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

PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, NY 10024 Number 3407, 17 pp., 6 ®gures May 22, 2003

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

RESUMEN Se describen cuatro nuevos taxa de la familia Calmoniidae del DevoÂnico Boliviano: Gemelloides delasernai, n. gen. y sp., Eldredgeia eocryphaeus, n. sp., Wolfartaspis liebermani, n.sp., y Granadocephalus hannibali, n.gen. y sp. El nuevo geÂnero Gemelloides es el grupo hermano de Vogesina Wolfart 1968. Eldredgeia eocryphaeus, registrada para la 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] 2 Principal Research Scientist, Australian Museum, 6 College Street, Sydney, NSW 2010, Australia. 3 266 Merrimon Avenue, Asheville, NC 28801-1218.

gap estratigraÂ®co/linaje duende en la historia temprana del grupo Metacryphaeus. TambieÂn parte de este grupo, la especie Wolfartaspis liebermani (FormacioÂn Icla, Kochis, Depar- tamento de Cochabamba) antecede a su uÂnico congeÂnere, W. cornutus. Entre los Calmon- iidae, Granadocephalus hannibali (FormacioÂn Icla del Departamento de Cochabamba) es caracterizada por una combinacioÂn de caracteres nueva para la familia. Este geÂnero mo- notõÂpico podrõÂa presentar como taxa maÂs cercanos miembros del grupo Calmonia.

INTRODUCTION which contributes to the taxonomic and stratigraphic record of Bolivian calmoniids. Calmoniid trilobites from the Devonian of Morphological terminology follows that used Bolivia have a long history of study. Since in the Treatise on Invertebrate Paleontology, the initial descriptive work by d'Orbigny Part O (Whittington and Kelly, 1997) as well (1842), numerous papers have documented a as the terminology of Eldredge and BranisÏa rich diversity in the Lower and Middle De- (1980). The term Large Eye Index was intro- vonian (e.g., Kozlowski, 1923; BranisÏa, duced by Wolfart (1968), calculated as the 1965; Wolfart, 1968; Eldredge and Ormiston, ratio between the exsagittal length of the eye 1979; Eldredge and BranisÏa, 1980; Lieber- and the sagittal length of the glabella exclud- man et al., 1991; Lieberman, 1993). Most ing S0. The chronostratigraphic scheme for Bolivian calmoniids occur in three forma- the Devonian of Bolivia is as summarized by tions localized in distinct geographic areas Adrain and Edgecombe (1996: ®g. 2), after (®g. 1): the BeleÂn Formation (BeleÂn-La Paz- Blieck et al. (1996). Specimens studied and Sicasica region of the West Bolivian Altipla- ®gured here are deposited in the collections no); the Icla Formation (Icla-Padilla region, of American Museum of Natural History Sierras Subandinas of central Bolivia), and (AMNH), Division of Paleontology (Inver- the Gamoneda Formation (Tarija region, tebrates), and Museo de HistoÂria Natural de southern Bolivia). The basal portion of all Cochabamba (MHNC), Bolivia. three formations is considered to be time- equivalent, falling within the ``Scaphiocoe- SYSTEMATIC PALEONTOLOGY lia-bearing beds'' (Isaacson, 1977) recog- FAMILY CALMONIIDAE DELO, 1935 nized as the Scaphiocoelia Assemblage Zone by Eldredge and BranisÏa (1980). Trilobites GEMELLOIDES, NEW GENUS occur both within and above this level. The DERIVATION OF NAME: Literally, ``like Ge- total stratigraphic range of Calmoniidae in mellus'', in reference to the name Dalmanites Bolivia is Late Silurian (PrÏõÂdolõÂ) (Edgecombe gemellus Clarke, 1890; gemellus (Latin), a and Fortey, 2000) to Middle Devonian (Giv- twin. etian). TYPE SPECIES: Gemelloides delasernai,n. It is now well established that calmoniid gen. and sp. trilobites are endemic to a Southern-hemi- REFERRED SPECIES: Dalmanites gemellus sphere biogeographic region that Clarke Clarke, 1890 (ϭ ``Vogesina'' gemellus (1913) ®rst characterized as an ``austral fau- (Clarke) ®de Lieberman et al., 1991) is pro- na'', which he was able to distinguish from visionally assigned. a ``meridional fauna''. Richter and Richter DIAGNOSIS: Cephalon twice wider than (1942) observed the endemic character of long, gently convex (tr. and sag.). All lateral this fauna and coined the term ``Malvinokaf- glabellar furrows well incised; apodemal part fric Province'', which corresponds generally of S1 abruptly shallowing, S1 faintly con¯u- to the Malvinokaffric Realm of Eldredge and ent with axial furrow; S2 effaced abaxially; Ormiston (1979). S3 steeply inclined exsagittally. Pygidium Smith and Edgecombe (1996) informally triangular in outline, relatively wide, lacking reported two new genera and four new spe- marginal spines or lappets; pygidial axis cies of Calmoniidae from Bolivia, but did not comprises 11 rings (®rst 5 separated by name or describe them. In this paper we pre- groovelike impression of distal part of ring sent a systematic description of this material, furrows, posterior rings progressively weak- 2003 CARVALHO ET AL.: CALMONIID TRILOBITES FROM BOLIVIA 3

Fig. 1. A, map of Bolivia, showing location of inset (map B); B, locations of trilobite collection sites in this work (indicated by ϫ) relative to major cities and towns. er); pleural ®eld moderately convex (tr.), proximately 16 mm. Axial furrows are very with at least six ribs. distinct, narrow, weakly divergent, slightly curved externally at L2. The widest point of Gemelloides delasernai, new species the glabella (across L3) is approximately 7 Figure 2A, B mm, and its posterior width is 4.9 mm. The occipital furrow (S0) is deep, moderately DERIVATION OF NAME: After Salvador de la Serna, ®eld companion to L. Smith for many wide, curving anteriorly (sag.). S1 is a deep, years, who helped to collect the material. wedge-shaped groove, widest mesially, DIAGNOSIS: Lateral glabellar furrows S2 sharply narrowing and shallowing distally, and S3 sharply de®ned; apodemal part of S1 faintly con¯uent with the axial furrow, pos- wedge-shaped; sculpture consisting of small, teromedian part positioned close to S0; S2 is strong pits widely distributed on cephalon, longer and narrower than S1 (tr.), slightly thorax, and pygidium, without coarse gran- convex forward, with weak posteromedial ules or tubercles. orientation, effaced well inward of axial fur- TYPES: Holotype: MHNC 8130, external row; S3 is straight, narrow, widest distally, mold of almost complete specimen (®g. 2A, strongly divergent toward the anterior gla- B), from the lower part of the Upper Member bellar margin (this margin is not preserved, of the BeleÂn Formation, layer of Wolfartaspis and it is uncertain whether S3 reaches it); the cornutus (Wolfart, 1968), late Emsian, BeleÂn strongly divergent arrangement of S3 gives area, La Paz Department, Bolivia (latex cast rise to a pattern in which the glabellar fur- AMNH 48073). Paratype: AMNH 48074, in- rows appear to radiate away from the center ternal mold of part of thorax and external of the glabella. The glabellar furrows do not mold of frontal glabellar lobe, from type lo- cross the median region of the glabella, cality. which is not in¯ated. The glabellar lobes DESCRIPTION: The cephalon is wider than likewise lack independent in¯ation; L1 is long, with length (sag.) of cranidium 7.2 mm narrow mesially, becoming wider near the and width (tr.) across posterior border ap- axial furrow; L2 is wider than L1 and is trap- 4 AMERICAN MUSEUM NOVITATES NO. 3407

