New Calmoniid Trilobites (Phacopina: Acastoidea) from the ... filela fauna de la biozona Scaphiocoelia en los departamentos de La Paz y Tarija, cierra un 1 Research Associate, Division

Copyright q American Museum of Natural History 2003 ISSN 0003-0082

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Number 3407, 17 pp., 6 figures May 22, 2003

New Calmoniid Trilobites (Phacopina: Acastoidea)from the Devonian of Bolivia

MARIA DA GLORIA PIRES DE CARVALHO,1 GREGORY D. EDGECOMBE,2

AND LEGRAND SMITH3

ABSTRACT

Four new taxa of Lower Devonian Calmoniidae from Bolivia are described: Gemelloidesdelasernai, n. gen. and sp., Eldredgeia eocryphaeus, n. sp., Wolfartaspis liebermani, n.sp., and Granadocephalus hannibali, n. gen. and sp. The new genus Gemelloides is sistertaxon to Vogesina Wolfart, 1968. Eldredgeia eocryphaeus, from the Scaphiocoelia Assem-blage Zone in La Paz and Tarija Departments, closes a stratigraphic gap/ghost lineage inthe early history of the Metacryphaeus group. Wolfartasapis liebermani (Icla Formation,Kochis, central Bolivia) predates its only congener, W. cornutus. A novel combination offeatures within Calmoniidae characterizes Granadocephalus hannibali, from the Icla For-mation in Cochabamba Department. This monotypic genus may have its closest relativesin the Calmonia group.

RESUMEN

Se describen cuatro nuevos taxa de la familia Calmoniidae del Devonico Boliviano:Gemelloides delasernai, n. gen. y sp., Eldredgeia eocryphaeus, n. sp., Wolfartaspis lie-bermani, n.sp., y Granadocephalus hannibali, n.gen. y sp. El nuevo genero Gemelloideses el grupo hermano de Vogesina Wolfart 1968. Eldredgeia eocryphaeus, registrada parala fauna de la biozona Scaphiocoelia en los departamentos de La Paz y Tarija, cierra un

1 Research Associate, Division of Paleontology, American Museum of Natural History. e-mail: [email protected] Principal Research Scientist, Australian Museum, 6 College Street, Sydney, NSW 2010, Australia.3 266 Merrimon Avenue, Asheville, NC 28801-1218.

2 NO. 3407AMERICAN MUSEUM NOVITATES

gap estratigrafico/linaje duende en la historia temprana del grupo Metacryphaeus. Tambienparte de este grupo, la especie Wolfartaspis liebermani (Formacion Icla, Kochis, Depar-tamento de Cochabamba) antecede a su unico congenere, W. cornutus. Entre los Calmon-iidae, Granadocephalus hannibali (Formacion Icla del Departamento de Cochabamba) escaracterizada por una combinacion de caracteres nueva para la familia. Este genero mo-notıpico podrıa presentar como taxa mas cercanos miembros del grupo Calmonia.

INTRODUCTION

Calmoniid trilobites from the Devonian ofBolivia have a long history of study. Sincethe initial descriptive work by d’Orbigny(1842), numerous papers have documented arich diversity in the Lower and Middle De-vonian (e.g., Kozlowski, 1923; Branisa,1965; Wolfart, 1968; Eldredge and Ormiston,1979; Eldredge and Branisa, 1980; Lieber-man et al., 1991; Lieberman, 1993). MostBolivian calmoniids occur in three forma-tions localized in distinct geographic areas(fig. 1): the Belen Formation (Belen-La Paz-Sicasica region of the West Bolivian Altipla-no); the Icla Formation (Icla-Padilla region,Sierras Subandinas of central Bolivia), andthe Gamoneda Formation (Tarija region,southern Bolivia). The basal portion of allthree formations is considered to be time-equivalent, falling within the ‘‘Scaphiocoe-lia-bearing beds’’ (Isaacson, 1977) recog-nized as the Scaphiocoelia Assemblage Zoneby Eldredge and Branisa (1980). Trilobitesoccur both within and above this level. Thetotal stratigraphic range of Calmoniidae inBolivia is Late Silurian (Prıdolı) (Edgecombeand Fortey, 2000) to Middle Devonian (Giv-etian).

It is now well established that calmoniidtrilobites are endemic to a Southern-hemi-sphere biogeographic region that Clarke(1913) first characterized as an ‘‘austral fau-na’’, which he was able to distinguish froma ‘‘meridional fauna’’. Richter and Richter(1942) observed the endemic character ofthis fauna and coined the term ‘‘Malvinokaf-fric Province’’, which corresponds generallyto the Malvinokaffric Realm of Eldredge andOrmiston (1979).

Smith and Edgecombe (1996) informallyreported two new genera and four new spe-cies of Calmoniidae from Bolivia, but did notname or describe them. In this paper we pre-sent a systematic description of this material,

which contributes to the taxonomic andstratigraphic record of Bolivian calmoniids.Morphological terminology follows that usedin the Treatise on Invertebrate Paleontology,Part O (Whittington and Kelly, 1997) as wellas the terminology of Eldredge and Branisa(1980). The term Large Eye Index was intro-duced by Wolfart (1968), calculated as theratio between the exsagittal length of the eyeand the sagittal length of the glabella exclud-ing S0. The chronostratigraphic scheme forthe Devonian of Bolivia is as summarized byAdrain and Edgecombe (1996: fig. 2), afterBlieck et al. (1996). Specimens studied andfigured here are deposited in the collectionsof American Museum of Natural History(AMNH), Division of Paleontology (Inver-tebrates), and Museo de Historia Natural deCochabamba (MHNC), Bolivia.

SYSTEMATIC PALEONTOLOGY

FAMILY CALMONIIDAE DELO, 1935

GEMELLOIDES, NEW GENUS

DERIVATION OF NAME: Literally, ‘‘like Ge-mellus’’, in reference to the name Dalmanitesgemellus Clarke, 1890; gemellus (Latin), atwin.

TYPE SPECIES: Gemelloides delasernai, n.gen. and sp.

REFERRED SPECIES: Dalmanites gemellusClarke, 1890 (5 ‘‘Vogesina’’ gemellus(Clarke) fide Lieberman et al., 1991) is pro-visionally assigned.

