The Anatomy of the Pelecypod Family Arcidae Author(s): Harold Heath Reviewed work(s): Source: Transactions of the American

Philosophical Society, New Series, Vol. 31, No. 5 (Aug., 1941), pp. 287-319 Published by: American Philosophical Society Stable URL: http://www.jstor.org/stable/1005609 . Accessed: 27/09/2012 02:43
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TRAII SACTIONS
OF THE

AMERICAN PHILOSOPHICAL SOCIETY

HELD FOR AT PHILADELPHIA USEFUL KNOWLEDGE PROMOTING

XXXI, PART V NEW SERIES-VOLUME AUGUST, 1941

THE ANATOMY OF THE PELECYPOD FAMILY ARCIDAE

HAROLD HEATH Hopkins Marine Station, PacificGrove, California

PHILADELPHIA:
THE AMERICAN PHILOSOPHICAL
FIFTH STREET

SOCIETY

104 SOUTH

1941

THE ANATOMY OF THE PELECYPOD

HAROLD HEATH

FAMILY ARCIDAE1

Hopkins Marine Station, Pacific Grove, California

ABSTRACT

The ancient and geologically importantfamilyArcidae is estimated to comprise upwards of eleven hundredspecies and subspecies. The greaternumberof these are in a fossilstate, and obviously have been classifiedon the basis of shell characters. The same is true of many living species, and where anatomical studies have been made they usually were limitedto some definite organ or system. While such a procedure resultsin supplyingthe morphologist with importantdata, it may lead to faultyconclusionson the part of the systematist. In the presentwork,forexample, there are two types of individuals with identical shells and declared to be of the same species, whereas their anatomy proves them to be distinct. Furthermore, there are subspecies where the shells are unlike, while the internal organization would scarcely warrant subspecific rank. And again thereare genera wherea definite organ or systemis ofdistinctdiagnosticvalue; in other genera it is of doubtful worth. Under such circumstancesit is apparent that comparisonsmust rest upon the entireorganizationof each species ratherthan upon one or a few selected elements. In the presentstudy the anatomy of thirty-two species and subspecies has been describedand figured in more or less detail. No attempt has been made to modifythe existingscheme of classificationbeyond pointingout in the finalparagraphs wherecertainchanges appear to be in order. It may be added that the relativelywide range of differences between species of certain genera suggestfurther revisions,but the importance of these variations can be determinedonly after a study has been made of a greaternumber of additional species.

Accordingto a recentreviewof the familyArcidae (Reinhart'35), it is conservatively estimatedthat not less than twelve hundredspecies are includedin the group. The majority of these are fossilforms,and obviously have been classifiedon the basis of shell characters. The same is true of many recentspecies. On the other hand numerousinvestigatorsare responsiblefor detailed studies of the urogenital,circulatory digestive or or systems ofvariousexternal features. Onlya veryfewspecieshave been studiedfrom the standpointof theirentireorganization. During the course of the presentinvestigation became evident that a scheme of it classification based upon shell charactersalone may lead to erroneousconclusions. And, speaking generally, was equally evident that comparisoRis it based upon a singleorgan or system also may prove to be untrustworthy.In one subgenus or genus fundamental differences may appear; in othercases theymay not. In such a studythe nervoussystemappears to be highly conservative, littlesubject and to change. The same apparentlyis trueofthe urogenital system. The circulatory system as a whole is unreliable,but wherethe pericardialcavityis singleor double or dividedby a dorsal or ventralseptum the situationis important, especiallywhen taken into considera1 Sincere thanks are extended to the followingpersons who have aided materiallyin the preparation of this paper; Dr. A. A. Wetmoreand Dr. Paul Bartsch of the National Museum forthe major portionof the species listed in the table; Dr. H. G. Schenck and Dr. P. W. Reinhartand Miss Myra Keen fornumerous suggestionsand guidance in matters relatingto synonomy; Dr. V. van Straelen forspecimens of Arca imbricata and several other species; Mr. and Mrs. E. C. Chace for carefullyprepared specimens of Anadara (A car) pernoides. 287

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species.

tion along with otherfeatures. The various types of abdominal sense organs frequently of and are of servicein determining relationships, the numberand arrangement the muscles an clue. The digestivesystemis highly the bridging visceralcavitymay afford important plastic, and oftenthereis a markeddegreeof variationeven in the membersof the same of subgenus. Neverthelessthe configuration the stomach,the position of the bile ducts factorsin tracingthe phylogeny the of and the lengthof the intestinemay be important group. In short,the most satisfactory must restupon evidence supplied by classification as the study of the entireorganism various authorshave maintained. is as Finally,it may be added, the presentinvestigation not presented a criticalscheme of classification, ratheras a contribution whichit is hoped may prove to be of service but of to studentswho are confronted withthe problemof tracingthe phylogeny this ancient and geologicallyimportantgroup. To this end the concludingparagraphs are devoted to a listing of the more importantresemblancesand differences between the different
METHODS AND MATERIAL

In the majorityof species a definite technique was employedin the preparationof materialfordissectionas well as forthe dissectionprocess itself. In most instancesthe shape of the shell is such that the removalof the body usually resultsin a seriousamount of damage to the softertissues and organs. Accordingly the shell was removedby dissolving it in dilute nitricacid. Furthermore, the majorityof the species the visceral in cavity, as usual, containsthe gonad, digestivegland and the largerpart of the digestive system,but unlikethe majorityof mollusksthese organsare bound together with connective tissue of extremetenacity. Even in those species where this condition is not so pronouncedthe digestivesystemis frequently such delicacy that no satisfactory of dissectionof the ordinary type is possible withoutthe consumption an inordinateamount of of time. Hence afterrepeated failuresand partial successes each specimenwas divided sagittallyalong the mid-line, and the dissectedstructures each halfwerethen combined of in the finalsketch. In some cases wherethe intestinalloops were of a morethan usually complicatedtype the coils of each half were modelledin plastic clay and the two halves were then combined. in The various species figuring the presentstudy are listed in the appended table. Arca imbricata was identified such by Dr. H. G. Schenck and Miss Myra Keen of the as Departmentof Palaeontology,StanfordUniversity. They also, togetherwith Dr. P. W. of Reinhart,are responsiblefor the identification Barbatia decussata,and for Scapharca sp. collectedoffthe coast of Senegal, Africa. The remaining in species were classified the National Museum.
ARCA

this subgenushas been the subject of investigaOwing to various primitive features, tions on the part of numerousauthorsduringthe past hundredand fifty years. Much of this workhas centeredin the arrangement and structure the circulatory of systemand its possible origin. The structureof the gills likewisehas received considerableattention, and Matthias ('14) has made an exhaustivestudyofthedigestivesystem and the circulation to a less extent.

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In the presentcollectionthis subgenusis represented six typical species, bouvieri, by imbricata, noae, pacifica,umbonata and zebra. From the standpointof externalappearance all of these species exhibita fairlyclose resemblanceto each other (compare the figures of the first fourand last plates). The palps are moreor less verticalin position,the byssus cavity is well developed, and the abdominal sense organs are related to the anal opening in essentiallythe same fashionthroughout. There is considerablevariationwith respect to the pedal retractors correlatedpossiblywiththe natureof the environment. They are relatively small in A. pacifica,of about average size in A. bouvieri, imbricata and umbonata and large in A. zebra. The digestivesystemvaries to a considerable degree,and yet thereare certaingeneral featurescommonto all. The mouth holds a positionadjacent to the lowerborderof the
TABLE Species Arca bouvieriFischer................................... Arca imbricataBruguiere ................................... Arca sp................................... Arca noae Linne ................................... Arca pacifica Sowerby ................................... Arca umbonataLamarek................................... Arca zebra Linne ................................... Barbatia barbataLinne ................................... Barbatia decussataSowerby ................................... Barbatia (Cucullaearca) candida Gmelin ................................... Barbatia (Acar) gradata Broderip & Sowerby ................................. Barbatia (Acar) pernoidesCarpenter ................................... Barbatia (Acar) reticulataGmelin ................................... ................................... Arcopsis adamsi Smith ................................... Arcopsis solida Broderip Scacchi................................... Bathyarcapectunculoides Trisodos tortuosa Linne ................................... Anadara bisenensisSchenck & Reinhart ................................... Anadara granosa Linne ................................... Anadara (Scapharca) concinna Sowerby ................................... Anadara (Scapharca) auriculata Lamarck................................... Anadara (Scapharca) transversa ................................... Say Anadara (Scapharca) sp................................... Anadara (Cunearca) chemnitzi Philippi................................... Anadara (Cunearca) incongruaSay ........................................... Anadara (Cunearca) perlabiataGrant & Gale. Argina pexata Say ................................... Noetia ponderosaSay ................................... Cucullaea granulosa Jones. americana DeFranco ................................... Glycymeris ................................... Glycymeris longiorSowerby Glycymeris migueliana Dall ................................... Linne ................................... Glycymeris pectiniformis subobsoleta ................................... Glycymeris Carpenter I Locality OffSenegal, Africa Fiji Ids. Fiji Ids. So. of Triest, Italy Gulf of California Porto Rico Porto Rico and Florida Florida OffSierra Leone, Africa Key West Gulf of California OffL. California VirginIds. Florida Gulf of L. California So. of Halifax Philippine Ids. Inland Sea, Japan China Gulf of California Florida Long Island Sound OffSenegal, Africa Porto Rico OffSo. Carolina OffPeru Long Island Sound Florida Suruga Gulf, Japan Porto Rico OffArgentine, Am. S. OffSo. California Philippine Ids. OffOregon, U.S.A.

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anterior shelladductor,and its relationto the palps follows usual plan. The esophagus the shortin A. bouvieri and A. zebra,of average lengthin A. umbonataand is comparatively comparatively long in A. noae, judgingfromthe singlespecimenin hand. In every case there is a dorso-ventral compression. Owing to the large size of all of the specimens, were made, and the presenceor absence of longitudinal no sections groovesand ridgesor of glandularelementswas not determined. was paid to any clearlymarkedindicationsof a special pharynConsiderableattention sectionof the digestivetractbetweenthe mouthopening geal cavityor of a clearlydefined and a definite esophagus,but such appears to be absent not only in Arca s.s., but in all of the otherspecies treatedin the presentwork. In several instancestherewere dilationsin this regionin the case of some of the specimens,whereasin otherindividualsof the same species these were absent. In A. zebra (P1. III, Fig. 2) therewas a markedenlargement of the esophagus,adjacent to the stomach,whichwas filledwith crystalline stile secretion or materialsecretedby the esophageal glands. In A. bouvieri (P1. IV, Fig. 2) the entire esophaguswas distendedwitha mixtureof some secretionand detritalmaterial. In other individualsthis sectionof the digestivetractwas of even caliberthroughout. It would be indeed to ally the pelecypodsin this respectwith the gastropods,forexample, interesting but so faras presentevidenceis concernedit is negative. The stomachpresentssuch a wide range of variationthat a detailed description out is on of of the question. A comparison the figures plates II, III, IV and XXII must suffice. thereis a fairly From theseit willbe seen that macroscopically distinctline of demarcation this last named organ comprisesthe betweenthe esophagus and stomach. Furthermore, well knowndivisions the dorsal stomachproper,and a clearlydefinedventralsection or this dorsal regionfindsits most simpleexpresMagendarmof List ('02). In A. umbonata sion; in A. zebrait becomes morecomplex,becomingstill more complicatedin A. bouvieri and A. noae. it In thisconnection shouldbe notedthat withrespectto the stomacha certainamount of variationoftenoccursamong individualsof the same species. In some instancesthis is as due to distension a resultof large amounts of detritalmaterialor crystalline stile secrecase ofthistypeis represented P1. III, Figs. 2, 3. Both specion tion. The mostextreme as the mens wereidentified A. zebra,and in each figure righthalfof the stomachis shown, in a portionof the leftwall being reflected Fig. 3. It will be noted that in Fig. 2 (from of North Carolina) the posteriorextremity the stomach (pars pylorica of Matthias) is slenderdiverticulum, whereasin Fig. 3 (fromPorto Rico) producedinto a comparatively voluminous. Also the course of the intestineis different the in this regionis much more two cases. It is possible that here we are dealing with distinctspecies, and not with a variationmuch widerthan othersdiscoveredin the courseof this work. The ventral section of the stomach resemblesthat of A. platei and A. (Barbatia) as barbata, describedby Matthias,wheretwo prominent ridges, separatedby a deep groove, are developed fromthe gastric epithelium. Accordingto this same author, the groove the while the largerchannel,with serves to transfer digestedmaterialinto the intestine, secretesthe crystalline stile. whichthe smallerone communicates, In a few of the drawingsthe position of some of the ducts fromthe digestivegland a (hepatopancreas)are indicated,but it is veryprobable that theyrepresent fraction only of those present. Matthias, forexample,notes that thereare 10 openingsin A. platei, 11

