Distribution Patterns of the Herbivorous Gastropods
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Benthos Research Vol.57,No.1:1-10(2002) BENTHOS RESEARCH The Japanese Association of Benthology Distribution Patterns of the Herbivorous Gastropods Chlorostoma lischkei and Omphalius pfeifferi pfeifferi(Trochidae)in Relation to the Algal Community on a Rocky Shore of the Oshika Peninsula, Northeastern Japan Akiyuki Ishida•õ,Koichi Sasaki,Michio Omori and Kazuya Taniguchi Graduate School of Agricultural Science,Tohoku University, 1-1Tsutsumidori-Amamiyamachi,Aoba,Sendai,Miyagi981-8555,Japan Abstract:The distribution of the herbivorous gastropods Chlorostoma lischkei and Omphalius pfeifferi pfeifferi in relation to the algal community was investigated on a rocky shore of the Oshika Peninsula,north eastern Japan.Chlorostoma lischkei was distributed chiefly in a shallower cobble area all year round while .p.pfeifferi lived always in the Eisenia bicyclis forest. Detailed microhabitat surveys showed that smallerO C.lischkei individuals inhabited the undersides of cobbles and the bottom beneath cobbles ,and that larger ones migrated to more exposed places. Larger O.p.pfeifferi individuals inhabited the laminae of E.bicyclis, medium-sized ones the holdfasts of E.bicyclis,and smaller ones the sea bottom under cobbles .Avoidance of predation is probably why smaller individuals of both species commonly lived beneath cobbles .With growth,O.p. pfeifferi utilized E.bicyclis not only as food but also as the main living space,but C.lischkei did not form such a relationship with algae.This indicates a fundamental difference between the two species in their relationships with algae and algal vegetation. Key words:algal zonation,Chlorostoma lischkei,distribution,habitat,Omphalius pfeifferi pfeifferi They are arranged in order from inshore to offshore:a INTRODUCTION kelp forest of larger perennial brown algae such as laminarians and fucoids,a zone dominated by the The trochid gastropods Chlorostoma lischkei and smaller perennial brown alga Dilophus okamurae,and a Omphalius pfeifferi pfeifferi are common throughout the flat of crustose coralline red algae.The rocky shore off waters around Japan from the southern part of Hokkaido Tomari-hama on the Oshika Peninsula in Miyagi Prefec to Kyushu,inhabiting rocky intertidal to subtidal areas. ture has an arrangement of benthos communities corre Both are herbivorous and similar in adult size and exter sponding to the algal zonation.There,O.p.pfeifferi in nal features. habits a kelp forest of Eisenia bicyclis,and C.lischkei a Subtidal seabeds on rocky shores along the Pacific cobble area shallower than the kelp forest.Both gastro coast from the Sanriku to Joban districts of northeastern pods are dominant species in their respective zones,each Japan commonly show a series of distinct algal zones, having a larger biomass than other gastropod species each dominated by a different alga(Taniguchi1991). (Omori et al.2000). Kelp forests of E.bicyclis on rocky shores have a Received July27,2001:Accepted January 28,2002 high annual productivity of20kg/m2wet weight and pro vide important habitats and food for herbivorous Present address:Institute of Environmental•õ Ecology, benthos,including commercially valuable animals such METOCEAN Co.Ltd.,1334-5 Riemon,Ooigawa, Shizuoka as sea urchins and abalone(Taniguchi&Kito1988). 421-0212,Japan Ishida et al. Seki(1997)showed that biological processes in rocky shore ecosystems include interactions among various or MATERIALS AND METHODS ganisms,including algae,and that animal populations Study area there might depend on such interactions as the pass through successive developmental stages.These findings The study was conducted in a small bay off Tomari suggest that the mode of life of animals in rocky shore hama on the northern coast of the Oshika Peninsula, ecosystems can only be understood in relation to algal Miyagi Prefecture(Fig.1).The U-shaped small bay vegetation.Although C.lischkei,which dominates the opens northeastwards to the open sea. It is exposed di shallow cobble area,and O.p.pfeifferi,which dominates rectly to wave action from offshore because of the ab in the kelp forest,are both inferred to intensely affect the sence of any islands or rocky reefs in front.The seabed dynamics of the rocky subtidal ecosystem,little is known slopes gently,reaching a depth of about8m at a distance about their interrelationships with the algal community of about200m from the coastline.The seabed down to from the juvenile to adult stages. 