DIAGNOSIS: Cephalon twice wider thanlong, gently convex (tr. and sag.). All lateralglabellar furrows well incised; apodemal partof S1 abruptly shallowing, S1 faintly conflu-ent with axial furrow; S2 effaced abaxially;S3 steeply inclined exsagittally. Pygidiumtriangular in outline, relatively wide, lackingmarginal spines or lappets; pygidial axiscomprises 11 rings (first 5 separated bygroovelike impression of distal part of ringfurrows, posterior rings progressively weak-

2003 3CARVALHO ET AL.: CALMONIID TRILOBITES FROM BOLIVIA

Fig. 1. A, map of Bolivia, showing location of inset (map B); B, locations of trilobite collectionsites in this work (indicated by 3) relative to major cities and towns.

er); pleural field moderately convex (tr.),with at least six ribs.

Gemelloides delasernai, new speciesFigure 2A, B

DERIVATION OF NAME: After Salvador de laSerna, field companion to L. Smith for manyyears, who helped to collect the material.

DIAGNOSIS: Lateral glabellar furrows S2and S3 sharply defined; apodemal part of S1wedge-shaped; sculpture consisting of small,strong pits widely distributed on cephalon,thorax, and pygidium, without coarse gran-ules or tubercles.

TYPES: Holotype: MHNC 8130, externalmold of almost complete specimen (fig. 2A,B), from the lower part of the Upper Memberof the Belen Formation, layer of Wolfartaspiscornutus (Wolfart, 1968), late Emsian, Belenarea, La Paz Department, Bolivia (latex castAMNH 48073). Paratype: AMNH 48074, in-ternal mold of part of thorax and externalmold of frontal glabellar lobe, from type lo-cality.

DESCRIPTION: The cephalon is wider thanlong, with length (sag.) of cranidium 7.2 mmand width (tr.) across posterior border ap-

proximately 16 mm. Axial furrows are verydistinct, narrow, weakly divergent, slightlycurved externally at L2. The widest point ofthe glabella (across L3) is approximately 7mm, and its posterior width is 4.9 mm. Theoccipital furrow (S0) is deep, moderatelywide, curving anteriorly (sag.). S1 is a deep,wedge-shaped groove, widest mesially,sharply narrowing and shallowing distally,faintly confluent with the axial furrow, pos-teromedian part positioned close to S0; S2 islonger and narrower than S1 (tr.), slightlyconvex forward, with weak posteromedialorientation, effaced well inward of axial fur-row; S3 is straight, narrow, widest distally,strongly divergent toward the anterior gla-bellar margin (this margin is not preserved,and it is uncertain whether S3 reaches it); thestrongly divergent arrangement of S3 givesrise to a pattern in which the glabellar fur-rows appear to radiate away from the centerof the glabella. The glabellar furrows do notcross the median region of the glabella,which is not inflated. The glabellar lobeslikewise lack independent inflation; L1 isnarrow mesially, becoming wider near theaxial furrow; L2 is wider than L1 and is trap-


Fig. 2. Gemelloides delasernai, n. gen. andsp. Holotype MHNC 8130, Upper Member of Be-len Formation, Belen, La Paz Department. A, dor-sal view of nearly complete specimen, latex castfrom external mold, scale 2 mm; B, detail of ce-phalon, scale 2 mm.

ezoidal, becoming wider laterally; L3 is alsotrapezoidal in outline and is larger than L2.The posterior median impression is a short(sag.), elongate pit on the frontal lobe, ante-rior to the tips of S3. The frontal lobe is notinflated or distinguished from the rest of theglabella. L0 is moderately arched (tr.), lon-gest sagittally, distinctly wider than L1, witha width of 5.5 mm. The posterior border fur-

row is straight and moderately wide acrossmost of its extent, gradually narrowing nearthe genal angle. The posterior border is verynarrow adjacent to the axial furrow but isconsiderably wider (exsag.) abaxially, withan evenly convex posterior margin and arounded genal angle. The palpebral lobe andsurrounding area of the fixigena are some-what swollen, sloping down laterally behindand around the eye. The palpebral lobe isnarrow, bounded by a faint palpebral furrow.The cephalon (glabella, L0, fixigenal field,and posterior border) is ornamented withsmall, widely distributed pits, but lacks anygranular or tuberculate sculpture.

The thorax comprises 11 segments, allweakly convex (tr.) with the axial regionsomewhat higher than the pleurae. Maximumaxial width is about one- third that of thethorax. The first four axial rings are faintlyconvex forward sagittally; this convexity isgradually accentuated in more posteriorrings. Axial furrows are shallow and narrowbut well defined. Thoracic pleurae are trans-verse and horizontal proximally, becomingweakly flexed posteriorly in the distal por-tion; this flexure is also more accentuatedposteriorly. Pleural furrows are deep, narrow,and run straight across the pleurae onto thearticulating facets. The margin of the anteriorpleural band is convex proximally; the pos-terior band is longer than the anterior (tr.)and is broader distally (exsag.).

The pygidium is moderately large (9.4 mmmaximum width) and triangular in outline.The axis is only slightly higher than the pleu-ral field, defined by shallow, narrow axialfurrows, and includes 11 discernible rings.The first four rings are gently flexed anteri-orly and are separated by broad ring furrows,which are deepest distally and impressed astransverse grooves; the following rings areprogressively fainter. The axial terminus isindistinctly defined, but a postaxial field, ifpresent, is short (sag.). The pleural field isgently arched (tr.) and comprises at least sixribs. The first three are gently curved poste-riorly, but the remaining ribs are more dis-tinctly flexed. Pleural furrows are narrow butmore distinct than the interpleural furrows,which are only weakly developed. Little ofthe pygidial margin is visible, but its exposedpart (at the ends of the anterior three ribs)


lacks marginal spines or lappets. The axialrings and pleurae of the thorax and pygidiumare ornamented with small pits like thosecovering the cephalon.

DISCUSSION: The close similarity betweenGemelloides and Vogesina Wolfart, 1968 in-dicates membership in the Malvinella group(sensu Lieberman et al., 1991), and the genuspossesses more general apomorphies of thatgroup, such as depression of the abaxial partof L1 beneath L2 (character 23 of Liebermanet al, 1991). Vogesina is united with Palpe-brops Lieberman et al., 1991 by a convex(sag.) glabella (Lieberman et al., 1991, node9, character 8), which is shared to a lesserdegree by Gemelloides. None of the threecharacters supporting node 10 in their phy-logeny (which pertain to the anterior part ofthe cephalon and the hypostome) can be de-termined in our material, but both characterssupporting their node 11 (Vogesina s.l., char-acters 5, 11) are present (weakly divergentcephalic axial furrows and steep inclinationof S3). The latter feature gives rise to a dis-tinctive radial pattern of glabellar furrowswhich is also weakly evident in Palpebrops,and may thus represent a synapomorphy ofVogesina, Gemelloides, and perhaps Palpe-brops.

