Growth and regeneration rates of the calcareous skeleton of the Caribbean coralline sponge Ceratoporella nicholsoni: a long term survey

11 Pages·1999·4.1 MB·

by  Philippe Willenz

GROWTH AND REGENERATION RATES OF THE CALCAREOUS SKELETON OF
THE CARIBBEAN CORALLINE SPONGE CERATOPORELLA NICHOLSON!: A LONG
TERM SURVEY
PHILIPPE WILLENZ AND W.D. HARTMAN
Willcnz, Ph.&Hartman, W.D. 19990630: Growthandregenerationratesofthecalcareous
skeleton ofthe Caribbean coralline sponge Ceratoporella nickolsoni'. a long term survey.
MemoirsoftheQueenslandMuseum 44: 675-685. Brisbane. ISSN 0079-8835.
The growth rate ofthe aragonitic skeleton ofthe Caribbean 'sclcrosponge' Ceratoporella
nicholsonihasbeenstudiedbyinsitustainingofspecimenswithcalceininareeftunnel,28m
depth, near Discovery Bay, Jamaica. Experiments were performed up to five times from
1984to 1997onapopulationof1 specimensrangingfrom 10-20cmmaximumdiameter.In
each experiment small skeletal samples were removed from the periphery ofsponges, and
specimens were left in place for furtherstudieson growth and regeneration. Perpendicular
sections, ground to a thickness of about lOum, were photographed by fluorescence
microscopy. Annual skeletalgrowthrateswerecalculated from measurementsofthe linear
extension between calcein stained lines along growth axes. Data indicate that although
average annual growth rates remained in the same range for different periods
(214.6±54.5-233.3±33.0u.m yr1), significant differences occurred from one individual to
another within the same period. The annual growth rate ofa given individual also varied
significantly in time (191.1±30.0-269.9±37.0um yr1). A second population of smaller
individuals, measuredafterasingle period ofoneyear, revealed astrikingly loweraverage
annual growthrate(124.4±35.0u,myr1). Regenerationoftheskeletonofinjuredspecimens
was also characterised by an initial slowergrowth rate. Nevertheless, afterthe firstyear, it
was comparableto normal growth, and exceeded it slightly thereafter. This first longterm
study ofCeratoporella provides important information on the variability in growth rates,
with implications on the use ofsclerosponges as paleoenvironmenta! proxies. OPorifera,
selerosponges, coralline sponges, growth rate, aragonite, skeleton, regeneration,
calcein,Ceratoporellanicholsoni.
Philippe Wittenz (email:[email protected]) Departmentofinvertebrates, RoyalBelgian
}
InstituteofNaturalSciences. 29Rue Vautier,B-1000Brussels,Belgium: W.D. Hartman, Yale
Universin: Peahodv Museum ofNaturalHistory, P.O. Box208118. NewHaven, CT. USA
06520-8118: 18January 1999'
Although the first specimen of the coralline aragonitic skeleton is still unknown, more than a
sponge Ceratoporella nicholsoni was dredged century after its discovery,
offCuba in 1878, anddescribed asanewalcyon- B„ot]. d.i.rect fin.i.ng and, i.nd,i.rect tec,hni.ques
arian coelenterate more than thirty years later ?
t/HThTait.ck.tshoisn,spi1ef9ct1ii1et)si,wimtawsarsedniostcuonvte+irlie+tduhe(Hmair-dat-m1ma9n^6n0&s h,CDaevumesl^been&uls/gae.dtthoeefvamolr3umae)tre tuohsreimcgeiarloacwleitizhnarrisanttaerienodls
Gsoproenague,(H19a6r6t)m,aann&d sGuobrseeaquu,en1t9l7y0)s.hAotwnthteosbaemea (UW£iUeann2J &zioHpabrtmcharnon1o9lgo5g)i.etshe(iBatetnearvbiadseesd o&n
time, thanks to the increased use of SCUBA Dmffeli 1986; Druffel & Benavides, 1986) or
diving, the extent of the diversity ot 'sclera- f0Cllsjng on carbon and oxygen isotope studies
sponges' became evident (Hartman, 1969; (joachimski et al., 1995; B6hm et al., 1996).
