Biology of the Antarctic Seas XXI PDF
Preview Biology of the Antarctic Seas XXI
Biology of the Antarctic Seas XXI Physical Sciences ANTARCTIC OCEANOLOGY Joseph L. Reid, Editor ANTARCTIC OCEANOLOGY II: THE AUSTRALIAN- NEW ZEALAND SECTOR Dennis E. Hayes, Editor ANTARCTIC SNOW AND ICE STUDIES Malcolm Mellor, Editor ANTARCTIC SNOW AND ICE STUDIES II A. P. Crary, Editor ANTARCTIC SOILS AND SOIL FORMING PROCESSES J. C. F. Tedrow, Editor DRY VALLEY DRILLING PROJECT L. D. McGinnis, Editor GEOLOGICAL INVESTIGATIONS IN NORTHERN VICTORIA LAND Edmund Stump, Editor GEOLOGY AND PALEONTOLOGY OF THE ANTARCTIC Jarvis B. Hadley, Editor GEOLOGY OF THE CENTRAL TRANSANTARCTIC MOUNTAINS Mort D. Turner and John F. Splettstoesser, Editors GEOMAGNETISM AND AERONOMY A. H. Waynick, Editor METEOROLOGICASLT UDIESA T PLATEAU STATION, ANTARCTICA Joost A. Businger, Editor OCEANOLOGYO F THE ANTARCTIC CONTINENTAL SHELF Stanley S. Jacobs,E ditor STUDIES IN ANTARCTIC METEOROLOGY Morton J. Rubin, Editor UPPER ATMOSPHERE RESEARCH IN ANTARCTICA L. J. Lanzerotti and C.G. Park, Editors THE ROSS ICE SHELF: GLACIOLOGY AND GEOPHYSICS C. R. Bentley and D. E. Hayes, Editors VOLCANOESO F THE ANTARCTICP LATE AND SOUTHERN OCEANS W. E. LeMasurier and J. T. Thomson, Editors MINERAL RESOURCES POTENTIAL OF ANTARCTICA John F. Splettstoessear nd Gisela A.M. Dreschhoff, Editors ANTARCTIC American Geophysical Union RESEARCH SERIES Biological and Life Sciences ANTARCTIC TERRESTRIAL BIOLOGY George A. Llano, Editor TERRESTRIAL BIOLOGY II BIOLOGY OF THE ANTARCTIC SEAS Bruce Parker, Editor Milton O. Lee, Editor TERRESTRIAL BIOLOGY III BIOLOGY OF THE ANTARCTIC SEAS II Bruce Parker, Editor George A. Llano, Editor BIOLOGY OF THE ANTARCTIC SEAS III George A. Llano and Waldo L. Schmitt, Editors ANTARCTIC ASCIDIACEA BIOLOGY OF THE ANTARCTIC SEAS IV Patricia Kott George A. Llano and I. Eugene Wa]]en, Editors ANTARCTIC BIRD STUDIES BIOLOGY OF THE ANTARCTIC SEAS V Oliver L. Austin, Jr., Editor David L. Pawson, Editor ANTARCTIC PINNIPEDIA BIOLOGY OF THE ANTARCTIC SEAS VI William Henry Burt, Editor David L. Pawson, Editor ANTARCTIC CIRRIPEDIA BIOLOGY OF THE ANTARCTIC SEAS VII William A. Newman and Arnold Ross David L. Pawson, Editor BIRDS OF THE ANTARCTIC AND SUB-ANTARCTIC BIOLOGY OF THE ANTARCTIC SEAS VIII George E. Watson David L. Pawson and Louis S. Kornicker, Editors ENTOMOLOGY OF ANTARCTICA BIOLOGY OF THE ANTARCTIC SEAS IX J. Linsley Gressitt, Editor Louis S. Kornicker, Editor HUMAN ADAPTABILITY TO ANTARCTIC CONDITIONS BIOLOGY OF THE ANTARCTIC SEAS X E. K. Eric Gunderson, Editor Louis S. Kornicker, Editor POLYCHAETA ERRANTIA OF ANTARCTICA BIOLOGY OF THE ANTARCTIC SEAS XI Olga Hartman Louis. S. Kornicker, Editor POLYCHAETA MYZOSTOMIDAE AND SEDENTIARIA OF BIOLOGY OF THE ANTARCTIC SEAS XII ANTARCTICA David L. Pawson, Editor Olga Hartman BIOLOGY OF THE ANTARCTIC SEAS XIII RECENT ANTARCTIC AND SUBANTARCTIC BRACHIOPODS Louis S. Kornicker, Editor Merrill W. Foster BIOLOGY OF THE ANTARCTIC SEAS XIV CONTRIBUTIONS TO ANTARCTIC RESEARCH I Louis S. Kornicker, Editor BIOLOGY OF THE ANTARCTIC SEAS XV Louis S. Kornicker, Editor BIOLOGY OF THE ANTARCTIC SEAS XVI Louis S. Kornicker, Editor BIOLOGY OF THE ANTARCTIC SEAS XVII Louis S. Kornicker, Editor BIOLOGY OF THE ANTARCTIC SEAS XVIII Louis S. Kornicker, Editor BIOLOGY OF THE ANTARCTIC SEAS XIX Louis S. Kornicker, Editor BIOLOGY OF THE ANTARCTIC SEAS XX Louis S. Kornicker, Editor ANTARCTIC Volume 52 RESEARCH SERIES Biology of the Antarctic Seas XXI Louis S. Kornicker, Editor (cid:127) AmericaGn eophysicUanl ion Washington, D.C. 1990 ANTARCTIC Volume 52 RESEARCH SERIES BIOLOGY OF THE ANTARCTIC SEAS XXI LOUIS S. KORNICKER, Editor Published under the aegis of the Board of Associate Editors, Antarctic Research Series Charles R. Bentley, Chairman Samuel C. Colbeck, David H. Elliot, E. Imre Friedmann, Dennis E. Hayes, Louis S. Kornicker, John Meriwether, and Charles R. Stearns Library of Congress Catalog Card Number: 86-647920 The Library of Congress has cataloged this serial title as follows: Biology of the Antarctic seas.