2m deep accumulates cobbles up to15-20cm in inter This study examines changes in the gastropods'dis mediate diameter,with some patches of Sargassum tribution patterns as they grow from settled juveniles to yezoense growing on the surface of the underlying rock adults,in relation to the algal community and the various exposed in places.The seabed2to7.5m deep consists microhabitats of each algal zone.The selection of habitat of black muddy rocks with scattered cobbles.The brown is discussed in relation to the change of feeding habits alga Eisenia bicyclis dominates to a depth of5m,and with growth.The species names of the gastropods used crustose coralline red algae in the range from5to7.5m. in this paper follow the taxonomic nomenclature of Dilophus okamurae forms a narrow band about1m wide Okutani(2000). more-or-less spanning the boundary between them.A sandy bottom with scattered cobbles occupies the area deeper than7.5m,where no dominant algae are found. Depth-related sampling The gastropods C.lischkei and O.p.pfeifferi were taken at intervals of about one to two months from August, 1997,through June,1998,in order to investigate their vertical distribution patterns and changes in size compo sition with depth.A transect150m long was set north eastwards from a starting point(St.0)at a depth of about1.5m.Sixteen stations were set along the transect at intervals of10m.The stations were designated St.0 to St.150in accordance with their increasing distance from the starting point.Apart from these stations,St.A was set in the cobble area1to2m deep,and St.B in a patch of S.yezoense,in order to collect C.lischkei from the shallower zone. At each station,a1m•~1m quadrat was placed and the snails within it were collected by hand from algal thallis and the sea bottom,including surfaces of cobbles as well as the sea bottom beneath cobbles.Then,another Sargassum zone Eisenia zone Dilophus zone smaller quadrat of0.5m•~0.5m was randomly placed crustose coralline algal zone inside the larger one.Organisms inside the smaller quadrat were scraped off the sea bottom and cobbles Fig.1.Bottom topography and profile of algal zonation in the with a metal scraper and collected by means of an air-lift study area,Tomari-hama,Oshika Peninsula,Miyagi Prefecture. sampler,which drew them upward through a suction Italic numerals show depths in meters.Solid circles denote sam pipe into a collecting bag of 1-mm mesh.Dominant pling stations.Stations0to150are set along a transect,16sta algae and bottom features within a5-m radius at each tions at10m intervals. station were also recorded.Organisms collected were 2 140145E•_•_”n•E•A•A•T‚î•A1•_•E•A•A•A‹T•A•A”••A‹T‚ç•œ•A89•A•A•A”n7•A‹T_34•B•_“¯'•Aƒm•_•A;bl•_"•A6‚Öil‚±,,,./ƒ€/1•c•œ'ƒ^'•_•A36‚¢8'‚ăt‚®ƒm•A•A•_•_•_•_•E•E‡`•A•A•E•A•æ•A'•A1•A•E‚µ•E•A•_•_•AŠd‡`"48"‚®0‚¢•A——•E•E•A•_•A•_•_li’È:˜K‹¿•ø“«•øli‘’i‘áƒjƒ“‚¬)1•Yˆê‹\•A“åƒm/œW Distribution of trochid gastropods immediately fixed in10%seawater formalin.Samples formalin solution in seawater.The length,width,and were not obtained at Sts.130and150in March of1998 height of each cobble were measured after the animals because of rough weather. had been removed from them,and the small algae grow Chlorostoma lischkei and O.p.pfeifferi were sorted ing on the cobble's surface were identified.The wet out and counted by species.The shell diameter(ShD)of weights of S.yezoense and E.bicyclis were recorded,and each individual was measured to the nearest0.01mm the number and arm length of lateral blades of E.bicyclis and the body weight was weighed to the nearest0.01g. were counted and measured.Both species of gastropod The smaller snails collected by air-lifting were sorted out were sorted out and measured in the same manner as for and their ShD was measured to the nearest0.01mm by the depth-related samples. using a stereo dissecting microscope equipped with an ocular micrometer. Other investigations Two supplementary investigations were conducted in Microhabitat-related sampling June and September,1999.The investigation in June fo To clarify and compare the micro-distribution of the gas cused on microhabitats in the Eisenia zone at Sts.3and tropods among zones exhibiting different conditions,six 4.At St.4,where E.bicyclis grew densely,a detailed stations(Sts.1to6)were established(Fig.1).Station distinction was made among three microhabitats:on 1was set in the cobble area1to2m deep,and St.2in algal laminae,around algal holdfasts,and on and beneath a patch of S.yezoense at a depth of about1m.Three sta cobbles.Collections of individuals of E.bicyclis and tions were set in the E.bicyclis zone:St.3in the cobbles,and drawing up of scraped-off organisms by the shoreward fringe about2m deep,St.4in the central part air-lift sampler were done in the same manner as for the 3-4m deep,and St.5 in the offshore fringe5-6m microhabitat-related samples.After the algae had been deep.Station6was in the crustose coralline red algal collected,a0.5m•~0.5m quadrat was placed on the zone at a depth of6-7m.Investigations were carried spot from which the algae had been cropped and organ out in September,November,and December of1998, isms were collected by the air-lift method.Then,three or when newly settled juveniles were expected to be ob four cobbles outside the quadrat were removed and the tained in the field.