MemoirsofMuseumVictoria63(1): 1-5(2006) iSSN 1447-2546(Print) 1447-2554(On-iine) http://www.museum.vic.gov.au/memoirs/index.asp A new species ofNeamia (Perciformes: Apogonidae) from the West Pacific Ocean Thomas H. Fraser^andGeraldR. Allen^ 'MoteMarineLaboratory, 1600KenThompsonParkway,Sarasota,Florida,34236USA([email protected]) ^DepartmentofAquaticVertebrates,WesternAustralianMuseum,FrancesSt.,Perth,WesternAustralia6000,Australia Abstract FraserT.H. andAllenG.R. 2006.Anew species ofNeamia(Perciformes: Apogonidae)fromtheWestPacificOcean. MemoirsofMuseumVictoria63(1): 1-5. A new species of apogonid fish, Neamia articycla, is described from Australia, Indonesia, Philippines and Fiji, bringingthetotalnumberofdescribedNeamiaspptothree.ItisdistinguishedfromNeamianotulainpreservationby: a circular, ocellateddarkspotratherthan onethatis oval-shapedandflattenedalong its anteriormargin, with anouter, narrow darkish edge clockwise dorsallyfrom abutting edge ofpreopercle aroundto meeting ventral abutting edge of preopercle,afaint,narrow,horizontallineofmelanophoresbelowtheeyereachingthepreopercleridge;inlifenodarkish marksbehindtheeyes,acircularspotwithpaleouterbandwithoutdarkedginginsteadofanovalspotflattenedalong preopercleedgewithadarkishedge;fivepredorsalscalesinsteadoffour;usually22pored,lateral-linescalesinsteadof 23 andalongerpectoralfin.NeamiaarticyclacanbedistinguishedfromNeamiaoctospinabyhaving 14pectoralrays insteadof17-21 andbyhavingsevenvisiblefirst-dorsalspinesinsteadofeight. Keywords Apogonidae,Neamiaarticycla,Neamianotula,cardinalfish,newspeciesApogonichthys,Foa,Fowleria. Introduction perforations (referred to as minipores) of the skin over the canal systems ofthehead. Thelatterare sometimes obscured Ichthyologistscollectingfrompoorlysampledhabitatsindepths of 10^0 m continue to find new species of cardinalfish. We bycongealedmucous. describehereanewNeamiawhichisassociatedwithHalimeda NeamiaSmithandRadcliffeinRadcliffe, 1912 bedsgrowingindeepercoralreefsystemsofthewesternPacific Ocean. The new species represents thethird known species in Typespecies.NeamiaoctospinaSmithandRadcliffeinRadcliffe, thisclade.Theremaybeothermaterialinmuseumsidentifiedas 1912. Fowleriabecauseofthesimilarphysiognomy.Neamiasppcan Diagnosis.Anapogoninewith3hypurals, 1 and2fused,3and4 bedistinguishedfromsimilarFowleriaspeciesbythefollowing fused,5free,hypurals3-1-4fusedtourostylarcentrum; 1 pairof characters:fusedhypuralsinthecaudal-finskeleton,completely reduced or slender uroneurals; 3 epurals; 3 supraneurals; 2 pored lateral-line scales in adults (versus partially pored and supernumerary dorsal-fin spines; smooth preopercle edges and then pitted, except one undescribed species of Fowleria), ridge, smooth post-temporal; no basisphenoid; a reduced, tiny cycloid scales on the head and anterior portions of the body supramaxilla; no palatine teeth; scales on cheek and opercle (weakly ctenoid scales on the nape for species of Fowleria, cycloid;lateral-linescalesextendingfrompost-temporaltobase elsewhere on the head, weak to strongly ctenoid scales and ofcaudal fin, all ctenoidwithsimplepores;roundedcaudal fin; ctenoidonentirebody). 