Noqvtates PUBLISHED by the AMERICAN MUSEUM of NATURAL HISTORY CENTRAL PARK WEST at 79TH STREET, NEW YORK, N.Y

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AMERICAN MUSEUM Noqvtates PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 Number 2982, 16 pp., 12 figs. August 9, 1990

Chonetacean Brachiopods of the "Pink Chonetes" Zone, Onondaga Limestone (Devonian, Eifelian), Central New York

PATRICK R. RACHEBOEUF1 AND HOWARD R. FELDMAN2

ABSTRACT The "Chonetes" Community of the Seneca ignated. Hallinetes is the most common chonetid Member, Onondaga Limestone (Devonian, Eife- found on the bedding planes (92%) while Longi- lian), is redefined. The chonetacean brachiopods spina and "Eodevonaria" are much less common found therein belong to three different taxa: (1) (5.4% and 2%, respectively). The Hallinetes Com- Hallinetes lineatus, n. gen., which is distinguished munity ofthe "Pink Chonetes" Zone corresponds from other chonetacean genera by its disymmetri- to the assemblage within the mudstone matrix cally arranged spines perpendicular to the hinge rather than to that on the bedding planes. It is a line and for which a neotype has been designated, low-diversity community representative ofa quiet (2) Longispina mucronata and "Eodevonaria" water environment, probably in a Benthic Assem- hemispherica for which lectotypes have been des- blage 4 position of Boucot (1975). INTRODUCTION The Onondaga Limestone in central New of the formation in Jamesville, New York York (fig. 1) can be divided into four mem- (AMNH Locality 3128) is 69.5 ft (Feldman, bers (Oliver, 1954; Feldman, 1980, 1985): 1985). In the eastern and western part of the from base to top, the Edgecliff, Nedrow, state, that is, away from the basinal axis, the Moorehouse, and Seneca. The total thickness total thickness approximately doubles. The

' Universite Claude Bernard, Centre des Sciences de la Terre, URA 1 1 du CNRS, 27-43, boulevard du 11 Novembre, F-69622 VILLEURBANNE Cedex, France. 2 Research Associate, Department of Invertebrates, American Museum of Natural History; Associate Professor, Biology Department, Touro College, New York, N.Y. 10036.

Fig. 1. Index map ofcollecting localities in the "Pink Chonetes" Zone, Onondaga Limestone, central New York. Numbers refer to AMNH Localities (see text for detail).

"Pink Chonetes" Zone (Oliver, 1954) in cen- free limestone unit with abundant "Stropho- tral New York is located in the Seneca Mem- mena (Chonetes)" lineata above the Comif- ber, 10 ft above the base of a greenish gray erous Limestone, which he referred to as the to ochre-colored volcanic ash layer, the Tioga Seneca Limestone. He considered the Seneca Bentonite (fig. 2) and is characterized in many to be a separate unit, as is evidenced by his localities by numerous stylolites. Overlying annual report (1839) and final report (1842). the Seneca in central New York is the Union Workers have studied the biostratigraphy of Springs Black Shale, a correlative ofthe Mar- the "Pink Chonetes" Zone (Oliver, 1954; cellus Shale in the Cherry Valley area, which Feldman and Lindemann, 1986), its com- represents a westerly advance of terrigenous munity structure (Feldman, 1980; Linde- sedimentation and relatively deep, poorly ox- mann and Feldman, 1981), the taxonomy of ygenated water (Feldman, 1980). its brachiopod fauna (Feldman, 1985), and Vanuxem (1839) recognized a nearly chert- even reasons for the "pink" coloration of its 1990 RACHEBOEUF AND FELDMAN: CHONETACEAN BRACHIOPODS 3

