Taphonomy of a Modern Shrimp: Implications for the Arthropod Fossil Record

Author(s): Roy E. Plotnick

Source: PALAIOS, Vol. 1, No. 3, Theme Issue: Taphonomy: Ecology's Loss is Sedimentology's
Gain (Jun., 1986), pp. 286-293
Published by: SEPM Society for Sedimentary Geology
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286 RESEARCHREPORTS

Taphonomyofa ModernShrimp:Implications
for
theArthropodFossilRecord
ROY E. PLOTNICK

ofGeological
Department ofIllinoisat Chicago,Box 4348, Chicago,Illinois60680
Sciences,University

PALAIOS,1986, V. 1, p. 286-293 tia,"is extremely poor,despitethegreatabundanceofthese

groupsin modernmarineenvironments (Schafer,1972). The
Everyliving organism representsa potentialfossil,although only fossilrecordofthe"Reptantia" (crabs,lobsters,spinylobsters,
a fewareeversuccessfully fossilized.Taphonomic studiesshould etc.) is betterbutstillquitelimited formostgroups.The critical
explainepisodesof bothfossilization and of non-fossilization. questionis this:Arethesedisparities inrepresentation "real"
Thispaperexamines someofthetaphonomic variablesthatmay (e.g., due to changesin taxonomicrichness),or are they
biasthearthropod fossilrecord. artifactsof differing preservation potentials? Studiesof the
The short-term preservation potentialof themoderncarid taphonomic processesaffecting arthropods are crucialforan-
shrimp, Pandalusdanae,was studiedin a variety oflaboratorysweringthisquestion.
andfieldsettings. Freshly killedspecimens were buried at two Exceptforthebasiccomparison ofchitinous versuscalcified
intertidallocalitieson San JuanIsland,PugetSound, Wash- exoskeletons (Brooks,1957;Schafer,1972;Glaessner,1969),
ington.Carcasseswereburiedat depths of5-20 cmforperiodsof themorphological andecologicalbases fordifferences inpres-
onedaytothree weeks.Destruction byscavengers (crabs?)wasthe ervationpotentialof different arthropod groupshave never
probable primary cause ofcarcassdestruction. Furtherbreak- beenexamined indetail.Thispaperreviewssomeoftheknown
downwas causedbybacterial decomposition and disturbance by factors in arthropod taphonomy andreportson a pilotexperi-
burrowing infauna. mentalstudyofthedestruction ofcarcassesofthecaridshrimp
Shrimpremainswereplacedin a seriesofglassjars in the Pandalusdanae.
laboratory.Jars differed in thepresenceand kindofenclosed
sediment and in thedegree ofaeration. Decomposition destroyed Arthropod Taphonomy
nearlyall softtissueswithin a periodoftwoweeks.Thecuticle
became extremely soft,resultingin lossofphysical integrityofthe Nearlyall studiesof arthropod taphonomy are a posterionr
remains. Differences betweenoxicandanoxicdecomposition were interpretations of fossiloccurrences,such as the excellent
minor. papersbyBishop(1981)andSpeyerandBrett(1985).It should
Theseresults,alongwitha consideration of thebiological, be recognized,however,thatthese interpretations are of
physical,and chemicaleffects ofbioturbation, suggestthatdis- somewhatlimited value,fortheyexplainonlysuccessfulfos-
turbance byscavengers orburrowing infaunais a majorfactor in silizationevents.The reasonsfor,andtherelativefrequency
thedestruction of buriedarthropod remains.The absenceor of,non-preservation remainunclear.
inhibitionofbioturbation maybe a necessary condition forar- Mostactualistic studiesoffossilization haveconcentrated on
thropod preservation. Thepreservation potential ofarthropods, heavily shelled,easily preserved organisms such as molluscs
and of othersoft-bodied forms,mayhave declinedsince the (e.g., Chave,1964; Driscoll,1970; Stanton,1976; Warmeet
Paleozoic. al., 1976). The onlyextensivediscussionof biostratinomic
processes affecting modernarthropodsis that of Schafer
(1972), whichdocumentsthe formation and breakdownof
arthropod carcassesandmoltsintheNorthSea.
INTRODUCTION We canidentify a number ofpossiblebiostratinomic andearly
The relativerepresentation ofdifferent taxonomic groupsin diageneticfactorsaffecting arthropod carcasses and molts.
the fossilrecordis, of course,highly variable.Tremendous These include1) physical disturbance, transport, anddestruc-
differences in thenumbersoflivingandfossilgeneraofdec- tion,2) predation and scavenging, 3) bacterialdegradation of
apodsbelonging to variousinfraorders canbe seen inTable 1. softtissuesandtheexoskeleton, and4) chemical breakdown of
The fossilrecordofthevarious"shrimp" groups,the"Natan- theremains. As discussedbelow,thesefactors arenotindepen-
Copyright? 1986, The Society of Economic Paleontologistsand Mineralogists 0883-1351/86/0002-0286/$03.00

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 OF A MODERNSHRIMP
TAPHONOMY 287

TABLE1-Fossil record of decapod crustaceans, based on carcassesinforestlitterinNorthCarolinaandGeorgia.They