Within Vogesina as recognized by Lieber-man et al. (1991), V. aspera and V. lacuna-fera are separated from ‘‘Vogesina’’gemellus(5 Gemelloides gemellus here) by 12 char-acters. Three of these cannot be observed inthe specimens of Gemelloides delasernai, n.sp., but this taxon definitely lacks two othercharacters (8, 18; strongly convex glabella;elongate but nearly effaced S2 and S3) andshares seven apomorphies with V. aspera andV. lacunafera (12, 20, 27, 29, 32, 45, 48; S3straight; absence of coarse spines on lateralglabellar lobe L2; L0 not elevated above pos-terior glabellar region; absence of spines onL0; palpebral furrow weak and palpebral rimlow; absence of spines on thoracic axial ringsand on pygidial axis). The well-developedlateral glabellar furrows in Gemelloides de-lasernai, n. sp. clearly distinguish this formfrom all Vogesina spp., which have S2 andS3 as shallow to indistinct, typically resem-bling thin pencil lines (Wolfart, 1968). Thisdifference between Gemelloides and Voge-sina is marked even on external molds of Vo-

gesina of equivalent size to the holotype ofG. delasernai. The glabella is considerablyless convex (sag.) in Gemelloides delasernaithan in Vogesina and is not significantly el-evated above L0. The pygidium of Gemel-loides delasernai resembles that of Vogesinain lacking marginal lappets, but has a rela-tively broader outline, with the distal part ofthe pleurae less steeply turned down than inVogesina. Collectively, these findings sug-gest that G. delasernai is the sister taxon toVogesina but can be differentiated as a dis-tinct taxon on the basis of cephalic as wellas pygidial characters.

‘‘Vogesina’’ gemellus (Clarke), from theMaecuru Formation in the Amazon Basin, isknown only from an isolated, poorly pre-served glabella (Lieberman et al., 1991: fig.8.5, 8.6), and the state of only one of theapomorphic characters uniting G. delasernaiwith V. aspera and V. lacunafera (Liebermanet al., 1991) could be determined. ‘‘Vogesi-na’’ gemellus has a similar glabellar profile(sag.) and arrangement of lateral glabellarfurrows to G. delasernai. We provisionallyassign it to Gemelloides based on these char-acters, although the generic diagnosis em-phasizes the better known G. delasernai.Specific distinction between G. gemellus andG. delasernai is made based on the widerglabellar furrows, more elongated longitudi-nal median groove on the glabella, and tu-berculate sculpture of the former species.

Eldredgeia Lieberman, 1993

TYPE SPECIES: Metacryphaeus venustusWolfart, 1968, from the Sicasica Formation(Givetian), La Paz Department, Bolivia. Alsoknown from the Upper Member of the BelenFormation, La Paz Department (Lieberman,1993).

Eldredgeia eocryphaea, new speciesFigure 3A–L

DERIVATION OF NAME: ‘‘Early Cryphaeus’’,being geologically an early representative ofthe ‘‘Metacryphaeus group’’.

DIAGNOSIS: Species of Eldredgeia with thefollowing unique character combination:frontal glabellar lobe gently inflated, withrounded profile sloping down to anterior ce-phalic margin; posterior median impression a


small pit when present. Eyes set above gla-bella; visual surface bearing at least 24 dor-soventral files with maximum of nine lensesper file. Genal angle blunt, lacking a spine.Hypostome with pronounced ovoidal macu-lae; posterior border only moderately long;two pairs of short marginal spines presentposterolaterally.

TYPES: Holotype: MHNC 8129, internalmold of cephalon (fig. 3D–G) with hypo-stome in situ (fig. 3D), from the LowerMember of the Belen Formation, Scaphio-coelia Assemblage Zone, Tikani, Estacion deBombeo, Sicasica, La Paz Department. Para-types: MHNC 12900, pygidium with partialthorax (fig. 3J–L) in same concretion as ho-lotype and probably belonging to same in-dividual; AMNH 47147, internal mold of ce-phalon and articulated thorax (fig. 3A–C),and AMNH 47146, internal mold of pygidi-um (fig. 3H, I), both from the GamonedaFormation, Cerro Picacho, 17 km S of Tarija,Tarija Department. Other trilobites collectedfrom the same stratigraphic interval at CerroPicacho are Tarijactinoides jarcasensis, Ko-zlowskiaspis (Romanops) sp., and Gamone-daspis scutata, all indicative of the Scaphio-coelia Assemblage Zone.

DESCRIPTION: The cephalon is nearly twiceas wide as long and is moderately convex(tr.), with a widely pointed and arched out-line. Some details of the anteriormost part ofthe cephalon can be observed only in AMNH47147 (fig. 3A–C), as this region is missingin the holotype. The anterior cephalic borderis narrow and extends medially into a shortmedian triangular frontal process. The frontallobe is subrhomboidal, with a rounded an-terior margin and a rounded profile (sag.)with independent convexity from the poste-rior glabellar region, sloping down to the an-terior cephalic margin. A pitlike posteriormedian impression is present on the frontallobe in the holotype (fig. 3G), well in frontS3, but is indistinct in the paratype (fig. 3A,C). The auxiliary impression system is ob-scured by the granulation covering the fron-tal lobe. The axial furrow is slightly diver-gent from S0 to S1, but becomes more di-vergent from S1 anteriorly. All three pairs oflateral glabellar furrows are well developedand reach the axial furrow on the internalmold. S1 is broad, deep, and distinctly con-

