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Contributions
to
Evolution
of reef-associated
particular reference
decapod
119-130
(2-3)
(2003)
The
Publishing bv,
Hague
through time, with
crustaceans
to the Maastrichtian
72
Zoology,
SPB Academic
type
area
NL-5283 WB
Boxtel,
René+H.B. Fraaije
Oertijdmuseum de Groene Poort, Bosscheweg 80,
Keywords:
Decapod
evolution, K/T boundary, biostratigraphy
crustacean
Abstract
prior
result
strata in
of
twenty
some
years
the Maastrichtian type
1,200 generally small-sized
The stratigraphical
date
from
shown
the
For the first
intensive collecting from
is
(Late Maastrichtian)
time, decapod
crustacean
tools
remains
local
on a
are
1887a-
1886;Forir,
1981) suffer from
lack of strati-
a
species
are
discussed.
now turn
control. A taxonomic revision of
of
most
to
out
to
regional scale.
to
1854;
den Binkhorst,
van
1881; Pelseneer,
1889; Mulder,
graphic
these
decapod assemblages
1857; Binkhorst
remains.
thirty-one species known
Formation
be useful biostratigraphic
brachyuran
by various authors (Bosquet,
1861; Noetling,
of more than
c,
of the
successive
collection
a
anomuran and
ranges
Maastricht
and five
of
area
1987
to
Binkhorst,
van
The
the Netherlands
Since
1987,
type
species
by Collins
from
the
et al.
1993), Collins
&
et al.
van
al.
et
(1995).
Maastrichtian
described and discussed
2002), Fraaye
a-c,
out
(1987), Feldmann
al. (1991,
al.
carried
new
area were
& Collins
Introduction
was
by Fraaye
(1990),
Jagt
et
(1995), Fraaye (1996
Bakel
(1998)
and
Jagt
et
(2000).
Rigid collecting from six key sections (see Collins
Brachyurans
utilize
a
broad array of
including deposit feeding,
filter
feeding types,
feeding, seaweed
grazing, scavenging and predation. They have been
a
major component
from the
played
rine
of many
Late Jurassic
an
marine communities
onwards and
the
important role in
probably have
evolution of
ecosystems. Unfortunately, they
described
by paleontologists
overlooked
by biologists
munities, especially
their
highest
in the
and
in
are
are
diversity (Zipser
to their
collections
&
Vermeij,
factors
and
small
relatively
chance of
being
•986; Plotnick
anomuran and
et
al.,
1990).
brachyuran
frss than 10
species
from
sizes,
com-
1978).
possible
mm.
the
often
in
due
and therefore the
overlooked in the
grained sediments of the
18
probably
is
The
field
(Bishop,
average
size of
crab remains in the fineMaastrichtian
Almost all
168, fig.
cades has resulted in
well-documented,
containing
over
Oertijdmuseum
an
the
1) during
past
two de-
extensive, stratigraphically
decapod,
collection
crustacean
1,200 specimens
and housed
de .Groene Poort, Boxtel
(the
at
the
Neth-
erlands).
frequently
marine
destruction after death, their
apparent scarcity
paleontological
p.
Carapace
size and
morphology through time
where crabs reach
tropics
Apart from paleoecological
1995,
only rarely
also
Recent
ma-
et al.
decapod
type
area
crustacean
Maastricht Formation described
Crab size is
sure
the
thought
(Vermeij,
majority
trolled
to be related to
predation
of Mesozoic
by predation
crabs
pressure
was
of
occurring cephalopods.
can
pressure
probably
con-
simultaneously
evolving reef teleost fish (Vermeij,
predation
pres-
1978). The relatively small size of
1978) and
co-
Apart from size selection,
also lead
to
avoidance strat-
egies, i.e., strategies devised
to minimize the risks
of
crabs,
being predated
age four
upon. For
strategies that play
ing their evolution:
1
-
a
we
may
envis-
continuous role dur-
to hide and live in
crevices;
120
2 -
R.H.B.
camouflage their
4
sediments;
carapace; 3
swim in open
-
in the first
Carapace morphology
erally quadratic
have been
successful
wards and have
small size.
