Particle-size utilization in the deposit-feeding polychaete Neanthes
succinea was examined. Gut analysis revealed that worms in San Francisco Bay
consumed a broad range of particle sizes (20-300mm diameter). Gut sediment
closely resembled size of surface sediment at the mouths of worm burrows
. indicating non-selective feeding. All size classes of worms consumed similar sized
particles. The flexibility in feeding modes and diet of nereids is discussed

Fong, Peter P.

Pacific Science

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Pacific Science (1987), vol. 41, nos. 1-4© 1988 by the University of Hawaii Press. All rights reservedParticle-Size Utilization in the Introduced Polychaete Neanthes succineain San Francisco Bay!P ETER P. FONG2ABSTRACT: Particle-size utilization in the deposit-feeding polychaete Neanth essuccinea was examined. Gut analysis revealed that worms in San Francisco Bayconsumed a broad range of particle sizes (20-300jlm diameter) . Gut sedimentclosely resembled size of surface sediment at the mouths of worm burrows. indicating non-selective feeding. All size classes ofworms consumed similar sizedparticles. The flexibility in feeding modes and diet of nereids is discussed.Neanthes succinea (FREY AND LEUCKART) is acommon, infaunal polychaete introduced toSan Francisco Bay with the Virginia oysterfrom 1860-1910 (Carlton 1975). Broadly dis-tributed in estuarine areas of both Americancoasts, Europe, and the Hawaiian Islands, itsabundance and large size make it a usefulorganism for the study of polychaete feedingbiology.A number of studies on N. succinea fromthe eastern United States and Europe havefocused on life history (Wilson 1892), repro-duction (Lillie and Just 1913; Kinne 1954),larval morphology (Banse 1954), occurrence(Smith 1963), ingestion rate (Cammen 1980),and growth (Neuhoff 1979). Since its intro-duction, very little research has been done onCalifornia populations, but some studies onsalinity tolerance (Hanson 1972; Kuhl andOglesby 1979) and spawning behavior (Car-pelan and Linsley 1961)have been done in theSalton Sea.The information on the diet of N . succineafrom the California coast is lacking, and fromthe eastern United States it is somewhat con-fused. Teal (1962) and Cammen (1980) callN. succinea a non-selective surface depositfeeder, but give no supporting data on particlesizes consumed. In contrast, Holland (per-sonal communication) has found juvenileamphipods, Corophium lacustre and spionid1 Manuscript accepted November 1985.2 Department of Biology, San Francisco State Univer -sity.37polychaetes in the guts of worms from Chesa-peake Bay.The present study focuses on the diet of N.succinea in San Francisco Bay. Preliminaryinvestigations on worms collected from thebay showed that N . succinea consumes mainlysediment. Therefore it was the aim of thisstud y to (1) determine the particle sizes ofsediment ingested by N. succinea, (2) examinepossible particle size selection by comparingingested particles with particles available foringestion at the sediment surface , and (3) mea-sure particle size ingested relative to wormlength.MATERIALS AND METHODSThe study was carried out from March 1983to February 1984 in an intertidal mudflat inSan Leandro Creek, one of many tidal creeksflowing into San Francisco Bay (Figure 1).Salinities are variable depending on seasonand tidal cycle, but can range from 18%0 in thesummer to freshwater in the winter. Watertemperature fluctuates from 13-20°C duringthe year. The mudflat proper is composedof very poorly sorted sediments (- 2.25<1».The flora is composed of a broad expanseof diatoms on the sediment surface over theentire mudflat. Macro-algae include the cos-mopolitan green algae Ulva and Enteromor-pha.The epifauna consists mainly of the mussels,Ischadium demissum and M ytilus edulis, andbarnacles. The under-boulder fauna includes38 PACIFIC SCIENCE, Volume 41, 1987were counted and measured at 100X. Possibleselective ingestion of different particle sizeswas examined by comparing size-frequencydistributions, and 95% confidence limits of380mean particle size of the sediment at themouth of worm burrows with that in wormguts . Electivity values were calculated usingthe formula ofIvlev (1961):E = (rj - pJ.