cave forward; S2 is straight, narrower (tr.),and shallower than both S1 and S3. S1 andS2 are gently directed forward medially,whereas S3 is more oblique, diverging an-terolaterally. S0 is weakly curved forwardmedially, of even width (sag., exsag.), witha U-shaped section and deep apodemal pitsdistally. The lateral glabellar lobes are welldefined and ornamented by coarse granules.L1 is narrow (exsag.) in comparison with L2and L3 and can be traced across the glabellaby faint impression of S1 medially. L2 ismore or less rectangular in outline; L3 ismore wedge-shaped. L0 has a uniform width(sag., exsag.) throughout; its posterior mar-gin is approximately transverse. In profile,L0 lies in almost the same plane as the cen-tral part of the glabella. The lateral borderfurrow is wide but very faint, and it does notdistinctly separate the lateral border from thegenal field. The posterior border furrow is U-shaped in cross-section, deepest near the ax-ial furrow, and becomes gradually wider be-fore curving forward and shallowing distally.The posterior border is narrower (exsag.)than L0 proximally but becomes longer dis-tally. Although the genal angles are not wellpreserved in either specimen, they are bluntand lack a spine (the cavity behind the ex-ternal mold of the genal doublure in fig. 3Ais not a genal spine). The palpebral lobe isswollen, gently inclined adaxially/posterior-ly, and is ornamented by granules. The pal-pebral furrow is narrow but relatively sharplyimpressed along its length. In frontal viewthe free cheek is almost vertical and also hasgranular sculpture. The eye projects higherthan the glabella, with its anterior margin lo-cated opposite the anterodistal corner of L3,adjacent to (but still separate from) the axialfurrow. The posterior margin of the eye ispositioned far from the axial furrow, oppositethe posterolateral corner of L1 (exsag.). Thevisual surface is not well preserved in eitherspecimen; the number of dorsoventral filescannot be determined in the holotype, butAMNH 47147 has 24 dorsoventral files onthe visual surface. In that specimen no morethan seven lenses are preserved per file, butthe top of the eye is damaged and the up-permost lenses are missing. In the holotypeas many as nine lenses are discernible in thelongest files, as was probably the case in


AMNH 47147. The anterior section of thefacial suture runs parallel to the axial furrow,and it reaches the anterolateral corner of thefrontal lobe without cutting across it.

The hypostome is preserved in situ be-neath the cephalon of the holotype, but hasbeen pushed forward below the doublure andagainst the anterior part of the glabella. Thus,the anterior morphology of the hypostome(including its anterior margin, border, andwing) are hidden from view. The middlebody is roughly circular and moderately con-vex (tr. and sag.). The maculae are very dis-tinct, forming a pair of ovoid protuberancespositioned abaxially, near to the junction ofthe middle furrow (which is faint) and thelateral border furrow (which is moderatelydeep); the border furrow maintains aboutequal depth posterolaterally and posterome-dially. The lateral border is narrow, slightlyconvex, and runs subparallel posteriorly, andthe posterior border is long (sag.) and slight-ly convex (sag.). The posterior margin isrounded, with at least one pair of smallspines posterolaterally and traces of a secondpair laterally. The posteromedian margin ap-pears to be gently curved backward, andlacks a median spine.

The thorax, in lateral view, is moderatelyconvex, with the axis raised above the pleu-rae, and is one-third of the thoracic width.The axial rings are spatulate, shortened (sag.)medially, with granular ornament concentrat-ed distally. Apodemal pits are developed in-ward of the anterolateral edges of the rings.The articulating half ring is located below itscorresponding axial ring. The proximal one-third of the pleural field is more or less hor-izontal and the remainder is gently flexedventrolaterally. Each pleural furrow is deepand wide, extending straight diagonallyacross the proximal two-thirds of the pleura.The anterior band is narrowest proximallyand longest (exsag.) at the fulcrum; con-versely, the posterior band is longest (sag.)proximally and tapers to the fulcrum. Thepleural furrow is effaced distally. The distalextremities of the pleurae are pointed. Gran-ular sculpture is present across most of thewidth of the pleurae, but is more prominentproximal to the fulcrum.

The pygidium is broadly triangular in out-line, wider than long (length approximately

four-fifths of pygidial width, excluding mar-ginal lappets). The axial furrow is moderate-ly deep and narrow; the anterior part of theaxis tapers strongly back as far as the sixthring, but the remainder is almost parallel-sid-ed and has a blunt termination that does notreach the posterior margin of the pygidium.The axis is composed of at least eight rings,of which the first six are well developed andthe following two are weaker but distinct; aninth ring is faintly defined in the poorly seg-mented terminal part of the axis. The firstaxial ring is markedly arched anteriorly andsomewhat spatulate distally; the second,third, and fourth rings are likewise spatulatedistally, but are less arched anteriorly. Thefirst five axial rings are separated by broad,deep ring furrows with apodemes distally;behind this, ring furrows are shallow andtransverse. The pleural field has six pairs ofribs defined by wide pleural furrows, sepa-rated by narrow and shallow interpleural fur-rows; the fourth interpleural furrow is the lastdiscernible on the internal mold. The ribsgradually increase in obliquity from front toback, and the sixth is strongly oblique andpositioned near to the axis. The anterior fivepleurae terminate as spinelike lappets with aconvex anterior margin and faintly concaveposterior margin. The terminal lappet(known only from impressions) is wide butits shape and length cannot be establishedfrom the available material.

DISCUSSION: The new species Eldredgeiaeocryphaea shares numerous detailed simi-larities with E. venusta, including a narrowanterior border to the cephalon, with a shortmedian triangular frontal process; the ante-rior part of the frontal lobe is rounded; theeye is relatively long (exsag.); the thoracicand pygidial axial rings have similar shapes;the anterior five pygidial pleurae terminate aspointed spines; the posterior border of hy-postome is relatively long (sag.); and coarsegranular ornament is widely distributed overthe cephalon, thorax, and pygidium. Thesefeatures collectively support inclusion of thenew species in the genus Eldredgeia.

Eldredgeia eocryphaea is most readily dis-tinguished from the younger E. venusta (thetype species, also from Bolivia) by its hy-postome having more pronounced maculae(fig. 3D) and a substantially shorter (sag.)



←

Fig. 3. Eldredgeia eocryphaeus, n. sp. A–C, H, I, Gamoneda Formation, Cerro Picacho, TarijaDepartment. All scales 5 mm. A–C, AMNH 47147, dorsal, lateral, and anterodorsal views of cephalonand articulated thorax, internal mold; H, I, AMNH 47146, dorsal and posterior views of pygidium,internal mold. D–G, holotype MHNC 8129, internal mold of cephalon, Belen Formation, Tikani, LaPaz Department. All scales 5 mm. D, ventral view of hypostome; E–G, lateral, anterior, and dorsal viewsof cephalon; J–L, MHNC 12900, dorsal, posterior, and lateral views of pygidium. All scales 5 mm.

posterior border (see Lieberman et al., 1991:figs. 3.6, 3.7 for E. venustus). In addition, thegenal angle is blunter (versus more spinelikein E. venusta) and the visual surface has alower number of dorsoventral files (24 versus26–27 in E. venusta), although the maximumnumber of lenses per file (nine) is the samein both species.