The
led
the
to
acterized
by
time. The
appear in
other three
Early
the
during
due to
antero-lateral
Cretaceous
spines.
It
global
is
crabs
sea
the Late
more
The
more
of clades
have
less
leading
1996b).
to
et
to
and
and
stands
new
clades
After
followed
ecological
more
leading
to
and
phology
fauna
the
ever
cycles, perfectly
diverse
on
matches
more
and
filled,
(in size,
feeding strategy) decapod
seen
rapid
the swim-
as
successfully
were
most
of
stable periods, in
period, however,
niches
recent,
more
evolutionary
pulses
of such groups
crabs. After each
ming
mor-
crustacean
earth.
sea
Cretaceous/Ccnozoic
to
level
forced
in
to
cides
tory
with
peto
Conse-
of
‘species
fall, newly
retreat and
predatory
communities,
To
the
put
are
type Maastrichtian
summarized in
of the type Maastrichtian
Fig.
lins
compared
all
age
(1983,
rapid evolution
of
in
probably had
are
1.
presently
As the
(1985: Recent,
decapod
crustacean
most
known. Studies used
Albian, Texas),
Hungary
French
regions
majority
is linked with
and
Polynesia)
diverse of
are
Bishop
Muller & ColMuller
(1991: Upper Eocene, Flungary),
Middle Miocene,
crus-
perspec-
from similar environments.
faunas represent the
1986: Lower
servations
benthic
decapod
some
decapods
reef-associated carbonates all
as
during
(semi-iso-
data for five other well-studied
tive, faunal
their
The
groups and others
evolutionary patterns
(1984:
Austria), Guinot
and
(Lower Kimmeridgian,
personal
southern
obGer-
many).
Brachyuran crab diversity increased in
1), implying
the
a
brachyuran
role
in
do
strong evolutionary
Late Jurassic onwards.
tain
the
groups
It is also
played
a
For
turnovers
(Fraaye,
and
instance,
clear that
more
cer-
prominent
than
they
and
the
Early Cretaceous, respectively.
evolutionary
(Fig.
the Dynomenidae
appeared during
radiation
time
radiation from
shallow marine ecosystems
currently.
Raninidae first
stasis,
with maximum
and thus also affected crab evolution.
taceans in the
and ages
(Taylor,
have thrived
effects upon the structure of all
possibly
of
seem to
lated) shallow-water habitats.
of these
in other preda-
fish and gastropods
These groups also
periods of transgression
diver-
common.
teleost
groups,
1981).
similar radiations
two
turn
by periods
crustacean
The Late Cretaceous diversification of crabs coin-
Further,
crabs
for crabs
as
decapod
evolution
1987).
during
rapid
very
referred
were
polyphyletic origin
faunas
produced by this type
1996).
the
and
30 million year
roughly
linked
leading
seas.
niches
probably
was
strong competition,
was
ah,
among
isolated,
extinction and faunal
This
are
the Albian-Ceno-
In isolated seas,
recently been
flocks’ (Yacobucci,
mixing led
(Haq
periods
transgressions. These
by high
evolved crab populations
carapace
Maastrichtian
of crabs
Jurassic,
ecospace
into existence.
monophyletic
event
or
latest
Eocene, the
Campanian-Maastrichtian,
shallow,
the
their
probable
stands
high
characterized
sification
in
the
adaptive radiations
the Eocene and Miocene
quently,
al.,
and with
long
coincidence that
diversity
diversification, mixing
profound
quadrupled
mere
level
during
are
et
swimming crabs
wider than
to their
not
Major explosive
the
in-
max-
the Ceno-
during
Recent
are
slower and stabilized
a
dominated. This model with
time
They
an
in shallow-water
lived
They appeared
of increase in size and
occurred
show
Mundlos, 1982). The
(Fraaye, 1996a). During
comparison
ancestors.
