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AMERICAN MUSEUM Norntates PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 Number 3192, 11 pp., 4 figures, 2 tables March 3, 1997

Devonian Calmoniid Trilobites from the Parnalfba Basin, Piaui State, Brazil

MARIA DA GLORIA PIRES DE CARVALHO,1 GREGORY D. EDGECOMBE,2 AND BRUCE S. LIEBERMAN3

ABSTRACT Two new species of calmoniid trilobites are de- genetic parsimony analysis is used to infer rela- scribed from the Middle Devonian of the Parnaiba tionships within the previously unresolved "Me- Basin, Piaui State, northeastern Brazil. One of tacryphaeus tuberculatus group," which is ex- these occurs in the lower Pimenteira Formation, tended to include the form from the Cabeqas For- and the other is found in the Passagem Member mation. The relationships of Metacryphaeus from of the Cabeqas Formation. The presence of the the Pimenteira Formation are less certain, but genus Metacryphaeus confirms faunal affinities some morphologic similarities suggest that it is with the Malvinokaffric Devonian fauna. Phylo- closely allied to M. curvigena Lieberman, 1993.

INTRODUCTION The Devonian System is well represented sented by the Upper Serra Grande Group, in the Parnaiba Basin, which is situated in i.e., Jaicos and Itaim Formations respective- northeastern Brazil, and extends across the ly, as indicated by new chitinozoan-based in- states of Ceara, Piaui, Maranhao, Para', and vestigations (Grahn, 1992). The Middle De- Tocantins (fig. 1). vonian is represented by the Pimenteira and Lower to lower Middle Devonian [Emsian lower Cabeqas formations (Melo, 1988). The (?) and early Eifelian] sediments are repre- upper Cabegas and Longa' formations are Up- ' Professora, Universidade Federal do Rio de Janeiro, Instituto de Geociencias, CCMN, Departamento de Geologia. Cidade Universitdria, Ilha do Fundao, CEP: 21. 949-940. Rio de Janeiro-RJ, Brazil. 2 Senior Research Scientist, Australian Museum, 6 College Street, Sydney South, NSW 2000, Australia. I Postdoctoral Fellow, Department of Organismic and Evolutionary Biology, Harvard University, Cambridge, MA 02138.

