A Middle Devonian Trilobite Assemblage from Venezuela

AMNH NOVITATES Tuesday Dec 11 2001 10:21 AM 2000 novi 00147 Mp_1 Allen Press • DTPro System File # 01cc

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PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, NY 10024 Number 3292, 15 pp., 5 ®gures March 16, 2000

MARIA DA GLORIA PIRES DE CARVALHO1 AND JOHN MOODY2

ABSTRACT

Three new species of trilobites are described from the upper level of the CanÄo del Oeste Formation (Middle Devonian) of Venezuela. These taxa include a phacopinine, Viaphacops venezuelensis, n. sp., and two asteropyginines, Rhenops odremani, n. sp., and Greenops perijaensis, n. sp. The biogeographic implications of these records are discussed. Viaphacops is somewhat cosmopolitan in its distribution. The presence of Rhenops in Venezuela represents a profound range extension beyond the Rhenish-Bohemia region of the Old World Realm. The Venezuelan occurrence of Greenops represents a southerly range extension within the Eastern Americas Realm, reinforcing faunal similarities previously recognized among Devonian brachiopods and trilobites of Venezuela and North America.

RESUMEN

Se describen tres especies nuevas de trilobites provenientes del nivel maÂs alto de la For- matioÂn CanÄo del Oeste (DevoÂnico medio) de Venezuela. Estos taxones incluyen un facopinino, Viaphacops venezuelensis, n. sp., y dos asteropigininos, Rhenops odremani, n. sp. y Greenops perijaensis, n. sp. Se discuten las implicancias biogeograÂ®cas de estos registros. Viaphacops tiene una distribucioÂn amplia. La presencia de Rhenops en Venezuela representa una extensioÂn signi®cativa de su distribucioÂn maÂs allaÂ de la regioÂn Rheno-Bohemia del Viejo Mundo. El registro de Greenops en Venezuela extiende su distribucioÂn hacia el sur dentro del Dominio del Este de las AmeÂricas, apoyando las similtudes reconocidas previamente, entre los bra- quioÂpodos y los trilobites devoÂnicos de Venezuela y AmeÂrica del Norte.

1Research Associate, Division of Paleontology, American Museum of Natural History. 2Professor, SeccioÂn de PaleontologõÂa, Museo de BiologõÂa, Universidade del Zulia, Facultad de Ciencias, Apartado 526, Maracaibo 4011, Zulia, Venezuela.

Copyright ᭧ American Museum of Natural History 2000 ISSN 0003-0082 / Price $1.90 AMNH NOVITATES Tuesday Dec 11 2001 10:21 AM 2000 novi 00147 Mp_2 Allen Press • DTPro System File # 01cc

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INTRODUCTION either destroyed in a ®re at MeÂrida or was lost during its relocation to Caracas, accord- The Devonian marine invertebrate fauna ing to Venezuelan ministry of®cials. from Venezuela has been known since the In the present work, new species of Via- beginning of the 20th century (Yeakel, 1924; phacops, Greenops, and Rhenops are de- Weisbord, 1926; Liddle et al., 1943), follow- scribed. This material was collected by one ing early expeditions to collect fossils from of us (J. M.) and is housed in the Museo de the type locality of the CanÄo del Oeste For- BiologõÂa, Universidad de Zulia (MBLUZ), mation (Middle Devonian; Eifelian±Give- Maracaibo, Venezuela. Casts of this material tian), which extends along the CachirõÂ river are also deposited in the fossil invertebrate in the PerijaÂ Mountains. Benedetto (1979, collection of the American Museum of Nat- 1984) subsequently described assemblages of ural History. pelecypods, trilobites, and brachiopods that The presence of Greenops in the Devonian were collected from several exposures of the of Venezuela supports the proposed biogeo- CanÄo del Oeste Formation in the area of graphic connection with the Eastern Ameri- CanÄo Colorado (®g. 1). The trilobites de- cas Realm, and the presence of Rhenops in scribed in the present work were also col- the Venezuelan assemblage suggests a link lected from this region. with the Rhenish-Bohemian subprovince of The ®rst Devonian trilobite described from Europe and Morocco (part of the Old World Venezuela (a fragmentary specimen lacking Realm; Boucot, 1988). Viaphacops is more the cephalon, deposited in the Cornell Uni- widespread and occurs in three different bio- versity Paleontological Laboratory) was re- geographic realms. The biogeographic sig- ferred to Phacops argentinus Thomas (Weis- ni®cance of these trilobites is discussed fur- bord, 1926). Eldredge and Ormiston (1979: ther below. 148) suggested that it may belong to Via- phacops, and according to Benedetto (1979: LOCALITIES AND STRATIGRAPHY 85) Weisbord's material was probably collected from the CanÄo Grande Formation, The CanÄo del Oeste Formation has long which underlies the CanÄo del Oeste Forma- been considered to be of Middle Devonian tion and is probably of Lower Devonian age. age (Barret and Isaacson, 1988; Benedetto, Additional trilobites from CanÄo del Oeste 1979, 1984; Weisbord, 1926). This forma- Formation were described by Benedetto tion, ®rst described by Liddle et al. (1943), (1979), who identi®ed a phacopinine (Pha- represents the middle beds of the Rio CachirõÂ cops sp. cf. P. rana; ibid., 96±97, pl. 4, ®gs. Group (Sutton, 1936), and consists princi- 1±7) and two asteropyginines (Greenops sp., pally of ®ne-grained, micaceous, quartzitic plus gen. and sp. indet.; ibid., 97±98, pl. 4, sandstones, siltstones, and shales. In the type ®gs. 8, 9). As discussed below, Benedetto's section, the formation may be up to 1000 m (1979) phacopinine apparently differs in sev- in thickness, but in the area from which the eral respects from the material described here present material was collected (a few kilo- as Viaphacops. Thus, the assemblage prob- meters south of the type section), it is less ably contains at least two different phacopi- than 400 m thick. The contact with the un- nines, although only Viaphacops is repre- derlying CanÄo Grande Formation (Early De- sented in the new sample (the identity of vonian) is faulted (Liddle et al., 1943; Hea Weisbord's original specimen is left an open and Whitman, 1960; Benedetto, 1984). The question). Two asteropyginine taxa are also Campo Chico Formation (Upper Devonian) described here, but it is not possible to com- appears to be transitional from a near-shore pare these with Benedetto's examples be- marine environment to one that is dominated cause critical characters cannot be deter- by terrestrial deposits (Benedetto, 1984; Ber- mined from his descriptions or illustrations. ry et al., 1993). Unfortunately, no direct comparison with any All of the specimens were collected from of his original material (which was deposited three localities along old and new parts of in the Ministerio de EnergõÂa y Minas collec- the Rio Socuy Road, between 1.2 and 2.5 km tion) is possible because that material was north of Hacienda El Reposo, and along the AMNH NOVITATES Tuesday Dec 11 2001 10:21 AM 2000 novi 00147 Mp_3 Allen Press • DTPro System File # 01cc