Eldredgeia venusta has also been reportedfrom the Middle Devonian (Eifelian) of Bra-zil and South Africa; in South Africa the spe-cies is even said to characterize an E. venustaZone (Cooper, 1982). Lieberman (1993: 554)suggested that the South African and twoBrazilian forms probably represent addition-al, distinct species of Eldredgeia. Eldredgeiaeocryphaea, n. sp. differs from the South Af-rican form in having a higher number of dor-soventral files on the visual surface (24 ver-sus 18), more lenses per file (nine; the SouthAfrican form has six or seven), the shape ofthe genal angles (blunt rather than pointed),and glabellar lobes (more inflated in the formfrom South Africa).

The Brazilian form attributed to Eldredg-eia from the Amazon Basin (E. paituna(Hartt and Rathbun) from the Erere Forma-tion; see Lieberman, 1993: 554) can be dis-tinguished from E. eocryphaea, n. sp. in hav-ing a longer (sag.) frontal lobe and S1 withan accentuated crescent shape. In the otherBrazilian form (from the Parnaıba Basin, Pi-menteira Formation; see Lieberman et al.,1991: fig. 2), S1 is more strongly curved(adaxially) and L0 is longer (sag.). We en-dorse Lieberman’s (1993) suggestion that theSouth African and Brazilian material repre-sents additional species of Eldredgeia, andthe new form described in the present workappears to be distinct from all of them.

Eldredgeia eocryphaea significantly ex-tends the range of Eldredgeia into earlierstratigraphic intervals than previouslyknown. It also represents the earliest occur-

rence of the Metacryphaeus morphotype(i.e., the dalmanitiform cephalic morphologyand five-lappeted pygidial form shared byMetacryphaeus and allied genera as revisedby Lieberman, 1993). This morphotype hasnot previously been known from the Sca-phiocoelia Assemblage Zone. On the basis ofa late Lochkovian occurrence of Parabou-leia, a member of the Malvinella subgroupof the Metacryphaeus group, in Argentina,Edgecombe et al. (1994) inferred that line-ages of the Metacryphaeus group predatedtheir known occurrences in Bolivia. Eldredg-eia was one of several lineages in the Me-tacryphaeus group that was inferred to havean unsampled range extension (or ghost lin-eage) that considerably preceded its first ob-served appearance (in late Emsian or Eifelianstrata in the Upper Member of the Belen For-mation). The discovery of Eldredgeia eocry-phaea in the lower part of the Lower Mem-ber of the Belen Formation and equivalentstrata in the Gamoneda Formation (both rep-resenting the Scaphiocoelia AssemblageZone) closes one of the stratigraphic gaps inthe Metacryphaeus group.

Wolfartaspis Cooper, 1982

TYPE SPECIES: Metacryphaeus cornutusWolfart, 1968, from the lower part of the Up-per Member of the Belen Formation (lateEmsian), La Paz Department, Bolivia.

Wolfartaspis liebermani, new speciesFigures 4A–F, 5A–F

DERIVATION OF NAME: For Bruce S. Lie-berman, who has made valuable contribu-tions to systematics and biogeography of cal-moniid trilobites.

DIAGNOSIS: Cephalic anterior border weak-ly pointed medially; ventral view of cephalontriangular in outline. Eyes less than one-thirdglabella length (Large Eye Index 31%); vi-


sual surface with 25–26 dorsoventral fileswith a maximum of 10 lenses per file. Oc-cipital ring strongly arched in cross section,wide medially, with pronounced convex an-terior margin and robust median spine taper-ing dorsally.

TYPES: Holotype: MHNC 8132, externalmold of an incomplete cephalon, Icla For-mation, equivalent to basal part of UpperMember at Icla type locality (see Discussionbelow), Kochis Hills (approximately 3 kmnorth of the Rio Grande, on the border be-tween Chuquisaca and Cochabamba Depart-ments), Bolivia. Paratypes: AMNH 47148,internal mold of thorax and pygidium;AMNH 47149, internal mold of a cephalon;AMNH 47150, external mold of pygidium;AMNH 47151, external mold of an incom-plete pygidium; AMNH 47401, hypostomein situ beneath cephalon; AMNH 47403, in-ternal mold of almost complete cephalon;MHNC 12749, internal mold of incompletepygidium. All type specimens are from theIcla Formation, Kochis, Cochabamba De-partment.

OTHER MATERIAL: Some additional topo-type specimens collected within the samestratigraphic interval as the types are referredto Wolfartaspis liebermani, but they are notwell preserved. Most are from broken con-cretions and include three internal molds andtwo external molds of pygidia, an internalmold of four or five thoracic segments, anexternal mold of some pleurae, and two in-ternal molds of cephala. In addition, how-ever, there is a single, well-preserved, smallpygidium in part and counterpart. This ad-ditional material is housed in the MHNC col-lection.

DESCRIPTION: The cranidial length (sag.) isabout 29.7 mm in AMNH 47403. The ante-rior cephalic border is narrowest medially,where the upturned doublure nearly contactsthe cranidial margin; the anterior cephalicborder projects only slightly in front to theglabella medially in dorsal view; the anteriorborder gently widens abaxially, but remainsnarrow (exsag.) in dorsal view, and is steepalong its entire width. The cephalic antero-lateral margin is nearly straight in dorsalview and is rounded medially. The axial fur-row is deep, wide, straight, slightly divergentforward. Glabellar length (sag., excluding