Hartman & Goreau, 1970, 1975). Consideringtheestimated slowcalcificationrate
A.mong t.,he nine ,known species of,rCari,b,bean coof^ntvheisnisp^enctifeosr,etlhuecildatatteirnt^gelcohnngiVqlueneislairnefotnhieiamtoisotn
coralline sponges, Ceratoporella nicholsoni
secretesthemostmassivebasalskeletono calcium ^ethosdcsa,ehsowofeVteenr,shoafve thetp(oftceennttiuarlieasd)v.aDnitraegcet
carbonate. Despite the fact that the ecology and ofrevea u dataongrowthrates forshortertime
eulxttreansstirvucetluyreinovfesCteirgaattoepdore(lLlaanghaevte anl.o,w1b9e7e5;n scaies (v,ears to decades),
Willenz& Hartman, 1989), thegrowth rateofits Calcein was firstused in invertebrates tomark
676 MEMOIRS OF THE QUEENSLAND MUSEUM
TABLE 1. Ceratoporellanichohoni. Successiveinsitu MATERIALS AND METHODS
labeling with calcein (*). Abbreviations: Tl,
9.VII.1984; T3, 15.11.1985; T4, 29.IV.1986; T5, EXPERIMENTAL DESIGN. Two sizecategories
l.V.1987;T6, 1.V.I997. ofCeratoporellanichohoniwerestudied inareef
tunnel at depths ranging from 25-29m at Pear
Specimen Experimentalperiod Tree Bottom, 5km EofDiscoveryBay,Jamaica.
number Tl T3 T4 T5 T6 The largestindividuals, 10-15cmdiameter,were
4 * * * * * labelled with calcein (Fluka 21030) in situ
7 * * * * * without being removed from their substrate.
8 * * * * Labelling was performed from 1984 to 1997 at
9 * * * * intervals given in Table 1. After the initial
10 * * * * labelling in July 1984 (Tl) a second incubation
11 * * * * was performed six days later (T2), to test the
14 * * * *
16 * * * * *
17 * * *
27 * *
2') * *
DURATION
T1-T3 221 days
T3-T4 438days
T4-T5 363days
T5-T6 1 years
the newly deposited calcium carbonate of the
basal skeleton of Ceratoporella (Willenz &
Hartman, 1985). Subsequently,thischemical has
beenusedtorecordcalcificationamongstawide
variety oftaxa such as brachiopods, bryozoans,
molluscs and echinoderms (reviewed in Rowley
& Mackinnon, 1995). More recently it has also
beenemployedinstudiesofthegrowthdynamics
ofcalcareous sponge spicules (Ilan et al., 1996),
orto estimate thegrowthrate ofthe Indo-Pacific
corallinespongesAcanthochaeteteswellsi(Reitner
& Gautret, 1996) and Astrosclera willeyana
(Worheide, 1998).
From these studies, calcein appears to be
permanently bound to calcium carbonate that
forms in the presence of the dye, although the
chemistry ofthe process has yet to be studied.
Calcein has the advantages of fluorescing
brightly under UV light and having only weak
toxicity.
FIG. 1. Ceratoporella nichohoni. Natural growth
Several specimens of Ceratoporella pattern. Ground section from specimen no. 16
nichohoni, includingthefourindividualsusedin sampled in May 1997, viewed by epifluorescence
the first experiment by Willenz & Hartman microscopy. Successive labelingevents with calcein
(1985), were repeatedly stained and sampled at are indicated at apex of walls separating
pseudocalicles, along aragonitic skeleton (A)
different intervals over 13 years, in order to extension axis. Living tissue (T) is brightly
evaluate potential growth rate variations among fluorescent. N=natural growth axis, S=surface of
the sponges during extended periods oftime. livingtissue. (Scale bar=500um).