2000 DE CARVALHO AND MOODY: MIDDLE-DEVONIAN TRILOBITES 3

Fig. 1. Locality map of the CanÄo Colorado Area, PerijaÂ Mountains, Zulia State, Venezuela, with insert map of northern South America. Approximate limit of PerijaÂ Mountains indicated by stronger dashed line.

high, eastern ¯ank of the upper valley from along the old road correspond to Level formed by CanÄo Colorado, a northern tribu- Co7 of Benedetto (1984), whereas the new tary of Rio Palmar in the PerijaÂ Mountains, road collecting locality, which was exposed Zulia, Venezuela (®g. 1). The collections after Benedetto made his collections, proba- AMNH NOVITATES Tuesday Dec 11 2001 10:21 AM 2000 novi 00147 Mp_4 Allen Press • DTPro System File # 01cc

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bly represent beds of his Level Co4, and pos- sibly slightly higher (®g. 2). Fragmentary trilobites have also been collected from additional beds of the CanÄo del Oeste Formation between Levels Co4 and Co7, so trilobites are probably present throughout the upper section. Morphological terminology follows that used in the Treatise on Invertebrate Pale- ontology, Part O (Struve, 1959), Eldredge (1972, 1973), and Lieberman and Kloc (1997).

SYSTEMATIC PALEONTOLOGY

CLASS TRILOBITA WALCH, 1771 ORDER PHACOPIDA SALTER, 1864 SUBORDER PHACOPINA STRUVE, 1959 SUPERFAMILY PHACOPOIDEA HAWLE AND CORDA, 1847 FAMILY PHACOPIDAE HAWLE AND CORDA, 1847 SUBFAMILY PHACOPINAE HAWLE AND CORDA, 1847 Genus Viaphacops Maximova, 1972

TYPE SPECIES: Phacops pipa Hall and Clarke, 1888; early Middle Devonian of North America (New York State, Michigan and Southern Ontario). GENERIC DIAGNOSIS (Maximova, 1972): Preoccipital lobe entirely reduced; preoccipital furrow merged with occipital furrow to form a deep broad furrow on site of preoccipital lobe. Occipital ring usually narrow. Vincular furrow deep, with prominent inner border. Eyes of moderate size, less frequently large. Pygidium usually wide, weakly seg- mented. Surface of glabella with fairly large nodes. SYSTEMATIC NOTE: Those phacopids that were originally placed in the genus Phacops from Central Kazakhstan and the Far Eastern part of the former Soviet Union were reex- amined by Maximova (1972), who erected a new genus, Paciphacops, with two subgen- era, P. (Paciphacops) and P. (Viaphacops), each containing several species. Elsewhere Fig. 2. Geological column for the CanÄo del Viaphacops has been elevated to generic Oeste Formation, CanÄo Colorado Area, PerijaÂ Mountains, Zulia, Venezuela (after Benedetto, rank (e.g., RamskoÈld and Werdelin, 1991; 1984), showing the approximate stratigraphic lev- Linsley, 1994) and that practice is also fol- els of the trilobites described in this study. lowed here. AMNH NOVITATES Tuesday Dec 11 2001 10:21 AM 2000 novi 00147 Mp_5 Allen Press • DTPro System File # 01cc

2000 DE CARVALHO AND MOODY: MIDDLE-DEVONIAN TRILOBITES 5

Fig. 3. Viaphacops venezuelensis, n. sp., holotype, MBLUZ P-33, internal mold of cephalon. (A) Dorsal view; (B) frontal view; (C) lateral view; (D) oblique dorsolateral view of right side.

Viaphacops venezuelensis, new species AGE: Middle Devonian (Eifelian/Give- Figure 3, A±D tian). DESCRIPTION: The cephalon is approxi- DIAGNOSIS: Cephalon broader than long, mately twice as broad as long, with a maxi- with posterior border slightly concave. Com- mum cephalic width of 34 mm (adjacent to posite glabellar lobe short and broad, rounded anteriorly, ¯attened dorsally, with mod- genal angles), and a maximum length of 16 erate inclination anteriorly. Glabellar surface mm (sagittally from front of glabella to pos- ornamentated with low, rounded tubercles, terior margin of occipital ring). Dorsally the each one somewhat separated from the oth- glabella is moderately in¯ated, ¯attened, and ers, and gradually diminishing in size ante- inclined forward, and anteriorly its frontal riorly. Occipital ring without spines or tu- surface is vertical. The length (sag.) of the bercles. Genal angles rounded. Eyes with 13 composite glabellar lobe is 12 mm (formed or 14 dorsoventral ®les, with 3 or 4 lenses by the union of the anterior, third, second, per ®le. Visual surface convex, raised some- and medial portion of the ®rst glabellar what above glabellar level. lobes; Eldredge, 1973: 316). The composite HOLOTYPE: MBLUZ P-33, cephalon (rep- glabellar lobe is covered by low, rounded tu- resented by an internal mold), from upper bercles, each one somewhat separated from level of the CanÄo del Oeste Formation, PerijaÂ the others and diminishing gradually in size Mountains, NW Venezuela. Cast of holotype toward the anterior region of the glabella. AMNH 46578. Tubercles are not present on the occipital ETYMOLOGY: Named after Venezuela, the ring or genae. The maximum anterior gla- country of origin. bellar width at the intersection of the axial AMNH NOVITATES Tuesday Dec 11 2001 10:21 AM 2000 novi 00147 Mp_6 Allen Press • DTPro System File # 01cc