dominant chonetid brachiopods (Zenger, Jemesville Quarry Stockbridge Falls, N.Y. 1967). Oliver (1954), in his classic study of AMNH Loc. 3128 AMNH Loc. 3129 the Onondaga Limestone in central New York, referred to the "Pink Chonetes" Zone "Pink Chonetes"Zone of the Seneca Member as Zone J. He described it as a very thinly bedded limestone, about 4 ft thick and composed almost entirely of shells of C. lineatus, many of which are stained pink. Oliver (1954) noted that a bed ofchert, similar to that ofZone I (8 ft thick), occurs at most localities about 6 in. below the top ofZone J in western New York. This chert bed, according to Oliver, includes the hyolithid Coleolus crenatocinctum and gastropod Loxonema sicula. Oliver (1954: 641) found the following brachiopod fauna in the "Pink Chonetes" Zone of western New York: Camarotoechia tethys Athyris spiriferoides Atrypa "reticularis" "Schuchertella" pandora A. spinosa Levenea lenticularis Coelospira camilla "Chonetes" lineatus Atlanticocoelia acutipli- Chonetes mucronatus cata Pentagonia unisulcata Elythafimbriata Meristella nasuta Acrospirifer duodenarius He also noted the presence of the trilobite Odontocephalus selenurus and fish teeth. LEGEND In central New York, Oliver (1954: 635) found the following brachiopod fauna in the "Pink Chonetes" Zone: LIMESTONE Coelospira camilla Levenea lenticularis I I Atrypa "reticularis" "Schuchertella" pandora MASSIVE LIMESTONE A. spinosa Chonetes "lineatus" Chonostrophia reversa Meristella sp. A He also found the corals Amplexiphyllum SHALE hamiltoniae and Heterophrentis sp. B. Feldman (1985) found the following additional brachiopods in the "Pink Chonetes" DARK CHERT Zone in central New York (AMNH locs. 3128A, 3129): Fig. 2. Measured stratigraphic sections of Megakozlowskiella rari- Pentamerella arata AMNH Loc. 3128 and 3129, Onondaga Lime- costa Athyris sp. A stone, central New York. Leptaena aff. "rhomboi- Megastrophia dalis" orthotetacids indet. Additional faunal constituents found by In this paper we revise the taxonomy of Feldman (1985) include the corals "Hetero- the chonetacean brachiopods found in the phrentis" and Amplexiphyllum, trilobites "Pink Chonetes" Zone of the Onondaga Phacops cristata and Odontocephalus, euom- Limestone in central New York and redefine phalacean gastropod fragments, and camer- the community based on new paleoecologic ate? crinoid columnals. and taxonomic information. 4 AMERICAN MUSEUM NOVITATES NO. 2982

ACKNOWLEDGMENTS DIAGNOSIS: Chonetid with vertical disym- metrically arranged hinge spines (4'- 1, 2, 3). We thank Drs. Arthur J. Boucot (Depart- Brachial interior with a median septum not ment of Zoology, Oregon State University), supporting the cardinal process. Alveolus well Paul Copper (Department of Geology, Lau- developed in juvenile brachial valves. An- rentian University), Neil Landman (Depart- deridia posteriorly fused with the elevated ment of Invertebrates) and David Grimaldi medial part ofthe inner cristae. No accessory (Department of Entomology) both of the septa. American Museum of Natural History, for REMARKS: Hallinetes is provisionally as- critically reading and commenting on the signed to the subfamily Devonochonetinae manuscript and offering valuable suggestions based on brachial interior morphology. The for improvement. lack of accessory septa links the new genus ABBREVIATIONS to the "Devonochonetes coronatus" group (see Racheboeuf, 1981: 141; Racheboeuf and Institutions and Localities Branisa, 1985: 1439 for discussion). Halli- AMNH American Museum of Natural netes can easily be distinguished from all oth- History, Department of Inverte- er chonetacean genera by its spines which are brates perpendicular to the hinge line and disym- AMNH Loc. American Museum of Natural metrically arranged. History locality number Morphology and arrangement of spines ANSP Academy of Natural Sciences, of Philadelphia closely resemble those the genus Johnson- NYSM New York State Museum, Albany etes Racheboeuf, 1987, from the Emsian Blue USNM United States National Museum Fiord Formation of the Canadian Arctic Ar- ofNatural History, Department of chipelago. However, Hallinetes lacks the me- Paleobiology, Smithsonian Insti- dian enlarged capilla and differs in its brachi- tution al interior, which is strongly pustulose with more divergent anderidia, and by the pres- Measurements ence of a large alveolus. (L) maximum length of shell Hallinetes resembles Aseptonetes Isaacson, (W) maximum width of shell 1977 from the Emsian Sicasica Formation of (T) maximum thickness of shell Bolivia, in its radial ornamentation as well mm millimeters as in spine morphology, but the latter pos- est. estimated b.v. brachial valve sesses symmetrically arranged spines; both p.v. pedicle valve genera lack accessory septa. b.p. bedding plane The posteromedian part ofthe brachial in- art. articulated terior of Hallinetes is somewhat similar to ext. exterior that ofthe genus Saharonetes Havlicek, 1984, from the Lower Carboniferous Ashkidah and SYSTEMATIC PALEONTOLOGY Marar formations of Libya, as the inner cristae medially bend posteriorly toward the in- PHYLUM BRACHIOPODA ner lobes ofthe cardinal process. But in Hal- ORDER STROPHOMENIDA linetes the inner cristae are much better SUPERFAMILY CHONETACEA BRONN, 1862 developed, spines are oriented vertically in- FAMILY CHONETIDAE BRONN, 1862 stead ofobliquely, and there are no accessory SUBFAMILY DEVONOCHONETINAE septa. MUIR-WOOD, 1962 Although Hallinetes resembles Johnson- Hallinetes, new genus etes, Aseptonetes, and Saharonetes there is no Figures 3-5 generic relationship evident between them, only morphological similarities. TYPE SPECIES: Strophomena lineata Con- Species questionably assigned to Halli- rad, 1839. netes, new genus: ETYMoLoGY: In honor of New York State paleontologist James Hall. Strophomena setigera Hall, 1843. 1990 RACHEBOEUF AND FELDMAN: CHONETACEAN BRACHIOPODS 5