CALCAREOUS SKELETON OF CERATOPORELLA 677
TABLE2.Ceratoporellamcholsoni.Annualgrowthratesduringthe concentration of100mg/l. Bags were
1 yearexperimentedperiod. A, Kruskal-WallisANOVAonranks removed from thespongesafter 12or
(H=379.5 with 9 degrees of freedom; P<0.0001). B, All 24hrs.
pairwise multiple comparison procedures (Dunn's Method). NS
Forlargespecimens,samplesofthe
indicates no significant difference and * indicates significant
(P<0.05)difference.Bothstatisticsindicateasignificantvariability skeleton, with attached living tissue,
betweenspecimens (PO.005). about l-3cm in volume, were
removedwithhammerandcoldchisel
A. Specimen Median 25% 75% Mean N fromtheperipheryofthesponge,each
4 236.0 228.0 242.0 234.6 42 specimenwas leftinplace forfurther
7 224.0 221.5 228.0 224.3 29 growth and regeneration. Small
9 280.0 275.5 284.0 279.4 33 specimens were sacrificed after one
10 219.0 210.0 224.0 216.9 36 year. Following dehydration in a
11 248.0 242.0 254.5 249.5 37 gradedseriesofalcohols,sampleswere
embeddedin Spurr'smedium (Spurr,
14 290.0 281.5 296.0 287.3 61
16 232.0 228.0 242.0 234.9 85 1969). Sections,cutwith a low speed
diamond saw (Bennet Labcut 1010)
17 215.0 212.0 222.0 215.8 18 weremechanicallygroundonaseries
27 217.1 214.7 219.6 :16.5 28 ofdiamond grinding disks (Buehler
29 173.7 170.4 178.2 173.6 59 ultra-prepTM)usingasemiautomatic
grinder (Buehler Minimet 1000) to a
thickness of 5-10uin and observed
B. Specimen 4 7 9 10 11 14 16 17 27 29
under epifluorescence microscopy
4 -
(NikonOptiphot-2microscope,excit-
7 NS -
ation filter 340-380nm, barrier filter
'-) * * . 420nm).
10 * NS * - Growth increments of the
11 NS * NS * - aragonitic skeleton were established
14 * * NS * * - by measuring the linear extensions
16 NS NS * * NS * _ (in micrometers) between stained
17 * NS * NS * * * _ lines along growth axes at the apical
27 * NS * NS * * * NS _ edges of the wall separating two
29 * * * * * * * NS NS - pseudocalices, or, forthe most recent
period, between stained lines andthe
surface ofthe skeleton.
sensitivity ofthemethod. Although distinctbands
could be detected at a distance of about 4urn DATA ANALYSIS. Statistical analyses were
(Willenz & Hartman, 1985), interval Tl-2 was performedusingSIGMASTATandSIGMAPLOT
omitted in this analysis because ofthe shortness (Jandel Scientific) data analysis and graphics
ofthe time period involved. Additional smaller software. All linear extension measurements were
normalised as annual growth rate prior to their
specimens (ll-25mm diameter) were removed
analysis. Non parametric Kruskal-Wallis analysis
from the substrate and cemented in situ to ofvariance(ANOVA)onrankswasperformedto
Plexiglas plates (5 specimens /12x12cm plate) test two null hypotheses. 1) Ho: there are no
withepoxyunderwaterpatchingcompound(Pettit differences in the average growth rates among
PaintCono. 7050 & 7055). Plates were stored in specimens of Ceratoporella within a given
Plexiglasracks placed on a ledge ofthe tunnel at
the depth ofcollection. TABLE 3. Ceratoporella mcholsoni. Comparison
between linear annual growth rate and regeneration
To label the sponges with dye, the large speci- growth rate. Unavailable data dueto bioerosion in a
menswere individuallyenclosed withinaplastic specimen are indicated (*).
bag (of4L volume) that was secured around the
bbaansdes.ofInthtehespcoasnegeofwipltahtensylbeoanricnogrdssmaolrlrsupbebceir- Period Linearratgerowth Rgergoewnetrhartaitoen Specimens
mens, the Plexiglas racks securing the plates T3-4 230.5+61.2 194.2+42.2 7-9-10-17*
wereenclosedinaplasticbag.Calcein,dissolved T4-5 232.0+59.6 238.4+38.8 7-0_ 10- 11 - 17
in sea water, was injected in each bag to reach a T5-6 244.4+25.10 272.2+36.9 9- 10- 11 - 16
678 MEMOIRS OF THE QUEENSLAND MUSEUM
FIG. 2. Ceratoporella nicholsoni. A, ground section offragment sampled atT5, view atbase ofpseudocalyx.