Chace (1951), Holthuis (1955), Hobbs (1974), Glaessner foundthatthe remainsdecomposedfairly slowly,fiveyears
(1969), and other sources. beingestimated as thetimenecessaryforcompletedisappear-
ance ofmillipede andcricketexoskeletons. Softtissueappar-
entlydisappeared within thefirst twoweeks(see alsoSeastedt
Modern andCrossley,1984).Okafor (1966b)studiedthedecomposition
Genera Total rateofburiedinsectwings.He was able to recoverrecogniz-
Modern w/Fossil Fossil ablefragments ofdesert-locust wingsafter300 days.
Infraorder Genera Record Genera The decomposition of marinecopepodswas examinedby
Harding(1973). Comparedwithterrestrial arthropods, the
Penaeidea 33 2 16 breakdown is rapid.Depending onwatertemperature, bacterial
Caridea 184 3 13 decomposition totally destroyed theremains within3-11 days.
Stenopodidea 7 0 0 Arthropod cuticleis composedof a complexof chitin(a
Uncinidea 0 0 1 mucopolysacharride) and proteinthatis frequently calcified
TOTAL "NATANTIA" 224 5 30 (Richards,1951). A numberof studieshave examinedthe
decomposition ofthechitin fraction.Okafor (1966a)determined
Anomura 92 9 24 thatthe maindecomposing organisms were fungi,bacteria,
Astacidea 45 6 31 actinomycetes, nematodes,and protozoa.Bacteriaand fungi
Brachyura 632 86 263 are also responsible forthedecomposition ofchitininmarine
Palinura 16 6 38 environments (Harding, 1973; Chan,1970; CampbellandWil-
Thallasinidea 20 6 10 liams,1951;Lear,1963;Seki,1966;SekiandTaga,1963a,b,c;
TOTAL "REPTANTIA" 805 113 366 Kohlmeyer, 1972). Mostofthese"chitinoclastic" bacteriaare
aerobic,although some are facultatively anaerobic(Campbell
andWilliams, 1951).
Chitinoclastic bacteriaare widelydistributed, beingfoundin
dent.In particular,theyare all eitherdirectly or indirectlysea water,sediments, inanimalguts,andonarthropods (Seki,
relatedto bioturbation. 1966; Chan,1970). Chan(1970) foundhighconcentrations of
chitinoclasticbacteriainintertidal andfresh-water sediments;
Predation and Scavenging thehighestdensitieshe observedwere on moltedarthropod
CoullandBell(1983)indicated thatpredation is animportantexoskeletons. Similarly, Lear (1963) discovered thatwhilethe
controlon the structure of arthropod assemblages.Known concentration ofchitinoclastic bacteriawas lowin pelagicwa-
predatorson largearthropods are fish,birds,cephalopods, ters,itwas quitehighon thezooplankton.
crocodilians,andcrustaceans (CoullandBell, 1983;Boyleand Seki (1966) examinedthedecomposition ofchitininmarine
Knoblach,1981; Nixonand Boyle,1982; Kneiband Stiven, sediments. The decomposition ofchitinis apparently farfaster
1982;Warner, 1977).Somecrabsareknown tobe cannibalisticin the upper,aerobiclayerof sediments;nearlyall chitin
(Warner,1977). Scavenging has been describedas a feeding breakdown occursinthislayer.A markedslowdown or cessa-
habitofcrabs,horseshoecrabs,sea urchins, and gastropods tionofbacterial activity underanoxicconditions was alsonoted
(Shuster,1982;Levinton, 1982).Schafer(1972)describedthe byChan(1970).
scavenging oflobstercarcassesbyworms,gastropods, star- Finally,Chan(1970) examined therateofchitin decomposi-
fish,fish,isopods,and otherdecapods.Onlyheavilycalcified tionas a function of particlesize. As can be expectedfrom
portionsof crab and lobsterbodiesare usuallyresistantto surface-area considerations, the decomposition rate of small
destruction. Schafer(1972,p. 128) suggestedthat"thebest particleswas farfasterthanthatoflargeparticles.This sug-
conditions forfossilpreservation of animalswithsegmented geststhatfragmented arthropod remains,suchas thosepro-
skeletonsare thosepartsofthedeep sea hostileto life." ducedbypredators andscavengers, shouldhavefasterratesof
To feedonbenthic infaunalorganisms andburiedcarcasses, breakdown thanwholearthropods.
thepredatoror scavengermustpenetratethe sedimentsur- A majorfactor inthedestruction ofmanyarthropod remains
face. This penetration strongly disturbsthe sedimentand is thedissolution ofthecalciteintheexoskeleton (Weidemann,
causesbothphysical andchemical changesinit(Woodin, 1983; 1972;BasanandFrey,1977).Aller(1982)found thatextensive
Thayer,1983; see below).Woodin's"weasel,""digging," and dissolution of calcareousmaterialtakes place near the sedi-
"burrowing" predators disturb
substantially thesediment. For ment-waterinterface,in the presence of carbonate-
example,bluecrabs(Callinectes) andhorseshoecrabs(Limu- undersaturated porewaters.Ratesofdissolution werelowest,
lus)producepitsranging indepthfrom 3-9 cm(Woodin, 1978; and shellpreservation best, in areas wherephysicaldisturb-
Botton,1984).The destruction ofburiedremainsbypredators anceprevented theestablishment ofa deep-burrowing infauna.
andscavengersis thuscloselytiedto sediment disturbance.
FIELD AND LABORATORYSTUDIES
Decomposition
Onlya few studieshave examinedthe decompositionof A pilotstudyofthepreservation ofPandalusdanae
potential
arthropodremainsinmarineor terrestrial
environments.
Sea- (Decapoda:Caridea)was conducted at theFridayHarborLab-
stedtand Tate (1981) examinedthebreakdownofarthropod oratoriesof the University
of Washington. The goal of the

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288 PLOTNICK

60 GARRISON
50 BAY

40 40- fALSE
GARRISON
BAY % 30 30- BAY
SAN JUAN mmmmmmmmmmm,. 20 % 20-
| ISLAND +_ 10 km
10 10-
0-10
-2 -1 0 1 2 3 4 5 -2 -1 0 1 2 3 4 5
4) ~~~~~~~~~4,
FIGURE2-Results of grain-sizeanalysis of sediment at burialsites
FALSEJ\
at GarrisonBay and False Bay.

