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flDemotrs of tbe flDuseum of Comparative Zoologp.

AT HARVARD COLLEGE.

Vol. XLIII. Part 1. AdvrfttUttS V ^Kt^f ^.£ 9kf.

THE AMERICAN CHARACIDAE.

BY

CARL H. EIGENMANN.

WITH SIXTEEN PLATES.

CAMBRIDGE, U. S. A.:

Iprtnteo for tbe flDuseum.

August, 1917.

Ha

rtDemotra of tbe /IDuaeum of Comparative Zob'lofl?

AT HARVAED COLLEGE.
Vol. XLIII. Part 1.

THE AMERICAN CHARACIDAE.

BY

CARL H. EIGENMANN

WITH SIXTEEN PLATES.

^sotiuS ,„

CAMBRIDGE, U. S. A.:

Iprlnteo for tbe flDuaeum.

August, 1917.

THE AMERICAN CHAEACIDAF, 1

INTRODUCTION.

The revision of the American Characidae, based on the collections of the
Nathaniel Thayer Brazilian Expedition in the Museum of Comparative Zoology
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 Zoological 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, 1 Rhoadsinae, Glandulo-
caudinae, Iguanodectinae, Stethaprioninae, and Stichanodontinae. All of the
then known species of these subfamilies, except the species of Stevardia, were
included by Gunther (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 Gunther (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, pi. 1-17.

SOURCES OF THE MATERIAL. 5

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 Gunther'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 Slevardia albipinnis considered below in the discussion of the new
subfamily, Glandulocaudinae.

In the last general review of Gunther, 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.

The 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 Magalhaes, 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, 1 865. Start up the Amazon. 20. Breves. 21. Tajapuru. 22. Gurupa.
23. Porto do Moz. 25. Montalegre. (Monte Alegre). 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-
Teffe. 15. Fonte B6a. 17. San Paolo. IS. Tabatinga. Bourget remains at Taba-
tinga. 20. Left Tabatinga. James and Talisman remain at San Paolo to ascend the lea.
20. Left Fonte Boa again. 21. Teffe again — take up residence on Lake Teffe. Sitio.
most remarkable catch in Forest Lake. October 17. James and Talisman returned from lea,
and Jutahy. About Oct. 21, left Teffe. 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, Jose-Fernandez,
Curupira, etc. 10. Dexter and Talisman returned from Rio Branco. Water too high.
Leave for Mauhes on war vessel. 14- 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. 18. Lago Maximo. 20. Leave Villa
Bella. 21. Obidos. 22. Santarem at the mouth of the Tapajos. 26. Monte Alegre.
Trip to Serra Erere. 29. Leave Monte Alegre. About 30. Porto do Moz on the Xingu:
Vinhas brings collections from above and below the cascades of the Xingu. February 1.
Gurupa. 2. Tajapuru. 4- Reached Para. March 26. Leave Para for Rio de Janeiro.
April 2. An inland excursion at Ceara. 6. Pacatuba and the region about it. 11.
Ceara. 16. Leave Ceara 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, 1 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 2 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. 3

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. John,
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

1 Note also another Rio Novo in Haseman's itinerary, p. 10.

2 Not the Porto Alegre, Rio Grande do Su] where von Ihering and Haseman collected.

3 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 Jacurpe and Posuca, localities 1
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 Goncallo, 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 Itapicuru
which empties into the ocean at Maranham. 1 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.
Eigenmann, 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,

p. 71.

Steindachneria. Amer. nat., 1S97," 31, p. 158-159.

New genera of South American freshwater fishes, and new names for some old

genera. Smith, misc. coll., 1903, 45, p. 144-148.
Eigenmann, 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 Zoology,

at Cambridge, Massachusetts. Proc. Cal. acad. of sci., 1888, ser. 2, 1, p. 51-7S.

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 Itapicuru 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,
pi. 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 catfishes. Occ. papers,
Cal. acad. sci., 1890, 1, p. 1-508, pi. 1.

A catalogue of the freshwater fishes of South America. Proc. U. S. N M
1891, 14, p. 1-82.
Garman, Samuel. On the species of the genus Chalcinus in the Museum of Comparative

Zoology 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, pi. 1.

The cyprinodonts. Mem. M. C. Z., 1895, 19, p. 1-180, pi. 1-12.
Jordan, D. S. and Eigenmann, C. H. A review of the Sciaenidae of America and Europe.

Rept. U. S. coram, fish., 1889, 14, p. 343-152, pi. 1-1.

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. 329^42.
Steindachner, Franz. Die siisswasserfische des sudostlichen Brasiliens. Sitz. Akad. wis-
sensch. Wien, 1875, 70, abth. I, p. 499-538, pi. 1-6. Separate, p. 1-10, pi. 1-6.

Beitrage zur kenntniss der chromiden des Amazonenstromes. Sitz. Akad.

wissensch. Wien, 1875, 71, abth. I, p. 61-137, pi. 1-8. Separate, p. 1-77, pi. 1-8.