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Fig. 3. Hallinetes lineatus (Conrad, 1839). A. Pedicle valve, original of Chonetes glabra Hall, .-;t D 1857, 1867, pl. 20, fig. 3d, AMNH 37235, x3. B. Pedicle valve, figured by Hall, 1867, pl. 20, fig. 3a, AMNH 37233a, x 3. C. Pedicle interior, Hall's original, 1867, pl. 20., fig. 3e, AMNH 37233c, x 3. D. Brachial exterior, Hall's original, 1867, pl. 20, fig. 3c, AMNH 37233B, x 3.

Chonetes cf. setigerus (Hall), Kindle, 1912: 71, pl. 3, fig. 17. Hallinetes lineatus (Conrad, 1839) Figures 3-5 Strophomena lineata Conrad, 1839: 64; Vanuxem, 1842: 139; Hall, 1843: 175, fig. 8. Fig. 4. Hallinetes lineatus (Conrad, 1839). A. Chonetes glabra Hall, 1867: 117, figs. 1-8. Pedicle valve, block 2, AMNH Loc. 3128, AMNH Chonetes lineata Hall, 1867: 121, pl. 20, fig. 3; 43693, x3. B. Pedicle valve, AMNH Loc. 3128, Hall and Clarke, 1892: pl. 16, fig. 34. AMNH 43694, x 3. C. Latex mold ofan immature "Chonetes" lineatus Zenger, 1967: 161, fig. 1. pedicle valve, AMNH Loc. 3128, AMNH 43695, "Chonetes" aff. lineata Feldman, 1985: 321, figs. x 3. D. Pedicle interior, AMNH 43695, x 3. E. 26-28 (non 25). Latex cast of a damaged immature brachial inte- NEOTYPE: Unsuccessful were rior, AMNH Loc. 3128, AMNH 43696, x3. F. attempts Latex cast ofa brachial interior, AMNH Loc. 3129, made to locate the genotype of Hallinetes at AMNH 43697, x 3. G. Latex cast ofa large, dam- various institutions (USNM, NYSM, ANSP, aged brachial interior, AMNH Loc. 3128, AMNH AMNH) for purposes of comparison. Since 43698, x 3. H. Brachial interior, latex, AMNH the genotype is apparently lost we are herein Loc. 3128, AMNH 43695, x3. designating a neotype, AMNH 37233a (fig. 3B herein). TYPE LOCALITY: Unknown; according to 1), and gave the first complete description of Hall, 1843: 175, this shell is abundant in Sen- the species. He considered Vanuxem as the eca County but rare farther west. Substitute author, and stated that the previously de- type locality herein designated as AMNH Loc. scribed species Chonetes glabra Hall, 1857 3128, Jamesville Quarry, Jamesville, New was a junior synonym of Chonetes lineata York. Vanuxem (Hall, 1867: 121). REMARKS: The species name lineata was EXTERIOR: Small shell with maximum first used by Conrad (1839), associated with width at hingeline in juvenile specimens but the genus name Strophomena. James Hall was at midlength in adults. Maximum length the first to establish the assignment of the about 9.0 mm; corresponding width about species lineata to the genus Chonetes (table 11.0 mm; length/width ratio decreasing from 6 AMERICAN MUSEUM NOVITATES NO. 2982

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0.95 to 0.75 during ontogeny. Longitudinal mens; length/thickness decreasing from 7.5 profile weakly concavoconvex in small shells, to a mean value of 2.4 during ontogeny. Ped- becoming strongly arched in larger speci- icle valve weakly flattened at the top. Umbo 1990 RACHEBOEUF AND FELDMAN: CHONETACEAN BRACHIOPODS 7