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furrows and anterior cephalic margin equal broad than long in V. venezuelensis); shape 20 mm. The posterior glabellar width is 10 of the composite glabellar lobe (pentagonal mm. The axial furrows are straight, broad, in V. nasutus, about 2/3 as long as wide, with deep, and strongly divergent anteriorly. The a produced anterior region; short and broad second and third glabellar furrows are shal- in V. venezuelensis); and genal angles (short, low, almost indistinct. A deep, broad trans- with a nodose genal spine in V. nasutus; glabellar furrow is apparently formed by rounded in V. venezuelensis). In both species con¯uence of the occipital furrow (S0) and the eyes are slightly higher than the glabella. ®rst pair of glabellar furrows (S1). All the Viaphacops canadensis (Stumm, 1954) paired glabellar lobes are incorporated into differs mainly in the shape of the genal an- the composite glabellar lobe. The distal por- gles (narrow, with sharp genal spine, angled tion of the ®rst glabellar lobe (L1) forms a obliquely upward in V. canadensis; rounded pair of lateral preoccipital lobes. The visual in (V. venezuelensis) and eye size (smaller in surface is supported by an eye socle, elevat- V. canadensis than in V. venezuelensis). Sim- ing the eye somewhat above the level of the ilarities between these two species include glabella. The palpebral lobes are well devel- the occipital ring, which lacks a spine or oped but are not so high as the visual surface. node, ornamentation of the glabella, and the The eyes have 13 or 14 dorsoventral ®les on shape of the cephalon. the visual surface (the posteriormost ®les are Viaphacops cristatus (Hall, 1861) differs somewhat indistinct, but there are certainly from V. venezuelensis in having the glabella no more than 14 ®les). In each ®le the num- more in¯ated and in possessing an occipital ber of lenses is no more than four lenses per spine and large genal spines. ®le (in the anteriormost two ®les there are Viaphacops orurensis (Bonarelli, 1921) only two or three lenses). The genae are al- from Bolivia differs from V. venezuelensis in most vertical and extend posteriorly behind the shape of the glabella (pentagonal and the occipital ring. The genal angles are strongly in¯ated in V. oururensis; moderately rounded. The occipital ring is narrow (long.), in¯ated and ¯attened in V. venezuelensis), without ornamentation or spine, slightly the form of the genal angles (with short curved anteriorly (sag.), slightly above the spines in V. oururensis; rounded in V. vene- glabellar lobe, with a width of 10 mm. The zuelensis), and the number of dorsoventral posterior margin of the cephalon and the oc- ®les and lenses per ®le in the visual surface cipital ring are both raised to form a promi- (17/5-7 in V. oururensis; 13 or 14/4 in V. nent margin. venezuelensis). The number of dorsoventral The thorax and pygidium are unknown. ®les is somewhat higher in V. oururensis than in all other Viaphacops spp. Both spe- MORPHOLOGICAL COMPARISON cies have the occipital ring narrow and raised WITH OTHER VIAPHACOPS SPECIES above the glabella. Three other species from Bolivia have also The specimen studied here differs from been referred to Phacops (Viaphacops)by Viaphacops pipa (Hall and Clarke, 1888; Pek and VaneÏk (1991). All three of these taxa type species of the genus) in the shape of its (V. multicinctus, spatiosus, and kozlowski) composite glabellar lobe (less in¯ated and show some differences from V. venezuelen- not protuberant in V. venezuelensis), genal sis, especially in the shape of the frontal gla- angles (rounded in V. venezuelensis; short, bellar lobe and the genal spines. A review of blunt spines in V. pipa), and eyes (above gla- Viaphacops from Bolivia is currently in bellar level in V. venezuelensis; below gla- preparation by Y. Iwasaki and no further bellar level in V. pipa). In both species, how- comparison with this material will be made ever, the eyes are set well forward on the here. cephalon a considerable distance from the posterior cephalic margin. REMARKS ON BENEDETTO'S (1979) Viaphacops nasutus (Stumm, 1954) differs PHACOPS SP. CF. P. RANA from V. venezuelensis in the shape of the Although the phacopine material referred cephalon (subtriangular in V. nasutus; more to Phacops sp. cf. rana by Benedetto (1979) AMNH NOVITATES Tuesday Dec 11 2001 10:21 AM 2000 novi 00147 Mp_7 Allen Press • DTPro System File # 01cc

2000 DE CARVALHO AND MOODY: MIDDLE-DEVONIAN TRILOBITES 7

was unavailable for comparison, from his of cephalon lacking the left genal spine, plus published description it can be excluded parts of ®rst three thoracic segments. Cast of from the genus Viaphacops by the following holotype AMNH 46579. Dimensions of the characters: (a) presence of preoccipital lobe holotype: cephalic length (sag.) approximate- (absent in Viaphacops); (b) strongly rounded ly 12.6 mm from occipital furrow to anterior glabella covered with dense tubercles almost margin of cephalon; width approximately con¯uent with each other (less rounded gla- 22.2 mm across posterior margin of cepha- bella, with fewer and more widely spaced tu- lon. bercles in Viaphacops); (c) 18 dorsoventral PARATYPES: MBLUZ P-985, internal mold ®les on the visual surface (only 13±14 in of pygidium; MBLUZ P-986, internal mold Viaphacops). of cephalon with part of the right genal spine, It is concluded that Benedetto's (1979) plus ®rst three thoracic segments; MBLUZ phacopine material does not represent the P-1079, small enrolled specimen with the py- same taxon we have, and there may be at gidium preserved (not ®gured). Casts of pa- least two different phacopine taxa represent- ratypes AMNH 46580, 46581, 46582. All ed in this assemblage. Weisbord's (1926) material of Rhenops odremani n. sp. is phacopine specimen lacks the cephalon and housed in the Museo de BiologõÂa, Universi- cannot be compared with our material. dad de Zulia (MBLUZ), Maracaibo, Vene- zuela, not in the Department of Geology, SUPERFAMILY ACASTOIDEA DELO, 1935 University of Rio de Janeiro, Brazil, as was FAMILY ACASTIDAE DELO, 1935 stated by Lieberman and Kloc (1997: 77). SUBFAMILY ASTEROPYGINAE DELO, 1935 Catalog numbers for the specimens illustrat- ed in Lieberman and Kloc (1997: ®g. 15) are Genus Rhenops Richter & Richter, 1943 as follows: Nos. 1, 2 ϭ MBLUZ P-1079; TYPE SPECIES: Cryphaeus anserinus Rich- Nos. 3±6 ϭ MBLUZ P-986; Nos. 7, 8 ϭ ter, 1916; Lower Devonian, Eifel (Germany). MBLUZ P-985. GENERIC DIAGNOSIS: See Struve, 1959: ETYMOLOGY: Named for Oscar Odreman, O483. who has made signi®cant contributions to the REMARKS: This genus is known from the paleontology of Venezuela. Lower Devonian (Emsian) of Europe (Ger- PROVENANCE AND AGE: CanÄo del Oeste many, France, and Spain) and Middle De- Formation, Middle Devonian (Eifelian/Gi- vonian (Eifelian) of Africa (Morocco). The vetian), PerijaÂ Mountains, NW Venezuela. presence of Rhenops in the Devonian of Ven- DESCRIPTION: The cephalon has a relatively ezuela was recently noted by Lieberman and elongate outline, tapering anteriorly, and thus Kloc (1997). There are still no records of this resembles a gothic arch (ogival shape), with genus in North America. the anterior cephalic border slightly pointed medially. The cephalic pro®le is moderately Rhenops odremani, new species convex. Axial furrows are narrow, well im- Figures 4, A±F; 5 A, B pressed, subparallel to S1 then becoming divergent anteriorly. Glabellar furrows are Rhenops, n. sp., Lieberman and Kloc, 1997: 76, weakly impressed and moderately wide. S1 ®g. 15, 1±8. is roughly crescent-shaped and deeper ad- DIAGNOSIS: Cephalon relatively elongate in axially; S2 is transverse, narrower than S1; outline, ogival, with anterior cephalic border S3 is faintly sinusoidal. Lateral glabellar slightly pointed medially; S3 faintly sinusoi- lobes (L1±L3) are ¯at sagittally. The ®rst dal, glabellar lobes not in¯ated; genal spines pair of lateral glabellar lobes (L1) is some- long and broad, slightly curved at their ex- what ¯exed forward, giving the appearance tremities, with a ridge on the dorsal surface of a transglabellar lobe; L2 is approximately extending from the cephalic posterior border rectangular and smaller than L3, which has to the tip. Eyes above glabellar level with a wedge shape. The frontal glabellar lobe is maximum of seven lenses per dorsoventral strongly expanded anteriorly and laterally, ®le. with its margin smoothly rounded in dorsal HOLOTYPE: MBLUZ P-987, external mold view, sloping gently anteriorly and laterally. AMNH NOVITATES Tuesday Dec 11 2001 10:21 AM 2000 novi 00147 Mp_8 Allen Press • DTPro System File # 01cc