Ueber einige neue Brasilianische siluroiden aus der gruppe der doradinen.

Sitz. Akad. wissensch. Wien, 1875, 71, abth. I, p. 138-151, pi. 1-4. Separate,

p. 1-14, pi. 1-4.

Die siisswasserfische des sudostlichen, Brasiliens. II. Sitz. Akad. wissensch.

Wien, 1875, 71, abth. I, p. 211-245, pi. 1-6. Separate, p. 1-35, pi. 1-6.

Beitrage zur kenntniss der der characinen des Amazonenstromes. Sitz. Akad.

wissensch. Wien, 1875, 72, abth. I, p. 6-24, pi. 1-2. Separate, p. 1-18, pi. 1-2.

Ichthyologische beitrage. IV. Sitz. Akad. wissensch. Wien, 1876, 72, abth.

I, p. 551-616, pi. 1-13. Separate, p. 1-65, pi. 1-13.

Ichthyologische beitrage". V. Sitz. Akad. wissensch. Wien, 1876, 74, abth.

I, p. 49-240, pi. 1-15. Separate, p. 1-190, pi. 1-15.

Die siisswasserfische des sudostlichen Brasiliens. III. Sitz. Akad. wissensch.

Wien, 1876, 74, abth., I, p. 559-694, pi. 1-13. Separate, p. 1-136, pi. 1-13.

Die siisswasserfische des sudostlichen Brasiliens. IV. Sitz. Akad. wissensch.

Wien, 1877, 76, abth. I, p. 217-230, pi. 1-2. Separate, p. 1-14, pi. 1-2.

tJber einige neue und seltene fischarten, etc. Denk. Akad. wissensch. Wien,

1878, 41, p. 1-52, pi. 1-9.

Beitrage zur kenntniss der flussfische Siidamerika's, I. Denk. Akad. wissensch.

Wien, 1879, 41, p. 151-172, pi. 1-3. Separate p. 1-24, pi. 1-4.

Ichthyologische beitrage. VIII. Sitz. Akad. wissensch. Wien, 1880, 80,

abth. I, p. 119-191, pi. 1-3. Separate, p. 1-73, pi. 1-3.

10 THE AMERICAN CHARACIDAE.

Beitrage zur kenntniss der flussfische Siidamerika's. II. Denk. Akad. wissensch.
Wien, 1881, 43, p. 103-146, pi. 1-7. Separate p. 1-46.

Beitrage zur kenntniss der flussfische Siidamerika's. III. Denk. Akad. wis-
sensch. Wien, 1881, 44, p. 1-18, pi. 1-5. Separate, p. 1-18, pi. 1-5.
Beitrage zur kenntniss der flussfische Siidamerika's. IV. Denk. Akad. wissen-
sch. Wien, 18S2, 46, p. 1-44, pi. 1-7. Separate, p. 1-44, pi. 1-7.

The Carnegie Museum Expedition to Central South 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 Branca, into Rio Itapicurii. 7. Rio Ipema, into Rio Itapicurii. Rio Lamaras, small
creek into Rio Itapicurii. Rio Zinga, small creek into Rio Itapicurii. 8. Rio Itapicurii
Grande, headwater of Rio Itapicurii. Rio Paiaia, into Rio Itapicurii. Rio de Jacohina, into
Rio Itapicurii. 10. Lagoa Salgado — Rio Salitre into Rio San Francisco. 11. Bom Fim,
Rio Amaratii, into Rio Itapicurii. 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. 21. Finca Amaratii,
Rio Itapicurii. Creek on farm emptying into Rio Itapicurii 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. Strome, 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-
curii. 4- Alagoinhas, Rio Catu. 5. Rio Itapicurii, 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. Silo Joao del Rey, Rio das Mortes, into Rio
Grande, into Rio Parana. 21 . 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.
IS, 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 Parana. 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 Tiete, into Rio Parana. 23. Piracicaba, Rio Tiete, into Rio Parana. 23.

SOURCES OF THE MATERIAL. 1 1

Sapina, Rio Tiete, into Rio Parana. Santos, coast of Sao Paulo. 25. Alto da Serra, Rio
Tiete, into Rio Parana. 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 Parana. Corrego de Joao de Deus. Twelve miles from Mogy Mirim.
14- Rio Paranahyba, into Rio Parana. IS, 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 Parana. 5-9. Piracicaba, into Rio Tiete", into Rio
Parana. Above and below big falls. 14- Salto de Avanhandava, Rio Tiete. 22. Salto
das Cruzes, Rio Tiete". 27. Itapura, Rio Tiete; Riberao Azul, twelve miles from Tiete.
October 8. Riberao Azul. 11. Miguel Calmon; twenty miles east of Miguel Calmon.
Bauhru, Rio Tiete". Salto Grande de Paranapanema, into Rio Parana. November 27.
Agua Quente, into Rio Ribeira da Iguape. 28. Cavernas das Areas. Sixteen miles south-
west of Iporanga in Serra do Mar. December 1. Iporanga, Rio Ribeira da Iguape. 5, 8.
Xiririca, Rio Ribeira da Iguape. 15, 16. Iguape, Rio Ribeira da Iguape. 22. Serrinha
Parana, Rio Iguassu, into Rio Parana. Rio das Mortes, into Rio Iguassii. Creek six miles
west of Serrinha, with numerous falls. 27-29. Porto Uniao da Victoria, Rio Iguassu.