-- --.Washington, D.C.: American Geophysical Union, v.: ill.; 28 cm. -- (Antarctic research series) ( :Publica- tion / National Research Council) ( : Publication / National Academy of Sciences) Began in 1964. Description based on: 11, paper 3; title from cover. Publisher's bound v. processeda fter Dec. 31, 1985, v. and parts of v. processedb efore Jan. 1, 1986 cataloged separately in LC. Vols. within the serial are issued either as complete publisher's bound v. or in unbound numbered parts (called paper or pers) within a v. 1. Marine biology--Antarctic regions--Collected works. I. American Geophysical Union. II. Series. III. Series: Publica- tion (National Research Council (U.S.)) IV. Series: Publication (National Academy of Sciences (U.S.)) QH95.58.B56 574.92'9 86-647920 AACR2 MARC-S ISBN 0-87590-761-X ISSN 0066-4634 Copyright 1990 by the American GeophysicalU nion 2000 Florida Avenue, N.W. Washington, DC 20009 Figures, tables, and short excerptsm ay be reprinted in scientificb ooksa nd journals if the sourcei s properly cited. Authorization to photocopyi tems for internal or personal use, or the internal or personal use of specificc lients, is granted by the American Geophysical Union for libraries and other users registered with the Copyright Clearance Center (CCC) Transactional Reporting Service, provided that the base fee of $1.00 per copy, plus $0.20 per page is paid directly to CCC, 21 CongressS t., Salem, MA 01970. 0066-4634/90/$01.00+0.20. This consent does not extend to other kinds of copying, such as copying for creating new collective works for resale. The reproduction of multiple copiesa nd the use of extracts, including figures and tables, for commercial purposesr equires specificp ermissionf rom AGU. Published by American Geophysical Union With the aid of grant DPP-85-20816 from the National Science Foundation December 21, 1990 Printed in the United States of America CONTENTS The AntarcticR esearchS eries:S tatemento f Objectives Board of Associate Editors ix The EunicemorphP olychaeteA nnelids From Antarctic and SubantarcticS eas With Addendat o the Eunicemorphoaf ArgentinaC, hile, New Zealand, Australia, and the Southern Indian Ocean Jos(cid:127) M. Orensanz AntarcticH alacaroidea(A cari): GeneraA gaue, Bradyagaue,a nd Halacarellus Ilse Bartsch 185 SomeL eeches( Hirudinea:P iscicolidaeo) f the SouthernO ceans Marvin C. Meyer and Eugene M. Burreson 219 The Antarctic Research Series: STATEMENT OF OBJECTIVES The Antarctic Research Series, an outgrowth of voted to papers in one or two of these areas. research done in the Antarctic during the Interna- Annual volumes, an innovation in 1990 to encour- tional GeophysicalY ear, was beguni n 1963w ith a age rapid publication, can contain papers in any grant from the National Science Foundation to the discipline.W hen needed, the seriesu tilizes special American Geophysical Union. It is a book series formats, such as maps. To reach the most appro- designed to serve scientists, including graduate priate community, papers approved for the Ant- students,a ctively engagedi n Antarctic or closely arctic Research Series may be published in any of related research, and others versed in the biologi- the monographso r serialsi ssuedb y the American cal or physical sciences.I t provides a continuing, Geophysical Union. authoritative medium for the presentationo f ex- Priorities for publication are set by the Board of tensive and detailed scientific research results Associate Editors. Preference is given to research from Antarctica, particularly the results of the manuscripts from projects funded by U.S. agen- United States Antarctic ResearchP rogram. cies, long manuscripts, and manuscripts that are Most Antarctic research results are, and will not readily publishablee lsewhere in journals that continue to be, publishedi n the standard discipli- reach a suitable reading audience. The series nary journals. However, the