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Fig. 4. Rhenops odremani, n. sp. (A±C) Cast made from holotype (an external mold), MBLUZ P- 987, in (A) dorsal view; (B) frontal view; (C) lateral view; (D-F) Paratype, MBLUZ P-986; internal mold of cephalon in (D) dorsal view; (E) frontal view; (F) lateral view.

The cephalic lateral border furrow is not dis- transverse line tangent to the anterior edge of cernible; the cephalic posterior border furrow the occipital ring. The visual surface has a is deep and ¯exed forward medially. The maximum of seven lenses per dorsoventral eyes are large and raised above the glabella, ®le. The palpebral region of the ®xigena occupying about 50% of glabellar length, slopes toward the glabella. Genal spines are and are positioned proximally and posteriorly relatively long and broad, curved at their ex- on the cephalon. Each eye is in contact with tremities, with a ridge on the dorsal surface the anterolateral corner of L3 anteriorly, and extending from the cephalic posterior border posteriorly is located a short distance from furrow to the tip. The internal margins of the the axial furrow. The posterior edge of the genal spines are not parallel to the sagittal visual surface is situated posteriorly to a line, but are de¯ected posteriorly at an angle AMNH NOVITATES Tuesday Dec 11 2001 10:21 AM 2000 novi 00147 Mp_9 Allen Press • DTPro System File # 01cc

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Fig. 5. (A, B) Rhenops odremani, n. sp. Paratype, MBLUZ P-985; internal mold of pygidium. (A) dorsal view; (B) lateral view. (C-E) Greenops perijaensis, n. sp., latex cast of external mold in the holotype, MBLUZ P-1448. (C) Dorsal view; (D) frontal view; (E) lateral view.

of approximately 15Њ. The length of the genal ent in two of the paratypes (MBLUZ P-985 spine cannot be determined accurately in any and MBLUZ P-1079). The following de- of the available specimens; it has approxi- scription is based mainly on MBLUZ P-985 mately the same length as the rest of the as the pygidium is better preserved than in cephalon. The occipital ring is not well pre- MBLUZ P-1079. The pygidium is small and served in the material and its morphology subelliptical in outline; in MBLUZ P-985 it cannot be determined. The preserved surface has a total length of approximately 5.6 mm of the cephalon is rough and has a granular (sag.) and maximum width about 9.6 mm (its appearance in places. In epoxy casts the sur- anteriormost region). In lateral view the pos- face appears to be tuberculose (®g. 4B), but teriormost region of the axis slopes to the this is an artifact of the casting medium. We rear. The axis is slightly raised above the have not found convincing evidence that tu- pleural ®eld, and includes at least 10 rings. bercles are present on the original specimen. The ®rst seven rings are easily distinguished, The pygidium is lacking in the holotype of but the remainder become progressively ob- Rhenops odremani, new species, but is pres- solete posteriorly and cannot be accurately AMNH NOVITATES Tuesday Dec 11 2001 10:21 AM 2000 novi 00147 Mp_10 Allen Press • DTPro System File # 01cc