odor). Both the False Bay and GarrisonBay sites supported

large populations of crabs (Cancer) and had locally abundant eel
grass (Zostera).
An initialset of experimentsinvolvedthe burialof shrimpin
fouradjacentholes at each site (Table 2). Holes were dug to a
depthof 10 cm, shrimpcarcasses placed in the holes at depths
FIGUREI-Map of San Juan Island, Puget Sound, Washington,indi- of 5 and 10 cm, and the holes refilled.The depthand place of
cating field localities. burialwere markedby layers of orange-paintedsand. All four
sites at False Bay and one of the sites at GarrisonBay were
experimentswas to document the short-termpreservation re-excavatedat the end ofthreedays. The remainingGarrison
potentialofburiedshrimpcarcasses inmarinesedimentsand to Bay sites were re-excavatedafterone week, two weeks, and
identifythe causes of carcass destruction.This investigation three weeks, respectively (Table 2). Sediment from re-
focusedon the earlieststages ofpost-burialhistory,specifically excavated holes was wet-sieved in the fieldto recover any
the first3-4 weeks followingthe death of the animal. The remains.
rationalewas thatanydestructionofremainsduringthese early It was found,at the end of the initialthree-dayperiod, that
stages would effectivelyprevent fossilization.Subsequent bioturbation had greatlydisseminatedthe coloredsand at False
studies willdeal withlongerperiods of time. Bay, makingprecise reconstruction ofthe burialdepthdifficult.
P. danae, known as the "coonstripe" or "dock" shrimp or impossibleto determinethe originalsource
It was difficult
(Butler, 1980), is commonin the waters of Puget Sound. It is animal for recovered materials. In addition,the wet-sieving
oftenfoundlivingadjacentto dockpilings,wherethe specimens techniquedisruptedthe decomposed remains.A second set of
utilizedin thisstudywere obtained.Shrimpwere sacrificedby experimentswas thusset up at False Bay, in whichthe shrimp
freezing,then thawed and weighed on an analyticalbalance. remainswere placed at the bottomofmedium-mesh(about 0.5
Weightsrangedfrom3.92-11.50 g. cm) chicken-wirebaskets (open at the top). The baskets
greatlyfacilitatedlocationand removalofburiedremains.Two
BurialStudies carcasses were placed in each basket and buriedat depthsof5,
10, 15, and 20 cm (Table 2). These baskets were carefully
Materials and Methods excavated, examined,and thenreburied,at 1, 2, 3, 9, and 14
Field studies examinedthe "survivability" of buried shrimp days afterinitialburial.
carcasses at two intertidallocalities(Figure 1). The firstsite, Because initial results suggested that scavenging by
at False Bay, was an extensivetidalflatwithlarge populations macrofaunamay be important,a singlespecimen (no. 25) was
ofinfaunal organisms,especiallythelugworm,Abarenicola,and buriedjust below the surfaceat GarrisonBay, fullyenclosed by
the thalassinidcrustaceans,Upogebiaand Callianasa (Hobson, wire mesh. It was re-excavatedafterone week.
1967; Brenchley,1981, 1982; Wilson,1981). The sedimentsin
this study area were well-sorted sands (Brenchley, 1981;
Figure 2). Based on color zonation,the depth of the redox Results
The results of the burial experimentsare summarizedin
potentialdiscontinuity (RPD) in the sedimentwas about 5 cm.
The second site, at GarrisonBay, was also dominatedby an Table 2. Sample sizes ofthese preliminary experimentsare too
infaunalcommunity.In this case, however, it was primarily smallforstatisticaltesting.Amongthe key results are:
molluscan(e.g., Clinocardium,Tresius,and Tapes). The sed- 1. The "survivaltime"ofshallowlyburied(5-10 cm) carcasses
imentswere very soft,poorlysorted muds withlarge matrix- was very shortat both localities.
supportedclasts (Figure 2). The clasts were primarily empty 2. Survival time appeared to increase with depth of burial,
bivalve shells. These shells were abundantjust beneath the especiallybelow 10 cm.
sediment surface and made diggingdifficult. The sediments 3. Short-termpreservationwas apparentlybetterin Garrison
became anoxic within1 cm of the surface(based on color and Bay thanin False Bay.

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 OF A MODERNSHRIMP
TAPHONOMY 289

SHRIMP SITE WEIGHT DEPTH TIME REMAINSFOUND TABLE2-Description and resultsofburialexperiments. In

