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Predation of the Upper Cretaceous spatangoid echinoid Micraster

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DOI: 10.1201/9781003077831-141

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 Echinoderms through Time, David, Guille, Feral & Roux (eds) a) 1 994 Balkema, Rofterdam, ISBN 90 5410 514 3

Predation of the Upper Cretaceous spatangoid echinoid Micraste r

Nigel E.Cross & Edward PERose
Department of Geologl; Royal Hollowtt-v, Universitl; of LondorL UK

ABSTRACT: Many specime ns of MicrLtster corttnguinum. from the Uppcr-Chalk (.M. coranguinum Zor.el of
Lorrg Bredv in southein England sho$' test clanrage which may be ascrjbed to the r'vork of-predators. Some
sho..i borings into ol through the test, of lr'hich most appear to be altachment sites of acrothoracic bzLrnacles
ancl r:lionicisponges or exiavations by zinnelid worms. br-rt others are comparahle r'vith holes fornled by
p1'esent-daypirasiticorpreclatorygastr:opods. Nearly 10%o.f over'l00.specinlensshor'vtestfracturesinthc
ipicalregiirn,inferrecitol-.ethcr,voikofapreclatorattackingthegonadsbeneath. Almostall ofthetestsshow
pbst-morlenr encrustation, notably by cementing bivalves andlhe sponge Porosphaera and lcss con-rmonly by
irtyo.oa trnd serpulids, r,,,hich never pre-dates the fiactr-rres. Some borings-and tlrese particular fractures are
therefbre inteqricted as eviclence of irue pledation on living animals. Predation has been claimeri to be thc
clriving force behind many evolutionary irends in echinoids. and it is likeilr to have been a contributot'y factot'
tavouring an infaunal adaptive strategy also in iilicruster'

I INTRODUCTION gastropocls in the Late Cretaceous. However, his

ideas have beer-r disputed, and, so far as rve are
The fossil heart-ulchin Micro.\ter (Figure 1a) is arvare, thele is little published evidence that
locally common in the Upper Cretaccous (Turonian Mic r a s e r suti'ered predation.
I

to Campanian) Chalk of Europe and rvestern North
Africa. Studies by a succession of ar,rthors fiom the
19th century to the present day have documented and
interpreted scrial changes within the genus,
particrrltrr'1y within the M. leskei - M. decipiens ' M.
corungrinum lineage, that havc rnade it one of the
rnost famous examples of microevolutionary change
described from the fossil record. Molphological
changes. u,hether interpreted in terms of phyletic
gradualisnr (Rorve 1899; Kermack 1954; Nichols
1959) or punctllated ecluilibrium (Smith 1984;
Daviil & Foulay L98zl) as rnodels tbr evolution. ale
norv widely interpreted in tertns of burrorving
el'ficiency, either as an adaptrition to increasing
burrowing depth (Nichols 1959) or replacement of
sancl-drvelling by mud-du,elling ecomorphotypes
(Srnith 1984; McNamara 1990).
NIcNamara has rccently ( 1990) repolted that the
same or similar changes can be seen in a nurnber of
spatangoid Iineages clescribed trom the Tertiary of
southern Australia. notably Sch.izoster' Pttroster,
Hemia.ster, PsephotLster and Pericosttttrs. He inl'ers
that tlrc cl'ranges ure heterocht'onic. and lhitl Nlitttisie r
and St:hizastet' shorv very similar development of a
nnmber ol' pet'amorphoclines. Predation prcssure is
argr-iedto be the driving force behind nranv
evolutionaly trcnds secn in echinoids, ancl
McNamara suggests thJ,t il[icroster may have been
displnc96 into a deeper-$atei' nichc as it
l'csponse to the aclaptive radiatiot-t of predatory
We here arnplif.v our pleliminaly accollnt (Rose &
Cross 1993) of predation tn M.corangui.num on the
basis of specimens fi'on.r the Upper Chtrik (Santonian
Stage,M. corunguinum Zone) at l,ong Bredy
Quarry. near Dorchester'. Dorset. in sor-rthern
Englancl. The quarry (grid ret-erence SY567917) is
curl'ently being operated as a face over 100 m long,
i,r,orked in three tcrraces to a combined height of
about 20 m. Over 400 specimens have been
collectcd, all easily clcaned of thcir soft unweathered
Cl'ralk matrix without appreciable damage to the test
surface.
2 BORT\GOFTHETEST
Four different types ol boring through or into the
test have been recognized in the Long Bredy
Micra.sier specinens:-
2.1 Round holes indicating gastropod attack (Figure
1b)
Five specimens show a few small (0.5 - 1.0 mm)
round holes near the apical region of the test
comparable in size and shape with the possible
gastropod drill holes reported by Kier (i981) from
an Upper Albian (Lower Cretaceous) spatangoid,
Hemiaster, although Smith (1984, p.13) notes that