Twopossibleorientationsofsectionplane indicatethatmeasurementcaneasilybebiasedwhensection isnot
parallel togrowthaxis(Tl'-T3' <T1-T3; T3'-T4' <T3-T4). T= livingtissue. (Scale bar=250um); B, ground
sectionoffragmentsampledatT6,viewatapexofwallseparatingtwopseudocalices.Narrowstructureofwalls
(arrow)preventsreadingerrors. Here,spacebetweentwowallshas been filledasspongegrewupward. (Scale
bar=250|am).
TABLE 4. Ceratoporella nicholsoni. Measurement period; 2) H : there are no differences in the
reproducibilitytest. AMann-Whitneytest indicates averagegrowthratesamongvariousperiodsfora
thatdifferencesobtainedfromtwodifferentslidesof givenspecimen(Sokal& Rohlf, 1981; Foxetal.,
the same sample arenotsignificantly different(P< 1994). Wherethe ANOVA onranksrejectedthe
0.005), except forspecimen 16 in periodT3-4.
null hypothesis the Dunn's all pairwise multiple
Specimen PeriodT3-4 PeriodT4-5 comparisonprocedurewasusedtodeterminethe
slideno. Mean n P Mean n P
groups that differed from each other (PO.005
7a 273.4 24 0.1120 287.6 24 0.0395
7b 260.9 20 266.6 21 level). A Mann-Whitney rank sum test was used
9a 268.1 35 0.0272 254.3 35 0.6680 where only two groups were to be compared.
9b 258.5 35 254.8 35 Significantdifferenceswere concludedat PO.005
10a 160.7 26 0.7190 204.8 26 0.7490 level. Datareproducibility was tested for fourof
10b 159.1 24 202.4 24 the largest specimens by comparing
16a 177.0 35 0.0054 211.6 35 0.7960 measurements from a second ground section
16b 189.7 35 211.2 35
prepared from the same fragment.
CALCAREOUS SKELETON OF CERATOPORELLA 679
350 35 i
—
300 300
250 250
— —
200 200
pm urn
—
150 150
— —
100 100
—
50
4 7 8 9 10 11 14 16
Specimens
FIG.3.Ceratoporellanicholsoni.Mean linearannual FIG. 4. Ceratoporella nicholsoni. Linear annual
growthrates(u,myr"')of4 specimensduringperiod growthrates(u.myr"')of8 specimensduringperiod
Tl-3 (221 days). Average of all specimens is T3-4(438 days). Conventions indicatedas in Figure
indicated. Numbers ofmeasurements are indicated 3. A Kruskal-Wallis ANOVA on ranks and all
withinbars.AKruskal-WallisANOVAonranksand pairwise multiple comparison test (Dunn's method)
all pairwise multiple comparison test (Dunn's indicateasignificantvariability between specimens
method) indicate a significant variability between (P<0.005),exceptbetweenspecimens 11 vs 14, 14vs
specimens (P<0.005), except between specimen 16, 8vs 16, 16 vs 10, and 10vs4.
numbers 4 vs 8.
Figures 4-6 present the same procedure for
RESULTS
periods T3-4 (438 days), T4-5 (363 days) and
T5-6 (lOyrs), respectively. Identical statistical
LINEAR ANNUAL GROWTH RATES. testsalsoindicateasignificantvariabilitybetween
specimens, except for pairs indicated in the
Observation ofground sections offragments of
captions.InperiodT4-5,abatchofsmallersamples
Ceratoporellanicholsoniinepifluorescentmicro- gave rise to an average annual value
scopy revealed successive labellingwith calcein
(Fig. 1 ). Figure 2A-B shows details ofthe bright (av1e2r4a.g4e±a3n5n.u0a|ligmroywr"th1)rarteeaocfhtihneglaorgnelryspheaclifmetnhse
aflnudoraetsctehnetabpaenxdsoaftwtahlelsbasseepaorfaatipnsgeutdwoocaulniictlse, m(2e3a3s.u3r±e4m5e.n0t(s.idmidynro"t1)r.evHeaolweavdeirre,cticnodrrievliadtuiaoln
respectively. It is shown that variations in the betweenthesizeofspongesandtheirgrowthrate.