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in A. noae and 10 in A. angulata. The count is based upon the fact that the basal section of each duct is a non-glandular continuation the gastricepithelium, of whichmoredistally unites withthe follicles the liver. This demands the use of sectionedmaterialwhichin of the present instance was utilized only in the case of various small species describedin succeedingsections. In all of the species the ventralsectionof the stomachis directedin an antero-ventral directionto a point adjacent to the pedal ganglia. From herethe intestine, a sweeping in curve, extends posteriorly, and on the rightside of the stomach makes its way beneath the heartto open on the ventralface of the posterior shell adductor. An exceptionto this rule occurs in A. imbricata wherethe intestine, upon leaving the stomach,extendsposteriorly beyond the forward borderof the pedal retractor beforerisingto the dorsal surface of the body (P1. XXII, Fig. 1). In A. pacifica (P1. III, Fig. 7) and A. bouvieri(P1. III, Fig. 6) the posteriormargin of the anal openinghas developed a small median flap which becomes a more prominent featurein some otherspecies, notablyA. (Barbatia) barbata(P1. V, Fig. 10). Whetherit serves as a sense organ or as a valve to close the anal openingremainsanotherof many unsolvedproblems. Thanks to the investigations the part of several authors,we possess a clear picture on of the circulatory systemin several species of Arca and theirallies. Accordingly the in presentstudy a superficial examinationonly has been carriedout. In everyspecies there is wide degree of separation of the pericardial cavities and the contained auricles and ventricles;and in most instances a single anterioraorta courses along the dorsal surface of the body to the rightof the mid-line(as in P1. II, Fig. 8). In the largernumberof species treated in the presentworkan effort been made has to study and illustratethe various transverse muscles spanningthe visceral cavity. But in the species of Arca we are considering here these structures surprisingly are small, and so delicate that, in the processof removing the gonad and digestivesystem,practicallyall of the muscle bundles were dislodged. Maceration methodswereunsatisfactory, furand therattemptsaccordingly were abandoned. No study was made of the details of the urogenitalsystemand, it may be added, that duringthe courseof dissectionconsiderableattentionwas paid to the nervoussystem, whichcloselyresemblesthat of Arca sp. describedin the next section. ARCASp. The collection kindly suppliedby Doctor van Straelenofthe BrusselsMuseum included a singlespecimenwithan attached label statingthat it was collectedin the Fiji Ids. on a reef. The shell was examined by Doctor Schenck who identified as Arca imbricata. it featureswhichindicatedthat ifindeed Subsequent dissection, however,revealed numerous this individualwas a normalone it certainly should not be includedin that genus. Upon request Doctor van Straelensupplied a second specimenfromthe same locality,whichhe had identified Arca imbricata, verdictwithwhichDoctor Schenckagreed. Its anatomy as a showedit to be a typicalArca, and imbricata doubtlessis the correct name. Until specific a further is investigation made this unique individual will be designatedin the text and explanationof plates as Arca sp.

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The chiefexternalfeaturesof this species are illustrated P1. I. From these figures on it will be seen that the palps of small, probably contracted,size are almost vertical in and the byssus cavity is of large dimensions. This last position,the gills are horizontal, named structure possesses a fairlyflat roof with low ridgesradiatingfroma sub-central position (Fig. 3). Originallyit containeda hornlikehighlycompact secretioncomposed of a seriesof many concentric layers. In keepingwiththe high degreeof development of are this gland, the pedal retractors of greatsize (Figs. 1, 7). The abdominalsense organsare shownon P1. II, Fig. 4. In a verybroad fashionthey resemblethose of membersof the precedingsection (P1. III, Figs. 6, 7, 9), and they very well may have evolved froma commontype. The mouth,holdingthe usual position(P1. I, Figs. 2, 7), is a slitlikeopening,and the its compressed. On P1. II, Fig. 1, esophagus throughout entireextentis dorso-ventrally a portionof the esophageal wall and the anteriorend of the stomach are reflected, giving the appearance of a prominent longitudinal ridge,but in reality,while ridgesare present, they are of small size and extendfromthe mouthto the stomach. is on The stomachitself fairly accuratelyrepresented P1. II, Figs. 1, 2; hencea detailed description appears to be unnecessary. It may be added that the openinginto the lower division of the stomach (Magendarm) is far to the rightside adjacent to the great fold developed in the ventralwall of the upper gastricdivision. and in addition thereare several other very Two bile duct openingsare represented, in additional openings. delicate depressions the same generalregionthat may represent In this particularindividualthe ventralsectionof the stomachwas filledwithcrystalline stile material that had become almost as hard as stone in the alcohol preservative. all Attemptsto remove it were unavailing,and accordingly that can be said is that this portionof the digestivetract possessed the usual form. The intestinealso followedthe same course as in the various species describedabove, coursingventral to the heart and shell adductor. openingbeneath the posterior For an animal ofthissize (84 mmlong) the visceralcavityis small,and it may be added slenderand delicate. As in the case of that the transversemuscles are correspondingly methodsplayed havoc, and all attemptsto determine Arca s. s., dissection theirdistribution wereabandoned. established. Regardingthe circulatory system,certainfeaturesappear to be definitely on P1. I, Fig. 1, the ventriclesare widely separated, and in this respect As may be seen they resemble Arca s. s. The vessels with which they connect, however,are unique. and By injectingair into these theircoursewas followedwithoutgreat difficulty, so faras the presentspecimenis concernedtheirpositionis believed to be accuratelyrepresented are in the figure. As in otherspecies,the two ventricles unitedacross the mid-line, but in theircourse two branchesare developed whichfollowa route toward the anteriorend of the animal. The one to the rightof the mid-line,the largerof the two, probably coraorta of A. bouvieri respondsto the anterior (P1. II, Fig. 8), forexample. Its mate, on the as was otherside ofthemid-line, tracedas farforward thepedal protractor whereit branched and became lost to view. The main aorta, upon reachingthe retractor, bent downward, gave offa small branch to the regionadjacent to the cerebralganglia, and then pursued the directcoursethrough visceralcavity,givingoff branchesby the way, to become a fairly lost betweenthe halves of the pedal retractor. More unusual is the fact that this same

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also gave riseto a medianvessel,the posterior transverse vessel unitingthe ventricles aorta, whichwas followedto a point whereit passed beneath the intestine. In the majorityof the species in thisfamilythe mantlepresentsa moreor less clouded appearance due to a glandulardevelopmentcomparableto that existingin many nuculids on (Heath '37). In A. imbricata, the otherhand, the mantleis crystalclear,and but little dissectionis needed to lay bare the main featuresof the nervoussystem. Upon removingthe anteriorshell adductor the supra-esophagealganglia are seen to be ellipsoidin form, and, whereganglioncells exist in noticeablequantity,are of a slightly yellowish tinge. These ganglia are prolongedinto the usual pedal and visceralconnectives are represented P1. I, Fig. 6. At the union of each cerebralganglionand its commissure in a strongnerve (m) is developed whichsupplies the mantleand the shell adductor. All of these branchessubdivideand form plexus whichprobablyextendsover the entiremantle. a In this particularindividual the strongestnerves fromthe supra-esophagealganglia extend along the mantle border,and become continuouswith the main branchesfromthe visceralganglia. A small nervearises fromthe ventralface of each cerebralganglion,and passes into the dense tissue surrounding mouth. It appears, thoughthisis not certain, the that these two branchesare destinedto enterthe palps. The visceral ganglia, attached to the ventral surfaceof the posteriorshell adductor, are conical in formand originatefivepairs of nerves(P1. II, Fig. 3). The median pair (g) extends into the tissue which supportsthe freetip of the gills, and althoughno sections were made it is probable that they innervatethe osphradia. The next most median pair (s) extends into the region of the rectum and evidentlyinnervatesthe abdominal sense organs shown in P1. II, Fig. 4. The remainingpairs are distributedto the mantle, the exceptionbeing a delicate pair (b) whichhave been traced as farforward the heart,and, as with the exceptionof a small divisionextending into the mantle,are confined the body to wall betweenthe mantleand the mid-lineof the body. In all of the other species describedin this paper the gonad comprisesa systemof relativelystubby follicularoutgrowthsessentiallythe same as those of Arca noae (P1. II, Fig. 5). On the other hand in this species the follicles, such they prove to be, are if extremely elongated,presenting appearance of a tangledmass of nematodeworms(P1. the II, Fig. 6). Unfortunately was not possible to keep this systemintact while carrying it on the customary processof dissection. Hence the connection the "follicles" or threads of withthe main duct was not determined, much less the relationof the gonoductand kidney. Some of the tubes, undisturbedin the process of dissection,were sectioned,and, althoughthe materialwas not especiallyfavorablefora carefulhistological examination, the following facts appear to be well established. Certain it is that these so-called threadsor folliclesare composed of a single layer of polygonal cells with centralnuclei. And it is equally true that these same threads are canals containingsmaller tubes of two sizes. Furthermore, appearances indicatethat thisindividualwas a male, and that spermatocytes are developed in the largerof these enclosed tubules (P1. I, Fig. 8), and are transported throughthe smallercanals. However, thereis no actual proofthat such is the case since no spermswerepresent. Obviouslythereare wide gaps in this description, and it is hoped that these may be filledby otherinvestigators.

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The two typical species of this subgenusare B. barbataand B. decussata. The first of these and last have been the objects of investigationon the part of several authors, notably Pelseneer,Theiler and Matthias. For the sake of comparison,and owing to a somewhatdifferent viewpointtogether withvarious additionalfacts,a moreor less cursory follows. description Externally these species agree fairlyclosely in their appearance Pl. V, Figs. 1, 2). The mantle exhibitsa slight glandular development,its appearance in B. barbatabeing represented P1. XX, Fig. 2. The byssus cavity is well developed,and the abdominal on sense organsare similarin B. barbata(P1. V, Fig. 10) and B. (Cucullaearca)candida. Owing to the state of preservation B. decussatatheirexact formcould not be determined. of The digestivesystemof B. barbatahas been the subject of a comprehensive study on the part of Matthias. His figures correspond fairly closelyto Figs. 6, 12 on P1. XV. One marked difference exists in the present case with respect to the ventral division of the stomach. Evidentlyin a normalstate two prominent ridgesin the liningepithelium form two troughs,the largercontainingthe crystalline stile, and a smallerone permitting the passage of nutritive material. In the two specimensof the presentcollectionthe crystalline stile secretion was extremely hard,and evidentlyabnormally swollento such an extent that it had flattened ridgescompletely. the The internalfeaturesof the stomachof B. barbatashow it to be unusuallysimplewith dilationsand ridgesreducedto a minimum (P1. XV, Figs. 6, 12). In B. decussatathe state ofpreservation farfrom was beingideal, and in additionthe gastricepithelium was dislodged to a large extentwiththe removalof the crystalline stile. So, whilefiner details probably are lacking,it is believedthat the figures the (P1. IV, Fig. 9, P1. V, Fig. 3) represent main characteristics, which,it may be added, are more complexthan in the foregoing species. The intestineis similarto that of typical arcids, A. noae being a good example. As otherinvestigators have observed,the gut passes beneaththe pericardialcavity. The circulatory systemof B. barbatahas been investigated Menegaux and by the by authors noted above. Accordingto Matthias, two separate pericardial cavities exist. of From the figure B. decussataby Pelseneer('11, P1. II, Fig. 10) it is clear that but one is presentin that species. musclesof the visceralcavity are shownon Pls. V, XVII. The chieftransverse Evidently no study has been made of the nervousand urogenitalsystemsby other continues since none of the speciesin the presentcollection authors,and thisstate of affairs was sectioned.
ACAR

This subgenusincludesthreespecies, A. gradata,A. reticulata and A. pernoides. The on chiefexternalfeaturesare represented P1. VI, Fig. 1, and P1. XIV, Fig. 2. On the first plate the figureof A. gradatashows a sharp fold in the externalgill plate, but this was of foundto be lackingin otherspecimens. There is a glandulardevelopment the external epitheliallayerof the mantle,scatteredgland cells'staining darklywith Delafield's haematoxylinbeing a conspicuousfeaturein A. reticulata. The abdominal sense organs of A. are shown on P1. V, Fig. 9. Those of A. gradataare similar. The byssus is a pernoides withits main features shownon P1. XII, Fig. 11. structure prominent