(ri + pJ'where E is the electivity value , ri , the percent-age of the ith particle size class in worm guts,and Pi' the percentage of the ith particle sizeclass of the surface sediment. E varies from- I to + 1, where negative values indicateselection against a certain size class, positivevalues indicate selection for a certain sizeclass , and zero indicates no selection.10IHILESFIGURE 1. Map of San Francisco Bay showing studysite near San Leandro Creek.12 30the crustaceans Hemigrapsus oregonensis (De-capoda), Anisogammarus confervicolus (Am-phipoda), and Tanais sp . (Tanaidacea).Infaunally, Neanthes succinea and the bivalveMya arenaria are the most common.Although no field observations of feedingbehavior were made, N. succinea in the labo-ratory constructed U-shaped burrows, andactive ly probed around burrow openings. Inthe field, individuals of N. succinea were col-lected by hand, immediately fixed in 10% for-malin and preserved in 70% ethanol. Samplesof the sediment surface were collected bygently scraping the uppermost I em of sedi-ment around the burrow openings with a plas-tic vial. All animals and sediment sampleswere transported to the laboratory for analysis.Individual worms were separated into threesize classes , 2-5, 5-8, and 8-12cm length,and defined as small , medium, and large sizerespectively. Prior to gut analysis, all wormswere rinsed of adhering particles in distilledwater. Guts were removed carefully, and con -tents placed in vials containing 15% HzOzfor 24 h to remove organic material. Surfacesediment was rinsed and treated similarly. Allsamples ofboth gut and surface sediment werethen rinsed again, and spread onto micro-scope slides. Using a phase contrast micro-scope with an ocular micrometer, the first200 particles encountered on each slideRESULTSParticles from the sediment surface in SanLeandro Creek are moderately sorted, andrange from 20-300/lm in width. Approxi-mately 82% of these particles were less than72 /lm, and 51% ranged from 20-50/lm. N.succinea inhabiting San Leandro Creek ingestthe identical particle size range of20-300 /lm,available to them at the surface, and 88% ofthe particles in the guts of these worms wereless than 72 /lm. Size-frequency distributionsof surface and gut sediment are given inFigure 2. For particle sizes combined overa one-year period, no significant differenceexists between surface and gut size-frequencydistributions (Kolmogorov-Smirnov test ; p >0.5). The 95% confidence limits and rangesfor mean particle size of surface and gut sedi-ment are presented in Figure 3. Confidencelimits overlap strongly in the months of June,August, October, and December, and areextremely close to overlapping in November,January, and February. Confidence limitscalculated over the entire year (annual mean)overlap strongly, and these data suggest thatthere is no difference between particle size ofsurface sediment and sediment ingested by N.succinea.Particle-Size Ut ilizat ion in Neanthes succinea- FoNG' 0039JAN5(,·!FEBMAR10 0APR MAY JUNE"~W I ~uz~ClZ:J 100lD~ AUG OCTJU LY~5010050NOV DEC ¥fAA....•..:'.PARTICLE WIDTH (",m)..:'.F IGURE 2 . Size-frequency distribut ions for particle sizes of surface sediment (open bars) and worm gut sediment(striped bars).Ivlev values for all worms examined aregiven in Table 1. Electivity values are negativefor large particle size classes, indicating avoi-dance of these particles. However, only the> 122Jlm size class has a sufficient negativevalue approaching significance. Similarly ,smaller particles (< 72 Jlm) have positive elec-tivity values, but these are no t large enough tosuggest significant selection for these parti-cles. Overall, in most particle size classes, elec-40 PACIFIC SCIENCE, Volume 41, 1987- - -JAN MAR1 FEB~1Qi 1.