difficulty and ex- serves to emphasize the U.S. Antarctic Research pense of conducting experiments in Antarctica Program, thus performing much the same function make it prudent to publish as fully as possiblet he as the more formal expedition reports of most of methods, data, and results of Antarctic research the other countries with national Antarctic re- projectss o that the scientificc ommunityh as max- search programs. imum opportunityt o evaluate these projects and The standardso f scientific excellence expected so that full informationi s permanentlya nd readily for the series are maintained by the review criteria available. Thus the coverage of the subjects is established for the AGU publications program. expected to be more detailed and extensive than is Each paper is critically reviewed by two or more possiblei n the journal literature. expert referees. A member of the Board of Asso- The seriesi s designedt o complementA ntarctic ciate Editors may serve as editor of a volume, or fieldwork, much of which is in cooperative,i nter- another appropriater esearcherm ay be appointed. disciplinaryp rojects. The Antarctic Research Se- The Board works with the individual editors of ries encouragesth e collectiono f paperso n specif- each volume and with the AGU staff to assure that ic geographica reasw ithin Antarctica. In addition, the objectives of the series are met, that the best many volumes focus on particular disciplines, possible papers are presented, and that publica- includingm arine biology, oceanology,m eteorolo- tion is achieved in a timely manner. gy, upper atmospherep hysics, terrestrial biology, Reseachers interested in submitting proposals geology,g laciology,h uman adaptability, engineer- for volumes or papers for consideration should ing, and environmentalp rotection. contact the AGU publication staff at 2000 Florida Topical volumes in the series normally are de- Avenue, N.W., Washington, D.C. 20009. BOARD OF ASSOCIATE EDITORS ANTARCTIC RESEARCH SERIES BIOLOGY OF THE ANTARCTIC SEAS XXI ANTARCTIC RESEARCH SERIES, VOLUME 52, PAGES 1-183 THE EUNICEMORPH POLYCHAETE ANNELIDS FROM ANTARCTIC AND SUBANTARCTIC SEAS With Addendat o theE unicemorphoa f Argentina, Chile, New Zealand, Australia, and the Southern Indian Ocean JOSl(cid:127) M. ORENSANZ Schoolo f Fisheries,U niversityo f WashingtonS, eattle,W ashington9 8195 The eunicemorppho lychaetefsr om Antarctica ndS ubantractisce asa rer eviewed,a ndn ew dataa ddingt o the knowledgeo f the eunicemorphfsr om New Zealand, Australia,C hile, and Argentinah ave been included. The systematicosf the orderE unicemorphias discusseda,n ds uggestionasr e madep ertinentt o the statusa ndd ef'mitiono f somef amilies. It is proposedto merget he Lysaretidaew ith the Lumbrineridae, and the Oenonidaew ith the Arabellidae,a nd to redefinet he Iphitimidaea s a family independenot f the Dorvilleidae. Abyssoninoen, ew genus,i s erectedf or lumbrineridsi n the "abyssorum"g roup of Lum- brineris sensul ato. The lumbrineridg enusE ranno Kinberg is revived and redefined,a nd Lumbrineris Blainville is restricted. Three new genera,M ammiphitimeP, inniphitime,a nd Palpiphitime,a re erected within the Iphitimidae. The following new speciesa re described: Leptoecia vivipara (Antarctic; = Leptoeciaa byssorursne nsuM onro, 1939), Hyalinoeciai ncubans(N ew Zealand),E unicea ustropacifica (Pacific-AntarcticR idge; = Eunicef rauenfeldi sensuH artman, 1967, (cid:127) part), Marphysa galluccii (Bounty-AntipodesP lateau),A rabella protornutans(M agellanic Province;= A. iricolor sensuM onro, 1930), Drilonereisv iborita (Chile), Pettiboneiah artmanae( Antarctic;= Dorvilleafurcata sensuH artman, 1978), Mammiphitimetr identata( MagellanicP rovince;= Ophryotrochac laparedeis ensuH artman,1 953, in part), andP inniphitimep innognatha( Antarctic;= Paractiusn otialiss ensuE hlers, 1913, in part). The diagnosiso f severals peciesis substantialleyx pandeda, ndn ew synonymieasr e presentedin many cases, largelyb asedo n the reexaminationo f collectionss tudiedb y othera uthors. CONTENTS Introduction ....................................................................... 