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A reconstruction the digestivesystemwas made by means of sectionsforeach of of the three species, and the results show that all are constructedupon the same general compressed;in the relativelysimple plan. In A. gradatathe esophagus is dorsoventrally othersit is circularin crosssection. In threespecimensof A. pernoides, preservedin Bouin's solutionand stained in Delaof field'shaematoxylin, thereare peculiaroutgrowths the esophagealwall. These comprise one is situated in the regionof the cerebralganglia (P1. XXII, Fig. threepairs. The first 6). The other two are closely associated, and are much closer to the stomach. At first with suggestedthat they were ducts communicating the appearance of these structures however, in some of the liverfollicles the immediateneighborhood. A closerexamination, of clearlyindicatedthat theyend in stubby outgrowths a unique character. Some of the cell elementscontain an abundant supply of a darkly stainingsecretion,and it may be as assumed that they have the same function similarthoughmore slendercells characteristic of the esophagus in general. The unique featureconcernsseveral large cells which formwartlikeprojectionsfromthe externalsurfaceof the duct (P1. XXII, Fig. 8). These whichstains but lightly. At the presenttimeit is futile granularsecretion containa finely to attemptto homologize withthe glands ofothermollusks. Provisionally thesestructures they may be termedesophageal glands since theirpositionscarcelywarrantsthe assumption that theyare of buccal or pharyngealorigin. numbernot less than thirteen. Two open on the right The bile ducts in A. reticulata over the ventral side ofthe stomach,two are on the leftface,whilethe othersare distributed surface(P1. VI, Fig. 9). In A. gradata(P1. XXII, Fig. 7) two bile ducts connectwiththe anteriorface of the stomach,the much largerone on the rightside extendingbeyond the mid-lineand ventrallyas far as the junction of the stomachand intestine. Anotherduct pair, and yet anotherentersthe right on the leftside is situated slightly dorsal to the first thereare threemain ducts, one attached to the ventralgastricwall, side. In A. pernoides anotherdorsal to it, and anotherfarther back on the rightside (P1. XXII, Fig. 6). is The course of the intestine much the same as in Arca s. s., and as in that subgenus the heart is in a dorsal position. A reconstruction the heart was made in the case of A. reticulata(P1. VII, Fig. 8). of Ventralto the heartthe pericardialcavityis continuous;dorsallyit is divided by a delicate valves are present. A very general sketch of the heart septum. Auriculo-ventricular on of A. gradatais shownon P1. XIII, Fig. 9; of A. pernoides P1. VI, Fig. 6. In this first is named species the pericardium continuousboth above and below the heart,whereasin pernoides thereare two distinctcavities. in The urogenitalsystemwas reconstructed the case of A. pernoides(P1. VI, Fig. 8) and of A. reticulata(P1. XV, Fig. 9). In both of these species the relationof the various shortcanal from kidneyuniteswith the elementsis essentially same. A comparatively the funnel entersthe kidney whilethe renopericardial the gonoductclose to theexternal opening, is a shortdistance away. This state of affairs practicallythe same as that describedby Odhner ('12) forArca glacialis. The gonad is typical in its position and its relationto the kidney. In A. reticulata the reproductivefollicleswere empty, and their epithelial lining was undeveloped, indicating that the reproductiveseason was over. The gonad of A. gradata contained a large numberof ova in various stages of development. Two specimensof A. pernoides

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were males; the thirdwas packed with developingspermsand ova which appeared to be herfullymatured (P1. XXII, Figs. 9, 10). Whetheror not this is a case of progressive must await an answer until an examination is made of maphroditismor proterandry seasons and of various sizes. individualscollectedat different fromsectionsin the case of A. reticulata(P1. The nervoussystemwas reconstructed named species the brain VI, Figs. 4, 5) and A. pernoides(P1. XXII, Fig. 5). In the first bilobed. It will be is typicalof othermembersof the family;in the second it is distinctly recalled that Stempell ('93) and Pelseneer ('11) emphasizedthe fact that a bilobed brain cerebraland and the two divisionswere believed to represent exists in the protobranchs, the origin and although pleural ganglia. This indeed may be the true interpretation, distributionof the related nerves casts some doubt upon their exact homologies (see Heath '37). on In A. pernoides, the otherhand, the pictureappears withgreaterdistinctness. The to ventrallobes, corresponding the cerebralganglia in gastropods,forexample,are united and connectiveslead offto the visceralganglia. Distinct conas usual by a commissure, each of whichsupplies a heavy nerve nectivesalso join themwith the dorsal components, to the mantleand a connectiveto the pedal ganglion. In otherwordsthe dorsal division resemblesthe pleural ganglion in many other mollusks. The cerebro-pedalconnective by may indeed be lacking or it may be represented fiberswhich are associated with the pleuro-pedalconnective. of Otocystsin the Arcidae are seemingly rare occurrence. Pelseneer ('11), evidently citingthe workof Carazzi ('02), states that theymay occur in Arca. I can findno other are in references the literature. They certainly presentin the threespecies of Acar in this theyare attachedto the muscles collection. In the onlysectionedspecimenofA. reticulata also they are in contact of the body wall opposite to the pedal ganglia. In A. pernoides or considerably behindthem. withthe body wall, and may be oppositeto the pedal ganglia In the only sectionedspecimenof A. gradataa slendermuscleband spans the body cavity posteriorto the pedal ganglia, and each otocystis imbedded in this band about halfway line of the body. betweenthe outerbody wall and the mid-ventral In a review of the genus or subgenus Acar by Reinhart ('39), several species are in the amongtheii- threerepresented the presentcollection. From thisaccount considered, A. some authorshave considered reticulata it appears that, on the basis of shell characters, and A. gradata to be conspecific. Reinhart,on the other hand, calls attentionto slight or in importance," differences the hingeswhichare "probably ofspecific at least subspecific is whenthe internalanatomyis taken into consideration. and this verdictcertainly correct there both above and below the intestine, In A. gradatathe pericardialcavityis continuous are fourmain bile ducts, and the otocystsare situated at a considerabledistancefromthe a body wall, to mentiononly a few details. In A. reticulata septum separates the peribile ducts, and the otocystsare cardiumdorsal to the gut, thereare not less than thirteen attached to the body wall. and althoughthere of Strong('32) has shownthat A. bailyiis a synonym A. pernoides, thissame authorconsiders to whentheshellsare compared, is a closeresemblance A. gradata, them to be distinct. Reinhart agrees with this conclusion,calling attentionto the fact in that "although gradata and pernoidesappear closely related, pernoidesdiffers being than gradata. For these reasons, distribution smallerand having a different considerably

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pernoidesshould be considereddistinctfromgradata,as Strong concluded." From the standpointof the internalanatomy this is indeed a legitimateconclusion. Acar pernoides is distinctly fromany otherspecies in the presentcollection. different
CUCULLAEARCA

This subgenus is represented a single species, C. candida. Externallyit bears a by fairlyclose resemblanceto B. barbata. The mantle lacks any visible glandular developupon much the same ment in the single available individual. The byssus is constructed plan as in B. decussata(P1. IV, Fig. 12). The regionadjacent to the abdominalsenseorgans conform that these sensorystructures is heavilypigmented, but it is evident,nevertheless, to the patternof those in B. barbata. The digestivesystem(P1. V, Figs. 4, 5) is somewhatmore complicatedin this species betweenthe two is no widerthan is known than in B. barbata, examplebut the difference for to occur betweenspecies of othersubgenera. There is a general resemblancebetween the muscles of the visceral cavity in this species and those of B. barbata;it is more remotewhen comparedto those of B. decussata (Pls. V, VII, XVII). In the figure representing generalappearance of this species fromthe dorsal side, the the shape of the heart and the originof the dorsal aorta are shown (P1. IV, Fig. 7). In this particularspecimenthe ventricles ventricular or pouches were gorgedwithcoagulated failed forthe fact that dye injected on one side of the mid-line, blood, whichmay account to cross over to the other side. Sections are needed to determineif the pericardiumis singleor double. betweenthis subgenus Various authorshave noted that thereis a distinctrelationship at and typical Barbatia species. Nevertheless there evidently are distinct differences, " Cucullaearca for least insofar shell charactersare considered, Reinhart('35) writesthat as noted above may comprisesa clearlyrecognizablegroup." Hence some of the differences be of diagnosticvalue. More extensiveand intensivestudy is needed beforethis point can be settled.
ARCOPSIS

This subgenusis represented two speciesA. adamsi and A. solida. In both ofthese by the entiremantle presentsan opaque appearance due to a glandulardevelopmentof the externalepitheliallayersuch as is knownto occur in the Nuculidae (Heath '37). Sections stained with Delafield's haematoxylin were made, and while the materialis not favorable fora detailed study it is certainthat the componentcells are slenderelementswithnuclei of adjacent to the externalsurface(P1. VIII, Fig. 13). The remainder each cell contains unaffected the dye. by finely granularsecretion of The arrangement the palps and gills in A. solida are shown on P1. VII, Fig. 10. From a reconstruction sectionsof A. adamsi it develops that the anterior of portionof the foot is traversedby a shallow longitudinalfurrow communicates which more posteriorly with a fairlydeep cavity characterizedby a median longitudinalfold borderedby a few othersof smallersize (P1. VIII, Fig. 9). Also, fromthe study of sections,it appears that in A. solida the same generalplan exists,the chiefdifference greater beingin the relatively of the glands. In both species a definitebyssus secretionis present. In development

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three specimensof A. solida the abdominal sense organs were unusually indistinct, the P1. VII, Fig. 4. Owing to the small size of A. adamsi clearest being the one shown on 7 but (approximately mm in length)theseorganswereeven moreindistinct, thereare some evidencesthat theyresemblethose of A. solida. The digestivesystemis essentiallythe same in both species. As may be seen in the section (P1. VII, Fig. 2) the, esophagus is devoid of any notedrawingof the longitudinal worthy features. In two sectionedindividualsof A. adamsi thisorganis almost circularin crosssection. In A. solida it is exceptionally thickwalled and is elliptical slender, relatively in outline. Without any clearly marked line of separation it unites with the stomach. This last named organ,represented P1. VIII, Fig. 8, is likewisea simplestructure on with heavy foldsin the ventralsectionas its most prominent feature. of The positionof the bile ducts is shownin a reconstruction the digestivesystemof A. adamsi (P1. VIII, Fig. 2). There it may be seen that the anteriorend of the stomach projects forward beyond its union with the esophagus,and into this pouch two unusually elongated ducts enter fromthe digestivegland. The appearance of these ducts in cross section is shown on P1. VII, Fig. 3. In a positionmore posteriorand nearerthe dorsal surface of the stomach a second pair of ducts have their origin. These in section are represented P1. VIII, Fig. 14. In A. solida the numberand situationof these ducts are on essentially the same. In A. adamsi the heart appears with exceptional clearness,and accordinglya reconstruction was made of this portionof the circulatory system. From P1. VIII, Fig. 1, it will be seen that the pericardium a singlecavity,and, as shownin Fig. 6 of this same is plate, the ventricular pouches are united dorsal to the intestineby a spacious transverse channel whichanteriorly gives rise to the dorsal aorta. In ventralview (P1. XV, Fig. 2) the vessels, which connect with the ventriclesand originatethe ventral aorta, are comparativelyslender,and theircavities are somewhatill-defined. In the onlysectionedspecimenof A. solida the pericardium a singlecavity,the venis tral aorta and its connections are clearlydefined, but the dorsal union of the ventricular to pouches is very difficult trace. Even in grossdissectionthis holds true. In one individual only was it possible to trace the dorsal union of the heart, and even thereit was tenuous indeed. Auriculo-ventricular valves are present. As to the muscularsystem,the adductorsare well developed,and the pedal musculature is at least of average size. On the other hand, the muscles spanning the visceral weak. Sectionsof A. adamsi show a fewstrandsin the neighborcavity are exceptionally hood of the mouth,and not more than fouror fiveelsewherein the cavity. In A. solida they appear to be somewhat more numerous,but their exact number and distribution remainuncertainowingto the factthat in tracingthe digestivesystemtheyweredislodged to a greatextent. The main featuresof the nervous system are essentiallythe same as in B. (Acar) reticulata. The urogenitalsystem of A. solida is a close counterpart this organ in Trisodos of tortuosa. The main outlineof the entiresystemwas not determined, the union of the but terminal sectionof the gonoductwiththe kidneyproperand withthe renopericardial canal is the same, the only marked difference being the somewhat longer and more tortuous canal. courseof the renopericardial

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withan average lengthof This genus is represented one species, B. pectunculoides, by 10 mm.; accordingly greaterportionof the following the description based upon sections is and reconstructions. on The externalappearance of the body is represented P1. IX, Fig. 6. From this it will be noted that the palps are comparatively short,and the ill-defined groovesare confinedto the moredorsalportionsoftheseorgans(P1. VIII, Fig. 4). From sectionsit also is evident that the greaterportionof the mantle is glandular,the cells closely resembling homologouselementsin Arcopsisadamsi (P1. VIII, Fig. 13). Pelseneer ('11) in his account of the anatomy of Bathyarca sinuata describes and figures two peculiar folds of the mantle in the posterior end of the body. Similarorgans exist in the presentspecies (P1. VIII, Fig. 15). Regardingtheirfunction, Pelseneer describes them as "une sort de tube inhalant dont les deux moities ne sont pas soudees." This appears to be a reasonable assumption,althoughthe termexhalentmay be a more accurate term. In supportof the view that these foldsmay function a siphonit is to be noted that as theyare well supplied withmuscles (P1. VIII, Fig. 5), and it may well be that on occasion they may take the positionindicatedby the dotted lines in P1. IX, Fig. 4, and so forma siphonal chamber. Immediatelyabove the attachmentof each scoop just describedthe epithelium lining the inner face of the mantle is developed into a clearly definedcirculardisc (P1. VIII, Fig. 5, d). While the preservation not the best, the component is cells appear to be of two main types, goblet cells of various sizes and interstitial supportingcells. The larger or goblet cells contain a yellowishsecretionaftertreatmentwith Delafield's haematoxylin. No reasonablesuggestion theirpossiblefunction of comes to mind. The byssus systemcomprises, with delicatelycorrugated walls a first, narrowfurrow that extendsalong the anteriorhalf of the foot. Posteriorly this grooveleads into a deep depressioncontaininga highmedian fold. The anteriorhalfof this foldis non-glandular; the posterior half,on the otherhand, is richlysuppliedwithgland cells that evidently form the byssus (P1. VIII, Figs. 7, 10). As indicatedon P1. VIII, Fig. 11, and P1. IX, Fig. 2, the esophagusis a simpletubular structure withoutany marked dilations or folds. As to its glandularactivityconditions vary. In one specimena very few cells only contain a secretion. In anotherindividual a large numberof cells (P1. IX, Fig. 1) containa darklystainingsubstanceaftertreatment with Delafield's haematoxylin,which closely resembles the chromatinof an ordinary resting cell. This variationmay be correlated withthe feeding intervals, sincethe stomach of the firstnamed specimen contained an abundance of detritaland nutritivematerial, while that of the second individualis nearlyempty. In the more glandularspecimenthe distribution the gland cells extends fromthe mouth to the stomach. No special agof gregationat any point indicatesa pharyngeal region. The muscular coat of the esophagus is comparativelyweak, consistingof delicate longitudinal fibers, slenderbundlesofcircular and muscles,whiletheregionabout themouth is slightlyheavier due to radiatingfibers whichfromtheirarrangement opening probably act as dilators.