orientation of sections can induce larger
measurementserrorsatthebase thanatthe apex. Table2presentstheresultsofaKruskal-Wallis
Consequently, only measurement at the apexes ANOVA on ranks on data available from the
were considered. longest interval (T5-6), showing a significant
variability between specimens (PO.005). An all
Figure 3 presents the means ofmeasurements pairwise multiple comparison procedure
of the four specimens of Ceratoporella
indicates in detail which pairs are significantly
successively markedduringthe firstperiod Tl-3
different.
(221 days), as well as the average growthrate of
the population. The results of Kruskal-Wallis Comparisonofthemeanannualgrowthratesof
ANOVAonrankstestindicatesignificantvariability specimens from one period to the other (Fig. 7)
betweenthemeans(PO.0001). However, aDunn's also indicates a significant variability
all pairwise multiple comparison procedure (191.1±30-269.9±37.0um yr 1), revealing that
determined that specimens no. 4 and 8 did not individual growth rate amongst Ceratoporella
differ from each other. was not steady either.
680 MEMOIRS OF THE QUEENSLAND MUSEUM
350
Average: 233.2+45.0 pm Average: 124.4+ 35.0 pm
—
300
250
200
pm
—
150
—
100
10 11 14 16 1 42 43 44 45 47 62 63 64 65 66 74 107 108 109
Specimens
FIG.5. Ceratoporellanichotsoni, 1 inearannualgrowthrales(umyr"')ol'A.9 largespecimensandB, 14smaller
onesduringperiodT4-5 (363days).Conventionsindicatedas in Figure3. A Kruskal-WallisANOVAonranks
andallpairwisemultiplecomparison test(Dunn'smethod)indicateasignificantvariabilitybetweenspecimens
(P<0.005),exceptbetweenspecimens4 vs 10, 8 vs 1 1,8vs 16, forthelargespecimensandspecimens42vs62,
45 vsM. 108 vs 109, 109VS 107. 107vs44 forthesmall ones. Bothpopulations havedistinctaverageannual
growth rates.
REGENERATIVE ANNUAL GROWTH
35C
RATES, Each sampling caused an injury to the
sponge, leaving a bare fracture ofthe skeleton 300
(Fig. 8A) Thelivingtissuerapidlyextendedover
—
this fracture within a few weeks and covered it 250
completely alter a month (observations reported
by a local diver). No detailed measurement was 2QQ
pm
done, but at most, after 200 days the naked
fracture was healed (shortest interval between 150
personal observations).
Subsequent labelling followed by re-sampling 100
in the healed zone (Fig. 8B) provided direct
observation on the regeneration pattern of the 50-
skeleton. Ground sections show that new walls
are progressively erected to form new pseudo-
calices perpendicularly oriented toward the
4 7 9 10 11 14 16 17 27 29
fracture(Fig. 9).
Specimens
Measurements ofthe extension ofthe skeleton
show that in the first period following an injury,
ttahhveeerasavageemreangoserpmeracelgiemlneiennresaatr(iFoginrgo.rwatt1he0,risTatalebolmweeear3s)t.uhraIenndttohhnee FITgKG5rr.uo-sw6kt6(a.h1l0-rCaWyetarrel)sa.ltitCousoprnnoAvryeNernlOt1li)VaooiAn'nsi1oc0ihnnsadlpirsecaocanrtiktemside.naassnLdiidnnuaerlFaliirngpguaraifenKrnw?u>i.asAle
subsequent periods, the regeneration rate multiplecomparison test(Dunn's method) indicatea
increased, exceeding progressively the normal significant variabilitybetweenspecimens(P<0.005),
average growth rate. except as indicated in Table2.
