
GENERAL
list) refer only
All of
them
tain
closer view of their
a
mentioned under
species
kept in terraria
were
(preceding the annotated
REMARKS
to the five
in the
in order
laboratory
spe-
(b) and (c).
to ob-
climbing behaviour.
DIET
In addition to
captivity,
of
mens
All
observations of
each
species
(except
species examined
tivity,
they eagerly
ever,
these additions
achs
contained
with
grains
were
feed
seem
only
or
were
some
of
riety
freshly
cruentata)
mainly
dead crabs
algae
wood,
black
rubripes,
tissue
those
or
speci-
examined.
rarely
field,
as
how-
all stom-
mostly macerated plant material together
and
brown
plant
algae.
opaque
of Metasesarma
Only
one
stomach
and 3 out of 9 in Sesarma roberti
material consisted of
In
two
rather uniform: The stomachs of Aratus
and
In the
to be obtained rather
insect remains. The
angiosperm
were
herbivorous. In cap-
fish.
or
the field and in
killed adult
of fine sand and indefinable detritus.
out of 13 in Metasesarma
yielded
Goniopsis
found to be
on
in
feeding crabs
the stomach contents of 9-13
pieces
species
pisonii yielded mainly
of
decayed
contained
rubripes
a va-
the remains
mainly
mangrove
very small
roots and various leaves.
Only Goniopsis
cruentata
been recorded to feed
1963 and
HARTNOLL,
grove roots
fallen to
attempts
species
in
it
as
1975)
well
and
ground (WARNER,
to catch
A.
was
seen
be
to
seems
truly omnivorous. It
young mangrove shots
1965)
(STONLEY,
the
In Tr 31 G. cruentata
thermore,
on
pisonii
was
as
to
on
(SCHÔNE
sponges
prey
on
&
growing
living
when
being
observed to prey
on
on
a
tree
on
Aratus
1967) though, apparently,
(Pl.
man-
pisonii
it
never
I, below).
Uca maracoani. Fur-
killing and eating lesser individuals of its
captivity (cf. frequent
has
SCHONE,
cannibalism in the related
own
Grapsus
grapsus ;
KRAMER, 1967).
30
BODY
The main
is
with the
on
low
the
species
mentioned
When
ground.
LOCOMOTION
brachyurans (LOCHHEAD,
able to walk
temporarily forward
A.
speed. Only
the crabs
of their
speed
though
pisonii)
normally
side-
body, i.e.
1961: 330), they
howare,
diagonally, especially
or
pisonii clearly prefers walking
substratum and in any
Aratus
(except
being there,
typical brachyuran orientation
all
ways. Like
ever,
of all
activity
performed
run
ORIENTATION DURING
still
being
forward
able to
on
at
any
walk side-
ways.
In
body orientation
climbing,
of inclination of
from
and
Aratus, walking sideways
times
and
the tree etc.
vertical trees
even
those with
tion of the
body
is
and
chosen
on
is abandoned
on
trunk
or
degree
big inclined and
Apart
some-
(Pl. I, above)
Normally
vertical trunks and
crab is forced to embrace the root,
to the
diameter.
horizontal branches
secondary twigs.
numerous
to its
according
is retained
on
according
this
as
orienta-
soon as
the
branch because of their
small diameter.
In
9a).
ascending,
When
taken forward
The four
When
9.
or
climbing
Body
(right)
in
from
cruentata
the
always
ground,
it
climbs
turns
backward (Fig.
after very
few steps
sideways.
remaining
orientate their
Fig.
Goniopsis
starting
up
a
body)
stick
prefer
in
to
go
experiment,
when the stick
orientation of
(a) Goniopsis
species
up
head first
they
will
is turned. But
they
tree-climbing grapsids. Ascending (left)
cruentata,
(b)
the
4
species
of the
(Fig.
turn
are
and
(i.e.
9b).
re-
able to
descending
subfamily Sesarminae.
31
ascend backward when chased after the
in
terity
Aratus
decreases in
however,
performance,
pisonii, Metasesarma rubripes,
None of
the
mal escape response therefore consists in
fleeing
the
up
tree.
loosen their hold and
trying
to
hide
close
On
jump
there
a
-
fall
on
tors
soft
however.
(e.g.
tioned
At
ground
Fig.
10.
escape
do
or
a
not
nearly
response
on
get
to
nor-
trunk
or
always
into the water
these responses
exposed
preying
crabs
ground
of
roberti.
hurt
of
tree-
remains
when
they
ground-living preda-
Aratus
pisonii
as
men-
above).
When
of
(drawn
cruentata
in
legs (left
black
joining the
(e.g.
tree crabs
Colomesus
being unmolested,
Goniopsis
set
one
cruentata
high tide, chasing
JI.
normally
but may become
Goniopsis
vide food for fish
Tr
Crabs
common
The effect
behind
hiding
approach
Dex-
following order:
great speed. The
the
fall to the
very
climbing crabs (see below).
ambiguous,
or
the
Sesarma ricordi, S.
is able to descend with
species
of the stick.
turning
white
and
black
ascending
and
not
lettered)
set,
psittacus)
all
a
species
-
will
the crab
move
as
are
Laguncularia
lettered)
which
behaves
into the water
is
next.
can
was
mean
experienced
During
supported by
L
in
able to climb down
tree.
independently. (Drawn
to pro-
5
movement
the other
(stippled),
from
a
instead
photograph).
of
set
of
32
for
considerable distance. This is
right
in
9a
on
the
to
descend backward
ward
seems
to be
few
a
normal
centimeters.
way
REYNE,
of
Aratus
perched
neath branches.
in
Birgus (HOLT-
discussion
a
once
down back-
of differ-
of
Use
OF
chelipeds
tions of their
to the
as
inverted
When
species
an
which
body
during
can
of
of
independent
The
(Fig.
climbing
(instead
(with
legs
They
as
-
tata)
R2
the
-
down.
underthe
all
of
gait
backward.
of
quartets)
L 3 -
Meta-
display
rota-
held close
are
the crab
tend to
posteriorly
as
well
stepping
normal-
moves
in
as
stretch
at
least
("froglike",
the
order
see
chelipeds
than
the
are
other
R4
latter set
-
ORDER
known for
through the change
or
and
down).
species
normal
as
descending
however,
are,
soon
its head
ambulatories
the
pisonii
pisonii the chelipeds
of
first
and
arboreal
perform
even
pair
But
locomotion there
of
hanging
10).
general type
forward
head
AMBULATORIES
Aratus
except
STEPPING
is retained
sit
position without
ascending
forward ascent.
head
pair
HINDMOST
Fig. 9b). These hindmost walking legs
peraeopods
to
found
were
during
In A.
orientation
ascending
AND
species
additional
hindmost
more
prefer
rubripes
support
as
for all
chelipeds.
carapace
used
all
able to walk in this
are
CHELIPEDS
rubripes,
sesarma
tree,
a
Both
compulsory
seems
their
on
chelipeds.
USE
LI
for
(drawings
observed
Coming
descending
1939: 300,
and Metasesarma
pisonii
aid of their
to
was
opinions).
When
ly
cf.
b). Only Sesarma roberti
for
the
1959b: 309;
HUIS,
ent
Fig.
done head first
always
are
of
as
the
(drawn
In
in
(LOCHHEAD,
chelipeds
normally
legs, namely
L5.
crabs
1961
:
331)
walking direction from sideways
Figure
Rl
10
black and
two
-
participate
can
alternating
L 2 -
R3
-
L 4 -
quintets
R5
(ascending Goniopsis
lettered)
is
in
and
cruen-
giving walking
33
whilst the former set
support,
Crabs
body
teresting
can
of
alter
cruentata
from
one
it
side
thus
first
stepping
head
1-3-5-2-4
dangerous
to
Goniopsis
pull
G.