NO. (g) (cm) (days) the firstset of experiments, at False Bay and at Garrison
Bay, shrimpwere buriedin fouradjacent holes, withthe
experiments
I. Orange-sand depthmarkedbya layerof orangesand. Twoshrimpwere
1 FB-1 6.95 5 3 no remainsfound buriedineach hole,at depthsof5 and 10 cm. Ina second
2 FB-1 6.09 10 3 no remainsfound set of experiments, at False Bay,the shrimpwere placed
3 FB-2 6.50 5 3 fragments: carapace, inopen-topwire-mesh basketsand buried,eliminating the
legs,antenna need for orange-sandmarkers.Depths of burial varied
4 FB-2 9.25 10 3 from5-20 cm. A single shrimpwas buriedin a fullyen-
5 FB-3 11.50 5 3 leg fragments closed cage at GarrisonBay. "Time"refersto the number
6 FB-3 10.34 10 3 leg fragment,antenna of days between initialburialand recovery;FB = False
7 FB-4 5.46 5 3 leg fragments Bay; GB = GarrisonBay.
8 FB-4 4.09 10 3 leg fragments
9 GB-1 5.56 5 3 antennaonly
10 GB-1 7.48 10 3 nearlycomplete
12 GB-2 5.10 5 7 4. Totalcaginggreatlyincreasedthesurvivaltimeofa shal-
11 GB-2 6.30 10 7 twoantennas lowlyburiedcarcass.
14 GB-3 4.40 5 15 no remainsfound 5. Shallowlyburiedcarcasseswere usuallyreducedto frag-
13 GB-3 4.74 10 15 intactwithsofttissues mentsoflegs,antennae,andtherostrum.
16 GB-4 3.96 5 21 6. Duringlongerperiodsofburial(3 days-21days)carcasses
15 GB-4 6.57 10 21 smallleg fragment becamesofterand muchmoreeasilyfragmented by any
physical
disturbance.
experiments
II. Wire-mesh The resultsof the longer-term experiments are probably
17 FB-4 5.06 5 1 bothpresent;complete biasedbythesusceptibilityofremains to disturbance produced
18 4.84 2 one missing;leg byrepeatedexcavation.
fragment on surface The majorprocessesactingto destroyor disturb remainsof
3 veryfragile,somewhat P. danae appearedto be 1) removalof shallowly buriedcar-
broken casses byscavengers,shortly afterburial,2) disturbance and
9 rostrum onlyfound scavenging bysmaller infaunal
animalssuchas polychaetes, and
19 FB-5 3.92 10 1 bothcomplete 3) slower-acting bacterialdecomposition, leadingto loss of
20 4.24 2 bothcomplete,some physicalintegrity of the remains.Scavenging,possiblyby
colorloss crabs,maybe themajorfactorindestroying shallowlyburied
3 one nearlyintact, carcasses. Crabs are knownto burrowto at least 10 cm in
otherbadly searchoffood(V. Galucci,pers. comm.;Freyet al., 1984).
fragmented Whendeadshrimp werefedto crabs,remainssimilar to those
9 rostrum w/onesideof foundin the burialexperiments were produced(i.e., legs,
carapace;antenna rostra,and antennae).Moreover,protection of buriedcar-
w/base casses fromlargeepifaunal scavengersby a barrierof shell
21 FB-6 6.77 15 1 bothpresent clastsmayhavebeenresponsible forthegreatersurvival times
22 4.82 2 bothpresent, observedat GarrisonBay. This increasein preservation po-
noticeablysofter tentialof buriedcarcassesdue to the accumulation of dead
3 bothpresent,soft, shells near the sedimentsurfaceis clearlyrelatedto the
easilybroken conceptof taphonomic feedback,developedby Kidwelland
9 both?,veryfragile, Jablonski(1983).
brokenup
14 smallfragments only
23 FB-7 7.66 20 1 bothpresent JarExperiments
24 7.47 2 bothpresent,softer Materialsand Methods
3 bothpresent,soft, The processofbacterialdecomposition was stud-
ofshrimp
easilybroken iedinthelaboratory.Eighteenshrimp carcasseswereplacedin
9 both,fragmentary separateglassjars(Table3). Halfthejarsweresealedtight,so
14 smallfragments thatfully
anoxicconditions coulddevelop.The remaining jars
only wereclosedbya screenof333-micron "Nitex"mesh,so that
theywererelatively oxygenated. mud(from
Sixjars contained
III. Cage Experiment GarrisonBay), six containedsand (fromFalse Bay), and six
25 GB-5 7.04 5 7 intact onlysea water.The sedimentswere sieved through1-mm
screensto removelargerclasts and infaunaand pickedto

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290 PLOTNICK

TABLE3-Set-up and resultsofjar experiments.Oxicjars duringgentlehandling. Colorswere stillsomewhatrecogniz-

covered by fine mesh, anoxic jars sealed withlid and able. A pronounced odorofdecaywas present.
Parafilm. Key to results: nc=no change; i=intact; Threeweeks Withtheexception oftheshrimp injars with-
i(b)= relatively
intact,some breakup,possiblydue to han- out sediment(bothopenand closed),theshrimp had disinte-
dling; b=broken up, usually into numerous pieces; gratedintonumerous largeandsmallpieces.Fragments were
o = noticeableodor; s = mostof exoskeletonsoft. Allcuticlewas verysoft,so thatdisintegration
stillidentifiable.
resultedfromeven the gentlestdisturbance. The odorwas
SedimentAerationShrimp Results(days) somewhatless noticeable.