S0) in AMNH 47403 is 23.4 mm, with thefrontal lobe having marked independent con-vexity from the posterior glabellar region,which has a flat sagittal profile. The frontalglabellar lobe is inflated, about 60% of gla-bellar length, and gradually expands anteri-orly and laterally, with its anteromedian mar-gin rounded. The anterior section of the fa-cial suture circumscribes but does not tran-sect the frontal glabellar lobe. The frontallobe has a welldeveloped circular to slightlyelongated posterior median impression (PMI)and auxiliary impression system (AIS) ofmuscle scars. The latter is most clearly pre-served on the left side and the median partof AMNH 47403 (fig. 5A) as small roundedpits on the internal mold, forming divergentrows. Medially, the AIS does not seem tohave a defined pattern. The lateral glabellarfurrows are well developed. S3 is wide, deep,oblique, straight, with its proximal partweakly bent posteromedially, lengtheningabaxially, and distinctly confluent with theaxial furrow. S2 is transverse, narrower andshallower than S1 and S3; it becomes abrupt-ly shallow distally and has only faint im-pression against the axial furrow on the ex-ternal cuticular surface (fig. 4A). S1 is deeperthan S2 and S3; it sharply narrows distallybut is distinctly confluent with the axial fur-row on the external mold; the posterior mar-gin of S1 is concave. L3 is wedge-shaped;L2 is approximately trapezoidal; L1 is slight-ly shorter (exsag.) than L2 and L3, with itsposterior margin markedly convex backwardand then continuing inward, forming a short(sag.) transglabellar lobe that is weaklycurved anteriorly; this feature makes the gla-bella slightly higher at L1 than at L3. S0 islong and relatively shallow (sag.), becomingshorter and deeper as it approaches the axialfurrow. Where the cephalic posterior furrowmeets S0, both have approximately the samewidth; the posterior furrow becomes wider(exsag.) toward the fulcrum, and is then gent-ly flexed forward and shallows distally. L0is strongly arched in its transverse section,wide medially, with a pronounced convexanterior margin, and bearing a robust medianspine that tapers dorsally; the median spineis weakly declined posteriorly. The eye ishigh, less than one-third glabellar length(Large Eye Index 31%). Its anterior edge is


Fig. 4. Wolfartaspis liebermani, n. sp. Icla Formation, Kochis, Cochabamba Department. All scales5 mm. A, holotype MHNC 8132, dorsal view of cephalon, latex cast from external mold; B, C, dorsaland lateral views of pygidium, AMNH 47150, latex cast from external mold; D, dorsal view of thoraxand pygidium, AMNH 47148, internal mold; E, dorsal view of pygidium and posterior part of thorax,AMNH 47151, latex cast from external mold; F, ventral view of hypostome in situ beneath cephalon,AMNH 47401, latex cast from external mold.


Fig. 5. Wolfartaspis liebermani, n. sp. Icla Formation, Kochis, Cochabamba Department. All scales5 mm. A–C, dorsal, lateral, and anterior views of cephalon and part of first thoracic segment, AMNH47403, internal mold; D–F, dorsal, anterior, and lateral views of cephalon, AMNH 47149, internal mold.

opposite the anterolateral corner of L3, andits posterior edge is opposite the posterolat-eral margin of L2 and at a considerable dis-tance from the posterior cephalic border fur-row (approximately 4 mm in the holotype).In anterior view, the visual surface is straightfrom top to bottom, gradually becoming wid-er toward its base, with 25–26 dorsoventrallens files, and a maximum of 10 lenses perfile. The interocular fixigena slopes upwardas it approaches the narrow, C-shaped pal-

pebral lobe; the palpebral furrow is moder-ately deep. The lateral cephalic border fur-row is faint. The posterior cephalic border isnarrow proximally, about equal in width(exsag.) to the posterior border furrow at thefulcrum, and widened distally; none of theavailable specimens preserves the genal an-gle.

The hypostome is preserved in situ be-neath the cephalon in one specimen (fig. 4F).Its anterior margin is largely missing, but its


course against the moderately wide anteriorwings suggests a transverse hypostomal su-ture. The middle body is roughly circular andgently convex (tr., sag.). The maculae arestrong, forming a pair of ovoid protuberancespositioned abaxially, near to the junction ofthe faint middle furrow and the lateral borderfurrow, anterior to the midlength of the mid-dle body. The border furrow is obscure an-terolaterally (against the wings), becomingmoderately impressed at the macula, main-taining moderate depth and width posterolat-erally and posteromedially. The lateral bor-der is narrow, with the lateral margins gentlyconverging between the anterior wing andfirst of two pairs of short, sharp posterolat-eral marginal spines; the inner pair of spines(on posterior border) is separated by slightlygreater than the distance between the twospine pairs. The posterior border is fairlylong and gently convex (sag.); the postero-median margin is faintly convex backwardbetween the inner pair of marginal spines.

The thorax consists of 11 segments. Therelative width of the axial rings and thoraxcannot be determined, as the distal ends ofthe pleurae are not preserved in any speci-mens, but in AMNH 47148 (fig. 4D) the ax-ial rings appear to be wider than the pleuralfields. The axial furrow is shallow and nar-row, weakly diverging back to the sixth ring,then gently converging. In lateral view, theaxial rings are faintly convex, raised wellabove the pleurae. The anteriormost axialrings are spatulate distally, but this is lessmarked in the more posterior rings. Small butdistinct apodemal pits lie between the axialrings abaxially. The pleural furrows are nar-row and moderately impressed adjacent tothe axial furrow, becoming wider distally,with a straight, oblique course across the rib;the anterior margin of the pleural furrows isgently convex anteriorly, while the posteriormargin of the furrows is straight.

The pygidium is triangular, moderatelyarched in lateral view, wider than long, witha pointed posterior tip. The axial furrow isnarrow and moderately deep. The axis isslightly waisted, the axial furrow moderatelyconvergent against the first five axial rings,weakly converging posterior to this; the axialterminus is blunt, lacking impression of theaxial furrow, lying well in advance of the

pygidial posterior margin. There are 10–11axial rings, but the last few are indistinct; thefirst five rings are gently arched anteriorly,but the subsequent ones are transverse orweakly bowed backward. The axial rings areseparated by wide ring furrows that becomeprogressively narrower posteriorly; the firstfive ring furrows have narrow, groovelikeapodemal pits distally. The pleural field in-cludes six pairs of strongly furrowed ribsplus a small seventh pair that parallels theaxis. The pleural furrows are deep and wide,though gradually narrowing on more poste-rior segments; they are moderately curved.The interpleural furrows are narrow, moder-ately deep proximal to the fulcrum betweenthe anterior few ribs, progressively shallowbetween more posterior ribs such that fifthinterpleural furrow is last distinct one on ex-ternal mold. The lateral margin of the pygid-ium is not well preserved in any of the ma-terial (fig.4E); the fifth lappet is small, flat,and wide (fig.4B), with a curved anteriormargin and straight posterior margin. Theposteromedian margin of the pygidium is anelongate, tapering, upturned spine (its basalpart is preserved in AMNH 47150; fig. 4B,C).