G.
1
-
the
down
species
It
can
in
head
(i.e.
legs
front
It is not
descending
the
sequence
be
may
more
“Mangrove Crab”
1802/3)
of this genus
tree-climbing
La-
down.
way
in
In
only
cautiously.
pic-
a
(when taking
legs leading.
use
legs
order
stepping
preferred.
to
seem
(Latreille,
1947: 466;
on
cruentata ascends backward
species
push oneself
cruentata
was
trunks and
(Table 2)
literature
1958:
GERLACH,
1963: 643;
seen
G.
cruentata is
(RATHBUN,
673;
1918:
RODRIGUEZ,
1967:
WARNER,
usually less than
ground.
On
within
hiding
1921:
RATHBUN,
Tree-climbing
and
greatest
about
night)
m
2
for
G.
from
but
m
G.
238;
1959:
322; STONLEY,
(WARNER,
were
of
the
perched
up to
4
observed
the mangrove
area
on
1969). Frequently
periodically
mangrove forest
individuals
the seaward
the individuals
Rhizophora
of
Laguncularia (Pl. I,
m
above the
climbing
LANG
(cf.
on
in
pelii).
This corresponds
grove swamp.
by counting
trunks
sometimes
crabs
in
and
branches of
cruentata occurred
flooding
of
on
these
stone walls
444,
of
number
100
even
occasions
two
prop roots
on
(rarely)
above),
(day
vertically
the
way up
in-
325).
on
and
descending
the
motion
mm
push itself "upstairs" with the
all
266; SCHONE & SCHONE,
and
the
on
themselves
Like the two other
DANSEREAU,
and
few 8
a
of the two
is
sequences
down)
as
of
their
direction. It is
sequence of
stepping
analysis
itself up with the hind
cruentata
well known
1975:
An
that
either
pulling
the
addition,
only). As
one
orientation of
same
3- 5- 2- 4 to 4-2-5-3
-
can
or
known whether
(likewise
1
the
lettered) is moving.
change the stepping
ascending
trunk shows
changed
down)
in
that,
during climbing.
guncularia
account
down with
used to
course,
to note
tures of G.
be
and
climbing up
are,
white and not
(left
fringe
to the
to
the
pattern
the floor
the crabs
was
at
during every
high
always
tide.
seen
interior of the
of
occurrence
during low tide
"wait" until
in
The
within
man-
found
Jamaica
the tide is
ac-
34
tually coming
the
in.
of the trees.
head
When
crabs
Juveniles
hidden
the
on
of
ground,
steep
vated
least
spend
A.
of
roots
individuals
Unlike A.
Aratus
pisonii
A.
M
.
being
Milne
(H.
a
mentioned its
climbing
but
truly tree-living
of
There
Pacific
deren
the tide has receded.
of
high
the
stay
juvenile
to
the
to
be
but
ground,
there
of crabs of different
to
seem
from the
be
algae
however,
was,
G.
cruentata
moti-
coat of
observed
no
on
never
“Tree
tree-climbing
in
seen
die auf die
It
benagt".
—
at
individuals
spend
others
most
is
keep
in
the
of
speaking of
Manglebusche
not
least
hide under
Crab”
crabs. F. MULLER
abilities when
fully corresponding
mangrove faunas.
was
1837)
species
and
however,
(occasionally
seems
always
overhanging
under
excursions
special
Blatter
mangrove trees
The adults,
trees
water
dead recumbent trees.
fully phytotelmic Metopaulias depressus
in the sea, and
spent
of
tree.
Edwards,
und
like in the
keep
rubripes,
on
steigt
a
trees
pisonii)
is the best known of all
already
neigh-
the burrows
into the open rather soon,
(see above). Feeding
(cf.
aggressive
an
climbing activity
zophora)
grove
and
I, below)
(Pl.
during high tide.
"allerliebsten, lebhaften Krabbe,
ground.
rising tide, sitting
hide in
periods
these vertical
and
pisonii
pisonii
tions
floor of
rush to the roots
only.
females when
(1864: 20)
is
the
clear of
river banks
some
Rhizophora
courting
return
or
under the bark of
areas
search of food
by
younger
an
they
considerable up and down
At
and
water
not reappear until
e.g.
open
the
low tide most individuals
During
the
do
found to
Individuals from
portions
human observer
a
Normally
individuals
some
sizes.
ahead of the
the intervals
during
into
jump
cordatus.
were
a
from
escaping
the
Ucides
is
down
of the
patches
well away from each other.
keeping
but
crowd the last
may
They ascend gradually
motionless and
bour
They
yet submerged, until they finally
not
swamp
only
(Rhi-
a
tree-
when adult. Unthe larval
period
to the lowest por-
the bark
their
vicinity
form in
or
time
of
on
on
the
man-
mangroves).
the Indo-West
35
We
are
well informed
and sexual
maturity
standing
detail than
on
leaves
habits
no
doubt that A.
pisonii
dad
(see above),
it becomes
crabs and
adult
ameter of root
activity
of
portions
aerial roots
with
for
noted
by
than 5
mm) only.
algae
main food of
climbers
cayed
117),
are
for
species
the
on
was
the
always
top
occasional
of
etc.;
sponges
This
feeding
Components
of
and the
individuals of
which the
those
grown
BACON,
see
(less
encrusted
see
with
tree
a
(Pl.
1958:
apparently
in
6
specimens from
detritus-
699,
and
forms
the
period.
at
least half of the crabs
bottom)
I,
m
above
climbing activity
(about
of
coat
GERLACH,
which
trunk
behaviour
1970,
also
data).
occurred
intraspecific
the lower
trunk)
as
m
high).
taking
up
tissue from wounded mangrove roots and in
faeces of other Aratus
a
feeding
but ascribed to smaller crabs
certain
the
(di-
not
was
branches within 2
a
tree
day,
a
from
well
as
of this kind
shaded
and
nearly
IV, centre),
(Pl.
of
When
population (about
from the
the low tide
feeding periods
the field.
receded
ones
names),
A. pisonii during
perched
bottom and
"bare"
barnacles,
(1965:
underneath the roots
there
the
(1965).
always "form
branches). Feeding
("bostrychietum",
Between two
man-
A continuous
leaves.
on
moist roots
All roots
47,
24 for
extending
from
and zonation
HARTNOLL
1970:
on
from Trini-
Rhizophora
one
only when the tide
(oysters,
names
BACON,
though
m) in Tr
found to feed
was
uncovered,
covered
rested
about 5
crowded the
epizoans
scientific
to
more
The examination
116).
of
ones)
young
(those dropping
just
outor
in
HARTNOLL
specimens
into observations in
the prop roots (those
various sizes
tide had
some
observed
was
and
the activities of the Aratus
system
single individual
pisonii
is able to feed
(1864)
1965:
(HARTNOLL,
continuously recording
20
A.
dubious whether leaves
contents fitted well
stomach
of
the stomach contents of the
food"
principal
MÛLLER
reported by
as
considering
its
locomotion
tree-climbing
to
respect
1967,
some
30.
p.
But
in
understand
to
attempt
features
specialized
There should be
grove
been little
morphological
consider the
growth,
dimorphism (HARTNOLL, 1965; WARNER,
has
1968), but there
breeding, larval development,
on
hanging
or
the
ground,
between the
Among
portions
picking
these
of
up
dethe
the bark of the tree.
behaviour in A.
pisonii
are
described
36
(1967,
WARNER
by
and
1970). Display
copulation
take
place
the
on
mangrove trees.
rubripes (Rathbun,
Metasesarma
The
S.
roberti)
Table
of
(1963: 201)
lower trunk
Climbing
and
during
or
other
pisonii),
females
others
on
able
species
all
roberti
flooding
are,
the
usually
entrance
of
dug
en-
oval-shaped
hidden under recumbent
ground.