size (g) 1 3 7 14 21 28 Fourweeks Alloftheshrimp inopenjarshadbeenreduced
to numerous tinyfragments. Someoftheshrimp inclosedjars
Mud anoxic 4.45 nc i,o i(b),o b,o b,o,s b,o,s withmudorsea wateralonewerestillrelatively intact,butvery
7.73 nc i,o i(b),o b,o b,o,s b,o,s
4.65 nc i,o i(b),o b,o b,o,s b,o,s easilybrokenbydisturbance. The odorwas evenless notice-
able. The experiment was terminated at thistime.
Mud oxic 6.23 nc i,s i,o,s b,o b,o,s b,o,s Thebulkofthedisintegration ofsofttissuestookplacewithin
5.66 nc i i,o,s i(b),o,s b,o,s b,o,s thefirsttwoweeks.The initial rateofdecomposition seemed
3.62 nc i i,o,s i(b),o,s b,o,s b,o,s to be somewhatfasterin the open jars, but no significant
Sand anoxic 8.27 nc i(b) i(b),o b,o,s, b,o,s b,o,s differencewas seen afterthefirsttwoweeks.
3.25 nc i i(b),o b,o,s b,o,s b,o,s Decomposition was accompanied byoverallsoftening ofthe
4.91 nc i i,o i(b),o,s b,o,s b,o,s cuticle,breakdown ofarthrodial
membranes, loss ofothersoft
Sand oxic 6.44 nc i i,o i,o i(b),o,s b,o,s tissues,anda resultingloss ofphysical
strength ofthecarcass.
5.43 nc i i,o i,o i(b),o,s b,o,s All specimenswere easily disarticulated by slightphysical
5.25 nc i i,o i,o i(b),o,s b,o,s disturbance,especiallyafterthefirst
week.Individual piecesof
None anoxic 5.53 nc i i,o i(b),o i(b),o,s i(b),o,s cuticle,however,generally remained intact.
Because shrimp in
3.73 nc i i,o,s i(b),o i(b),o,s i(b),o,s thesea-water jars couldbe examined without openingthejars
6.18 nc i i,o i(b),o i(b),o,s i(b),o,s and manipulating the specimens,theyremainedintactfor
longerperiodsof time.Withthisexception,the processof
None oxic 4.83 nc i i,o,s i,o,s i(b),o,s b,o,s
4.82 nc i i,o,s i(b),o,s i(b),o,s b,o,s
decomposition appearedto be onlyslightly fasterwhensedi-
5.41 nc i i,o,s i,o,s i(b),o,s b,o,s mentwas present.
DISCUSSION
EffectsofBioturbation
andScavenging
removesmallinfauna. Natural porewaterswereretained. The The impactofbioturbation andscavenging ontheprocessof
shrimp werebarelycoveredbysediment. has not been considered,withthe exceptionof
fossilization
The jars wereplacedina largetank,through whichrough- briefdiscussionsby Thayer(1983) and Boucot (1981). In
(3-mmscreen),naturalsea waterwas passed. Water
filtered contrast,a floodofrecentliteraturehasdealtwiththeinfluence
temperature remained at or near120 C forthedurationofthe ofbioturbation on sediment chemistryandphysical properties
experiment. Blackplasticwas usedtomaintain darknessforall and on the organization of benthiccommunities (Larsonand
runs.Jarswereopened,examined, andresealedafter1, 7, 14, Rhoads,1983;Thayer,1983;Aller,1982;Woodin,1983;Tev-
21, and28 days. esz and McCall,1983). Bioturbation mixesthe top layerof
sediment, whichleadsto 1) reducedphysical stabilityofthetop
Results layerofsediment, withcorrespondingreduction inload-bearing
The resultsof thejar experiments are summarized below capacity andincreasedease ofresuspension; 2) increasedrates
andinTable3. ofcarbonate andmovement
3) increasedirrigation
dissolution;
One Day Afterone day,no noticeablechangeswere ob- of particlesleadingto the depressionof the RPD, so that
served,withtheexception ofa slight
fading colors aerobicbacteriacanliveat greaterdepthsinthesediment;
ofthebright and
oftheshrimp. Therewas a slightclouding ofthewater. 4) changesincommunity structure, theexclusionof
especially
Threedays Allshrimp withcolorsfadedin epifauna.
werestillintact,
some shrimp.Decay productswerevisiblein the water,ac- In the fieldand laboratory experimentsdiscussedabove,
companied in somecases byodor. shrimpcarcassesremainedintactin the absenceof external
Oneweek Allshrimp werestillintact, soft- disturbance
withnoticeable (e.g., sieving).Freshremainswere resistantto
eningof thecarapacecuticleof the shrimpin openjars. All roughhandling. As decomposition proceeded,thebreakdown
shrimpwere easilybrokenby handling, withthe carapace ofarthrodial membranes andinternalconnective tissuesled to
separatingfromtheabdomen.Decay appearedto be further disarticulation of the exoskeletonby even moderatedisturb-
advancedintheopenjars, especially inmud. ance (suchas thatcausedbyscavenging macrofauna andother
Two weeks Shrimphad begunto disintegrate noticeably, bioturbators).
withcarapacesseparatedfromabdomensandlegs brokenoff. These results,combinedwiththe knowninfluencesof
The cuticlewas verysoftinalljars andagain,mostso inthe bioturbation onsediment chemistry, andphysical
biology, prop-
was observed erties, suggestthatbioturbation
openjars. Littleor no resistanceto disturbance and scavengingof buried