Copyright (C American Museum of Natural History 1997 ISSN 0003-0082 / Price $1.70 2 AMERICAN MUSEUM NOVITATES NO. 3192 per Devonian and Lower Carboniferous (Fa- genetic analysis of the taxa we believe be- mennian to Tournaisian; Quadros, 1982; Lo- long in this group following the formal de- boziak et al. 1992). Further details of the scription of the new Brazilian material. stratigraphy of the Parnafba Basin are pro- Metacryphaeus (in the restricted sense of vided in Melo (1988: 676). Lieberman, 1993) is now known from the Although the deposits of this basin have Parnal'ba Basin of Brazil (Carvalho et al., been studied since the early 20th century 1994, and this paper), as well as from De- (Small, 1914), its fossils are less well known vonian assemblages of Bolivia (Belen, Icla, than in other Devonian Brazilian areas and and Sicasica formations; Wolfart, 1968; Falk- there is a paucity of information concerning land Islands (Fox Bay Formation; Edgecom- its paleontology. There are only a few papers be, 1991) and Parana' Basin (Ponta Grossa describing or illustrating fossil species, Formation and Chapada Group; Clarke, among them Krausel and Dolianiti (1957), 1913; Castro, 1968; Carvalho et al., 1987). Suarez-Riglos (1967), Castro (1968), Melo Alleged members of the "M. tuberculatus (1985), Caldas et al. (1987), and Fonseca and group" occur in all these areas. The only oth- Melo (1987). Castro (1968) and Caldas et al. er previous record of Metacryphaeus in Bra- (1987) in particular presented systematic zil is M. paituna (Hartt and Rathbun, 1875), treatments of Devonian trilobites from the from the Erere Formation, Amazon Basin. Parnaiba Basin. That form has been reassigned by Lieber- The Malvinokaffric Realm (sensu Eld- man, 1993, to a new genus, Eldredgeia. redge and Ormiston, 1979) is characterized Morphological terminology follows that by a highly endemic trilobite fauna known used in the Treatise on Invertebrate Paleon- from austral localities. Members of the fam- tology, Part 0 (1959), and in Lieberman et ily Calmoniidae (Delo, 1935) represent al. (1991). Specimens figured herein are de- three-quarters of the generic diversity (Eld- posited in the Department of Geology, Sec- redge and Ormiston, 1979). Recently pub- tion of Paleontology of University Federal do lished studies among these austral trilobites Rio de Janeiro, RJ, Brazil; Department of In- (Lieberman et vertebrates, American Museum of Natural al., 1991; Lieberman, 1993) History (AMNH), NY; Departamento Na- recognized a complex phylogenetic pattern cional de Produqao Mineral (DNPM), Sec- requiring the erection of new genera for tion of Paleontology, Rio de some and the restriction of others to just a Janeiro, Brazil. few species. The genus Metacryphaeus Reed, INSTITUTIONAL ABBREVIATIONS 1907, has proven particularly problematic; Lieberman (1993) retained nine with- AMNH American Museum of Natural His- species tory, New York. in this genus. The relationships of five of DGM,DNPM Divisao de Geologia Mineralogia, these species were completely resolved, but Departamento Nacional de Produ- not those within the "M. tuberculatus sao Mineral, Rio de Janeiro, Bra- (Kozlowski, 1923) species group" (compris- zil. ing M. tuberculatus, caffer, allardyceae, and MCT Museu Ciencias da Terra, Setor de australis). Paleontologia, DNPM, Rio de Ja- Two new species of calmoniid trilobites neiro, Brazil. UCGM University of Cincinnati, Geologi- from the Middle Devonian of the Parnaiba cal Museum. Basin (Brazil) were referred to Metacry- UFRJ Universidade Federal do Rio de Ja- phaeus by Carvalho et al. (1994), but were neiro, Rio de Janeiro, Brazil. not described; this paper presents compre- hensive descriptions of this material. One of SYSTEMATIC PALEONTOLOGY these taxa is known from only a few poorly FAMILY CALMONIIDAE DELO, 1935 preserved specimens, but its closest relations can be broadly determined. The other species SUBFAMILY CALMONIINAE DELO, 1935 is known from numerous specimens that pro- Genus Metacryphaeus Reed, 1907 vide sufficient data to suggest placement TYPE SPECIES: Phacops caffer Salter, 1856. within the "M. tuberculatus group" of Lie- [By subsequent designation of Rennie berman (1993). We have appended a phylo- (1930); revised Cooper (1982)]. 1997 CARVALHO ET AL.: TRILOBITES FROM BRAZIL 3