607
 a

Figure 1. a. Micraster coranguinum (Leske): aboral surface. Natural size. b. Round holes indicating
lQ mp c. Lozenge-shaped borings (arowed) indicating
gastropod attack. Scale ba1 uttu"f,-""t.
Scale bar 1 mm. d. Galleried borings (arrowed) of the sponge ? iona. Scale bar 2 nirn. "iriip"J"
frUrta. Uoiirg.
(anowed) of annelids . Natural size. f . Micraster coraiguiiun, showing hole broken thr;rth;d;i*;",
".
of the test. Natural size. (Atl illustrations are from speci"mens oi M. ,oringrinum froml_on! ni6a/
Dorset.) Qru-..y,

608
 evidence of gastropod predation is rare in fbssil
echinoicls, and that the earliest undoubted predation
borings that he has seen havc been in Uppcr'
Clletirceous (Coniacian) M ic r u.y t e r t ur o ne
Salthe in France. He notes aiso that parasitic
gastropods may attack sea urchins. but they
generally do not kill their hosts. The holes produced
by these parasites are quite dift'erent from those of
predators in that they tend to be less regular. and
characteristically have abnolral stereom deposition
:iround thc rvound. One of the Long
specirnens with scveral borings on the apical sulface
show's srvelling of the stereorn and developn'rent of
secondary tubercultrtion on the inner margins of the
hole. indicative of growth after the attack, iind
provinr that it took place on the living echinoid.
Fossil gastropods are not found in association lvitlt
spccinrens of Micrastcr at Long Brecly. but this ma1,
be a leaturc of their lor.v preserv;Ltion potential:
gastropods have an aragonite shel1 rnineralogy. and
this is rarely preserved in the British Upper Chtrlk
(Hancock 197(r).
2.2 Lo:enge-sha.ped boritt.gs indicutir.g cirripede
trtkrlmrtnt (Figure I c)
Many specimens show scatterecl srnall (0.5 nirn)
holes which are not round but lozerrge-shaped.
compalable in size and shape with the attachment
cavities excavated by.iuvenile acrothoracic barnacles
in carbonate. conimonly shell sr.rbstrates (Seilacher'
1969) . Most borin-9s are situated on the apical
surtace, where they appeal to penetl'ate to the test
interior. It is not^ however, possible yet to confirn'r
r.vhether these borings were into the test of a living
or a clead aninial- Scanning E,lectron Microscope
studies reveal no growth :nodification to indicate ljfe
continlled after test penetration. The concentration
of borings on the upper surface is consistent wilh
attack on ar-i echinoicl in life orientation. but
pediccilariae normally prevent lalval settlement on
living specirnens. Barnacles bore tbr attachment
only, not as predators. Trvo different lozense-
shaped boring n-rolphologies are recognisable on
these specimens and may indicate attack b.v t$,o
diflerent species of acrotholacic ciripede.
'2.3 Entobitr: .shttlloy,-gulleried borin,q.s of' tlte
sponge Cliona (Figure ld)
Borings on several specimens are stenomorphic.
following the constraints of space available in the
test, and when exhurncd show gallelies typical of
Clionu. Spongcs bole by the chemical removal oi
isolated silt sizecl elernents of sr-rbstrate. producing a
unicpre microsculptnre on the w,alls of their gallerics
(Ekdale, Brornley & Peurberton 198,1, p.1 16). Such
borings arc located on both apical and oral surface.
and clearlv represent sites of post-mortem
attachment.
2.4 Tubular borings of ntnel.id u;orms (Figure l e)
Long, almost straight to strongly curved single ol
r t si s fr orn bri,nching tubes, generaliy less than 1 mm in
diameter, are excavated within the test parallel to its
sulface in sevcral specimens. These are conparable
r.vith annelid borings rvidely illustrated from the
tbssil lecord (e.g. Carneron 1969), and well-known
fron-r dead tests of Recent echinoids.
Bredv Thus of four types of borings recognized in these
spccimens. three alc evidence only of post-mortem
attachment but one is evidence of some pledatory or
parasitic attack by gastropods on living inclividuals.
3 FRACTUREOFTHETEST
Thiltv tbur of over .100 specimens (ne;rrly 10%)
collccted frorl Long Breiiy in the last few ycars
shou' a hole broken through the test, approximateiy
10 mm in diameter. centred on or close to the apical
system (Fi-qure 1fl. Since the Chalk matrix in rvhich
these specimens are irnbedded is soft and has been
ver1, easill,but carefullv cleaned away, the fractures
are demonstrabiv not an artefact of preparation but a
pre-burial t-eature.
Since these holes are consistently- situated at the
rpie,rl lceion. it apperus rhlt thel wire ibrrned when
the animai rvas orientated in life position and
attacked flom above. The object of the attack was
presumablr.to break through to the gonads beneath,
the gonads being the majol structure of edible value
u ithin an echinoid test. Size of hole appears to be
generally ploportional to the size of test; large tests
appear to have largc holes and smaller tests smaller
ho1es.
The ed_ue s of the hoies norrlally show, a clean break
through the test. rvith no clue to the causative
animal. HorveVeL. a few tests show sclatch miuks
(Figure La) associated u,ith test fi'actr-ue, and a ferv
other specimens also show scratch marks (Figule
lb) r.r'hich are bettcr preserved since the test has not
been broken away. These parailel scratches
seeminslv indicate the pincer action of a crustacean
clarv or possibly the tooth marks of a fish, but
u'hether tl'ris animal was a predator trying to break
tllough the test, or merely rasping o1T epibionts later
settlinq on the clead test, is not entireiy clear.
Manl' of the specimens also exhibit damage in the
region of the peristome, rvhele the labrum is
comnonly damaged or absent. so this area may also
har.e been exploited by a pledator to gain entry to the
test. bllt there is as yet no proof that the damage was
biornechanical in origin and inflicted on a living
animal. rathel than through post-mortem pre-burial
sedimentary transportation. This part of the test is
very easily broken.
Smith (1984), following Moore (1966), has nored
that post-larva1 echinoids are preyed upon by a wide
range o1'animals including other echinoids. starfish,
gaslr'opods, decapod crustaceans, octopuses, fish,
birds, otters and man. There is no clue flom the
associated fauna at Long Bredy as to the identity of a
609
 -,'