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 OF A MODERNSHRIMP
TAPHONOMY 291

remains couldleadto 1) resuspension oftheremains, leadingto Byers(1984) criticized some oftheseideas, showingthatat
theirphysicaldestruction or to scavenging by epibenthic and least some Paleozoictrace-fossil communities were diverse,
pelagicforms;2) movement ofarthropod carcassesandmolts abundant, andcapableofburrowing to substantial depths.
intothezoneabovetheRPD, wheretheyare subjectto rapid Miller(1984)discussedthehistory ofbioturbation inmarine
decomposition by aerobicbacteria;3) directdigestionand andmarginal-marine settings. She indicated thatbioturbation in
breakdown of remainsby scavengers,withtheresulting re- thePaleozoicdecreasedfrom marine to fresh-water sediments
ductionin the averageparticlesize of the remainsgreatly andwas essentially absentinthelatter.Millersuggestedthat
enhancing therateofbacterial decomposition; 4) disarticulationfresh-water sedimentswere not bioturbated untilthe Mes-
andscattering ofdecomposing exoskeletons; and5) enhanced ozoic.
ratesofdissolution ofcalciticexoskeletons. In brief,bioturba- Consideration oftheeffects andhistory ofbioturbation sug-
tionshouldvastlyenhancetheratesofphysical andbiological geststhefollowing hypothesis:
breakdown ofarthropod remains.The absenceor inhibition of
If the absence or inhibition of bioturbation duringearly
bioturbation, as underanoxicbottomwatersor verydeep
withinthe sedimentmaybe a necessarycondition for the taphonomic stages is a necessary condition forarthropod
ofarticulated arthropods. preservation, and if rates of bioturbation have increased
preservation
It shouldbe notedthatthe degreesof sedimentand exo- through the Phanerozoic, then the preservation potentialof
skeletondisruption dependdirectly on thesize anddensity of arthropods andofother"soft-bodied" formsmayhave de-
scavengersandbioturbators. Smallburrowing organisms (rel- creasedovertime.
ativeto thesize oftheremains)maynotbe capableofsignif- This hypothesis has the following corollaries:1) Arthropod
icantlydisturbing theremains.In addition, processessuchas remainsshouldgenerally be restricted to environments where
earlyconcretion formation,whichmayoccurveryrapidly after bioturbation is excludedandshouldbe rareorabsentinheavily
burial,couldsignificantly influence theability ofanyinfauna to bioturbated sediments, and2) The fossilrecordofnon-heavily
disturb theremains.These factorsmayaccountforthepres- calcified and non-infaunal arthropods shouldbe betterin the
ervation ofarticulated crabsinCretaceoussediments thathad PaleozoicthanintheMesozoicor Cenozoic.
beenburrowed andpelleted(Bishop,1981). Ifitcanbe shownthatpreservation potential has decreased
through time,rather thanincreasedorremained constant,then
Bioturbation andtheArthropod FossilRecord:A Hypothesis studies of Phanerozoic diversity history will have to be reas-
sessed (e.g., Bambach,1977).
If the intensityof bioturbation has remainedconstant
throughout thePhanerozoic, thenits impacton preservation
willalso havebeenconstant (allotherfactorsbeingequal). A
numberof studies,however,strongly indicatea monotonic
increaseinbioturbation intensity overtime. SUMMARYAND CONCLUSIONS
Thedirectrecordofbioturbation is provided bytracefossils. The degreeto whichthearthropod fossilrecordhas been
Seilacher(1977) describeda generalincreasein trace-fossilbiased by preservational factorshas neverbeen accurately
diversity through the Phanerozoic.This increase,however, assessed. Potentialdestroyers of arthropod remainsinclude
occurred onlyinthedeep-marine "flysch" faciesandnotinthe bacterial andchemical decomposition, scavenging, andphysical
shallow-marine "neritic"facies.The increasein deep-water disturbance. These processesare eitherdirectly or indirectly
bioturbators is especiallystriking fortheUpperCretaceous. relatedto sediment disturbance bybioturbation.
Sepkoski(1982) andSepkoskiandBambach(1979) consid- Processes responsible fordestruction of the remainsof a
ered thecommonoccurrence offlat-pebble conglomerates in caridshrimp werestudiedinboththefieldandlaboratory. The
thelowerPaleozoictobe duetothelackofanabundant infauna. preservation potentialof the shrimpremainswas apparently
Similarly, LarsonandRhoads(1983)documented a progressive relatedtodepthofburialandtotheresistance topenetration of
changein sedimentary fabricsfromthe Ordovicianto the the overlyingsediment.Scavengingby crabs and other
Devonian,withDevoniansediments beingmoreextensively macrofauna mayhavebeenthemajorcauseofcarcassdestruc-
reworked. tion.Bacterialdecomposition anddisturbance bysmallburrow-
The mostcompletestudyofbioturbation intensity is thatof inginfauna wereapparently themajorcauses ofdestruction of
Thayer(1983).Thispaperconstituted a testofThayer'searlier deeper-buried or better-protected carcasses.
(1979) claimthatthe increasein sedimentdisturbance by Bacterialdecomposition was studiedina seriesofjar exper-
"biological bulldozers" led to a declinein the immobile soft- iments.Shrimp remains wereplacedinglassjarsandmonitored
sediment epifauna. He inferred an increasein all measuresof overa periodoffourweeks.Jarsdiffered in thepresenceor
bioturbation intensity withtime:innumber oftaxawithrapid- absenceofoxygenandinthepresenceorabsenceofsediment.
or deep-reworking abilities;in meanreworking depth;andin The carcassesdisintegrated slowlyover the duration of the
totalreworking. The basic patternshowsa slow but steady experiment. Decay of softtissues and reductionin cuticle
increasein bioturbation through thePaleozoic,followed by a rigidityledtotheloss ofphysical integrity ofthespecimens.As
more rapidand sustainedincreasein reworking intensity a result,the remainswere easilydisaggregated by physical
through theMesozoicandCenozoic.Thislatterincreasegen- disturbance.
erallycorrespondsto the "Mesozoicmarinerevolution" of It has been suggestedthatbioturbation, and the resulting
Vermeij (1979).SheehanandSchiefelbein (1984)andMillerand disturbance of sediment,has increasedthroughthe Phan-