Climbing
prolonged
G.
seen
arboreal
inundation of
of the climbers
rapax). Many
(cf.
was
of
cruentata and
were
A.
courting
seen
in G.
not the
only situations causing
cruentata, there
was
a
consider-
tide.
M.
rubripes
comes
closest
to the
pisonii.
Milne
hides
into
with
high
able to
are
largest numbers
of
of
periods
crabs
Some males
however,
at low
A.
and the
branches.
mentioned,
grapsids
galleries
(cf.
Uca
during
the
or
during periods
to climb trees. As
Sesarma roberti H.
S.
the
trees
(Pl. IV, left), mostly
burrows
the
on
though
remained active.
climbing dexterity of
within
occurred
self-dug
crabs
Paspalum
grass
ground.
motionless head down
climbing activity
From
above the
night,
at
as
the
under
mentioned in
Laguncularia
Apparently,
rains
Laguncularia
Periods of
this
m
material
heavy
sitting
well
as
briefly by
(see below,
localities
ascended
the banks of rivers
counted
were
found
all
and coconut trees
6
their
tides
PEARSE
daytime
frequently
plant
high
river banks after
were
as
within
near
normal
individuals
far
regularly.
not
submersion
entrances in
trees
as
activity
though
In
of different sizes
of Bactris
region
rarely
up to 3 m,
in
frequently
Tr 26.
mentioned
already
probably by
stems and branches of
serpentinum, the
dure
in
and
sexes
rubripes,
and
observed
was
1, especially
both
water
of M.
tree-climbing
RODRIGUEZ
1897)
Edwards,
under
steep
the burrows
logs
clayey
1853
and stones
banks
on
the
of rivers
being oval-shaped.
To
ground
and
reach
or
ditches,
the
en-
37
trance the crab often has to climb up
HOBBS
by CHACE &
(1969:
almost vertical surface
In the two
these crabs
close to
or
the stumps of
the
cut
bigger
and had
11, below)
m
node.
Several others
and
canes
As
Tr
TO).
seventh from
climbing
examination of
third of the 9
It
specimens.
can
fragments
be
no
Sesarma ricordi
1853
During
this
daytime,
water
S.
mark,
ricordi
At
J
)
or
more, the
roberti.
2
)
is
It is
utilized
dornys
the beach
from
it
carapace
the
were
trees". But
seems
They excellently
by roosting
gliroides
entrance holes
and
seen
that
as
fit
the
in
an
loose smaller
opened
of
in
that the bamboo
time. 2 )
This
the tidal
roberti
crab
of
M.
refer
rubripes
species.
in
about
Tr 32
over
"had burrows
to S. roberti.
fit those of
the
The
two
the
among
Further-
subquadrate
instead.
the internodal space of intact bamboo
and
the
be
may
zone
"walked
crabs
actually
Tylonycteris)
1969).
from
cryptic-coloured
only.
PEARSE do not
longicaudatus).
(MEDWAY,
yielded
(all
“Beach Crab”
that these
his notes
Malaya
up to
inter-
stones and coconut husks.
Colombia S.
proportions
(two species
Hapalomys
themselves
forming
contents
same
and
foraging
given by
note that in
bats
open
top
less than 8-10 different
the
night
he remarks
dubious whether
measurements
interesting to
on
of
to
access
thus
the
some
maze
along and somewhat above high
at
cover
A
little is known of the behaviour of this
(1915: 553) reported
climbed in
the mangroves"
S.
crabs
night the
PEARSE
roots
far
why relatively
reason
relatively small
driftwood, leaf litter,
among
moves
on
Edwards,
10
of the stream.
therefore,
at
keeps usually hidden
fact that
lateral slit in
stomach
of
31)
10,
in Tr
only
were
specimens
concluded,
Milne
( Tr
facilitating
among
provided moist hiding places and food
H.
by the
a
the action
in
the
)
taken out of these stumps
were
seen
One male contained
1
seen
and
hide within the stems.
filled with rain water
exposed by
one
but
was
cliff
in Trinidad
Several stumps
above,
fragments
insect
were
upon roots
mentioned
insect
a
bridge.
forest,
11, above)
(Pl.
partly
above the lowest roots,
a
considerable time ago.
a
stems.
phytotelmae. Six individuals
2
bamboo
climb up and to
about
lay
canes
a
a
have been induced
may
bamboo had been
loose smaller
(Pl.
of the cement wall of
found to
were
This local behaviour
of the
the wall of
186) scaling
main localities of observation
S. roberti lived within
S. roberti
vertically.
nesting
latter,
tree-mice
however,
canes
(Chiropo-
gnaw
their
38
climbing (1-2
and
m
ground) in large numbers the branches
above
of dead trees and shrubs washed ashore
and
Sesarma
the surface of the stems and branches.
nocturnally ascending living
Sesarma
S.
S.
was
not
ricordi
hid
found to
phytotelmic
LER
live in
ascending
seen
& SATTLER
for synonymy
crab
the
(1965),
night.
in
trees
those
S.
KENNY
2
deep
*)
A
now
)
by
A
single
)
The
meliads
S.
this
SATT-
by
(1969;
marine
bromeliads
boulders in each of
on
male
from
(1972
of the
husks
1918:
Dana
lost
Parahyba
as
found "on
of
324)
so
do
that
the
collected
(ABELE,
1972a)
to
way
near
in
to
Ger-
remark
"on
of
a
roberti).
Brazil
obviously
with the
mouth
inside the walls of
with S.
But
pisonii
angustipes
the
(USNM 25712)
miersii” and later identified
as
this male
same
was
collecting
mangroves"
may
.S.
only
data
be
men-
angustipes
after-
(USNM
only
a
de-
species.
at these
a, b)
their
Trinidad S.
Norte, Cabedello,
habitat of this latter
crabs
in
leaves
palm
mangroves".
of Aratus
by ABELE (1972
that
(together
“Sesarma
apparently,
possibly belonged
on
collected from
was
the
wanting.
sure
and
Tr 31
specimens
typical
stated
angustipes
Rio
was
9
conformity
was
be
only
(1918: 304)
a, b)
found crabs in
and,
(Cocos Bay)
The crabs he collected
single specimen
25713, RATHBUN,
2
can
coconut
wards isolated from
scription
The Brazilian
unfortunately, they got
we
RATHBUN
ABELE
From
"nickel-size
as
growing
were
Trinidad
abandoned well in
tionedby
finally
(1975).
ZAHL
confirmation is still
under
rivers.
and
1972b)
(personal communication)
in
trees.
on
But,
that
so
Until
Brazil.
“Sesarma ricordi ”
as
is
species recently
)
J.
growing
occurs
reported
others)
cruenta and
species.
many
of bromeliads in
but
Trinidad,
this
“Sesarma miersii ”by MCWILLIAMS
ABELE,
see
tanks of bromeliads
this
as
on
seen
mentioned.1
cases
Prof.
tanks
was
Sesarma)” by
(genus
(Neoregalia
the
habitat it
never
and climbed
during daytime
nevertheless taken into account here, because
was
was
1852
angustipes Dana,
angustipes
from
particles
mangrove trees.
always
the walls of the terrarium at
twigs and
on
these crabs
captivity
small
on
Though sometimes found
within the mangrove area,
sandy ridges
In
(cf. Metopograpsus latifrons
2). They fed
Table
guttatum,
sites
with
those
living
who finally cleared
the
up
and examined
part
of
my
in
Brazilian bro-
taxonomic
material from
status
of
Trinidad,
39
Though
true bromeliad
demic to
bromeliads in
HARTNOLL,
1964).
lasting only
water
ambulatories.
a
S.
individual of
S.