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REMARKS: For a more complete discussion rior branch of facial suture has a weakly sin- of Metacryphaeus, see Lieberman et al. uous anterolateral course between E and the (1991) and Lieberman (1993). lateral border furrow, and is sharply flexed backward across the lateral border. Anterior Metacryphaeus kegeli, new species branch of facial suture circumscribes frontal Figure 2A-H lobe, crossing axial furrow at the cephalic anterolateral margin, and with a nearly Asteropyge sp. Kegel, 1953. straight posterodorsal course to -y. S3 are Metacryphaeus sp. Melo, 1988: 681, pl. 25, figs. 9-15. broad, deeply incised, straight, weakly diver- Metacryphaeus kegeli Carvalho et al., 1994: 119 gent, and connect with the axial furrows; to- (nomen nudum). pographic separation between frontal lobe and posterior glabellar region is produced by DIAGNOSIS: A Metacryphaeus with the a change in convexity. S2 are transverse, de- frontal glabellar lobe steeply inclined anter- veloped as apodemal grooves, shallowing omedially; cephalic anterior margin and fron- distally, and weakly confluent with axial fur- tal lobe of gentle even convexity; broad, rows. SI are broad, crescent shaped, and be- deep cephalic axial furrows; small eyes well come shallow adjacent to axial furrows. SO offset laterally from axial furrows; librigena is long (sag.), shallow, weakly arched for- steeply inclined. ward medially, becoming deeper and narrow- ETYMOLOGY: In honor of Wilhelm Kegel, er distally, connecting with the axial furrow. who made important geological studies on L2-L3 are flattened (tr.) medially; L3 lateral the Devonian of the Parnafba Basin and who lobes are wedge shaped, weakly convex (tr.), collected these specimens. and gently declined outward. L2 are convex HOLOTYPE: DNPM N.6133-I, an internal anteriorly, concave posteriorly. LI are cres- mold of a cephalon, from the base of the Pi- cent shaped, short (exsag.), depressed below menteira Formation, nearly 11 km south of L2 exsagittally, and moderately arched (tr.) Pimenteira, Piaui State, Brazil. medially. The occipital ring is uniformly PARATYPES: DNPM N.6131-I, an internal long (sag.), and deflected backward (sag.). mold of a rolled up small specimen; 6132-I, Large Eye Index (LEI) is 0.33; the visual sur- an internal mold of a thorax and pygidium face has 23 dorsoventral lens files, with a of a small specimen; 6134-I, an external maximum of six lenses per file. Interocular mold of a small cephalon. fixigena is elevated to height of palpebrum; OCCURRENCE: Lower Pimenteira Forma- librigenal field and extraocular fixigena have tion; Eifelian (Fonseca and Melo, 1987), re- shallow pits. Palpebral furrow is narrow and gion of Pimenteira (sensu Kegel, 1953), Par- moderately deep. Eyes are well offset later- naiba Basin, Piaui State, Brazil. ally from axial furrow, nearly overhanging DESCRIPTION: Cephalon moderately convex lateral border furrow in dorsal orientation, (tr.), with length (sag.) 50% of width. Axial and eye ridge weakly connects with axial fur- region projects beyond arc of genae. Glabella row. Cephalic lateral border furrow is shal- is truncated anteriorly, with broad, deep axial low and broad. Cephalic posterior border fur- furrows, weakly diverging to the anterior row is deep, broad, with a gentle posterolat- margin. Width (tr.) of occipital ring is 80% eral inflection to behind outer (exsag.) edge of width of frontal lobe. Cephalic anterior of the eyes, then is flexed forward distally, border is obscured medially. In dorsal view, where it is shallow but continuous with lat- its anterior margin is smoothly rounded, eral border furrow. Posterior border is short formed by frontal glabellar lobe medially. (exsag.) adjacent to axial furrows, becoming Frontal glabellar lobe is depressed beneath longer abaxially, and genal margin is mod- L1-L2 (sag.); L1-L3 are nearly flat (sag.) erately convex. and gently declined forward. Posteromedian DISCUSSION: Kegel (1953) noted differ- part of the frontal lobe is weakly convex ences between the samples here referred to (sag.). Frontal lobe is 60% of length (sag.) Metacryphaeus kegeli, n. sp. (= Asteropyge, of glabella. Posterior median impression n. sp., of Kegel) and Metacryphaeus aus- (PMI) is well developed and circular. Poste- tralis (Clarke, 1913), from the Ponta Grossa 1997 CARVALHO ET AL.: TRILOBITES FROM BRAZIL 5

A B

F G H I Fig. 2. Metacryphaeus kegeli, n. sp., Pimenteira Formation, Parnaiba Basin. A: Holotype DGM, DNPM 6133-I. Internal mold of cephalon, dorsal view. Scale bar = 10 mm. B-D: dorsal, frontal, and dorsolateral views of holotype. E: Paratype DGM, DNPM 6132-I, internal mold of thoracopygidium. Scale bar for B-E = 10 mm (below E). F-H: Paratype DGM, DNPM 6131-I, internal mold of small rolled-up specimen: (F) dorsal view of cephalon; (G) lateral view; and (H) dorsal-posterior view of thorax and pygidium. Scale bar for F-H = 10 mm (next to H).