-- ~... 1 1 1-'~... [II::.- -E APR MAY JUNE.::t j...Qi I. - f ~ - ~ ~~!:.'...a: 0:E JULY AUG OCT.::z:...tQ ~i l• - - 1 ~~ t!:.'... ~a::- NOV I.IYEA"! DEC:z: ~...Qil.· ~ - 51-...-' ~ 1~... ~II:fI ISURFACE GUT SURFACE GUT SURFACE GUTF IGURE 3. Range and 95% confidence limits for means of surface sediment and worm gut sediment .Particle-Size Utili zation in Neanthes succinea-FoNG 41TABLE 1ELECTIVITYVALUES CALCULATED FROMPARTICLE-SIZE DISTRIBUTIONS OFSURFACE ANDGUTSEDIMENT SAMPLESFROM Neanthes succinea (n = 30) FEEDING ON SEDIMENTS IN SAN LEANDRO CREEKSize Clas s (Jtm) 20-50 51-72 73-102 103-122 > 122Eleetivity 0.017 0.009 - 0.066 - 0.020 - 0.255TABLE 2MONTHLY SAMPLING OF G UTPARTICLE SIZE OF Neanthes succinea FROMSAN LEANDRO CREEK.Partiele size valu es are in JtmWORM LENGTH x:MONTH n (WORMS) n (PARTICLES) (em) (PARTICLESIZE) S2 HIGH LOWMarch 2 200 4.7 63.55 1543.2 39.28 326.4 20.41983 5.0Ap ril 5 200 6.0 58.96 794.8 28.19 265.2 20.4200 6.0 54.26 684.6 26.16 255.0 20.4200 7.0 50.49 613.5 24.77 153.0 20.4200 7.5 60.99 1120.8 33.48 214.2 20.4200 9.5 50.23 571.9 23.91 132.6 20.4May 2 200 7.0 49.15 598.2 24.45 153.0 20.4200 8.5 46.46 516.2 22.72 132.6 20.4June 2 200 3.0 49.77 456.4 21.37 132.6 20.4200 7.0 51.10 397.3 19.13 153.0 20.4July 3 200 5.0 67.32 819.7 28.63 173.4 20.4200 7.5 44.06 471.3 21.71 122.4 20.4200 13.0 57.88 920.0 30.33 204.0 20.4August 2 200 7.0 63.34 506.7 22.51 142.8 20.4200 9.0 61.30 551.0 23.47 153.0 20.4October 3 200 8.0 47.84 376.8 19.41 153.0 20.450 10.5 52.84 949.9 30.82 204.0 20.4200 10.5 53.86 529.2 23.00 153.0 20.4November 2 200 9.0 46.46 470.2 21.46 142.8 20.4200 9.0 37.79 183.7 13.55 91.8 20.4December 3 207 2.5 28.67 89.7 9.42 71.4 20.4100 5.0 30.29 112.3 10.60 71.4 20.4101 10.0 33.12 252.6 15.80 102.0 20.4January 3 180 6.0 35.93 264.4 16.26 112.2 20.41984 200 8.0 29.78 118.5 10.88 71.4 20.4200 13.5 34.98 275.5 16.60 132.6 20.4February 3 200 3.5 34.07 256.6 16.02 122.4 20.4200 6.0 31.52 204.0 14.28 122.4 20.4200 18.5 39.42 239.1 15.40 102.0 20.4tivity values are close to zero, indicating thatN. succinea is not selecting specific particlesizes.A tabulation of the one-year gut particle-size survey of N. succinea is given in Table 2.The three worm size classes have similar meanparticle sizes (45.5-50.0 Jim) and all three95% confidence limits overlap (Figure 4) sug-gesting that particle size ingested is indepen-dent of worm length.DISCUSSIONThough nereids are generally omnivorous,only a few species are known to be depositfeeders (Fauchald and Jumars 1979). Hence ,very few papers have focused directly on nereidparticle feeding or their effects on the benthos.In San Francisco Bay, Neanth es succinea isa strict surface deposit feeder (Fong 1985).The results of the present study indicate that42E-=-s:;:;~ 50'"~:-Q.2-5 5- 8 8-12PACIFIC SCIENCE, Volume 41, 1987cinea precludes any comparison with the pre-sent study. Furthermore, nereids in generalshow extreme flexibility in feeding modes.For example, Nereis virens is herbivorous atWoods Hole, but omnivorous at other loca-tions (Theede et al. 1973). Nereis diversicoloris omnivorous, but also capable of filterfeeding using a mucous cone (Fauchald andJumars 1979). Therefore, generalized feedingmodes for certain species may not be veryvaluable. Additional studies from differentlocations should be attempted to consolidatewhat litt le information exists for N. succineaand other nereids .W orm Size Class (em)FIGURE 4. Range and 95% confidence limits for meanparticle width of gut sediment relative to worm length .this po lychaete consumes a wide range of par-ticle sizes. Moreover, the sediment sizes inworm guts closely resemb le sizes of surfacesediment