and
Mesozoic but today are,
of the
largest swimming
came
through
deep-water organisms.
possess carapaces that
manian,
increase
strong competition
ancestors
which
of reef decapods through
char-
are
diversity through time, with
predominantly
zoic, mainly
The
prosopids
Evolution
-
strategies, which
Cretaceous
and Eocene (Forster &
probably
of
the Late Cretaceous (Feldmann
environments
many
ex-
niches
of the third group.
examples
are
in size and
Raninidae
riods
an
ecological
new
considerable size
a
the
during
with
with
strongly elongate and burrowing Raninidae
(frog crabs)
size in
on-
relatively
a
seas
evolved during the Cretaceous,
probably
1996)
Such crabs
Jurassic
diversification
rapid
1985). The
(Forster,
ima
of shelf
area
offered various
transgression
crease
Late
retained
consistently
vast
is gen-
group
elongate.
from the
during
pattern
of redundant bioherms after the Callovian
pansion
and
somewhat
or
burrow into loose
-
waters.
Fraaije
during
Late Jurassic
After
the Late
a
rapid
Cretaceous,
Contributions
Fig.
to
Zoology,
I. Reef-associated
72 (2-3)
decapod
-
crustacean
121
2003
faunas
through
time.
122
Fig.
R.H.B.
2.
Stratigraphic
ranges
of decapod
crustacean
they dramatically declined from the
The
to
a
same
may
hold true for the
somewhat
Raninidae
lesser
sequent niche
from
Eocene onwards.
Calappidae, although
extent. The
possibly corresponds
biotic radiation of
genera
decline of the
to the
synchronous
clypeasteroid echinoids and sub-
competition
Other groups, such
as
and
however, display
a
nidae from
rapid evolutionary
as
do the
Evolution
cessful
radiation in
Ocypodidae and Pilumsuc-
of reef decapods through
however,
is
that of the Xanthidae.
Having originated during
the
evolutionary adaptations
resulted in the
diverse
time
area.
group,
group of
Early Cretaceous,
brachyurans
in
by
their
far most
reef-associated
faunas.
Decapod
Cancridae,
the Miocene onwards. The most
-
Maastrichtian type
replacement.
the Portunidae,
Majidae, Leucosiidae, Parthenopidae and Grapsidae,
post-Eocene times,
Fraaije
most
&
crustaceans are, and
important
food
source
for
tion pressure
hunting
on
crabs
in the
the
1997). The preda-
by relatively fast swimming
ammonites such
Sphenodiscus
were,
cephalopods (Fraaye
Jager, 1996; Jager & Fraaye,
and
probably
as
Placenticeras and
Campanian and Maastrichtian
Contributions
to
Zoology,
(2-3)
72
2003
-
123
era
which
ary
some
apparently do
of them in
For
deposits.
The
possibly
a
occur
classification and
possible evolutionary
of
Glyptodynomene
in younger
and
subsequent
is uncertain at the
offspring
extinction
bound-
the K/T
cross
Stephanometopon
two genera,
Aulacopodia, generic
moment.
not
reality may
Graptocarcinus
and
correspond
to
seems
to
niche-displacement by the apparently better adapt-
ed
Dromiopsis
1996b).
(Fraaye,
spp.
Of the fifteen genera known from the Danian of
Denmark and Sweden,
3.
Fig.
Number of
for the
Middle
Danian
may have led
xanthids
decapod
to
crustacean
of Fakse
new
the introduction of the
and abundant food
In the
Maastrichtian
(central
Poland)
of
mens
more
Scaphites
as
a
less
chambers,
regular
are
The known
chyuran
unscathed,
day,
in
&
pers.
in
their
body
decapod
1996; Fraaye,
of the
body
crus-
and the
the result
evolved
adaptations,
relatively
breaking
and
(Taylor, 1981; Zipser
development
of
&
crushing
mol-
Vermeij, 1978)
larvae.
teleplanic
Such
long planktonic larval phase is known
tropical Recent representatives
1997).