Formation, Parana Basin. Metacryphaeus ke- incised axial furrows, with deeply impressed geli, n. sp., has some distinctive derived lateral glabellar furrows. Metacryphaeus ke- characters shared with M. curvigena Lieber- geli nevertheless shows a more marked to- man, 1993, from the Icla Formation (Emsian) pographic break (sag.) between the frontal of Bolivia. These include the shape of the lobe and posterior glabellar region than M. frontal glabellar lobe (broadened subrhom- curvigena. Another possible difference be- boid, slightly bulging anterior to the genal tween these two species (the form of the gen- arc, steep anteromedially), and broad, well al angles) cannot be determined at present. 6 AMERICAN MUSEUM NOVITATES NO. 3192

Genal spines are not preserved in specimens (sag.) equal to width across frontal lobe. of M. kegeli, although the holotype may have Frontal lobe is moderately convex (sag.) and an eroded spine base. raised above posterior glabellar region. An- terior branch of each facial suture has a Metacryphaeus meloi, new species straight course from -y to just outside antero- Figure 3A-I ventral corner of frontal lobe. Posterior branch of these sutures has a straight anterior Metacryphaeus australis (Clarke). Castro, 1968: inflection between E and lateral border fur- 485, plate III, figs. 2-5. rows, than is sharply flexed backward across Metacryphaeus cf. australis (Clarke). Caldas et al., 1987: 539, phot. 01-03. lateral border; E posterior to w. Small tuber- Metacryphaeus meloi Carvalho et al., 1994: 119 cles are scattered on LO, the glabella, and (nomen nudum). interocular fixigenae. Frontal glabellar lobe is broadly ovate in outline, 60% length (sag.) DIAGNOSIS: Cephalon subtriangular, with of glabella. Axial furrows are narrow, short median frontal process; glabella dense- straight, and moderately incised. Occipital ly covered with small, relatively subdued tu- ring (LO) is 60-70% width (tr.) of frontal bercles; genal angles bluntly pointed; eyes lobe. Lateral glabellar furrows are well indistinctly separated from axial furrows, mod- cised; S3 are faintly convex posteriorly; S2 erately oblique (exsag.). Pygidial pleurae ter- are slightly shallower than S1, with very minate as blunt spines; axial rings convex weak incision distally; S1 are deep adjacent forward medially. to axial furrow, and crescent shaped. Median ETYMOLOGY: In honor of Jose Henrique region of L1-L3 is flat (sag.) and inclined Goncalves de Melo, who has studied the Bra- anteriorly. L3 lobes are wedge-shaped and zilian Devonian and has worked with the se- expanded distally. L2 lobes are smaller than nior author in the Parnal'ba Basin, where L3, and are rectangular; L2-L3 are gently these specimens were collected. arched (tr.). LI lobes are almost transverse. HOLOTYPE: MCT N. 6 822-I, an internal Occipital furrow (SO) is moderately incised mold of a cephalon, from the Cabecas For- (sag.), longest medially, and impressed as mation (Passagem Member), near the village transverse apodemes abaxially. Occipital ring of Oiti, a few kilometers from the junction (LO) is gently and evenly convex (sag.), of the road Valen,a do Piaui to Pimenteiras, moderately arched transversely, with con- Piaui State, Brazil. stant length (sag., exsag.), and weakly PARATYPES: MCT N. 6 823-I, 6824-I, curved forward distally. Posterior median UFRJ N. 047-Tr, and 048-Tr, AMNH 45352 impression (PMI) is obscure to moderately (Fossil Invertebrates Collection). defined. Large Eye Index (LEI) is 0.29- OCCURRENCE: Cabecas Formation (Passa- 0.34. Palpebral lobes are moderately oblique gem Member), type locality, also by road BR (exsag.), gently inflated, with their anterior 316, km 305, near Picos, Piaui State, Brazil margin distinctly separated from axial fur- (see Caldas et al., 1987, fig. 1) and the vil- rows. Number of dorsoventral lens files in lage of Barreiro Branco (fig. IC). The trilo- visual surface is 26, with a maximum of 8 bites are preserved as disarticulated and lenses per file. Extraocular fixigenae are mostly fragmentary molds in a fine- to me- strongly pitted, with postocular region gently dium-grained micaceous sandstone. sloping backward to border furrow. Librigen- DESCRIPTION: Cephalic length (sag.) is al fields are steeply declined outward, and about 55% of width across the genal angles. faintly convex. Lateral border furrows are Anterior cephalic border is obsolete medial- shallow, becoming deeper posteriorly. Pos- ly, becoming evenly lengthened abaxially. terior border furrow is transverse, deeper ad- Anterior cranidial border is short (exsag.) jacent to axial furrows, broadening and shal- and visible in dorsal view. Angular antero- lowing distally, and distinctly continuous median process is short (sag.). Cranidial an- with lateral border furrows. Lateral cephalic terolateral margin is nearly straight. Pregla- borders are narrow anteriorly, broadening bellar furrow is narrow, sharply impressed. slightly posteriorly, and convex. Genal mar- Glabella is expanded anteriorly, with a length gins are gently and evenly convex laterally. 1997 CARVALHO ET AL.: TRILOBITES FROM BRAZIL 7