row is straight and moderately wide across most of its extent, gradually narrowing near the genal angle. The posterior border is very narrow adjacent to the axial furrow but is considerably wider (exsag.) abaxially, with an evenly convex posterior margin and a rounded genal angle. The palpebral lobe and surrounding area of the ®xigena are somewhat swollen, sloping down laterally behind and around the eye. The palpebral lobe is narrow, bounded by a faint palpebral furrow. The cephalon (glabella, L0, ®xigenal ®eld, and posterior border) is ornamented with small, widely distributed pits, but lacks any granular or tuberculate sculpture. The thorax comprises 11 segments, all weakly convex (tr.) with the axial region somewhat higher than the pleurae. Maximum axial width is about one- third that of the thorax. The ®rst four axial rings are faintly convex forward sagittally; this convexity is gradually accentuated in more posterior rings. Axial furrows are shallow and narrow but well de®ned. Thoracic pleurae are transverse and horizontal proximally, becoming weakly ¯exed posteriorly in the distal portion; this ¯exure is also more accentuated posteriorly. Pleural furrows are deep, narrow, and run straight across the pleurae onto the articulating facets. The margin of the anterior pleural band is convex proximally; the posterior band is longer than the anterior (tr.) and is broader distally (exsag.). The pygidium is moderately large (9.4 mm maximum width) and triangular in outline. The axis is only slightly higher than the pleural ®eld, de®ned by shallow, narrow axial Fig. 2. Gemelloides delasernai, n. gen. and furrows, and includes 11 discernible rings. sp. Holotype MHNC 8130, Upper Member of Be- The ®rst four rings are gently ¯exed anteri- leÂn Formation, BeleÂn, La Paz Department. A, dor- orly and are separated by broad ring furrows, sal view of nearly complete specimen, latex cast which are deepest distally and impressed as from external mold, scale 2 mm; B, detail of ce- transverse grooves; the following rings are phalon, scale 2 mm. progressively fainter. The axial terminus is indistinctly de®ned, but a postaxial ®eld, if ezoidal, becoming wider laterally; L3 is also present, is short (sag.). The pleural ®eld is trapezoidal in outline and is larger than L2. gently arched (tr.) and comprises at least six The posterior median impression is a short ribs. The ®rst three are gently curved poste- (sag.), elongate pit on the frontal lobe, anteriorly, but the remaining ribs are more dis- rior to the tips of S3. The frontal lobe is not tinctly ¯exed. Pleural furrows are narrow but in¯ated or distinguished from the rest of the more distinct than the interpleural furrows, glabella. L0 is moderately arched (tr.), lon- which are only weakly developed. Little of gest sagittally, distinctly wider than L1, with the pygidial margin is visible, but its exposed a width of 5.5 mm. The posterior border fur- part (at the ends of the anterior three ribs) 2003 CARVALHO ET AL.: CALMONIID TRILOBITES FROM BOLIVIA 5 lacks marginal spines or lappets. The axial gesina of equivalent size to the holotype of rings and pleurae of the thorax and pygidium G. delasernai. The glabella is considerably are ornamented with small pits like those less convex (sag.) in Gemelloides delasernai covering the cephalon. than in Vogesina and is not signi®cantly el- DISCUSSION: The close similarity between evated above L0. The pygidium of Gemel- Gemelloides and Vogesina Wolfart, 1968 in- loides delasernai resembles that of Vogesina dicates membership in the Malvinella group in lacking marginal lappets, but has a rela- (sensu Lieberman et al., 1991), and the genus tively broader outline, with the distal part of possesses more general apomorphies of that the pleurae less steeply turned down than in group, such as depression of the abaxial part Vogesina. Collectively, these ®ndings sug- of L1 beneath L2 (character 23 of Lieberman gest that G. delasernai is the sister taxon to et al, 1991). Vogesina is united with Palpe- Vogesina but can be differentiated as a dis- brops Lieberman et al., 1991 by a convex tinct taxon on the basis of cephalic as well (sag.) glabella (Lieberman et al., 1991, node as pygidial characters. 9, character 8), which is shared to a lesser ``Vogesina'' gemellus (Clarke), from the degree by Gemelloides. None of the three Maecuru Formation in the Amazon Basin, is characters supporting node 10 in their phy- known only from an isolated, poorly pre- logeny (which pertain to the anterior part of served glabella (Lieberman et al., 1991: ®g. the cephalon and the hypostome) can be de- 8.5, 8.6), and the state of only one of the termined in our material, but both characters apomorphic characters uniting G. delasernai supporting their node 11 (Vogesina s.l., char- with V. aspera and V. lacunafera (Lieberman acters 5, 11) are present (weakly divergent et al., 1991) could be determined. ``Vogesi- cephalic axial furrows and steep inclination na'' gemellus has a similar glabellar pro®le of S3). The latter feature gives rise to a dis- (sag.) and arrangement of lateral glabellar tinctive radial pattern of glabellar furrows furrows to G. delasernai. We provisionally which is also weakly evident in Palpebrops, assign it to Gemelloides based on these char- and may thus represent a synapomorphy of acters, although the generic diagnosis em- Vogesina, Gemelloides, and perhaps Palpe- phasizes the better known G. delasernai. brops. Speci®c distinction between G. gemellus and Within Vogesina as recognized by Lieber- G. delasernai is made based on the wider man et al. (1991), V. aspera and V. lacuna- glabellar furrows, more elongated longitudi- fera are separated from ``Vogesina''gemellus nal median groove on the glabella, and tu- (ϭ Gemelloides gemellus here) by 12 char- berculate sculpture of the former species. acters. Three of these cannot be observed in the specimens of Gemelloides delasernai,n. Eldredgeia Lieberman, 1993 sp., but this taxon de®nitely lacks two other TYPE SPECIES: Metacryphaeus venustus characters (8, 18; strongly convex glabella; Wolfart, 1968, from the Sicasica Formation elongate but nearly effaced S2 and S3) and (Givetian), La Paz Department, Bolivia. Also shares seven apomorphies with V. aspera and known from the Upper Member of the BeleÂn V. lacunafera (12, 20, 27, 29, 32, 45, 48; S3 Formation, La Paz Department (Lieberman, straight; absence of coarse spines on lateral 1993). glabellar lobe L2; L0 not elevated above posterior glabellar region; absence of spines on Eldredgeia eocryphaea, new species L0; palpebral furrow weak and palpebral rim low; absence of spines on thoracic axial rings Figure 3A±L and on pygidial axis). The well-developed DERIVATION OF NAME: ``Early Cryphaeus'', lateral glabellar furrows in Gemelloides de- being geologically an early representative of lasernai, n. sp. clearly distinguish this form the ``Metacryphaeus group''. from all Vogesina spp., which have S2 and DIAGNOSIS: Species of Eldredgeia with the S3 as shallow to indistinct, typically resem- following unique character combination: bling thin pencil lines (Wolfart, 1968). This frontal glabellar lobe gently in¯ated, with difference between Gemelloides and Voge- rounded pro®le sloping down to anterior ce- sina is marked even on external molds of Vo- phalic margin; posterior median impression a 6 AMERICAN MUSEUM NOVITATES NO. 3407 small pit when present. Eyes set above gla- cave forward; S2 is straight, narrower (tr.), bella; visual surface bearing at least 24 dor- and shallower than both S1 and S3. S1 and soventral ®les with maximum of nine lenses S2 are gently directed forward medially, per ®le. Genal angle blunt, lacking a spine. whereas S3 is more oblique, diverging an- Hypostome with pronounced ovoidal macu- terolaterally. S0 is weakly curved forward lae; posterior border only moderately long; medially, of even width (sag., exsag.), with two pairs of short marginal spines present a U-shaped section and deep apodemal pits posterolaterally. distally. The lateral glabellar lobes are well TYPES: Holotype: MHNC 8129, internal de®ned and ornamented by coarse granules. mold of cephalon (®g. 3D±G) with hypo- L1 is narrow (exsag.) in comparison with L2 stome in situ (®g. 3D), from the Lower and L3 and can be traced across the glabella Member of the BeleÂn Formation, Scaphio- by faint impression of S1 medially. L2 is coelia Assemblage Zone, Tikani, EstacioÂn de more or less rectangular in outline; L3 is Bombeo, Sicasica, La Paz Department. Para- more wedge-shaped. L0 has a uniform width types: MHNC 12900, pygidium with partial (sag., exsag.) throughout; its posterior mar- thorax (®g. 3J±L) in same concretion as ho- gin is approximately transverse. In pro®le, lotype and probably belonging to same in- L0 lies in almost the same plane as the cen- dividual; AMNH 47147, internal mold of ce- tral part of the glabella. The lateral border phalon and articulated thorax (®g. 3A±C), furrow is wide but very faint, and it does not and AMNH 47146, internal mold of pygidi- distinctly separate the lateral border from the um (®g. 3H, I), both from the Gamoneda genal ®eld. The posterior border furrow is U- Formation, Cerro Picacho, 17 km S of Tarija, shaped in cross-section, deepest near the ax- Tarija Department. Other trilobites collected ial furrow, and becomes gradually wider be- from the same stratigraphic interval at Cerro fore curving forward and shallowing distally. Picacho are Tarijactinoides jarcasensis, Ko- The posterior border is narrower (exsag.) zlowskiaspis (Romanops) sp., and Gamone- than L0 proximally but becomes longer dis- daspis scutata, all indicative of the Scaphio- tally. Although the genal angles are not well coelia Assemblage Zone. preserved in either specimen, they are blunt DESCRIPTION: The cephalon is nearly twice and lack a spine (the cavity behind the ex- as wide as long and is moderately convex ternal mold of the genal doublure in ®g. 3A (tr.), with a widely pointed and arched out- is not a genal spine). The palpebral lobe is line. Some details of the anteriormost part of swollen, gently inclined adaxially/posterior- the cephalon can be observed only in AMNH ly, and is ornamented by granules. The pal- 47147 (®g. 3A±C), as this region is missing pebral furrow is narrow but relatively sharply in the holotype. The anterior cephalic border impressed along its length. In frontal view is narrow and extends medially into a short the free cheek is almost vertical and also has median triangular frontal process. The frontal granular sculpture. The eye projects higher lobe is subrhomboidal, with a rounded an- than the glabella, with its anterior margin lo- terior margin and a rounded pro®le (sag.) cated opposite the anterodistal corner of L3, with independent convexity from the poste- adjacent to (but still separate from) the axial rior glabellar region, sloping down to the an- furrow. The posterior margin of the eye is terior cephalic margin. A pitlike posterior positioned far from the axial furrow, opposite median impression is present on the frontal the posterolateral corner of L1 (exsag.). The lobe in the holotype (®g. 3G), well in front visual surface is not well preserved in either S3, but is indistinct in the paratype (®g. 3A, specimen; the number of dorsoventral ®les C). The auxiliary impression system is ob- cannot be determined in the holotype, but scured by the granulation covering the fron- AMNH 47147 has 24 dorsoventral ®les on tal lobe. The axial furrow is slightly diver- the visual surface. In that specimen no more gent from S0 to S1, but becomes more di- than seven lenses are preserved per ®le, but vergent from S1 anteriorly. All three pairs of the top of the eye is damaged and the up- lateral glabellar furrows are well developed permost lenses are missing. In the holotype and reach the axial furrow on the internal as many as nine lenses are discernible in the mold. S1 is broad, deep, and distinctly con- longest ®les, as was probably the case in 2003 CARVALHO ET AL.: CALMONIID TRILOBITES FROM BOLIVIA 7

AMNH 47147. The anterior section of the four-®fths of pygidial width, excluding mar- facial suture runs parallel to the axial furrow, ginal lappets). The axial furrow is moderate- and it reaches the anterolateral corner of the ly deep and narrow; the anterior part of the frontal lobe without cutting across it. axis tapers strongly back as far as the sixth The hypostome is preserved in situ be- ring, but the remainder is almost parallel-sid- neath the cephalon of the holotype, but has ed and has a blunt termination that does not been pushed forward below the doublure and reach the posterior margin of the pygidium. against the anterior part of the glabella. Thus, The axis is composed of at least eight rings, the anterior morphology of the hypostome of which the ®rst six are well developed and (including its anterior margin, border, and the following two are weaker but distinct; a wing) are hidden from view. The middle ninth ring is faintly de®ned in the poorly seg- body is roughly circular and moderately con- mented terminal part of the axis. The ®rst vex (tr. and sag.). The maculae are very dis- axial ring is markedly arched anteriorly and tinct, forming a pair of ovoid protuberances somewhat spatulate distally; the second, positioned abaxially, near to the junction of third, and fourth rings are likewise spatulate the middle furrow (which is faint) and the distally, but are less arched anteriorly. The lateral border furrow (which is moderately ®rst ®ve axial rings are separated by broad, deep); the border furrow maintains about deep ring furrows with apodemes distally; equal depth posterolaterally and posterome- behind this, ring furrows are shallow and dially. The lateral border is narrow, slightly transverse. The pleural ®eld has six pairs of convex, and runs subparallel posteriorly, and ribs de®ned by wide pleural furrows, sepa- the posterior border is long (sag.) and slight- rated by narrow and shallow interpleural fur- ly convex (sag.). The posterior margin is rows; the fourth interpleural furrow is the last rounded, with at least one pair of small discernible on the internal mold. The ribs spines posterolaterally and traces of a second gradually increase in obliquity from front to pair laterally. The posteromedian margin ap- back, and the sixth is strongly oblique and pears to be gently curved backward, and positioned near to the axis. The anterior ®ve lacks a median spine. pleurae terminate as spinelike lappets with a The thorax, in lateral view, is moderately convex anterior margin and faintly concave convex, with the axis raised above the pleu- posterior margin. The terminal lappet rae, and is one-third of the thoracic width. (known only from impressions) is wide but The axial rings are spatulate, shortened (sag.) its shape and length cannot be established medially, with granular ornament concentrat- from the available material. ed distally. Apodemal pits are developed in- DISCUSSION: The new species Eldredgeia ward of the anterolateral edges of the rings. eocryphaea shares numerous detailed simi- The articulating half ring is located below its larities with E. venusta, including a narrow corresponding axial ring. The proximal one- anterior border to the cephalon, with a short third of the pleural ®eld is more or less hor- median triangular frontal process; the ante- izontal and the remainder is gently ¯exed rior part of the frontal lobe is rounded; the ventrolaterally. Each pleural furrow is deep eye is relatively long (exsag.); the thoracic and wide, extending straight diagonally and pygidial axial rings have similar shapes; across the proximal two-thirds of the pleura. the anterior ®ve pygidial pleurae terminate as The anterior band is narrowest proximally pointed spines; the posterior border of hy- and longest (exsag.) at the fulcrum; con- postome is relatively long (sag.); and coarse versely, the posterior band is longest (sag.) granular ornament is widely distributed over proximally and tapers to the fulcrum. The the cephalon, thorax, and pygidium. These pleural furrow is effaced distally. The distal features collectively support inclusion of the extremities of the pleurae are pointed. Gran- new species in the genus Eldredgeia. ular sculpture is present across most of the Eldredgeia eocryphaea is most readily dis- width of the pleurae, but is more prominent tinguished from the younger E. venusta (the proximal to the fulcrum. type species, also from Bolivia) by its hy- The pygidium is broadly triangular in out- postome having more pronounced maculae line, wider than long (length approximately (®g. 3D) and a substantially shorter (sag.) 8 AMERICAN MUSEUM NOVITATES NO. 3407 2003 CARVALHO ET AL.: CALMONIID TRILOBITES FROM BOLIVIA 9 posterior border (see Lieberman et al., 1991: rence of the Metacryphaeus morphotype ®gs. 3.6, 3.7 for E. venustus). In addition, the (i.e., the dalmanitiform cephalic morphology genal angle is blunter (versus more spinelike and ®ve-lappeted pygidial form shared by in E. venusta) and the visual surface has a Metacryphaeus and allied genera as revised lower number of dorsoventral ®les (24 versus by Lieberman, 1993). This morphotype has 26±27 in E. venusta), although the maximum not previously been known from the Sca- number of lenses per ®le (nine) is the same phiocoelia Assemblage Zone. On the basis of in both species. a late Lochkovian occurrence of Parabou- Eldredgeia venusta has also been reported leia, a member of the Malvinella subgroup from the Middle Devonian (Eifelian) of Bra- of the Metacryphaeus group, in Argentina, zil and South Africa; in South Africa the spe- Edgecombe et al. (1994) inferred that line- cies is even said to characterize an E. venusta ages of the Metacryphaeus group predated Zone (Cooper, 1982). Lieberman (1993: 554) their known occurrences in Bolivia. Eldredg- suggested that the South African and two eia was one of several lineages in the Me- Brazilian forms probably represent addition- tacryphaeus group that was inferred to have al, distinct species of Eldredgeia. Eldredgeia an unsampled range extension (or ghost lin- eocryphaea, n. sp. differs from the South Af- eage) that considerably preceded its ®rst ob- rican form in having a higher number of dor- served appearance (in late Emsian or Eifelian soventral ®les on the visual surface (24 ver- strata in the Upper Member of the BeleÂn For- sus 18), more lenses per ®le (nine; the South mation). The discovery of Eldredgeia eocry- African form has six or seven), the shape of phaea in the lower part of the Lower Mem- the genal angles (blunt rather than pointed), ber of the BeleÂn Formation and equivalent and glabellar lobes (more in¯ated in the form strata in the Gamoneda Formation (both rep- from South Africa). resenting the Scaphiocoelia Assemblage The Brazilian form attributed to Eldredg- Zone) closes one of the stratigraphic gaps in eia from the Amazon Basin (E. paituna the Metacryphaeus group. (Hartt and Rathbun) from the ErereÃ Forma- tion; see Lieberman, 1993: 554) can be dis- Wolfartaspis Cooper, 1982 tinguished from E. eocryphaea, n. sp. in hav- TYPE SPECIES: Metacryphaeus cornutus ing a longer (sag.) frontal lobe and S1 with Wolfart, 1968, from the lower part of the Up- an accentuated crescent shape. In the other per Member of the BeleÂn Formation (late Brazilian form (from the ParnaõÂba Basin, Pi- Emsian), La Paz Department, Bolivia. menteira Formation; see Lieberman et al., 1991: ®g. 2), S1 is more strongly curved Wolfartaspis liebermani, new species (adaxially) and L0 is longer (sag.). We en- Figures 4A±F, 5A±F dorse Lieberman's (1993) suggestion that the South African and Brazilian material repre- DERIVATION OF NAME: For Bruce S. Lie- sents additional species of Eldredgeia, and berman, who has made valuable contribu- the new form described in the present work tions to systematics and biogeography of cal- appears to be distinct from all of them. moniid trilobites. Eldredgeia eocryphaea signi®cantly ex- DIAGNOSIS: Cephalic anterior border weak- tends the range of Eldredgeia into earlier ly pointed medially; ventral view of cephalon stratigraphic intervals than previously triangular in outline. Eyes less than one-third known. It also represents the earliest occur- glabella length (Large Eye Index 31%); vi-