Sesarminae
thus
could not
Rathbun
be
or
digging
a
ground,
even
epiphytic
angustipes
shallow
However,
identified with
tufts
and
ad-
are
of hair be-
dense
development
When
dig.
in
as
of
the
the tufts
in
kept
excavated the
ter-
a
under
ground
pit.
at
Like
the
least
a
third
of
species
Metopaulias depressus,
only specimen known
certainty
S. cookei Hartnoll ;
1961 ;
its whole
the third and fourth
long
as
able to
in bromeliads.
Jamaica.
en-
non-feeding phase
a
i.e.
third and
mentioned that there is
occurring
found in
the
where maximum
ricordi,
is
(LAESSLE,
m
phytotelmic throughout
nearly
are
is
It should be
was
uptake
angustipes
hiding log
which
and
S.
the
by
1964).
from
These hairs
related S.
an
above 250
condensed into
of the second and
coxae
is reached.
depressus
reached
striking adaptations of S. angustipes, however,
to
the
rarium
.
is
stages being
bromeliads,
of
saxicolous
large
area
days (HARTNOLL,
3
The most
aptations
closely
M
tanks
adaptation
depressus Rathbun,
restricted to
the limestone hill
the zoeal
cycle,
tween
and
the
the state of
crab, Metopaulias
Jamaica
in
to live
is very far from
angustipes
life
able
apparently
until
now
cf. HARTNOLL,
so
it
far
( Sesarma jarvisi
1964:
164,
and
1971
:
259).
Pachygrapsus
P.
gracilis
among
a
gracilis (De Saussure,
was
local
collected
by the author in Tr
of
aggregation
mangrove area, associated
species
is
gracilis
in
grove
sent to
part
by
included
which
Washington by
of the material
Miss
(now
Prof. L. B. HOLTHUIS
(1968).
hang
the
31
only. It
seaward
HARTNOLL
algae
and
of
fringe
114)
the
This
(Gibbes).
(1965:
sponges
found
was
saw
covering the
P.
man-
into the water".
M.
STONLEY
in Leiden
Museum)
J.
at
the
with Petrolisthes armatus
Jamaica "among
roots
stones
because
here,
1858)
(Leiden)
in
(ABELE,
was
1972a:
already
168;
identified
1966 and mentioned under this
1972b:
as
S.
123).
Another
angustipes
name
by
VON
Dana
HAGEN
40
OCYPODIDAE
The two
rather
are
species of Uca (vocator
related.
closely
They
rapax)
and
belong
both
to be mentioned here
to the
"jerker group"
of New World fiddler crabs. A third member of this group, U. brevi-
frons,
was
found
U.
species,
on
390)
1970b:
cordatus,
Uca vocator
(Herbst,
area.
not
is
In
contrast
feeding
crabs
fig.
They
9).
to
were
Uca
that
species,
found
floating
likewise
ascended
was
CHACE &
only
captivity
transferred from
HOBBS
Incorporation
species
be
can
driftwood
on
below,
(1970).
most other
Males
compulsory.
in
into
(1969),
a
of the
one
1804)
of the
one
A fourth
(see Table 2).
branches
Ocypodidae by
still wanted.
vocator
CRANE
ascended
TÙRKAY
adopted by
by
mentioned
subfamilies is
U.
tree
and is omitted here.
briefly
the Gecarcinidae to the
view also
a
Rathbun,
thayeri
(VON HAGEN,
Ucides
perched
seen
on
invade the
possession
displaying
ditches
waving
on
above
of
the
the
lested,
up
or
tree,
a
When
ground.
of
1970b:
of
stump
these males hid in slits and holes of the wood
cm
herds
(VON HAGEN,
they
20
burrow is
a
among
which
to
freshwater
mo-
underneath
the bark.
females
Ovigerous
around full and
of
nearby
larvae.
females
canals
are
gather
edge
and
molested
ascend
Laguncularia
Tr
until
the
a
(especially
on
Laguncularia
the banks
hatching
few males.
by egg-stealing poeciliid
(VON HAGEN,
of
their
As
these
fishes and
1969), they keep close
and
Rhizophora
to the
mangroves,
III).
individuals of U.
in
periodically
large numbers
normally joined by
are
(Pl.
pools
in
tidal influence
in Tr 23 and 26
A few
below.
stagnant
alternately
when the tide rises
of
from
moon)
with
The females
grackles
water's
new
31
vocator
are
that
were
mentioned
observed
together
on
with
the stems
U.
rapax
41
Uca rapax
(Smith,
1870)
Displaying males of this and other species of Uca tend
small elevations of the ground
trance of their burrows
males of
32)
not
More
inland
branches
the
rain in
June.
covered the
was
than
more
1
had
cm
On
in
an
period of heavy
a
and
rapax
U.
higher ground
a
few
U.
(Pl. IV, right).
the total
high
cm
However,
ashore
continuously flooded
above water level
50-60
en-
ground.
was
U.
the
In two instances
the inhabitants of
by
of
to select
near
washed
during and after
ground
m.
been
above the
achieved
individuals
found up to 40-50
were
pellets
shrubs instead.
use
7> 31, which
the river
Numerous
the water
height
grove
from
water
cm
were
in
to
(which
than about 5
higher
Laguncularia
vocator
observed
were
of mud
heaps
special courting places.
as
impressive heights
by fresh
As
rapax
ascended
they
Tr
U.
or
rapax
climbing
ascended
tall grass.
"arboreal
The
crabs
fed
on
motionless,
in
some
the so-called
head first
with
just
its
as
the
males
on
the water
Ucides
still
cordatus
Some
within
was
approach,
display
perched
of
the
in their sockets
still
leaped
big
reappeared
moving
came
perched
They
after
tried to
slowly
down
dodge
around
water.
In
preceded this
usually
a
(i.e.
ascended
fiddlers behaved
down into the
claw
the
individuals sat
while.
On
the
third
high.
(Linnaeus,
flooded
above). All crabs
the
Some of
days.
perched crabs had drastically decreased though
medium-sized
the
molested,
mentioned above.
threat
the number of
the
One male
backward.
When
closer
two
only
most of
eyestalks laid down
(more rarely)
and,
for
condition"). Crabs
"sleeping
Some individuals
retreat.
day
bark,
grapsids
a
but
with the
head first.
the trunk
the
or
lasted
period"
moist
1763)
individuals
mangrove
seen
kept
a
of
area
U.
cordatus
together
with
ascended
U.
rapax
trees
(see
few centimeters above water level.
42
XANTHIDAE AND
GECARCINIDAE,
brachyurans Cardisoma guanhumi
The
baeus
well
as
tioned
the
as
because all information
together here,
haviour
was
in Colombia and
in
perched
Florida.
CHACE
fig. 20-22)
to 3
m on
&
of
a
reported
(1962: 219)
HOBBS
(1969:
men-
climbing be-
males and
by
females
(1915:
PEARSE
and STEPHENS
found this
197)
temporary bridge
in
(1965:
species
Dominica. A
to climb
more
(1975:
HENNING
284,
regularly
Prosopis juliflora feeding
trunks and
reed, palm
observed
recently been given by
detailed account has
who
was
GIFFORD
by
the braces
on
their
1825
Latreille,
guanhumi
of C.
Tree-climbing
201)
clypeatus
on
are
obtained from second hand.
Cardisoma guanhumi
553)
carib-
Hexapanopeus
and
Coenobita
crab
anomuran
COENOBITIDAE
on
up
leaves
and insect remains.
