Bulletin of the American Museum Of'natural :History Volume 89: Article 1 Nw York~: 1947,

BOBB SCHAEFFER

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- | - | -| - l - CRETACEOUS AND TERTIARY ACTINOPTERYGIAN FISHES FROM BRAZIL

CRETACEOUS AND TERTIARY ACTINOP- TERYGIAN FISHES FROM BRAZIL

BOBB SCHAEFFER Assistant Curator of Fossil Vertebrates

BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY VOLUME 89: ARTICLE 1 NEW YORK: 1947 BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY Volume 89, article 1, pages 1-40, text figures 1-6, plates 1-7, tables 1, 2 Issued April 30, 1947 CONTENTS

INTRODUCTION...... 7

GEOLOGIC OCCURRENCE ...... 8

SYSTEMATIC DESCRIPTIONS ...... 12

Order Semionotoidea ...... 12

Family Semionotidae ...... 12

Lepidotus Agassiz...... 12

Lepidotus piauhyensis Roxo and Lofgren ...... 12

Order Isospondyli...... 13

Suborder Clupeoidea ...... 13

Family Leptolepidae ...... 13

Leptolepis Agassiz...... 13

Leptolepis bahiaensis, new species ...... 13

Family ?Elopidae...... 14

Lastroichthys, new genus...... 14

Lastroichthys longipectoralis, new species...... 14

Family Chirocentridae ...... 15

Chiromystus Cope ...... 15

Chiromystus mawsoni Cope...... 15

Chiromystus alagoensis Jordan...... 15

Family Chanidae ...... 16

Dastilbe Jordan...... 16

Dastilbe crandalli Jordan...... 16

Family Clupeidae...... 17

Knightia Jordan...... 17

Knightia branneri (Jordan)...... 17

Diplomystus Cope ...... 20

Diplomystus longicostatus Cope ...... 20

Preliminary Analysis of the Double-armored Clupeids ...... 21

Order Ostariophysi . 24 Suborder Cyprinoidea 24

Family Characinidae . 24 Eobrycon Jordan . 24 Eobrycon avus (Woodward). 24

Eobrycon branneri Eigenmann ...... 25 Eobrycon ligniticus (Woodward). 25 Order Acanthopterygii. 28 Suborder Percoidea ...... 28 Family Serranidae. 28

Percichthys Girard ...... 28 Percichthys antiquus Woodward. 28

Family Cichlidae ...... 29 Aequidens Eigenmann and Bray. 29 Aequidens pauloensis, new species. 29 DISCUSSION. 30 Analysis of Fossil Assemblage. 30 The Origin of the South American Fresh-water Fish Fauna 30 Marine Distribution ...... 33 Land Bridge Dispersal ...... 34 Distribution and Continental Drift. 35 Holarctic Continental Distribution ...... 35 REFERENCES 36

INTRODUCTION FISHES FROM THE CRETACEOUS OF BRAZIL lutionary and zoogeographical studies. Until were first recorded by Spix and Martius in critical analyses, based on most of the known 1828. Since that time collections of Creta- available material, are made, particularly of ceous and Tertiary actinopterygian fishes -certain genera with many species, such as from various localities have been described by Lepidotus, it would appear that a conserva- Agassiz, Cope, Woodward, Jordan, Maury, tive approach, at least towards the creation of and Roxo. additional related genera, is advisable. The The collection that forms the basis for this designation of new species based on slight and study was sent to the American Museum of possibly statistically invalid differences seems Natural History in 1931 by Dr. Euzebio de inevitable, however, unless critical studies Oliveira, former Director of the Geological are possible and the range of variation within Survey of Brazil. It includes material from such species is determined. A similar situa- localities in the States of Sao Paulo, Alagoas, tion exists for the genus Leptolepis, as noted Piaui, Baia, and Sergipe. Dr. Louis Hussakof by Dunkle (1942, p. 64) and confirmed by the partially examined this collection at the time present writer. In this case it may prove of its reception and made some preliminary necessary to revise the entire family Lepto- identifications, but unfortunately circum- lepidae. stances prevented a detailed study until The problem of the origin of the Recent undertaken by the present writer. Excepting South American fresh-water fish fauna is still the absence of material from the rather fa- unsolved. An attempt is made in the discuss- mous localities of the Serie Araripe, Cear6, this ion, however, to evaluate the evidence pre- collection is representative of most of the sented by the fresh-water fossil fishes thus well-known localities. In addition, unde- far discovered in Brazil. It is obvious that scribed material from several new localities this question will require far more fossil evi- is herein included. dence for a solution as satisfactory as that A satisfactory study of fossil fishes is lim- obtained for the origin of the Tertiary mam- ited to a great degree by the usual rather poor malian fauna of South America. preservation of the material. Fossilization, The writer wishes to acknowledge his grati- such as occurs in the Green River and Mount tude to Dr. W. K. Gregory, Dr. G. G. Simp- Lebanon shales and certain localities in son, Dr. C. M. Breder, Jr., and Dr. E. H. Europe, is exceptional, but highly desirable Colbert for critically reading the manuscript not only for morphological but also for quan- and for helpful suggestions. He is also in- titative studies. Very often important diag- debted to Mr. J. T. Nichols for assistance in nostic skeletal characters are irretrievably certain problems related to recent ichthyology smashed or obscured during preservation and to Dr. L. I. Price of the Division of and, as is well known, dissociated fish remains Geology and Mineralogy, Ministry of Agri- are far more common than complete or par- culture, Brazil, for information on the geology tially complete skeletons. This situation has and the collecting localities. Dr. M. de Oli- naturally led to differences of opinion in veira Roxo, Director of the Division of Geol- identification and certainly to an extreme ogy and Mineralogy, very kindly supplied state of taxonomic inflation in many groups, several useful maps. The author is obligated such as in the genus Lepidotus, with over 90 to Mr. J. Le Roy Kay for the loan of speci- described species. Failure to compare new mens from the Carnegie Museum. material with the genotype has also led to The following abbreviations have been much confusion, as formerly existed in the used in referring to catalogued specimens: case of the actinistian Coelacanthus. These' problems are all too evident to anyone who A.M.N.H., American Museum of Natural History has worked with fossil fishes and are here B.M.N.H., British Museum (Natural History) mentioned merely to emphasize certain in- C.M., Carnegie Museum herent difficulties in dealing with this type of Col. Pad. Serv. Geol. Min., Collecgao Padrao do material, particularly in connection with evo- Serviro Geologico e Mineralogico of Brazil 7 GEOLOGIC OCCURRENCE

TABLE 1 represents an attempt to assign a fortunate that the fish fauna cannot be reli- definite age to most of the localities in Brazil ably used for making definite determina- from which fossil fish have been obtained. tions. It is based mainly on the recent work of The Lower Cretaceous is represented by Maury (1930, 1936), E. P. de Oliveira (1939), deposits in the States of Pernambuco, Ser- and A. I. de Oliveira and Leonardos (1943). gipe, Bafa, and Rio Grande Do Norte. The The last-named work has been particularly S6rie Baia extending northward from Baia helpful in that it represents a synthesis of de Todos os Santos, and the Formagao most of the work accomplished since early in Almada, a localized deposit north of Ilheus, the last century. There is still some difference Bala, were formerly placed in the Lower Cre- of opinion regarding the age of several of taceous. Roxo (1936) considers the S6rie the localities and formations, and it is un- Baia to be Upper Cretaceous on a faunistic TABLE 1 Period or Epoch Stage Locality Nature of Sediments Unknown Taubate and Tremembe, Sao Shale, lacustrine-fluvial Paulo ?Pliocene Unknown Nova York, Maranhao Shale, probably lacustrine- Cd fluvial Formario Maria Farinha, Limestone, marine ?Lower Sparnacian or Pernambuco Eocene Thanetian S6rie Alagaos in vicinity of Bituminous shale, estu- Riacho Doce, Alagaos arine Maestrichtian Rio Gramame Limestone, Limestone, marine Paraiba ?Senonian (formerly Serie Bala exposed along east- Shales and sandstones, es- considered Lower ern shore of Baia de Todos tuarine Neocomian) os Santos Fomagao Almada, Bafa Shale, estuarine Upper U2 S6rie Aranrpe o a. Formaglo Santana, Concretions in sandstone Cear& and limestone, marine Turonian or Ceno- U manian (S6rie GrajaCi) b. Isolated deposit, east of Limestone, marine Floriano, Piaui c. Barra do Corda, Maran- ? hfo Lastro Oolitic Limestone Oolitic limestone, marine Lower Middle or Upper Al- (S6rie Sergipe, Grupo Gan- bian hamaroba), Rio Ganhama- roba, Sergipe 1-947 SCHAEFFER: ACTINOPTERYGIAN FISHES FROM BRAZIL 9 basis, and a copy of the "Atlas Geologico Do showing faunal relationships with the Serie Brasil" (E. P. de Oliveira, 1939) recently re- Araripe (Rego, 1923). ceived has been annotated by Dr. M. J. de The Rio Gramame Limestone, considered Oliveira Roxo, placing both the Serie Baia to be Maestrichtian (Maury, 1930), is repre-; and the Formagao Almada in the Upper sented by an isolated deposit in eastern Cretaceous (? Senonian). This determination Parajba. From it have been obtained a pyc'. is more in agreement with the Upper Creta- nodontid and a stomiatoid. ceous nature of the fish fauna, particularly The Serie Alagoas is generally considered with the presence of a diplomystid and an to be an Eocene estuarine deposit consisting amiid. The Serie Baia is an estuarine deposit of bitumino,us shales. It is exposed along the of shales and sandstones and has been de- coast of Alagoas, in some places only at low scribed most completely by Woodward and tide, at about six localities from a point south Mawson (1907). Most of the fossil fishes from of Macei6 northward to Maragogi and is this series have actually been found in the overlain by thick beds of probably Miocene- shales exposed along the eastern shore of the Pliocene age. The only locality that has bay, north of the city of Salvador. The For- yielded a fossil fish fauna is the village of magdo Almada has produced lepidotid re- Riacho Doce, described by Jordan (1910). mains and a single species of clupeid. The Formago Maria Farinha, consisting The Grupo Ganhamaroba, more specifi- of a series of localized marine limestones cally the Lastro Oolitic Limestone of the along the coast of Pernambuco, is considered Serie Sergipe, has yielded several specimens to be basal Eocene, equivalent to the Midway from beds exposed along the Rio Ganham- group of Texas by Maury (1930, p. 33). Oli- aroba, a branch of the Rio Sergipe. This veira and Leonardos (1943, p. 425) consider deposit is considered to be Middle or Upper the formation to be Upper Cretaceous (Dan- Albian and, with the assignment of the ian), but do state that it may be Paleocene Serie Baia and Formagao Almada to the (op. cit., p. 622). It is considered as basal Upper Cretaceous, this is the only recorded Eocene in the table. A fragment of a stomia- Lower Cretaceous deposit containing fish toid was described by Cope from this forma- remains. tion. The Upper Cretaceous deposits are rela- A lacustrine-fluvial deposit of unknown tively very extensive, particularly in Mar- Tertiary age is present in the southeastern anhao, western Baja, and Matto Grosso, with portion of Sao Paulo. Two localities, near the smaller exposures in Ceara, Rio Grande Do towns of Taubate and Tremembe, have Norte, Minas Gerais, and Goyaz. The richest yielded a small fauna first described by fossil fish fauna and probably the best known Wood'ward in 1898. Woodward (1939) has is that from various localities in the Formaglo described two fossil fishes from an unrecorded Santana of the Serie Araripe, Ceara. This is Tertiary formation at Nova York on the Rio a marine formation considered to be Turon- Parnalba in Maranhao, and he believes the ian and consisting of shales and sandstones fossils indicate early Tertiary age, possibly with the fossils occurring in concretions which Eocene. are found for the most part in the shales. The wide separation of fossiliferous locali- While Spix and Martius first reported fossils ties of similar age would seem to indicate from here in their "Reise in Brasilien" pub- that the actinopterygian fauna, as it is now known, represents but a small sample of the lished in 1828, the most complete studies of material that might be obtained with more the fauna are those of Jordan (1908 and 1921). intensive collecting. As will be discussed The most distinctive feature of the Serie later, fossil fishes from fresh-water deposits, Araripe is the large preponderance of elopids, in particular, are of primary importance in a group probably represented in but one other throwing some light on the unsolved problem known Brazilian locality. of the dispersal of the Recent South American Upper Cretaceous beds near Floriano, fresh-water fishes. The fauna thus far dis- Piaui, and Barra do Corda, Maranhao, have covered unfortunately offers little more than yielded a small amount of material, the latter a few tantalizing suggestions on this problem. 10 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL.- 89

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Scale 1:5,000,000

ALMADA Qo KEY m Tertiary L Upper Cretaceous ILower Cretaceous

FIG. 1. Map of northeastern Brazil showing the Cretaceous and Tertiary deposits and localities from which fossil fishes have been obtained. This map is based on the Geological Map of South America published by the Geological Society of America (1945) and on various other sources mentioned in the text. 1947 SCHAEFFER: ACTINOPTERYGIAN FISHES FROM BRAZIL 11