← Fig. 3. Eldredgeia eocryphaeus,n.sp.A±C, H, I, Gamoneda Formation, Cerro Picacho, Tarija Department. All scales 5 mm. A±C, AMNH 47147, dorsal, lateral, and anterodorsal views of cephalon and 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, BeleÂn Formation, Tikani, La Paz Department. All scales 5 mm. D, ventral view of hypostome; E±G, lateral, anterior, and dorsal views of cephalon; J±L, MHNC 12900, dorsal, posterior, and lateral views of pygidium. All scales 5 mm. 10 AMERICAN MUSEUM NOVITATES NO. 3407 sual surface with 25±26 dorsoventral ®les S0) in AMNH 47403 is 23.4 mm, with the with a maximum of 10 lenses per ®le. Oc- frontal lobe having marked independent con- cipital ring strongly arched in cross section, vexity from the posterior glabellar region, wide medially, with pronounced convex an- which has a ¯at sagittal pro®le. The frontal terior margin and robust median spine taper- glabellar lobe is in¯ated, about 60% of gla- ing dorsally. bellar length, and gradually expands anteri- TYPES: Holotype: MHNC 8132, external orly and laterally, with its anteromedian mar- mold of an incomplete cephalon, Icla For- gin rounded. The anterior section of the fa- mation, equivalent to basal part of Upper cial suture circumscribes but does not tran- Member at Icla type locality (see Discussion sect the frontal glabellar lobe. The frontal below), Kochis Hills (approximately 3 km lobe has a well- developed circular to slightly north of the Rio Grande, on the border be- elongated posterior median impression (PMI) tween Chuquisaca and Cochabamba Depart- and auxiliary impression system (AIS) of ments), Bolivia. Paratypes: AMNH 47148, muscle scars. The latter is most clearly pre- internal mold of thorax and pygidium; served on the left side and the median part AMNH 47149, internal mold of a cephalon; of AMNH 47403 (®g. 5A) as small rounded AMNH 47150, external mold of pygidium; pits on the internal mold, forming divergent AMNH 47151, external mold of an incom- rows. Medially, the AIS does not seem to plete pygidium; AMNH 47401, hypostome have a de®ned pattern. The lateral glabellar in situ beneath cephalon; AMNH 47403, in- furrows are well developed. S3 is wide, deep, ternal mold of almost complete cephalon; oblique, straight, with its proximal part MHNC 12749, internal mold of incomplete weakly bent posteromedially, lengthening pygidium. All type specimens are from the abaxially, and distinctly con¯uent with the Icla Formation, Kochis, Cochabamba De- axial furrow. S2 is transverse, narrower and partment. shallower than S1 and S3; it becomes abrupt- OTHER MATERIAL: Some additional topo- ly shallow distally and has only faint im- type specimens collected within the same pression against the axial furrow on the ex- stratigraphic interval as the types are referred ternal cuticular surface (®g. 4A). S1 is deeper to Wolfartaspis liebermani, but they are not than S2 and S3; it sharply narrows distally well preserved. Most are from broken con- but is distinctly con¯uent with the axial fur- cretions and include three internal molds and row on the external mold; the posterior mar- two external molds of pygidia, an internal gin of S1 is concave. L3 is wedge-shaped; mold of four or ®ve thoracic segments, an L2 is approximately trapezoidal; L1 is slight- external mold of some pleurae, and two in- ly shorter (exsag.) than L2 and L3, with its ternal molds of cephala. In addition, how- posterior margin markedly convex backward ever, there is a single, well-preserved, small and then continuing inward, forming a short pygidium in part and counterpart. This ad- (sag.) transglabellar lobe that is weakly ditional material is housed in the MHNC col- curved anteriorly; this feature makes the gla- lection. bella slightly higher at L1 than at L3. S0 is DESCRIPTION: The cranidial length (sag.) is long and relatively shallow (sag.), becoming about 29.7 mm in AMNH 47403. The ante- shorter and deeper as it approaches the axial rior cephalic border is narrowest medially, furrow. Where the cephalic posterior furrow where the upturned doublure nearly contacts meets S0, both have approximately the same the cranidial margin; the anterior cephalic width; the posterior furrow becomes wider border projects only slightly in front to the (exsag.) toward the fulcrum, and is then gent- glabella medially in dorsal view; the anterior ly ¯exed forward and shallows distally. L0 border gently widens abaxially, but remains is strongly arched in its transverse section, narrow (exsag.) in dorsal view, and is steep wide medially, with a pronounced convex along its entire width. The cephalic antero- anterior margin, and bearing a robust median lateral margin is nearly straight in dorsal spine that tapers dorsally; the median spine view and is rounded medially. The axial fur- is weakly declined posteriorly. The eye is row is deep, wide, straight, slightly divergent high, less than one-third glabellar length forward. Glabellar length (sag., excluding (Large Eye Index 31%). Its anterior edge is 2003 CARVALHO ET AL.: CALMONIID TRILOBITES FROM BOLIVIA 11

Fig. 4. Wolfartaspis liebermani, n. sp. Icla Formation, Kochis, Cochabamba Department. All scales 5 mm. A, holotype MHNC 8132, dorsal view of cephalon, latex cast from external mold; B, C, dorsal and lateral views of pygidium, AMNH 47150, latex cast from external mold; D, dorsal view of thorax and 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. 12 AMERICAN MUSEUM NOVITATES NO. 3407