Figure 2. a. Scratch rnarks associiited \Yith test tl'acture. b. Scratch rnarks on unfracturecl lest' c Cementing
biialve: Atretu nilsortnl. d. Encrusting sponge: Poro.sphttera. e. Cheilostome bryozoan. f. Serpulidworm
tubes: Seryrzla (Cvckt.serpul.a). (A11 ifiuitrati.-ons are 1i'om specirnens of M. t:ortmguitttnt fro:nt Long Bredy
Quarry, Dorset. Scate bai l0 rul on photomicrographs a.bl I mm on SEM photographs c,d,e'f')

610
potential predator. Otters ancl men. and indeed
appropriate birds, evolved long atter the Santonian
so can be ercluded flom consicleration. Starllsh c1o
not bleak open the test: gastropods and also
octopuses drill a smaller, neater hole than seeir in the
lractured Long Bredy specimens; most fish dcvotu
echinoids whole: othel fish and crabs and lobsters
tend to attack the peristomial region (at least in
regular echinoids) where spine protection is minimal:
stan,ing regr-rlar (and therefore jawed) echinoids
have been known to cherv an irregular hole in the
apical region of other regular echinoids
(Himmelman & Steele 1971) or attack the rnargins of
sand dollars. but have not been reported to attack
spatangoids. Even large gastl'opods such as some
cassids cut relatively srnall (c. 2 mm) r'egular holes
(McNamara 1991; McClintock & Marion 1993).
The srvimming crab Liocarcittus puber has been
observed (Crump, pers. co11111. I993) Lo feed on
regular sea nrchins (.ParttcentrotL.ts lividus) at Loch
Ine in Scotiancl bv cutting off aboral spines rouncl
the anus and bi'eaking the test by using a claw
Vertically like a hamrner, but a close prescnt-day
aralogue fol the test fractule observed in Micra.ster is
not yct knorvn.
.1 ENCRLTSTATION OF THE TEST
Nearly all of the ,100 specimcns show encrustation
of the test exteriol by onc or rnore orgurisms. The
most common encrusters are:-
1. Cernenting bivalves (Figure 2c). Valves of
Atreta nilsonnl, as small inclividuals (<5 tun) but
commonly in ltirge numbers, ol P -\cnodonte
rnutu.bilis, as larger inclividuals (c. l0 rnm) but
u5urll) ir) .nt:ril rrunthc|s. ocLur on n)osl \fecimen\.
2. Thc sponge Porosphoero (Figure 2d). This is a
ve.rvcolllron encrllsting organism, but it has a verv
delicate structure and has cornmonly sr-rffered pre-
burial :Lbrasion to a thin sheet covering parts of the
te st.
Other' less comnon encrusting organisrns include
cheilostome bry'ozoans (Figurc 2e), setpulici *'orms
(FigLrre 21). and the octocoral Molkia. A11 clear11'
settled on the test alier dcath of the animal and decrLr'
of thc surfacc sott tissues. notably the trlbe teet ancl
the rnuscles attachirrg spiucs to tubercles. since tht'r'
enc.rList dilectll, across the pores of the ambuiacra
arrd the tubercles ol the test in general. There is litt1e
jndication of pr:eferred encrustation of particular test
:ircas. :Lncl this together rvith the high densitv oI
encrlrstation sllggests that thc tcsts were exposecl t,,r