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292 PLOTNICK

erozoic.Thissuggeststhatthepreservation potentialof"soft- BUTLER,T. H., 1980, ShrimpsofthePacificcoast ofCanada: Canad. Bull.

bodied"organisms has declinedthroughthePhanerozoic. Fish. Aquat. Sci., v. 202, p. 1-280.
This studymarksonlythe firststage in unraveling the CAMPBELL, L. L., and WILLIAMS, 0. B., 1951, A study of chitin-
decomposing micro-organisms ofmarineorigin:Jour.Gen. Microbiol.,
relativerolesplayedbyvarioustaphonomic variables.Results v. 5, p. 894-905.
obtained forlightly
skeletonizedshrimpmayhaveonlylimited CHACE,F. A., 1951, The numberof species of decapodand stomatopod
to formswithmoremassiveand resistantexo-
applicability Crustacea:Jour.Wash. Acad. Sci., v. 41, p. 370-372.
skeletons.In addition,taphonomic processes may be very CHAN,J. G., 1970,The occurence,taxonomy,andactivityofchitinoclastic
dependenton factorssuch as cuticlecomposition, sediment bacteriafromsediment,waterandfaunaofPugetSound[unpubl.Ph.D.
composition, andbioticdiversityandabundance.It is vitalto dissert.]:Seattle,Univ.Wash.
K. E., 1964,Skeletaldurability andpreservation,
in IMBRIE,J.,and
expandthese studiesto longertimescales, to use larger CHAVE, NEWELL, N. D., eds., Approachesto Paleoecology:New York,John
sampleswitha greatervarietyof taxonomic groups,and to Wileyand Sons, p. 377-387.
examinethese processesin different environments. Future COULL, B. C., andBELL, S. B., 1983,Bioticassemblages:Populationsand
studiesshouldleadto an improved understandingofthepres- communities, in VERNBERG,F. J., and VERNBERG,W. B., eds., The
ervationalprocessesthatshapethefossilrecord.Ifsuccessful, Biologyof Crustacea,Volume7, Behaviorand Ecology: New York,
thesemaychangetaphonomy from ananecdotaltoa predictive AcademicPress, p. 283-319.
DRISCOLL, E. G., 1970, Selective bivalve shell destructionin marine
science. environments, a fieldstudy:Jour.Sed. Petrology,v. 40, p. 898-905.
ACKNOWLEDGMENTS FREY, R. W., CURRAN,H. A., and PEMBERTON,S. G., 1984, Tracemaking
activitiesofcrabsand theirenvironmental The inchnoge-
significance:
Thisresearchwas conducted at theFridayHarborMarine nusPsilonichnus:Jour.Paleont.,v. 58, p. 333-350.
LaboratoriesoftheUniversity ofWashington andsupported by GLAESSNER,M. F., 1969, Decapoda, in MOORE, R. C., and TEICHERT,C.,
a University at ChicagoFacultySummerFellowship.
ofIllinois eds., Treatiseon Invertebrate PartR: Geol. Soc. Amer.
Paleontology,
and Univ.Kansas Press, p. 399-533.
Technicalsupportwas receivedfrom Tomasz Baumiller,Julie HARDING,G. C. H., 1973, Decompositionof marinecopepods: Limnol.
Wulff,andGabrielle Toutonghi.VinceGalucciis thankedforhis Oceanog., v. 18, p. 670-673.
assistanceat GarrisonBay. Detailedand fruitful discussions HOBBS, H. H., JR., 1974. Synopsisofthefamiliesandgeneraofcrayfishes
were held withMichaelLaBarbera.The designof the jar (Crustacea:Decapoda): Smiths.Contr.Zool., v. 164, p. 1-32.
experiments was developedafterconsultation withRobertC. HOBSON, K. D., 1967, The feedingand ecology of two NorthPacific
Aller.J.JohnSepkoski,Jr.provided compilation
theinitial of Abarenicolaspecies (Arenicolidae, Polychaeta):Biol. Bull., v. 133, p.
decapodgenericranges.DeborahJ.Stewartprovided technical 343-354.
HOLTHUIS, L. B., 1955, The recentgenera of the carideanand steno-
assistanceandadvice,andeditedthemanuscript. The editorial podidean shrimp(Class Crustacea, Order Decapoda, Supersection
comments of RogerD. K. Thomasand two anonymous re- Natantia), with keys for determination:Zool. Verhand., v. 26,
viewersare gratefullyacknowledged. p. 1-157.
KIDWELL,S. M., andJABLONSKI, D., 1983, Taphonomic feedback:Ecolog-
REFERENCES ical consequences of shell accumulation,in TEVESZ, M. J. S., and
MCCALL,P. L., eds., BioticInteractions in Recentand Fossil Benthic
muds:
innearshoreterrigenous
ALLER, R. C., 1982,Carbonatedissolution
New York,PlenumPress, p. 195-248.
Communities:
The role of physicaland biologicalreworking: Jour.Geol., v. 90, p.
75-79. responsesto
KNEIB, R. T., and STIVEN,A. E., 1982, Benthicinvertebrate
size and densitymanipulations ofthe commonmummichog, Fundulus
BAMBACH,R. K., 1977,Species richnessinmarinebenthic habitatsthrough
heteroclitus,in an intertidal
saltmarsh:Ecology,v. 63, p. 1518-1532.
the Phanerozoic:Paleobiology,v. 3, p. 152-167.
and neoichnol- KOHLMEYER,J., 1972, Marinefungideteriorating chitinof hydrozoaand
BASAN, P. B., and FREY, R. W., 1977,Actual-paleontology
keratin-likeannelidtubes: Mar. Biol., v. 12, p. 277-284.
ogyofsaltmarshesnearSapelo Island,Georgia,in CRIMES,T. P., and
LARSON, D. W., and RHOADS, D. C., 1983, The evolutionof infaunal
HARPER,J. C., eds., Trace Fossils 2: Liverpool,Seel House Press, p.
41-70. communities andsedimentary fabrics,in TEVESZ, M. J.S., andMCCALL,
P. L., eds., BioticInteractions in Recentand Fossil BenthicCommu-
BISHOP, G. A., 1981, Occurrenceand fossilization of the Dakoticancer
nities:New York,PlenumPress, p. 627-648.
assemblage,UpperCretaceousPierreshale, SouthDakota, in GRAY,
J.,et al., eds., CommunitiesofthePast: Stroudsburg, Pa., Hutchinson LEAR, D. W., 1963,Occurrenceandsignificance ofchitinoclasticbacteriain
Ross, p. 383-413. pelagicwatersand zooplankton, in OPPENHEIMER,C. H., ed., Sympo-
siumon MarineMicrobiology: Springfield,Ill., CharlesC. Thomas,p.
BOTTON, M. L., 1984, The importance ofpredationby horseshoecrabs,
Jour.Mar. 594-610.
Limuluspolyphemus, sand flatcommunity:
to an intertidal
LEVINTON,J. S., 1982,MarineEcology:EnglewoodCliffs, N. J., Prentice-
Res., v. 42, p. 139-161.
Hall, 526 p.
BOUCOT,A. J., 1981, Principlesof BenthicMarine Paleoecology:New
MILLER, M. F., 1984, Distribution of biogenicstructuresin Paleozoic
York,AcademicPress, 463 p.
nonmarine and marine-margin sequences: An actualisticmodel:Jour.
BOYLE, P. R., andKNOBLOCH,D., 1981,Hole boringofcrustaceanpreyby Paleont.,v. 58, p. 550-570.
theoctopusEledonecirrhosa(Mollusca:Cephalopoda):Jour.Zool. Lon- MILLER, M. F., and BYERS, C. W., 1984, Abundantand diverse early
don,v. 9, p. 1-10. Paleozoicinfauna indicatedbythestratigraphic record:Geology,v. 12,
BRENCHLEY,G. A., 1981, Disturbanceand community structure:An ex- p. 40-43.
perimental studyofbioturbationin marinesoft-bottom environments: NIXON,M., and BOYLE, P., 1982, Hole drilling in crustaceansbyEledone
Jour.Mar. Res., v. 39, p. 767-790. cirrhosa(Mollusca: Cephalopoda):Jour.Zool. London., v. 196, p.
BRENCHLEY,G. A., 1982, Mechanismsof spatialcompetition in marine 439-444.
soft-bottom Jour.Exp. Mar. Biol. Ecol., v. 60, p. 17-33.
communities: OKAFOR,N., 1966a, Ecologyof micro-organisms on chitinburiedin soil:
BROOKS,H. K., 1957, Chelicerata, Trilobitomorpha, Crustacea(exclusive Jour.Gen. Microbiol.,v. 44, p. 311-327.
of Ostracoda) and Myriapoda:Geol. Soc. Amer. Mem., v. 67, p. OKAFOR,N., 1966b,The ecologyofmicro-organisms on, and the decom-
895-930. positionof,insectwingsin the soil: Plantand Soil, v. 25, p. 211-237.