9dorsal-finspines,8thspinevisibleorhiddenbyskin,dorsalfin Methods for counts and measurements were described in deeply notched and considered separate, the 9th spine at mdetmail by Fraserand Lachner, 1985. All measurements are in beginning of 2nd dorsal fin; dorsal-fin rays 9; anal-fin rays 8; to the nearest 0.1. Percentages are of standard length. pectoral-finrays 13-21;palestomach,intestineandperitoneum. InstitutionalacronymsfollowEschmeyer, 1998andLevitonet ah, 1985. Field station numbers are listed for additional Neamiaarticyclasp.nov. collectioninformation,forexample,VGS69-23.Radiographs (X-rays)havebeentakenfromtypematerial.Acameralucida Figures 1-4 attachment on a Wild microscope and a needle blowing air Holotype. AMS 1.25121-005; 1, 35.5; Australia, Great Barrier were used to make the diagrammatic drawings ofthe pores. Reef,Qld, 14°41'S, 145°30'E,eastofLizardL;31 Jan 1982;36-38m, The description of pores refers to larger and to smaller J.Leis. 2 ThomasH.FraserandGerald R.Allen FUmiSmguNrSMeL13.(7pN0re2ea9se1mr,ivae2d3)a.,r5t,Aiucmsytcmlraa:lSiLAa,,.PQHhloidll,oitpeypapisente,so,AfMMLiiSznad1ra.dn25aL.1o2B2L.1,-P0ma0or5da,itf3yi5pe.e5d, Fhmsloiailgngoudhtrityelbpyue2l,.tairlCNteeepdpoa.hrmaei.Cl.iaec.Vapleroantsttitercreayarclliliosarv.isaeryAtws.itceouDmlfoarrodsfepanolptraoer.rvyei-fse.awrp.otoinscBtut.-lhtaeerLm,ahpteoedarr.daallvoepfnvotirtreeha.wle fromcolourimageofafreshspecimenbyJ.T.Williams.C.Paratype, USNM 209665, 32.4 mm SL (preserved), Indonesia, Moluccas, Ceram 1. D. Paratype CAS 223500, 32.9 mm SL (preserved), Fiji, VitiLevu,Nananui-i-cake1. Figure3.Post-ocularmelanophorepatternsonthecheek(faintline) andcircularspotwithpaleintervalandouteredgingofmelanophores startingalongposterioredgeofpreopercle(POP).USNM209665 paratype,32.4mmSL(preserved). AnewspeciesofNeamia(Perciformes:Apogonidae)fromtheWestPacificOcean 3 Paratypes.PacificOcean:Indonesia:USNM209665;8, 14.2-32.3; teethonvomer;noneonpalatine,ectopterygoid,endopterygoid Ceram,PirnBay;VGS73-6;10Jan1973;8m.USNM209929;5,22.5- orbasihyal. 27.7;Saparua;VGS73-14; 18Jan1973;9m;X-ray.Fiji:CAS223500; Vertebrae 10-1-14. 3 hypurals 1-1-2 fused, 3-1-4 fused to 71116,.4;4530°2631.62955.;-3M001a'07Er4°,;192b.0e410,6t22w;'3eS4e,.1n3-;13L7i8m&z°,a1r4dA.c2oMI5l.0oS&'uEr,Y1to.Nr4ana0nng6sae6pna5uRr-iee0-en0ifc5-,y;c.aGkrA1eeu,saLtt,r1a7BV.lai0irt;ari,iLe1Qre4lv°Rdu3e:;8e.fA2G;00M'22SS-6, pueraporusrhtayylpluarcraoluc.ledn3tnrosuutmp,rbae1neipudareianrltsiu,frioe2ndesufrurapolemsr,ntuh2meelrraaradgreiyogesrppauiprnhae)ls,s oa(anf3r1resdet Jan2001;27m.AMS1.40666-012;1,31.1&AMS1.40666-013;1,19.0; dorsal pterygiophore, 8thdorsal-fin spinehiddenas anubbin. 14°38.16'S, 145°32.16'E,betweenLizardI. &outerbarrierreef. Great Basisphenoidstatusunknown,lowerportionapparentlyabsent, Barrier Reef; 29 Jan 2001; 25 m, colour digital photos. Philippines: but unclear on X-ray. Status of suspensory pharyngeal USNM370291; 1,23.5; 13°10.04'N, 120°35'12'E,MindanaoL;MinOO- unknown. Supramaxilla present. Posttemporal smooth on 62;3Jun2000;30m;X-ray,colourdigitalphotos. posteriormargin.Preoperclesmoothonverticalandhorizontal Non-type material. Indonesia: USNM 209847; 3, 10.0-20.4; Ceram, margins. Infraorbitaledges smooth. Pirn Bay; VGS 73-5; 9 Jan 1973; 5-7 m, WAM P.32771-002; 23.4; Cephalic pored sensory system (fig. 2) - all larger pores 01°36.252'S, 135°24.553'E,TanjungWoka,YapenL,Papua,Indonesia; bilateral.Novariationforthefollowingpores: 2nasalporesat 13Feb2006;34m.Philippines:USNM369983;2,18.0-27.4; 13°45'N, anteriorendofsupraorbital canal system, 1 medial totubular 120°55'E,Anilao, Batangas; 26Apr 1980; 22 m. USNM 268318; 1, anteriornare, 1 slightlyanteriorandmedialtoposteriornare, 32.6;9°04'38'N, 123°16'44'E,ApoL;SP78-36;7Jun1978;39m. 1 near edge of orbit on interorbital; 4 infraorbital pores at Other material. Fowleria sp. AMS 1.23585-001; 1, 45.0; 14°34'S, anterior part ofinfraorbital canal, 1 between ventral edge of 145°34'E,south-westofCarterReef,GreatBarrierReef,Australia;31 lachrymal and flat posterior nare, 2 on ventral edge of Jan1982;30m,J.Leis.ApogonichthysnafaeHolotype,USNM62947; lachrymal, 