1
CALCAREOUS SKELETON OF CERATOPORELLA 68
350
300
—
250
200
100
—
50
T1-3T3-4T4-5T5-6 T1-3T3-4T4-5T5-6 Periods T1~3 T3"4 T4-5 T3-4 T4-5 T5-6
AAGR: 193.3+45.7 AAGR: 269.9+37.0 AAGR:211.5+43.4 AAGR: 263.0+24.2
350 350
10
300
—
250
200-
urn
—
150
100
50
T3-4 T4-5 T5-6 T3-4 T4-5 T5-6 Periods T3-4 T4-5 T5-6 T1-3T3^T4-5T5-6
AAGR: 191.1 +30.0 AAGR:266.3+25.5 AAGR: 268.1 +30,0 AAGR: 269.9+37.0
Fig. 7. Ceratoporella nichohoni. Growth rate variability within samples, from one experimental period to
another. A Kruskal-Wallis ANOVA on ranks and all pairwise multiple comparison test (Dunn's method)
indicate a significant variability between means (P<0.005), except for the following specimens and periods:
specimen4(T4-5vsT5-6),specimen7(Tl-3vsT4-5;T3-4vsT4-5),specimen lO(T4-5vsT5-6),specimen 1
(T3-4 vs T 4-5), specimen 16 (Tl-3 vs T4-5). Specimen numbers are indicated within each graph. AAGR-
Averageannual growthrate.
Obvious but unexpectedmarks from sampling skeleton, was abandoned. This was decided in
were noticed on several specimens when revis- considerationofthepotential effectstheantibiotic
iting them in May 1997 (Figs 8A-8B). Measure- might have on the abundant intercellular
ment of the extension of those particular symbiotic bacteria present in sponse tissues
regeneration areas indicates that someone had (Willenz& Hartman, 1989; Hartman & Willenz,
shown interest in our samples exactly one year 1990). Calcein, first used then on invertebrates,
before. Luckily, only one specimen had disapp- wasconsideredasamoreappropriatebenchmark
eared (specimen no. 8). tomeasure 'growth sincemarking'. Atthattime,
REPRODUCIBILITY OF THE MEASURE- there was no indication ofthe permanence ofits
strongfluorescenceforlongterm measurements,
MENTS. In order to test the reliability of the whereasthis study clearly showed that calcein is
method, measurements of pairs of sections stable enough to mark aragonite for at least 13
prepared from the same fragment forfour ofthe
years.
largest specimens were compared. A Mann-
Whitney test indicates that differences obtained Based on calcein-labelling experiments, the
from two different slides were not significant, average annual growth rates of Ceratoporella
except forone specimen (Table 4). nichohoni were shown to remain in the same
rangethroughoutthedifferentexperimentalperiods
DISCUSSION (214.6±54.5-233.3±33.0|im yr). Measurements
made over the largest time interval (lOyrs) are
Both tetracycline and calcein were used to obviouslymostindicativeofaverageannualgrowth
initiate this study (Willenz & Hartman, 1985). rates (233.3±33.0u.m yr"1). These values are
Firstexperimentsfoundthattetracyclinefailedto comparabletootherstudiesbasedeitheronindirect
labelthe skeleton ofCeratoporella. Althoughno radiometric methods [270|xm yr-1 using C, and
harm was apparent to the organism, further 220um yr"1 using 210Pb (Benavides & DrutTel,
experimentationinanattempttoadjusttheconcent- 1986; Druffel & Benavides, 1986), 180-260um
ration ofthe dye, to improve its recovery in the yr"1 (Joachimski et al., 1995), and 220|im yr"1
682 MEMOIRS OF THE QUEENSLAND MUSEUM
FIG. 8. Ceratoporellanicholsoni.A,Specimenno. 14afteraparticularlyseveresamplinginMay 1987.Arrow
indicatesfracturedaragoniticskeleton.Encircledzonewasfoundmissingin 1997. (Scalebar=5cm);B, same
specimen as inA, seen inMay 1997. Zone fractured in 1987 has healed and shows round edges created by
skeletonregeneration.Arrows indicatesharpedgeofamorerecentunexpectedinjury.(Scalebar=5cm).
CALCAREOUS SKELETON OF CERATOPORELLA 683
growth rate of a given individual also varied
significantly with time.
ItfiA
ill Variations in growth rate were shown to occur
in two other circumstances, suggesting that
young tissues produce aragonite at a slower rate
thantissuesofolder individuals. Firstly, measure-
ments on a second population of smaller sized
individuals, revealed a striking lower average
annual growth rate (124.4±35.0|im yr" ).