Hexapanopeus
This
caribbaeus
small xanthid crab,
and stones
in
crustations
on
the
This
species
where
it
hides
people reported
to
in the
during
have
corresponds
in
(PHILLIPS,
simply speaks
other small
found in
of
Trinidad "in the
STONLEY
area
day
these
of
and
1964:
near
5,
(1975:
en-
38).
north-eastern Trinidad
is
active
"Soldier
to similar observations
trees,
plants
coastal
the
seen
by
from under driftwood
1791)
(Herbst,
trees. This
Florida
was
the mangrove roots"
occurs
1871)
usually collected
tidal zone,
Coenobita clypeatus
only,
(Stimpson,
HAZLETT,
HAZLETT writes of
the shore".
in
at
Crabs"
on
litt.).
the
night. Local
perched
same
While
on
species
PHILLIPS
"mangrove rootlets and
43
DISCUSSION AND
In order to
ing
get
Table 2.
the earlier
are
certain
a
without
reports
on
1963:
94,
references
It
can
climbing
other
lack
of
in
Trinidad
members.
species
1939)
remarks of HARMS,
2,
that all
crabs which
systematic
to
(see
Most
were
tata)
were
however,
of
and
observed
these
forms.
appears to be
climbing habits
mals involved
some
are
of observers in
names
to
trees.
by
the word
noun
on
Finally
groups
these
Goniopsis
In
species
species
cruen-
"trees",
for rather different kinds
"mangroves" is, of
considerations
necessary.
certain
Some of
various authors. The term
collective
a
are
The
tree-climbing.
to
Sesarma rotundatum and
pisonii,
named "tree crabs"
Therefore
omit-
containing
introduction)
predispositions
grades of affinity
vegetation (though
ing).
of
1929) could
were
groups
only terrestrial
or
many
addition, the accumulation of
(especiallv Aratus
were
many further
Certainly
names.
Typically
could share certain motives
different
in
compiled
the Trinidad
prevalence of Sesarminae (15 species) is impressive. All these
reveals
(in-
tree-climbing crabs (e.g. BERRY,
many
belong
contain
that
groups
trees)
were
of relevant references.
on
species
read from Table
trees
climbing
mostly
accounts
climb-
the author's attention.
escaped
be
than
synopsis,
repetition
"grapsoids", and
on
included for
other
on
data from literature
Birgus (reviewed by REYNE,
intentionally. Other
not be
typical, pertinent
vegetation
on
achieve
To
included, though
ted
impression whether the observations
an
crabs in Trinidad
cluding climbing
CONCLUSIONS
different
some
course,
phyletic
prevail-
routes
to
consequences to the ani-
should be discussed.
MOTIVES
It will
at all
in
not be discussed
(see PEARSE,
1950)
here
why animals emigrate from the
and that intertidal and coastal
vegetation
general favour this evolution. We shall restrict ourselves
tain motives
apparent
sea
to
cer-
from the behaviour of the crabs observed in
Trinidad and elsewhere.
44
1)
ESCAPE
tidal
FROM
and
zone
burrows
flooded. The
West
rubripes,
sarma
many
for
190
of the
hours;
in
come
trees
under
air
the
trapped
salinities
ESCAPE
guanhumi
vegetation
FOOD.
1902).
eral
Uca
Other
species
patches
of
moss
Aratus
of
or
feeds
284)
1967:
on
feed
on
rapax
and
in
this)
or
thids mentioned
718).
climbing
predators
they
are
not
inundations. Cardi-
Ucides
lichens"
the
cordatus,
how-
Irom low
Escape
case.
(observed
roots.
in
Go-
of material of the
bark. Even blossoms
reported
as
food of
of the Marshall Islands
or
rotundatum
epizoic
were
cover
(the
Goniopsis
(HOLTHUIS,
locally
gracilis
of still other encrustations
may eat parts
1953:
on
on
and
Sev-
"green
from
bostrychietum)
at least
Pachygrapsus
climbing
(SCHNEE,
encrustations.
taken
bread fruit trees
algae
Cardisoma
(observed
conspecifics
frequently consists
on
in
332).
epiphytic
Sesarma
Rhizophora
1958:
others,
hand,
by prolonged
have been
lily)
and stems of mangroves,
growing
the other
PREDATORS
The food
on
that
reason.
TIDE
tall
hiding is
as
themselves to
expose
On
crabs, probably Coenobita,
individuals
roots
a
well
GERLACH,
: leaves, fruits, decayed wood,
Pandanus and
hermit
33).
OF
itself
("Flutgâste",
(and only
1975:
HENNING,
;
SEARCH
(of
this
LOW
water
Sesarma mederi
(e.g.
as
species.
sea
aperiodic freshwater floods within its bur-
be the main
FROM
conditions
(Metase-
possibilities
of other
those
242) climbing
burrows
284).
in
the two
Sesarma mederi and
vocator).
own
1975:
are
chrysospilos
species
survive submersion in
burrows,
cruentata; WARNER,
niopsis
3)
in
Uca
climbing
may
tide
endures
(HENNING,
ever, start
can
crab
of
of Indo-
species
by predators (fishes, portunid crabs)
Aratus,
hiding
(see
by Goniopsis
1930:
rising
within their
guanhumi
soma
2)
the
Some
ground is
the
Periophthalmus
advantage
experimental
induced
Goniopsis,
when
vegetation
1972).
take
forms
VERWEY,
instead of
from
row
with
like
used
climbing
probably mainly
Crabs
POLUNIN,
the
inhabiting
during low tide, either hide in
behave much like certain
Goniopsis)
long periods
for
186,
burrows
though the
As
air
mud-skippers, especially
1968:
Crabs
GROUND.
into the
upwards
latter crabs
Pacific
(MACNAE,
THE
active in
being
climb
or
OF
FLOODING
the
sponges
the
on
xan-
these
45
of
of
Capture
roots.
stomach
Metasesarma
perched
4)
SEARCH
(1929:
1964:
Geograpsus
vocator and
5)
SEARCH
OF
is not
(Metopaulias,
in
WATER
Sesarma
habitats.
dry
a
The
main motive
climbs
is not
for
(like
Typ-
1953:
roberti)
Se-
29),
clypeatus
(PHIL-
Goniopsis
were
phytotelmata (see 5)
angustipes,
introduction)
1961),
genus Coenobita
USE
OF
LOW
compared
(GROSS,
to
the
perhaps
because it
two
has
species
habits
climbing
a
sea
the
the "fish"
or
sap
water
water in
the lowest
of
of
correct statement
says that
water to
and
micro-pools
blood
closely
capcon-
related
1964).
BRANCHES
by displaying
is
these
something (whether
1955)
to
supply by phytotel-
but also for land crabs
roberti),
Birgus prefers fresh
1932, GROSS,
when
S.
earliest reference
see
Water
inhabitants of
the coconut trees to drink
centration
OUT
frog.
Sesarma
Aratus and
PHYTOTELMATA.
(MCNEILL,
quite clear).
tivity (HARMS,
6)
IN
only important
Birgus (SOLEYMAN,
on
identified)
tree
shelter function, too.
a
mata
a
UNDER TREE BARK.
were
Coenobita
67),
juvenile
cicada
17-year
(not
eating
found to hide under the bark of trees. Of course,
have
tree-climbing
a
live
crinipes (HOLTHUIS,
1956:
(LAIRD,
Uca
5).
HOLES AND
a
crab
a
and
collected from trees
species
rotundatum
sarma
preying
Sesama roberti and
figured
on
mentioned
261)
OF SHELTER IN TREE
found in tree holes:
LIPS,
4)
flooded mangrove thicket
a
several
ically,
(1964:
remains
arthropod
in
as
Examination
rarely.
more
yielded
insects
(possibly Aratus)
HARMS
on
reported
Metopaulias
PHILLIPS
rubripes.