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No especial attempt was made to reconstruct the internalfeaturesof the stomach, and its generalshape only is indicated (P1. VIII, Fig. 11). The exact arrangement the of bile ducts likewise is somewhat diagrammatic,but the numberand general position of these is fairlyaccurate (P1. VIII, Figs. 3, 11). A gastricshield is present. The intestine followsessentiallythe same course as in Arca (P1. VIII, Fig. 8), and passing ventrallyto the pericardium opens at the end of a slenderstalk hangingfreely the mantle cavity it in (P1. IX, Figs. 2, 6). A reconstruction the heart shows that it closelyresemblesthat of Barbatia barbata of as describedby Menegaux ('90). It also is similarto that of Barbatia (Acar) reticulata (P1. VII, Fig. 8). A reconstruction the nervoussystemalso indicatesthat it is built upon essentially of the same plan as that of Barbatia (Acar) reticulata (P1. VI, Figs. 4, 5). The general configuration the urogenitalsystemclosely followsthat describedby of Odhner ('12) forArca glacialis, and of Barbatia (Acar) reticulata the presentwork (P1. in XV, Fig. 9). The gonoductterminates what may be termedan atrium,which,on the in other hand, communicatesdirectly with the kidney. The reno-pericardial canal also appears to connectwitha diverticulum the kidney,but its exact relationsare uncertain. of TRISODOS This genus is represented a singlespecies, T. tortuosa. In certaingeneralrespects by it resemblesa typical Arca, yet at the same time it presentscertainotherfeatureswhich are unique so far as the presentcollectionis concerned. Of these unusual charactersthe most striking its asymmetry. Scapharca, as is well known,possesses a shell in whicha is slightdifference existsin the size of the valves of the shell,but in Trisodosthis inequality is carriedout to a highdegree. Furthermore, these two typesalso differ that the asymin metryof Trisodosdoes not affect entireshell,but it occursin the regionposterior the the to umbones,that is froma point slightly anterior the heart (P1. X, Fig. 13). to The palps are more inclinedthan in Arca, theirposteriorextremities being located nearerthemid-region thebody. Sectionsalso indicate(P1. X, Fig. 5) thatthe customary of palp ridgesof otherspecies are here throwninto a seriesof minorfolds. Anotherunique featurerelatesto the abdominalsense organs(P1. XI, Fig. 1). In threespecimens examined on this point the asymmetry the posterior in regionsof the body has includedthese structures,the one on the rightside being markedlylargerthan that on the left. The byssus cavity (P1. XI, Fig. 5) is well developed,but in no specimenwas therea trace of a secretionalthoughgland cells were much in evidence. The anteriorfourthof the footis traversedby a shallow longitudinal groovewhichmoreposteriorly continued is into the main largerand deeper depression. The lateral walls of the posterior half of this last mentioned sectionare dorsallyfashioned into a seriesof delicate foldswhichdisappear more ventrally, theirplace being taken by a band of gland cells. In the posterior half of the cavity the ridgesof the walls are more pronounced, and at the extremeposterior end of the cavitytheyform markedclusterreminiscent the state of affairs A. (Barbatia) a of in for decussata, example. In this species the digestivesystemis excessivelydelicate, and the connectivetissue bindingtogetherthe various organsof the visceralcavity is so highlytough and resistant that ordinarymethodsof dissectionwere fruitless. Accordingly the anteriorhalf of the

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body was sectioned, and the followingdescriptionis based in large measure upon the reconstruction sections. of The mouth in surfaceview exists as an elongatedtransverseslit, resembling Arca sp. (P1. I, Fig. 2) in its shape and relationto the palps. From sectionsit may be seen that thereis a fairly sharp line of demarcationbetweenthe smoothepitheliumof the palps and the grooved walls of the esophageal tube (P1. 10, Fig. 3). In a more or less irregular fashionthese groovesand ridgesextend to the stomach withoutany othermarkedpeculiarities, and it may be added that throughoutits entire extent the esophagus is dorsoventrallycompressed. Certainauthorshave noted that in a fewspecies of arcids the esophagusshades gradually into the stomach,and in the presentcase it is difficult determine to the limitsof the two. On P1. X, Fig. 1, it will be seen that at a point approximately halfwaybetweenthe mouthand bile ducts thereis a distinctbend in the digestivetube. At this point thereis no markedchange in the natureof the liningepithelialcells,althoughit is to be noted that the tube posteriorto the bend is of smaller caliber,and the ridgesare distinctly fewerin number(compare Figs. 2, 4 of P1. X). Despite the fact that the liningcells on eitherside of the "bend" resembleeach other to a markeddegree,it nevertheless appears probable that at this point the esophagusends and the stomach begins. As a matterof fact thereis but little change in the character of the epitheliumbetween the mouth and the bile ducts. The digestive tube enlarges considerablybetween the bend and liver ducts, but otherwisethere are no especially distinctivefeatures. However, it is importantto note in this respect that in all of the Arcidae there is a distinctsection between the bile ducts and the undoubted esophagus. This interval,to be sure, varies in extent,but invariablyit is present. In this species, it therefore, appears that the most reasonablesolutionof the problemis to assume that the union of the esophagus and stomachis located at the point wherethe bend appears. Posteriorto the bile ducts the stomachbecomes a more complicatedorgan withwalls composed of cells of varyingheightsand fashionedinto ridgestoo complicatedto reconstruct. The lower divisionof the stomach is providedwith ridgeswhichmay correspond to those responsibleforthe secretionof the crystalline stile in otherspecies,but here they never are prominent (see rightside of stomachon P1. IX, Fig. 8). An unusual developmentexistsin the gastricepitheliumcoveringapproximately onethirdof the rightside of the lowerdivisionof the stomach (P1. IX, Fig. 8). Throughout this area the componentcells are low cubical elementsin marked contrastto their surroundingneighbors. Anteriorly is borderedby a distinctridgewhichextendsfromthe it to the junctionof the two divisionsof the stomach. Posteriorly shades graduintestine it ally into the usual type of epithelium. The configuration the upper divisionof the stomach is determined an unusual of to degree by ducts from the digestive gland. At not less than thirtypoints the gastric epitheliumformsdistinctdiverticula,each of which unites with secretingfollicles. On the rightside of the stomachthese are relativelyfewin number(P1. XI, Fig. 6), and thus are in marked contrastto those extendingacross the ventralsurfaceof the stomach and up its leftface. This last named groupis shownunitingwith the stomachin a successive series of sections (P1. X, Figs. 9, 10, 12). Also in diagrammatic fashionthese are shown on P1. X, Fig. 1.

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The intestineat a point nearest to the mouth is slightlymore complicatedthan in Arca, but otherwiseits course is essentiallythe same. Unlike Arca, however,the heart surrounds the gut (P1. XI, Fig. 10). Throughoutits entireextenta typhlosoleis present in the intestine, generalformbeing represented P1. IX, Fig. 8. As usual the anal on its openingis on the ventralsurfaceof the posterior adductor. Attention already has been called to the fact that the gut penetratesthe heart,and in a more detailed fashionthis fact is indicated on P1. XI, Fig. 10. As is there indicated, thereis but a single pericardialcavity, and distinctvalves guard the auriculo-ventricular give riseto a large anterior openings. Anteriorly ventricle ventricular the or outpouchings aorta. This same figure, togetherwith that of P1. XV, Fig. 1, also shows that thereis a similarbroad union ventralto the intestine. Evidentlythis unionis due to a fusionof the for outpouchings and not to the enlargement the ventralvesselsresponsible the formation of ofthe posterior and hold essentially aorta. As may be seen thesevesselsare clearlydefined, the same positionas in otherspecies represented the same plate. on The urogenitalsystem in this species is exceptionallyclearly defined,the greater numberof its principalfeaturesappearing in a single section (P1. X, Fig. 8). Here the externalopeningleads into an atriumwhich on one hand leads into the genital duct and canal and a passage into the at the same level connectswith the ciliated reno-pericardial the main kidneycavity as well. No attemptwas made to reconstruct shape of the entire in kidney,but a cross section (P1. XI, Fig. 3) shows the organ to be somewhattriangular roofto the secreting folliclesof the outline with a thin-walled, apparentlynon-glandular othertwo sides.
ANADARA

In this genus A. bisenensisand A. granosa are regardedas typical species. As may the the be learnedfromthe figures, externalfeaturesof both are essentially same (compare P1. XIII, Fig. 1, and P1. XIV, Fig. 7). The mantle in A. bisenensisis highlyglandular, the affects size of the palps whichin part much less so in A. granosa. Anotherdifference the may be due to shrinkage. In A. bisenensis abdominalsense organsvary to a considerable degreeas a comparisonof P1. XII, Fig. 5, and P1. XIII, Fig. 10, will indicate. This difference may again be due to the action of the preservative. Evidently the Fig. 10 the normal state of affairssince it is almost the exact copy of these drawingrepresents organsin A. granosa. The byssuscavityin bothspeciesis ofsmall size withwalls delicately ridgedin A. bisenensis(P1. XII, Fig. 9). The digestivesystem of A. bisenensisis shown on P1. XIII, Figs. 5, 6, that of A. it granosaon P1. XII, Fig. 10, and P1. XIII, Fig. 2. From these figures will be seen that there are no unusual featuresattaching to the esophagus, and in neitherspecies is the in different. In each a markeddiverticulum the gastricwall occursto stomachstrikingly its the rightof the esophageal opening,and, judging fromgross dissection, apex is firmly the attached to the body wall. Furthermore, removal of the connectivetissue fromthe visceral cavity doubtlessdestroyssome of the bile ducts, yet it is certainthat, at least in A. bisenensis, ducts are attached to the antero-ventral gastricwall, and one or more open on the rightside. is subfamily Arcinae, The intestine markedly longerthanin any speciesoftheforegoing concernsthat althoughin its generalpositionthereis an agreement. The chiefdifference section of the gut adjacent to the pedal ganglia which is throwninto several loops. It

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as passes to the ventralside of the pericardium it makes its way to the under side of the posterior shell adductor. The general appearance of the heart in both species is typified the sketchof this by into organ in A. bisenensis(P1. XIV, Fig. 3). A dye injected by means of a micropipette the pericardialcavity on one side of the mid-line passed readilyto the oppositeside in this species, indicatinga single cavity. The same methodindicatesthat in A. granosaa connection exists, but evidentlyit is relativelyof much smaller caliber. In A. bisenensis thereis a well definedflapwhichmay act as a valve at the junctionofthe ventricle and the anterioraorta (P1. XII, Fig. 6). The muscles of the visceral cavity of A. granosaare represented P1. XI, Fig. 12, on ofA. bisenensis P1. XII, Fig. 6. In thislast named speciesthereare two unique muscles on in whichare inserted thetissuesurrounding mouthopening. Both are unpairedstrands, the and the larger,extending across the visceralcavity,attachesa fewofits fibers the gastric to of wall in the neighborhood the bile ducts (P1. XII, Figs. 6, 8). The main bundle continues on to the junctionof the two divisionsof the stomachwhereit attaches to the gastricwall and to one of the ordinarytransverse muscles. The smallermate likewiseextendsacross the visceral cavity,and is attached to one of the smallertransverse muscles. of to a Regardingthe function thesemuscles,it is difficult form reasonableexplanation. Those fibers stomachwall may producea dilation,possiblyallowing attachedto the forward of a freerflowof digestivefluidthroughthe ducts, and a contraction the main muscle of may also dilate a portionof the stomach. Also those fibersattached to the transverse muscles may serve to enlarge the mouth opening or regulate in part the movementsof the palps. No especial study was made of the nervous,urogenitalor circulatory systems.
SCAPHARCA