Secondly, injuries made to large specimens
inducedhorizontal regeneration zones which also
Ti appearedtoreducegrowthrate. Inthislattercase,
however, afteroneyear,growthwasrecordedbut
it was never higher than 18% of the normal
..:, growthfigure.Theimpressivelateralregeneration
growth rate (102-154 times faster) reported by
&
Lehnert Reitner (1997) corresponds to an
ordinary extension ofthe living tissue produced
to repair a damaged zone ofthe sponge prior to
calcification. However, these authors did not
record growth rates ofthe skeleton itself.
— R Z^H The present study examined populations of
Ceratoporella,whereasmostpreviousreportson
A growth rates were based on measurements of
single specimens. These studies assumed a
constant growth rate during the lifetime of the
FIgGr.owt9.hpCaettreartn.opGorroeulnldasencitcihoonlfsroonmuspReecgiemneenranot.io1n sponge. In this work we found that statistically
sampled from its regeneration area in May 1997. significant variations are consequential, espec-
Fracture, indicatedbyarrowsoccurred in May 1986, ially in researches relating growth rates and
when a fragment was sampled. T6 indicates surface marine paleoenvironmental conditions such as
ofskeleton marked with calcein 363 days later. R= watertemperatureorsalinity,usingskeletalchem-
regeneration growth axis. Epifluorescence istry ofsclerosponges.
microscopy. Captions as in Figure 1. (Scale bar=
500p.n0- ACKNOWLEDGEMENTS
(Bohmetal., 1996)].orondirectmethods(Dustan We are indebted to Roland Baron, Department
& Sacco, 1983; Willenz & Hartman, 1985). No of Internal Medicine and Cell Biology, Yale
details of analytical methods were reported by University School of Medicine, for his cordial
Dustan & Sacco (1983), who used alizarin red reception and forsuggestingtheuse ofcalceinto
staining; their data remain approximate us. During the many years that this work has
((0W.i1l-l0en.z2m&mHyarr't1)m.anI,nit1ia9l85v)awlueerse sulsiignhgtlycallocweeirn wpirtohcetehdeeads,siusntdaenrcewaotfetrhewfoorlklowwaisngmbauddedypodsisviebrlse:
Jeremy Woodley, Peter Gayle, Robin Hall,
(184.2±19.4um yr"1) than in the present study Debbie Santavy and Pierre Van de Steen. Field
becausemeasurementsatthe infillingzoneatthe work assistance by Sylvain and Donat Willenz
base ofpseudocalices were included in the latter has been much appreciated. We are indebted to
study.Suchartifactswereavoidedherebyrejecting MrEric Regoutformakingspecial arrangements
measurements made inthis zone, becausethey are with his company to lend us a Nikon Optiphot-2
too sensitive to minor deviations in the orient- epifluorescent microscope on several occasions,
ation ofskeletal sections. priortoitspurchase. Thisworkwassupportedby
National Science Foundation Grant
This study also provides clear evidence that BSR-8317690 to Yale University, and by the
statistically significant differences occur in the following to one of us (Ph.W): two NATO
growthratefrom oneindividual toanother, within science fellowships, two Fondation Agathon de
the same period of time. Moreover, the annual Potter subventions, a Lerner-Gray Fund for
684 MEMOIRS OF THE QUEENSLAND MUSEUM
400
Averages: YZX T3~4 194-2-42-2 v™
-
350 T4-5 238.4 + 38.8 pm
300 - X T5-6 272.2 + 36.9 pm
I
250
x
*"" 200 -
150 1
%
I +& *t 4$ g 44
-
100
—
50
10 11 16 17
Specimens
FIG. 10. Ceratoporellamcholsoni. Regenerativeannualgrowthrate(umyr1)of6specimensafterinjurycaused
by sampling. Conventions indicated as in Figure 3. A Kruskal-Wallis ANOVA on ranks and all pairwise
multiplecomparisontest(Dunn'smethod)indicateasignificantgrowthrateincrementwithinagivenspecimen,
from one period to the nextone.
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