2 Addendum3 : Additionst o the eunicemorph Generali ntroductionto the systematicasn d faunao f the ArgentineB iogeographiPc rovince.. ...1 40 phylogenyo f thee unicemorppho lychaete.s.. ...............3 Addendum4 : Note on Eunicef rauenfeldi ................1 43 Systematicos f Antarctica nd Subantarctiecu nicemorph Biogeographicco nsiderations.. ....................................1..4 3 polychaetes.. ...........................................................1..5.. Supplementto theb iogeographisce ction FamilyO nuphidaeK inberg,1 865 .............................1..5 Appendix1 : Locationo f typem aterialso f Family EunicidaeS avigny,1 818 ................................6. 0 eunicemorphs peciesd escribedfo r Antarctica nd Family LumbrineridaeM almgren,1 867 .....................7. 2 Subantarctic waters .................................................... 156 Family OenonidaeK inberg, 1865. ............................1. 01 Appendix2 : Maxillary formulaer eportedf or Family DorvilleidaeC hainberlin1, 919 ....................1. 10 onuphida nde unicids peciesfr om Antarctic FamilyI phitimidaeF auchald,1 970 ..........................1 16 and Subantarctic waters ............................................. 158 Addendat o the systematics ection. ................................1 29 Appendix3 : Eunicemorphcsi tedf or the Argentine Addendum1 : Eunicemorppho lychaetefsr om B iogeographicP rovince( Uruguaya nd Argentina, Australia and New Zealand in the collections northo f 45 o $) ..........................................................1. 59 of the USARP ........................................................ 129 Appendix4 :, Stationsfo r whiche unicemorph Addendum2 : Noteso n somee unicemorph polychaetehsa veb eenr eportedin Antarctica nd polychaetefsr om CentralC hile .............................1 36 Subantarctic waters ..................................................... 161 Copyrigh1t 990b y theA mericanG eophysicaUln ion 2 BIOLOGY OF THE ANTARCTIC SEAS XXI INTRODUCTION stored in several institutions: United States National Museumo f Natural History (USNM), SmithsonianIn - General Remarks stitution (Washington, D.C.), Naturhistoriska Riksmusee(tS tockholm)Z, oologiskM useum( Copen- The Antarcticp olychaetefa unah asb eent he subject hagen), Zoologisches Museum der Humboldt- of many systematica ndd istributionasl tudiess incet he Universitat( ZMHU) (Berlin), Allan HancockF ounda- middle of the past century. Hartman's [1964, 1966] tion (Los Angeles), Museo Argenfino de Ciencias monographssu mmarizedth ek nowledgee xistingu p to Naturales( BuenosA ires), MuseoN acionald e Histofia that time. Those sourcesa re still the most important Natural (Montevideo), British Museum of Natural andc omprehensivree ferencet o Antarcticp olychaetes. History (London),Z oologischesM useumu nd Staat- Among recent additions,t he more substantiawl ere sinsfitut (Hamburg), Zoological Museum of the those by Hartman [1967a, 1978] and Averincev WroclawU niversity,Z oologicalI nstituteo f the USSR [1972]. In all, about5 0 publicationsd eal directlyw ith Academy of Sciences( Leningrad), and Centre Oc- Antarctico r Subantarctice unicemorphp olychaetes. 6anologiqudee Bretagne(B res0. Somem aterialss up- The Antarctic and Subantarctic,a s defined here, in- posedly stored at the Mus6um National d'Histoire clude the following areas: (1) the AntarcticO cean Naturelie( Paris)a re apparentlyu ncatalogueidf they southo f the Antarctic Convergence(,2 ) the Suban- exist at all and unfortunatelyw ere not availablef or tarcftc areas south of 45øS in the African and thiss tudy. Americanq uadrantsa, nd (3) the Subantarctica reas For each speciesa completel ist is given of all southo f 50øSin theP acifica ndA ustraliaqnu adrants. records,b oth those that I personallye xamineda nd In biogeographicte rms, thoseb oundariesc