This subgenus,embracingS. auriculata,S. concinnaand S. transversa, been estabhas in lished on the basis of a slightdegreeof asymmetry the valves of the shell. This state of affairs, however,is not correlatedwith any otherstriking differences betweenthis subgenus and typical species of Anadara. The mantle shows a weakly developed glandular differentiation it adjacent to the umbonesin S. auriculataand S. transversa; is more pronounced in S. concinna. Other externalfeaturesof the body are sufficiently represented to requireno further comment(Pls. XI to XIV). The byssus cavity is weaklydeveloped (P1. XI, Fig. 11) with the exceptionof an unidentified species fromthe coast of Senegal, Africa,in which this organ is more complicated (P1. VI, Fig. 10). The abdominal sense organs bear a close resemblanceto those of Anadara s. s., a typicalexample being that of S. auriculata (P1. XII, Fig. 7). The digestivesystempresentsno particularcharacterswhich separate this subgenus from typical anadaras. In every instance the esophagus is a simple tubular structure withoutany markeddilationsor glandulardevelopments. The stomachlikewiseis rather plain-walledin S. auriculata (P1. XI, Figs. 4, 7) and S. concinna(P1. XIII, Fig. 3). It is somewhatmore complicatedin S. transversa XV, Fig. 5). (P1. The intestinedevelops several loops in the neighborhood the pedal ganglia, but of possibly owingto theirsmallerbody size these are less complicated(as in P1. XII, Fig. 4) than in Anadara s. s. The gut passes by the ventral side of the pericardiumto open

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beneath the posterior adductor. The main musclesof the visceralcavity are represented on P1. XII, Figs. 2, 3, and P1. XIV, Fig. 9. No sectionswere made of any of these species, but it has been determined that the is pericardium a singlecavityin S. concinna.
CUNEARCA

On the basis of shell charactersthe threespecies, C. chemnitzi, perlabiata,and C. C. incongrua,are credited with a close relationship. From an anatomical standpoint this classification appears to be correctso far as the firsttwo are concerned,but, as will be pointedout later on, thereare certaindifferences whichappear to throwsome doubt upon the status of C. incongrua. Externallyall of these species exhibitthe same generalappearance (P1. XIV, Fig. 4, P1. XVI, Fig. 2, P1. XVII, Fig. 5). The palps are nearlyverticalor but slightly inclined, a statementeven more accurate with respect to the upper half of the gills. A section throughthe palp of C. perlabiatais shown on P1. XV, Fig. 8. The byssus is small in C. chemnitzi XI, Fig. 11), and somewhatlargerin C. perlabiatawherea definite (P1. secretion has been developed (P1. XIV, Fig. 4). Elsewherein this paper attentionis called to the fact that in a genus or subgenusthe various species exhibit a complex of characterswhich fundamentally are similar. And in this connection is importantto note that this also holds true withrespectto the abit dominal sense organs. In this subgenus Cunearca the three species possess these organs fashionedaccordingto two patterns,and, as will be seen, there are two corresponding types of digestivetractsand of the transverse musclebundlesin the visceralcavity. The first type of abdominal sense organis presentin C. incongrua(P1. XVII, Fig. 3) wherethe anal openingand the regionimmediately anteriorto it are borderedby delicate foldswhichmay be sensoryin character. The second type is presentin C. chemnitzi (P1. XV, Fig. 11) and C. perlabiata(P1. XIV, Fig. 6). Here a fold,in the formof an arc, and located immediatelybehind the anal opening,has developed a conspicuous yet slender finger shaped projectionin the mid-line. The esophagus is comparatively and C. perlabiata(compare P1. shortin C. chemnitzi XIII, Fig. 8, and P1. XV, Fig. 7). Also the stomach of these two species followsmuch the same pattern. In each a foldof the gastricwall is situated dorsallyto the esophageal openingand one of the folds,coursingthroughout lengthof the ventraldivisionof the the stomach,terminates dorsallyin a conspicuously developedknob. A verydifferent picture appears in the case of C. incongrua(P1. XVI, Fig. 1). Here the dorsal gastricfold holds a different position,and, althoughthe ventralfold ends in a knob, the gastricepithelium presentsan appearance unlikethat in the othertwo species. do From these same figures will be seen that the relative lengthsof the intestines it not followthe rule accordingto whichthey should be of the same lengthin C. chemnitzi and C. perlabiata. As a matter of fact this organ is relativelyshort in the firstnamed species,whereasin the otherit is of approximately same lengthas that of C. incongrua. the In all threespecies the gut passes ventrally the pericardium it makes its way to the to as openingbeneath the posterior adductor. The patternofthe musclesbridging visceralcavityis muchthe same in C. chemnitzi the (P1. XVI, Fig. 6) as in C. perlabiata(P1. XVII, Fig. 1). The resemblancein the case of

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C. incongrua more remote(P1. XVI, Fig. 3), the bundles in the regionof the heart and is those forming posteriorwalls of the cavity being heavier than in any otherspecies of the the presentcollection. As just noted, the heart is dorsal to the intestine. The pericardialcavity in C. perlabiata was found by the use of sectionsto be singleand auriculo-ventricular valves were present. No sections were made of C. chemnitzi, but in the single individual examined on this point no dye injected on one side of the mid-line passed to the otherside. In C. incongrua the generalappearance of the heart is practicallythe same as that of C. perlabiata(P1. XIV, Fig. 11), and as in that species the pericardium a single cavity. is No sectionswere made of the othertwo species of this subgenus,but in C. incongrua the organs in the visceral cavity were imbedded in connectivetissue so tough and resistant that a small specimen,19 mm in length,was sectioned. In this mannerit was discovered that the connectivetissue "packing" was penetratedby a system of capillaries rather than by sinusesor lacunae of the usual type. Whetherthe termcapillaryis justified may be open to question. However, these vessels are clearly defined,and appear to possess is definite walls. In any event each liverfollicle moreor less surrounded one or moreof by these channels,a condition that is unique, at least so far as the present collectionis concerned. The urogenital systemof C. perlabiatawas studiedby means of sections,and, although it the materialwas not well preserved, appears that the generalplan is similarto that of Acar reticulata(P1. XV, Fig. 9), forexample. There is no doubt that the duct fromthe kidneyunites in the same fashionwith the gonoduct,and close to this point of union the funnelappears to enterthe kidney. reno-pericardial
ARGINA

of on Argina pexata is the sole representative this genus, the individual figured P1. XVIII, Fig. 3, having a length of 52 mm. The mantle, especially in the region of the 1 umbones,is highlyglandular. The pedal furrow, averagingapproximately mm in depth, is smooth-walledthroughoutwithout any sign whatever of a byssus. The abdominal sense organsare shownon P1. XVIII, Fig. 5. withan innerlining caliberthroughout, The esophagus,as usual, is a tube of uniform thrown into delicate longitudinal folds. The characteristicsof both divisions of the stomach are sufficiently represented(Pls. XVII, Fig. 8, XVIII, Fig. 4, XIX, Fig. 3) to showthe complexity the intestinal of comment. Also thesesame figures demand no further coils. Owing to the resistantcharacterof the connectivetissue in the visceralcavity,the are bile ducts were detached, and hence their numberand distribution unknown. It is on certain,however,that the gut penetratesthe heart,and its outer openingis figured P1. XVIII, Fig. 5. In the drawingof the muscularsystemspanningthe visceral cavity it is to be noted that, with the removal of the digestivesystemand the associated connectivetissue, the more delicate muscles were displaced. Thereforein the figure(P1. XVIII, Fig. 6) only musclesare represented. the more prominent As noted previously, heart surrounds the intestine, and gives rise to the unpaired the anterior (P1. XIX, Fig. 1) and posterioraortae. The auricles ventrallyare distinctly separated by a thin median partition. This partition,however,is incomplete,the two

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division of the pericardial cavity communicating a small foramena short distance by to anterior the pedal retractors. No attempt was made to trace the circulatory systembeyond this point, nor were the nervousor urogenitalsystemsinvestigated.
NOETIA

This genusis represented a singlespecies,N. ponderosa, generalexternalappearthe by on ance of whichis represented P1. XVIII, Fig. 1. The mantle is opaque and relatively thickdue to the usual glandulardevelopment. The pedal furrow comparatively is shallow, and posteriorly median terminates a byssus cavity of small size with a small irregular in roof. The abdominalsense organsare shownon P1. XIX, Fig. 2. foldand corrugated The esophagusis flattened dorsoventrally, otherwise unmodified. The stomach, but is on the other hand, is unusually complicated (P1. XVII, Fig. 11, and P1. XVIII, Fig. 2). A gastricshieldis present, withthe dorsal foldnear the anterior havingits main attachment end of the stomach. Also in this particularspecimenthe stomachwas entirely filledwith secretion whichobscuredthe finer stile details of the gastricepithelium. crystalline the Bile ducts open through antero-ventral gastricwall, and also on the rightat about the same level. At these points thereare small conical papillae (P1. XVII, Fig. 11), but the ducts actually open through whether themcould not be determined satisfactorily. of the heartis shownon P1. XVII, Fig. 7. A definite The generalappearance septum dorsal to the intestine, and dye injected into the cavity did not separates the pericardium cross the mid-ventralline. It thus appears that two cavities exist. The dorsal aorta to courses anteriorly the rightof the mid-lineas far as the pedal protractor, and, after a small branchto the cerebralganglia,it ramifies supplying the throughout visceralcavity. of The musculature the visceralcavity is represented P1. XXI, Fig. 9. From this on it is evidentthat in thisrespectthe genus Noetia does not appear to be verycloselyrelated to otherspecies in the presentcollection. No seriousattemptwas made to trace the various branchesof the nervoussystem,or the relationsof the urogenitalsystem. It may be added, however,that an to determine with the reproductive unusual featureoccursin connection systemwherea portionof the of in gonad occupies spaces among the fibers the pedal retractor the regionof the "heel."
CUCULLAEA

in Cucullaea granulosais represented the presentcollectionby a single individual 70 features represented P1. XVI, Fig. 4. The mantle mm in length. Its main external are on and evidentlylacks the glandulardevelopmentcharacteristic is unusuallytransparent, of the average arcid. The abdominal sense organs are illustratedon P1. XX, Fig. 9. The footis traversedby a longitudinal the groovethroughout greaterextentof its ventralface. Near the "heel" of the foot the wall of the byssus cavity is fashionedinto a small yet distinctmedian fold bounded externally three othersof much smallersize (P1. XVII, by withPelseneer'sdescription Fig. 10, and P1. XIX, Fig. 5). This is in essentialagreement in and figure the Siboga report. The esophagus is a relativelysimple tube, thick walled, and internallyis provided with a series of longitudinalfolds. The stomach likewisepresentsa comparativelyunmodifiedappearance both externallyand internally(P1. XIX, Fig. 6, P1. XXI, Fig. 8).

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In the presentspecimenthe dorsal gastricdivisionwas laterallymuch compressed, and to the rightof the esophageal openingtherewas a deep lateral pocket the blind end of which was firmly anchored to the body wall. The gastric epitheliumof the lower division of the stomachwas foldedinto seven longitudinal ridges. It maybe added thatboth divisions of the stomach were completelyfilledwith crystalline stile secretion. Also one bile duct opened on the leftface of the stomach (P1. XXI, Fig. 8), two were more ventrallyplaced near the mid-lineof the body, while a thirdwas situated about the middle of the right gastricwall. and in the neighborhood The intestine, upon leaving the stomach,coursesanteriorly of the pedal ganglia turnssharplyupon itselfand, in close contact with the rightside of whereit is surrounded the heart. Its the stomach,makes its way to the pericardium by on openingupon the ventralsurfaceof the shell adductoris represented P1. XX, Fig. 9. The main featuresof the musculaturein the greaternumberof species treatedin the present work conformto a broad general plan. Hence, in order to avoid unnecessary repetition, muscularsystemof Cucullaea granulosais herewith the describedin some detail, of and only exceptionsto the rule will be noted in the description otherspecies. and the pericardium, The dorsal wall of the body, between the pedal protractors is reinforced a series of more or less transversemuscle bundles (P1. XX, Fig. 12). In by dissectionsthese are seen to be continuedlaterally,and to share in the formation the of in surface view are several muscle bundles which lateral body wall. Less conspicuous are insertedin the dorsal body wall close to the mid-dorsalline. These ventrallybreak up into a series of radiatingfibers, and, extendingacross the visceral cavity, also forma portionof the lateral body wall. Owing to the large size of this species, a more than usual amount of care has been taken to examinethe muscularsystemin the regionofthe mouth. As usual, the esophagus itself is supplied with circular and longitudinalfibers. The dorsal palp appears to be fromthe pedal protractor, a operated by, first, delicate offshoot and, secondly,thereis a well developed muscle which on one hand radiates, and attaches to the body wall into is in whilethe otherextremity insertedin the regionimmediately whichit is incorporated, frontof the palp, at the same time sendinga small numberof strandsinto the palp itself. of and thusperhaps Evidentlythe main function thesemusclesis to elevate the oral region, the musclesin forcing food onwardtowardthe stomach. cooperatewiththe esophageal filaments In additionto the above describedmusclesthereare numerous from radiating the esophagus in the oral regionsome of whichare insertedin the body wall, whileothers extend in among the folliclesof the digestivegland whereevidentlythey are anchoredto the connectivetissue. Obviously the functionof these strandsis to increase the caliber of the esophagus. the on musclesbridging visceralcavityare represented P1. XX, Fig. 12. The transverse in shape and situated immediately behind the It is to be noted that one, highlyirregular oral region,gives rise to a single strand (not figured)that, as in Anadara bisenensis(P1. some ofthe fibers attach XII, Fig. 8), makes its way to the stomachwall. In A. bisenensts wall whilethe main cordpasses on to the junctionofthe two divisions the anterior to gastric of the stomach. In the presentcase it is not certainthat any fibers pass to the anterior the two species are similarin this respect. regionsof the stomach,but otherwise