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counted. The anterior margins of the ®rst ®ve nops, and their basic conclusions were not rings are slightly arched anteriorly; after the affected. ®fth ring they are straight. The axis tapers posteriorly as far as the ®fth ring, after which COMPARISON WITH OTHER RHENOPS the axial furrows are roughly parallel back to SPECIES the terminus of the axis. The pleural region consists of ®ve pairs of pleurae of approxi- The rounded anterior margin of the cep- mately equal width, and with equally elevat- halon and the position of the eyes (extending ed anterior and posterior pleural bands; the to the lateral border furrow of the cephalon) pleural furrows are deeper than the interpleu- distinguish Rhenops anserinus (Richter, ral furrows. The ®rst three pleurae are gently 1916) from the Venezuelan species. Tuber- curved posteriorly, and the last two are cles are present on the pygidial axial rings in curved more abruptly. The ®fth pleura is R. anserinus but are absent in Rhenops odre- much shorter than the others and it is very mani. close to the pygidial axis. Each pleura ter- Rhenops babini Morzadec, 1983 has a minates with a long, broad, and ¯at lappet, maximum of eight lenses per dorsoventral but its distal tips are not well preserved. The ®le on the visual surface, whereas rhenops external lateral margin of each lappet is odremani has a maximum of seven. The smoothly curved and the inner margin is ap- shape of the genal spine is also distinct; in proximately straight. The terminal lappet is R. babini the spine is developed medially as slightly broader than the posterior part of the a ¯attened ledge, while in the Venezuelan axis, but its terminal part is missing. No or- species there is a ridge along its surface. The namentation is present on the pleurae or py- ®fth pair of pygidial lappets is twice as long gidial axial rings. The only ornamentation as the other lappets in R. babini, but it is observed in these specimens is ®ne granu- roughly of the same length as the others in lation on the lappets, but other details are Rhenops odremani. Rhenops circumapodemus Smeenk, 1983 dif®cult to ascertain as both the available py- is distinguished from Rhenops odremani in gidia are weathered. having a wide ogival cephalon shape; L3 and L2 slightly swollen; a maximum of nine lens- CHARACTERIZATION OF es per dorsoventral ®le on the visual surface; VENEZUELAN RHENOPS BY long pygidium with 14 or 15 axial rings; ®ve LIEBERMAN AND KLOC (1997) pairs of sickle-shaped pygidial lappets, with the ®fth pair longer and more pointed than Lieberman and Kloc (1997) included the the others; terminal pygidial lappet reduced Venezuelan material described here in their to rounded posterior border. phylogenetic analysis of the subfamily As- In Rhenops index Richter and Richter teropyginae, which used 39 taxa and 66 char- 1943, the terminal pygidial lappet is longer acters. Some of their codings for this material than the other ®ve pairs (its length cannot be are considered questionable here (mostly for accurately determined in R. odremani, but it preservational reasons), including characters seems not to be elongated), and tubercles are 24, 26, 40, 42, 44, 57, and 58 (adult proso- present on the axial rings of the pygidium pon; length of genal spine; shape of pygidial (absent in R. odremani). lappet; lateral margins of pygidial lappets; Rhenops lethaeae (Kayser, 1889) has a distal tips of pygidial lappets; number of py- very distinct pygidial morphology, with two gidial axial rings; margin of terminal pygid- small rounded cavities on the ®rst ®ve py- ial lappet). We would also recode character gidial axial rings, located abaxially but not 39 (ornament on medial part of pygidial and in contact with the axial furrows (cavities ab- thoracic axial rings) from state 1 (tubercles) sent in R. odremani); furthermore, the distal to 0 (absent). Even after these changes, how- tips of the pygidial lappets are developed as ever, reanalysis using an identical procedure blunt triangles, but the shape of the tips of did not alter the phylogenetic position of the the pygidial lappets cannot be determined in Venezuelan material within the genus Rhe- Rhenops odremani. In both Rhenops anseri- AMNH NOVITATES Tuesday Dec 11 2001 10:21 AM 2000 novi 00147 Mp_11 Allen Press • DTPro System File # 01cc

2000 DE CARVALHO AND MOODY: MIDDLE-DEVONIAN TRILOBITES 11

nus (Richter, 1916) and R. lethaeae (Kayser, approximately the same size as their respec- 1889) the visual surface is expanded later- tive pleurae and in contact with each other ally, reaching the lateral border furrow of the proximally for at least 30% of their length; cephalon. In Rhenops odremani the visual tips of ®rst three, pointed; fourth and ®fth, surface is less expanded laterally and does somewhat rounded. not reach the lateral border furrow. HOLOTYPE: MBLUZ P-1448 A and B, part Rhenops rodesianus Morzadec, 1981, is and counterpart of an almost complete spec- readily distinguished from all other members imen with total length of approximately 29 of the genus (including the Venezuelan form) mm, from upper level of the CanÄo del Oeste by its genal spines, which are longer than the Formation, PerijaÂ Mountains, NW Venezue- entire thorax; as in R. circumapodemus la. Cast of holotype AMNH 46583. Smeenk, the ®fth pair of pygidial pleural lap- ETYMOLOGY: Named for PerijaÂ Mountains, pets is longer than the others. of NW Venezuela. It is concluded from the above comparison DESCRIPTION: The cephalon is semicircular that the form described here differs from spe- in outline, moderately convex, a little more cies of Rhenops described from Europe and than twice as wide as long, with length (sag.) North Africa (Morocco), and these differenc- approximately 8.7 mm and an inferred width es are regarded as suf®cient to erect a new (across the posterior border of the cephalon) species on the basis of a unique character of 19 mm (the right side is incomplete). The combination. anterior cephalic border is pointed medially. Inclusion of these Venezuelan fossils in The axial furrows are moderately wide; they Rhenops, by Lieberman and Kloc (1997), are more divergent anteriorly from S1. The and here is of considerable biogeographic in- glabella has three distinct pairs of lateral gla- terest, because it represents a signi®cant ex- bellar furrows. S1 is moderately wide, de- tension of the genus geographic range. veloped as a smooth convex curve posteriorly, becoming deeper proximally and in Genus Greenops Delo, 1935 contact with the axial furrows. S2 is roughly transverse, with its anterior margin weakly TYPE SPECIES: Cryphaeus boothi Green, convex, and almost reaching the axial fur- 1837; Middle Devonian of North America rows. S3 is widest distally and slightly con- (Pennsylvania). vex anteriorly. The frontal glabellar lobe is EMENDED DIAGNOSIS: As for Lieberman expanded anteriorly and laterally, declines and Kloc (1997: 77) except for: six or seven smoothly forward, has a rounded anterior lenses in dorsoventral ®le on visual surface; margin, and represents about 70% of total genal spine developed as moderately long glabellar length. The glabellar lobes are ¯at- ¯ange extending back to ®fth or sixth tho- tened, their height decreases posteriorly racic segment. (from L3 to L1), with the medial region ¯at. REMARKS: This genus has been rediagno- L3 is wedge-shaped, lengthening distally; L2 sed and discussed by Lieberman and Kloc is approximately rectangular, and L1 is nar- (1997), along with the designation of a neo- rower than L2. S0 is shallower than S1, mod- type for Greenops boothi. erately wide, convex anteriorly (sag.). The occipital lobe (L0) is longest sagittally, high- Greenops perijaensis, new species er medially, slightly above the median area Figure 5, C±E of glabella; becoming lower and narrower DIAGNOSIS: Cephalon semicircular in out- distally, without ornamentation. The eyes are line, with anterior border pointed; frontal gla- raised slightly above the glabella, with a bellar lobe expended anteriorly and laterally, maximum of seven lenses per dorsoventral genal spine ¯at, smooth, extended to sixth ®le on the visual surface. The anterior edge thorax segment; eyes with maximum of sev- of the visual surface contacts the axial furrow en lenses per dorsoventral ®le; visual surface at the anterolateral corner of L3; the posterior in dorsal view lies partly within a sagittal line edge of the visual surface is located posterior drawn tangentially to the frontal lobe; pygid- to a transverse line tangent to the anterior ium with ®ve pairs of triangular lappets all edge of L0. In dorsal view the visual surface AMNH NOVITATES Tuesday Dec 11 2001 10:21 AM 2000 novi 00147 Mp_12 Allen Press • DTPro System File # 01cc