an be thosem adeb y previousa uthorsa nd not verifiedb y defined with referencet o the schemed evelopedb y me (referredt o as otherr ecords).A ppendix1 contains Knox [1960a] for shallow watersa nd that developed informationo n the location of type materials of by Vinogradova[ 1979] for the deeps ea: eunicemorphs peciesd escribedfo r Antarctica nd Sub- antarcticre gions. I. Shallow,s helf,a nd slopea reas. The collectionsf rom the USARP contained,i n ad- a. All the Antarctic water masses (Antarctic dition to the Antarcticm aterials,s everali nteresting Province). samplesfr om areaso ff SouthA ustralia,N ew Zealand, b. Transitional zones between Antarctic and Sub- andc entralC hile. Resultsc oncerningth esea nd some antarcticc old-temperatew ater masses( South Geor- othern on-Antarcticm aterialsa re presentedin four ad- gianP rovince). dendat o the systematics ection. c. Subantarcticzo nes( AntipodeanK, erguelenian, I haved evoteda s mucha ttentionto geographidci s- and Magellanic provinces), except their northern tribufionsa s to morphologicadl escriptionsF. or that edges. reason, already published extensive collectionso f II. Deep sea: The entireA ntarcticd eep-seare gion. widely distributedn ominals peciesw ere reexamined. Unusualo r particularlyw idespreadd istributiongse n- The text is organizedin to five sectionsA: general erallyt urnedo ut to reflecti dentificatione rrors. introductiont o the systematicso f the order, a sys- Some biogeographicth oughtsa re summarizedi n tematic study of Antarctic and Subantarctics pecies, the sectiono n biogeographicco nsiderationsI .t riedt o addendat o the systematics ection,a biogeographic move away from the traditional considerationo f discussiona, nd a supplementto the biogeographic provincialb oundariesa nd presenta n assemblagoef section. explanatorhyy pothesetsh at,a lthoughu nquestionably The systematicos f the eunicemorphp olychaetesis crude,m ight servet o focusf urther inquiries,a s well currently being reexamined by several authors. as to link biogeographica nd phylogenetic-systematic Modificationsa re expectedt o be introducedin the fu- discussions. ture,m ostlya s thek nowledgeo f fossilf ormse xpands. Generally,I found that the knowledgeo f Antarctic Since the treatmenti n virtually all textbooksa nd andS ubantarctiecu nicemorphws ase venl esss atisfac- monographsi s unsatisfactory( mostly because of toryt hanI hads upposeadt the outseot f thiss tudy. being outdated)I, presenta n introductorys ectioni n The completiono f the study raised more problems which several systematica nd phylogeneticg eneral thanc ouldb e solved. This, of courseI, seea s a posi- questionsa re reassessed. tive accomplishment. The systematics ectioni s mostlyb asedo n collec- tions obtainedb y the U.S. AntarcticR esearchP ro- Conventions Followed in the Text gram( USARP) andr eceivedt hroughth e $mithsonian OceanographiSco rtingC enter( SOSC). In addition,I Recordsa rei dentifiedb y codesm akingr eferenceto have been able to examine important collections thee xpeditioonr collectorC. odesc onsisot f a ORENSANZ: EUNICEMORPH POLYCHAETE ANNELIDS 3 of letters, symbols,a nd numbers,a s in the following mandibular structures,u sually known as Scoleco- examples: donta. A completed escriptiono f the maxillaryp arts is beyondt he scopeo f thisp aper. Kielan-Jaworowska SWAE/62 [1966], Jansoniuasn d Craig [1971], Oug [1978], and HERO715/905 Wolf [1980] give completea ccountso f the morphol- HERO712/71-2-32 ogy andn omenclatureT. ext Figures1 and 2 synopti- BERG/A callyd isplayt hem aint ypesf oundi n eachf amily. Kinberg [1865] proposed the classificationo n Charactersto the left of the slashi dentify the survey whicht he moderns ystemo f the extante unicemorphs or collector. The first four lettersi dentifyt he vessel, is based. Its subsequendte velopmenht as been sum- expedition, or collector; in the examples above, marized by Mcintosh [1910] and Hartman [1944]. HERO means 'RV Hero,' SWAE means 'Swedish Both