This content downloaded from 195.78.109.52 on Wed, 11 Jun 2014 10:49:20 AM

All use subject to JSTOR Terms and Conditions
 OF A MODERNSHRIMP
TAPHONOMY 293

RICHARDS, A. G., 1951,The Integument ofArthropods: Minneapolis, Univ. and Environmentally StressedAnimals:New York,AlanR. Liss, Inc.,
Minnesota Press, 441 p. p. 1-52.
ScHAFER, W., 1972, Ecologyand Paleoecologyof MarineEnvironments: SPEYER,S. E., andBRETT, C. E., 1985, Clusteredtrilobite assemblagesin
Chicago,Univ. ChicagoPress, 568 p. the MiddleDevonianHamiltonGroup:Lethaia,v. 18, p. 85-103.
SEASTEDT, T. R., and CROSSLEY, D.A., JR., 1984, The influenceof Ar- STANTON, R. J., JR.,1976, Relationship of fossilcommunities to original
thropodson ecosystems:Bioscience,v. 34, p. 157-161. communities of livingorganisms,in ScoTT, R. W., and WEST,R. R.,
SEASTEDT, T. R., andTATE,C. M., 1981,Decomposition ratesandnutrient eds., Structureand Classification of Paleocommunities: Stroudsberg,
contentsofarthropod remainsinforestlitter:Ecology,v. 62, p. 13-19. Pa., Dowden, Hutchinson and Ross, p. 107-142.
SEILACHER, A., 1977,Evolutionoftracefossilcommunities, in HALLAM, A., TEVESZ,M. J. S., and MCCALL, P. L., eds., 1983, BioticInteractionsin
ed., PatternsofEvolution:Amsterdam, Elsevier,p. 359-376. RecentandFossilBenthicCommunities: New York,PlenumPress, 837
SEKI, H., 1966, Microbiologicalstudieson the decomposition of chitinin
P.
marineenvironment. X. Decomposition ofchitinin marinesediments:
THAYER, C. W., 1979, Biologicalbulldozersand the evolutionof marine
Jour.Ocean. Soc. Japan,v. 21, p. 265-269. benthiccommunities: Science, v. 203, p. 458-461.
SEKI,H., and TAGA,N., 1963a, Microbiological studieson the decompo-
THAYER, C. W., 1983, Sediment-mediated biologicaldisturbanceand the
sitionof chitinin marineenvironment. I. Occurrenceof chitinoclastic
v. 27-34. evolution ofmarinebenthos,in TEVESZ,M. J. S., and MCCALL, P. L.,
bacteria inthe region:
neritic Jour. Ocean. Soc. Japan, 21, p.
eds., Biotic Interactions in Recent and Fossil BenthicCommunities:
SEKI,H., and TAGA,N., 1963b,Microbiological studieson the decompo-
New York,PlenumPress, p. 480-625.
sitionof chitinin marineenvironment. II. Influenceof some environ-
mental factors on the growthand activity of marine chitinoclastic VERMEU, G. J., 1977, The Mesozoic marinerevolution:Evidence from
bacteria:Jour.Ocean. Soc. Japan,v. 19(2), p. 35-38. snails,predatorsand grazers:Paleobiology,v. 3, p. 245-258.
SEKI,H., and TAGA,N., 1963c, Microbiological studieson the decompo- WARME, J.E., EKDALE,A. A., EKDALE,S. F., andPETERSON, C. H., 1976,
sitionof chitinin marineenvironment. III. Aerobicdecomposition of Raw materialofthe fossilrecord,in SCOTT,R. W., and WEST,R. R.,
chitinbytheisolatedchitinoclasticbacteria:Jour.Ocean. Soc. Japan,v. eds., Structureand Classification of Paleocommunities: Stroudsberg,
19(3), p. 23-31. Pa., Dowden,Hutchinson, and Ross, p. 143-170.
SEPKOSKI, J.J.,JR., 1982, Flat-pebbleconglomerates, stormdeposits,and WARNER,G. F., 1977,The Biologyof Crabs: New York,Van Nostrand
the Cambrianbottomfauna,in EINSELE,G., and SEILACHER, A. eds., ReinholdCompany,202 p.
Cyclicand EventStratification: Berlin,Springer-Verlag, p. 371-385. WEIDEMANN, H. U., 1972, Shelldepositsand shellpreservation in Quater-
SEPKOSKI, J.J.,JR.,and BAMBACH, R. K., 1979,The temporalrestriction naryandTertiaryestuarinesedimentsinGeorgia,U.S.A.: Sed. Geol.,
offlat-pebble conglomerates: An exampleofco-evolution oforganisms v. 7, p. 103-125.
and sediments[abstr.]:Geol. Soc. Amer.Abstr.w. Progr.,v. 11, p. interactionsin a densely
WILSON,W. H., JR.,1981, Sediment-mediated
256. assemblage:The effectsofthe polychaeteAbareni-
populatedinfaunal
SHEEHAN, P. M., and SCHIEFELBEIN, D. R. J., 1984, The trace fossil
cola pacifica:Jour.Mar. Res., v. 39, p. 735-748.
ThalassinoidesfromtheUpperOrdovician oftheeasternGreatBasin:
WOODIN, S. A., 1978, Refuges,disturbance, and community structure:A
Deep burrowingin the early Paleozoic: Jour. Paleont., v. 58, p.
440-447. marinesoft-bottom example:Ecology,v. 59, p. 274-284.
SHUSTER, C. N., JR.,1982, A pictorialreviewof the naturalhistoryand WOODIN,S. A., 1983, Bioticinteractions in recentmarinesedimentary
ecologyofthe horseshoecrabLimuluspolyphemus, withreferenceto environments, in TEVESZ,M. J. S., and MCCALL,P. L., eds., Biotic
otherLimulidae,in BONAVENTURA, J.,BONAVENTURA, C., andTESH,S., Interactions in Recent and Fossil BenthicCommunities:New York,
eds., Physiology and Biologyof Horseshoe Crabs: Studies on Normal Plenum Press, p. 3-38.

Strictobservationdoes not,or should not,lead to manyerrorsin work,whereas the interpretation of

facts must inevitablybe erroneous at times. But the fear of errormust not deter,once the facts are
straightand ifthe workerhas any faithin his own judgment,and it is by sequences of errorand
correctionthatthe truthis approximated.

-G. G. Simpson, 1933 (age 31)

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