1 onventral edgeof2ndinfraorbital; dentarywith 22.1;Japan,Okinawa,Naha. 3 pores atanteriorend ofmandibularcanal, 1 ontip oflip, 1 Diagnosis. A species of Neamia with 7 visible first-dorsal behind lip, and 1 lateral; 7 articulo-preopercular pores near spines, 8th spine hidden by skin, darkish circular spot on the posterior edge along the preopercular canal. Variation in operclebelowopercularspinewithinapalearea; inlife,body numberandlocationofminipores: supraorbitalwithnoneor1 uniformly reddish to brownish, all fins pale, 13-14 pectoral porenearedgeoforbitoninterorbitalarea;pairedporesalong rays, 5 predorsal scales, eye diameter 10-12%, pectoral fin upperandlowerportions ofinfraorbitalsbelow orbit; dentary 25-31%andbodydepth37-40%ofstandardlength. andarticularwith4pairs(innerandouter) ofposteriorpores, with several unpaired pores on dentary and articular; Description. For general body shape see fig. la-d. Holotype preopercular minipores along narrow upper portion of proportions and characteristics first with range of values in preopercle. Minipores variably present posterior to eye on parenthesesforparatypesandothermaterial.Proportionsas% upper portion ofinfraorbitals, on nape and above preopercle of standard length: greatest body depth 37.4 (34-40); head endingonthepost-temporal. length 44.6 (41-46); eye diameter 10.0 (11-12); snout length All scales cycloid on head, nape and breast. Scales 8.3 (8-10);bonyinterorbitalwidth3.7(4-6);upperjawlength becoming ctenoid above breast and behind pelvic fin. Scales 20.3 (20-22); caudal peduncle depth 15.7 (15-17); caudal above lateral line ctenoid. All lateral-line scales ctenoid, peduncle length 20.9 (19-22); 1st dorsal-fin spine length 1.7 except first. Pored lateral-line scales simple (with 1 opening (2-4); 2nd dorsal-fin spine length 6.6 (6-10); 3rd dorsal-fin aboveandbelowmainpore),extendingfrompost-temporalto spinelength 16.9(16-21);4thdorsal-finspinelength 16.0(16- baseofcaudalfin. 19); spinein2nddorsal fin9.7(9-12); 1stanal-finspinelength 2.0 (2-5); 2nd anal-fin spine length 10.0 (8-12); pectoral fin Life colours. Colours based on photographs are from fresh length28.9(25-31);pelvicfinlength25.7(24-30). dead specimens shortly after collection. From B. M. Carson- DorsalfinVII(I)-I,9,8thspinehiddenunderskin,3rdand Ewart photograph (AMS 1.40666-012) -body, pelvic, caudal 4th spines aboutthe samethickness; anal fin11,8;pectoral fin anddorsalfinsorangishred,flankgradingtowhiteonabdomen; 14;pelvicfin1,5;principalcaudalrays9-1-8;caudalfinrounded; head without marks extending from orbit edge, lips whitish; pored lateral-line scales 22 (11-12 for specimens 17-20 mm circular spot on opercle yellowish with faint outer margin of SL, 18 for specimens 20-24 mm SL, 22-23 in specimens melanophoresdorsally,largeinnercircleofdarkmelanophores greaterthan 26 mm SL); transverse scale rows above lateral aboutsizeofpupil;iriswhitishwithnarrowinnerringoflight line 2; transverse scale rows below lateral line 6; median yellowsurroundingpupil. predorsal scales 5; circumpeduncular scale rows 12 (5-I-2-I-5). FromB. M. Carson-Ewartphotograph (AMS 1.40666-013) Totalrudiments and gillrakers 15 (13-15), 2-I-1-5-I-7 (1-2-1-1-5- -body and caudal fins a pale orangish red to more yellowish 6-I-5-7), well developed gill rakers 6 (6-7), upper arch 1(1), posteriorly,withbrownishmarksincaudal fin, flankgradingto lowerarch5(5-6).Rudimentsonlowerarchasflattenedtooth white on abdomen; head without marks extending from orbit patches. 