but is
cham-
(Vermeij,
ranges
type
of all
genera
presented
area are
anomuran
rare
or
absent in taxa
a
in
of these families,
inhabiting cooler
seas
1978).
area, ten
do not extend be-
survives
Paguristes,
studied
crusta-
for instance
here) and partly because of
effort.
and Sweden
lesser
to
Leptoides
decapod
outcrops (as
Except
for the fauna of
(Collins & Jakobsen,
1994),
degree those from Greenland (Collins
1992), Argentina (Feldmann
and Antarctica (Feldmann
et
ah,
Therefore,
overall poor
it is
to
be
Percentages
shown
sen
members in
show
a
Fig.
the
their ‘mobile homes’,
are
plotted and
Meers-
Paguridae (hermit crabs)
increased
an
empty
diversity, di-
availability
of
shells.
In
gastropod
the Emael Member, the Raninidae and
Calappidae
dominate. In the Nekum and Meerssen
members,
the
Calappidae
remain
important component
come
a
more
or
whereas the
less
constant,
Raninidae be-
proportionally less dominant. The Callianas-
1993),
gen-
families
Emael, Nekum and
4. From the base of the Emael
rectly correlated with
but
ten
for the
clear increase in number and
the
the
in the
diversity
area
decapod
Member up section,
sidae is the
control.
for
separately
ah,
biostratigraphic
expected that of
Maastriehtian type
et
crustacean data base for the Danian is rather
meager, with
Substrate and decapod
and bra-
poorly known, partly because
few accessible
Rasmussen,
decapod
feeding
Calappidae,
are
comm.).
of which,
of collection
1995)
paired,
have
two, Cretachlorodius and
region
a
probably
speci-
of
boundary, whereas thirteen pass this
and
Denmark
and to
and Xanthidae
predatory
boundary
faunas of the
luscan shells
this interval, ten
one
relatively
luck
crossing
Recent
3.
and
ization of the chelae for
place
stratigraphic
cean faunas are rather
the
of their
in
of the K/T
valley
gave rise to Recent forms. Danian
m
Fig.
The successful
Vistula River
genera known to date from this
appear
yond the K/T
of
Fakse,
shown in
are
during the Late Cretaceous, especially the special-
Fig. 2. Of the twenty-three
Ihis
1994). Percentages
1996).
punctures
present in the Maastrichtian
first
the Cre-
(Westermann,
constrictus
same
am-
in
known from the Upper Cretaceous of USA
(N.H. Landman,
m
planktonic
high percentage
(Radwahski,
Identical holes in the
bers
in
Jakobsen,
appear
based upon field observations,
Carpiliidae
1997).
&
(Collins
swimming crabs found
assumed to be the result of
tacean attacks
taceous
ten first
of faunal elements in the Middle Danian of
persistance
spp.
Hoploscaphites
or
swimming
1996a,
source
of the
very
percentages
In
1991; Fraaye,
(Bishop,
monites such
and
(Denmark).
their turn these
well-adapted
a
species
rapidly
commonest
group
decline in the
nommids reached their
ber and
overlying member. Tory-
acme
rapidly declined
Member, probably
as
a
in the Nekum Member,
in
in the Nekum Memthe lower Meerssen
result of
competition with
124
R.H.B.
4.
Fig.
Percentages
presumably
the
for
decapod
crustacean
families
efficient swimmers
more
the Xanthidae (Fraaye,
1997);
some
in
the three
amongst
of these latter
Fraaije
highest
members
Dynomenidae and Flomolidae display the same trend
The
Tethyan
as
(Fraaye,
pagurids going up-section
1996a).
from the Nekum to
the Meerssen members. The Galatheidae and Car-
pi I iidae make their first
Member
(Fig. 5), but
families.
sive
in the Meerssen
appearance
are
far outnumbered by other
Within the interval
decapod assemblages
follows
five
studied,
may
be
succes-
recognised,
as
sen
diversity (Fig.
dead shell material grew
spectrum
by
a
was
an
up
increase
section. The
accumulation)
as
(ecologiin benthic
‘taphonomic feed-
Kidwell & Jablonski (1983). The
predominantly
Meers-
Accumulations of
markedly
referred to
et
Meerssen
mem-
ah, 1980; Voigt, 1981).