DISCUSSION: The monotypic Wolfartaspiswas erected by Cooper (1982) as a subgenusof Metacryphaeus, but recent usage (Lieber-man, 1993) recognizes it at the generic rank.According to Lieberman, Wolfartaspis dif-fers from Metacryphaeus in having an en-larged occipital spine and an upwardlycurved pygidial spine. The material de-scribed here includes both of these features,and it is therefore assigned to the genus Wol-fartaspis. We place our material in a newspecies, based on the following differencesfrom W. cornutus: (1) the ventral view of thecephalon has a rounded shape in W. cornu-tus, but is more pointed in our form; (2) theLarge Eye Index in W. cornutus is 34%, butis less in our material (31%); (3) the occipitalspine in W. cornutus (Lieberman, 1993: fig.2.8) is stouter than in our material (fig. 5E,F) and does not taper from its base. Differ-ences are also noted in the stratigraphic lev-els of the two forms. Wolfartaspis cornutusoccurs in the basal part of the Upper Memberof the Belen Formation, in strata overlyingthe layer of Francovichia branisi (5 layer of


Odontochile branisi sensu Wolfart, 1968, inthe upper part of Lower Member of BelenFormation). At Kochis, Francovichia occursstratigraphically above the beds with Wolfar-taspis liebermani.

Granadocephalus, new genus

DERIVATION OF NAME: Combination oftype locality (Villa Granado, CochabambaDepartment) and ‘‘cephalon’’.

TYPE SPECIES: Granadocephalus hanniba-li, n. gen. and sp.

REFERRED SPECIES: Monotypic.DIAGNOSIS: Cephalon moderately convex

(tr.), subsemicircular in outline; ventral mar-gin with moderate anterior arch. Profile ofglabella evenly curved, steepening anterior-ly; frontal lobe not distinctly separated fromrest of glabella; posterior median impressionweakly developed. Axial furrow moderatelyimpressed, divergent especially from S2,with small, rounded fossular depression nearanteromedial corner of palpebral lobe. Gla-bellar furrows not reaching axial furrow oninternal mold; S3 lightly impressed, narrow,faintly sinuous; S2 shallow but deeper thanS3, weakly directed backward at its inner tip;S1 deeper, weakly concave anteriorly, direct-ed anteromedially; S0 with strongly convexanterior margin. L3 and L2 not inflated; L1about two- thirds length (exsag.) of L2. L0slightly shortened abaxially. Anterior sectionof facial suture not transecting anterolateralcorner of glabella. Palpebral area slopesdownward abaxially, lying entirely below in-terocular fixigena; palpebral furrow weak.Librigenal field relatively strongly pitted.Posterior cephalic border furrow straight. En-tire cephalon covered by very fine granularornament on internal mold.

DISCUSSION: We were hesitant to create anew genus and species on the basis of sucha limited sample. However, the material dis-plays a unique combination of characterswithin the Calmoniidae (e.g., ventral marginwith moderate anterior arch; a peculiar con-vex [sag.] profile of the glabella; PMI weaklydeveloped; distinctive shape and develop-ment of the glabellar furrows and lobes; an-terior section of the facial suture does nottransect the frontal lobe; visual surface low,lying entirely below the interocular fixigena)

that does not permit it to be placed in any ofthe described calmoniid genera. We considerthat the pygidium and the cephalon includedhere belong to a single species. Both werecollected at the same locality and stratigraph-ic level, they correspond in size, the pygidi-um has appropriate proportions of the axisand pleurae relative to the cephalon, theyhave similar depth and width of the pleuralfurrows on the thorax (fig. 6E) and pygidium(fig. 6F), and the pygidium cannot be as-signed to any other calmoniid taxon. Still,since the association of the pygidium is notcertain, the generic diagnosis is restricted tocephalic characters.

The affinities of Granadocephalus are dif-ficult to determine. Of suprageneric groupsidentified in previous work (Eldredge, 1979;Eldredge and Ormiston, 1979; Eldredge andBranisa, 1980), Granadocephalus lacks thediagnostic characters of either the Probolopsgroup or the Malvinella subgroup, and mem-bership in the Calmonia group is most prob-able. In the Metacryphaeus group, the onlyobvious comparison is with the topologicallyprimitive genus Plesioconvexa Lieberman,1993, which has an evenly convex glabellarprofile reminiscent of Granadocephalus(compare Plesioconvexa praecursor in Lie-berman, 1993: fig. 3.8 with G. hannibali,here fig. 6C). However, Plesioconvexa, likeother members of the Metacryphaeus group,has much deeper incision of S2 and S3 thandoes Granadocephalus. In Plesioconvexa, S2has the characteristic transverse, apodemalimpression shared by other members of theMetacryphaeus group. Furthermore, Plesio-convexa, and indeed all members of the Me-tacryphaeus group except for deeply nested,highly modified taxa such as Vogesina, hasa much deeper palpebral furrow than doesGranadocephalus. Character states such aslight impression of S2 and S3, the formerbeing weakly convex forward and the lattersinuous, as well as a shallow palpebral fur-row, are shared by Granadocephalus andmembers of the Calmonia group. These sim-ilarities, however, may be primitive charac-ters for Calmoniidae. Eldredge and Branisa(1980) noted that axial morphology of theCalmonia group displays conservative char-acters for Calmoniidae, whereas the variousgenera of that group are more variable in


Fig. 6. Granadocephalus hannibali, n. gen. and sp. Icla Formation, 5 km east of Villa Granado,Cochabamba Department. All scales 10 mm. A–E, holotype MHNC 8131, cephalon with a few artic-ulated thoracic segments (internal mold): (A) dorsal; (B) ventral; (C) lateral; (D) anterodorsal; and (E)oblique posterodorsal views (orientations with respect to cephalon); F, G, dorsal and lateral views ofpygidium, MHNC 9451, internal mold.

their features of the exoskeletal margin (e.g.,genal spines, thoracic pleural spines, pygidialmarginal spines). The state of these charac-ters is not known for Granadocephalus.

Granadocephalus hannibali, new speciesFigure 6A–G

DERIVATION OF NAME: The species nameexpresses our appreciation to Dr. Joseph T.Hannibal (Cleveland Museum of NaturalHistory), who has made significant contri-butions to the knowledge of Bolivian pale-ontology, and who collected the holotype.