TABLE 1 TABLE 2 Measurements (in millimeters) of Chonetes lineata Distribution of Hinge Spines on Hall, 1867 (originals of Chonetesglabra Hall, 1867) Hallinetes lineatus No. No. spines 4' 1 2 3 costae Distance from beak per Original (mean, mm) 3.63 0.32 0.94 2.46 Specimnen (L) (W) (T) mm specimen No. observations 5 3 23 16 AMNH 37235 8.2 - 4.1 4 Hall, 1867, p1. pl. 20, fig. 3d AMNH 37233a 5.8 8.0 - 5 Hall, 1867, lectotype p1. 20, fig. 3a cle field with subcircular adductor scars and AMNH 43707 5.8 8.0 2.2 5 subtriangular, anteriorly rounded diductor AMNH 43708 3.0 4.0 0.7 5 scars. Muscle field extending anteriorly up to AMNH 43709 6.8 8.5 2.8 5 one-third ofthe valve length. Vascular trunks AMNH 43710a 5.8 8.2- - originating at the anterior margin of adduc- AMNH 43711 5.3 7.4 1.8 5 tors. Posterior ridges relatively low, wide, and AMNH 43712 3.4 4.6 0.8 5 rounded, anteriorly divergent at about 1000. AMNH 43713 7.5 10.0 3.0 5 Teeth stout, transversely elongate, subpar- AMNH 43714 6.5 8.3 2.6 5 allel to hingeline. Visceral cavity deep, a Pedicle interior. smooth, well defined anterolaterally. Pos- terolateral parts of valve coarsely pustulose; anterolateral margins with radially arranged endospines. relatively small, narrow, strongly arched lon- BRACHIAL VALVE INTERIOR: Posteromedi- gitudinally. Ears small, triangular, often ill an part of valve deeply depressed between defined and smooth. Ornamentation of low, cardinal process, median septum, and prox- rounded radial costellae with wider rounded imal part of inner cristae. Median septum interspaces; costellae increasing by interca- forms low, rounded ridge extending progres- lation on the pedicle valve and by bifurcation sively toward anterior margin during growth; on the brachial valve; on the pedicle valve length about 1/4 valve length in small shells, the first intercalation occurs about 1.2 mm reaching 2/3 valve length in largest specimens. from the beak. Two mm anterior of beak, Outer cristae reduced to very low and narrow costellae number 5 per mm; along anterior linear ridges along hingeline. Inner cristae margin they number 5-6 per mm. Costellae strongly developed as two stout rounded progressively narrower from beak to anterior ridges almost parallel to hingeline, not fusing margin as intercalations increase. Total num- medially with anterior part of cardinal pro- ber of costellae between 72 and 84 for shells cess. Posterior edge of inner cristae deeply which are more than 6.0 mm in length. Due notched by dental sockets. Cardinal process to difficulties in preparing the material the internally bilobed, each lobe fusing antero- pseudodeltidium and chilidium have not been laterally with proximal part of inner cristae. observed. Pedicle interarea concave and ap- Myophore quadrilobed. Anderidia anteriorly sacline; brachial interarea almost linear and divergent at about 50-55°, as straight, nar- hypercline. In largest shells, three spines are row, and elongated ridges posteriorly fusing inserted on the right side of the beak, only with inner cristae. Adductor scars deeply im- one on left side. Spines are straight, perpen- pressed in the valve floor of largest shells. dicular to hingeline, of orthomorph type. Brachial platform strongly convex, elevated Their distribution is: 4'- 1, 2, 3 (see fig. 4A) above median septum, pustulose in juvenile (table 2). specimens, smooth in adult shells. Postero- PEDICLE VALVE INTERIOR: Visceral cavity lateral part ofvalve between inner cristae and well defined. Stout myophragm extending an- brachial platform depressed. Inner surface teriorly to midlength or two-thirds ofthe valve covered with relatively strong endospines, ra- in largest specimens; myophragm narrows dially arranged, except for the inner cristae, anteriorly, dividing a deeply impressed mus- muscle field and brachial platform. 8 AMERICAN MUSEUM NOVITATES NO. 2982