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The pericardiumand heart of this species are represented P1. XV, Fig. 10, and on P1. XIX, Fig. 4. The ventricle surrounds intestine, the and the aortae originate normal in fashion. Each auricle,it may be added, extendsto the mid-ventral ofthe pericardium, line in thus terminating a cul-de-sac. The details of the nervousand urogenitalsystemswere not determined.
GLYCYMERIS

The intestineis comparativelyshort, and in most instances its formand location in that occuring Arca. Its mostcomplicateddevelopment resemble appears in G. americana (P1. XXI, Fig. 5). In everycase the heartsurrounds the gut. The musculaturepresentsno noteworthy featureswith the possible exceptionof the transversemuscle bands spanning the visceral cavity. In the two smaller species, G. these are relativelyslender and delicate, whereas in G. migueliana and G. subobsoleta, americana and G. Iongiorthey are unusually well developed. In another connection of (Heath '37) it was suggestedthat by the contraction this set of musclesthe volume of the visceral cavity is decreased, and that a portionof the blood in the visceral sinusesis thus forced into the foot. Hence these muscles serve indirectlyas pedal protractors. Their development, therefore, may be correlatedwith body size and weightand possibly withthe natureof the sea bottomover or through whichthe animal progresses.

position.

This genus is represented fourspecies, G. americana,G. migueliana,G. longior by and G. subobsoleta. These forma closely related group,the only differences appearing to be those relatingto minordetails. In every instance eye spots occur on the mantle border the half. The footis well developed,and is providedwitha shallow throughout posterior plain walled groove. No microscopicalsections were made, but macroscopicallythere are no evidences of a definite glandular development. The palps differ considerablyin in relativesize, due possiblyto contraction the preservative, and the characteristic grooves are fainteven in stained preparations. The digestivesystemin all of the speciesis constructed the upon essentially same plan. In some individualsthereare slightvariations,such as the presenceor absence even in the same species,of the delicate radiatinggroovesin the upper divisionof the stomach (as in P1. XIX, Figs. 7, 8), but otherwise thereare no important differences. The esophagus is compresseddorso-ventrally, the liningepitheliumis fashioned and into a series of delicate longitudinal folds. At no point is therea noticeableenlargement or any indicationof glandular developmentsuggestiveof the pharyngealcavity in other classes of mollusks. The generalappearance of the stomach of the various species is represented Pls. on XIX, XX, XXI. In G. subobsoleta ventraldivisionof the stomach is smoothwalled, the or is provided with one or two very small longitudinalfolds. In the other species these foldsusually are more conspicuous,but even here the variationmay be considerable. In mostinstancesthe lowergastricsectioncontainedan abundance ofcrystalline stilesecretion. One bile duct, ventrallysituated anteriorto the junction of the two divisionsof the a stomach,is universallypresent. In G. subobsoleta second duct opens behind the union of the esophagus and stomach. In G. iongiora single duct opens at approximately the centerof the rightside of the stomach. In G. americanathereare two ducts in the same

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At thisstage attentionis called to the factthat severalspecies in the presentcollection possess muscles,usually extremely delicate whichare attached to the stomach,and usually are insertedin the body wall. However, owing to theirdelicacy,and the tenacious connectivetissue of the visceral cavity,it usually is impossibleto carryon ordinary dissection withoutdislodging these strands. In G. subobsoleta connectivetissue is weak, and the the major portionof these gastricmusclesis represented P1. XXI, Fig. 4. They doubtless on act as dilators. The abdominal sense organsvary to a slightdegree. What appears to be the typical conditionoccurs in G. americana (P1. XXI, Fig. 6) and G. migueliana(P1. XX, Fig. 10). In both of these species the rectumextendsbeyondthe level of the sense organs,a state of affairs fromthat of G. Iongior(P1. XX, Fig. 5). differing RE'SUME Accordingto Reinhart's classification('35), the familyArcidae includes three subfamilies. The first, Arcinae,embraces,so far as the presentcollectionis concerned, Arca, Barbatia, Arcopsis, Bathyarca and Trisodos. The second subfamily,Anadarinae, is represented Anadara, while Noetia is includedin the subfamily by Noetiinae. In an attemptto determine what extentthisschemeof classification to actually repreit sentsphylogenetic relationships has becomeincreasingly evident,as Pelseneeremphasized to years ago, that we must carryour investigations a greaterlengththan existsat present. In this family,the Arcidae, not over a dozen species have been investigatedup to the theirdevelopment. Neverpresenttime,and nowhereis thereany information regarding of thelessvarioustheorieshave been proposedto accountforthe phylogeny thisand related and while most of these theories families,or forthe originof certainanatomical features, may be discardedat a laterdate theyare valuable aids in focussing attention upon the more importantproblemsin need of solution. As Weismann remarked,"the way to truthis oftenthroughinevitableerror,"and with this dictumin mind various data are presented in the following paragraphswhichit is hoped may lead to a truerknowledgeof the group. During the courseof the presentworkit became evident (as it has to otherinvestigaon tors) that classification the basis of a single characteror individual systemof organs may lead to faultyconclusions. Obviouslywhereonly fossilremainsare available classicannotbe otherwise, wherelivingrepresentatives available, shell characters fication but are and the internalanatomyshouldbe treatedas a unit. The validityofthisline ofreasoning is indicated in the case of Arca imbricata, example. Here is a species,represented for by two individuals,the shell characters whichare declaredto be identicaland to be typical of species of the genus. From the standpoint of their internalanatomy they are unlike. Also fromconchologicalcriteria threespecies have been includedin the subgenusCunearca. Anatomicallyit is certainthat one of these is incorrectly included. In the same category is Acar pernoideswhichdiffers materially fromotherspecies of the genus. But whilethereis no obvious objectionto the plan of including studyof the internal a anatomy togetherwith the shell characters, thereare certaindifficulties.As all students of the subject are well aware, some systemsof organsare moresubject to modification than others. And what adds to the difficulty the fact that the range of variationin one subis genus or genus is greaterthan in another. Hence, as usual, the interpretation the data of and the formulation a schemeof classification of become mattersof personaljudgment.

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However, it appears to be fairlywell establishedthat in the fanmily Arcidae the urogenitalsystemis constructed upon muchthe same plan throughout. Also it seems equally certainthat the nervoussystemis highlyconservative, undergoing slightchanges only in the various included species. Possibly Acar pernoidesmay prove to be an exception. But while this may be the case it apparentlydoes not apply to some of the sense organs which it supplies. Pelseneer ('11) in his study of the "eye spots" of various pelecypods has shown that, at least broadly,these structures aid in the formulation a scheme of of in classification. Furthermore, several instancesin the presentstudy, wherethe general internalanatomyindicatesthe genericor subgeneric rank of certainspecies,the abdominal sense organsexhibita close resemblance. To what extentthese structures may be relied remains in doubt at present,but most emphaticallythey upon in tracingrelationships meritfurther investigation. considerabledoubt exists at present with respect to the byssus as a Furthermore, featurein classification. In some instancesit appears to be of value when distinguishing it is consideredalong withotherorgansof the body. In othercases its value is doubtful. The question calls for a broader survey beforeits true worthcan be determined. The same line of reasoningapplies to the transverse musclesspanningthe visceral cavity. In some species these are of such a distinctive size, or theirdistribution such that together is withotherorgansof the body theydoubtlessshould be taken into consideration. A buccal or pharyngealcavity may have been present in the ancestral pelecypod, its but so far as the presentcollectionis concerned existenceis doubtfulindeed. In some individuals there were slight dilations in the region of the mouth or at varyingpoints betweenthe mouthopeningand the stomach,but in otherspecimensof the same species the no trace of these existed. Furthermore, epithelialliningthroughout was of uniform character,and where sectioned material was available the same types of cells occurred were throughout. Where gland cells existed these, with the exceptionof Acar pernoides, or evenlydistributed, wherethey were moreabundant in the mouthregiontheirnumbers gradually lessened as the stomach was approached. In short,no definite buccal region was distinguished. In the present work various other featuresof the digestive system have been investigated,but it should be noted that the resultslargelyare based upon grossdissection. Where histological methods are employed the results may disclose resemblancesand but at presentthis systemappears to be relatively of differences value in classification; markeddifferences. However, plastic, some species even in the same subgenusexhibiting thereare indicationsthat the lengthof the intestinemay serve,in some instancesat least, to distinguishsubfamilies. For example, this organ is comparativelyshort throughout the entiresubfamilyArcinae, whereas it is of greaterlength and more contortedin the Anadarinae. of of The numberand distribution the bile ducts likewisemay aid in the classification or are subfamilies even subgenera. In grossdissectionthese structures so delicate and so readily dislodged that in a few species only have their exact numberand position been determined. In a fewcases a gastricshield is knownto exist,and in severalothersit appeared to be stile secretioncoated the lining present. In the majorityof the specimensthe crystalline of ofthe dorsaldivision the stomachto such an extentthat ifany shielddid existits presence

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could be detected only after sectioning. The value of this structurein classification is therefore unknownquantity. an In this familythe pericardium and heart have been the subject of investigation and discussion to a greaterextent,excludingthe shell, than any other organs of the body. From the time of Poli (1795), who discoveredthe paired heartsof Arca, until the present time a very considerablenumberof investigators have published studies along this line, so that now we have a fairlyaccurate pictureof these organs in approximately dozen a species. The whole subject is adequately reviewedin the worksof Theiler ('07) and Matthias ('14). A few additional items included in the next paragraphsare based upon findings duringthe courseof the presentwork. That Arca noae,forexample,has a double heartno one denies. But thereare numerous species in which it is difficult decide whetherthe heart is a single chamber with to lateral dilationsof the ventricle whether actually is double. Theilercuts the Gordian or it knot betweenthe stage wherethereare two pericardialcavities on one hand and a single one on the other. A paired pericardium indicates a double heart,a single cavity an unpaired heart. And this appears to be the best line of cleavage that any one has proposed. No doubt exists as to the dorsal position of the heart with reference the intestine to in those speciesof Arcidae witha singleheart,and in species,such as Arca noae or Barbatia barbata,the heart is still consideredto be dorsal althougha ventricular bridgeis lacking. In thislattercase the junctionis accomplished theunionoftwo aortae the fusion which of by in the mid-lineresultsin the formation the anterioraorta; and in the same fashionthe of ventralaorta is formed the union of two aortae. by in those more highlyspecialized species the dorsal aortae appear unFurthermore, questionablyto be arterialin charactersince they resembleundoubted arterieselsewhere in the body. On the otherhand, the ventralaortae are not so sharplydifferentiated.As various authors have noted, this ventralsystem,as it leaves the ventricles ventricular or pouches,is oftendifficult trace since the lumina are crossedby musclefibers to the general appearance and arrangement whichsuggestthat actuallytheyare ventricular of elongations ratherthan definite blood vessels. And this appearance on occasion is heightenedwhen the vessels are gorgedwith blood. In Arcopsis adamsi, forexample, the ventral vessels in one specimenwere so distendedwith blood that it appeared to be certainthat the gut penetratedthe heart. In two otherindividualssubsequentlyexamined the heart was of the usual type (P1. VIII, Fig. 8). In this connection is interesting note the state of affairs Trisodostortuosa. In it to in this speciesthe pericardium a singlecavity (P1. XI, Fig. 10), and the heartsurrounds is the intestine, and, it may be added, the dorsal bridgeunitingthe ventricular pouches is of the same size as the ventralunion shown of P1. XV, Fig. 1. Furthermore, is a significant it fact that the ventricularpouches have fused ventral to the gut, and in the process the roots of the ventral aorta have retainedtheirindividuality. In other words,it appears to be certainthat these last named vessels are actual arteries. At the presenttime it appears to be a futileundertaking account forthe double to heart in Arca, or to decide whetherit represents primitive a state or not. It is generally agreed that it occurs in those species in which the pedal retractors have become greatly the widthof the body. During this processthe gills and their developed,thus increasing attendantsinuses have been drawn farther and farther away fromthe mid-line. In such