1981), fora-
(Voigt,
1981) and calcareous algae
communities
erosion within the
preservation
its
distinctive and,
are
and other
if
they
forms of associated
biotope, they stand a high chance
1975).
(Brasier,
ability
Sea
is
grass
no-
influence the character of
to
sediment substrate.
The
dense
plant growth
reduced current velocities,
whereas the
faunal filter feeders, e.g., Protocallianassa faujasi,
of live/dead interactions
cal consequences of shell
communities
6).
area.
rhizomes stabilised the accumulated sediments. In-
linked with
intimately
type
during deposition of the
bryozoans
as
channeling
probably
calappid/dynomenid
of substrate
from
escape
the
members is
back’
grass
raninid/diogenid/calappid
diversity increase between the Emael and
entire
sea
(II)
(V)
the
(Brasier, 1975). Sediments which accumulate around
table for
(IV) calappid/xanthid/raninid
The
such
minifera (Sprechmann,
raninid
callianassid/calappid/torynommid
in
the colonisation of small
grasses enabled
sea
(I)
(III)
1978; Felder
(Liebau,
epizoans
of
(in ascending order):
Formation
Maastricht
uppermost Emael, Nekum and lower
bers
represent
of the
time
of reef decapods through
of seagrass fields
currence
The
invaders
Evolution
-
soft-bottom
change
dwelling
com-
thrived in the nutrient-enriched sediments around
sea
grasses,
plant
making
material
(Sven
dead hard parts
use
et
of
things such
of the
decaying
facili-
grass-epibionts
sea
tated the colonization of other
sponges, corals,
as
ah, 2001). The accumulated
epizoans such
as
fungi,
brachiopods, serpulids, lunulitiform
bryozoans, boring and encrusting bivalves, sessile
gastropods and cirripedes.
Member (units IVf-1 to
-4)
the lower Meerssen
finally
munity in the lower Emael Member (and underly-
led
ing members),
vertically and laterally alternating with hard grounds
to the firm
dwelling communities
members,
was
ground and shell-gravel
in the Nekum and Meerssen
probably triggered by
the
mass
oc-
to
(Voigt,
the
In
this colonization
development of small-sized bioherms,
1974;
van
fillings (Zijlstra,
den
Elsen,
1985)
1995). Cavities
in
and relief in-
hardgrounds
Contributions
Fig.
5.
Decapod
to
Zoology, 72 (2-3)
crustacean
species
range
-
125
2003
chart in
the
Maastrichtian type
area.
R.H.B.
126
Fig.
6.
Major palaeoenvironmental
played
an
systems
The
important role in
as
refugia
ber and size of
Kudla,
on
of holes
own
Maastrichtian type
(Voigt,
and
eco-
1959).
crevices
positively correlated with the
in
num-
decapod crabs and shrimps (Reaka-
1990). These decapod
their
the
Late Maastrichtian
and domiciles
number and size
reef substrates is
factors in
crustaceans each prey
and each others’
juveniles,
forcing
Fraaije
area
-
which
Evolution of reef
influenced
the larvae
inshore at
(caused
to
settle in
larger
by
decapod
sizes.
decapods through
crustacean
communities,
deeper habitats and migrate
The
infrequent availability
storm disturbance and
occupation
competitors) of holes and adult-juvenile
interactions
tion of
over
was
decapod
time.