available for ingestion (Figure 2).Ivlev values indicate no selection for any sizeclass below 122Jlm . These data alone are no tsufficient enough to conclude non-selectivefeeding since worms may be living under opti-mal sediment conditions (Hammond, 1982).Worms did not feed readi ly in the labora tory,therefore experiments to test particle sizepreference were impossible. However, wormsin the field consume not only sediment ofextremely broad range, but also a consider-able amount of alloch thanous material. Ifworms were feeding selectively, these materials(main ly small bits of terrestrial plants) wou ldnot be present in worm guts . Moreover, thereappears to be no relationship between wormbody length and sizes of particles ingested .These results and observations lead to theconclusion that N. succinea is a non-selectivesurface deposit feeder in agreement with theearlier reports of Teal (1962) and Cammen(1980).Many cases for non-selective deposit feed-ing have been reported in capitellids, ophe-liids, orbiniids, and cirratulids (Fauchald andJumars 1979). But, the small amount of re-search on particle-size utilization by N. sue-R&LITERATURE CITEDBANSE, K . 1954. Uber morphologie und lar-valentwicklung von Nereis (Neanthes) sue-cinea (Leuckart) 1847 (Polychaeta errantia).Zool. Jahrb. Anato. Ontog. Tiere. 74 : 160-171.CAMMEN, L. M . 1980. A method for measur-ing ingestion rate of deposit-feeders and itsuse with the polychaete Nereis succinea .Estuaries 3(1) :55-60.CARLTON, J . T . 1975. In troduced invertebra-tes. In: R. 1. Smith and J. T. Carlton (eds.)Light' s Manual: Intertidal invertebrates ofthe central California coast. University ofCalifornia Press , 716p.CARPELAN, L. H ., and R. H . LINSLEY. 1961.The spawning of Neanthes succinea in theSalton Sea. Ecology 42(1): 189- 190.FAUCHALD, K ., and P. A. JUMARS. 1979. Thediet of worms: A study of polychaete feed-ing guilds. Oceanography Marine BiologyAnnual Review 17: 193-284.FONG, P. P. 1985. Particle-size analysis of thedeposit-feeding polychaete Neanthes sue-cinea (Frey and Leuckart) with observa-tions on its reproduction. Master's thesis ,San Francisco State University, 78 p.HAMMOND, L. S. 1982. Analysis of grain -size selection by deposit-feeding holo -thurians and echinoids (Echinodermata)from a sha llow reeflagoon, Discovery Bay,Jamaica. Marine Ecology Progress Series8 :25-36.Ii;;;'"Particle-Size Utilization in Neanthes succinea-FoNG 43HANSON, J. A. 1972. Tolerance ofhigh salinityby the pileworm Neanthes succinea, fromthe Salton Sea, California. California Fishand Game 58: 152-154.IVLEV, V. 1961. Experimental ecology of thefeeding of fishes. Yale University Press,New Haven, 302 p.KINNE, O. 1954. Uber das schwarmen unddie 1arvalentwicklung von Nereis succineaLeuckart. Zoo\. Anz. 153: 114-126.KUHL, D ., and L. C. OGLESBY. 1979. Repro-duction and survival of the pileworm Nereissuccinea in higher Salton Sea salinities. Bio-logical Bulletin 157: 153-165.LILLIE, F. R., and E. E. JUST. 1913. Breedinghabits of the heteronereis form of Nereislimbata at Woods Hole, Massachusetts.Biological Bulletin 24: 147-168.NEUHOFF, H. G . 1979. Influence of tempera-ture and salinity on food conversion andgrowth of different Nereis species (Poly-chaeta, Annelida). Marine Ecology Pro-gress Series I: 255-262.TEAL, J . M. 1962. Energy flow in the saltmarsh ecosystem in Georgia. Ecology 43:614-624.THEEDE, H., J. SCHAUDINN, and F. SAFFE.1973. Ecophysiological studies on fourNereis species of the Kiel Bay. Oikos(suppl) 15:246-252.WILSON, E. B. 1892. The cell-lineage ofNereis.J. Morphology 6: 361-466.

Particle-Size Utilization in the Introduced Polychaete Neanthes succinea in San Francisco Bay

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Particle-Size Utilization in the Introduced Polychaete Neanthes succinea in San Francisco Bay

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