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BIOTROPICA30(4): 604-608 1998
Andrew L. Mack2
Suite 200, 2501 M St., N. W., Washington,DC 20037, U.S.A.
ConservationInternational,
ABSTRACT
Althoughlargeseeds mightbe moreattractive and apparentto seed predators,
largeseed size could enable tolerance
of seed predators.If seeds are largeenough to sustaindamage thatwould kill smallerseeds yetstillproduceviable
seedlings,investment above the minimumby the maternalplant could be advantageous.I testedthishypothesis by
removing0-80 percentof the cotyledonsof fourlarge-seeded(4-180 g) treespeciesfromPapua New Guinea and
monitoringgermination and seedlinggrowthforeightmonths.All speciesshowedlittlenegativeeffecton seedling
size with up to 50 percentremovalof cotyledonsand the largerspeciesshoweda less seriouseffecton growththan
smaller-seeded speciesabove 50 percentremoval.Large-seededspeciesclearlyhave morethanthe minimum-required
cotyledonary reserves.Observationsof viable seedlingswith heavilydamaged cotyledonssuggestthat thesespecies
withstandattackby rodentsand beetlesby virtueof theirlargesize.
Keywords: cotyledons;
germination;
herbivory;
maternalinvestment; seedpredation;
Papua New Guinea;rodents; seedsize;
tolerance.
604
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Advantages of Large Seed Size 605
sp 1
Cryptocarya sp2
Cryptocarya
,= 100' . 100
,= 80 , * 80
60 *60
4O 40
20 20
0.0 0.2 0.4 0.6 0.8 1.0 0.0 0.2 0.4 0.6 0.8 1.0
0.0 0.2 0.4 0.6 0.8 1.0 0.0 0.2 0.4 0.6 0.8 1.0
tissueremoved
proportion tissueremoved
proportion
FIGURE 1. Plotsof percent of maximum heightof seedlings of cotyledon
againstproportion removed priorto
Alllinearregressions
germination. shownweresignificant(P < 0.001).The regression foreachspeciesare:
equations
sp. I y = -72x + 67, R2 = 0.33; Cryptocarya
Cryptocarya sp. 2 y = -48x + 52, R2 = 0.15;Aglaia y = -50x +
76, R2 = 0.40; Endiandrasp. y = -28x + 66, R2= 0. 11.
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606 Mack
150
evidentin the lowerrightportionof the contour
*
125 - surface(Fig. 3).
75- DISCUSSION
50
Studies of seed predationhave sometimesfound
25-
-s
ioo that rodentsprefer"large" seeds (Mittelbach&
Gross 1984, Westobyet a!. 1992). Alternatively,
-25
large seeds could preventpredationby smallseed
predatorssuch as ants (Thompson 1987, Wes-
0 - 25 -50 -75 > 75 tobyeta!. 1992). Studies of the benefitsof large
oftissueremoved
Percentage seeds have focused on the size and vigor of re-
sulting seedlings (e.g., Zhang & Maun 1993).
FIGURE 2. Hi-lo plots of percentof maximumseed- However, the literatureon this topic (reviewed
ling heightamong seedlingswith different amountsof
theircotyledonsablated.The centralhorizontalline rep-
by Westobyet al. 1992) mostlydeals with tem-
resentsthe mean, the box represents? 2 SE and the peratefloraswherevirtuallyall specieshave seeds
whiskerrepresents + 2 SD. Those categorieswithiden- weighing less than 5 grams (e.g., Baker 1972,
tical symbols(* or #) did not differsignificantly
(SNK Mazer 1989, Westobyet a!. 1992, Juradoet a!.
test;see Results). 1991). Little attentionhas been devoted to the
possibilitythat large seeds enable plants to tol-
erate damage (Rosenthal & Kotanen 1994),
effectdue to cotyledonablation.The effectofpro- probablybecause fewstudieshave examinedflo-
portionremovedon seedlingheightwas stronger ras wheretrulylarge seeds occur.
in the smallest-seeded species(the largestnegative Largecotyledonsor endospermcan protectthe
regressioncoefficient) but the othersmall Crypto- embryobysatiatingpotentialseed predatorsbefore
caryaspecieshad a regression coefficient similarto theseagentsdamage the embryo."Satiation"here
thatof themuchlargerAglaiaseeds(Fig. 1). How- includesboth physically fillingthe rodent'sgut or
ever, removal of large amounts of cotyledon reachingsome maximumtolerableamountoftoxic
(>50%) caused greatermortality in thetwo small- secondarycompoundscontainedin the consumed
er-seededCryptocarya speciesthanin thetwolarge- tissues.In the largestspeciesstudied,up to 141 g
seededspecies(x2 = 11.25, df = 1, P < 0.00 1; Fig. of cotyledonscould be removedwithoutkillingthe
1). seed. This amount could certainlysatiate most
I pooled thedata forall speciesand dividedthe seed-gnawing rodentsat thestudysite.Elevenspe-
seedsintofiveclasses:thosewith0, > 0 and ? 25, cies of rodenthave been recordedat the site (D.
> 25 and ? 50, > 50 and ? 75, and > 75 percent D. Wright,pers. comm.). Nine of thesetypically
removal.A one-wayANOVA revealedthat there weighless than 120 g, one speciesaverages212 g,
was a significant effecton the dependentvariable, and one species weighs over 500 g on average
relativegrowth,due to amount of removal(F = (Flannery1995). Viable,partially-gnawed seedsare
6.48, SS = 1.97, df = 4, P < 0.0001). Student- commonlyfound (Mack, pers. obs.). Likewise,
Newman-Keuls tests showed that the relative largeseed size makesit unlikelythatrodentswill
growthamong all seeds with0-50 percentof the carryseeds to a cache site. In an Australianrain
cotyledonremoveddid not differ but forest,verylargeseeds are less likelyto be moved
significantly,
thatthosewith > 50 percentremovalhad signifi- by rodentsthan smallerseeds (Osunkoya et a!.
cantlysmallerseedlingsthanthosewith< 50 per- 1994). Aftergrazingon a seed,it is likelya rodent
cent (P < 0.01; Fig. 2). will abandon it. Establishedseedlingsof the tWo
The overallpatternis evidentin a three-dimen- largestspeciesin this studyare commonlyfound
sionalplot of thedata (Fig. 3). A surfacewas fitted growingfrompartially-gnawed seeds (Mack, pers.
to the plot using the distance-weightedleast obs.).
squaresmethodwithtensionrelaxedto 0.15 (SYS- The situationis similarwherebeetlesminethe
TAT, Wilkinson1990). The upturnedsurfacefor storagetissue.If the storagetissueis largein vol-
large seeds indicatesthat the effectof cotyledon ume, tunnellingscarabbeetlescan extensively feed
ablationon largeseedsis less pronouncedthanfor and tunnelthroughthistissuewithoutlethallyen-
small seeds and the drop in growthand survivor- counteringthe seed embryo.Heavily-mined coty-
ship occurringaround 50 percentremovalis also ledons were commonlyobservedon young seed-
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Advantages of Large Seed Size 607
0' N
IV 0'3
-o
FIGURE 3. Three-dimensional plot of initialseed size (z axis), proportionof seed ablated (x axis) and percentof
maximumheightof seedlings(y axis). Data forall fourspeciesare pooled (N = 117) and a contoursurfacefitted
leastsquaresprocedure.
witha distance-weighted
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608 Mack
LITERATURECITED
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