“Sesarma crab"
in Florida.
in
147), probably
1964:
(HARTNOLL,
prey is
agile
contents
AND
TREE
STUMPS
AS
A
SORT
OF
LOOK-
Uca males.
GRADES
With respect
the
ground)
categories
:
to time
most
spent
of the
on
species
the tree
can
be
(compared
placed
to
periods
into the
on
following
46
a)
and
temporary
sporadical,
Callinectes,
e.g.
Ucides, Ocypode,
Uca,
Sesarma ricordi;
b)
and
temporary
bita, Cardisoma,
c)
and
temporary
the
tidal
Sesarma
except
including
It
is
interesting
sarminae
Superimposed
in
on
this
characterized
gradients,
marine
( Ocypode)
Metopaulias),
seaward
nautes),
and
roots
grass
fringe
reed
(Pachygrapsus)
to
nevertheless
the
more
do not
tanks
of
At
least
apparent
the
not
and
as
even
-
sea
in the hills.
tipes
the first
huge
branches
on
bromeliads.
two
Se-
several
are
completely terrestrial
( Sesarma roberti,
(Potamo-
(Potamonautes),
trees
( Aratus)
case.
every
trees than Aratus.
Metopaulias
The latter is
former, which normally
Such
other
(examples given
discrepancies
can
hides
be
ex-
routes.
phyletic
ROUTES
routes
from
sea
to
tree
are
quite
:
i) Estuary
near
to
terrestrial than the
large
subfamily
to inland forest
coincide in
PHYLETIC
is
to almost
(Goniopsis)
topmost
plained by different phyletic
there
extremes
freshwater-adapted
permanently living
more
scale,
following
( Callinectes)
These different scales
e.g., is
to
of the
category.
basic
the
latifrons,
etopograpsus
mederi;
grapsid crabs
each
rough
by
Meta-
juveniles, Metopaulias.
parenthesis) : aquatic ( Callinectes)
(Birgus),
in
in
to
(corresponding
juveniles, Aratus;
larvae and
represented
are
M
S.
elegans,
note that
to
Birgus, Coeno-
e.g.
mostly periodical
larvae and
d) permanent
e)
permanent
and
Goniopsis,
e.g.
rhythmic,
not
rotundatum;
frequent
rhythm),
rubripes,
sesarma
but
frequent,
Sesarma
freshwater
-
epiphytic
The last
two
-
tanks of saxicolous bromeliads
bromeliads
stages
are
Metopaulias, respectively.
clearly phytotelmic
less
phytotelmic
as
and
growing
on
trees
represented by
The former
the latter.
inhabits
a
and
growing
high
species, however,
A third
species,
somewhat
up
Sesarma angus-
S.
is
roberti,
different niche.
47
In
the
were
2)
Indo-West
S.
Pacific
mangrove trees
the Sesarminae
etation is
mangrove roots
-
individuals
young
of other
but, obviously,
mentioned in
living,
burrowing
see
tation. Metasesarma
holes.
Aratus
burrows in
rubripes
terrestrial
Trinidad,
the
is
a
final
a
skilful
The
same
the
seems
climber
mangrove
taller trees
to hold true
ascend
though
stage
(cura-
str.
s.
never
still
vege-
digging
digging
never
is reached within the
stage
area
listed in
typical ground-
Sesarma
evolutionary
this
route of
examples
are
1977) that
mangrove thicket;
fringe of
the
there
subgenus
HAGEN,
ground. Typically,
shaded and humid
casionally.
VON
represents
the
of
species
and rectum ;
caoense
phyletic
groups too: mangrove veg-
half of
than
more
trunks and branches
-
This is another
(inland trees).
-
Table 2. In the
mangrove belt of
the
only
found between the leaves of Crinum asiaticum.
Ground of intertidal zone
of
rotundatum,
for
growing beyond
ascended
are
only
oc-
the Indo-West Pacific
Sesarma mederi and others.
j) It
is
however,
possible,
rotundatum)
that
via
progressed
some
also the route of the Coenobitidae
least
at
of the
some
from mangroves
to CHACE &
A
4)
This
sus
fourth
phyletic
gracilis,
occurring
and
on
grove roots,
seems
ment where
the
encrustations.
portions
A
)
On
1
)
among
the roots
crab.
that
are
the confusion of habitat data of
(1918, following JARVIS, 1897)
see
S.
been
However,
also recorded
were
begun
crabs
grapsid
stones
at
(algae
displaced
xanthid
rock
may
Birgus.
rocky
on
of
the
of the xanthids mentioned.
encrustations
The
(e.g.
have
( Coenobita clypeatus according
could have
some
closely related
displaced
of
one
to be
Sesarma
126).
route
and of at least
mangroves
another
1969:
in
Coenobitidae
and cliffs
to be the route of
seems
Grapsini
HOBBS,
of
which
culminating
tree-climbing
(Table 2)
species
beaches,
sandy
P.
the
and
seaward
rocky
transversus
Hexapanopeus
these
submerged
these two
HARTNOLL
of
fringe
of
on
caribbaeus
during
the
man-
marine environ-
still lives
crabs
species
(1965).
subfamily
Pachygrap-
sponges) covering
from the
Both
shores.
live
on
most
of
similar
could
the
the
Pachygrapsus by
be
lower
tidal
RATHBUN
48
But
cycle.
hide in occasional
likely
This
Grap sus type.
walls
stone
evolved
have
to
"air-climber"
the
even
its
within the mangrove
from
directly
would
Goniopsis with
rock-living
of
the lack
explain
to
tendency
vegetation,
of
ancestors
is
the
digging abilities
in
Goniopsis.
Affinities of
rock
climbers
discussed in
predisposition
to
the
climbing
tree
following
lead
the
to
idea of
section.
PREDISPOSITION
When
its habitat
shifting
from encrustations
Predisposition
to
the
nature of
tics
similar to this
of
the
habitat
of
nautes
see
sp.
Table
bank of
a
may
by
DIETERLEN in
preadaptations
BLISS,
living
and
the
situation
Birgus
with
kind
are
"aware"
provided by
tree holes
as
individual of Potamoforest
(eastern Congo,
Parinari tree
huge
dangers
not
of
to
for
the
steep
is
is
preadaptive
possible
which
The shell
desiccation and
a
specializations
shell is
1930, EDNEY,
typical
are
given by
crabs,
prédation
in
handicap
to the
air
1960,
Sesarminae,
(Ocypodidae,
superior
for
for most groups
of hermit
a
certain
though
any
for
re-
terrestrial
a
climber.
species of Coenobita
climbing.
Preadaptations
Is
e.g.
1969), is nevertheless
possessing
respect
1968)
be
can
(see VERWEY,
environments
Coenobitidae).
(REESE,
themselves,
economy
MACNAE,
climbing
to tree
crabs
semi-terrestrial
Gecarcinidae,
ducing
of the
water
1968 and
in
Irangi
given by
characteris-
scarcely
locally take
the
on
to lower
river.