12 AMERICAN MUSEUM NOVITATES NO. 3292

lies partly within a sagittal line drawn tan- triangular lappets, all approximately the gent to the frontal lobe. The ®xigena slopes same size as their respective pleurae. The from the axial furrow to the palpebrum; the distal tips of the ®rst three lappets are point- librigena is moderately declined toward the ed, but the fourth and ®fth are somewhat lateral cephalic border, whose margin forms rounded. The lappets make contact with each a broad continuous arch. The posterior ce- other proximally, for at least 30% of their phalic border furrow is nearly transverse and length, but are separated distally. The lateral broadest distally, and the posterior cephalic margins of the lappets are curved posteriorly, border widens abaxially. Genal spines are ¯at whereas their inner margins seem to be and smooth, and extend back to the sixth tho- straight. The lappets are covered by small, racic segment (their exact length is not very dense granules. The terminal lappet is ap- clear on the specimen). No ornamentation is proximately equal in length (sag.) to the ®fth observed on the cephalon. lappet, although its end is not very clear in The thorax contains 11 moderately convex the specimen. segments, with the axis occupying about one third the width of the thorax (maximum tho- DISCUSSION rax width 17.6 mm; axis width 6.1 mm). Ax- ial furrows are shallow but well de®ned. The Liebermann and Kloc (1997) diagnosed axis is somewhat higher than the pleural Greenops on the basis of 18 characters: 6, 8, ®eld. Each axial ring is of uniform length 21, 26, 28, 29, 31, 33, 34, 35, 42, 45, 47, 49, (exsag., sag.) with the medial portion convex 50, 51, 57, and 66 (see their data matrix and anteriorly and higher than the distal extrem- character states in tables 1 and 2), of which ities, which have an almost straight anterior 12 are discernible in the Venezuelan material margin and a convex posterior margin. The (21, 28, 29, 31, 33, 34, 42, 45, 47, 49, 50, articulation furrows are moderately wide and and 51); the remaining characters cannot be deep. The proximal part of the pleura is determined for preservational reasons. The transverse, approximately horizontal, where- new taxon differs from their generic diag- as the distal part is strongly ¯exed down- nosis in two aspects; there are seven lenses ward. Pleural furrows are moderately deep, in the dorsoventral ®le in the visual surface wide, and almost straight. instead of six; and the genal spine extends to The pygidium is broad, semicircular in the sixth thoracic segment instead of the ®fth outline, approximately 9.4 mm long (sag.) (characters 6 and 26 of Lieberman and Kloc, and 16.3 mm wide (maximum width anteri- 1997). We have opted here to include the Ve- orly). The axis is raised slightly above the nezuelan form within an emended concept of pleural ®elds, and is well de®ned by narrow Greenops (see diagnosis), rather than to erect and shallow axial furrows. Only the ®rst sev- a new genus for our unique specimen. en axial rings are distinct, plus the anterior Greenops perijaensis differs from G. boothi part of the eighth. Axial rings are delimited (Green, 1837; the type species of the genus) by shallow and moderately wide ring fur- in having S1 less deep and not as crescent- rows. The ®rst ®ve axial rings are gently shaped as in G. boothi; L0 without medial ¯exed anteriorly at the midline and are of tubercle (present in G. boothi); shape of nearly equal length throughout; the more pleural lappets (separated in G. boothi, posterior rings are almost straight. There are joined proximally for at least 30% of their ®ve pairs of pleurae; the ®rst three are gently length in the new species; terminal lappet of curved backward and the last two are more equal length to ®fth lappet in G. perijaensis, abruptly ¯exed; pleurae become slightly wid- (shorter in G. boothi); lappets covered by er distally. Interpleural furrows are obsolete; dense granulation in Greenops perijaensis the pleural furrows are shallow and relatively (no lappet ornamentation in G. boothi). wide. The ®rst three pleural furrows are more Greenops perijaensis is readily distin- conspicuous than the posterior two. The an- guished from Greenops barberi Lieberman terior and posterior bands of the pleural seg- and Kloc, 1997 by the absence of ornamen- ments lie at the same level. The border fur- tation on the glabella, genal ®eld, and genal row is well de®ned. There are ®ve pairs of spines (®ne tubercles are present in G. bar- AMNH NOVITATES Tuesday Dec 11 2001 10:21 AM 2000 novi 00147 Mp_13 Allen Press • DTPro System File # 01cc