encrustation and overturning on thc original sea lloor
tbr a significant length of time. That sorne of the
encrusting bivalr,es were thelnselves bolecl by Iater
organisms is further er,idence fol prolonged
exposllre. or rervorking of the tests.
5 CONCLUSION
Encrustation appears never to pre-clate supposed
predatory fracture or boring of the test. lor
611
fractures/bolings never cut through any encrustirlg
organism- In a few cases, the converse can be
observed: an encrusting orgilnism grows acloss the
edge of the apical fi'acture, proving fracture to be the
earlier event. Several specimens showing apical
fi'acture have no encrusting organisms whatsoever,
presumably indicatin-e they became buried relatively
cluickly after death, and that fracture caused death oi
took place verv soon afterwards. If fracture at the
apical region was to gain access to the edible gonads
beneath, as seeills probable. a similar timing is
implicit.
Sr-rch predation on Iiving M. corangLtinutn is
perhaps unlikel-v if Nichols'(1959) interpretation of
the species as a potentially deep-burrowing form is
correct. It is, horvever, consistent with its
interpretation b-v Srnith (1984) and McNamara
(1990) as a shallou,-burrowing but mud-dwel1ing
ecomorphotype. Goldring & Stephenson (1972)
had ear'lier drawn attention to thc Lrck of Micraster
burrows observable in the Chalk, and Bromley (e.g.
1990) has also noted the poolpreservation potential
of shallow-tier as opposed to deep-tier trace fossils,
so the weight of evidence is now increasingll, in
favour of a shallou' rather than deep burlowing
adaptation.
Arguably, Mi.crttsrer may thlls provicle yet another
example of an echinoid evolutionary trend inlluencecl
by predation presslrle. its species adapting to a
cryptic, bulrou,ing habit not only in order to eat but
to avoid being eaten.
ACKNOWLEDCEMENTS
Although the authors are responsible for the views
expressed here, they have benefiteri from helpfnl
comment tlom Drs A B Smith, P D Taylor, and NJ
Nlorris of the Natural History Museum, London.
Prot-essor J NI Ltrwrence of the Univelsitl, of South
Floricla. and Dr R Crump of the Orieiton Field
Centre. Pembroke. Jane Pickard of Royal Hollor.vay
tvped the draft manuscript. The Roval Societ-v is
thanked for a travel grant to EPFR l'hich funded
prescntation of this paper at thc Sth International
Echirrodelm Conference.
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612
PROCEEDINGS OF THE EIGHTH INTERNATIONAL ECHINODERM CONFERENCE
DIJON / FRANCE / 6- IO SEPTEMBER 1993

Echinoderrns through Time

Edited by

Bruno David
Universitd de Bourgogne, Dijon, France

Alain Guille & Jean-Pierre F6ral

Ob s e nt at o ire O c d ano I o gique, B any ul s /M e r, F ranc e

Michel Roux
Universit€ de Reims, France

OFFPRINT

A.A. BALKEMA / ROTTERDAM / BROOKFIELD / 1994

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