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Fig. 3. Metacryphaeus meloi, n. sp., Cabeqas Formation (Passagem Member). A: Holotype MCT 6822-I. Internal mold of cephalon, dorsal view. B: Detail of the holotype, showing the granulation on the cephalon. C, D: Frontal and dorsolateral views of holotype. Scale bar for A-D = 10 mm. EG: Paratype MCT 6823-I, internal mold of pygidium: dorsal, lateral, and posterior views, respectively. Scale bar for E-G = 10 mm. H, I = Metacryphaeus tuberculatus. J: Metacryphaeus rotundatus. Scale bar for H-J = ,10 mm. 8 AMERICAN MUSEUM NOVITATES NO. 3192

Posterior border lengthens abaxially, and is (PMI) in Metacryphaeus meloi, n. sp., is a gently flexed forward distally. Cephalic dou- distinctive difference separating it from other blure is weakly convex, longest medially, species of the "Metacryphaeus tuberculatus and narrowing abaxially, with a vincular fur- group." Other reports of this group in Brazil row. Hypostomal suture is faintly convex are M. australis (Clarke, 1913) from the Pon- forward. ta Grossa Formation and Chapada Group, Pygidium is triangular, with a length about Parana Basin (Carvalho and Edgecombe, 55% of width. Its axial furrows are shallow, 1991) and M. tuberculatus (Kozlowski, weakly converging posteriorly, with a de- 1923), although the concretion with M. tub- creased angle of convergence behind fifth ax- erculatus (illustrated here in fig. 3H) is lith- ial ring. Axis is moderately convex (tr.), ologically different from the typical concre- about 35% of pygidium width anteriorly, and tions that occur in the Pimenteira Formation, is composed of eight or nine rings plus an and its provenance is questionable. indistinctly segmented axial terminus, round- Metacryphaeus meloi, n. sp., can be dis- ed posteromedially. First five rings of axis tinguished from M. tuberculatus (fig. 3H-I) are well defined and slightly shortened sag- by its less inflated frontal glabellar lobe, nar- ittally, with anterior ring furrows deep and rower cephalic axial furrows, and convex py- gently arched forward. Narrow (tr.) apode- gidial axial rings. In addition, in M. tuber- mes are slightly medial to axial furrows in culatus the lateral glabellar lobes are more four anteriormost ring furrows. First three strongly defined. Another potential differ- pygidial lappets are gently curved posterior- ence is the Large Eye Index (LEI), which is ly, but last two are more strongly flexed. 0.26-0.30 in M. tuberculatus and 0.29-0.34 Pleural furrows are almost straight, narrow, in M. meloi, although these values weakly moderately deep, narrowing slightly adjacent overlap. Metacryphaeus tuberculatus never- to axial furrows. Interpleural furrows are theless shares several synapomorphies with shallow, with first three distinct. Pleural lap- M. meloi, n. sp., and we consider them to be pets have a straight or gently curved trun- related. cation, and they are pointed posteriorly. Post- closely axial region is short (sag.) and blunt, extend- Metacryphaeus meloi, n. sp., differs from ing back to tips of fifth pair of pleural lap- M. australis in the form of its anterior ce- pets. phalic margin, which bears a distinct median DISCUSSION: All material referred to this process. In M. meloi, the anterior edges of species is from the Cabe,as Formation, from the eyes are well-separated from the axial a restricted stratigraphic and geographic furrows, and their sculpture is more coarsely range. Some of the observed variation can be tuberculate. The lp furrows (S1) are crescen- accounted for by differences in preservation. tic in M. meloi whereas they are almost For example, specimens preserved as internal straight in M. australis. The bluntly pointed molds in medium-grained sandstone lack ev- genal angles, the pygidial axial rings that are idence of exoskeletal tuberculation like that convex anteriorly (sag.), and blunt pygidial preserved on the holotype. Characters which pleural tips are other characters that distin- vary continuously within the sample, such as guish M. meloi from M. australis. depth of incision of the PMI and convexity Specimens from the Pimenteira Formation (sag.) of the frontal lobe, are regarded as in- (Picos area) described by Castro (1968) as traspecific variation. Metacryphaeus australis (Clarke) are closest Metacryphaeus meloi, n. sp., is referred to to M. meloi from the Cabeqas Formation. In- the "Metacryphaeus tuberculatus group" on deed, our restudy of some of Castro's collec- the basis of exoskeletal tuberculation (albeit tion suggests that the Pimenteira form may weak in M. meloi), triangular anteromedian be conspecific with our M. meloi material cranidial border extending beyond the gla- that is stratigraphically somewhat higher. An bella, rounded frontal glabellar lobe, and the apparent difference is that cephala from the anterior edge of the eye well separated from Pimenteira Formation show a less pro- the axial furrows. nounced cranidial anteromedian process; the The weak posterior median impression cranidial anterolateral margin is convex out- 1997 CARVALHO ET AL.: TRILOBITES FROM BRAZIL 9