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.; j E $. Fig. 6. Longispina mucronata (Hall, 1843). A. lhla. Lectotype, pedicle valve with spines. Hall's orig- Fig. 7. Longispina mucronata (Hall, 1843). A. inal, 1867, pl. 21, fig. 1, AMNH 37246, x3. B. Pedicle valve with spines, AMNH Loc. 3128, Brachial exterior, original of Chonetes laticosta AMNH 43699, x 5. B, C. Pedicle valve, lateral Hall, 1857, figured in Hall, 1867, pl. 20, fig. 2, and ventral views, AMNH Loc. 3123128, AMNH AMNH x 3. C.Brachial 37244, interiors, original 43700, x 3. D. Latex cast of a brachial valve, of Chonetes scitula Hall, 1857, figured in Hall, AMNH Loc.3123129, AMNH 43701, x 3. E. La- 1867, pl. 21, fig. 4f, AMNH 37254, x3. tex cast of an incomplete pedicle valve, AMNH Loc. 3123128, AMNH 43706, x 3. MATERIAL: Several hundred isolated valves and articulated shells, all from two localities Union Springs Black Shale. The Seneca here in the "Pink Chonetes" Zone, central New is a fine-grained, jointed limestone, medium York. to thinly bedded, heavily weathered and STRATIGRAPHIC OCCURRENCE: The shells muddy on weathered surfaces but dark gray were collected from two localities, both with- on fresh surfaces. in the Seneca Member, Onondaga Lime- stone. SUBFAMILY RETICHONETINAE The first (AMNH Loc. 3129A) is in the MUIR-WOOD, 1962 Jamesville Quarry #3 pit which is the most GENUS LONGISPINA COOPER, 1942 complete section ofthe Onondaga in the central part ofNew York. Collecting in this quar- Longispina mucronata (Hall, 1843) ry is difficult since it is very active. Not only Figures 6-8, 12 is it hard to find weathered bedding surfaces but the rocks are continually being blasted Strophomena mucronata Hall, 1843: 180, fig. 3. and removed, making it difficult to locate and Chonetes laticosta Hall, 1857: 119. recover fossil material. The Seneca Member Chonetes mucronata Hall, 1867: 124, pl. 20, fig. here is 14 ft thick, with the Tioga Bentonite 1; pl. 21, fig. 1. at the base. Ten feet above the Tioga is found Chonetes mucronata Hall and Clarke, 1892: pl. the "Pink Chonetes" Zone which consists of 16, figs. 6, 7; 1894, pl. 20, fig. 3. a fine-grained limestone with a fresh dark TYPE SPECIES: Chonetes emmetensis Win- gray surface. There are wavy contacts be- chell, 1866: 92. tween bedding planes and stylolites present. LECTOTYPE: Pedicle valve, AMNH 37246, The second (AMNH Loc. 3129) is located figured in Hall, 1867 (pl. 21, fig. la-c) (fig. just west of Stockbridge Falls, New York, 6A herein; table 3). along Stockbridge Falls Road and Oneida TYPE LOCALITY: Ontario County, New York Creek. Here, 10-15 ft ofthe Seneca Member State (more specific information unavailable; is exposed, with an upper contact with the Hall, 1857: 120, described the locality as a 1990 RACHEBOEUF AND FELDMAN: CHONETACEAN BRACHIOPODS 9

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limestone a few miles southeast of Buffalo of the beak of largest specimens; spines typ- and in shales ofthe Hamilton Group on Can- ical for the genus and symmetrically ar- andaigua Lake). Substitute type locality herein ranged. designated as AMNH Loc. 3128, Jamesville PEDICLE VALVE INTERIOR: The myophragm Quarry, Jamesville, New York. does not exceed one-third valve length. Mus- EXTERIOR: Small shells, moderately con- cle field deeply impressed on valve floor with cavoconvex with maximum width at hinge- well-defined diductor scars. Adductors nar- line. Anterior margin regularly rounded; lat- row, posteriorly situated. Vascular trunks eral margins parallel or slightly divergent strongly developed, anteriorly divergent from posteriorly. Maximum length about 6.0 mm; anterior end of myophragm. Visceral cavity corresponding width about 7.5 mm. Length/ well defined, peripheral margin of valve im- width ratio between 0.75 and 0.8 mm. Umbo pressed by external radial costae. Teeth not relatively small, weakly overlapping poste- observed on material available. rior edge of ventral interarea. Pedicle inter- BRACHIAL VALVE INTERIOR: Typical for the area concave and anacline; brachial interarea genus in every aspect ofits morphology, with linear. Pseudodeltidium and chilidium not a low and narrow median septum supporting observed. Ornamentation consists of rounded, radial costae; along margins costae number 12 to 24 during growth; costae usually TABLE 3 simple, progressively widening from umbo to Measurements (in millimeters) of margins. Costae wider than interspaces on L?ngispina mucronata (Hall, 1843) pedicle valve while narrower than interspaces on brachial valve. Along anterior margin cos- Total no. tae number 3 per 2 mm. Costae crossed by Specimen (L) (W) costae thin and regular concentric growth lines. Ac- cording to Hall (1867: 125) some costae orig- AMNH 37246 lectotype, p.v. 6.6 8.5 20 inate by bifurcation or by intercalation; this AMNH 37247 p.v. 3.1 4.2 13 AMNH 37244 was not observed on the specimens from the original Chonetes laticosta, p.v. 5.9 6.8 17 Seneca Member. Three spines on each side 10 AMERICAN MUSEUM NOVITATES NO. 2982