2ndarchwith 1 short, nob-like rakerin angle,upper edge,lipswhitish;circularspotonopercleyellowishwithouter arch with 1-2 flattened tooth patches, lower arch with 5-6 margin ring (^-270°) ofmelanophores except posteriorly, large nob-likerakersfollowedbyflattenedtoothpatches. innercircleofdarkmelanophoresaboutsizeofpupil;iriswhitish Villiform teeth in a band of about 5 rows becoming 2-3 withnarrowinnerringoflightyellowsurroundingpupil. rows on side of premaxilla; villiform band of about 5 rows FromD. W. Greenfield photograph (CAS 223500) -body becoming 2-3 rows on side ofdentary;3-4rows ofvilliform andcaudalfindarkbrownish,headwithoutmarkingsextending 4 ThomasH.FraserandGerald R.Allen from orbit edge, lowerhalf ofhead and breasttannish white edgeoutsideofthepaleareainlife,fourpredorsalscalesinstead extending posteriorly to opercular spot and then above offive,usually22pored,lateral-line scalesinsteadof23 anda pectoral-fin base descending to near anal base; pelvic fins longerpectoralfin.Neamiaarticyclacanbedistinguishedfrom whitish; spot on opercle whitish tan with inner darkish ring Neamiaoctospinabyhaving 14pectoralraysinsteadof17-21 smaller than eye diameter; iris tannish white with narrow andbyhavingsevenvisiblefirst-dorsal spinesinsteadofeight. innerringoflightyellow surroundingpupil. Fraser and Allen, 2001 discussed the relationships of FromJ. T. Williams’ photograph (USNM370291) -body Neamia with other apogonid genera. Neamia articycla does generallyuniformbrownish,with2rows ofsmall spots along nothavecharactersthatalterthatdiscussion.Neamiaarticycla caudal peduncle, about5pairs,3 spots extended anteriorlyin is more similar to N. notula based on the presence of the the lower row with 2 spots below the 3 spots; head without opercle spot, the reduced, hidden, eighth dorsal spine, larger markings extending from orbit edge; all fins generally pale eye andthelowerpectoral fin-ray countthantoN. octospina. with numerous small melanophores in caudal fin; round spot Specimens of N. articycla less than about 25 mm SL have onoperclewithcompletebrownishouteredge ofbodycolour incompletely pored lateral-line scales (the number of pored around pale area with large dark inner spot; narrow whitish scales varies with size) and may therefore be confused with barsonlowerlip;irisorangish. Fowleria, most species of which have an incomplete lateral line-seethefigureofFowlerianafae,Snyder, 1909inSnyder, Colour in ethyl alcohol. Holotype uniform light tannish in 1912 one of the nominal species originally described as alcoholwithtinymelanophoresonhead,body,bothdorsalfins, Apogonichthysnafae. denser on membranes of 2nd dorsal fin, melanophores absent SomesimilaritiesexistbetweenN.articyclaandspeciesof on anal, pelvic and pectoral fins; large rounded dark spot on Fowleria.Theocellus-likeopercularspot,tubularanteriornare, opercle,surroundedbypalearea,melanophoresatedgeofpale lack of palatine teeth and body shape are characteristics N. zoneforming aboutahalf-circle;faintseries ofmelanophores articyclashareswithmostspeciesofFowleria.However,these inahorizontallinefromlowerorbittopreopercularridge(fig. genera can be distinguished by fused caudal skeletal 3).Palestomach,intestineandperitoneum.Paratypesandnon- characteristics inNeamia, completely poredlateral-line scales typematerialuniformbrownishtotannishinalcoholwithtiny in adult Neamia, (number of pitted scales in N. articycla melanophores on head, body, both dorsal fins, denser on decreases posteriorly with growth, unlike Fowleria), and membranes of 2nd dorsal fin, melanophores absent on anal, cycloid scales on the head and anterior portions of the body pelvic and pectoral fins; large rounded dark spot on opercle, (weakly ctenoid scales on nape for species of