TABLE 1 NN(P Characters Used in Phylogenetic Parsimony Analysis, with Various Character States in Parentheses (0) plesiomorphic state, (1) apomorphic state. See table 2 for taxon character matrix Character 0: Dorsoventral height of pygidium-(O) gradually decreases posteriorly; (1) sharply reduces posterior to the posteriormost apodemal ring. Character 1: Anterior margin of cephalon-(O) rounded point at the midline; (1) with triangular median pro- Fig. 4. Single most parsimonious tree; length cess. 12 steps, consistency index 0.91, retention index Character 2: Frontal glabellar lobe-(O) moderately in- 0.88, showing the phylogenetic relationships of flated; (1) strongly inflated. of to or the "Metacryphaeus tuberculatus group," pro- Character 3: Anterior edge eye-(O) adjacent duced by analysis of the character data in table 2. in contact with axial furrow; (1) displaced from axial furrow. Character 4: Width of cephalic axial furrows (O) nar- ward (vs. straight in Formation row; (1) broad. Cabeqas Character 5: Shape of Sl-(O) straight to slightly specimens). curved; (1) crescentic shape. Species from South Africa, [like M. caffer Character 6: Cephalic tuberculation-(O) absent or (Salter, 1856)], and from the Falkland Islands weak; (1) coarse. [M. allardyceae (Clarke, 1913)] show signif- Character 7: Genal angle-(O) spinose; (1) blunt. icant differences from M. meloi including the Character 8: Pygidial margin-(O) lappets ending in placement of the eyes and more subdued ex- pointed spines; (1) lappets shorter and blunter oskeletal tuberculation. Additionally, M. caf- spines. fer differs from the new species in having Character 9: Shape of anterior 3-4 pygidial rings-(O) spinose genal angles, a pygidium with trans- with spatulate distal terminations; (1) without spatulate distal terminations. verse rings, and longer pygidial pleural lap- Character 10: Pygidial terminus-(O) short triangular pets. spine; (1) blunt triangular point. PHYLOGENETIC ANALYSIS The Hennig86 program of Farris (1988) was used to perform a cladistic parsimony analysis on six species of the "Metacry- phaeus tuberculatus group," employing 11 TABLE 2 characters (fig. 4; tables 1, 2). In this analy- Character State Distributions for Taxa Used in sis, M. rotundatus (Kozlowski, 1923; see fig. Phylogenetic Parsimony Analysis taxon Characters and alternative states are as listed in 3J) is held to be exemplar for the sister 1 of the "M. tuberculatus group" of Lieber- table man (1993). Analysis with this outgroup 0 1 2 3 4 5 6 7 8 9 10 coding produced a single most parsimonious Metacryphaeus rotundatus tree of length 12 steps, with a consistency 0 0 0 0 0 0 0 0 0 0 0 index of 0.85 and