DIAGNOSIS: As for genus.TYPES: Holotype: internal mold of cephal-

on with a few displaced, articulated thoracicsegments, MHNC 8131 (cast AMNH 47150)(fig. 6A–E). Paratype: internal mold of py-gidium, MHNC 9451 (fig. 6F, G). Both fromthe Icla Formation, layer of Francovichiabranisi (Emsian, inferring equivalence tolayer of F. branisi sensu Wolfart (1968) inthe Belen Formation in the Altiplano), on ahillside 5 km east of Villa Granado, Cocha-bamba Department, Bolivia, near the high-way leading from Aiquile to Santa Cruz dela Sierra.

DESCRIPTION: The cephalon is moderatelyarched (tr.), subsemicircular in outline. Itsmaximum width is 45 mm (across the pos-


terior border) and its sagittal length is 30mm. The ventral margin is moderatelyarched anteriorly, with at least two pairs oflow, widely spaced protuberances. The an-terior cranidial border is uniformly narrow,set off by a sharp anterior cranidial borderfurrow. The anterior cephalic border is alsonarrow, especially medially. The axial furrowis moderately impressed, moderately wide,divergent forward especially from S2, with asmall, rounded fossular pit situated oppositeS3 (exsag.) and near the anteromedial cornerof the eye. The anterior section of the facialsuture circumscribes the frontal glabellarlobe, passing around its anterolateral cornerwithout transecting it. Glabellar length (sag.)is 25 mm (excluding S0) and its maximumwidth is 27 mm. In longitudinal profile, theglabella is curved, without a break in slopebetween the frontal lobe and S0 medially; thefrontal lobe is steepest anteriorly. The gla-bellar median region is almost flat (tr.). Theposterior median impression (PMI) is weaklydeveloped as a faint elongate groove that ex-tends back nearly opposite the inner tips ofS3. The three pairs of glabellar furrows arenot connected with the axial furrow. S3 isnarrow, lightly impressed, and faintly sinu-ous; S2 is shallow but deeper than S3, weak-ly convex anteriorly, in part by a gentle pos-terior curvature at its inner tip; the overallcourse of S2 is approximately transverse, ef-facing well inward of the axial furrow; S1 isdeeper impressed as gently concave, antero-medially directed apodemal grooves, abrupt-ly shallowing distally, at most faint imme-diately adjacent to axial furrow. S0 is welldeveloped, with a strongly convex anteriormargin and a gently convex posterior margin,such that S0 is longest sagittally; medial partof S0 moderately deep, distal part incised asfaintly concave apodemal grooves that par-allel apodemes in S1. L2 and L3 lack inde-pendent inflation; L3 is wedge-shaped, ex-panded distally; L2 is more conspicuous thanL3, and slightly narrowed medially. L1 isabout two- thirds the length (exsag.) of L2.L0 is of even length across much of itswidth, with slight shortening distally, behindthe apodemal part of S0; its width is 15 mm.The gena is subtrigonal, moderately convex.The palpebral area slopes downward laterallyto the border furrow. The eye, though not

well preserved, is relatively small, with thevisual surface about 8 mm long and not el-evated, lying entirely below the interocularfixigena; midlength of the palpebral furrowis opposite S2, its posterior edge opposite theposterolateral corner of L2. The anterior edgeof the palpebral lobe is very close to the axialfurrow, but the distance between them in-creases posteriorly such that the posterioredge of the eye is equidistant between theaxial furrow and the posterior cephalic bor-der furrow. The librigenal field slopes steeplydown to the lateral cephalic border furrow,with the field bearing numerous strong pits.The lateral border of the cephalon and thegenal angle are not preserved. The posteriorcephalic border furrow is narrow, deep, andtransverse; its anterior margin is straight. Theposterior border gradually widens (exsag.)distally, more strongly widening distal to thefulcrum. The cephalon is ornamented withfine granulation, which is especially dense onthe glabella.

Only an internal mold of a laterally incom-plete pygidium is known. The axial furrowis shallow, narrow, and rather strongly con-verging posteriorly. The axis appears to beslightly wider than the inferred width of thepleurae (excluding marginal spines, if pre-sent). In lateral view the axis is gently raisedabove the pleural field. Five axial rings arestrongly developed, plus parts of a sixth. Thefirst two rings are moderately arched anteri-orly and the succeeding ones are more gentlyarched. Medially the rings are widely sepa-rated from each other, with a pseudoarticu-lating half ring in the first two ring furrows;these rings are shortest sagittally and distinct-ly spatulate distally. Apodemal pits are de-veloped in the distal parts of the ring fur-rows. The axis terminates far in advance ofthe posteromedian margin of the pygidium,with the postaxial region defined by a changein convexity rather than distinct impressionof the axial furrow posteromedially. Fivepleurae are preserved on the right side, butonly parts of four are present on the left. Thepleurae are gently arched (tr.), the ribslengthening abaxially; the anteriormost ribsare gently and evenly curved laterally, butthe last ones bend back more abruptly. Thefirst interpleural furrow is narrow but sharplyimpressed, the succeeding furrows are simi-


larly narrow, but shallow; the fourth is thelast discernible interpleural furrow on the in-ternal mold. The pleural furrows are deepand relatively wide, maintaining their im-pression to the inner edge of the doublure.The lateral and posterior margins are not pre-served, though the even outer margin of thedoublure beneath the fourth and fifth ribssuggests that marginal spines or lappets wereprobably lacking on those segments at least.

ACKNOWLEDGMENTS

Yongyi Zhen (Australian Museum) photo-graphed the material and assembled the plates.For assistance in the field we thank Joseph T.Hannibal (Cleveland Museum of Natural His-tory), Salvador de la Serna (Cochabamba, Bo-livia), and Mr. Angel. Susan Klofak (AMNH)and Gary Chilson (Plantation, Florida) assist-ed with preparation and casting of specimens.Suggestions by the reviewers are appreciated.M.G.P.C. thanks John Maisey (AMNH) forhis support and encouragement, and AdrianaAquino (AMNH) for helping with the Spanishabstract.

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Recent issues of the Novitates may be purchased from the Museum. Lists of back issues of theNovitates and Bulletin published during the last five years are available at World Wide Web sitehttp://library.amnh.org. Or address mail orders to: American Museum of Natural History Library,Central Park West at 79th St., New York, NY 10024. TEL: (212) 769-5545. FAX: (212) 769-5009. E-MAIL: [email protected]

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