Fowleria, surrounded by pale area, melanophores at edge of pale zone elsewhereonheadweakto stronglyctenoidscales andctenoid forming about a half-circle; faint, darkish, horizontal line of on body). Bergmann, 2004 noted some variation in the pore melanophoresfromlowerorbittopreopercularridge;lipswith pattern on the dentaries for the four species of Fowleria afewnarrowpalebarsorlipsindistinct. examined.NoneappearedtobesimilartoN.articyclainspatial Etymology. The Greek words artios, complete orperfect and organization,withF.vaiulaeperhapshavingafewofitsslightly cyclus,circleasarticyclaanadjectivereferringtothedarkspot largerminipores spacedandlocatedinasimilarfashion. ontheoperclewithitsedgeasacompleteorperfectcircle. IdaandMoyer, 1974, fig. 3 comparedthecephaliclateralis andfree neuromasts offive generaofcardinalfishes (including Distribution. Thenew species is apparentlywidespreadinthe three subgenera), Apogon {Ostorhinchus), Apogon (Apogon), western Pacific from Sumatra to Fiji, and eastern Australia Fowleria,Cheilodipterus,Rhabdamia(Verulux),andPseudamia. northtothePhilippines(fig.4).Althoughithasbeencollected Bergmann, 2004 provided a more extensive survey of these at relatively few locations, future collections will no doubt systemsforthese samegeneraandanother30apogonidgenera expandtheknowndistribution. Duetotheircryptichabits,the and subgenera in two subfamilies. Species of Neamia differ species ofNeamia are seldom observedby divers, ratherthey most obviously from illustrations of Fowleria, Foa, and adraetacsolulgegcetsetdNw.iatrhtiiccyhctlhayofcriedqeuse,ntdsretdhgee1s0a-n4d0smleddse.ptCholzloencet,ioinn Apogonichthys in having fewer pores on the dentary. A comparison of the descriptions of Neamia suggest that N. association with beds of a calcareous algae, Halimeda or octospinaandtheholotype ofN. articycla show similarities in calcareous rubble, sometimes mixed with the pocilloporid coralSeriatoporahystrix.TheURLhttp://www.amonline.net. thethree slit-likepores alongtheventral edgeofthelachrymal au/fishes/about/research/halimeda.htm has more information (firstinfraorbital)andsecondinfraorbital,threesuborbitalpores andthepaireddentarypores.Differenceswereapparent:location aboutotherfishes collectedin suchHalimedabeds and alink of the anterior-most supraorbital pore back near the anterior tocolourphotographsofthisspecies. nare inN. articycla, slit-like inN. octospina located along the Remarks and discussion. Neamia articycla has an ocellated margin with the premaxilla; larger pores along the posterior opercularspotthatinlifelacksanouterdarkmarginposteriorly, edge of the subopercle-preopercle in N. articycla than for N. butinpreservationhasadarkishedgefromdorsaledgeabutting octospina, and no mini-pores just posterior to the preopercle preoperclearoundtoventraledgeabuttingpreopercle.Ithasno ridge in N. articycla versus numerous minipores; few paired darkishmarks behindthe eyesinlife, butinpreservationdoes pores on the dentary and articular in N. articycla versus haveafaint,narrow,horizontallineofmelanophoresbelowthe numerous paired pores; no minipores in N. articycla versus eyereachingthepreopercleridge. Incontrast,N. notulahasan manyminiporesonsnoutandinterorbit;andfewerminiporeson oval spot flattened along the preopercle edge with a darkish thenapeinN.articyclaversuswidespreadandnumerous. AnewspeciesofNeamia(Perciformes:Apogonidae)fromtheWestPacificOcean 5 IdaandMoyer, 1974failedtoindicatethetaxonomicvalue Greenfield,J.M.Leis(takenbyB.M. Carson-Ewart)andJ.T. of pore characteristics, although their fig. 3 was suggestive. Williams provided colour images ofthe new species. Useful