a retention index of 0.91. M. tuberculatus Metacryphaeus meloi and M. tuberculatus 0 1 1 1 1 1 1 1 1 1 1 are resolved as sister taxa, and the most to- M. caffeer pologically derived members of the "M. tub- 1 1 0 0 0 0 0 0 0 0 1 erculatus group." Metacryphaeus allardy- M. allardyceae ceae and M. caffer are successive sister taxa 0 1 0 0 0 0 0 1 1 0 1 to M. meloi, n. sp., plus M. tuberculatus, and M. australis M. australis is topologically basal. 0 0 0 0 0 0 0 0 0 1 1 Bolivian species and M. tuberculatusiM. M. meloi 0 1 0 1 0 1 0 1 1 1 1 meloi material of the Parnaiba Basin are 10 AMERICAN MUSEUM NOVITATES NO. 3192 more closely allied to taxa from the Falkland ACKNOWLEDGMENTS Islands and South Africa than to species from We thank N. Eldredge (AMNH) for per- the Parana Basin. mitting study of specimens under his care, providing access to appropriate facilities, and for reviewing this manuscript. The senior au- BIOGEOGRAPHY thor is grateful to J. G. Maisey (AMNH) for help in preparing this paper. D. R. B. Campos Eldredge and Ormiston (1979) contended arranged for loan of specimens housed at that the family Calmoniidae is exclusively DNPM (Departamento Nacional de Produqco Malvinokaffric. The presence of the calmon- Mineral). Assistance with preparation and iid genus Metacryphaeus in the Parnaiba Ba- photography was provided by S. Klofak and sin is therefore biogeographically important. A. Modell (AMNH, Department of Inverte- Discovery of this Malvinokaffric form in the brates) and L. Meeker (AMNH, Department of Vertebrate Paleontology). The senior au- Parnafba Basin significantly extends the geo- thor thanks the Departamento de Geologia da graphic range of the family. Universidade Federal do Rio de Janeiro The associated brachiopods of this basin (UFRJ) for permitting her to take a sabbatical probably represent a mixed fauna, with forms semester in order to conduct this project. Her typical of the Old World and the eastern research was supported in part by an Amer- Americas (Copper, 1977; Fonseca and Melo, ican Museum of Natural History (AMNH) 1987; Melo, 1988). This mixture of endemic collections study grant which is gratefully and more cosmopolitan taxa suggests that the acknowledged. Independent reviews of the Parnaiba Basin represents a biogeographic manuscript were kindly provided by J. H. G. boundary area which contains faunal ele- de Melo (Petrobras, Cenpes, Rio de Janeiro), ments of the Old World, eastern American M. R. Cooper (University of Durban-West- and Malvinokaffric biogeographic regions ville, Durban, South Africa) and D. Brezinski (Melo, 1988). (Maryland Geological Survey).

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