Fig. 5. Wolfartaspis liebermani, n. sp. Icla Formation, Kochis, Cochabamba Department. All scales 5 mm. A±C, dorsal, lateral, and anterior views of cephalon and part of ®rst thoracic segment, AMNH 47403, internal mold; D±F, dorsal, anterior, and lateral views of cephalon, AMNH 47149, internal mold. opposite the anterolateral corner of L3, and pebral lobe; the palpebral furrow is moder- its posterior edge is opposite the posterolat- ately deep. The lateral cephalic border fur- eral margin of L2 and at a considerable dis- row is faint. The posterior cephalic border is tance from the posterior cephalic border fur- narrow proximally, about equal in width row (approximately 4 mm in the holotype). (exsag.) to the posterior border furrow at the In anterior view, the visual surface is straight fulcrum, and widened distally; none of the from top to bottom, gradually becoming wid- available specimens preserves the genal an- er toward its base, with 25±26 dorsoventral gle. lens ®les, and a maximum of 10 lenses per The hypostome is preserved in situ be- ®le. The interocular ®xigena slopes upward neath the cephalon in one specimen (®g. 4F). as it approaches the narrow, C-shaped pal- Its anterior margin is largely missing, but its 2003 CARVALHO ET AL.: CALMONIID TRILOBITES FROM BOLIVIA 13 course against the moderately wide anterior pygidial posterior margin. There are 10±11 wings suggests a transverse hypostomal su- axial rings, but the last few are indistinct; the ture. The middle body is roughly circular and ®rst ®ve rings are gently arched anteriorly, gently convex (tr., sag.). The maculae are but the subsequent ones are transverse or strong, forming a pair of ovoid protuberances weakly bowed backward. The axial rings are positioned abaxially, near to the junction of separated by wide ring furrows that become the faint middle furrow and the lateral border progressively narrower posteriorly; the ®rst furrow, anterior to the midlength of the mid- ®ve ring furrows have narrow, groovelike dle body. The border furrow is obscure an- apodemal pits distally. The pleural ®eld in- terolaterally (against the wings), becoming cludes six pairs of strongly furrowed ribs moderately impressed at the macula, main- plus a small seventh pair that parallels the taining moderate depth and width posterolat- axis. The pleural furrows are deep and wide, erally and posteromedially. The lateral bor- though gradually narrowing on more poste- der is narrow, with the lateral margins gently rior segments; they are moderately curved. converging between the anterior wing and The interpleural furrows are narrow, moder- ®rst of two pairs of short, sharp posterolat- ately deep proximal to the fulcrum between eral marginal spines; the inner pair of spines the anterior few ribs, progressively shallow (on posterior border) is separated by slightly between more posterior ribs such that ®fth greater than the distance between the two interpleural furrow is last distinct one on ex- spine pairs. The posterior border is fairly ternal mold. The lateral margin of the pygid- long and gently convex (sag.); the postero- ium is not well preserved in any of the ma- median margin is faintly convex backward terial (®g.4E); the ®fth lappet is small, ¯at, between the inner pair of marginal spines. and wide (®g.4B), with a curved anterior The thorax consists of 11 segments. The margin and straight posterior margin. The relative width of the axial rings and thorax posteromedian margin of the pygidium is an cannot be determined, as the distal ends of elongate, tapering, upturned spine (its basal the pleurae are not preserved in any speci- part is preserved in AMNH 47150; ®g. 4B, mens, but in AMNH 47148 (®g. 4D) the ax- C). ial rings appear to be wider than the pleural DISCUSSION: The monotypic Wolfartaspis ®elds. The axial furrow is shallow and nar- was erected by Cooper (1982) as a subgenus row, weakly diverging back to the sixth ring, of Metacryphaeus, but recent usage (Lieber- then gently converging. In lateral view, the man, 1993) recognizes it at the generic rank. axial rings are faintly convex, raised well According to Lieberman, Wolfartaspis dif- above the pleurae. The anteriormost axial fers from Metacryphaeus in having an en- rings are spatulate distally, but this is less larged occipital spine and an upwardly marked in the more posterior rings. Small but curved pygidial spine. The material de- distinct apodemal pits lie between the axial scribed here includes both of these features, rings abaxially. The pleural furrows are nar- and it is therefore assigned to the genus Wol- row and moderately impressed adjacent to fartaspis. We place our material in a new the axial furrow, becoming wider distally, species, based on the following differences with a straight, oblique course across the rib; from W. cornutus: (1) the ventral view of the the anterior margin of the pleural furrows is cephalon has a rounded shape in W. cornu- gently convex anteriorly, while the posterior tus, but is more pointed in our form; (2) the margin of the furrows is straight. Large Eye Index in W. cornutus is 34%, but The pygidium is triangular, moderately is less in our material (31%); (3) the occipital arched in lateral view, wider than long, with spine in W. cornutus (Lieberman, 1993: ®g. a pointed posterior tip. The axial furrow is 2.8) is stouter than in our material (®g. 5E, narrow and moderately deep. The axis is F) and does not taper from its base. Differ- slightly waisted, the axial furrow moderately ences are also noted in the stratigraphic lev- convergent against the ®rst ®ve axial rings, els of the two forms. Wolfartaspis cornutus weakly converging posterior to this; the axial occurs in the basal part of the Upper Member terminus is blunt, lacking impression of the of the BeleÂn Formation, in strata overlying axial furrow, lying well in advance of the the layer of Francovichia branisi (ϭ layer of 14 AMERICAN MUSEUM NOVITATES NO. 3407

Odontochile branisi sensu Wolfart, 1968, in that does not permit it to be placed in any of the upper part of Lower Member of BeleÂn the described calmoniid genera. We consider Formation). At Kochis, Francovichia occurs that the pygidium and the cephalon included stratigraphically above the beds with Wolfar- here belong to a single species. Both were taspis liebermani. collected at the same locality and stratigraphic level, they correspond in size, the pygidi- Granadocephalus, new genus um has appropriate proportions of the axis and pleurae relative to the cephalon, they DERIVATION OF NAME: Combination of have similar depth and width of the pleural type locality (Villa Granado, Cochabamba furrows on the thorax (®g. 6E) and pygidium Department) and ``cephalon''. (®g. 6F), and the pygidium cannot be as- TYPE SPECIES: Granadocephalus hanniba- signed to any other calmoniid taxon. Still, li, n. gen. and sp. since the association of the pygidium is not REFERRED SPECIES: Monotypic. certain, the generic diagnosis is restricted to DIAGNOSIS: Cephalon moderately convex cephalic characters. (tr.), subsemicircular in outline; ventral mar- The af®nities of Granadocephalus are dif- gin with moderate anterior arch. Pro®le of ®cult to determine. Of suprageneric groups glabella evenly curved, steepening anterior- identi®ed in previous work (Eldredge, 1979; ly; frontal lobe not distinctly separated from Eldredge and Ormiston, 1979; Eldredge and rest of glabella; posterior median impression BranisÏa, 1980), Granadocephalus lacks the weakly developed. Axial furrow moderately diagnostic characters of either the Probolops impressed, divergent especially from S2, group or the Malvinella subgroup, and mem- with small, rounded fossular depression near bership