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Fig. 9. "Eodevonaria" hemispherica (Hall, 1857). A-C. Lectotype, pedicle valve in ventral (A), lateral (B), and posterior (C) views. Hall's original, 1857, p. 116, fig. 1; 1867, pl. 20, fig. 6b-d, AMNH 2825, x 2. D-G. Decalcified articulated damaged shell, brachial interior (D), pedicle interior (E), latex cast of the brachial exterior (F), and interior (G), AMNH 37224, x 2. the cardinal process and extending anteriorly to Oliver (1954, 1956). It is also found in the up to two-thirds the valve length; posteriorly overlying Hamilton Group, specifically in the elongated and narrow cardinal process; a pair Marcellus Shale. of well-developed accessory septa and an- MATERIAL: A total of 105 specimens from deridia anteriorly divergent at 700. the Hallinetes Community, mostly complete DIscussIoN: Strophomena mucronata Hall, juvenile shells; only one brachial valve and 1857, is a typical representative of the genus two pedicle valves were available for pre- Longispina Cooper 1942, as attested by the paring the interiors; AMNH Locs. 3128,3129. shell shape and spine morphology, as well as interiors ofboth valves. The original of Cho- FAMILY EODEVONARIIDAE SOKOLSKAYA, 1960 netes laticosta Hall, 1857, figured in 1867 GENUS EODEVONARIA BREGER, 1906 (AMNH 37244, pl. 20, fig. 2) is a brachial "Eodevonaria" hemispherica (Hall, exterior or L. mucronata, as stated by Hall 1857) (1867: 125). Two dorsal interiors (AMNH Figures 9-12 37254), one of which was figured by Hall Choneteshemispherica Hall, 1857: 116; 1867: 118- (1867: pl. 21, fig. 2f) as Chonetes scitula, are 119, pl. 20, fig. 6A-D. undoubtedly representatives of the genus Chonetes hemisphaerica Hall and Clarke, 1892: Longispina; according to our own observa- pl. 16, fig. 14. tions they must be assigned to the species L. TYPE SPECIES: Chonetes arcuata Hall, 1857. mucronata, but L. mucronata and "Cho- LECTOTYPE: Pedicle valve AMNH 2825 netes" scitula are distinct species. figured in 1857 (p. 116, fig. 1), 1867 (pl. 20, STRATIGRAPHIC OCCURRENCE: Longispina figs. 6b, c), 1892 (pl. 16, fig. 14). mucronata is a common species of the Ned- TYPE LOCALITY: Schoharie Grit (Hall, 1857: row, Moorehouse, and Seneca members of 116). Substitute type locality herein desig- the Onondaga Limestone; it is rare or very nated as AMNH Loc. 3128, Jamesville Quar- rare within the Edgecliff Member according ry, Jamesville, New York. 1990 RACHEBOEUF AND FELDMAN: CHONETACEAN BRACHIOPODS I1I

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Fig. 10. "Eodevonaria" hemispherica (Hall, 1857). A. Pedicle valve, latex cast, AMNH Loc. 3128, AMNH 43702, x 3. B. Juvenile pedicle valve with spines, AMNH Loc. 3128, AMNH 43703, x 3. C. Pedicle interior, AMNH Loc. 3128, AMNH 43704, x 3. D. Articulated shell in dorsal view, AMNH Loc. 3129, AMNH 43705, x 3. E, F. Damaged pedicle interior, posteroventral and ventral views, AMNH Loc. 3128, AMNH 43706, x3.

EXTERIOR: Large shell, strongly concavo- length. Diductor scars ill defined, poorly im- convex with maximum width at hingeline. pressed on valve floor, wide, radially grooved, Maximum length about 20 mm; length/width occupying posterior half of visceral cavity. ratio about 0.66; length/height ratio about Adductor scars relatively large, semioval in 0.5. Umbo widely rounded, strongly arched outline, almost semicircular; length about 4 dorsally, posteriorly overhanging hingeline mm and corresponding width about 3 mm. by more than 2 mm in largest specimens. Ears Two narrow and deeply impressed vascular triangular, strongly convex and well differ- trunks (vascula media) originate anterior to entiated from the body of the pedicle valve. adductor scars. Whole inner surface im- Ornamentation oflow, rounded radial costae pressed by external ribbing except for an- increases by bifurcation on the pedicle valve terolateral margins of valve where endo- and by intercalation on the brachial valve. spines remain distinct, radially arranged. Along anterior margin costae number 3 to 4 Hingeline strongly denticulate. per mm. At 2 mm anterior to the beak the total number of costae is about 15 while at 5 mm from the beak the number increases TABLE 4 to 35. Maximum number of costae 70 (esti- Measurements (in millimeters) of mated) in largest shells. On pedicle valve first "Eodevonaria" hemispherica (Hall, 1857) bifurcation occurs at 2.5 mm from the beak. No. Costae separated by narrower spaces. Total costae Pseudodeltidium reduced to small triangular, no. per convex plate at apex ofdelthyrium; chilidium Specimen (L) (W) (T) costae mm not observed. Pedicle interarea strongly con- AMNH 2825 cave and catacline, almost perpendicular to lectotype, p.v. 19.7 +30 10 70a 3-4 commissural plane. At least 6 spines on each AMNH 37224 side of beak; spines high angled and oblique b.v. ext. 18.5 24a - 60a 3 at their base. AMNH 37225 PEDICLE VALVE INTERIOR: Only one spec- p.v. 13.1 20a 7 64a 3 imen (AMNH 37224) was available for study. AMNH 37226 Muscle field longitudinally divided by stout art. 12a 16+ - - 3 myophragm extending anteriorly up to mid- a Estimate. 12 AMERICAN MUSEUM NOVITATES NO. 2982