Secondly, predisposition
breathing
the
be
firstly
may possess
of this
Likewise,
adapt
could remain similar.
therefore
that
roberti)
have taken
to
that the animal is
examples
Sesarma
their burrows.
observed
2)
so
Further
being displaced.
can
habitat. Trees
original
species ( Uca vocator,
equivalents
climbing
tree
habits
feeding
salinities. Its locomotion and
mainly
had
Pachygrapsus gracilis
mangrove roots,
rock to those
on
can,
that not
to
tree
and down within the
of
course,
only
the
climbing,
more
include behavioural
ability
but
vertical
of
also
scaling
the
portions
of
habit of
a
components.
rocks
(see above)
climbing
burrow.
up
Burrowing
49
is
in
common
the
and Gecarcinidae
Ocypodidae
CONSEQUENCES
The book
adaptations
could be
thors
spines
classified
as
with
dealing
bromeliads.
under S.
into
to be
and
served in
falling
1930:
as
well
(see
tendency
present
thus
The
uses
only
specialized
locomotor
an
of
above,
leaping
to
the
or
preda-
different
(unre-
is
known
from
it
ob-
any
noise of
flooded
and
453,
was
man-
VERWEY,
mederi, respectively;
Eriocheir).
generally
the
chelipeds
activation
first
praying
two
when
and S.
au-
cara-
leaves
(see
splashing
noise of
1921:
are
clearly specialized.
not
the
legs
additional
walking legs
a
locomotor
pattern
of
of
pair
mantis
join
insect
peraeopods
as
one
of
still
is not
might
as
believe.
( Mantis religiosa),
these
is
The
as
posterior pairs
prothoracic
action
ocypodids.
elegans
the
Trinidad,
The
locomotor function
the
the
trait of
In
flat
very
observer
response
RATHBUN,
that the
original
is found in
parallel
normally
as
finally
close
a
1314)
All
too.
here,
176).
characteristic
on
use
indicating
remote from its
A
to
of
bent
1955:
Metopaulias
and
escape
1932:
in
Sesarma
The locomotion patterns
interpreted
of
a
like
well.
as
between
spaces
behavioural
This
in
cf. PETERS e.a., 1933: 84,
be
that it has
tree
a
above)
special
on
characters
by BALSS,
to
(see
stress
presence
as
a
LANG
183, concerning
common
(referred
of
(HARMS,
to be
seems
grove swamp
Meta-
as
"how the
on
be said
can
Some
ontogeny
trunk
climbers.
grapsids
crabs.
narrow
common
Coenobita
crabs
long chapter
a
very little
should be mentioned
a
of
species
Birgus
the
into the water in
or
the
THE ANIMALS
preadaptations
behind the
seems
lated)
Coenobita
specialized
angustipes)
Dodging
ground
The
in
Metopaulias,
fitting easily
pace
Until now,
arboreal life
in
TO
(1950) contains
changed".
to
with
dactyli
PEARSE
by
animals have
in
and Aratus must have evolved.
rubripes
sesarma
can
also
ancestral Sesarminae from which the expert climbers
more
tor,
and
which
legs during walking.
posterior
climbing
a
legs
vertical
in
their
surface
(ROEDER, 1937:363).
Most
of the
species
listed
in
Table
2
are
still
mainly
ground-
50
living and
be
expected
climbing
to
turn
out
worth
and behaviour should
morphology
best
at
between needs
compromise
a
The
other activities.
and
are
ever,
Their
burrowing.
even
of
specializations
of
Aratus, how-
closer examination.
a
Summary
An
climbers,
in
Grapsidae
on
yielded
these
The
leaves.
further
belong
trees
to
Some
systematic
are
of
groups
of
and
diet
feed
mainly
tree-climbing
in about half
on
crabs.
or
of all
algae
from
and
of
cases.
re-
climbing
decayed wood,
(Table 2)
most
Trinidad,
semiterrestrial
only
treeis
behaviour
literature
in
Like
of
species.
Motives, grades, phy-
of the habit of
consequences
well-known
are
peculiar
crabs
of the
Some
the locomotion
data
many
tree-climbing
species names).
this
and
pertinent
comprising
mangroves
1 for
ricordi)
the
to
found
names
predispositions
S.
on
synopsis
(1)
anomuran
Table
Goniopsis cruentata)
roberti,
data
was
A
species
ascended
letic routes,
pisonii,
Sesarma
and
(12)
presented (see
Aratus
given.
are
30
(e.g.
first time.
mangrove
brachyuran
is
Aratus
(e.g.
others
for the
corded
of the
(West Indies)
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list
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3
17,
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main
MENTIOED
SPECI S
TRINDAIN
TABLE
31 31
31
13,
and
31
9,
24, 26,
31)
g,
13, 23,
6,
5,
32 10
31)
26,
31, 23, 31 26,
(Tr
SYNOPSI
3
(56)
(15.9) (26) 22. 24.
sites
)
S
(Tr
colecting
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maximu
)
1
cb
mm
3
m
6
4
1
23. (27) (34) 36.
91.8 (113) (11.5)
are
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author.
the
1
OF
2)
(Tr Tr Tr Tr Tr Tr (Tr (Tr
Tr
Tr Tr
32
cruenta
speci s
gracil s rubripes
pisonii
angustipes ricordi roberti
Goniopsi Pachygrapsu Aratus Metas rma Sesarma Sesarma Sesarma
r
rapax vocator
Uca Uca
cordatus
Ucides
guanhumi
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Cardisoma Hexapnoeus
those
clypeatus literature.
Coenobita
parenthesi
sedis
Grapsinae Sesarminae
group
0
BRACHYURA
Ocypodinae
incertae
OCYPODIAE, GECARINDAE XANTHIDAE ANOMURA COENBITDAE
only). localites trees,
Coenobita
figures for
car pace,
of
GRAPSIDAE,
observed
given but
from
in
r
climbing
not
(including
SYTEMAIC
below,
by
width
=
cb
!)
(used
car pace
of
of
ascend
list
to
to
literature
refer
figures from
Identifcaon
known
Length species
))
2
3
crab
56
other
and
and
boulders
m)
TRE S ascende
ground)
(6
(4
veg taion above
TO
KNOWN
2
BRACHYUN
trunk
mangroves
m)
(1.5
trunks other
Mesrchmida
mangroves
m)
(2
tree
mangroves mangroves mangroves roots
bromeliads Rhizophora growingRica)
Pisonia mangroves
t
r
u
nk
Trinda :
recumbent
m
a
n
g
r
o
v
e
s
epi hytic Asia: bromeliads (Costa
Islands Neotr pics Estuary: Neotr pics: mangroves Que nsla d: dead Rhizophora bushes Islands
East Island
grass
Marshal Eastern Congo Western Java: North Ethiopia amaica Germany Neotr pics Marshal argentea Ven zuela trees, amaica trees South Brazil Cocos
tree
height
and
on
:
of
and
:
:
:
geographic
m)
(climbing
m) and
(6
:
(2
:
:
:
:
:
THAT
J
SPECIES
prep.).
(in
J
(e.g.
HOLTHUIS
1969)
443)
92)
(s)
84,
192
29)
author
(
RATHBUN (1918: (1953:
(1953:
ANOMURAN
240), 126),
HOLTHUIS text LANG RATHBUN WEYL
in
see
?
33)
(unpubl.) 305), 436) litt.)
(1933:
(1929: (1967:
al.
HAGEN
VON
HARMS MACNAE
and
cf.
syno ym
167)
1964;
(1953:
(in
LEWINSOHN
see
others READ,
1:
CRAB
boulders
roots
on
(1892) (1968:
litt.)
et
text
see
PETERS
for
(in
text
see
HOLTHUIS
text
see
ANDREWS HARTNOL, MACNAE text ABELE
see
(Latreil )",
ANDCLIMB
TABLE
ARE
region,
trees
m)
OF
SYNOPSI
Dana
species
(Latreile)
(Lockington)
)
1
(Herklots)
YURA
crinipes cruenta pulchra
pelii
BRACH
GRAPSIDAE Geograpsu Goniopsi Goniopsi Goniopsi
(White) (Owen)
Saus ure)Edwards Edwards)
(De
latifrons thuk har gracils
Milne
H.
sinensi
Edwards
M.
H.