in the Calmonia group is most prob- anteromedial corner of palpebral lobe. Gla- able. In the Metacryphaeus group, the only bellar furrows not reaching axial furrow on obvious comparison is with the topologically internal mold; S3 lightly impressed, narrow, primitive genus Plesioconvexa Lieberman, faintly sinuous; S2 shallow but deeper than 1993, which has an evenly convex glabellar S3, weakly directed backward at its inner tip; pro®le reminiscent of Granadocephalus S1 deeper, weakly concave anteriorly, direct- (compare Plesioconvexa praecursor in Lie- ed anteromedially; S0 with strongly convex berman, 1993: ®g. 3.8 with G. hannibali, anterior margin. L3 and L2 not in¯ated; L1 here ®g. 6C). However, Plesioconvexa, like about two- thirds length (exsag.) of L2. L0 other members of the Metacryphaeus group, slightly shortened abaxially. Anterior section has much deeper incision of S2 and S3 than of facial suture not transecting anterolateral does Granadocephalus.InPlesioconvexa,S2 corner of glabella. Palpebral area slopes has the characteristic transverse, apodemal downward abaxially, lying entirely below in- impression shared by other members of the terocular ®xigena; palpebral furrow weak. Metacryphaeus group. Furthermore, Plesio- Librigenal ®eld relatively strongly pitted. convexa, and indeed all members of the Me- Posterior cephalic border furrow straight. En- tacryphaeus group except for deeply nested, tire cephalon covered by very ®ne granular highly modi®ed taxa such as Vogesina, has ornament on internal mold. a much deeper palpebral furrow than does DISCUSSION: We were hesitant to create a Granadocephalus. Character states such as new genus and species on the basis of such light impression of S2 and S3, the former a limited sample. However, the material dis- being weakly convex forward and the latter plays a unique combination of characters sinuous, as well as a shallow palpebral fur- within the Calmoniidae (e.g., ventral margin row, are shared by Granadocephalus and with moderate anterior arch; a peculiar con- members of the Calmonia group. These sim- vex [sag.] pro®le of the glabella; PMI weakly ilarities, however, may be primitive charac- developed; distinctive shape and develop- ters for Calmoniidae. Eldredge and BranisÏa ment of the glabellar furrows and lobes; an- (1980) noted that axial morphology of the terior section of the facial suture does not Calmonia group displays conservative char- transect the frontal lobe; visual surface low, acters for Calmoniidae, whereas the various lying entirely below the interocular ®xigena) genera of that group are more variable in 2003 CARVALHO ET AL.: CALMONIID TRILOBITES FROM BOLIVIA 15

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 articulated 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 of pygidium, MHNC 9451, internal mold. their features of the exoskeletal margin (e.g., on with a few displaced, articulated thoracic genal spines, thoracic pleural spines, pygidial segments, MHNC 8131 (cast AMNH 47150) marginal spines). The state of these charac- (®g. 6A±E). Paratype: internal mold of py- ters is not known for Granadocephalus. gidium, MHNC 9451 (®g. 6F, G). Both from the Icla Formation, layer of Francovichia Granadocephalus hannibali, new species branisi (Emsian, inferring equivalence to Figure 6A±G layer of F. branisi sensu Wolfart (1968) in DERIVATION OF NAME: The species name the BeleÂn Formation in the Altiplano), on a expresses our appreciation to Dr. Joseph T. hillside 5 km east of Villa Granado, Cocha- Hannibal (Cleveland Museum of Natural bamba Department, Bolivia, near the high- History), who has made signi®cant contri- way leading from Aiquile to Santa Cruz de butions to the knowledge of Bolivian pale- la Sierra. ontology, and who collected the holotype. DESCRIPTION: The cephalon is moderately DIAGNOSIS: As for genus. arched (tr.), subsemicircular in outline. Its TYPES: Holotype: internal mold of cephal- maximum width is 45 mm (across the pos- 16 AMERICAN MUSEUM NOVITATES NO. 3407 terior border) and its sagittal length is 30 well preserved, is relatively small, with the mm. The ventral margin is moderately visual surface about 8 mm long and not el- arched anteriorly, with at least two pairs of evated, lying entirely below the interocular low, widely spaced protuberances. The an- ®xigena; midlength of the palpebral furrow terior cranidial border is uniformly narrow, is opposite S2, its posterior edge opposite the set off by a sharp anterior cranidial border posterolateral corner of L2. The anterior edge furrow. The anterior cephalic border is also of the palpebral lobe is very close to the axial narrow, especially medially. The axial furrow furrow, but the distance between them in- is moderately impressed, moderately wide, creases posteriorly such that the posterior divergent forward especially from S2, with a edge of the eye is equidistant between the small, rounded fossular pit situated opposite axial furrow and the posterior cephalic bor- S3 (exsag.) and near the anteromedial corner der furrow. The librigenal ®eld slopes steeply of the eye. The anterior section of the facial down to the lateral cephalic border furrow, suture circumscribes the frontal glabellar with the ®eld bearing numerous strong pits. lobe, passing around its anterolateral corner The lateral border of the cephalon and the without transecting it. Glabellar length (sag.) genal angle are not preserved. The posterior is 25 mm (excluding S0) and its maximum cephalic border furrow is narrow, deep, and width is 27 mm. In longitudinal pro®le, the transverse; its anterior margin is straight. The glabella is curved, without a break in slope posterior border gradually widens (exsag.) between the frontal lobe and S0 medially; the distally, more strongly widening distal to the frontal lobe is steepest anteriorly. The gla- fulcrum. The cephalon is ornamented with bellar median region is almost ¯at (tr.). The ®ne granulation, which is especially dense on posterior median impression (PMI) is weakly the glabella. developed as a faint elongate groove that ex- Only an internal mold of a laterally incom- tends back nearly opposite the inner tips of plete pygidium is known. The axial furrow S3. The three pairs of glabellar furrows are is shallow, narrow, and rather strongly con- not connected with the axial furrow. S3 is verging posteriorly. The axis appears to be narrow, lightly impressed, and faintly sinu- slightly wider than the inferred width of the ous; S2 is shallow but deeper than S3, weak- pleurae (excluding marginal spines, if pre- ly convex anteriorly, in part by a gentle pos- sent). In lateral view the axis is gently raised terior curvature at its inner tip; the overall above the pleural ®eld. Five axial rings are course of S2 is approximately