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2 4 i6 8 lb 12 14 16 18 20 22 4 2628 30 32 WIDTH and THICKNESS (mm)

Fig. 1 1. "Eodevonaria" hemispherica (Hall, 1857). Scattergram showing length/width and length/ thickness. Circle denotes the lectotype.

BRACHIAL VALVE INTERIOR: Known from spherica (Weisbord), and E. inca Isaacson one incomplete internal mold only (AMNH show very close morphologies and similar 37224). A strong median septum extends an- ornamentation but more detailed informa- teriorly up to midlength; septum progres- tion is needed for better comparisons. How- sively widens posteriorly, developing a flat- ever, they would probably represent a group tened triangular platform. Anderidia of globose, coarsely costate Eodevonariids posteriorly fused with the platform, ante- distinct from Chonetes arcuatua Hall, 1857, riorly diverging at 50°. Cardinalia and other type species ofthe genus Eodevonaria Breger. structures unknown. Inner surface covered Specimens from the "Pink Chonetes" Zone with radially arranged distinct endospines. are globose, coarsely costate eodevonariids DISCUSSION: The species hemispherica (ta- which sporadically occur within this bio- ble 4) undoubtedly belongs to the family Eo- stratigraphic zone. They are here assigned to devonariidae as evidenced by its shape and the species "Eodevonaria" hemispherica. denticulate hingeline as well as other char- While mostly represented by fragmentary, acters. However, the lack of complete bra- isolated valves, this material has the char- chial interiors precludes any generic assign- acteristics of Hall's species. However, it dif- ment. For this reason Chonetes hemispherica fers from the type material in its smaller size; Hall is here assigned to "Eodevonaria" ac- shell shape ornamentation and spines are cording to recent recommendations (Rache- similar. Specimens from the "Pink Cho- boeuf, 1986). netes" Zone are probably juveniles and the The coarse radial ornamentation of this size difference does not preclude suggested species is very similar to that of several taxa specific assignment. The smaller size may also described from Central and South America. be due to environmental conditions. Eodevonaria imperialis (Caster), E. subhemi- STRATIGRAPHIC OCCURRENCE: According 1990 RACHEBOEUF AND FELDMAN: CHONETACEAN BRACHIOPODS 13

A J25 2 -J -r2 Co20- Co20- oU- ~~~~~~~~U-0 a:15 c15 co ~~~m ClLEGT (m)LNGH(m D 10- Z)10

5 5

0 12 4 67 0 2 4 6 8 1 LENGTH (mm) LENGTH (mm) Fig. 12. Histograms for 55 pedicle valves of Longispina mucronata (A) and 85 pedicle valves of Hallinetes lineatus (B) from the Hallinetes Community within the dark mudstone matrix of the "Pink Chonetes" Zone. to Oliver (1956: 1452, table 1) Chonetes most abundant species (92%); Longispina hemisphericus? is very rare in normal facies mucronata may be listed as common (5.4%) of the Edgecliff fauna and rare in the central while "Eodevonaria" hemispherica is rare facies of the Lower to Middle Moorehouse (2%). After tabulating the above results, based fauna in the Richfield Springs area (1956: on a count of 907 specimens, it became ap- table 3, p. 1462). parent that the relative abundance ofthe taxa MATERIAL: Two articulated shells; 17 bro- on bedding planes (both top and bottom) on ken, or crushed pedicle valves. one hand, and within the dark mudstone matrix on the other hand reveals significant dif- THE HALLINETES COMMUNITY OF THE ferences. "PINK CHONETES" ZONE On the studied bedding planes only very The "Chonetes" Community, which oc- few specimens of Longispina mucronata (7) curs in the mudstones ofthe Seneca Member, and "Eodevonaria" hemispherica (8) were 10 ft above the Tioga Bentonite, was de- counted together with 630 isolated valves and scribed by Feldman (1980), and was defined articulated shells of Hallinetes lineatus. The as a low-diversity assemblage in which "Cho- specimens of L. mucronata occurring on the netes" aff. lineata is highly dominant (> 99%) bedding planes are among the largest ob- with a high ratio of living shells (76%) ac- served in the material studied. cording to the concave-up position of the Within the dark mudstone matrix, 41 small specimens. While other faunal constituents shells of L. mucronata and 1 1 of "E." hem- are similar to those listed previously (Feld- ispherica have been counted together with man, op. cit.: table 12, p. 40), the community 209 specimens of Hallinetes lineatus; most needs redefinition since the chonetacean bra- of the specimens are articulated shells and chiopods belong to three different taxa. the shells of L. mucronata are very small to Among these, Hallinetes lineatus is by far the small juveniles. 14 AMERICAN MUSEUM NOVITATES NO. 2982