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fashionvarious authorsaccount forthe gradual lateral extensionof the heart and its final separationinto two distinctdivisions. Accordingto this view the double heart has been derived froma single one. On the other hand there are others,notably Milne-Edwards and Thiele, who considerthat the paired heartis the moreprimitive. In the presentcollectionthereis a definite relationin Arca s. s. betweenthe size of the retractors and the degreeof separationof the ventricles. The largerthe musclesthe wider in the gap. And it appears reasonableto go a step farther directing attentionto the fact of that the size of these musclesis correlated withthe development the byssus. Furthercondimore,it scarcelycan be arguedthat in mollusksat least a sessilestate is a primitive it tion. Accordingly would logicallyfollowthat the pedal retractors were of smallersize in the ancestraltypes,and the widthof the body was relatively less. From whichit may be argued further that the type of heart in Barbatia, forexample,more closelyresembles an ancestralconditionthan it does in a typicalArca such as A. noae. However,these are pure assumptions;we need facts,many morethan we have at present. Matthias has discussed exhaustivelythe various theoriesto account for the phyloof geneticbeginnings the arcid heart, and the presentdata add little if anythingto the subject. All that we knowat the presenttimeis based upon not morethan a dozen species, withotherclasses of mollusksor annelids and also we have no assurancethat comparisons are legitimate. Moreover, we have no assurance that the developmentof a freshwater a followsthe same species, such as Cyclas corneaof Nucula delphinodonta, protobranch, course as that of Arca the developmentof which is unknown. Until we have such inand a widerview of filibranch is anatomy,it would appear that our theorizing formation, littlemorethan a mentalexercise. differences The more significant between the various species are included in the folof and lowingparagraphs. The most striking these occurs in the case of Arca imbricata A. sp., as detailed in the main body of the text. The fourother species in the present collectionare typicalarcids. The genus Barbatia is represented two typicalspecies,B. barbataand B. decussata. by reticulata and pernoides, includedin the subgenusAcar. Accordare Three species,gradata, ing to some authors the firsttwo are believed to be conspecific. Reinhart ('39) argues which against this point of view, and in the presentwork (page 296) evidenceis presented thereare thatin thecase ofA. pernoides supportshisclaim. This same authoralso maintains betweenits shelland that ofA. gradata,and that theyare distinctspecies. slightdifferences fromany other for is is This certainly a legitimateconclusion, A. pernoides quite different species in the present collection. Reinhart ('35) also states that Cucullaerca,fromthe standpoint of shell structure," comprises a clearly recognizable group." Its internal that of B. barbata. organization, however,resembles and distinctfromother genera in the family. The genus Arcopsisis clearlydefined, is Bathyarca,represented B. pectunculoides, equally sharply demarcated. The genus by and unusual type of stomach and heart,is even Trisodos,with its remarkableasymmetry more distinctand remotefromothergenerain the family. The subfamilyAnadarinae is represented the genus Anadara of which there are by in two typical species, A. bisenensisand A. granosa. Owing to a slightasymmetry the valves of the shell, the subgenusScapharca has been establishedwith its membersin the present collection comprisingS. auriculata, S. concinna and S. transversa. However,

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from the standpoint of their internal anatomy these three species ordinarilywould be consideredto be typicalmembersof the genus. The subgenus Cunearcais creditedwiththreespecies in the presentlist, C. chemnitzi, two are typical,but the status of C. perlabiataand C. incongrua. Anatomicallythe first betweenthe two groupsare detailed is incongrua open to doubt. Fundamentaldifferences in the description the subgenus. of Noetia ponderosais the sole species belongingto this genus. It does not appear to be closelyallied to otherspecies in the family. At the suggestionof Doctor H. G. Schenck, Cucullaea granulosaand a few members to descriptions. Their relationship of the genus Glycymeris included in the foregoing are the Arcidae is obscure.
ZI CARAZ 1902 Contributo all'istologia e alla fisiologiadei Lamellibranchi. Intern. Monatsschr. f. Anat. u. Physiol., Bd XX. DREW 1902 The life historyof Nucula delphinodonta. Quart. Journ.Mic. Sci., Vol. XLIV. GROBBEN 1893 Zur Kenntnis des Baues von Cuspidaria, nebst Beitrachtungenuiberdas System der Lamellibranchiaten. Arb. a. d. Zool. Inst. Wein, Bd. X. HEATH 1937 The anatomy of some protobranchmollusks. Memoirs du Mus. Roy. d. Hist. Nat. de Belgique, Ser. 2, fasc. 10. LIST 1902 Die Mytiliden. Fauna u. Flora d. Golfes v. Neapel. Monograph XXVII.
MATTHIAS

BIBLIOGRAPHY

1914 Vergleichendanatomische Untersuchungenuiberden Darmkanal und das Herz einigerArcaceen. Jen. Zeitschr.f. Naturw., Bd. LII. MENEGAUX 1890 Recherches sur la Circulation des Lamellibranchesmarins. Besangon. ODHNER Morphologische und phylogenetischeUntersuchungenuiber die Nephriden der Lamellibranchien. Zeitschr.f. wiss. Zool., Bd. 100. PELSENEER 1891 Contribution'a 1'tude des Lamellibranches. Arch. de Biol., T. XI. 1911 Les Lamellibranchesde l'Expedition du Siboga. Monograph Lllla. REINHART 1935 Classification of the pelecypod family Arcidae. Bull. Mus. Roy. d'Hist. Nat. de Belgique, T. XI. 1939 The Holotype of Barbatia (Acar) gradata (Broderip & Sowerby). Trans. of San Diego Soc. of Nat. Hist., Vol. IX. STEMPELL 1898 Beitrage zur Kenntnis der Nuculiden. Zool. Jahrb. Suppl., Bd. IV. THEILER 1907 Zur Anatomie und Histologie des Herzens von Arca. Jen. Zeitschr.f. Naturw., Bd. XLII. THIELE der 1891 Die Stammesverwandtschaft Mollusken. Jen. Zeitschr.f. Naturw., Bd. XXV. Die abdominalen Sinnesorganeder Lamellibranchier. Zeitschr.f. wiss. Zool., Bd. XLVIII.
ZIEGLER

1885 Uber die Entwicklungvon Cyclas cornea Lam.

Zeit. f. wiss. Zool., Bd. XLI.

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EXPLANATION
PLATE I

PHILOSOPHICAL

SOCIETY

OF PLATES

FIG. FIG. FIG. FIG. FIG. FIG.

nerves.
FIG.

1. 2. 3. 4. 5. 6. 7.

Arca sp., length84 mm. Arca sp., ventral view of anteriorend of body. Arca sp., byssus cavity, X 1.5. Arca sp., lateral view, X 1. Arca sp., section throughgonad. Arca sp., dorsal view of cerebro-pedal system; a, border of shell adductor sheath; m, mantle Arca sp., righthalf of body, lateral view. Arca sp., section throughone of the gonad tubules.
PLATE

FIG. 8.

II

1. Arca sp., righthalf of digestivesystem, X 5. FIG. 2. Arca sp., lefthalf of digestivesystem, X 5. FIG. 3. Arca sp., visceral ganglia and nerves, X 4; b, brancheson dorsal body wall; g, pair of nerves to gills; m, mantle nerves; s, nerve to abdominal sense organ. FIG. 4. Arca sp., posteriorend of body. FIG. 5. Arca noae, folliclesof gonad. FIG. 6. Arca sp., gonad tubules. FIG. 7. Arca sp., nervous systemof leftmantle lobe. FIG. 8. Arca bouvieri, dorsal view, X 2.
FIG.

PLATE III

1. FIG. 2. FIG. 3. FIG. 4. FIG. 5. FIG. 6. FIG. 7. FIG. 8. FIG. 9.

FIG.

Arca zebra, X 1.5. The anteriorend is slightlywidened. Arca zebra,rightside of digestivesystem, X 4; r, pedal retractor. Locality, North Carolina. Arca zebra,rightside of digestivesystem, X 4. Locality, Porto Rico. Arca zebra,leftside of body, length,60 mm. Arca noae, leftside of stomach, X 4. Arca bouvieri, ventral view of anal region, X 5. Arca pacifica,ventral view of anal region, X 6. Arca noae, rightside of digestivesystem, X 4; r, pedal retractor. Arca zebra,ventral view of anal region, X 3.
PLATE IV

1. FIG. 2. FIG. 3. FIG. 4. FIG. 5. FIG. 6. FIG. 7. FIG. 8. 2, P1. iv. FIG. 9. FIG. 10. FIG. 11. FIG. 12.
FIG.

Arca pacifica,lateral view, length,82 mm. Arca bouvieri, righthalf of digestivesystem, X 3; r, pedal retractor. Arca pacifica,dorsal view. Arca umbonata, righthalf of stomach, X 5; r, pedal retractor. Arca umbonata, lefthalf of stomach, X 5. Arca umbonata,dorsal view, X 2. Barbatia candida, dorsal view, X 2. Arca bouvieri, left side of digestive system, X 4; r, pedal retractor. Anotherspecimen Fig. Arca pacifica,rightside of stomach, X 5. Barbatia decussata,dorsal view, X 2. Arca bouvieri, byssus cavity, X 3. Arca zebra,byssus cavity, X 4.
PLATE V

FIG. FIG.

1. Barbatia decussata, X 2. 2. Barbatia barbata, X 2.

HEATH: ANATOMY OF PELECYPOD

FIG.

FAMILY ARCIDAE

315

3. FIG. 4. FIG. 5. FIG. 6. FIG. 7. FIG. 8. FIG. 9. FIG. 10.

Arca pacifica,leftside of stomach, X 5. Barbatia candida, rightside of stomach, X 4. Barbatia candida, leftside of stomach, X 4. Anadara granosa, X 2. Abdominal sense organs. Barbatia barbata,dorsal view, X 2. Barbatia decussata,muscles of visceral cavity, rightside, X 4. abdominal sense organs, X 5. Barbatia pernoides, Barbatia barbata,abdominal sense organs, X 5.

PLATE VI FIG. 1. Barbatia reticulata, lateral view; length,13 mm. FIG. 2. Barbatia decussata,lateral view of digestivesystem, X 8. FIG. 3. Barbatia decussata,righthalf of body, X 3. FIG. 4. Barbatia reticulata, dorsal view of cerebro-pedalnervous system,a reconstruction;a, nerve to anteriorshell adductor; m, mantle nerves; o, otocyst. FIG. 5. Barbatia reticulata, dorsal view of visceral ganglia and nerves, a reconstruction;d, nerve to dorsal surfaceof the body; g, nerve to gill; m, mantle nerves. FIG. 6. Acar pernoides, dorsal view, X 5. FIG. 7. Acar pernoides, section throughstomach and bile ducts posteriorto P1. VII, Fig. 1, and along line b, P1. VII, Fig. 11. FIG. 8. Acar pernoides,longitudinal section through kidney and gonoduct; a, pericardial cavity; s, gonad. of FIG. 9. Barbatia reticulata, reconstruction digestivesystem. FIG. 10. Barbatia decussata,byssus cavity, ventral view, X 4. PLATE VII FIG. 1. Acar pernoides, section throughstomach and bile ducts anteriorto P1. VI, Fig. 7, and along line a, P1. VII, Fig. 11. FIG. 2. Arcopsis adamsi, median longitudinalsection. FIG. 3. Arcopsis adamsi, section throughbile ducts. FIG. 4. Arcopsis solida, abdominal sense organs, X 10. FIG. 5. Barbatia decussata,dorsal view, X 2. FIG. 6. Arcopsis solida, dorsal view, X 3. FIG. 7. Arcopsis solida, lefthalf of stomach. FIG. 8. Barbatia reticulata, reconstruction heart, dorsal view. of FIG. 9. Barbatia candida, main muscles of visceral cavity, X 5. FIG. 10. Arcopsis solida, lateral view, X 4. FIG. 11. Acar pernoides, median longitudinalsection throughanteriorend of body. a, b correspond to P1. VII, Fig. 1, and P1. VI, Fig. 7. PLATE VIII FIG. 1. Arcopsis adamsi, section throughheart; i, intestine. FIG. 2. Arcopsis adamsi, reconstruction digestive system; a, indicates position of Fig. 3, P1. VII; of b, that of Fig. 14, P1. VIII. of FIG. 3. Bathyarcapectunculoides, reconstruction stomach and bile ducts, ventral view. FIG. 4. Bathyarcapectunculoides, cross section throughpalp region; g, gonad; p, palp. FIG. 5. Bathyarca pectunculoides, cross section in region of rectum; d, circular disc of modifiedepithelium,possibly glandular; f,mantle fold shown in Fig. 15; r, rectum. of FIG. 6. Arcopsis adamsi, reconstruction heart and sub-intestinalloop. cross section throughanteriorend of foot. FIG. 7. Bathyarcapectunculoides, FIG. 8. Arcopsis solida, lateral view of digestivesystem. FIG. 9. Arcopsis solida, cross section throughbyssus region.