time
by
predatory
probably important in the evolucrustaceans
in reef communities
Contributions
to
Zoology,
Abele (1974,
ber of
2003
-
127
habitat is
a
num-
function of the
a
complexity (= number of substrates) of
that habitat The
also
(2-3)
1982) documented that the
decapods in
structural
72
author
same
concluded that
similar numbers of
(Abele,
1976,
habitats
comparable
1982)
support
decapods, regardless of size of
the overall available
to this
species pool. Referring
fragility of decapod
trast to
out
some
ticular raninids
preserved
best groups suited
in
situ,
hypothesis, the habitat of the Middle Danian fauna
preservation
be attributed further to the low
most
closely approximates that
of the Nekum Member in the Maastricht
of
occurrence
Raniliformis baltica in
The
area.
both
settings
the
of
high pH
decapod
remains
crustacean
organic
of the warm-water
ronment, which would buffer
A second bloom of Raninoides
well.
as
The crab
fauna also
points
to
the
environmental similarities between the Nekum and
ferences in
Geulhem
material from the Gronsveld and
members in
servation backed
The
comm.).
the
the Maastricht area,
by crinoid data (J.W.M. Jagt,
most
depositional
environment of
a
substantial
ture, indicated by the drastic
all
bottom-dwelling
the Nekum and
drop
in tempera-
decline of bioherms
fossil
started
1986).
This
the
groups, with
possible exception of echinoids (Felder,
den Elsen,
pers.
important difference between
Geulhem members is
and
ob-
an
1981;
van
drop of temperature already
in the upper half of the Meerssen Member
(sections IVf-5 and -6) where decapod
remains
extremely
are
rare
or
crustacean
absent.
fected
in
morphology
bers, shows this species
striking
comparison
Valkenburg
and
tus
are
the
ing members, whereas in
mented
smaller
the
Nekum and
highly
occurs.
sculp-
underly-
Meerssen Member
sized and but
morphotype
and
of Eumorphocorystes
only known from
significantly
More
partially
et
are
ah, work
ontogenetic
under
or
a
orna-
specimens
needed to determine whether these different
photypes
to
mem-
depth within the
type Maastrichtian. Thus far, large-sized
specimens
in
dif-
to have been the least af-
by changes in substrate
ornamented
ak, 1988).
et
quadrispinosus
lower Meerssen Member, without
carapace
and
depositional envi-
produced by soft-tissue decay (Plotnick
ronment
may
content
the effects of acids
Den-
depositional
the
and Meerssen
envi-
similar
par-
evolutionary rela-
mark)
a
in
amongst
(Danian of Limburg [Geulhem Member] and
might suggest
con-
Naturally, the excellent
area.
of Fakse
(Denmark)
are
tionships within the Emael, Nekum
members in the
type
and
crabs,
document
to
in
fragments, would rule
Therefore,
reworking.
any
crustacean carapaces,
of their claw
are
mor-
phylogenetic (van Bakel
way).
Zijlstra (1995) documented storm-induced sedimentation
ates. The
cross
and
ot
cycles in the type Maastrichtian carbon-
highest degree of sedimentary,
e.g.,
trough
Acknowledgements
beds, channels, spillover and tempestite sheets,
taphonomic features,
the
e.g.,
serpulid Pyrgolopon
bioclasts, indicating high
parallel orientation
and
elongated
other
I wish
for
deposits,
ber (sections
that
occur
in the middle Meerssen Mem-
This is
supported by
of
preservation potential
the near-uniform
carapaces in the
Hinte prepared
position is the
1951). The successive change
orphology (ornament
and
&
convexity)
many
most
crabs
in carapace
within
the
van
Raniliformis and Eumorphocorystes (Fraaye
Bakel,
°f such
a
1998) is indicative of
depositional environment
bonarily significant period
G.
and Boreal
der Zwaan
of this
improvement
to
figures. This
van
Meulenkarap (Utrecht)
is a
paper.
contribution to
T.
IGCP
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Sedimentology
of the
Early Tertiary (Tuffaceous) Chalk
Geol.
Western
Symp. Cephalo-
Fraaije
119:
1-192.
Late
of
Cretaceous
Northwest
Eu-
Received:
15
March
2003
Crushing
Exp.
Mar.
behavior
Biol.
31:
of
time
tropical
155-172.