TABLE 5 ding surfaces, even if shells in a concave-up Counts of Hallinetes lineatus on B edding Planes position are common (see table 5). It is prob- and Within the Matri, able that the number of articulated shells in % con- the has been underes- Con- Con- concave-up position cave- vex- % cave-up timated; in some cases the brachial valve of up up artic-4I articu- such oriented specimens appears to have been Speci- posi- posi- ulatedI lated weathered off. In any case, the relative fre- Block no. mens tion tion shellsII shells quency of articulated shells in a concave-up 1 b.v. 66 22 position would not exceed twice the total Top b.p. p.v. 89 42 8.3 7.0 number of specimens found. art. 17 3 Most of the shells lying on the bedding Total 172 67 planes have broken spines while spines are 3 b.v. 10 6 nicely preserved, and always found, on shells Bottom b.p. p.v. 29 7 11.0 10.0 encased within the matrix. This fact is further art. 6 1 support for the interpretation that bedding Total 45 14 plane assemblages are post mortem accu- 4 b.v. 15 12 mulations. Top b.p. p.v. 45 27 9.1 2.7 In conclusion, from the new observations art. 3 7 of the chonetaceans from the "Pink Cho- Total 63 46 netes" Zone, the true Hallinetes Community 6 b.v. 6 15 corresponds to the assemblage within the dark Top b.p. p.v. 25 32 8.2 2.3 mudstone matrix rather than to the shells art. 5 2 distributed on the bedding plane surfaces. The Total 36 49 Hallinetes Community of the Seneca Mem- 14 b.v. 9 11 ber is a dominated" Top b.p. p.v. 30 20 6.6 6.6 low-diversity, "highly art. 5 (although not monospecific) community Total 44 31 within a quiet water environment. This in- 14 b.v. 15 6 terpretation is supported by the coexistence Bottom b.p. p.v. 25 14 6.7 3.1 oflong-spined chonetaceans as well as by the art. 2 1 reduced number of spines on the shells Total 42 21 (Racheboeuf, in prep.). I b.v. 8 14 The Benthic Assemblage position (cf. Bou- Middle bed p.v. 44 26 2.1 2.1 cot, 1975) is more difficult to pinpoint but in art. 2 this case, a Benthic Assemblage 4 position is Total 54 40 most probable since it is below the lower limit 4 b.v. 11 6 for active photosynthesis and reef building Middle bed p.v. 21 26 9.8 5.6 activity (Yu et al., 1987: 6) as evidenced by art. 4 3 a lack of reef building corals and scarcity of Total 36 35 solitary corals (Feldman, 1980: 40). A normal marine environment is indicated by the presence of some bioturbation, which increases Counting shows that the same chonetacean dramatically at the top ofthe formation. The taxa occur on the bedding planes and within fine-grained limestone matrix and domi- the mudstone matrix, but their rcclative abun- nance of Hallinetes are consistent with low dance is different. Relative frecluencies are, (to intermittent) current activity and lower respectively, as follows: Longis,pina mucro- than normal oxygen, but not quite dysaerobic nata 1 and 15%; "Eodevonaria' hemispher- conditions since trilobites, gastropods, and ica 1.2 and 4.2%; Hallinetes linieatus 97 and crinoids are present, although rare. The cause 80%. The fact that only large sheIlls lie on the of relatively low diversity here is uncertain. bedding planes probably indicaites that the tinyjuvenile shells ofL. mucronaita have been REFERENCES winnowed away. This conforms with the iso- Boucot, A. J. lated valves ofH. lineatus foundI on the bed- 1975. Evolution and extinction rate controls. 19901990RACHEBOEUF AND FELDMAN: CHONETACEAN BRACHIOPODS 15

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1987. Community paleoecology as a geologic Zenger, D. H. tool: the Chinese Ashgillian-Eifelian 1967. Coloration ofthe "Pink Chonetes" (bra- (latest Ordovician through early Middle chiopod) of the Onondaga Limestone, Devonian) as an example. Geol. Soc. New York. J. Paleontol. 41: 161-166. Am. Spec. Pap. 211: 1-100.

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