Milne
(H.
pisonii
rouseauxi
cruenta
(Rathbun) Rathbun
Man)
(De
rubripes depres us minut m
Dana Smith
“Goni psi
as
angustipes angustum
Metopgrasu Metopgrasu Pachygrapsu Eriocheir Aratus Metas rma Metas rma Metopaulis Nanosearm Sesarma Sesarma
mentio ed
!)
57
ascende
veg taion
mangroves
ground)
above
region, height
)
man-
indvi uals
ferns ferns dead
mangrove
and and
of
tree
(SCHNE )
Islands HOLTH.),
trees
:
Congo Java: Etiopia
:
:
:
ava ava ava
J
of
(2
bushes (1.5
trunk
branches bambo Marshal asi tcum
m,
(climbing Estuary mangroves branches co nut mangroves mangroves
geographic
:
m)
J
J
mangrove
(4.5
m)
(1.5
trees
:
and
roots
and
:
Trinda : groves Trinda Ellice other Crinum
tree
Rica:
cm)
(20
mangrove
(3
see
reed
syno ym
Lagunclari mangroves Pacifc mangroves mangroves
and
:
trees
for
:
Bombay Costa Trinda Trinda : Indo-West Trinda : Col mbia: Florida:
on
TESCH
m),
m)
:
dead
(1917).
grass
breadfruit young
recumbent
recumbent
roots
and
(lm
(1940).
(SCHNE ),
Man"
453)
(s)
author
183)
(1921:
245) litt.)
RATHBUN (1930:
(in
306),
63-65)
594),
180, 180)
406),
(1900:
(1929: (1930: (1930:
LEWINSOHN text
LANG VERWEY HARMS VERWEY VERWEY
in
see
fig.)
131)
33,
(1953:
67)
(1904: (1956:
text
see
(1959:
(1941:
text text
see
see
THESING text
see
text
see
)
3
species Herklots
Man Milne
Milne
Man
Milne
Milne
(Pal as)
Hess
elegans eumolpe gut atum mederi meinerti ricordi roberti rotundatum
ceratoph lmus
Sesarma Sesarma Sesarma Sesarma Sesarma Sesarma Sesarma Sesarma
OCYPDIAE Ocypode
De A.
De
H.
H.
oceanica
for
White",
taeniolat
2
Edwards Edwards
see
syno ym
(1912:
)
Edwards Edwards
De
103,
177)
BORADILE SCHNE HOLTHUIS LAIRD ALTEVOGT CRANE
TWE DIE
H.
(Stimpson)
Latreile
(L.)
(Smith) (Herbst)
brevifrons rapax vocator cordatus
sp.
Uca Uca Uca Uca
Ucides
guanhumi
GECARINDE Cardisoma
“Sesarma
as
“S.
or
(BOR .)
Boradile"
gardineri
“S.
as
mentioed mentio ed
))
2
3
58
roots
TÛRKAY
and
ascende
ground)
veg taion above
region,
geographic
height
(climbing
trees
m)
:
mangroves mangroves
(West
roots
Rolas
mangrove Rhizophora Rhizophora
:
trunk
Parinari
S.
grass
Rhizophora
Spartina
mangrove
see
trees
and
trees
syno ym
other
m)
(20
(6
islands
on
trees
Congo
buildngs Trinda Ven zuela: Panama Eastern
(USA)
Asia
Coast
East
South
Gulf
Pacif c
Trinda
mangroves
Indo-West
Florida
Java
author
and
(1882:
GRE F
DAWSON
litt.)
(in
&
text
see
(RATHBUN,
(80
184)
many
201),
(1963:
cm)
Edwards"
(1975:
26)
1918),
and
:
347)
(s)
for
:
and
:
:
:
:
:
Thomé
mangroves
(40
(20
Africa)
and
m)
cm)
RODIGUEZ MCOSCER DIETRLEN
167)
128)
1932,
1975)
figs,
307-308, 180,
H.
7)
DAVIS,
252, (1930:
(1705: HARMS, 1959b, (1929:
(1968:
(1967:
&
(e.g.
MACNAE
AB OT
Milne
VERWEY
RUMPHIUS others HOLTHUIS, ALTEVOGT HARMS 327),
?
lagostoma
text
“G.
or
see
(GRE F )
)
4
2)
(conti ued
2
Table
species
(Sendler)
weileri
(L.)
(Stimpson) (Gould)
Haan)
Rathbun
carib aeus haris i
sp.
sapidus
Gecarinus XANTHIDAE Hexapnoeus Rhitropan eus POTAMIDAE Potamnutes PORTUNIDAE Calinect s
(De
RA
U
OM
AN
longitarsu
Stimpson (Herbst)
ruricola
“G.
as
(L.)
DIOGENDA Clibanrius CoENOBITDAE Birgus
cavipes
clypeatus
latro
Coenobita Coenobita
mentio ed
)
4
(1973).
59
he
and
ascende
veg taion
ground) bushes
above
region, height
geographic
spp.
bushes
(climbing
Hibscus
Pand us
and
:
of
:
trees
Neotr pics:
Islands)
trees
Guinea:
(Maldive
Coenobita
m)
Islands) trees
as
(mentio ed
Dana
list
The
(6
(Tuamotu co nut bushes
peltata
name.
brevimanus
C.
:
and
Sudan
Western New Minikoi Herna de Mangareva tilaceus Port
species
no
and
Edwards
gives
Milne
72)
)
5
92)
(s)
1:
r
o
h
t
u
271)
litt.)
(in
a
(1972:
BALL
190
238-239)
PESTA
(
AILE
AD
HOLTHUIS BORR
(1927:
(1904:
in
BORADILE
Coenobita,
of
SEURAT WERNER
?
habits
H.
rugosu
C.
Edwards,
Milne
H.
Edwards
tre-climbng perlatus
Edwards
1969).
LEWINSOHN,
see
explantion
(for
Heler"
signatus
)
Milne
species
H.
6
Man
De
Milne
H.
compresu longitarsi perlatus
(Forskal)
the
C.
of
“Clibanrius
comprise Latr.").
s
p
e
a
k
i
n
g
scaevola
When clypeatus mentio ed
as
68)
(p.
Coen bita Coenobita Coenobita
Coenobita
)
5
found “C.
)
6
PLATE
Above.
—
Goniopsis
cruentata
3
Below.
— G.
cruentata
low
(right)
perched
m
above
and
tide. Both
on
a
branch
of
a
Laguncularia
tree,
I
about
ground (Tr 3I).
Aratus
pisonii (left)
species prefer
to rest
on
a
Rhizophora
head down.
trunk
at
PLATE
II
— Sesarma
Above.
roberti
(arrow) climbing
within
a
maze
of loose
bamboo
canes
(Tr I0).
Below.
—
The
legs
of S. roberti
rain
are
water
visible
within
(thus forming
a
a
bamboo
stump partly
phytotelma).
filled with
PLATE
Above
and below.
ascending
—
the lower
Ovigerous
portions
females
of
and
a
few
Laguncularia
males
trees
(below right)
ahead
of the
of
rising
Uca
tide
III
vocator
( Tr 27).
PLATE
Left.
IV
— Metasesarma
rubripes
flooded
Centre.
—
Four
on
grass
individuals
of
the
trunk
Aratus
male
of
Uca
flooded
on
rapax perched
after
a
coconut
pisonii feeding,
(Crassostrea rhizophorae)
Right. — A
of
tree,
1.5
m
above
the
Paspalum serpentinum (Tr 26).
heavy
rains.
Rhizophora
on
a
at
low
Laguncularia
(From
VON
tide,
among
oysters
(Tr 24).
roots
tree
HAGEN,
in
an
1970b).
area
(Tr 3I)