2000 DE CARVALHO AND MOODY: MIDDLE-DEVONIAN TRILOBITES 13

beri); the shape of pygidial lappets (very (1) Viaphacops. Localities from which pointed in G. barberi;inG. perijaensis the Viaphacops is recorded are geographically ®rst three are pointed, and the last two are widespread, including Kazakhstan, North rounded); and relative length (sag.) of the America, and South America. These occur- terminal lappet (smaller than the ®fth lappet rences also fall within several different bio- in G. barberi; equal size in G. perijaensis). geographic areas. Maximova (1972) included Greenops grabaui Lieberman and Kloc, Kazakhstan within a large ``Paci®c paleogeo- 1997 is distinguished from Greenops peri- graphic region,'' but that concept is rather jaensis by its ornamentation (®ne tubercles generalized. According to Meyerhoff et al. in G. grabaui, mainly on the glabella), pres- (1996), Kazakhstan falls inside the Balkhash- ence of circular fenestrae on thoracic axial Mongolia-Okhotsk region of the Old World rings (absent in G. perijaensis); presence of realm. The North American and Venezuelan one transverse row of circular fenestrae on occurrences of Viaphacops lie within the the anterior band of thoracic pleural segment Eastern Americas realm (sensu Boucot, (absent in G. perijaensis); and shape of the 1988), and its Bolivian occurrence is within pygidial lappets (distal tips developed as the Malvinokaffric realm (sensu Eldredge blunt triangles in G. grabaui;inG. perijaen- and Ormiston, 1979). Viaphacops is thus a sis the ®rst three are pointed, the fourth and relatively cosmopolitan genus and it occurs ®fth are somewhat rounded. in both warm- and cold-water assemblages, Greenops chilmanae Stumm, 1965 differs suggesting it was eurytopic. from G. perijaensis in having a much nar- In Bolivia Viaphacops is known from the rower glabella posteriorly, and has tubercles Lower Devonian (Pragian, Gamoneda For- covering the glabella; tubercles are also pres- mation, its earliest known occurrence; also ent on the palpebral lobes, the axial part of Emsian, BeleÂn and Icla Formations). In the occipital ring, and the genal spines. North America its geological range extends Greenops widderensis Lieberman and from the Lower to Middle Devonian (Em- Kloc, 1997 differs from G. perijaensis in sian-Eifelian). In Kazakhstan Viaphacops is having the anterior cephalic border rounded; also known from the Lower and Middle De- tubercles on glabella and L3; visual surface vonian, although in this area its stratigraphic in dorsal view not located within a sagittal range extends farther, into the Givetian (its line drawn tangent to the frontal lobe; a sin- youngest known occurrence). The distribu- gle row of circular fenestrae on the anterior tion of Viaphacops does not therefore char- band of thoracic pleural segments, and all acterize any particular biogeographic prov- pygidial lappets pointed. ince, and its distribution pattern is relatively uninformative. Although the range of Via- BIOGEOGRAPHIC CONSIDERATIONS phacops extended into the southerly cold- water Malvinokaffric realm, no characteris- The trilobite assemblage from the Middle tically Malvinokaffric trilobites have been Devonian of Venezuela (CanÄo del Oeste For- found in the Venezuelan assemblage (El- mation) includes two representatives of the dredge and Ormiston, 1979). family Phacopinae. One of these represents (2) Rhenops. Until its discovery in Vene- a new species of Viaphacops (described zuela, the genus Rhenops was known only here), and the other was referred by Bene- from within the Rhenish-Bohemian region of detto (1979) to Phacops. The assemblage the Old World (Boucot, 1988), with occur- also includes two new species of the acastid rences in Europe (upper Emsian of northern subfamily Asteropyginae, one referable to Spain, the Armorican Massif, and Germany) the genus Rhenops and the other to Gree- and Africa (Eifelian of Morocco). The pres- nops. Some commentary seems worthwhile ence of Rhenops in Venezuela (®rst noted by regarding the biogeographic signi®cance of Lieberman and Kloc, 1997) extends this these occurrences. The Venezuelan phacopi- range to encompass part of the Eastern ne material described previously by Weis- Americas realm much farther westward than bord (1926) and Benedetto (1979) is too previously supposed, although its distribu- poorly known to be of biogeographic value. tion is disjunct. The Middle Devonian paleo- AMNH NOVITATES Tuesday Dec 11 2001 10:21 AM 2000 novi 00147 Mp_14 Allen Press • DTPro System File # 01cc

14 AMERICAN MUSEUM NOVITATES NO. 3292

position of the Eastern Andean Terrane is un- san Klofak assisted with preparation and certain (it may represent an exotic block casting of specimens, and the illustrations from Caledonian and Hercynian chains of were prepared by Chester Tarka and Lorraine North America or Europe; Restrepo and Meeker. We thank the anonymous reviewers Toussaint, 1998). Given the progressively for their very helpful comments and sugges- younger age of Rhenops occurrences from tions. east to west, it is tempting to suggest a pro- gressive westward migration of the genus REFERENCES from a center of endemism in Europe, but Arbizu, M. the fossil record of the genus is too fragmen- 1979. Asteropyginae (trilobite) du DeÂvonien tary to formulate a more rigorous biogeo- des Montagnes Cantabriques (Espag- graphic hypothesis. ne). Bull. Soc. Geol. Mineral. Bretagne, (3) Greenops. Traditionally, Greenops was seÂr. C. 9(2): 59±102. recognized in Emsian-Eifelian assemblages Barret, S. F., and P. E. Isaacson of Europe and Africa (Arbizu, 1979; Gandl, 1988. Devonian paleogeography of South 1972; Morzadec, 1983; Smeenk, 1983), but America. In N. J. McMillan, A. F. in North America it is known only from the Embry, and D. J. Glass (eds.), Devo- Givetian (Hamilton Group: Delo, 1935, nian of the World. Can. Soc. Pet. Geol. Mem. 14(1): 655±667. 1940; Lieberman and Kloc, 1997; Stumm, Benedetto, J. L. 1953, 1965, 1967). These records were 1979. La fauna de la FormatioÂn CanÄo del spread across two different biogeographic Oeste (DevoÂnico) en la regioÂn de CanÄo realms: the Old World realm (Rhenish-Bo- Colorado, Sierra de PerijaÂ, Venezuela. hemian Region, including northern Africa; P. I: Mollusca y Trilobita. Bol. Geol. Boucot, 1988) and the Eastern Americas (Caracas) 13(25): 81±111. realm. In their revision of Greenops, Lieber- 1984. Les brachiopodes DeÂvoniens de la Si- man and Kloc (1997), rede®ned the genus to erra de PerijaÂ (Venezuela). Systemati- include only North American species, and all que et implications paleÂogeographi- the Old World taxa formerly assigned to ques, V.1. Brest: Coll. Biostrat. du Pa- leoz. Univ. Bretagne Occidentale. Greenops were excluded from that genus. Berry, C. M., J. E. Casas, and J. M. Moody The Venezuelan occurrence of Greenops rep- 1993. Diverse Devonian plant assemblages resents a southerly extension of its distribu- from Venezuela. Doc. Lab. GeÂol. Lyon, tion within the Eastern Americas realm, and 125: 29±42, 4 ®gs., 1 table. reinforces the faunal similarities noted by Boucot, A. J. Benedetto (1984) and Boucot (1988) be- 1988. Devonian biogeography: an update. In tween the Devonian brachiopods of Vene- N. J. McMillan, A. F. Embry, and D. J. zuela and those of eastern North America; Glass (eds.), Devonian of the World. and by Eldredge and Ormiston (1979) within Can. Soc. Pet. Geol. Mem. 14(3): 211± the trilobites. 227. Delo, D. M. 1935. A revision of the phacopid trilobites. J. ACKNOWLEDGMENTS Paleontol. 9(5): 402±420. 1940. Phacopid trilobites of North America. The authors thank Ascanio RincoÂn and Geol. Soc. Am. Spec. Pap. 29: 135 pp. Reina Rosales for assistance with ®eld col- Eldredge, N. lections. This study was undertaken in the 1972. Systematics and evolution of Phacops Department of Invertebrate Paleontology at rana (Green, 1832) and Phacops io- the American Museum of Natural History, wensis Delo, 1935 (Trilobita) from the and we are especially grateful for the support Middle Devonian of North America. and encouragement provided by Neil Land- Bull. Am. Mus. Nat. Hist. 147: 45±114. 1973. Systematics of Lower and Lower Mid- man and Niles Eldredge. M.G.P.C. thanks dle Devonian species of the trilobite John Maisey (AMNH) for discussions about Phacops Emmrich in North America. paleogeography and for reviewing the man- Ibid. 151: 289±337. uscript. Inez Horovitz and Jesus Alvarado Eldredge, N., and A. R. Ormiston kindly helped with the Spanish abstract. Su- 1979. Biogeography of Silurian and Devoni- AMNH NOVITATES Tuesday Dec 11 2001 10:21 AM 2000 novi 00147 Mp_15 Allen Press • DTPro System File # 01cc