316

TRANSACTIONS

OF THE AMERICAN PHILOSOPHICAL

SOCIETY

FIG. 10. Bathyarcapectunculoides, cross section throughbyssus representedby heavy irregularline; i, intestine;r, gonad. FIG. 11. Bathyarcapectunculoides, reconstruction digestivesystem. of FIG. 12. Bathyarcapectunculoides, cross section throughabdominal sense organ, s, and r, rectum. FIG. 13. Arcopsis adamsi, externallayer of mantle, showingglandular development. FIG. 14. Arcopsis adamsi, cross section of stomach and bile ducts, correspondingto line b, Fig. 2, P1. VIII. FIG. 15. Bathyarcapectunculoides, ventral view of posteriorend of body. PLATE IX

FIG. 1. Bathyarcapectunctuloides, section throughmiddle of oesophagus. FIG. 2. Bathyarcapectunctuloides, median longitudinalsection throughbody. FIG. 3. Bathyarca pectunculoides, cross section of pedal furrowbetween Figs. 7 and 10, PI. VIII; i, intestine;s, gonad. FIG. 4. Bathyarcapectunctuloides, cross section near posteriorend of body. FIG. 5. Trisodostortuosa, reconstruction digestivesystem,righthand face. of FIG. 6. Bathyarcapectunculoides, lateral view, X 8. FIG. 7. Trisodostortuosa, lateral view, X 1.3. FIG. 8. Trisodostortuosa, cross section near middle of stomach. b, dorsal aorta; c, liver; d, bile ducts; g, gonad.
PLATE X

FIG. 1. Trisodos tortuosa,reconstructionof digestive system; letters indicate position of sections shown in other figures. FIG. 2. Trisodostortuosa, cross section of stomach along line c, Fig. 1. FIG. 3. Trisodostortuosa, cross section of oral opening,m, and palps, p. FIG. 4. Trisodostortuosa, cross section of oesophagus along line b, Fig. 1. FIG. 5. Trisodostortuosa, cross section of palps. FIG. 6. Bathyarcapectunculoides, cross section of heart; i, intestine. FIG. 7. Bathyarcapectunculoides, cross section of stomach and bile ducts. FIG. 8. Trisodos tortuosa, cross section at level of external urogenital opening; a, auricle; c, pericardium; r, kidney; s, blood sinus. FIG. 9. Trisodos tortuosa,firstof three successive cross sections through bile ducts along line d, Fig. 1, P1. X. FIG. 10. Trisodostortuosa, section throughbile ducts immediatelybehind Fig. 9. FIG. 11. Trisodos tortuosa,cross section through kidney slightly anterior to Fig. 8; a, auricle; c, pericardium;g, gonoduct; n, cerebro-pedalconnective; r, kidney; s, blood sinus. FIG. 12. Trisodostortuosa, cross section of stomach immediatelybehind Fig. 10, P1. X. FIG. 13. Trisodostortuosa, dorsal view, X 1.3.
PLATE XI

FIG. 1. Trisodostortuosa, ventral view of rectumand abdominal sense organs. FIG. 2. Anadara auriculata,lateral view, X 1.3. FIG. 3. Trisodostortuosa, cross section of main portionof kidney. FIG. 4. Anadara auriculata,left half of digestivesystem, X 4. FIG. 5. Trisodostortuosa, ventral view of byssus cavity, X 4. FIG. 6. Trisodos tortuosa, cross section throughstomach and bile ducts along line e, P1. 10, Fig. 1. FIG. 7. Anadara auriculata,righthalf of digestivesystem, X 4. FIG. 8. Anadara concinna,dorsal view, X 2. FIG. 9. Anadara auriculata,dorsal view, X 2. FIG. 10. Trisodostortuosa, the gut; auriculo-ventricular cross section throughventriclesurrounding valves present; cerebro-visceral connectives,gonoductsand foot shown ventral to heart. FIG. 11. Anadara concinna,byssus cavity, X 4. FIG. 12. Anadara granosa,main muscles of visceral cavity, X 3.

HEATH: ANATOMY OF PELECYPOD

PLATE XII
FIG. FIG. FIG. FIG. FIG. FIG. FIG. FIG. FIG. FIG. FIG.

FAMILY ARCIDAE

317

1. 2. 3. 4. 5. 6. 7. 8. 9. 10. 11.

Anadara concinna,lateral view, X 2.5. Anadara concinna,muscles of visceral cavity, X 5. Anadara auriculata, muscles of visceral cavity, X 5. Anadara auriculata,intestinalloops, leftside. Anadara bisenensis,rectumand abdominal sense organs, X 4. Anadara bisenensis,muscles of visceral cavity. Anadara auriculata, rectumand abdominal sense organs, X 4. Anadara bisenensis, intestinalloops, bile ducts, b, and muscle, m, attached to gastric wall. Anadara bisenensis, byssus cavity, X 5. Anadara granosa,left half of stomach, X 4. Barbatia gradata,byssus cavity, rightside, X 4.
PLATE XIII

1. FIG. 2. FIG. 3. FIG. 4. FIG. 5. FIG. 6. FIG. 7. FIG. 8. FIG. 9. FIG. 10.
FIG.

Anadara granosa,lateral view, X 2. Anadara granosa,digestive systemviewed fromleft side, X 4. Anadara concinna,digestivesystemviewed fromleftside, X 5. Anadara concinna,rectumand abdominal sense organs, X 4. Anadara bisenensis,digestivesystem, X 3. Anadara bisenensis,lefthalf of stomach, X 3. Anadara transversa, dorsal view, X 3. Cunearca perlabiata,digestivesystem, X 2. Anadara gradata,length,22 mm. Anadara bisenensis,abdominal sense organs, X 3.
PLATE XIV Anadara transversa, 2. X Barbatia gradata,length,24 mm. Anadara bisenensis,X 1.5. Anadara perlabiata, X 1. Anadara transversa, abdominal sense organs, X 6. Anadara perlabiata,abdominal sense organs, X 4. Anadara bisenensis,X 1. Barbatia reticulata, cross section throughanteriorportionof foot. Anadara transversa, main muscles of visceral cavity, X 4. Barbatia reticulata, section throughbyssus cavity; i, intestine. Anadara perlabiata, X 2. PLATE XV

1. FIG. 2. FIG. 3. FIG. 4. FIG. 5. FIG. 6. FIG. 7. FIG. 8. FIG. 9. FIG. 10. FIG. 11.
FIG. FIG.

1. T'risodos tortuosa, ventral view of heart, reconstruction. FIG. 2. Arcopsis adamsi, ventral view of heart and sub-intestinal loop; a reconstruction. FIG. 3. Anadara perlabiata,dorsal view of heart and aortae, a reconstruction. FIG. 4. Anadara perlabiata,cross section of oesophagus. FIG. 5. Anadara transversa, lateral view of digestivesystem. FIG. 6. Barbatia barbata,intestineand righthalf of stomach, X 5. FIG. 7. Anadara chemnitzi, intestineand righthalf of stomach. FIG. 8. Anadara perlabiata,section throughpalp. FIG. 9. Barbatia reticulata,diagram of kidney and connections; e, external opening; p, reno-pericardial canal; r, opening of kidneyinto common chamber with gonoduct; s, gonad and gonoduct. Ventral view. FIG. 10. Cucullaea granulosa,ventral view of heart; i, intestine. FIG. 11. Anadara chemnitzi, abdominal sense organs, X 8. FIG. 12. Barbatia barbata,left half of stomach, X 5. Fig. 13. Barbatia gradata,righthalf of stomach, X 10.

318

TRANSACTIONS
FIG. FIG. FIG. FIG. FIG. FIG.

OF THE AMERICAN PHILOSOPHICAL

PLATE XVI

SOCIETY

1. 2. 3. 4. 5. 6.

Anadara incongrua,intestineand righthalf of stomach, X 3. Anadara incongrua, actual length,60 mm. Anadara incongrua, main muscles of visceral cavity, X 3. Cucullaea granulosa,actual length,70 mm. Cunearca incongrua, liver folliclesand capillary net; connectivetissue omitted. Anadara chemnitzi, main muscles of visceral cavity, X 3.
PLATE XVII

FIG. 1. Anadara perlabiata,main muscles of visceral cavity, X 3. FIG. 2. Anadara chemnitzi, dorsal view, X 2. abdominal sense organs. FIG. 3. Anadara incongrua, FIG. 4. Barbatia barbata,main muscles of visceral cavity, X 6. FIG. 5. Anadara chemnitzi, 2. X FIG. 6. Barbatia gradata,lefthalf of stomach, X 10. FIG. 7. Noetia ponderosa,X 2. Anterior end below. FIG. 8. Argina pexata, lefthalf of stomach, X 3. FIG. 9. Anadara incongrua, lefthalf of stomach. FIG. 10. Cucullaea granulosa,section throughbyssus cavity at level of line a, P1. XIX, Fig. 5, X 5. FIG. 11. Noetia ponderosa,right half of digestive system, X 5. Reversed with referenceto P1. XVIII, Fig. 2.
PLATE XVIII
FIG. FIG.

PI. XVII.
FIG. FIG. FIG. FIG.

1. Noetia ponderosa, length,63 mm. 2. Noetia ponderosa,left half of digestive system, X 5. 3. 4. 5. 6.

Reversed with referenceto Fig. 11,

Argina pexata, X 1.5. Argina pexata, intestineand righthalf of stomach, X 3. Argina pexata, anal region, X 3. Argina pexata, main muscles of visceral cavity, p, pericardium,X 3.
PLATE XIX

1. FIG. 2. FIG. 3. FIG. 4. FIG. 5. FIG. 6. anteriorly,X FIG. 7. FIG. 8. FIG. 9.

FIG.

Argina pexata, heart, dorsal view, X 2. Noetia ponderosa,abdominal sense organs, X 5. Argina pexata, intestine,leftface. Cucullaea granulosa,dorsal view of heart, X 5. Cucullaea granulosa,lateral view of byssus cavity, X 2. Cucullaea granulosa,righthalf of stomach, a, line of section, P1. XVII, Fig. 10; arrow points 4. intestineand righthalf of stomach; shell length,27 mm. Glycymeris longior, intestineand righthalf of stomach. Glycymeris subobsoleta, Glycymeris migueliana,digestivesystem,leftface; lengthof shell 17 mm.
PLATE XX

FIG. FIG. FIG. FIG. FIG. FIG. FIG. FIG.

1. 2. 3. 4. 5. 6. 7. 8.

FIG. 9.

Glycymeris americana,lateral view of digestivesystem. mantle epithelium,outer surfaceto left. Barbatia reticulata, muscles of visceral cavity, X 3. Glycymeris longior, Glycymeris longior,X 5. Glycymeris longior,ventral view of anal region, X 6. Glycymeris americana,lefthalf of stomach. Glycymeris longior,X 2.5. Glycymeris migueliana,righthalf of stomach, X 10. Cucullaea granulosa,anal region, X 4.

HEATH: ANATOMY OF PELECYPOD

FIG.

FAMILY ARCIDAE

319

10. Glycymeris migueliana,anal region, X 4. FIG. 11. Glycymeris migueliana,lefthalf of stomach, X 10. FIG. 12. Cucullaea granutlosa, muscles of visceral cavity, X 2. PLATE XXI americana, X 1. Glycymeris main muscles of visceral cavity slanted forsake of clearness, X 6. Glycymeris subobsoleta, Glycymeris americana,main muscles of visceral cavity; p, pericardial cavity, X 2. lateral view of stomach with attached muscles and bile ducts (b), X 2. Glycymeris subobsoleta, Glycymeris americana,intestineand righthalf of stomach. Glycymeris americana,abdominal sense organs, X 4. lefthalf of stomach. Glycymeris longior, Cucullaca granulosa,lateral view of digestivesystem, X 2 Noetia ponderosa,muscles of visceral cavity, X 3.

FIG. 1. FIG. 2. FIG. 3. FIG. 4. FIG. 5. FIG. 6. FIG. 7. FIG. 8. FIG. 9.

PLATE XXII FIG. 1. Arca imbricata. Lateral view of righthalf of anteriorend of body, X 6. FIG. 2. Cross section of stomach of Acar gradata. FIG. 3. Cross section of stomach of Acar pernoidesat about the same level as Fig. 2. FIG. 4. Anal regionof Arca imbricata,X 5. FIG. 5. Reconstructicnof circumesophagealnerve ring of Acar pernoides. FIG. 6. Reconstructionof digestive system of Acar pernoidesviewed fromleft side; a, b lines corresponding respectivelyto Figs. 9, 10 on this plate. FIG. 7. Reconstructionof digestivesystemof Acar gradata viewed fromleftside. FIG. 8. Cross section of the most dorsal pair of esophageal glands. FIG. 9. Cross section of Acar pernoidesthroughmost ventral pair of esophageal glands and cerebral ganglia; along line a, Fig. 6. FIG. 10. Cross section of Acar pernoidesthroughmost dorsal pair of esophageal glands and pedal ganglia; along line b, Fig. 6.

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