FIG. 2. Map of Sao Paulo and adjacent states to show particularly the Tertiary deposits around Tremembe and Taubate. The scale, key, and sources of this map are the same as in figure 1. SYSTEMATIC DESCRIPTIONS ORDER SEMIONOTOIDEA Santos (1945). They agree in relative size FAMILY SEMIONOTIDAE and arrangement with the typical lepidotid GENUS LEPIDOTUS AGASSIZ, 1832 condition as described by Dechaseau (1943) (For generic synonomy and generic diag- for L. elvensis. There appears to be a reduc- nosis, see Woodward, 1895, p. 77.) tion, however, in the number of postorbitals Blain- with a single element under the eye rather GENOTYPE: Lepidotus elvensis (De than a series of four or five. ville). Lepidotus temnurus, described by Agassiz Lepidotus piauhyensis Roxo and Lofgren, 1936 (1841) from the Serie Araripe, has scales that Lepidolus piauhyensis RoXO AND LOFGREN, are characteristically marked with three 1936, p. 7, figs. 1, 2. coarse ridges, and the lateral line scales are Lepidotus piauhyensis, SANTOS, 1945, p. 1, pls. distinctly elevated. Lepidotus mawsoni was 1-4. founded by Woodward (1888) on detached TYPE: Col. Pad. Serv. Geol. Min. No. 3447. fragments from the Serie Bala. The scales Complete fish excepting imperfectly pre- show occasional faint posterior radiating served fins. grooves with some crenulations but are other- HOlUZON AND LOCALITY: S6rie Araripe wise smooth. They are considered to be un- (Grajau), Formagao Santana, Upper Cre- usual in the great thickness of the laminated taceous, near Floriano, Piaui. bony base, but unfortunately no measure- REVISED DIAGNOSIS: Length about 56 cm., ments of this layer are given to make com- greatest depth 20 cm. Head about one- parison possible. The bony layer of the scales quarter total body length. Dorsal outline of L. piauhyensis in cross section appears to elevated anterior to dorsal fin. Dermal bones be of a thickness characteristic of the genus. of skull tuberculated, distribution and con- L. piauhyensis resembles L. mawsoni in the centration of tubercles not constant. Parietals presence of discontinuous enamel in the flank- about one-third length of frontals, maximum scale area and in the development of peculiar width of opercular somewhat more than one- rounded excrescences, possibly developed half greatest length. Scales smooth, without from the bonv base of the scale, although this ornamentation in any region of body. Scale cannot be demonstrated in ground sections. enamel usually lacking in some areas. Dorsal 9, One specimen in this collection (pl. 1, fig. 1) pectoral 12, pelvic 8-9, anal 8. Origin of has two such protuberances in the flank area. dorsal fin about halfway between origin of Lepidotus souzai was described by Wood- pectoral and origin of anal fins. ward (1908) from beds probably referable to REMARKS: The two specimens of Lepidotus the Formagao Almada. The bones of the piauhyensis in the present collection, A.M. skull are tuberculated as in L. piauhyensis, N.H. No. 10012 and A.M.N.H. No. 10013, but the scales are covered with thickened were obtained in 1913 from a locality near enamel and have six to eight radiating fur- Riacho Olhos d'Agna, Floriano, Piarui. The rows. The ridges between the furrows are type description is based on a third specimen covered with tubercles. L. souzai and L. found between the towns of Amarante and piauhyensis are both short and relatively Floriano on the right bank of the Rio Par- stout species and in this respect resemble L. naiba. The latter specimen is deposited in mantelli from the Lower Cretaceous of Eu- the collection of the Servico Geologico e Min- rope. Weiler (1935) considers a lepidotid from eralogico, Brazil. the Upper Cretaceous of Egypt to have an Lepidotus (pl. 1, fig. 1) is obviously dis- affinity with L. souzai because of similar scale tinct from the three previously described characters. species of the Brazilian Cretaceous, although There is thus reasonably good evidence, it does share certain characters in common based mainly on distinctive scale characters, with several European forms. The dermal that four species of Lepidotus occur in the elements of the skull have been studied by Cretaceous formations of Brazil. Of the four 12 1947 SCHAEFFER: ACTINOPTERYGIAN FISHES FROM BRAZIL 13 species described from North America, scale TYPE: A.M.N.H. No. 10014. Complete characters are again relied upon almost en- fish with somewhat imperfect fins. tirely and usually through necessity. East- HORIZON AND LOCALITY: ?Serie Baia, man (1917), for instance, compared L. wal- Upper Cretaceous, Bacuparytuba, Bafa. cotti from the Kiowa Shales of Kansas with (Exact location of Bacuparytuba unknown to L. gallineki from the Rheatic of Upper Silesia Brazilian Survey.) with which he considered it to be closelyre- SPECIFIC DIAGNOSIS: Fish of small size, lated, presumably on the basis of scale mor- ranging between 4.4 and 5.7 cm. in length. phology. There is a very close resemblance Head somewhat less than one-quarter total between the scales of L. piauhyensis and length of body. Origin of pelvic fin opposite those of L. congolensis Hussakof (1917) from origin of dorsal fin, origin of anal fin halfway the possibly Upper Triassic Lualaba Beds between origin of pelvic and origin of caudal of the Belgian Congo, and, for that matter, fins. Vertebrae about 38, of which approxi- with several of the North American and mately 15 are caudal. Maxillary a wide European species. curved plate, dentary of characteristic shape Many of the 90 odd described species of with robust coronoid process. this world-wide genus are based on scale REMARKS: This is the first record of Lepto- characters alone, and it is very tempting to lepis from Brazil, excepting an otolith from see genetic affinities in species with similar the Serie Araripe considered to be leptolepid scale morphology. While it is true that scale by Erasmo (1938). L. australis has been de- characters may turn out to be as constant sciibed from Argentina (De Saez, 1939), rep- and reliable in the Holostei as they appear to resenting apparently the one other described be in the Teleostei on the basis of the work species from South America. of Cockerell and David, the significance of The skull of L. bahiaensis (pl. 2, fig. 2) close similarity, for instance in Lepidotus, appears to conform closely to the typical between the scales of species from widely leptolepid pattern. The premaxillary is char- separated regions and from different ages is acteristically small, apparently represented yet to be determined. The role played by par- by an enlarged area at the distal end of the allelism is, for instance, unknown. maxillary. The supramaxillaries are not well The key worked out by Woodward (1895, preserved and cannot be properly distin- p. 82) for about 16 species of Lepidotus in- guished from each other or from the maxil- volves the relative length of the frontals and lary. Dermal elements of the upper jaw as a parietals, relation between breadth and depth whole resemble closely those found in L. of the opercular, tooth form and scale mor- sprattiformis from the Upper Triassic of phology. Unfortunately, such a key cannot be Solenhofen. developed for a majority of the species, as in- The circumorbital series is missing in the formation on most of these characters is not type, revealing a rather badly crushed palatal available. complex that resembles very closely Rayner's restoration (1937) of this region in the geno- ORDER ISOSPONDYLI type L. coryphaenoides. The articulation be- SUBORDER CLUPEOIDEA tween the quadrate and articular is clearly FAMILY LEPTOLEPIDAE defined, indicating a suspensorium swung GENUS LEPTOLEPIS AGAssIz, 1832 forward to about the same degree as in the genotype. The opercular elements are also (For generic synonomy and generic diag- somewhat crushed but appear to be typical nosis, see Woodward, 1895, p. 501). in relative size and arrangement. GENOTYPE: Leptolepis coryphaenoides About four branchiostegal rays are present (Bronn). below the ventral margin of the opercular Leptolepis bahiaensis,' new species region, and are typically broad and flat. A portion of the left hyoid arch is preserved. 1 From Bahia (also spelled Bafa), the state in which The distal ossification of the ceratohyal ap- the specimens were found. pears to be hourglass-shaped with possibly a 14 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 89 bar of bone running between the proximal Leptolepis on the basis of resemblance in the and distal portions along the dorsal border vertebrae and dentary. Although leptolepids as in L. coryphaenoides. have been reported from Cuba and Mexico, Most of the dermal portion of the pectoral the remains thus far described are too frag- girdle is preserved, but separate elements mentary for comparison. cannot be recognized. The cleithrum has a Leptolepis schoewei Dunkle (1942), from rather prominent longitudinal ridge as in the the probable Upper Triassic of Colorado, is genotype, while the posterior expansion at a much more primitive type with an uncon- the lower end is relatively greater than in stricted notochord and ring-like centra. L. the genotype and the anterior expansion nevadensis, from the probable Lower Cre- relatively smaller. taceous of Nevada, is about the same size but The postcranial skeleton (pl. 2, fig. 1) is has a larger number of vertebrae (50). quite typical. The vertebrae are well ossified cylinders with two lateral, longitudinal lam- FAMILY ?ELOPIDAE inae. The notochordal canal is relatively large LASTROICHTHYS,1 NEW GENUS and easily demonstrated in hemisected verte- GENOTYPE: Lastroichthys longipectoralis, brae. The neural arches are delicate and at- new species. tenuated with intermuscular bones lying GENERIC DIAGNOSIS: Trunk relatively across them anteriorly. The dual nature of short, head approximately one-quarter total the former cannot be satisfactorily demon- mouth extending strated. The ribs are moderately robust with body length. Gape of wide, a longitudinal groove on the lateral side. behind orbit. Sclerotic ossified. Suspensorium The relative positions of the fins have been vertical. Suborbital region wide. Centra cu- indicated in the diagnosis. The dorsal has boidal, constricted, with single lateral lamina. about 10-12 rays, pectoral 12, pelvic about 9, Neurapophyses and haemapophyses long and and anal about 9. The pelvic girdle appears to delicate. Anal extended. be an elongated and exceedingly thin plate Lastroichthys longipectoralis,2 new species which makes it very difficult to uncover TYPE: A.M.N.H. No. 10017. Incomplete satisfactorily. The hypural elements of the fish with poorly preserved skull, dorsal fin caudal fin appear to be less expanded than and posterior portion of caudal region miss- those of L. coryphaenoides and more like ing. those in L. voithi or L. dubius. The detailed HORIZON AND LOCALITY: Lastro Oolitic structure of the caudal fin is not clearly dem- limestone, Grupo Ganhamaroba, Serie Ser- onstrated in any of the specimens, but it is gipe, Lower Cretaceous, Rio Ganhamaroba, evident that the homocercal condition is be- Sergipe. ing approached as in L. nevadensis (David, SPECIFIC DIAGNOSIS: Length to end of 1941). There are but five or six vertebrae caudal fin approximately 34 cm. Head about contributing to the structure of the tail. 12 cm. in length. Width of operculum about There is no evidence of the ray-like fulcural two-thirds of depth with tuberculated stria- scales along the dorsal border of the epaxial tions radiating from point of attachment. lobe of the caudal as in the genotype. Articular with longitudinal striations. Total The genus Tharrhias was established by number of vertebrae unknown, abdominal Jordan and Branner (1908) for a leptolepid about 18-20. Pectoral fin 14-15, pelvic fin from the Serie Araripe. It is distinguished about 10, anal approximately 25. Origin of from Leptolepis by having a relatively larger dorsal fin midway between origin of pectoral operculum, which is four times deeper than fin and origin of pelvic fin. Pectoral fin in- the suboperculum, a greater number of ver- serted on ventral border and extending be- tebrae, and by a larger body size. Haplo- yond origin of pelvic fin. Origin of pelvic fin spondylus, a Lower Cretaceous leptolepid midway between origins of pectoral and anal from Argentina (Cabrera, 1927), appears to resemble Tharrhias more closely in the oper- 1 From Lastro, the name of the limestone deposit in cular region, although Erasmo (1938) con- which the specimen was found. siders it to be possibly synomymous with 2 In allusion to elongated pectoral fin. 1947 SCHAEFFER: ACTINOPTERYGIAN FISHES FROM BRAZIL 15 fin. Pelvic rays extending to origin of anal Enchodontidae and to certain of the Iniomi, fin. Extended anal fin with long rays. but it appears to be based on such characters REMARKS: Apparently Lastroichthys (pl. as those listed in the preceding paragraph, 1, fig. 2) is the second Cretaceous vertebrate characters that are common to several sub- to be discovered in the State of Sergipe, the orders. other being Palaeobalistumflabellatum (Cope). Lastroichthys longipectoralis is thus assigned The locality from which the latter was col- tentatively to the Elopidae, until additional lected is unknown, but the horizon is prob- material makes a more detailed study pos- ably Middle Albian (Maury, 1936) and possible. sibly the same as that for Lastroichthys. With diagnostic portions of the skeleton FAMILY CHIROCENTRIEDAE missing, it is difficult to determine the affini- GENUS CHIROMYSTUS COPE, 1885 ties of this new genus. The width of the area Chiromystus COPE, 1885, p. 4. between the orbital region and the anterior of GENOTYPE: Chiromystus mawsoni Cope. border the preopercular indicates that the REVISED GENERIC DIAGNOSIS: Trunk elon- suborbitals were large as in the Elopidae. gate as in other chirocentrids. Mouth large, The wide gape, probably small teeth, ossified oblique. Maxillary narrow, extending beyond sclerotic ring, and striated opercular further posterior border of eye. Small styliform teeth suggest association with the Elopidae, par- in shallow pockets on maxillary, premaxil- ticularly with such forms as Thrissopater or lary, and dentary; those at mandibular sym- Pachyrhizodus from the Upper Cretaceous. physis enlarged. Pectoral fin large, anterior The elongated fins and rather delicate skele- rays broad and flat, pelvic and dorsal fins ton with thin dermal skull bones further sug- anal fin extended and gest a tendency towards a deep-sea habitus, small, lacking anterior a trend likewise found to some extent in the acuminate lobe. Caudal fin deeply forked, elopids. lobes narrow, pointed at ends. Vertebrae with The vertebrae are not characteristically three prominent lateral laminae. Ribs mod- elopid, having constricted centra externally erately robust, with longitudinal groove. and a single lateral lamina. The typical elopid Scales cycloid, delicate. vertebrae do not have centra constricted Chiromystus mawsoni Cope, 1885 externally and usually have two or three Chiromystus mawsoni COPE, 1885, p. 4. lateral fossae with intervening bars. The Chiromystus mawsoni, WOODWARD, 1888, p. 134. condition found in Lastroichthys resembles Chiromystus mawsoni, WOODWARD, 1901, p. 90. very closely that found in the Clupeidae, and Chiromystus mawsoni, Roxo, 1936, p. 67. on the basis of the vertebrae alone this genus TYPE: Status unknown; should be in the might very reasonably be assigned to that American Museum of Natural History but family. The Cretaceous elopid Pachyrhizodus cannot be located. does have somewhat similar vertebrae, al- HORIZON AND LoCALITY: Serie Bafa, Upper though the single lamina is much thicker. Cretaceous, eastern shore of Baia de Todos Relationship with the Characinidae is os Santos, near Agua Comprida, Baia. possible in view of the evidence here pre- DIAGNOSIS: (Based on Cope and Wood- sented favoring a Cretaceous marine habitus ward.) Length about 15-40 cm. Vertebrae for this family. The enlarged suborbitals, 50: abdominal 28, caudal 22. Anal fin 16. elongated pectoral, extended anal, character Small dorsal opposing anal, origin of anal of the vertebrae, and finally the wide gape are equidistant between pelvic and caudal. all present in this family. As these are not in themselves distinctive characin characters, Chiromystus alagoensis Jordan, 1910 it is not deemed advisable to refer Lastroich- Chiromystus alagoensis JORDAN, 1910, p. 32, thys to this family. A more detailed knowl- pls. 12, 13. edge of the skull must be available before TYPE: C. M. No. 5246/100. Almost com- assignment to the Characinidae could be plete skull and portion of pectoral fin. considered. HORIZON AND LOCALITY: S6rie Alagoas, There is likewise some resemblance to the ?Lower Eocene, near Riacho Doce, Alagoas. 16 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 89 REVISED SPECIFIC DIAGNOSIS: Length site the pelvic. Were it not for this forward ranging from about 20 to 35 cm. Skull about location of the dorsal, Woodward (1901, p. one-fifth total body length. Vertebrae about 108) would refer this genus, which may be 48: abdominal 21-23, caudal 25. Dorsal fin world-wide, to Ichthyodectes. 16, pectoral fin 12, pelvic fin about 6, anal REFERRED SPECIMENS: A.M.N.H. Nos. fin 21. Origin of dorsal fin opposite first third 10015, 10016. of anal fin, origin of pelvic midway between front of head and origin of caudal fin. Anal FAMILY CHANIDAE fin about equal to skull length. GENUS DASTILBE JORDAN, 1910 REMARKS: Jordan (1910, p. 33) considered Dastilbe JORDAN, 1910, p. 29. C. alagoensis (pl. 3, fig. 1) to differ from C. mawsoni (never figured) on the basis of a GENOTYPE: Dastilbe crandalli Jordan. supposed difference in the relative size of the REVISED GENERIC DIAGNOSIS: Moderately pelvic fin. In both species, however, this fin elongate fishes with somewhat rounded ven- is "small," and no specific differences are in- tral border. Dorsal and ventral scutes absent. dicated. There appears to be a smaller number Mouth small, oblique and jaws subequal. of vertebrae in C. alagoensis and a larger Opercular large, finely striated. Subopercular number of anal rays. The known material distinct, preopercular not expanded. Verte- for each species does not, however, permit an bral centrum with small notochordal canal, accurate count of the vertebrae. The size hourglass-shaped. Caudal portion of verte- range is about the same as are also the observ- bral column of stegurous type with last able details of the skull. Were it not for the vertebra elongated into a urostyle. Ribs short difference in the age of the two species, it and with wide shallow groove on outer sur- would be very reasonable to consider them face. synonymous, and this may prove desirable, particularly if the S6rie Alagoas proves to be Dastilbe crandalli Jordan, 1910 Upper Cretaceous rather than Lower Ter- Dastilbecrandalli JORDAN, 1910, p. 30, pl. 10, fig. tiary. 9. The skull fragment, presumably C. maw- TYPE: C. M. No. 5247/91. Complete fish soni, figured by Allport (1860) appears to with head imperfectly preserved. agree very closely with the type of C. alagoen- HORIZON AND LOCALITY: S6rie Alagoas, sis. Further preparation of the latter speci- ?Lower Eocene, Riacho Doce, Alagoas. men (pl. 3, fig. 2) has revealed certain details REVISED SPECIFIC DIAGNOSIS: Length of the skull not observed by Jordan, but they ranging from 3.50 to 9.40 cm., greatest depth are unfortunately of little diagnostic value. from 0.40 to 1.60 cm. Head one-quarter to The lacrimal is relatively large and the maxil- one-fifth total length of body. Opercular lary is relatively narrow as in Chirocentrus and about two-fifths of head length. Vertebrae in contrast to the condition found in Portheus. about 30: abdominal 17, caudal 13. Pelvic 8, The supramaxillary and the circumorbital anal 12, dorsal 12. Origin of pelvic fin behind plates are relatively large, agreeing in this origin of dorsal. respect with Ichthyodectes and Portheus. The REMARKS: Jordan (1910) considered Das- supraoccipital crest was apparently elevated tilbe (pl. 4, fig. 1) to be most nearly related as in the last-named genera. to the clupeid Halecopsis from the Lower The characters shared in common with Eocene of Europe. There are obvious differ- Platinx from the Eocene of Europe include ences, however, such as the expanded pre- the small teeth set in close series, tendency opercular and relatively smaller opercular towards enlargement of the pectoral fin, and in the latter. The resemblances between these lack of acuminate lobe on the anterior por- genera would appear to be due more to the tion of the anal fin. It is very possible that fact that they are both relatively primitive Chiromystus is related to this genus. generalized Isospondyli than that they are Cladocyclus gardneri Agassiz, a chirocen- actually near relatives. trid from the S6rie Araripe, differs from Chiro- Arambourg (1935b) transferred Dastilbe mystus in the position of the dorsal fin oppo- to the Chanidae, a family considered to be in 1947 SCHAEFFER: ACTINOPTERYGIAN FISHES FROM BRAZIL 17 some respects more primitive than the Clu- the Clupeoidea and several of the derived peidae and more closely related to the Al- suborders such as the Salmonoidea. bulidae. It includes the genus Chanos, which REFERRED SPECIMENS: A.M.N.H. Nos. occurs in the Eocene and Miocene of Europe, 10018-10045. and the Recent "milk fishes." Arambourg emphasizes the persistent notochordal canal FAMILY CLUPEIDAE and the stegurous type of terminalization (pl. GENUS KNIGHTIA JORDAN, 1907 4, fig. 2) for the vertebral column as two im- Diplomystus COPE, 1877, P. 811 (in part). portant retained primitive characters. These, Knightia JORDAN, 1907, p. 136. he believes, together with the observable Ellipes JORDAN, 1910, p. 24 (Brazilian species skull characters and scale morphology, indi- only). cate that Dastilbe should be placed in the Ellimma JORDAN, 1913, p. 79 (new name for Chanidae. EUipes Jordan, 1910, preoccupied). This author has further pointed out that a Ellimma, JORDAN AND GILBERT, 1919, p. 26. series of leptolepids such as the Lower Ju- GENOTYPE: Knightia eocaena Jordan. rassic L. coryphaenoides, Upper Jurassic L. REVISED GENERIC DIAGNOSIS: Double- sprattiformes, Upper Jurassic L. brodiei, and armored clupeids of varying body depth. the Lower Cretaceous L. formosus, together Cleft of mouth moderately oblique. Teeth with Dastilbe and the related Lower Creta- very small or absent. Vertebrae 31-36, cau- ceous Parachanos, represent a structural series dals with double lateral laminae. Origin of demonstrating the morphological transition, pelvic behind origin of dorsal. Anal fin small, particularly in the structure of the caudal fin, 12-14 rays. Dorsal scutes ovate with median from the paleoniscid stage to a stage repre- carina and with or without extended poste- sented by the clupeids. In such a series, it is rior median spine. Ventral scutes of typical possible to observe a reduction in the size of clupeoid type. Scales large, thin, smooth. the notochordal canal, a change from the REMARKS: The inclusion of Ellimma bran- stegurous to homocercal type of caudal fin, nri Jordan within the genus Knightia has and a gradual loss of ganoine on the scales. necessitated a revision of the generic diag- While the presence of a persistent noto- nosis of Knightia, particularly in regard to chordal canal in the leptolepids and certain the nature of the dorsal scutes. As the latter of the Chanidae may be considered as a genus is here understood, it includes the fol- truly primitive character, a hold-over from lowing species: eocaena, alta, copei, branneri, the holostean stage, it would appear that this brasiliensis, and, tentatively, elmodenae and character is of questionable phylogenetic barbarae. The last two species, from the Mio- value in more specialized groups in which it cene of California (Jordan and Gilbert, 1919), appears. Its occurrence in characins such as have not been available for study. The valid- the Pliocene Eobrycon and the Recent Brycon ity of copei (Tanner, 1925), described from a and even in certain carangids suggests an in- single specimen, is open to question as it may dependent retention of an embryonic condi- very well fit into the range of alta. tion. The presence of a notochordal canal in adult actinopterygians must be considered ghta branneri (Jordan), 1910 as a case of neoteny, and it may or may not Ellipes branneri JORDAN, 1910, p. 25, p1. 8, fig. 3. be a primitive character, depending on the Ellipes riacensis JORDAN, 1910, p. 28, p1. 10. phylogenetic position of the group under con- EUimma branneri, JORDAN, 1913, p. 79. sideration. TYPE: C. M. No. 5249/1. Complete speci- In adult actinopterygians, the stegurous men excepting imperfectly preserved fins. condition of the caudal fin likewise represents HORIZON AND LOCALITY: Serie Alagoas, the retention of an embryonic character that ?Upper Eocene, Riacho Doce, Alagoas. is very pronounced in the larval stages. Un- REVISED SPECIFIC DIAGNOSIS: Standard like the character discussed above, however, length ranging from 2.20 to 8.70 cm., depth there is little reason for believing that neot- from 1.05 to 4.00 cm. Head about one- eny has caused the retention of the stegurous quarter total body length. Opercular with caudal fin in any of the teleost groups beyond striations radiating ventrally from point of 18 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 89

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40en't- q1 qAp l 20 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 89 attachment. Vertebrae 31-33: abdominal 15- REFERRED SPECIMENS: A.M.N.H. Nos. 16, caudal 17. Dorsal fin 12, pectoral fin 8, 10046-10062. anal fin 12. Origin of pelvic fin behind origin of dorsal fin and midway between origin of GENUS DIPLOMYSTUS COPE, 1877 pectoral and origin of anal fins. Dorsal scutes Diplomystus COPE, 1877, p. 811. widely ovate, lacking posterior spine. REMARKS: This species (pl. 5, figs. 1, 2) GENOTYPE: Diplomystus dentatus Cope, was placed in a separate genus by Jordan be- 1877. (For generic diagnosis, see Thorpe, cause he considered its squamation to differ 1938, p. 280.) from that found in either Diplomystus or Diplomystus longicostatus Cope, 1885 Knightia. He did state, however, that it pos- Diplomystus longicostatus COPE, 1885, p. 3. sesses characters found in both these genera. Diplomystus longicostatus, WOODWARD, 1888, p. The scales are well preserved in a number of 134. specimens, and they appear to agree very Diplomystus longicostatus, WOODWARD, 1895, p. closely in size and shape with those of K. 2, pl. 1. eocaena and alta, being much larger than the Diplomystus longicostatus, WOODWARD, 1901, P. scales of Diplomystus. The dorsal scutes 143. (fig. 5) lack the lateral wings typical of Diplo- EUlipes longicostatus, JORDAN, 1910, p. 29, p1. 11. mystus and agree in their ovate configuration EUlimmichthys longicostatus, JORDAN, 1919, P. with those of Knightia, with the exception 27. that the posterior border is not produced into TYPE: Status unknown; should be in Amer- a long spine. ican Museum of Natural History but cannot The general body form is very similar to be located. that found in K. alta, although the dorsal HORIZON AND LOCALITY: S6rie Bafa, Upper border, in mature individuals of K. branneri, Cretaceous, between Plataforma and Ita- tends to rise to a slight degree to an apex at carAnha, along eastern shore of Bafa de Todos the point of origin of the dorsal fin. A similar os Santos, Bafa. dorsal outline is characteristic of Diplomystus REVISED DIAGNOSIS: Standard length 8.50 longicostatus and D. goodi, and affinity with to 10.30 cm., greatest depth from 5.80 to these species might be considered were it not 6.15 cm. Head about one-third total body for the differences in the shape of the dorsal length. Dorsal outline rising sharply to apex scutes and in the squamation. at origin of dorsal fin with equally sharp de- Jordan (1910) recognized a second species scent posteriorly. Body very deep with ventral of the genus Ellipes from Riacho Doce, which border markedly concave. Vertebrae about he named E. riacensis. He considered it to 36: abdominal 24, caudal 10-12. Dorsal fin differ from branneri in having a relatively 10, pectoral fin 9, pelvic fin 6-7, anal fin 8. longer head, shorter mandible and ribs, more Origin of pelvic fin opposite origin of dorsal, elongate body, and broader opercular. A origin of anal midway between origin of dor- restudy of Jordan's specimens does not reveal sal and origin of caudal fins. Pelvic fin smaller any consistent differences between the two than pectoral. Dorsal scutes with lateral species. The skull is not relatively larger in wings, posterior border not denticulated. individuals of the same size, and the other Ventral scutes enlarged behind pelvic fin. proportions appear to be identical in speci- REMARKS: Although D. longicostatus (fig. 5) mens formerly assigned to each species. E. is not included in the collection under con- riacensis is therefore considered to be a syno- sideration, it is represented by two specimens nym of K. bronneri. on a single slab of unknown origin from the Woodward (1939) has described Knightia Newberry Collection, A.M.N.H. No. 734. brasiliensis from Tertiary deposits at Nova A reconsideration of its systematic position York, Maranhao. It resembles K. alta with is desirable as Cope (1895) considered it to a flattened dorsal border and rather deep be allied to Knightia (D. humilis), presum- body. The dorsal scutes are not described ably because the dorsal scutes are cordate. excepting that they lack lateral wings. The dorsal scutes of one specimen show lateral PLATES 1 - 7 BULLETIN AMER. Mus. NAT. HIST. VOL. 89, PLATE 1

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1. Leptolepis bakiaensis, new species. A.M.N.H. No. 10014. Type specimen. X2/1 2. Leptolepis bahiaensis, new species. Outline drawing of skull as preserved in type. Abbretiations: Brstg, branchiostegals; Clt, cleithrum; Dn, dentary; Ecpt, ectopterygoid; Enpt, entopterygoid; Fr, frontal; Hy, hyoid arch; Mpt, metapterygoid; Mx, maxillary; Op, opercular; Pas, parasphenoid; Pop, preopercular; Qu, quadrate; Sop, subopercular. X6 BULLETIN AMER. Mus. NAT. HIST. VVOL. 89, PLATF, 3

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1. Chiromystus alagoensis Jordan, A.M.N.H. No. 10015. X2/3 2. Chiromystus alagoensis Jordan, type. C.M. No. 5246/100. Skull and pectoral fin. Abbreviations: Dn, dentary; Mx, maxillary; Pas, parasphenoid; P1, palatine; Qu, quadrate. X1/1 BULLETIN AMER. Mus. NAT. HIST. VOL. 89, PLATE 4

Na,41 *.5n,7nI- A \ 1. Dastilbe crandalli Jordan. A.M.N.H. No. 10034. Positions of the fins well indicated. The skull is poorly preserved in all available specimens. X3/2 2. Dastilbe crandalli Jordan. A.M.N.H. No. 10028. The stegurous terminalization of the vertebral column is clearly demonstrated in this specimen. X4/3 3. Aequidens pauloensis, new species. A.M.N.H. No. 10095. Type specimen. The dorsal fin is missing and the anal fin not completely preserved. X3/4 BULLETIN AMER. Mus. NAT. HisT. VOL. 89, PLATE 5

1. Knightia branneri (Jordan). C.M. No. 5248/4. This specimen is the type Ellipes riacensis Jordan. X4/5 of 2. Knightia branneri (Jordan). A.M.N.H. No. 10057. There is no indication in this specimen of the elevation of the dorsal border present in figure 1. This character is highly variable within the species and may be partly a result of poor x 1/1 preservation, BULLETIN AMER. Mus. NAT. HIST. VOL. 89, PLATE 6

4' ' '> £XX93{iw 1. Eobrycon avus (WVoodward). A.M.N.H. No. 10070. Pelvic fin restored. x4/5 2. Eobrycon branneri Eigenmann. A.M.N.H. No. 10089. Type specimen. Fpf, fontanelle. X4/5 fronto-parietal 3. Eobrycon ligniticus (Woodward). A.M.N.H. No. 10067. X3/2 BULLETIN AMER. Mus. NAT. HIST. VOL. 89p PLATE 7

1. Percichthys antiquus Woodward. A.M.N.H. No. 10091. Portion of the preopercular is preserved showing the serrated posterior border. X 1/2 2. Percichthys antiquus Woodward. A.M.N.H. No. 10092. Impression of the ventral surface of the skull. Abbreviations: Op, opercular; Pal, palatine; Pas, parasphenoid; Pmx, premaxillary; Vo, vomer. X3 1947 SCHAEFFER: ACTINOPTERYGIAN FISHES FROM BRAZIL 21 wings, a fact pointed out by Woodward study of the fish fauna of the Green River (1901), indicating that this species, now con- shales. sidered to be Upper Cretaceous rather than In figure 3 the standard length is plotted Lower, should be included in Diplomystus. on the horizontal axis against maximum body Jordan (1910) included D. longicostatus in depth on the vertical axis. In figure 4 the Ellipes but failed to state the reason and horizontal axis represents the length of the later (1919) placed it in a separate genus, base of the anal fin and the vertical the length Ellimmichthys, principally because of the high of the skull from the anterior border of the elevation of the dorsal border. As the latter premaxillary to the caudal border of the oper- is a character apparently occurring in both culum along a line represented by the para- Knightia and Diplomystus, it cannot be con- sphenoid. The species considered are Diplo- sidered of generic value. mystus dentatus, analis, and pectorosus from The resemblance between D. longicostatus the Green River Middle Eocene, D. longi- and D. goodi (Eastman, 1912; Weiler, 1923a, costatus from the Upper Cretaceous of Brazil, 1923b) from the Tertiary of West Africa is D. brevissimus from the Upper Cretaceous of very close, and they can be distinguished Syria, D. goodi from the early Tertiary of mainly by a probable significant difference in West Africa, Knightia eocaena and alta from the shape of the dorsal scutes (difficult to de- the Green River, and K. branneri from the termine in the case of D. goodi) and a differ- possible early Tertiary of Brazil. ent vertebral count. It would appear from inspection of the regressions that D. dentatus is quite distinct PRELIMINARY ANALYSIS OF THE from the other species considered. The rather DOUBLE-ARMORED CLUPEIDS large collection of diplomystids from the Cope long ago recognized the need for Green River shales contains no individuals separating Diplomystus into two sections, that are intermediate in size between D. mainly because of a difference in the shape of dentatus and D. analis, a fact which is rather the dorsal scutes. Jordan (1907), in agree- difficult to explain. The positively identified ment, elevated one of these sections to the specimens of D. dentatus are all character- rank of the separate genus Knightia. In order istically large, with apparently no younger to determine further the validity of assigning stages represented. Although there very well the species formerly grouped under Ellimma may be some young of this species included to Knightia, Ellimmichthys longicostatus back in the D. analis scatter, these cannot be sepa- again to Diplomystus, and also to investigate rated with any degree of certainty, particu- the relationships between the various species larly since the vertebral counts of the two of the two genera here recognized, four char- species overlap. There is some evidence, how- acters in nine species have been studied quan- ever, of differences in the details of the dorsal titatively. scutes, and it may be possible to separate As in the case of the scatter diagrams for further specimens of the same size by count- Eobrycon to be discussed later, the regression ing the annular rings of the scales. for each species of Diplomystus and Knightia The separation between the regressions of represents essentially an ontogenetic series D. dentatus and analis may be narrowed by rather than the range of variation within a the study of a larger series, although a survey population sample of adult individuals. An of the literature indicates that this is not overlap between separate regressions does likely. Several possible explanations might indicate similarity in body form, although be offered, for instance, a restriction of the the trends in some cases may be in somewhat young of D. dentatus to a particular type of different directions. This analysis is more or environment such as the running water of less tentative and is based only on the ma- streams or shallow water near the edge of the terial available in the American Museum and Green River lakes. Individuals segregated some from the Carnegie Museum. The use of under such conditions either might not have much larger samples and additional char- been preserved or have not thus far been col- acters is contemplated in connection with a lected. 22 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 89

FIG. 5. Tentative phylogeny of the double-armored herrings. Not all of the species represented in the distribution map are included in the phylogeny. The dorsal scutes, where known, are outlined. On the basis of the graphs (figs. 3 and 4) and the shape of the dorsal scutes, it is possible to recognize three phylogenetic trends. The first includes D. brevissimus, D. longicostatus, and possibly D. goodi; the second includes the Knightia group; and the third includes the Green River diplomystids. D. birdi may possibly be ancestral to the Green River forms through the D. brevissimus stock. The dorsal scutes of D. vectensis and D. elmodenae have not been described, and the position of these species is questionable. 1947 SCHAEFFER: ACTINOPTERYGIAN FISHES FROM BRAZIL 23 The regressions of the other species of both A tentative phylogeny of the double- Diplomystus and Knightia are clustered to- armored clupeids (fig. 5), based on the pres- gether for both sets of characters studied. In ent study, indicates the changes in body figure 3 the regressions for Knightia alta and form that have occurred since the Lower eocaena run imperceptibly into one another, Cretaceous. The first true generalized clu- bearing out Thorpe's statement (1938) that peids occur in the Lower Cretaceous, and it in a large series it is very difficult to separate would appear that the rough-backed herrings these species. Their status will be further were differentiated from the primitive clu- considered in the Green River fauna study. peoid stock at about that time. The extreme The more deep-bodied individuals, identified depth of the body in D. longicostatus further as alta, overlap the regressions of Diplo- suggests that a tendency in the deep-bodied mystus analis, pectorosus, and brevissimus. direction appeared about as early as the This situation is not surprising since the body transformation of the middorsal scales into form in the two genera is almost identical, scutes. (Woodward, 1942, p. 908, considers particularly in species with short anal fins. the dorsal and ventral ridge scutes to be a In the Green River diplomystids only, the direct "ganoid" inheritance.) The Creta- anal fin is relatively long and the ventral ceous diplomystids are all fairly deep bodied border in the caudal region is hence quite as are all but one species of Knightia. The straight, resulting in a rather distinctive body tendency towards a fusiform body outline outline. D. longicostatus and D. goodi represent seems to represent more of an end develop- the extreme in body depth as their regres- ment, appearing in the Eocene K. eocaena, sions demonstrate. the Oliogocene D. vectensis, and the Recent In figure 4, in which the length of the base Hyperlophus, Potamolosa, and Ethmidium. of the anal fin is plotted against skull length, Diplomystus brevissimus could very well the relatively long anal fin and shorter skull represent the generalized type from which distinctly separate the Green River diplo- Knightia, the other species of Diplomystus, mystids from the other species of this genus and the Recent double-armored herrings are and also from Knightia. The regressions for derived. The body is moderately deep, the Knightia branneri and K. eocaena plus alta dorsal scutes have poorly developed lateral are completely superimposed, further sup- wings, and the anal fin is of moderate length. porting the assignment of branneri to this It is likely, however, that the two types of genus. The shorter anal fin and longer skull dorsal scutes developed in the Lower Cre- combination has also developed within the taceous, although Knightia is unknown be- diplomystids, accounting for the proximity of fore the Lower Tertiary or Upper Cretaceous D. longicostatus and D. goodi with Knightia. of Brazil (Serie Alagoas). The difference in A feature of the body form that is difficult scale size, mentioned above, also must have to present quantitatively is the elevation of developed at this time. the dorsal border, a character that has ap- The dorsal scutes of the Oligocene and parently developed in both genera. This ele- Miocene diplomystids have not been ade- vation is associated with the relative increase quately described. Woodward (1901) states in skull length and decrease in anal fin length. that those of D. vectensis, from the Oligocene Diplomystus birdi from the Upper Creta- of the Isle of Wight, are "narrow," while the ceous of Syria, with its hypsisomatic body dorsal scutes of K. elmodenae (Jordan, 1919) form, may represent a side-line developed are "simple." It is very possible that D. vec- from the early diplomystid stock. Its extreme tensis should be transferred to Knightia. depth is analogous to that found in such a Smirnov (1935, 1936) has described Diplo- form as Zanclus, and as is characteristic of mystus brevicaudus and D. alter from the such types, the dorsal and anal fins are oppo- north Caucasus, but unfortunately the pub- site one another. The dorsal scutes are strong- lished descriptions have not been available ly denticulated and rather resemble those of for comparison with the other members of D. dentatus. D. brevissimus from the same this genus. It is becoming evident, however, locality, on the other hand, is far more gen- that the double-armored herrings had a world- eralized with dorsal scutes that have a smooth wide distribution probably from the Creta- posterior border. ceous through the greater part of the Tertiary. 24 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 89 During this long period they inhabited fresh, D. longicostatus and D. brevissimus, for in- brackish, and salt water and consequently stance, have dorsal scutes with a smooth pos- cannot be considered as being typically asso- terior border and wide, rounded, lateral ciated with any of these habitats. wings. Although the term cordate has been Three Recent genera are assigned to this employed to describe the dorsal scutes of group. Hyperlophus is known from the coastal Knightia, it more nearly fits the shape of waters of Australia and is probably most these plates in most species of Diplomystus. nearly related to Knightia. The dorsal scutes Those of the Green River species and of D. of several specimens of Hyperlophus spratel- birdi with narrow lateral wings and a denticu- lides in the American Museum collection are lated posterior margin can be considered only ovate with a spinous carina projecting some- as modifications of a basic cordate type. A what beyond the posterior border. The scutes possibly more appropriate term for the shape resemble those of K. eocaena in general out- of the dorsal scutes in Knightia is ovate, as line, and the shallow, fusiform body is very they lack the posterior notch which in effect similar to that found in this species. The scales produces the lateral wings. As is evident, are likewise large and thin. such finely drawn distinctions become almost Potamalosa, presumably closely related to meaningless when the less specialized diplo- Hyperlophus, occurs in fresh water and is mystids are compared with Knightia, and apparently restricted to the coastal rivers of there is no doubt that the differences be- New South Wales. Ethmidium inhabits the tween Diplomystus and Knightia are of much coastal waters of Peru and Chile and is very less significance than they were considered similar in body form to the genera from to be by Jordan. A comparison of these gen- Australia. These Recent forms can be con- era from the Green River shales only by no sidered only as isolated remnants of a group means presents the true picture. that was undoubtedly universally distributed until the later Tertiary. ORDER OSTARIOPHYSI In many respects it appears desirable to establish a separate family for the double- SUBORDER CYPRINOIDEA armored herrings. The presence of the dorsal FAMILY CHARACINIDAE scutes certainly sets this group apart from GENUS EOBRYCON JORDAN, 1907 the other clupeids, which otherwise may be Tetragonopterus Cuvier, 1817, (in part) WOOD- divided into two sections according to the WARD, 1898, p. 66. presence or absence of the ventral scutes. Eobrycon JORDAN, 1907, p. 140. The taxonomy of the Diplomystus-Knightia GENOTYPE: Tetragonopterus avus Wood- complex is, however, still unsatisfactory. In ward. many cases, the specific differences disappear REVISED GENERIC DIAGNOSIS: Fusiform, when large series of specimens are available, moderately for instance between K. eocaena and K. alta. deep-bodied fishes with rounded Even the distinction between these two gen- ventral border as in recent Brycon. Length of era is not sharply defined because of inter- anal fin base about three-fourths of skull grading forms such as K. branneri. It is length. Fronto-parietal fontanelle present. hoped that the large series of specimens of Premaxillary teeth with three cusps arranged all species to be employed in the study of the in linear order as in Brycon. Vertebral cen- Green River fauna will aid in resolving, trum with single lateral ridge. Notochordal through a further quantitative analysis, this canal relatively larger than in Brycon. Ribs rather confusing picture. robust, not quite reaching ventral border, The presence of a denticulated posterior with single longitudinal groove. Anal fin border on the dorsal scutes of the Green River extended. Fork of caudal fin shallow. Dorsal diplomystids has been generally considered fin short based. Pectoral rays extend beyond as a generic character of some importance origin of pelvic. (Thorpe, 1938). It is evident, however, that Eobrycon avus (Woodward), 1898 by no means all of the species assigned to Tetragonopterus avus WOODWARD, 1898, p. 66, Diplomystus possess this character. Both fig. 3. 1947 SCHAEFFER: ACTINOPTERYGIAN FISHES FROM BRAZIL 25 Tetragonopterus avus, WOODWARD, 1901, p. 298, exposed at Taubat6 and Trememb6, Sao pl. 17, fig. 1. Paulo. Lacustrine-fluvial. Eobrycon avus, JORDAN, 1907, p. 140, fig. 31. REVISED SPECIFIC DIAGNOSIS: Length of Eobrycon avus, EIGENMANN AND MYERS, 1929, skull plus vertebral column ranging from p. 513. 4.45 to 8.60 cm., maximum depth from 1.07 TYPE: B.M.N.H. No. P. 9222. Complete to 2.20 cm. Body similar to E. avus and E. fish with imperfectly preserved skull. branneri. Opercular not striated. Orbit rela- HORIZON AND LOCALITY: ?Pliocene beds tively large, about one-third of skull length. exposed at Taubat6 and Trememb6, Sao Teeth as in preceding species. Vertebrae 38: Paulo. abdominal 18, caudal 20. Dorsal 8, pectoral REViSED SPECIFIC DIAGNOSIS: Standard 11, pelvic about 7, anal about 30. Origin of length ranging from 8.60 to 17.60 cm., great- dorsal above origin of anal. est depth from 1.85 to 4.05 cm. Skull about REFERRED SPECIMENS: A.M.N.H. Nos. one-fourth total body length. Vertebrae 38- 10067-10069. 42: abdominal about 20, caudal about 22. REMARKS: The taxonomic history of E. Dorsal fin 10-11, pectoral fin 9-11, pelvic ligniticus represents an interesting compari- fin 8-9, anal fin 22-23. Origin of dorsal fin son between the approach of paleoichthyol- midway between origin of pectoral and origin ogists and neoichthyologists to the problem of anal fins. of the relative value or importance of single REFERRED SPECIMENS: A.M.N.H. Nos. characters in a fossil teleost. Woodward, 10063-10066, 10070-10088. representing the former group, placed E. avus and E. ligniticus in the same genus, con- Eobrycon branneri Eigenmann, 1929 sidering the differences to be of specific value Eobrycon branneri EIGENMANN, 1929, p. 514, pl. only. Eigenmann and Myers, representing 81. the latter, considered E. ligniticus to be worthy of separate generic rank on the basis TYPE: A.M.N.H. No. 10089. Complete fish of the position of the dorsal fin, but admitted with well-preserved fins. Type and paratype that in other respects it resembles Eobrycon. (A.M.N.H. No. 10090) donated to the Ameri- Discussion with a neoichthyologist confirms can Museum of Natural History by Indiana the attitude of the latter authorities, namely, University. that the position of the dorsal may be a char- HORIZON AND LOCALITY: ?Pliocene beds acter of generic significance in Recent fishes exposed at Taubate, Sao Paulo. and that a difference in position, separating REVISED SPECIFIC DIAGNOSIS: Length of two genera, is usually accompanied by other skull plus vertebral column in type and para- diagnostic differentiation, for instance in type (only known specimens) 13.2 and ± 14.7 squamation, which is usually observable only cm., greatest depth 4.00 and 5.15 cm. Skull in the living fish. Hence the two groups of more than one-fourth total length of body. students, using a somewhat different ap- Vertebrae about 38: abdominal about 20, proach, are prone to place different values on caudal 18. Ray counts about as in E. avus. the available differentiating characters in the Origin of dorsal midway between origin of fossil forms. Where accurate, conclusive, pelvic and origin of anal. quantative work is not possible because of the nature of the material, a more conserva- Eobrycon ligniticus (Woodward), 1898 tive approach would appear desirable, even Tetragonopterus ligniticus WOODWARD, 1898, p. though such "lumping" of species may still be 67, figs. 4, 5. quite subjective. Tetragonopterus ligniticus, WOODWARD, 1901, P. In the genus Eobrycon,l as here recognized, 298, pl. 17, figs. 2, 3. the three species avus, branneri, and ligniticus Lignobrycon ligniticus, EIGENMANN AND MYERS, I Santos has recently described Lignobrycon altus 1929, p. 513. (1946, Notas Prelim. Estudos, Div. Geol. Min., Rio de Janeiro, no. 36, pp. 1-3, pls. 1-2) from the Lower TYPE: B.M.N.H. No. P. 9012. Complete Tertiary locality at Nova York, Maranhao. He consid- fish. ers it to differ from L. ligniticus in having a greater body HORIZON AND LOCALITY: ?Pliocene beds depth and a smaller number of rays in the anal fin (26). 26 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 89 6 IC Key * Eavus 05 * E.branneri

-j4 . A E.I igniticus U. qa U ..i f U Un A i.I.b U a a c 1 A A A-

6T 54+ I. X 4 am. *. .J 3.- Ua U a A .= 2 - a V. A B 0

t 04 a 0 lb on so N % E 3 w1 U a A U E 2'.1- A . xI t. I. C

v 3 4 5 6 7 8 9 10 11 12 13 14 15 Standard Length FIG. 6. A. Graph of anal fin base length and standard length in centimeters for the three species of Eobrycon. B. Graph of skull length and standard length in centimeters. C. Graph of maximum body depth and standard length in centimeters. 1947 SCHAEFFER: ACTINOPTERYGIAN FISHES FROM BRAZIL 27 (p1. 6), in the order named, represent a series been possible to confirm an indication of the demonstrating three different positions of adipose fin in the type of Eobrycon branneri the dorsal fin from one essentially above the as reported by Eigenmann and Myers. pelvic fin to one above the anterior portion of Eobrycon, being so nearly related to a well- the anal fin. The location of the dorsal fin in defined Recent genus, offers no information ligniticus, therefore, would appear to lose its as to the origin of ancestry of the Characini- possible generic value. The orbit in E. ligniti- dae. The two characins described by Piton cus is apparently not relatively larger than in (1938) from the Middle Eocene lacustrine- E. avus. An analysis of skull length compared fluvial beds at Menat, France, Prohydrocyon with orbit length indicates that the orbit in- pellegrini and Procharacinus arverniensis, creases in size at a greater constant rate than throw no light on the problem. The former the skull through E. ligniticus and E. avus. has a small mouth, bordered above mostly A graphic comparison of the variation in by the premaxillary. The teeth are styliform total skull length, length of the base of the and are arranged in two rows on the upper caudal fin and maximum body depth (vertical jaw and in a single row on the lower jaw. The axis) with the standard length (horizontal general pattern and relative size of the cheek- axis) is made in figure 6. As previously bones do not indicate a close relationship pointed out, the distribution within each with the recent Hydrocyon. Procharacinus species represents essentially an ontogenetic likewise has a small mouth, bordered above series, and it cannot be considered as indi- entirely by the premaxillary. The arrange- cating the range of variation in a population ment of the cheek bones appears very similar of adult individuals. It is evident, however, to that found in Prohydrocyon, but the num- that for the characters studied E. ligniticus ber of teeth is much smaller than in this grades into or overlaps E. avus, while E. genus. branneri overlaps E. avus at the other end of In our present state of knowledge regarding its range. This analysis is simply another the phylogenv of the characins, it is difficult indication of the close affinity of the three to determine the relationships of these gen- species of Eobrycon. Eobrycon branneri may be era. That they are not close to Eobrycon is distinguished from E. avus by the position of quite certain. the dorsal fin and possibly a greater maxi- Cockerell (1919) and David (1946) have mum body length. The small number of described a characin, Erythrinolepis, from the known specimens of the former species makes marine Upper Cretaceous of California and it impossible to determine with any degree of Wyoming, and Cockerell (1921) has described accuracy the amount of intraspecific varia- another form, Characilepis, from beds at tion and the degree of overlap with E. avus. Huacho, Peru, which are probably marine E. ligniticus may be distinguished from the Cretaceous. Both genera are based on isolated other two by its smaller body size, lack of scales only. Erythrinolepis is probably related radiating striations on the opercular bone, to the subfamily Erythrininae, an American and, of course, the position of the dorsal fin. group including the Recent Erythrinus and The resemblance between Eobrycon and Lebiasina, but this relationship is not abso- Brycon is very marked and appears to indi- lutely certain. Characilepis may be included cate a close affinity, if not synonymy, as noted in the subfamily Cheirodontinae of Gregory by Eigenmann and Myers (1929, p. 514). and Conrad (1938) or Tetragonopterinae of Besides close resemblance in tooth form, Eigenmann (1929) as the scale pattern bears nature of the fronto-parietal fontanelle and a very close resemblance to that of the Recent body shape, there is a very similar cheek- Bryconamericus. The nature of the relation- bone pattern in the two genera. They appear ship is, however, again uncertain. to differ in that Eobrycon has a narrowing of Although the fossil characin evidence is the operculum dorsally, a wider horizontal anything but illuminating, there is good arm on the preopercular, and finally no indi- reason for believing that the Characinidae cation of the anterior and posterior plate-like were undergoing a radiation by Upper Cre- expansions of the interneurals, particularly taceous time and that the subfamilies were in those anterior to the dorsal fin. It has not established by the early Tertiary. If the scale 28 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 89 identifications can be considered as reliable, According to all recent taxonomic treat- there is further evidence that this character- ments of this genus, it should be included in istically fresh-water family had marine mem- the Serranidae rather than the Percidae. bers and possibly a marine origin in Creta- Woodward so placed it in his original descrip- ceous time, a problem that will be outlined fur- tion of P. antiquus (1898), although he in- ther in the discussion. cluded it in the Percidae later (1901), the family Serranidae not being recognized in ORDER ACANTHOPTERYGII the latter work. Percichthys bears a close re- SUBORDER PERCOIDEA semblance to the percids, and Jordan (1923) FAMILY SERRANIDAB was undoubtedly cognizant of this when he created a separate family, the Percichthyidae, GENUS PERCICHTHYS GIRARD, 1854 which he considered to be intermediate be- Percichthys GIRARD, 1854, p. 197. tween the Percidae and Moronidae or white GENOTYPE: Perca trucha Cuvier and Valen- basses. Von Ihering's contention (1898) that ciennes. (For generic diagnosis, see Bou- Percichthys is not a serranid but a cichlid is lenger, 1895, p. 118.) denied by Eigenmann (1909, p. 336). The cichlids have, among other differences, but Percichthys antiquus Woodward, 1898 one nostril on each side, while serranids, in- Percichthys antiquus WOODWARD, 1898, p. 68, cluding Percichthys, have two. figs. 6, 7. Although its resemblance to the percids Percichthys antiquus, WOODWARD, 1901, p. 514. may be due to its generalized percomorph TYPE: Senckenberg Museum, Frankfurt, nature, Percichthys does appear to bridge Germany. Number and status unknown. Im- the small morphological gap between the perfect fish lacking portion of dorsal and percids and serranids in most respects. A caudal. Figured by Woodward, 1898 (fig. 6). comparison of the Eocene Serranidae and HORIZON AND LOCALITY: ?Pliocene beds Percidae (Schaeffer, 1947) indicates very few, at Taubate and Trememb6, Sao Paulo. if any, distinguishing differences between REVISED DIAGNOSIS: Probable length rang- these families. The number of anal spines, ing from 11.20 to 20 cm. Greatest depth no which has been employed rather arbitrarily less than 2.5 cm. Skull somewhat more than to separate the early serranids and percids, one-third total length of body. Posterior is by itself a character of questionable value. border of preopercular finely serrated. Ver- The number of vertebrae in P. antiquus and tebrae 32-35: abdominal 14-15, caudal 18-20. the Recent species of this genus is within the Anterior dorsal fin X; posterior dorsal fin 12; Recent percid range of 30 to 48. P. hondoen- pectoral fin 12-13; pelvic fin I, 5; anal fin sis, however, has 24-26 vertebrae, the prin- III, 8-9. Origin of anal fin nearly opposite cipal observable difference between it and P. origin of posterior dorsal. Caudal fin truncate. antiquus, but in agreement with the vertebral Ctenoid scales with about seven ridges radiat- number for most Eocene perches and basses. ing apically. One character that favors placing Percichthys REMARKS: The Recent species of the genus in the Serranidae is the presence of a ser- Percichthys are discussed by Eigenmann ranid-like subocular shelf, observed in a (1909, p. 284; 1927, p. 61). P. antiquus (pl. specimen of P. trucha, but not recognizable 7) agrees very closely in observable char- in either fossil species. acters with P. trucha, a species that occurs in The assignment of Percichthys to the Ser- both Patagonia and Chile. The presence of ranidae is hence more or less arbitrary, at P. antiquus in the fresh-water Tertiary of least as far as the osteological characters are Brazil indicates that this genus must have concerned. This fact has been noted by sev- had a much greater distribution than it does eral ichthyologists including Hills (1934, p. at the present time. It is also evident that 170) in a discussion of the affinities of Per- the genus was restricted to fresh water by the calates. Although Percalates is assigned to the Eocene with the discovery of P. hondoensis Moronidae, Percalates antiquus from the in the Lower Eocene of Patagonia (Schaeffer, Paleogene of Australia appears to resemble 1947). Percichthys very closely. This can only be 1947 SCHAEFFER: ACTINOPTERYGIAN FISHES FROM BRAZIL 29 considered as further evidence of the merging in the single available specimen. A. pauloensis of the Percidae, Serranidae, Moronidae, and has 16 caudal vertebrae while the other spe- related groups in the early Tertiary. cies have but 13. There are approximately 29 REFERRED SPECIMENS: A.M.N.H. Nos. spines and rays in the dorsal fin according to 10091-10094. a count of the interneurals. This number falls FAMILY CICHLIDAE at the extreme upper range among the species of Aequidens and approaches the number in GENUS AEQUIDENS EIGENMANN AND BRAY, 1894 the related genus Astronotus. Until more and Acara HECKEL (in part), 1840, p. 338. better preserved material is available in Aequidens EIGENMANN AND BRAY, 1894, p. 616. which the vertebrae and the spines and rays GENOTYPE: Acara tetramerus Heckel. (For of the dorsal fin can be accurately counted it generic diagnosis based on osteological char- would appear advisable to assign pauloensis to acters, see Regan, 1905, p. 330.) Aequidens rather than to create a new genus. An indeterminable species of Acara has Aequidens pauloensis,1 new species been described by Woodward (1898) from a TYPE: A.M.N.H. No. 10095. Nearly com- fragment collected at Taubate. It agrees in plete specimen with dorsal, anal, and tip of all noted characters with the specimen de- caudal fin missing. scribed above excepting a smaller dorsal fin HORIZON AND LOCALITY: ?Pliocene beds at formula XIV, 8-9. The number of vertebrae Tremembe, Sao Paulo. in this form is unknown. Woodward (1939) DIAGNOSIS: Length to end of vertebral has also described another cichlid, Macracara column about 13.75 cm, greatest depth about prisca, from presumably early Tertiary beds 7.5 cm. Skull about one-third total length of exposed at Nova York, Maranhao. It differs body. Dentition as in A. tetramerus. Cheek from the Aequidens group in having 36 verte- and opercular bones scaled, excepting pre- brae but is otherwise very similar. opercular. Vertebrae about 26-28: abdominal The Cichlidae are not known from the about 10-12, caudal 16. Dorsal fin with ap- Tertiary of North America. Regan (1908) proximately 29 spines and rays; pectoral fin and Myers (1937) are of the opinion that 13-14; pelvic fin I, 5; anal fin III, 8-10. Ori- Priscacara and Cockerellites, considered to be gin of anal fin opposite posterior third of dor- cichlids by Cope, Haseman, and Pearson, are sal. Scales finely ctenoid. more closely related to the sunfishes, and REMARKS: Regan (1905) thoroughly re- Romer (1945) has placed them in the Poma- viewed the osteological characters of Acara centridae. One genus, Cichlasoma, is found tetramerus and related species, but did not clear in the Miocene of the West Indies and also up the taxonomy of Acara (see Eigenmann, occurs today in South America. Three genera, 1910, p. 470). A. pauloensis (pl. 4, fig. 3) two of which are still in existence, are found agrees with the other species of Aequidens, fossil in Africa. Palaeochromis from the Ter- such as A. tetramerus, in the relative length tiary of North Africa is considered by Sau- of the ascending process of the premaxillary vage (i907) to be related to Acara. Such resem- which just extends to the orbit. The total blance, however, is probably due to the gen- number of vertebrae appears to be the same, eralized cichlid pattern shared by both, which although it cannot be definitely determined may also account for the resemblance be- 1 From State of Sao Paulo in which specimen was tween many of the Recent South American found. and African cichlids. DISCUSSION ANALYSIS OF FOSSIL ASSEMBLAGE

THE FOSSIL ACTINOPTERYGIAN FISHES that (1935), the resemblance between the Creta- have been described from the Cretaceous and ceous fauna of Brazil and that of the Gabon Tertiary deposits of Brazil are listed in table Basin in West Africa is very striking. Diplo- 2. Until the ages of the various formations mystus longicostatus from the S6rie Baia and discussed herein are accurately determined, D. goodi from the Gabon Basin are very simi- it appears advisable to consider the known lar and, as pointed out earlier, apparently actinopterygians together, particularly since belong to the same phyletic line. The simi- there appears to be no sharp faunal break larity further includes closely related chan- between the Cretaceous and Tertiary, and ids, the chirocentrids Chiromystus and ?Chiro- the position of the Serie Alagoas is still ques- centrites, Enchodus, Leptolepis, and a lepi- tionable. dotid with squamation resembling that of L. As in similar faunal assemblages from other mawsoni. This faunal resemblance, however, parts of the world, this one demonstrates the does not necessarily imply a close age corre- rise to dominance of the teleosts and, in this lation between the beds in Africa and Brazil. case, the introduction into South America of The deposits of the Gabon Basin are consid- its typical fresh-water fish fauna. The Cre- ered by the above authors to be Middle Cre- taceous assemblage, while not very diversi- taceous (Lower Cenomanian or Albian), fied, is typical of the period, and, as will be while the beds in Brazil yielding this assem- pointed out below, shows a great similarity blage are considered to represent a much to another fauna of similar age. With the last greater portion of the Cretaceous (Middle occurrence of Lepidotus and Palaeobalistum Albian through the Maestrichtian). Until the in the Upper Cretaceous, the rather meager age of the formations involved is better un- evidence indicates that the Holostei disap- derstood, the significance of this resemblance, peared from South America. Possibly the except in a general way, largely remains a amiids persisted into the Teritary, although moot question. the single specimen thus far discovered be- The questionable early Tertiary deposits in longing to this group comes from the Upper Baia, regarded at present by the Brazi!ian Cretaceous. Survey as possibly Upper Cretaceous in age, The Upper and Lower Cretaceous assem- contain a fauna that unfortunately offers blage is made up of holosteans and teleosteans little assistance in settling this problem, as from marine and estaurine beds. Continental many of the genera represented are known Cretaceous deposits are rare in Brazil, if not to occur in formations of both periods. The entirely missing, although they are found in later Tertiary deposits in Sao Paulo, possibly the western part of South America. In any Pliocene, are entirely lacustrine-fluvial and case, teleosts from such beds are likewise rare, contain a fresh-water fauna that is Recent in and the Brazilian picture further emphasizes character, indicating that this assemblage Romer's suggestion (1945) that the teleosts was already in existence by no later than mid- very probably had a marine origin. Tertiary time. As stated by Arambourg and Schneegans THE ORIGIN OF THE SOUTH AMERICAN FRESH-WATER FISH FAUNA The origin of the fresh-water fish assem- evidence by Haseman, Myers, and Gosline, blage of South America has been the subject and these authors are well aware of its im- of much speculation that is based, for the portance. As the problem is essentially a most part, on information obtained from historical one, the ultimate solution can be studies of the Recent fauna. There has been revealed only by paleontological and geo- some consideration of the paleontological logical data. 30 1947 SCHAEFFER: ACTINOPTERYGIAN FISHES FROM BRAZIL 31 TABLE 2 FossIL ACTINOPTERYGII FROM THE CRETACEOUS AND TERTIARY OF BRAZIL Cretaceous Tertiary

0 K.J 0 o~~~~~~~~~~eC

a) c a UbO H 3- 0 $ 0 03- Cd ~ % ~0 Cd Cd C 0.a)a~~~~~~~~~~~ a) ~~~~~~~ ~~~0da)

0 0 0 Cl) (iT) ~~ (iT) c HOLOSTEI Semionotoidea Semionotidae Lepidotus temnurus Agassiz, 1841 X Lepidotus mawsoni Woodward, 1888 X Lepidotus souzai Woodward, 1908 X Lepidotus piauhyensis Roxo and L6fgren, 1936 ?X Pycnodontoidea Pycnodontidae Palaeobalistumflabellatum (Cope, 1885) X Palaeobalistum dossantosi Maury, 1930 X Aspidorhynchoidea Aspidorhynchidae Vinctifer comptoni (Agassiz, 1841) X Belonostomus (?) carinatus Woodward 1907 X Amioidea Amiidae Urocles mawsoni (Woodward, 1902) X TELEOSTEI ISOSPONDYLI Clupeoidea Leptolepidae Tharrhias araripis Jordan and Branner, 1908 X Tharrhias rochae (Jordan and Branner, 1908) X Leptolepis bahiaensis, new species ?X Elopidae Brannerion vestitum (Jordan and Branner, 1908) X Calamopleurus brama (Agassiz, 1841) X Rhacolepis buccalis Agassiz, 1841 X Rhacolepis latus Agassiz, 1841 X Rhacolepis de-fiorei D'Erasmo, 1938 X Anaedopogon tenuidens Cope, 1871 X Enneles audax Jordan and Branner, 1908 X Ennelichthys derbyi Jordan, 1921 X Lastroichthys longipectoralis, new genus

and species X-- 32 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL.' 89 TABLE 2-Continued Cretaceous Tertiary ... .~ ~~~~~~~~~~~ ~~ ~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~ 0 C0 '4a 0 0 '.4 Sz . cd Sd S Ho o Se (L) E- aL) E 0 CS C,) 0- co Cfj W. 0 ca) S. '4 <4 . Sd E 40J CS ,0 Cd Cd U) ':4 &. 0 0 0 PQ 0 E 4-9 04 a) aL) a) . 0d Ei 4 Ei -40-4 ',4 '.4 '.4 HE '4 U) .-q 'a) 'a)%V CQ Cl) 1:4 Cl) z Chirocentridae Cladocyclus gardneri Agassiz, 1841 x Chiromystus mawsoni Cope, 1885 x Cliromystus alagoensis Jordan, 1910 x Chanidae Dastilbe crandalli Jordan, 1910 x x Clupeidae Scombroclupeoides scutata (Woodward, 1908) x x Diplomystus longicostatus Cope, 1885 x Knightia branneri (Jordan, 1910) x x Knightia brasiliensis Woodward, 1939 X Stomiatoidea Enchodontidae Enchodus oliveirai Maury, 1930 x Enchodus subequilateralis Cope, 1885 x Cimolichthys n. sp. Williston, 1902 x OSTARIOPHYSI Cyprinoidea Characinidae Eobrycon avus (Woodward, 1898) Eobrycon branneri Eigenmann, 1929 X Eobrycon ligniticus (Woodward, 1898) Siluroidea X Ariidae Arius iheringi Woodward, 1898 Xx ACANTHOPTERYGII Percoidea X Serranidae Percichthys an.tiquus Woodward, 1898 xX Cichlidae Acara sp. Woodward, 1898 Aequidens pauloensis, new species Macracara prisca Woodward, 1939 x

There are probably but four possibilities tribution. It is furthermore probable that the that might be considered to explain the dis- various elements of the fauna employed dif- persal of a fresh-water fish fauna into South ferent or several means of dispersal, although America (or Africa): (a) by marine radiation, there is no way of demonstrating this at the (b) by land bridges, (c) by direct contact of present time. While each of these alternatives South America and Africa prior to drifting has received much attention, the paleontolog- apart, and. (d) by Holarctic continental dis- ical evidence that has accumulated in recent 1947 SCHAEFFER: ACTINOPTERYGIAN FISHES FROM BRAZIL 33 years has not been considered. These data are positively recognizable fossil cichlids have still very fragmentary, but they do suggest been found in North America (the prisca- certain conclusions such as the early arrival carids possibly being related to the sunfishes of the characins and the possible late appear- or close to the base of percoid radiation) or ance of the osteoglossids, and they offer a Europe. It is interesting to note, however, basis for future investigation along possibly that Myers places this family in the second- less speculative lines. ary division, and Dr. C. M. Breder has in- formed the writer that a species of Tilapia MARINE DISTRIBUTION lived for a number of years in sea water at Myers (1938) has analyzed the Recent the New York Aquarium. Hence a marine South American fresh-water fish fauna from dispersal is again a very reasonable possi- the standpoint of its origin. In this connec- bility, particularly with Simpson's discussion tion, he separates all fresh-water fishes into (1939) of the Galaxias problem in mind. two categories: (1) a primary division con- The amiids, placed in the primary division, sisting of fishes that are always restricted to were spread rather widely over the Northern fresh water, and (2) a secondary division in- Hemisphere by the beginning of the Creta- cluding those fishes that are able to enter the ceous. As the Jurassic members of this family sea for a short period of time. Of the various occur in marine formations, a marine dispersal fishes which he places in each division, only is a reasonable possibility. It is therefore not those that have been found as fossils in Brazil particularly surprising that an amiid, Urocles will be considered here. mawsoni, should occur in the Serie Baia. It is The distinctness of the Recent South rather more surprising that amiid remains American fresh-water fish fauna centers are not more common in South America and around the presence of characins and cichlids, have not been found in Africa. both also occurring as fossils and Recent The siluroid genus Arius is semimarine, types in Africa. The Characinidae are placed and the Tertiary A. iheringi from the lacus- by. Myers in the primary division and the trine-fluvial deposits at Taubate was prob- Cichlidae in the secondary. As pointed out ably a landlocked species. The evidence re- in the section on Eobrycon, the identification garding the dispersal of the catfishes is care- -of characins from scales in Upper Cretaceous fully summarized by Gosline (1944) and will marine beds in California, Wyoming, and not be discussed here. It will suffice to state Peru and their presence in lacustrine Eocene that Cretaceous siluroids are unknown, and beds in France indicate a far wider distribu- the Tertiary and Recent forms do not offer tion for this family than has been generally even a tentative solution to this problem. realized. It is therefore within the realm of Gosline suggests, but considers improbable, possibility that the characins were distrib- that the African and the South American uted to the continents by way of the sea -catfishes are all descended from the Ariidae. during the late Mesozoic and were restricted The presence of the serranid Percichthys to- fresh water by the early Tertiary. It is in the Taubate deposits, in the Lower Eocene further possible that the members of this of Patagonia, and at the present time in the group became extinct in North America and streams of Chile and Patagonia suggests that Europe during the Tertiary because of tem- this genus has been restricted to fresh water perature or other environmental changes. since the early Tertiary, as has likewise been While such conclusions can be substantiated the Australian Percalates (Hills, 1934). The only by further discovery, they offer an alter- world-wide distribution, by the early Tertiary, native explanation for the distribution of the of such transitional percid-serranid types characins without the creation of dubious again suggests a marine distribution in the late South Atlantic and Antarctic land bridges. Mesozoic. It is becoming increasingly evident A clue regarding the dispersal of the cich- that the key to fresh-water fish dispersal lies lids is still to be found. As in the case of the -in the Cretaceous. By the beginning of the characins, the known fossil representatives Tertiary, the various fresh-water groups from South America and Africa are closely were restricted to their present habitat and related to Recent species, and thus far no differentiated to almost the same degree as 34 BULLETI-N AMERICAN MUSEUM OF NATURAL HISTORY VOL. 89 their Recent descendants. Gosline's studies order has never had a greater distribution appear to support this conclusion. than at present, and that the fossil types are Fossil osteoglossids have thus far not been all related to genera living today. In this con- 'discovered in South America. They occur pos- nection, the evidence for the Holarctic pres- sibly as early as the Cretaceous in North ence of the characins will be recalled. It is America (?Eurychis), certainly by the Eocene true, however, that the record of ostario- in both this continent (Phareodus) and in physian evolution is still very fragmentary, North Africa and Europe (Brychetus). and most of the fossil types known are closely Phareodus is also present in the Oligocene of related to Recent genera. As pointed out Australia in a fresh-water deposit (Hills, earlier and agreed to by Gosline, the ostario- 1934). Myers places the osteoglossids in his physids undoubtedly reached South America primary division, although the early mem- during the Mesozoic. The scarcity of conti- bers of this group again were probably ma- nental Mesozoic beds in South America might rine, which could account for their very wide -account for our ignorance concerning them dispersal by the Eocene or Oligocene. The during this era, or the scarcity of their re- resemblance between the recent Australian mains in Cretaceous marine beds, the period genus Scleropages and Phareodus is not sur- during which they may have evolved, could prising, as the latter existed in the fresh-water be another factor. Furthermore, it may prove streams of Australia prior to the isolation of very difficult or impossible to distinguish the that continent. There are several conceivable primitive ostariophysids from the clupeid- explanations for the lack of fossil osteoglos- like fishes of the Mesozoic, a case in point sids in South America: first, that they invaded being the possible origin of the Order Ostario- this continent from the sea in the Cretaceous physi from the Jurassic-Cretaceous Lycop- and their remains have as yet not been found teridae. Lastly, Gosline raises the question in either marine or continental beds, and sec- of why, if the Ostariophysi could cross a ond, that if they were entirely restricted to marine barrier, they have not been distribut- fresh water by the Eocene or Oligocene, they ed to a greater extent by this means. If fur- were unable to enter South America until the ther discovery demonstrates that the Ostario- end of the Tertiary when the connection with physi had a marine dispersal, the question North America was reestablished. This prob- more or less answers itself, and also a physio- lem can, again, be solved only by future dis- logical and structural restriction to fresh covery. water by the Eocene would certainly tend to Gosline (1944) has also dwelt upon the reduce or eliminate marine distribution after problem of the origin of the South American that time. fresh-water fish fauna with particular refer- The somewhat scanty evidence here pre- ence to the derivation of the Ostariophysi. He sented would appear to offer an alternative considers, for instance, the question of salt- hypothesis for the origin of the South Ameri- water tolerance and the possiblity of the can fresh-water fauna without the necesssity Weberian apparatus being particularly suited of creating a land bridge-between that con- or evolved for fresh water. Unfortunately, tinent and Africa. no amount of work on recent members of this order regarding either of these problems can LAND BRIDGE DISPERSAL do more than increase or decrease the prob- With no apparent other possibilities, the ability of a marine origin or marine dispersal. land bridge hypothesis has been generally The millions of years involved in such a hab- proposed as the one credible solution to the itus change might very well obscure changes problem. That fresh-water fishes are able to in the histological structure of the kidney, migrate across a land bridge is demonstrated such as the relative number of the glomeruli by the northward movement of the cichlids in fresh- and salt-water types, and also result into Texas and the characins into New Mex- in many minor changes in the Weberian ap- ico by way of the Panamanian connection paratus. (the possibility that the cichlids and chara- Regan has stated that the imperfect past cins -once existed in North America and are history of the Ostariophysi indicates that this now restricted, by temperature, to the South- 1947 SCHAEFFER: ACTINOPTERYGIAN FISHES FROM BRAZIL 35 west must also be considered). A migration and cichlids, although their paleontological of this sort, excluding transportation by history is unknown. In any case, it is rather birds, cyclonic storms, etc., could be accom- surprising that, with the two continents plished probably in only two ways, either by united according to the above theory, even stream capture or by passage through the these more ancient types did not enter South sea from one river mouth to another. The America. As Myers infers (1938, p. 353), how- latter method might be employed by fishes ever, a filter bridge between the two conti- of Myer's second division, such as the cich- nents might have excluded them but permitted lids. There is evidence, of course, that Recent exchange of the other groups. fresh-water fishes do use both means of extending their distribution. It is logical, there- HOLARCTIC CONTINENTAL DISTRIBUTION fore, that fishes of this type could cross a Finally, a Holarctic continental distribu- land bridge, given enough time, between tion of the various groups under discussion South America and Africa. The northward remains to be considered. In spite of the pres- dispersal of the characins and cichlids must ence of marine amiids in the Jurassic, it is have occurred in a relatively short period, entirely possible that they were restricted probably beginning no earlier than the end of to fresh water in the Northern Hemisphere the Tertiary when the connection between during the Cretaceous, entering South North and South America was reestablished. America by way of the Panamanian bridge It may hence be concluded that if land before it was cut off. The indication for bridges did exist, with a series of originally this is, however, not particularly convinc- separate drainage systems, the migration of ing because of the lack of any positive fresh-water fishes could take place across paleontological evidence. The same sort of them. But the fact still remains that there is reasoning might be developed for the osteo- little geological evidence for the past exist- glossids and the characins on the basis of the ence of oceanic land bridges (excepting the meager paleontological evidence. The case Siberian-Alaskan bridge), or, for that matter, for the osteoglossids is possibly on a sounder of the extensive orogenic disturbances of the footing with the positive identification of the ocean floor that would be necessary to pro- North American Eocene Phareodus, even duce such bridges. To create them with no though this particular genus is apparently geological basis, just to satisfy the needs of not closely related to either Osteoglossum or zooge6graphers, though commonly done, Arapaima. Haseman (1912) developed a num- seems unsound. ber of valid arguments against the land bridge theory and in turn supported a north- DISTRIBUTION AND CONTINENTAL DRIFT ern origin for the South American fresh- Although the hypothesis of continental drift water fish fauna. He considered the known has -generally fallen into disrepute in recent paleontological evidence at some length, rely- years, it is interesting to consider the possibil- ing rather heavily on the much-worked Green ities offered for distribution. According to one River fishes. As Myers (1937) has pointed theory, South America and Africa started to out, however, this fauna does not clearly sup- separate in the early Cretaceous, thus requir- port a northern origin, and this is particu- ing a fresh-water characin exchange by no larly true in the case of the cichlids. later than this time. Myers (1938) points out The confusing picture presented by the the very interesting fact that none of the five Recent distribution of vertebrates in South primitive autochthonous African families of America and Africa can, as has been demon- fishes (bichirs, pantodonts, phractolaemids, strated in the case of the mammals, be re- kneriids, and mormyrids) are found in South solved only by paleontological discovery America. These families, he believes, were backed by reasonably probable geological present in Africa before the characins, carps, evidence. REFERENCES The literature on the fossil actinopterygians of COUTO, CARLOS DE PAULA Brazil is widely scattered and for that reason an 1938. A paleontologia do Brazil. Rev. Inst. attempt has been made to make this list as com- Hist. Geogr. Rio Grande do Sul, ano plete as possible. 