Bergmann, 2004 found four characteristics of the cephalic comments wereprovidedby D. W. Greenfield,J. M. Leis and lateralis which were shared by all cardinalfishes examined. ananonymousreviewer. Neamia articycla has these four characteristics: a terminal lachrymal pore, a pair ofventral lachrymal pores, an anterior References dentary pore in the lower lip and a mental (ventral dentary pore) immediately behind the lower lip. Bergmann, 2004 Bergman,L.M.R.2004.Thecephaliclateralissystemofcardinalfishes identifiedeightcharacteristicsthathadvariation,andproposed (Perciformes: Apogonidae) and its application to the taxonomy that these various states had phylogenetic value within and systematics of the family. Unpublished PhD Dissertation, cardinalfishes. Neamia articycla has: (1) the terminal end of UniversityofHawaii,i-vi+1-226,56figs, 1tab.,3app. t(2h)estuhpera-loartberiatlalmcaarnaglinendoifngthienasulparrag-eoprobirteanleacranaanlternieoarrnatrhe;e Esche9md5e.8.yeCprpa,.taWl.ogN.o,ff1i9s9h8e.s.InVtorloduumcetiIo.nC.alPipf.o1r6n-i2a2Aicn:adWe.mNy.oEfsScchimeenyceers,, posteriornarewithasinglelargepore;(3)thelateralmarginof Eraser,T.H.andG.R.Allen.2001.ANewSpeciesofCardinalfishin the supra-orbital canalmidwayoninterorbitwithalargepore, Neamia (Apogonidae, Perciformes) with a Review of Neamia but also with an additional smaller anterior pore (variably octospina.RecordsoftheWesternAustralianMuseum20(2):159- present) apparently intermediate to having multiple canal 165,2figs, 1tab. projections;(4)thepostorbitporeswithsimplecanalprojections Eraser,T. H. andE.A. Lachner. 1985.Arevisionofthecardinalfish with minipores; (5) the supratemporal canal anterior margin subgeneraPristiapogon andZoramia ofthe Indo-Pacific region smooth;(6)thesupratemporalcanalposteriormarginwithfew (Teleostei: Apogonidae). Smithsonian Contributions toZoology canal projections withminipores; (7) thelateral margin ofthe 412:1-47,20figs,4tabs. mandibular canal with relatively large and simple pores; and Ida,IHs.hiagnakdi-J.JiTm.a,MoJyaeparn.,1w9i7t4h.dAespcorgioptniiodnsfoifshaesneowfMspieycaikese.-JJaipmaaneasned (8) a terminal mandibular pore (here called the posterior JournalofIchthyology,21(3):113-128,5figs,3tabs. articular pore) near the boundary with the preopercle. The Leviton, A. E., R. H. Gibbs, Jr., E. Heal and C. E. Dawson. 1985. usefulness of these eight characteristics and their character Standards in Herpetology and Ichthyology: Part I. Standard states will befurtherdevelopedthrough examination ofmany symbolic codes for institutional resource collections in species not yet reported in the literature. Many of the other HerpetologyandIchthyology.Copeia, 1985(3):802-832. minipores had some variation in number and location among Radcliffe, L. 1912. Descriptions of fifteen new fishes ofthe family individualsofN.articycla. Cheilodipteridae from the Philippine Islands and contiguous waters. Proceedings of the United States National Museum, Acknowledgments 41(1868):431-446,pls.34-38. Snyder,J. O. 1909. Descriptions ofnew generaand species offishes WethankD.G. Smith,J.T.Williams(USNM),DavidCatania fStraotmesJNaaptainonaanldMtuheseRuimu,K1i6n(1I6s8la8n)d:s5.97P-r6o1c0e.edings ofthe United andDavidW.Greenfield(CAS),andMarkMcGrouther(AMS) Snyder,J.O. 1912.JapaneseshorefishescollectedbytheUnitedStates who aided in curatorial processes and loan of fishes. D. G. Bureau of Eisheries steamer ‘Albatross’ expedition of 1906. SmithprovidedX-rays.L.M.R.Bergmannallowedustheuse ProceedingsoftheUnitedStatesNationalMuseum,42(l909):399- of illustrations and information from her dissertation. D. W. 450,pis.51-61.