transverse, ef- strongly developed, plus parts of a sixth. The facing well inward of the axial furrow; S1 is ®rst two rings are moderately arched anteri- deeper impressed as gently concave, antero- orly and the succeeding ones are more gently medially directed apodemal grooves, abrupt- arched. Medially the rings are widely sepa- ly shallowing distally, at most faint imme- rated from each other, with a pseudoarticu- diately adjacent to axial furrow. S0 is well lating half ring in the ®rst two ring furrows; developed, with a strongly convex anterior these rings are shortest sagittally and distinct- margin and a gently convex posterior margin, ly spatulate distally. Apodemal pits are de- such that S0 is longest sagittally; medial part veloped in the distal parts of the ring fur- of S0 moderately deep, distal part incised as rows. The axis terminates far in advance of faintly concave apodemal grooves that par- the posteromedian margin of the pygidium, allel apodemes in S1. L2 and L3 lack inde- with the postaxial region de®ned by a change pendent in¯ation; L3 is wedge-shaped, ex- in convexity rather than distinct impression panded distally; L2 is more conspicuous than of the axial furrow posteromedially. Five L3, and slightly narrowed medially. L1 is pleurae are preserved on the right side, but about two- thirds the length (exsag.) of L2. only parts of four are present on the left. The L0 is of even length across much of its pleurae are gently arched (tr.), the ribs width, with slight shortening distally, behind lengthening abaxially; the anteriormost ribs the apodemal part of S0; its width is 15 mm. are gently and evenly curved laterally, but The gena is subtrigonal, moderately convex. the last ones bend back more abruptly. The The palpebral area slopes downward laterally ®rst interpleural furrow is narrow but sharply to the border furrow. The eye, though not impressed, the succeeding furrows are simi- 2003 CARVALHO ET AL.: CALMONIID TRILOBITES FROM BOLIVIA 17 larly narrow, but shallow; the fourth is the bites from the Argentine Precordillera; new last discernible interpleural furrow on the in- taxa of the Bouleia group, and remarks on the ternal mold. The pleural furrows are deep tempo of calmoniid radiation. Geological Mag- and relatively wide, maintaining their im- azine 131: 449±464. pression to the inner edge of the doublure. Eldredge, N. 1979. Cladism and common sense. In J. Cracraft and N. Eldredge (editors), Phy- The lateral and posterior margins are not pre- logenetic analysis and paleontology: 165±198. served, though the even outer margin of the New York: Columbia University Press. doublure beneath the fourth and ®fth ribs Eldredge, N., and L. BranisÏa. 1980. Calmoniid tri- suggests that marginal spines or lappets were lobites of the Lower Devonian Scaphiocoelia probably lacking on those segments at least. Zone of Bolivia, with remarks on related species. Bulletin of the American Museum of Nat- ACKNOWLEDGMENTS ural History 165: 181±289. Eldredge, N., and A.R. Ormiston. 1979. Bioge- Yongyi Zhen (Australian Museum) photo- ography of Silurian and Devonian trilobites of graphed the material and assembled the plates. the Malvinokaffric Realm. In J. Gray and A. J. For assistance in the ®eld we thank Joseph T. Boucot (editors), Historical biogeography, plate Hannibal (Cleveland Museum of Natural His- tectonics, and the changing environment: 147± tory), Salvador de la Serna (Cochabamba, Bo- 167. Corvallis: Oregon State University Press. livia), and Mr. Angel. Susan Klofak (AMNH) Isaacson, Peter E. 1977. Devonian stratigraphy and brachiopod paleontology of Bolivia. Part and Gary Chilson (Plantation, Florida) assist- A. Orthida and Strophomenida. Palaeontogra- ed with preparation and casting of specimens. phica Abteilung A Palaeozoologie-Stratigraphie Suggestions by the reviewers are appreciated. 155: 133±192. M.G.P.C. thanks John Maisey (AMNH) for Kozlowski, R. 1923. Faune deÂvonienne de Boli- his support and encouragement, and Adriana vie. Annales de Paleontologie 12: 1±112. Aquino (AMNH) for helping with the Spanish Lieberman, B.S. 1993. Systematics and biogeog- abstract. raphy of the ``Metacryphaeus group'' Calmon- iidae (Trilobita, Devonian), with comments on REFERENCES adaptive radiations and the geological history of the Malvinokaffric Realm. Journal of Pale- Adrain, J.M., and G.D. Edgecombe. 1996. De- ontology 67: 549±570. vonian aulacopleurid trilobites of the Malvino- Lieberman, B.S., G.D. Edgecombe, and N. Eld- kaffric Realm. Geobios 29: 417±436. redge. 1991. Systematics and biogeography of Blieck, A, P.-Y. Gagnier, F.P. Bigey, G.D. Edge- the ``Malvinella group'', Calmoniidae (Trilobi- combe, P. Janvier, S. Loboziak, P.R. Rache- ta, Devonian). Journal of Paleontology 65: boeuf, T. Sempere, and P. Steemans. 1996. New 824±843. Devonian fossil localities in Bolivia. Journal of Richter, R., and E. Richter. 1942. Die Trilobiten South American Earth Sciences 9: 295±308. der Weismes±Schichten am Hohen Venn, mit BranisÏa, L. 1965. Los foÂsiles guõÂas de Bolivia I. Bemerkungen uÈber die Malvino-caffrische PaleozoÂico. Servicio Geologico de Bolivia, Provinz. Senckenbergiana 25(1/3): 156±179. Boletin 6: 1±282. Smith, L., and G.D. Edgecombe. 1996. Sinopsis Clarke, J.M. 1913. FoÂsseis devonianos do ParanaÂ. descriptiva de cuatro nuevos calmoniidos, Tri- ServicËo GeoloÂgico e MineraloÂgico do Brazil. lobita, DevoÂnico. Memorias del XII Congreso Monogra®a 1: 1±353. Geologico de Bolivia. Tarija, Bolivia: 321±322. Cooper, M.R. 1982. A revision of the Devonian Whittington, H.B., and S.R.A. Kelly. 1997. Mor- (Emsian-Eifelian) Trilobita from the Bokkeveld phological terms applied to Trilobita. In R. Group of South Africa. Annals of the South Kaesler (editor), Treatise on invertebrate pale- African Museum 89: 1±174. ontology. Part O. Arthropoda 1. Trilobita, re- d'Orbigny, A. 1842. Voyage dans l'Amerique vised: O313±O329. Lawrence: University of meÂridionale, 1826±1833, 2, 4 (PaleÂontologie). Kansas Press. Paris: Bertrand. Wolfart, R. 1968. Die Trilobiten aus dem Devon Edgecombe, G.D., and R.A. Fortey. 2000. Siluri- Boliviens und ihre Bedeutung fuÈr Stratigraphie an trilobites from the El Carmen Formation, und Tiergeographie. In R. Wolfart and A. Vo- Bolivia. Senckenberg. Lethaea 79: 329±355. ges (editors), BeitraÈge zur Kenntnis des devon Edgecombe, G.D., N.E. Vaccari, and B.G. Wais- von Bolivien. Beihefte zum Geologischen Jahr- feld. 1994. Lower Devonian calmoniid trilo- buch 74: 5±241. Recent issues of the Novitates may be purchased from the Museum. Lists of back issues of the Novitates and Bulletin published during the last ®ve years are available at World Wide Web site http://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]

a This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper).