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BOLENEN SSS ce wha i — al —“ Ww at yi) = YWiw rs = bite ~eeuvre. <~ Suveuyvaulccuerce AAAS I Ce J I 4 DM asa VS US We Swe DAB Ape RAG we S wag Beery Vv PICS ly AL Tes, CX } vi, PT A ~/ Tp NN 1 "| yg UUs eel SNE RADAPAA LER “a ah A Wi VWuss w due wwe ee) \ SAS we ee Syvvuwyv ~ ww ot 3 a a ‘we win ve Vv ‘set hf he Be te asc ily BS ISIS SAA led eh OO hd bi eee, OV SOE o, ay ©, —_ | | | pws Ti, 5 et 1De ine Mt O; ei ogy l S ATES. WITH SIXTEEN PL Parr 1. of Comparative Zod ARD COLLEGE. I. yuseum ARY CAN CHARACIDAE OL. z LERT ‘CARL H. EIGENMANN. . 1917 CAMBRIDGE, U. S. A. Aueust, PRB cs SO a aire MR ne RE em Y ye d@emoirs of the Auseum of Comparative Zodlogy M/ 2% AT HARVARD COLLEGE. ; S E34 Vou. XLII. Parr 1. 1417) v.43 pr Fishes THE AMERICAN CHARACIDAE. BY CARL H. EIGENMANN. 276 16%, WITH SIXTEEN PLATES. CAMBRIDGE, U. S. A.: Printed for the Museum. Aueust, 1917. THE AMERICAN CHARACIDAE.’ INTRODUCTION. THE revision of the American Characidae, based on the collections of the Nathaniel Thayer Brazilian Expedition in the Museum of Comparative Zodlogy was commenced by Mrs. Rosa Smith Eigenmann and myself in 1888. A little more than a year later the work was temporarily suspended. The published results, so far as obtained, are enumerated on p. 8-10. Work was resumed in 1903 with the collections of Indiana University and of the United States « National Museum, and a monograph completed in 1906. Types in several museums in London, Paris, and Vienna were examined in 1906 and 1907. In 1907 the study of the collections made by the Thayer Expe- dition was renewed. This material made it necessary to rewrite the entire monograph, for, forty years after it was gathered, it was still the most impor- tant collection of South American fresh-water fishes, and contained a large number of undescribed species and genera. The revised monograph, through the Tetragonopterinae, was finished in the spring of 1908. During the preparation of this revision it became apparent that the material at my command, from several regions of South America, was deficient. To obtain material from one of these regions, I spent the autumn of 1908 in British Guiana. Volume 5 of the Memoirs of the Carnegie Museum contains a detailed account of this Guiana Expedition. In May, 1909, my position of Curator of Ichthyology in the Carnegie Mu- seum placed me in charge of another large series of South American fresh-water fishes. This collection was made under my general direction, between November 6, 1907 and January 10, 1910, by Mr. John D. Haseman, one of my former students. For the most part it came from areas not covered by the Thayer Expedition. The Director of the Carnegie Museum, Dr. W. J. Holland, has kindly allowed me to incorporate in the present monograph the results obtained from a study of the Carnegie collections. 1 Contribution from the Zoélogical Laboratory of Indiana University, No. 98. 4 THE AMERICAN CHARACIDAE. Still later (December 1911—April 1912) I made extensive collections in Colombia; and in the early part of 1913 two of my students, Mr. Arthur M. Henn and Mr. Charles Wilson, gathered many fishes in the streams of western Colombia. Mr. Henn remained in South America till 1914 and extended the explorations to western Ecuador. Smaller collections were made by Miss Lola Vance in the streams about Tarma, Peru, and by Dr. Ellis and Dr. William Tucker in British Guiana. The collections of the Yale-National Geographic Society Expedition to the Urubamba River have also been available in the preparation of this monograph. These different collections admirably supplement each other. They are the most extensive collections that have ever been brought together from South America. In the number of specimens at my disposal they far exceed the com- bined collections recorded in all other museums of the world. The number of American species of Characins exceeds six hundred; and as they offer some of the best material to demonstrate several facts of evolution it is hoped to illustrate the species quite thoroughly. To provide for the illus- trations of the Tetragonopterinae I am indebted to the Trustees of the Eliza- beth Thompson Science Fund for an appropriation of $250.00, and to the Trustees of the Bache Fund for a similar amount. The Carnegie Museum and Mr. H. W. Fowler have granted the reproduction of a number of figures. The portion of the monograph dealing with Hemigrammus, Hyphessobrycon, and Hasemania was prepared with the collaboration of Marion Lee Durbin, now Marion Durbin Ellis. This Memoir deals with the Tetragonopterinae,! Rhoadsinae, Glandulo- caudinae, Iguanodectinae, Stethaprioninae, and Stichanodontinae. All of the then known species of these subfamilies, except the species of Stevardia, were included by Giinther (Cat. fishes Brit. Mus., 1864, 5) in the Tetragonopterinae. There are recognized in this Memoir fifty-two genera, and three hundred and twelve species. Over half of these were described during the course of the prepa- ration of the Monograph. The subfamily Tetragonopterinae, at present the dominant group of tropi- cal American fishes, was defined by Giinther (loc. cit.) to include Piabucina, Alestes, Brachyalestes, Chirodon, Chalceus, Brycon, Chalcinopsis, Chalcinus, Gasteropelecus, Piabuca, and Agoniates, besides the genera listed below. Elimi- nating these genera, which are now relegated to various separate subfamilies 1 The portion of the Monograph dealing with the Cheirodontinae appeared recently in the Memoirs Carnegie Museum, 1916, 7, p. 1-99, pl. 1-17. SOURCES OF THE MATERIAL. ) or which (Alestes and Brachyalestes) are geographically extra-limital, there remain: — Tetragonopterus, with thirty-two species; Scissor, with one species; Pseudochalceus, with one species; Bryconops, with two species; Creagrutus, with one species; making a total of five genera, with thirty-seven species. Of the Iguanodectinae, which were included in Giinther’s Tetragonopterina, he recorded two species belonging to the genus Piabuca. He recorded four species of Corynopoma and placed them in his Erythrinina. They are the Stevardia albipinnis considered below in the discussion of the new subfamily, Glandulocaudinae. In the last general review of Giinther, there were a grand total of seven genera and forty-three species, as compared with the fifty-two genera and over three hundred species, known today. SOURCES OF THE MATERIAL. Tue NATHANIEL THAYER BRAZILIAN EXPEDITION. In January 1807, Karolina, Archduchess of Austria, was married to the Crown Prince of Brazil. In the retinue accompanying the young couple to Brazil went Johann Natterer of Wien. Natterer remained in Brazil for eighteen years and collected extensively from Rio de Janeiro to Cuyaba, thence down the Madeira to the Amazon and up the Rio Negro and Rio Branco. Taking advan- tage of the opportunity, the King of Bavaria sent two naturalists, Johann Bap- tist von Spix and Carl Friedrich von Martius with the bridal party. They also collected natural history specimens. The routes of travel of Natterer and of Spix and Martius are shown on Plate 1. The fishes collected by Natterer were described by Heckel, Kner, and Steindachner. Those collected by Spix and Martius were being described by Spix when he died. Louis Agassiz, then a student at the University of Munich, was selected to complete the work of Spix. Agassiz’s work resulted in a sumptu- ous folio, and a desire to personally inspect the fauna of Brazil, a desire fulfilled forty years later. In 1865, the generosity of Mr. Nathaniel Thayer made it possible for Agassiz to undertake his journey to Brazil. ._ The assistants of the Thayer Expedition were James Burkhardt, artist, J. G, Anthony, conchologist, C. F. Hartt, and Orestes St. John, geologists, J. A. Allen, ornithologist, and George Sceva, preparator. Besides these assistants 6 THE AMERICAN CHARACIDAE. several volunteers accompanied the Expedition: these were Edward Copeland, Newton Dexter, Walter Hunnewell, William James, Stephen Van Rensselaer Thayer, and Thomas Ward. Agassiz had the further assistance of the Brazilian Government through the Emperor, Dom Pedro II. Among Brazilians who joined the Expedition or aided in other ways were Major Coutinho, Messrs. Vinhas, Bourget, Talisman, Dr. Justa, Couto de Magalhaés, and others. The Expedition started late in March, 1865, and landed at Rio de Janeiro on April 22nd. Three months were spent about Rio. On July 25th Agassiz with Coutinho, Burkhardt, Bourget, Hunnewell, and James, joined later by Dexter and Thayer, went along the coast to Bahia (July 28th), Pernambuco (31st), Parahyba do Norte (August 2nd), Ceara (4th), Maranham (6th) and Para (11th). On August 20th the party started up the Amazon. It was divided in various ways. One or two of the assistants were left, or sent to some place to collect for longer or shorter periods to rejoin the main party at times. The itinerary up the Amazon was:— August 20,1865. Startupthe Amazon. 20. Breves. 2/. Tajapuru. 22. Gurupa. 23. Porto do Moz. 25. Montalegre. (Monte Alégre). 26. Santarem (Dexter, James, and Talisman, a young Brazilian start up the Tapajos from here; Bourget and Hunnewell remain at Santarem. They rejoin the Expedition at Manaos). September 1. On an arm of the Rio Ramos, connecting the Amazons, through the Mauhes, with the Madeira. 3. Return to steamer at Villa Bella. 4. Manaos. 10. Leave Manaos. 13. Coari. 14. Teffé. 15. Fonte Béa. 17. San Paolo. 18. Tabatinga. Bourget remains at Taba- tinga. 20. Left Tabatinga. James and Talisman remain at San Paolo to ascend the Iga. 20. Left Fonte Béa again. 2/. Teffé again — take up residence on Lake Teffé. Sitio. most remarkable catch in Forest Lake. October 17. James and Talisman returned from Ie¢a, and Jutahy. About Oct. 21, left Teffé. Verge of rainy season. Bourget returns from Tabatinga. 23. Reach Manaos. 27. Go to Lake Hyanuary. 29. Return to Manaos for a six weeks’ stay. December 6. Thayer returns from Lago Alexo, Bourget and Thayer from Cudajas, James from Manacapuru, Coutinho from Lake Hyanuary, José-Fernandez, Curupira, ete. 10. Dexter and Talisman returned from Rio Branco. Water too high. Leave for Mauhes on war vessel. 74. Mucaja-Tuba. 16. Mauhes. 20. Leave Mauhes. 25. Manaos. 27. Leave Manaos to ascend the Rio Negro. 29. Pedreira. 31. Manaos. January 16, 1866. Villa Bella and Mauhes again. 7/8. Lago Maximo. 20. Leave Villa Bella. 27. Obidos. 22. Santarem at the mouth of the Tapajos. 26. Monte Alégre. Trip to Serra Erere. 29. Leave Monte Alégre. About 30. Porto do Moz on the Xingu: Vinhas brings collections from above and below the cascades of the Xingu. February 1. Gurupa. 2. Tajapuri. 4. Reached Para. March 26. Leave Para for Rio de Janeiro. April 2. An inland excursion at Ceard. 6. Pacatuba and the region about it. 1/7. Ceara. 16. Leave Cearaé for Rio. July 2. Sail for home. During Agassiz’s trip up the Amazon with his immediate assistants, the rest of his party was divided to collect in the rivers of eastern Brazil from Rio de Janeiro to Para. “One object was kept constantly in view throughout this SOURCES OF THE MATERIAL. 7 Expedition,— namely, that of ascertaining how the freshwater fishes are dis- tributed throughout the great river systems of Brazil.” The routes of the separate parties are indicated on Plate 1. Hartt and Copeland collected in the lower courses of the rivers from the San Francisco to Rio de Janeiro. They left Rio de Janeiro on June 19. They visited the lower courses of the Parahyba collecting at Campos and San Fidelis and ascended the tributary Muriahy. From San Fidelis they crossed the divide to the Itabapuana and descended that stream to its mouth going thence via the Rio Itapemerim, Rio Novo,’ and Guarapary to Victoria. On a subsequent journey from Rio they ascended the Rio Doce for ninety miles to the first fall at Porto de Souza and collected also at Linhares in the Rio Doce and the lake and river Juparana. From Linhares they went to San Matheus and the Rio Mucury in which they collected at its mouth, Porto Alegre? and some distance inland at Santa Clara. From Santa Clara they went to Philadelphia collecting on the way in the Rio Urucu and thence to Calhoa and Minas Novas both on the Arassuahy. They separately descended the Rio Jequitinhonha, three hundred and sixty miles to the sea. They collected next at Cannavierias and ascended the Rio Pardo to its fall. Collections were also made at Belmonte, Santa Cruz and Porto Seguro on the coast south of the Rio Jequitinhonha. They then visited Bahia and returned to Rio de Janeiro.* St. John, Allen, Ward, and Sceva went from Rio de Janeiro to the Rio Parahyba and Juiz de Fora, across the Serra do Mantequeira to Barbacena. They collected at Rodeio, Rio Macacos, a tributary of the Santa Anna, State Rio de Janeiro. Ward left the party at Barbacena and went over Ouro Preto and Santa Barbara to Diamantina “passing from one river-basin to another in order to examine as many of the tributaries of the Rio Doce and the Jequitin- honha as possible.” Ward then crossed the San Francisco at Januaria and went to the Tocantins which he followed to its junction with the Amazon. St. J ohn, Ward, and Allen continued via Logoa Dourada and Pradas across the Rio Carandahy and Rio Paraopeba to Sabura (Rio Macacos, into Rio San Fran- cisco), Santa Luiza, Lagoa Santa, Sete Lagoas and Gequitiba. Sceva remained about Lagoa Santa and later went via Rio de Janeiro to Canto-Gallo. St. John and Allen went down the San Francisco to Januaria. Here ‘ Note also another Rio Novo in Haseman’s itinerary, p. 10. * Not the Porto Alegre, Rio Grande do Sul where von Ihering and Haseman collected. ‘Collections were made by Hartt in the Rio San Francisco below the fall either at this time or on a later trip. Hartt and Copeland sent in specimens recorded as from J acurpe and Posuca, localities I have not been able to locate. 8 THE AMERICAN CHARACIDAE. Allen, on account of his health, was compelled to leave the Expedition. St. John followed the San Francisco to Villa do Barra then went by land through the valley of the Rio Grande, a tributary of the San Francisco to Santa Rita, “thence to Mocambo and across the table-land separating the basin of the Rio San Francisco from that of the Rio Paranahyba.” He collected in the Basin of the Rio Paranahyba at Paranagua, Manga, San Gongallo, and Therezina. Near the latter place he collected in the Rio Poty or Puty a tributary of the Rio Paranahyba. From Therezina he crossed the divide to Caxias on the Itapicurt which empties into the ocean at Maranham.' He reached the latter place on January 8, 1866. The collection of fresh-water fishes made by Louis Agassiz and his assist- ants is the largest ever brought together by a single expedition. The hope of Agassiz to complete the work on the fishes was not realized, the very wealth of material making a comprehensive report almost herculean. Dr. Franz Steindachner, Hofrat and Intendant of the K. K. Naturhistorisches Hofmuseum, Wien, who has written extensively concerning the collections made by Natterer, obtained leave of absence from May 1870 until June 1873. During this time he accompanied Professor Agassiz on the Hassler Expedition, December 1871 to August 1872, and devoted more than two years to the study of the fishes secured during the Thayer Expedition which he considered “without any exaggeration the richest and most complete in the world.” The following papers are based wholly or in part on the Thayer Brazilian collection: Agassiz, Louis and Mrs. E.C. A journey in Brazil. Boston, 1868. E1genMANN, C. H. The evolution of the catfishes. Zoe, 1890, 1, p. 10-15. On the presence of an operculum in the Aspredinidae. Amer. nat., 1892, 26, joe (ll Steindachneria. Amer. nat., 1897; 31, p. 158-159. New genera of South American freshwater fishes, and new names for some old genera. Smith. mise. coll., 1903, 45, p. 144-148. Ergenmann, C.H.and R.S. A list of the American species of Gobiidae and Callionymidae, with notes on the specimens contained in the Museum of Comparative Zodlogy, at Cambridge, Massachusetts. Proc. Cal. acad. of sci., 1888, ser. 2, 1, p. 51-78. American Nematognathi. Amer. nat., 1888, 22, p. 647-649. Preliminary notes on South American Nematognathi, I. Proc. Cal. acad. sci., 1888, ser. 2, 1, p. 119-172. Preliminary descriptions of new species and genera of Characinidae. West Amer. sci., 1889, 6, p. 7-8. 1 There is another river Itapicuri emptying into the ocean between the bay of Bahia and the mouth of the San Francisco, in which Dr. Haseman made extensive collections, (p. 10). SOURCES OF THE MATERIAL. 9 Description of new nematognathoid fishes from Brazil. West Amer. sci., 1889, 6, p. 8-10. Preliminary notes on South American Nematognathi, II. Proc. Cal. acad. SCi., 1889, ser. 2, 2, p. 28-56. A review of the Erythrininae. Proc. Cal. acad. sci., 1889, ser. 2, 2, p. 100-116, pl. 1. A revision of the edentulous genera of Curimatinae. Ann. N. Y. acad. sci., 1889, 4, p. 409-440. A revision of the South American Nematognathi or eatfishes. Occ. papers, Cal. acad. sci., 1890, 1, p. 1-508, pl. 1. A catalogue of the freshwater fishes of South America. Proc. U. S. N. M., 1891, 14, p. 1-82. Garman, Samuet. On the species of the genus Chalcinus in the Museum of Comparative Zodlogy at Cambridge, Mass., U.S. A. Proc. Essex inst., 1890, 22, p. 1-7. On the species of Gasteropelecus. Proc. Essex inst., 1890, 22, p. 8-10. On the species of Cynopotamus. Proc. Essex inst., 1890, 22, p. 11-14. On the species of the genus Anostomus. Proc. Essex inst., 1890, 22, p. 15-23. On a genus and species of the Characines (Henochilus wheatlandii, gen. n. et sp. n.). Proc. Essex inst., 1891, 22, p. 49-52, pl. 1. The cyprinodonts. Mem. M. C. Z., 1895, 19, p- 1-180, pl. 1-12. Jorpan, D. S. and Ercenmann, C. H. A review of the Sciaenidae of America and Europe. Rept. U. S. comm. fish., 1889, 14, p. 343-452, pl. 1-4. A review of the genera and species of Serranidae found in the waters of America and Europe. Bull. U. S. fish comm., 1888, 8, p. 329-442. STEINDACHNER, Franz. Die siisswasserfische des siidéstlichen Brasiliens. Sitz. Akad. wis- sensch. Wien, 1875, 70, abth. I, p. 499-538, pl. 1-6. Separate, p. 1-40, pl. 1-6. Beitriige zur kenntniss der chromiden des Amazonenstromes. Sitz. Akad. wissensch. Wien, 1875, 71, abth. I, p. 61-137, pl. 1-8. Separate, p. 1-77, pl. 1-8. Ueber einige neue Brasilianische siluroiden aus der gruppe der doradinen. Sitz. Akad. wissensch. Wien, 1875, 71, abth. I, p. 138-151, pl. 1-4. Separate, p. 1-14, pl. 1-4. Die siisswasserfische des siidéstlichen, Brasiliens. II. Sitz. Akad. wissensch. Wien, 1875, 71, abth. I, p. 211-245, pl. 1-6. Separate, p. 1-35, pl. 1-6. Beitriige zur kenntniss der der characinen des Amazonenstromes. Sitz. Akad. wissensch. Wien, 1875, 72, abth. I, p. 6-24, pl. 1-2. Separate, p. 1-18, pl. 1-2. Ichthyologische beitriige. IV. Sitz. Akad. wissensch. Wien, 1876, 72, abth. I, p. 551-616, pl. 1-13. Separate, p. 1-65, pl. 1-13. Ichthyologische beitrige. V. Sitz. Akad. wissensch. Wien, 1876, 74, abth. I, p. 49-240, pl. 1-15. Separate, p. 1-190, pl. 1-15. Die siisswasserfische des siidéstlichen Brasiliens. III. Sitz. Akad. wissensch. Wien, 1876, 74, abth., I, p. 559-694, pl. 1-13. Separate, p. 1-186, pl. 1-13. Die siisswasserfische des siidéstlichen Brasiliens. IV. Sitz. Akad. wissensch. Wien, 1877, 76, abth. I, p. 217-230, pl. 1-2. Separate, p. 1-14, pl. 1-2. Uber einige neue und seltene fischarten, etc. Denk. Akad. wissensch. Wien, 1878, 41, p. 1-52, pl. 1-9. Beitriige zur kenntniss der flussfische Siidamerika’s, I. Denk. Akad. wissensch. Wien, 1879, 41, p. 151-172, pl. 1-3. Separate p. 1-24, pl. 1-4. Ichthyologische beitrige. VIII. Sitz. Akad. wissensch. Wien, 1880, 80, abth. I, p. 119-191, pl. 1-3. Separate, p. 1-73, pl. 1-3. 10 THE AMERICAN CHARACIDAE. Beitrige zur kenntniss der flussfische Siidamerika’s. II. Denk. Akad. wissensch. Wien, 1881, 43, p. 103-146, pl. 1-7. Separate p. 1-46. Beitriige zur kenntniss der flussfische Siidamerika’s. III. Denk. Akad. wis- sensch. Wien, 1881, 44, p. 1-18, pl. 1-5. Separate, p. 1-18, pl. 1-5. Beitriige zur kenntniss der flussfische Siidamerika’s. IV. Denk. Akad. wissen- sch. Wien, 1882, 46, p. 1-44, pl. 1-7. Separate, p. 1-44, pl. 1-7. Tue Carnecige Museum EXpebITIon To CENTRAL SoutTH AMREICA. A detailed account of this Expedition is given in volume 9 of the Annals of the Carnegie Museum. During this Expedition Mr. Haseman collected at the following localities :— November 6, 1907. Rio Coite, into the Rio Salitre, into Rio San Francisco. 6. Rio Aqua Branea, into Rio Itapicuri. 7. Rio Ipema, into Rio Itapicuri. Rio Lamaras, small creek into Rio Itapicuri. Rio Zinga, small creek into Rio Itapicuri. 8. Rio Itapicurti Grande, headwater of Rio Itapicuri. Rio Paiaia, into Rio Itapicuri. Rio de Jacobina, into Rio Itapicuri. 10. Lagoa Salgado — Rio Salitre into Rio San Francisco. 7. Bom Fim, Rio Amaratti, into Rio Itapicuri. 12. Saa Thome, Rio Salitre, into Rio San Francisco. 14. Rio Salitre, into Rio San Francisco. Baixa Grande, Rio Paqui into Rio Salitre, into Rio San Francisco. Rio Paqui, into Rio Salitre near Baixa Grande. 2/. Finca Amaratt, Rio Itapicurti. Creek on farm emptying into Rio Itapicuré Mirim. 27, 28. Joazeiro, Rio San Francisco. December 6. Barra, fork of Rio San Francisco and Rio Grande. December 12,18. Januaria, Rio San Francisco. 15. Cachoeira de Pirapora, Rio San Francisco. 23. Lagoa de Joao Pereira Barra, Rio San Francisco. 24. Lagoa de Porto, near Barra, Rio San Francisco. January 4, 1908. Lagoa Barreiras, Rio San Francisco. 6-9. Boqueirao, Rio Grande of Rio San Francisco Basin. 16. Lagoa Parnagua or Paranagua, Paranahyba Basin. 24. Santa Rita de Rio Preto, into Rio Grande, into Rio San Francisco. 27. Rio Preto, ten miles below fork of Rio Sapao. February 4. Cachoeira da Velha, Rio Novo, into Rio Somno, into Tocantins. 6. Stromé, Rio Somno. Headwaters. 11. Rio Sapao, into Rio Preto, into Rio San Francisco. Near Prazer. 15. Rio Preto, into Rio Grande, into Rio San Francisco. 23, 24. Barra, Rio San Francisco. March 2. Queimadas, Rio Itapi- eurt. 4. Alagoinhas, Rio Catu. 5. Rio Itapicurt, 12 miles from Timbo. 11, 13. About Bahia; 22, Penedo, mouth of Rio San Francisco. 30. Propria, Rio San Francisco. April 2. Penedo. 6. Maceio, on the coast. 7. Penedo. 10. Acaraju. 14. Cachoeiro, Rio Paraguassu. May 4. Sete Lagoas, into Rio das Velhas, into Rio San Francisco. 10, 11, 13. Creek, ponds, mountain rills near Rio das Velhas. 14. Miguel Burnier, headwaters of Rio das Velhas and tributary of Paraopeba. 19. Sao Joao del Rey, Rio das Mortes, into Rio Grande, into Rio Paranda. 2/. Sitio, Rio das Mortes. 22. Serraria, Rio Parahybuna, into Rio Parahyba. 24-28. Creeks, pools, river at Rio Doce. June 1-3. Entre Rios, Rio Para- hyba. 13-15. Campos, Rio Parahyba. 16. Lagoa Feia, south of mouth of Rio Parahyba. 18, 19. Rio Itapemerim and swamp near Munez Freire. 23, 24. Sao Joao da Barra, Rio Parahyba. July 5. Barra da Pirahy. 7, 8. Bom Jardin, above and below falls, Rio Grande, into Rio Parand. 9, 10. Santa Rita de Jacutinga, Rio Preto, into Rio Parahyba. 12, 13. Barra da Pirahy, Rio Parahyba. 14,15. Jacarehy, Rio Parahyba. 17,20. Mogy das Cruzes, Rio Tieté, into Rio Parand. 23. Piracicaba, Rio Tieté, into Rio Parana. 23. SOURCES OF THE MATERIAL. 11 Sapina, Rio Tieté, into Rio Parané. Santos, coast of Sa0 Paulo. 25. Alto da Serra, Rio Tieté, into Rio Parand. 26, 28. Mogy, into Santos Bay. Creek at base of mountains, ten miles from Santos. 29. Piassaguera, near Santos. 29. Santos. 31. Rio Pilao, fifteen miles southwest of Santos and Cubatao, Rio Cubatao. Clear, swift and rocky creeks seven miles west of Santos. August 7. Mogy Mirim, into Rio Mogy Guassu, into Rio Grande, into Rio Parané. Corrego de Joao de Deus. Twelve miles from Mogy Mirim. 14. Rio Paranahyba, into Rio Parané. 18, 19. Jaguara, Rio Grande, into Rio Parana. 25, 26. Mogy Guassu, Rio Mogy Guassu, into Rio Grande, into Rio Parana. September 1. Bebedouro, near Rio Grande, and Rio Parané. 4-9. Piracicaba, into Rio Tieté, into Rio Parand. Above and below big falls. 14. Salto de Avanhandava, Rio Tieté. 22. Salto das Cruzes, Rio Tieté. 27. Itapura, Rio Tieté; Riberao Azul, twelve miles from Tieté. October 8. Riberao Azul. 11. Miguel Calmon; twenty miles east of Miguel Calmon. Bauhru, Rio Tieté. Salto Grande de Paranapanema, into Rio Parana. November 27. Agua Quente, into Rio Ribeira da Iguapé. 28. Cavernas das Areas. Sixteen miles south- west of Iporanga in Serra do Mar. December 1. Iporanga, Rio Ribeira da Iguapé. 4, 8. Xiririca, Rio Ribeira da Iguapé. 15, 16. Iguapé, Rio Ribeira da Iguapé. 22. Serrinha Parana, Rio Iguassti, into Rio Parana. Rio das Mortes, into Rio Iguassti. Creek six miles west of Serrinha, with numerous falls. 27-29. Porto Uniao da Victoria, Rio Iguasst. January 2-4, 1909. Morretes, on Marunby, into Rio Nhundiaquara, into ocean at Pranagua. 17-24. Porto Alegre, Rio Grande do Sul, Rio Guahyba. 26-27. Cachoeira Rio Jacuhy, into Lago dos Patos at Rio Grande do Sul. 29. Santa Maria, Rio Vaccacahy- Mirim, into Rio Guahyba into Rio Jacuhy. 37. Cacequy, Rio Ibacuhy, into Rio Uruguay. February 1. Cacequy. 4. Uruguayana, Rio Uruguay. Rio Negro, Uruguay or Paso de los Torros, into Rio Uruguay. 17. Arroyo Miguelete, Montevideo. 25, 27. San Juan, Argentina. March 4. Rio Colorado, Argentina. Choel-choel, tributary of Rio Negro. 5. Rio Negro. 6. Muddy ponds twenty miles east of town of Colorado. 1/7. Buenos Aires, Rio de Prata. 23. Asuncion, Paraguay. 30. Cerro de Lambaré, five miles below Asuncion in saline swamp. 3/. Bays in front of and near Asuncion. April 2. Sapucay, Paraguay. In mountain rills. 5. Arroyo Pond near Sapucay. 7, 8. Arequa, Laguna Ipacary. 11, 13, 14. Villa Hays. 27. Urucum Mountains, 25 miles back of Corumba. 28. Corumba. May 2. Urucum Mountains. 6, 7. Puerto Suarez, 15 km. from Corumba. 9. Corumba. 23-27. Sao Luiz de Caceres, Matto Grosso. June 2,3. Cam- pos Alégre, Rio Jauru, into Rio Paraguay. San Matias, Bolivia, into Rio Paraguay. 10. Rio Sao Francisco, into Rio Paraguay. 12. Rio Santa Rita, into Rio Paraguay. 13. Rio Petas, Bolivia, into Rio Paraguay. 16. Rio Boa Ventura, into Rio Guaporé. 21, 27. Posada, into Rio Guaporé, about 40 miles south of Villa de Matto Grosso. 26, 28. Bastos, Rio Alegre, into Rio Guaporé. July 8. Below Rio Paragahu, in Rio Guaporé. 23. Sixty miles above San Antonio de Guaporé. July 29-August 13. San Antonio de Guaporé. August 28. Rio Machupo, Bolivia, into Rio Guaporé. Twenty miles below San Joaquin. September 4-7. San Joaquin, Bolivia. Rio Machupo. Lake one mile west of town and mud-hole near town. 14, 16. Berlin, Rio Mamoré. 19. Rio Mamoré, below mouth of Rio Guaporé. 28. Guaja-ussu, Rio Madeira. 30. Palo Grande, Rio Mamoré. In rapids under stones. October 5. Villa Bella, Bolivia, Rio Beni, into Rio Madeira. 13. Cachoele de Theotonio, Rio Madeira. Whirlpools. 17. Cachoele de Ribeirao, Rio Madeira. 26. Cachoele de Girao, Rio Madeira. Whirlpools. November 2,3. Sao Antonio de Rio Madeira. 15, 19, 25, 27-29. Manaos, mouth of Rio Negro. 30. Igarapé de Cachoeira Grande, two miles out of Manaos. December 2. Manaos. 4. Ten miles above Manaos on Rio Negro. 6, 8. Santarem, Rio Tapajos. 7. Swampy pools of Amazon and Rio Tapajos opposite Santarem. 9. Upper end of island, Amazon, four miles above Santarem. 10. Tapajos, in 12 THE AMERICAN CHARACIDAE. hollow logs — Amazon, one mile above Santarem. 1/7, 19. Igarapé de Jrura entering Rio Tapajos, two miles above Santarem. 12. Igarapé de Maica, four miles below Santarem. 14. Tapajos in front of Santarem. 15. Island in Amazon, three miles above Santarem. 20. Rio Tapajos at Santarem. 2/. Amazon. Half-way between Santarem and Para. 24. Para market. 27. From Gran Para between Belem and Salinas. 29. Braganea, Rio Caete. 16 kilometers from ocean, 162 from Para. January 1, 1910. Salt water, mouth of Rio Caete. 10. Aleobaca, Tocantins. Below first falls. 15-22. Para market. Tue GUIANA EXPEDITION. The joint Expedition of the Indiana University and the Carnegie Museum, led by myself, collected in British Guiana, between September 9 and December 1, 1908. A detailed account of the results of this Expedition is published as volume 5 of the Memoirs of the Carnegie Museum. The localities, enumer- ated from east to west are:— Maduni Stop-Off, Lama Stop-Off, Cane Grove Corner, the Georgetown Trenches, Morawhanna, Mora Passage, and Koriabo and Issorora Rubber Plantations, all in low tidal land; Christianburg, Wismar, and Malali, the latter about one hundred miles from its mouth, on the Demerara River and Bartica, Rockstone, Gluck Island, Crab Falls, Konawaruk, Warra- puta Cataract, and Packeoo Fall, all on the middle course of the Essequibo River. Characidae Grit, Mem. Nat. acad. sci., 1893, 6, p. 131. Proce. U.S. N. M., 1895, 18, p. 206. = Characinidae BoULENGER, Poissons bassin Congo, 1901, p.132. Cambridge natural history. Fishes 1904, p. 575. Cat. freshwater fishes Africa, 1909, 1, p. 174. = Characidae E1GENMANN, Rept. Princeton univ. exped. Patagonia, 1909, 3, p. 252; 1910, p. 420. = Characiformes Reaan, Ann. mag. nat. hist., 1911, ser. 8, 8, p. 15. > Characidae Reaan, Loc. cit. < Erythroides VALENCIENNES, Hist. nat. poissons, 1846, 19, p. 480. = Hrythrinidae Ricuarpson, Encyel. Brit., 1856, ed. 8, 12, p. 250. > Erythrinoidet BueeKer, Enum. sp. piscium Archipel. Indico, 1859, p. 31. > Erythrinidae Gru, Ann. Lye. nat. hist. N. Y., 1858, 6, p. 410. Mem. Nat. acad. sci., 1893, 6, p. 131. Proc. U.S. N. M., 1895, 18, p. 206. ; > Erythrinidae Corr, Proc. Amer. assoc. adv. sci., 1872, p. 333. > Erythrini Frraincer, Sitzungsb. Akad. wiss. Wien, 1873, 67, abth., p. 37. > Myletidae ApAMs, Man. nat. hist., 1854, p. 108. > Gastropelecidae Reaan, Ann. mag. nat. hist. 1911, ser. 8, 8, p. 19. > Xiphostomatidae Rraan, Loc. cit., p. 20. > Anastomidae REGAN, Loc. cit., p. 20. > Hemiodontidae Ruaan, Loc. cit., p. 21. > Citharinidae Reaan, Loc. cit., p. 21. ll ZOOLOGICAL Position. The Characins are a family of Ostariophysi which, with the other families of this superorder, are now at their prime. They are the dominant family of fresh-water fishes in Tropical America and they play a prominent réle in Africa. The Ostariophysi, which include most of the fresh-water fishes of the world, are distinguished from all other fishes by the peculiar arrangment of a series of 18 THE AMERICAN CHARACIDAE. ossicles for placing the air-bladder in communication with the auditory appa- ratus. The first four vertebrae are modified. The first vertebra lacks the superior arch which is replaced by the “‘claustrum” and ‘“‘scaphium”’ of the Weberian apparatus; the principal ossicle of the series, the ‘‘tripus,” is associated with the third vertebra consisting of the rib and parapophysis of the third vertebra. The ‘‘intercalarium,” representing the neural arch of the second vertebra, is imbedded in the ligament extending from the tripus to the secaphium. Very frequently the air-bladder comes in close contact with the skin, forming a pseudotympanum above the pectorals. In the Characins the area is not unfrequently marked by a humeral spot, a gathering of pigment cells from contiguous areas. This spot may become shifted away from the tympanum. The air-bladder is usually connected with the intestine by a duct. The pectoral girdle is suspended from the skull by a long posttemporal; the mesocoracoid is present and the ventral fins are abdominal. The orders and suborders of the Ostariophysi may be separated by the following key:— a. Maxillary bone usually a toothless vestige carrying a barbel; no subopercle or symplectic; no seales; supraoccipital and parietals codssified; mouth usually with teeth; ribs attached to trans- verse apophyses; skin naked or covered with bony plates. Usually an adipose dorsal. NEMATOGNATHI or SILUROIDEA. aa. Maxillary usually well developed, not forming the base of a barbel, but sometimes one or more small barbels at or near its end; subopercle and symplectic present; parietals distinct from the supra- occipital; thoracic vertebrae without parapophyses; ribs mostly sessile; naked or with scales. PLECTOSPONDYLI Or CYPRINOIDEA. b. Lower pharyngeals falciform, parallel with the gill-arches; jaws toothless; brain-case produced between the orbits; basis cranii simple; two superior pharyngeals; mouth without teeth, more or less protractile; no adipose fin...:.........:....... (EVENTOGNATHI or CYPRINIFORMES). bb. Lower pharyngeals not falciform; three basal branchihyals; basis cranii double, sometimes with myodome; one to four superior pharyngeals; mouth usually not protractile, usually with teeth. c. Anus submedian; body variously shaped, never eel-shaped; dorsal and usually an adipose fin present; ventralsvabdominally..../.124-' ere ee eeenie (HeTrRoGNaTHt or CHARACIFORMES). cc. Anus at throat; body eel-shaped; dorsal absent, or reduced; ventrals absent... (GyMNONOTI). The Eventognathi do not enter the Neotropical realm. The Gymnonoti are confined to it, the Heterognathi, as stated, occur in Africa and South America. The Nematognathi have an all but universal distribution. Sagemehl (Morphologische jahrbiich, 1884, 10) pointed out the similarity of the Erythrinoids to Amia, and suggested the derivation of the family from the Holostei and more particularly from the Cycloganoidea. Boulenger (Poisson bassin Congo, 1901) considers the Characidae, more particularly the Erythrins, THE CHARACIDAE. 19 as the primitive Ostariophysi from which the Cyprinidae (Gymnonoti) and Nematognathi have been derived. These families seem to him to represent three states in the evolution of Teleosteans, the Cyprinidae and Siluridae being derived apparently from a common ancestor, very near the Characins, these latter ranging themselves to the holostean Ganoids through the Erythrins. Later Boulenger (Cambridge natural history. Fishes, 1904, p. 575) refers to them as ‘‘a very generalized type, although perhaps not directly derived from the bony Ganoids.’”’ The relationship recognized by him he represented as follows: Eventognathi Nematognathi | Gymnonoti Heterognathi Rowntree (Trans. Linn. soc. Lond., 1903, ser. 2, 9, p. 78) concludes his extensive account of the visceral anatomy of Characins with the following words :—‘‘ With the possible exception of the indications of a cellular air-bladder, there appears to be nothing in the visceral anatomy of the Characinidae which strengthens the deductions made from the skull as to the Amioid affinities of the group. In opposition to such deductions are especially the cystoarian ovaries, the asymmetric ductus pneumaticus, the presence of pyloric appendages and the absence of all trace of a valvular conus and of an intestinal spiral valve.” In view of the observations of every naturalist who has studied the Char- acins, notably Boulenger with the African species, and the author with the American forms, it is doubtful whether the similarity of the Erythrins with the Ganoids is anything more than one of the innumerable radial adaptations this plastic family has undergone. RapIAL ADAPTATION. Boulenger’s remarks (Poissons bassin Congo, 1901, p. 132-135) concerning the African representatives may be translated as follows:— This is a very natural family whose internal organization shows a great affinity with the Cyprinidae but whose exterior appearance, due to adaptation to various modes of life, varies so much that the beginner in African ichthyology will mistake its various representa- tives as near the Salmonids, pikes, and roaches of Europe. It is also impossible to state in a 20 THE AMERICAN CHARACIDAE. few words the marks distinguishing the members of this family from such other fishes with abdominal ventrals and soft rayed fins as the Siluridae, Mormyridae, and Nototeridae. The presence of an adipose dorsal behind the rayed dorsal, although absent in some non-African genera, is a common character of the African fauna which it shares only with the Siluridae, which differ by their naked skin [one of the South American Characins is naked]. The Salmonids which also possess this adipose fin belong, on account of their general structure, to the same great group as the Clupeids which lack it and are strangers to Africa south of the Atlas. One of the prime characters used in separating the Characinidae from its nearest related families is the structure of the mouth described as bordered by the premaxillaries in the middle and the maxillaries on the sides, while in the Cyprinidae and Siluridae the upper jaw is said to be bordered by the premaxillaries only. While this definition is valid for the most of the representatives of this family we must not forget, as Segemehl has impressed on us, that there are numerous exceptions which destroy the diagnostic value of this character. Thus the maxillary is so reduced in Serrasalmo, Citharinus, Eugnatichthys, Phago, being confined to the angle of the mouth that it is proper to question if this bone really forms part of the border of the mouth and very recently I have been able to determine that it is absolutely excluded in Ichthyoborus and Neoborus. On the other hand, among Siluridae (Nematognathi), whose premaxillaries are greatly reduced the mouth is bordered laterally by the maxillaries. Chaca has a large maxillary bordering the mouth, Diplomystes and Eutropichthys have a toothed maxillary, and among the Cyprinidae Catostomus shows us the premaxillary and maxillary together forming the mouth border. There are otherwise very great differences in the structure of the jaws among the fishes which are justly united in one family, the Chara- cinidae. Thus the premaxillary, while never protractile, [I have since found Bivibranchia, a new genus, with protractile premaxillaries] is often vertically mobile (like the lower jaw), Ichthyoborinae; the mavillaries are either ankylosed to the premaxillaries or mobile on them, and in the majority of genera they do not directly articulate with the cranium, the only African exception being Sarcodaces. The character given by other authors to distinguish the Characins from the Cyprinids, i. e. the non prolongation of the brain capsule between the orbits in the former is not more constant, for in Citharinus and Xenocharax, for example, it extends forward to the nasal region. There are all degrees between this and the more usual one from which the objection- able definition is drawn. The orbitosphenoid, undivided and often membranaceous, forms a large interorbital septum in front of the cerebral cavity or below its anterior part. We have here an important character although it may not be constant, a specialization in this reduc- tion of the anterior part of the basis cranii and in its replacement by a thin interorbital parti- tion. The presence of teeth in the jaws does not distinguish the Characins from the Cyprinids because both African and American genera lack them. The teeth vary enormously in struc- ture and furnish valuable characters to distinguish numerous genera of this family. They are found on the premaxillary, the lower jaw, sometimes they are also present on the maxil- laries. They are but rarely found on the palate and none of the African genera offer any examples. The branchiostegal rays number only 3 to 5. There are four branchial arches. Pseudo- branchia are lacking or are rudimentary and glandular. The coracoid often forms a ventral ridge. The scapulars, epicoracoid, and postclavicle are distinct. The ventral rays number 10 to 13. The first pectoral, the dorsal, and anal rays are but rarely ossified and never form a formidable spine as in the Siluridae. Vertebrae in the African species vary from 17-33 +- 13-24 = 33-57. THE CHARACIDAE. 21 The scales, large or small, are ciliated in certain genera — a character very rare among the Physostomes. As Valenciennes says of Distichodus, the free part of the scales is truly ctenoid, the rest cycloid. The head always lacks scales, a rare thing in fishes. The lateral line organ is represented on the opercle by a branch from the suborbital. The air-bladder is always divided into two parts by a constriction. The anterior part is much smaller than the posterior. In some, Alestes, the air-bladder may be prolonged along the right side of the hemapophyses and interneurals as far as the posterior end of the anal. There are generally 10-25 coeca on the stomach and this number may be raised to 35 or 40 (Hydrocyon, Citharinus). The intestine in the carnivorous genera is short and makes but one loop; on the other hand, it is excessively long and with many windings in the herbivorous types. In looking over this review of the characters of the family of Characins it is seen that there is none that taken alone and allowing for the exceptions, justifies its separation from the Cyprinids. We are forced to content ourselves with a combination of characters, any one of which taken by itself is insufficient. These words apply almost verbatim to the American members of the family. The American Characins range from the border of the United States to some distance south of Buenos Aires. They form about one third of the entire South American fresh-water fauna and have diverged in adaptation to diverse food, diverse habitat, and diverse enemies to fill nearly every niche open to fishes. The ends of the three lines of adaptation to different food give us mud- eating forms, with long intestinal tract and no teeth,' flesh-eaters with shear- like teeth that are able to cut their way out of nets, attack large fishes, horses, and bathers, and conical-toothed forms with sharp, needle-like teeth and com- paratively huge fangs. Greater diversity could scarcely be imagined, and one is led to suspect that some of the forms are over-adapted. In their divergence in form they have reached almost every conceivable shape, and have approached or paralleled many members of the diverse families of North American fresh- water fishes. Our shads and fresh-water herrings have their counterparts in Elopomorphus, Potamorhina, and Psectrogaster, our salmons are paralleled by Salminus and Catabasis, and our minnows by Astyanax and its relatives. It takes but a slight flight of the imagination to detect the striking similarity of Luciocharax to our gar pikes; our mullets are duplicated by Prochilodus; our top minnows are mimicked by Nannostomus. Bivibranchia, a recent dis- covery, shows a close similarity to Albula, and even our festive darters are duplicated by members Characidium of this most remarkable family. This plasticity of the family in both America and Africa, and the apparent if not real duplication of forms in the two continents, is the more remarkable 1 The toothless forms are not represented in Africa where members of the Cyprinidae replace them. 22 THE AMERICAN CHARACIDAE. when we consider the very probable long separation of the African and Ameri- can sections of the family, and that the present forms in Africa and America have probably independently evolved in the two continents from a common rudiment. As von Ihering first pointed out for animals in general, and as I have more particularly shown for the fishes, there has probably been no inter- migration between the two continents since or previous to the early Tertiary. Before the Tertiary von Ihering has postulated an Archhelenic continent between Africa and South America from which the two continents probably derived the ancestors of the Characins among other elements of the present fauna. It is known that the Characins have inhabited South America since the early Tertiary at a time when the tropical part of the continents probably con- sisted of two islands, one occupying the Guianas, the other the plateau of eastern Brazil. From this small area and from the primitive group of Characins inhabiting it, the continent and family grew contemporaneously to their present proportions. They spread from these centers over the developing continent till they met unfavorable climatic conditions in the south, and high mountains or contestants in the north. They spread southward in decreasing numbers to and beyond the barren areas of Argentine to the edge of Patagonia, and they spread northward decreasing rapidly in numbers at Panama, only one of them reaching the United States, three southern Mexico, and three the Motagua River. In South America they have always flourished, their only competitors in the rivers of the growing continent being the indigenous fishes contemporaneously developing from similar small rudiments. They have never been hampered or affected by intrusive elements. Their territory has from the first been unap- proachable to foreign fresh-water fishes, and the marine species that have accli- mated themselves have not diverged to any extent from their marine relatives. The African section, on the other hand, came into competition with emi- grants from India, and, according to Boulenger, the affinity of the entire Afri- can fauna with Asia, ‘‘is much greater than with America which is emphasized by the fact that the genera in the two first named regions are identical, while America possesses genera very closely related but not identical with those of Africa.” The American Characins have diverged, as stated before, in the most amaz- ing manner in almost all possible ways towards diverse forms assumed by fishes. Some are long and slender, some nearly as deep as long, some spindle-shaped, others compressed. It is, however, in the teeth and alimentary canal that we find the greatest diversity, as has already been pointed out, and as will be THE CHARACIDAE. 23 described in detail. The fact that different authors have associated different members of the Characidae with the herring, trout, cyprinids, and poecilids indicates in a measure their versatility. LITERATURE. The earliest observations on Characins are recorded on some ancient monuments of Egypt. Heckel (Die fische Aegyptens chronologisch der zeitfolge ihrer ersten und spiteren wissenschaftlichen kenntnissnahme nach geordnet, p. 213 of Abbildungen und beschreibungen der fische Syriens, 1843) identifies Distichodus niloticus and Citharinus geoffroyi from mural decorations of graves near the Pyramids. Boulenger (Fishes of the Nile, 1907, p. 156) finds that Citharinus citharus Geoffroy is represented on the mural paintings of the tombs at Giza and Sapara, at Deir el Gebrawi, and on the tomb of Ti at Sakkara. The American Characins were first brought to the notice of naturalists by Marcgrav, who, in 1648 in his Historiae rerum naturalium Braziliae, 4, described the following :— Curimata, p. 156 = Prochilodus argenteus (Agassiz). Tareira do Rio, p. 157 = Hoplias malabaricus (Bloch). Piraya, p. 164 = Pygocentrus piraya (Cuvier). Maturaque, p. 169 = Hoplias malabaricus (Bloch). Piabucu, p. 170 = Piabucus dentatus (Koelreuter). Piaba, p. 170 = An Anostomus (?) Ten years later, in 1658, Piso followed this account in his Historiae naturalis et medicae Indiae Occidentalis libri quinque, which is but a second edition of the former work, with figures and descriptions of the same species :— Piabucu, p. 66, Piaba, p. 67, Maturaque, p. 67, Tareira II do Rio, p. 68, Piranha, p. 69, Curimata, p. 70. The detailed history of the African section of the family will not be given here.'| The number of known African species is far smaller than the number of American species which exceeds six hundred. In Seba’s Locupletissimi verum naturalium thesauri accurata descriptis, 1 Boulenger (Fishes of the Nile, 1907, p. 117) indicates that the Salmo niloticus Linné, Syst. nat., ed. 12 is not the Salmo niloticus of Hasselquist. The date, 1757, of Hasselquist, however, being pre- Linnean, the name Salmo niloticus must be applied to the species of Linné ed. 10. The S. niloticus of the Systema ed. 12, which is the same as that of ed. 10, is, according to Boulenger, identical with Myletes baremose Joannis, and the latter = Alestes baremose of Boulenger, p. 117, should stand as Myletes nilo- ticus. Further according to Boulenger, p. 141, the Salmo niloticus of Hasselquist is Salmo aegyptiacus of Gmelin. The name Salmo niloticus being a synonym of Myletes niloticus cannot be used for any other species and the oldest name after Hasselquist should be applied to his species. This oldest na me is Salmo aegyptiacus and since this is a Distichodus the species should be Distichodus aegyptiacus inste ad of Distichodus niloticus as given by Boulenger, p. 141, 24 THE AMERICAN CHARACIDAE. 1758, over the signature of Artedi, but according to Gill! by another author, the name Tetragonoptrus, coined by Klein and originally intended for entirely dif- ferent fishes, was applied to the species now known as Tetragonopterus argenteus. In 1754 and 1756 Gronovius in his Museum ichthyologicum defined the following genera: — Charax, 1754, p. 19. Gasteropelecus, 1756, p. 7, pl. 7, fig. 5. Anostomus, 1756, p. 13, pl. 7, fig. 2. Erythrimus, 1756, p. 6, pl. 7, fig. 6. The two species referred by Gronovius to Charax were subsequently in- corporated in the genera Charax (gibbosus) and Astyanax (fimaculatus). The Gasteropelecus is the Gasteropelecus sternicla of present authors. His Anos- tomus is Anostomus anostomus and Erythrinus is Erythrinus salmoneus. In 1777 Scopoli (Introductio ad historiam naturalem * * *) adopted these genera into the binomial system with Gronovius species as the types. Linné abandoned all of these genera and distributed the species known to him in the genera Clupea, Cyprinus, and Salmo. The species described or recognized by Linné in his tenth and twelfth editions and in Gmelin’s the thir- teenth edition of the Systema Naturae are given in the following table modified from that of Gill (Proc. U. S. N. M., 18, p. 213): 1758 1766 1788 Page No. Page No. Page No. Salmo argentinus * 2 511 12 1372 12 Piabucus dentatus gibbosus 311 19 513 20 1384 20 Charax gibbosus. notatus 513 21 1385 21 Astyanaz fasciatus bimaculatus 311 20 513 22 1385 22 Astyanax bimaculatus immaculatus 312 21 513 23 1385 23 (?) cyprinoides 514 25 1385 25 Curimatus cyprinoides niloticus 312 22 514 26 1386 26 Myletes niloticus aegy ptiacus 1386 49 Distichodus aegyptia- cus & rostratus pulverulentus 312 23 514 27 1386 27 Astyanax ? rhombeus 514 28 1386 28 Serrasalmo rhombeus anostomus 312 24 514 29 1387 29 Anostomus anostomus ?Clupea sima * 319 6 524 Ui ? ? ~ sternicla 319 7 524 3 1384 48 Gasteropelecus sternicla Cyprinus cephalus part? 322 7 527 7 1417 6 Erythrinus cephalus dentex 325 531 26 1383 47 Myletes dentex & bare- mose 1 Proc. U.S. N. M., 1895, 18, p. 225-227. * The generic names are those used in the tenth edition. § Clupa sima is credited to Asia by Linné. It is placed in the synonymy of sternicla by Bloch, p. 418, I know nothing further about it. THE CHARACIDAE. 25 The total number of species known to Linné is thus seen to have been from eight to ten in 1758 and from twelve to fourteen in 1766, to which but a single species had been added at the time of the publication of the 13th edition in 1788. Linné apparently knew none of the species described by Maregrayv except Marcgrav’s piabucu which is Salmo argentinus Linné. In 1794 Bloch in his Ausliindische fische added a number of species without, however, recognizing any distinct Characinid genera. In Schneider’s edition of Bloch’s Systema ichthyologiae published in 1801 the following species were recognized: — Synodus malabricus Bloch, p. 397. (tafel 392 of Bloch). Synodus erythrinus Bloch & Schneider, p. 397. Synodus tareira Bloch & Schneider, p- 398, pl. 79. Synodus palustris Bloch & Schneider, p. 398. Salmo friderici Bloch, p. 403. (tafel 378 of Bloch). Salmo fasciatus Bloch, p. 403. (tafel 379 of Bloch). Salmo argentinus Linné, p. 403. (tafel 382, fig. 1 of Bloch). Salmo rhombeus Linné, p. 404. (tafel 383 of Bloch). Salmo falcatus Bloch, p. 404. (tafel 385 of Bloch). Salmo odoe Bloch, p. 405. (tafel 386 of Bloch). Salmo pulverulentus Linné, p. 406. Salmo dentex Linné, p. 407. Salmo edentulus Bloch, p. 412. (tafel 380 of Bloch). Salmo melanurus Bloch, p. 412. (tafel 381, fig. 2 of Bloch). Salmo unimaculatus Bloch, p. 412. (tafel 381, fig. 3 of Bloch). Salmo bimaculatus Linné, p. 413. (tafel 382, fig. 2 of Bloch). Salmo anastomus Linné, p. 414. Salmo niloticus Linné, p. 414. Salmo cyprinoides Linné, p. 414. Salmo curimata Bloch & Schneider, p. 417. Salmo aegyptius-Gmelin, p. 418. (aegyptiacus of Gmelin). Salmo gasteropelecus Gmelin, p. 418. Salmo immaculatus Linné, p. 419. Salmo gibbosus Linné, p. 419. In 1802 Lacépéde (Histoire naturelle des poissons) created for Salmo rhom- beus Linné the genus Serrasalmo and adopted the Charax of Gronovius as Characinus for piabuca, dentex, gibbosus, notatus, bimaculatus, immaculatus, cyprinoides, niloticus, nefasch, pulverulentus, anostomus, friderici, fasciatus, melanurus, and odoe. The real advance in our knowledge of the relationships of the Characins did not begin until several years later when Cuvier (1817) published his Régne animal, and a series of articles in the Memoires Museum @histoire naturelle. In the Régne animal (p. 174) he recognized Erythrinus which he placed in his Clupes and the following genera which were included in 26 THE AMERICAN CHARACIDAE. the family of Salmones: — Characinus, Curimatus, Anostomus, Serrasalmo, Piabucus, Tetragonopterus, Myletes, Hydrocynus, Citharinus, and Gastero- pelecus.' In 1829, Memoires Museum d’histoire naturelle, 4, he further defined Chalceus and in 1819, 5, apparently substituted Hydrocyon for Hydrocynus. A great advance towards a knowledge of the South American Characinid fauna was made by Spix and Agassiz in the Selecta genera et species piscium Brasiliensis, 1829. They defined Prochilodus Agassiz (= Pacu Spix), Anodus Spix, Lepori- nus Spix, Schizodon Agassiz, Salminus Agassiz, Hiphorhynchus Agassiz (= Acestrorhynchus Eigenmann), Rhaphiodon Agassiz (= Cynodon Spix), and Xiphostoma; they also described many new species. Up to this time and for several years later the Characins were distributed among the Salmonids and Clupeids and the peculiar parallelism between some of the genera of these families and the genera of the Characidae made such an association seem natural. In 1842 Johannes Miiller in his treatise on the air- bladder of fishes (Monatsb. Acad. wiss., Berlin, June 1842 and Arch. anat. u. phys., 1842, p. 307) described the genera Macrodon (= Hoplias) and Hemiodus and united all of the Characins in his new family Characinidae. In 1844 Miller and Troschel published a synopsis of the known genera (Wiegmann’s archiv, 1844, 1, p. 81) and defined the new genera Chilodus, Distichodus, Alestes, Brycon, Exodon, Epicyrtus, Hydrolycus, Pygocentrus, Pygopristis, Catoprion, and Myleus. They followed this in 1845 by the first monograph on the Characinidae, Die familie der Characinen (Horae Ichthyo- logicae, 1, 2). Here all of the then known genera, including the new genus Agoniates are described and the known species enumerated. This work by Miller and Troschel was up to that time by far the most important as well as the most comprehensive work on the Characins. It is the first of three general accounts that have appeared. In it were recognized thirty-one genera and eighty-eight species. 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