extant and fossil forms were put into a unified AntarcticE xpedition,' and BERG refers to samples framework( mostlyb asedo n maxillarys tructuresb)y collectedb y C. Berg. If, in addition,t herea re some Kielan-Jaworowska[ 1966], who erected several fossil numbers,t hesei dentify the cruisen umber;i n the ex- familiesa nd generaa nd elaboratedo n the interpreta- ample, HERO715 means 'RV Hero, cruise 715.' tion of homologies.U nfortunatelyt,h e work of pal- Characterst o the fight of the slash identify an eontologisthsa s not receivedf rom neontologisttsh e oceanographisct ationo r a collections ite;i n the exam- attentiont hat it deserves. No systemo f the extant ples above, 62 means 'station 62,' 71-2-32 means forms shouldi gnoret he insightst hat can be gained 'station 71-2-32,' and A means 'site A.' Codes and from comparisonws ith the extinctt axa. data for stations and collection sites are summarized In this section,I summarizes omer ecentd evelop- in Appendix4 . mentsp ertinentt o our understandingo f the relation- Many characterasr e sized ependentU. nfortunately, shipsa monge unicemorphfa milies,a nd discusss ome mostp olychaetes pecimensin oceanographisca mples aspectst hat have been overlookedo r were the source are incomplete.F or thatr easonit is necessartyo util- of misinterpretationAst. thee ndI discussth e systems ize somek ind of sizei ndext hat,i deally,s houldb e in- proposed by Hartman [1944], Kielan-Jaworowska sensitivet o the degree of contractiono f preserved [1966], Kozur [1970], and Tzetlin [ 1980] and reassess specimens.I made use of a size index which is the themi n the light of thosec onsiderations. product between the length of the anterior end (measuredf rom the tip of the prostomiall obe to the posteriorb ordero f the tenth setiger)a nd the maxi- 1. Morphological GradesB ased mumd iametero f the samep art of the body( parapodia on Maxillary Structure excluded),b oth measuredi n millimeters. In a few cases,o ther size indices were used out of convenience. When this is the case,i t is explicitlym entionedin the Ehlers [1864-1868] introduced the idea that text. eunicemorpmh axillaec ouldb e groupedin to architec- tural types( grades). He recognizedtw o types,l abid- ognatha nd prionognath.T his typologyw as revived GENERAL INTRODUCTION TO THE by Hartman [1944] and expandedb y Kielan-Jawo- SYSTEMATICS AND PHYLOGENY OF THE rowska [1966], who added two more grades, the EUNICEMORPH POLYCHAETES ctenognath(i nto which the living dorvilleidsf all) and the placognath( composedo nly of fossil forms). The extante unicemorphp olychaetesc onstituteo ne Mierzejewskia nd Mierzejewska[ 1975] describedA r- of the best defined and most homogeneous chaeoprionq uadricristatusf rom the Ordovician of suprafamiliala ssemblageso f the class Polychaeta. Polanda nd ascribedit to a new grade,t he xenognath The mostc onspicuouusn ifyingc haracteris a complex type. Interestinglyt,h is grader esemblesth e hypo- maxillomandibulaprh aryngeaal rmature. As pointed theticals tructureo f primitive eunicemorphja ws as out by Purschke[ 1987] for Ophryotrocha,t he ele- proposedin the speculationos f other authors. Thus, mentso f the maxillarya pparatusa re not 'free': "the five basic eunicemorphg radesa re currentlyd istin- apparatusis built up of stripsl ying on the maxillary guished.B oundariesb etweent hema re sometimesu n- folds, and the individual plates are folded areas of clear( kalloprionidsf,o r example,a re saidt o be transi- these strips." Although varying from family to tionalb etweenla bidognatahn dp rionognatthy pes). It family, it is composedo f elementsw hoseh omologies mustb e stressedth at gradess houldn ot be equated can be traced across most of the genera. One with a systematihc ierarchys, incet hey do not neces- transcendentafal ct in eunicemorphs ystematiciss the sarily imply considerationosf monophylya nd com- existenceo f fossil remainso f these maxillary and mon
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