18, trim. 3, pp. 113-141, figs. 1-8. CUVIER, GEORGES L. C. F. D. AGASSIZ, L. 1817. Le regne animal. Paris, Deterville, vol. 1841. On the fossil fishes found by Mr. 2, xviii+532 pp. Gardner in the Province of Ceari, in DAVID, LORE ROSE the north of Brazil. Edinburgh New 1941. Leptolepis nevadensis, a new Cretaceous Phil. Jour., vol. 30, pp. 82-84. fish. Jour. Paleont., vol. 15, no. 3, pp. 1884. Sur quelques poissons fossiles du Bresil. 318-321, figs. 1-2. Compt. Rendus Acad. Sci. Paris, vol. 1946. Upper Cretaceous fish remains from 28, pp. 1007-1015. the western border of the San Joaquin ALLPORT, SAMUEL Valley, California. Carnegie Inst. Wash- 1860. On the discovery of some fossil remains ington Publ., no. 551, pp. 81-112, figs. near Bahia in South America. Quart. 1-11, pls. 1-3. Jour. Geol. Soc. London, vol. 16, pt. 1, DECHASEAUX, COLETTE pp. 263-266, pls. 14-17. 1943. Contribution a l'etude du genre Lepi- ARAMBOURG, CAMILLE dotus. Ann. Paleont., vol. 30, pp. 3-13, 1935a. Contribution a l'etude des poissons du pls. 1-3. Lias superieur. Ann. Paleont., vol. 24, DE SAEZ, MATHILDE DOLGOPOL pp. 3-32, pls. 1-3. 1939. Noticias sobre peces f6siles Argentinos. 1935b. Observations sur quelques poissons fos- Notas Mus. La Plata, vol. 4, Paleont. siles de l'ordre des halecostomes et sur no. 19, pp. 423-432, figs. 1-5. l'origine des clupeid6s. Compt. Rendus DUNKLE, DAVID HOSBROOK Acad. Sci. Paris, vol. 200, no. 25, pp. 1940. The cranial osteology of Notelops brama 2110-2112. (Agassiz), an elopid fish from the Cre- ARAMBOURG, CAMILLE, AND D. SCHNEEGANS taceous of Brazil. Lloydia, vol. 3, pp. 1935. Poissons fossiles du bassin s6dimentaire 157-190, figs. 1-9. du Gabon. Ann. Paleont., vol. 24, pp. 1942. A new fossil fish of the family Lepto- 137-160, pls. 18-20. lepidae. Sci. Publ., Cleveland Mus. Nat. CABRERA, ANGEL Hist., vol. 8, no. 5, pp. 61-64, pl. 6. 1927. Sobre un pez f6sil del lago San Martin. EASTMAN, CHARLES R. Rev. Mus. La Plata, vol. 30, pp. 317- 1912. Tertiary fish-remains from Spanish 319, 1 fig. Guinea in West Africa. Ann. Carnegie COCKERELL, T. D. A. Mus., vol. 8, no. 2, pp. 370-378, pls. 1919. Some American Cretaceous fish scales, 23-24. with notes on the classification and dis- 1917. Fossil fishes in the collection of the tribution of Cretaceous fishes. Prof. United States National Museum. Proc. Papers, U. S. Geol. Surv., no. 120, pp. U. S. Natl. Mus., vol. 52, pp. 235-304, 165-188, pls. 31-37. pls. 1-23. 1921. Some fossil fish scales from Peru. Proc. EIGENMANN, CARL H. U. S. Natl. Mus., vol. 59, pp. 19-20, 1909. The fresh-water fishes of Patagonia and figs. 1-7. an examination of the Archiplata- COPE, EDWARD DRINKER Archhelenis theory. Rept. Princeton 1871. On two extinct forms of Physostomi of Univ. Exp. Patagonia 1896-1899, vol. the- neotropical region. Proc. Amer. 3, pt. 3, pp. 225-374, pls. 30-37. Phil. Soc., vol. 12, no. 86, pp. 52-55. 1910. Catalogue of the fresh-water fishes of 1877. A contribution to the knowledge of the tropical and south temperate America. icthyological fauna of the Green River Ibid., vol. 3, pt. 4, pp. 375-511. shales. Bull. U. S. Geol. Surv., vol. 3, 1927. The fresh-water fishes of Chile. Mem. art. 34, pp. 807-819. Natl. Acad. Sci., vol. 22, menu 2, pp. 1885. A contribution to the vertebrate paleon- 1-63, pls. 1-16. tology of Brazil. Proc. Amer. Phil. Soc., EIGENMANN, CARL H., AND W. L. BRAY vol. 23, pp. 1-21, 1 pl. 1894. A revision of the American Cichlidae. 36 1947 SCHAEFFER: ACTINOPTERYGIAN FISHES FROM BRAZIL 37 Ann. New York Acad. Sci., vol. 7, pp. JORDAN, DAVID STARR 607-624. 1907. The fossil fishes of California with sup- EIGENMANN, CARL H., AND GEO. S. MYERS plementary notes on other species of 1929. The American Characidae. Mem. Mus. extinct fishes. Univ. California Publ., Comp. Zool., Harvard College, vol. 43, Bull. Dept. Geol., vol. 5, no. 7, pp. 95- pt. 5, pp. 429-558, pls. 1-101. 144, pls. 11-12. ERASMO, GEREMIA D' 1910. Description of a collection of fossil fishes 1938a. Ittioliti cretacei del Brasile. Atti Accad. from the bituminous shales at Riacho Sci., Napoli (3), vol. 1, no. 3, pp. 1-44, Doce, State of Alag6as, Brazil. Ann. pls. 1-6. (Not available.) Carnegie Mus., vol. 7, no. 1, pp. 23-34, 1938b. Ittioliti cretacei del Brasile (Sunto della pls. 5-13. memoria inserita nel, vol. 1, ser. 3, degli 1913. ERimma, a genus of fossil herrings. Proc. Atti). Rendiconto Accad. Sci. Fis. Biol. Soc. Washington, vol. 26, p. 79. Math., Naples, ser. 4, vol. 8, p. 119. 1919. New genera of fossil fishes from Brazil. GARDNER, GEORGE Proc. Acad. Nat. Sci. Philadelphia, vol. 1841a. Fossil fishes of Ceari. Edinburgh New 71, pp. 208-210. Phil. Jour., vol. 30, pp. 75-81. 1920. The genera of fishes. Part IV. Leland 1841b. On the geology and fossil fishes of north Stanford Jr. Univ. Publ., univ. ser., pp. Brazil. Rept. Brit. Assoc. Adv. Sci. for 411-576. 1840, Trans. of Sections, pp. 118-120. 1921. Peixes cretaceos de Ceark e Piauhy. 1842. Peixes petrificados que se acham na Monogr. Serv. Geol. Min. Brasil, Rio Provincia do Ceari. Jor. Commercio, de Janeiro, vol. 3, pp. 1-97, pls. 1-16. Rio de Janeiro, no. 95. (Not available.) JORDAN, DAVID STARR, AND JOHN CASPER BRAN- NER GOSLINE, WILLIAM A. 1908. The Cretaceous fishes of Ceari, Brazil. 1944. The problem of the derivation of the Smithsonian Misc. Coll., vol. 5, pt. 1, South American and African fresh- pp. 1-29, pls. 1-8. water fish faunas. An. Acad. Brasileira JORDAN, DAVID STARR, AND JAMES ZACCHEUS Cien., vol. 16, no. 3, pp. 211-223. GILBERT GREGORY, WILLIAm K., AND G. MILES CONRAD 1919. Fossil fishes of southern California. II. 1938. The phylogeny of the characin fishes. Fossil fishes of the (Miocene) Monterey Zoologica, vol. 23, pt. 4, pp. 319-360, formations of southern California. Le- figs. 1-37. land Stanford Jr. Univ. Publ., univ. ser., HASEMAN, JOHN D. pp. 13-60, pls. 8-28. 1912. Some factors of geographical distribu- LINNAEUS, CAROLUS tion in South America. Ann. New York 1758. Systema naturae.... Editio decima, re- Acad. Sci., vol. 22, pp. 9-112, pls. 2-16. formata. Stockholm, Laurentii Salvii, HECKEL, J. J. 824 pp. 1840. Johann Natterer's neue Flussfische MCCULLOCH, ALLAN R. Brasilien's nach den Beobachtungen 1917. Studies in Australian fishes. Rec. Aus- und Mittheilungen des Entdeckers be- tralian Mus., vol. 11, no. 7, pp. 163-188, schriehen. Abth. 1. Die Labroiden. Ann. figs. 1-2, pls. 29-31. Wiener Mus. Naturg., vol. 2, pp. 325- MAURY, CARLOTTA JOAQUINA 470, 1 fig. 1930. 0 Cretaceo da Parahyba do Norte. HILLS, EDWIN SHERBON Monogr. Serv. Geol. Min. Brasil, Rio de 1934. Tertiary fresh water fishes from south- Janeiro, vol. 8, pp. 1-305, pls. 1-35. ern Queensland. Mem. Queensland 1936. 0 Cretaceo de Sergipe. Ibid., vol. 11, pp. Mus., vol. 10, pt. 4, pp. 157-174, pls. 1-283, pls. 1-28. 18-25. MAWSON, JOSEPH, AND A. S. WOODWARD HUSSAKOF, LOUIS 1907. On the Cretaceous formation of Bahia 1917. Fossil fishes collected by the Ameri- (Brazil), and on the vertebrate fossils can Museum Congo expedition. Bull. collected therein. Quart. Jour. Geol. Amer. Mus. Nat. Hist., vol. 37, art. Soc. London, vol. 63, no. 250, pp. 128- 27, pp. 761-767, figs. 1-7, pl. 88. 139, 1 fig., pls. 6-8. IHERING, H. VON MYERS, GEORGE S. 1898. Observag6es sobre os peixes fosseis de 1938. Fresh-water fishes and West Indian Taubate. Rev. Mus. Paulista, Sao zoogeography. Smithsonian Rept. for Paulo, vol. 3, pp. 71-75. 1937, publ. 3465, pp. 339-364, pls. 1-3. 38 BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY VOL. 89 OLIVEIRA, AVELINO IGNACIO DE, AND OTHON Amer. Mus. Novitates, no. 1331, pp. HENRY LEONARDOS 1-9. 1943. Geologia do Brasil. Serv. Inf. Agr., SIMPSON, GEORGE GAYLORD Ministerio Agr., Rip de Janeiro, 813 1939. Antarctica as a faunal migration route. pp., 202 figs., 1 map. (Contains exten- Proc. Sixth Pacific Sci. Congr., pp. 755- sive bibliography.) 768. OLIVEIRA, EUZEBIO PAULO DE SMIRNOV, V. P. 1939. Atlas geologico do Brasil. Ministerio, 1935. [Fishes of the north Caucasus Oligocene Agr., Dept. Nac. Prod. Min., Serv. Geol. (Chernaya Rechka region).] Trav. Sci. Min., Rio de Janeiro, pp. 1-18. Univ. Usbekistan, vol. 1, pp. 41-115, PITON, L. figs. 1-18. (Not available.) 1938. Les Characinidae fossiles de Menat. 1936. [Fishes of the north Caucasus Oli- Rev. Sci. Bourbonnais, nos. 3-4, pp. gocene.] Ibid. (1936), pp. 1-96, figs. 1- 98-104, figs. 1-2. 23, pls. 1-13. (Not available.) RAYNER, DOROTHY H. Spix, J. B. VON, AND C. F. P. VON MARTIUS 1937. On Leptolepis bronni Agassiz. Ann. Mag. 1828. Reise in Brasilien, 1817-1820. Munich, Nat. Hist., ser. 10, vol. 19, pp. 46-74, I. J. Lentner, vol. 2, pp. i-viii, 415-884. figs. 1-14. TANNER, VASCO M. REGAN, CHARLES TATE 1925. Notes on the collection of fossil fishes 1905. A revision of the fishes of the South contained in the University of Utah American cichlid genera Acara, Nan- collection, with the description of one nacara, Acaropsis, and Astronotus. Ann. new species. Bull. Univ. Utah., vol. 15, Mag. Nat. Hist., ser. 7, vol. 15, pp. no. 6, pp. 1-13, pls. 1-3. 329-347. THORPE, MALCOLM RUTHERFORD 1908. Pisces. In Godman, Frederick D., and 1938. Wyoming Eocene fishes in the Marsh 0. Salvin [editors]. Biologia Centrali- collection. Amer. Jour. Sci., vol. 36, pp. Americana. London, pt. 193, xxxii+203 279-295, figs. 1-8. pp., 26 pls., 2 maps. VOIGHT, EHRHARD ROMER, ALFRED SHERWOOD 1934. Die fische aus der mitteleoziinen Braun- 1945. Vertebrate paleontology. Chicago, Uni- kohle des Geiseltales. Nova Acta Leo- versity of Chicago Press, x+687 pp., poldina, new ser., vol. 2, pp 21-144, 377 figs., 4 tables. figs. 1-23, pls. 1-14. Roxo, MATHIAS G. DE OLIVEIRA WEILER, WILHELM 1936. Revalidasao do Thoracosaurus bahiaensis 1923a. Die Fischreste aus den bituminosen e consideraoes sobre a edade da Serie Scheifern von Ibando bei Bata (Spanish da Bahia. Bol. Mus. Nac., Rio de Guinea). Paleont. Zeitschr., vol. 5, pp. Janeiro, vol. 12, nos. 3-4, pp. 59-72, 1 148-160, figs. 1-5. chart. 1923. Nachtriagliche Bernerkungen zur Ab- Roxo, MATHIAS G. DE OLIVEIRA, AND AXEL handlung iiber die Fischreste aus den LOFGREN bituminosen Schiefern von Ibando. 1936. Lepidotus piauhyensis, sp. nov. Notas Ibid., vol. 5, p. 209. Prelim. Estudos, Serv. Geol. Min., Rio 1935. Ergebnisse der Forschungsreisen. Prof. de Janeiro, no. 1, pp. 7-12, figs. 1-2. E. Stromers in den Wusten Agyptens. SANTOS, RUBENS DA SILVA II. Wirbeltierreste der Baharije-Stufe 1945a. Estudo morfologico de Lepidotus pia- 16. Neue Untersuchungen an der Fisch- uhyensis. Notas Prelim. Estudos, Div. resten. Abhandl. K. Bayerischen Akad. Geol. Min., Rio de Janeiro, no. 28, pp. Wiss., Math.-Nat. Abt., new ser., vol. 1-18, pls. 1-4. 32, pp. 1-57, figs. 1-4, pls. 1-3. 1945b. Revalidaglo de Aspidorhynchus comp- WILLISTON, SAMUEL W. toni Agassiz do Cret'aceo- do Cearfi, 1902. Cimolichthys n. sp. Bull. Geol. Soc. Brasil. Ibid., no. 29, pp. 1-12, pl. 1. Amer., vol. 13, pp. 44-45, fig. 1. SAUVAGE, H. E. WOODWARD, ARTHUR SMITH 1907. Sur des poissons de la famille des 1887. On the fossil teleostean genus Rhacolepis, cichlides trouves dans le terrain ter- Agassiz. Proc. Zool. Soc. London, pp. tiaire de Guelma. Compt. Rendus Acad. 535-542, pls. 46-47. Sci., Paris, vol. 145, pp. 360-361. 1888. Notes on some vertebrate fossils from SCHAEFFER, BOBB the Province of Bahia, Brazil, collected 1947. An Eocene serranid from Patagonia. by Joseph Mawson, Esq., F.G.S. Ann. 1947 SCHAEFFER: ACTINOPTERYGIAN FISHES FROM BRAZIL 39 Mag. Nat. Hist., ser. 6, vol. 2, no. 8, Part IV. London, British Museum pp. 132-136. (Natural History), xxxviii+636 pp. 1890. Vertebrate paleontology in some Ameri- 1902. On an amioid fish (Megalurus mawsoni, can and Canadian museums. Geol. Mag., sp. n.) from the Cretaceous of Bahia, ser. 3, vol. 7, pp. 390-395. Brazil. Ann. Mag. Nat. Hist., ser. 7, vol. 1891. On some upper Cretaceous fishes of the 9, no. 50, pp. 87-89, pl. 2. family Aspidorhynchidae. Proc. Zool. 1907. Notes on some upper Cretaceous fish- Soc. London (for the year 1890), pp. remains from the provinces of Sergipe 613-616, pls. 54-55. and Pernambuco, Brazil. Geol. Mag., 1895a. Catalogue of the fossil fishes in the new ser., decade 5, vol. 4, no. 5, pp. British Museum (Natural History). 193-197, pl. 7. Part III. London, British Museum 1908. On some fossil fishes discovered by (Natural History), xlii+544 pp. Prof. Ennes de Souza in the Cretaceous 1895b. On two deep-bodied species of the clupe- formation at Ilheos (State of Bahia), oid genus Diplomystus. Ann. Mag. Nat. Brazil. Quart. Jour. Geol. Soc. London, Hist., ser. 6, vol. 15, no. 85, pp. 1-3, pi. vol. 64, art. 20, pp. 358-362, pls. 42-43. 1. 1939. Tertiary fossil fishes from Maranhao, 1898. Considerag6es sobre alguns peixes Terci- Brazil. Ann. Mag. Nat. Hist., ser. 11, arios dos schistos de Taubat6, Estado de vol. 3, no. 16, pp. 450-453, pl. 16. S. Paulo, Brazil. Rev. Mus. Paulista, 1942. The beginning of the teleostean fishes. Sao Paulo, vol. 3, pp. 63-70, pls. 1-7. Ibid., ser. 11, vol. 9, no. 60, pp. 902-912. 1901. Catalogue of the fossil fishes in the figs. 1-7. British Museum (Natural History).

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