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an trilobites of the Malvinokaffric mentologie, paleÂontologie, stratigrap- Realm. In J. Gray and A. J. Boucot hie). Les Trilobites. MeÂm. Soc. GeÂol. (eds.), Historical biogeography, plate MineÂral. Bretagne 24(313): 279±291. tectonics, and the changing environ- Pek, I., and J. VaneÆk ment: 147±167. Corvallis: Oregon 1991. On some Silurian and Devonian trilo- State Unvi. Press. bites of Bolivia. Acta Universitatis Pa- Gandl, J. lackiana Olomucensis, Fac. Rer. Nat. 1972. Die Acastavinae und Asteropyginae Geogr.-Geol. XXX, 103: 75±104. (Trilobita) Keltiberiens (NE-Spanien). Pillet, J. Abh. Senckenb. Natforsch. Ges. 530: 1958. Contribution aÁ l'eÂtude de quelques As- 1±184. teropyginae. Bull. Soc. GeÂol. France Green, J. (6)8: 3±20. 1837. Description of several new trilobites. RamskoÈld, L., and L. Werdelin Am. J. Sci. (1)32: 343±349. 1991. The phylogeny and evolution of some Hall, J., and J. M. Clarke phacopid trilobites. Cladistics 7: 29± 1888. Descriptions of the trilobites and other 74. Crustacea of the Oriskany, upper Hel- Restrepo, J. J., and J. F. Toussaint derberg, Hamilton, Portage, Chemung, 1998. Terranes and continental accretion in and Catskill Groups. Natural History of the Colombian Andes. Episodes 11(3): New York. N.Y. Geol. Surv., Paleontol. 189±193. V.7. Smeenk, Z. Hea, J. P., and A. B. Whitman 1983. Devonian trilobites of the southern 1960. EstratigrafõÂa y petrologõÂa de los sedi- Cantabrian Mountains (northern Spain) mentos pre-cretaÂcicos de la parte norte- with a systematic description of the As- central de la Sierra de PerijaÂ, Estad Zu- teropyginae. Leidse Geol. Meded. 52: lia, Venezuela. Mem. Tercer Congr. 383±511. Geol. Venez. 1: 351±376. Caracas: Ed- Struve, W. itorial Sucre. 1959. Asteropyginae. In R. C. Moore (ed.), Liddle, R. A., G. D. Harris, and J. W. Wells Treatise on invertebrate paleontology, 1943. The Rio Cachiri section in the Sierra de part O, Arthropoda 1: 447±483. Perija, Venezuela. Bull. Am. Paleontol. Lawrence: Univ. Kansas Press. 27: 269±375, Ithaca. Stumm, E. C. Lieberman, B. S., and G. J. Kloc 1953. Trilobites of the Devonian. Contrib. 1997. Evolutionary and biogeographic pat- Mus. Paleontol. Univ. Michigan, 10(6): terns in the Asteropyginae (Trilobita, 101±157. Devonian) Delo, 1935. Bull. Am. Mus. 1954. Lower Middle Devonian phacopid tri- Nat. Hist. 232: 1±127. lobites from Michigan, southwestern Linsley, D. M. Ontario, and the Ohio Valley. Ibid. 1994. Devonian paleontology of New York. 11(11): 201±221. Paleontol. Res. Inst. Spec. Publ. 21. Maximova, Z. A. 1965. Two new species of trilobites from the 1972. New Devonian trilobites of the Phaco- Middle Devonian Silica Shale of poidea. Paleontol. J. 1: 78±83. North-Western Ohio. Ibid. 19(13): 163± Meyerhoff, A. A., A. J. Boucot, D. M. Hull, and 166. J. M. Dickins 1967. Devonian trilobites from north-western 1996. Phanerezoic faunal and ¯oral realms of Ohio, northern Michigan, and western the Earth: the intercalary relations of New York. Ibid. 21(6): 109±122. the Malvinokaffric and Gondwana fau- Sutton, F. A. nal realms with Tethyan faunal realm. 1946. Geology of Maracaibo Basin, Venezue- Geol. Soc. Am. Mem. 189: 1±69. la. Bull. Am. Assoc. Pet. Geol. 30(10): Morzadec, P. 1621±1741. 1983. Trilobites du Devonien (Emsien - Fa- Weisbord, N. E. mennien) de la rade de Brest (Massif 1926. Venezuelan Devonian fossils. Bull. Armoricain). Palaeontographica (A) Am. Paleontol. 11(46): 223±272. 181: 103±184. Yaekel, S. W. Morzadec, P., F. Paris, and P. Racheboueuf. 1924. Notas provisionales sobre la zona de El 1981. La TrancheÂe de la Lezais Emsien Su- Tigre, Estado Zulia, Distrito Mara. Ma- peÂrieur du Massif Armoricain (seÂdi- racaibo: Venez. Atl. Re®ning Co. AMNH NOVITATES No RH this page!! Tuesday Dec 11 2001 10:21 AM 2000 novi 00147 Mp_16 Allen Press • DTPro System File # 01cc

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