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. 31 . Cacequy, Rio Ibacuhy, into Rio Uruguay.
February 1. Cacequy. 5. 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. 11. Buenos
Aires, Rio de Prata. 23. Asuncion, Paraguay. SO. Cerro de Lambare, five miles below
Asuncion in saline swamp. 31. Bays in front of and near Asuncion. April 2. Sapucay,
Paraguay. In mountain rills. 5. Arroyo Pona 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 Alegre, 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 Guapore". 21, 27.
Posada, into Rio Guapore, about 40 miles south of Villa de Matto Grosso. 26, 28. Bastos,
Rio Alegre, into Rio Guapore. July 8. Below Rio Paragahu, in Rio Guapore. 23. Sixty
miles above San Antonio de Guapore. July 29-August 13. San Antonio de Guapore. August
28. Rio Machupo, Bolivia, into Rio Guapore. 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, 15. Berlin, Rio Mamore. 19. Rio Mamore, below mouth of Rio
Guapore. 28. Guaja-ussu, Rio Madeira. 30. Palo Grande, Rio Mamore. 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. Igarape 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. //, 19. Igarape de Jrura entering Rio
Tapajos, two miles above Santarem. 12. Igarape 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. 21. Amazon. Half-way between Santarem and Para.
24. Para market. 27. From Gran Para between Belem and Salinas. 29. Braganca,
Rio Caete. 16 kilometers from ocean, 162 from Para. January 1, 1910. Salt water, mouth
of Rio Caete. 10. Alcobaca, Tocantins. Below first falls. 15-22. Para market.

The 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 6 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. A special effort was made to get a complete series from the Potaro
River both above and below the seven hundred and forty-one foot Kaieteur
Fall. Collections were made between October 6 and November 4, 1908, at
Aruataima, Holmia, and Savannah Landing above the Kaieteur and at Shrimp
Creek, Tukeit, Waratuk, Amatuk, Erukin, Kangaruma, Potaro Landing, and
Tumatumari below the Kaieteur. Mr. William Grant my Indian guide sent
in additional collections from the Rupununi and the Ireng Rivers. The first
series of specimens of this Expedition is in the Carnegie Museum, the second
series in Indiana University. Other series are in the Museum of Comparative
Zoology, the U. S. National Museum, the Field Museum, Stanford University,
the British Museum, the Museums at Amsterdam, Berlin, Vienna, and George-
town, British Guiana.

The Gimbel Expedition.

Through the generosity of Mr. Jake Gimbel of Vincennes, Indiana, Dr.
Max Mapes Ellis and Dr. William M. Tucker were enabled to go to British
Guiana primarily to gather material for a monograph of the Gymnotidae. 1

1 The gymnotid eels of Tropical America. Memoirs Carnegie museum, 1913, 6, p. 109-195, pi.
15-23.

SOURCES OF THE MATERIAL. 13

They started from New York in August 1910 and went direct to Georgetown.
After collecting in the Demerara and along the coast at Georgetown and con-
ducting experiments in the regeneration on various species of the Gymnotidae
they ascended the Demerara to canal Number Two. They went through the
canal to a tributary of Hubabu Creek, descended to the Demerara and returned
to Georgetown. They also ascended the Demerara to Wismar and crossed over
to the Essequibo at Rockstone. After collecting on Gluck Island in the Esse-
quibo and in a tributary of the Essequibo at the railway crossing between
Rockstone and Wismar they returned to the coast.

The Colombian Reconnaissance.

Preliminary notes of my reconnaissance in Colombia have been published
in Indiana University Studies 16 and 18. A detailed account will appear in
the Memoirs of the Carnegie Museum. I entered Colombia at Cartagena in
December 1911, and left from the same port in April 1912. From Cartagena
I went to Soplaviento on the Dique and to Calamar on the Magdalena; from
Calamar up the Magdalena River by steamer to La Dorado, collecting at various
stopping places, Barbosa, El Blanco, Canaletal, Puerto Wilches, Penas Blancas,
Puerto Berrio. From La Dorado, the route was by rail to the upper part of
the Magdalena, collections being made on the way at Honda, especially in Bernal
Creek. The upper part of the Magdalena was followed to Girardot, where exten-
sive collections were made. From Girardot, the route led first over the western
rim of the plain at an elevation of about 8800 feet to Bogota, on an elevated
plain among the eastern Cordilleras; collections were made on the plain near
Chapinero, north of Bogota, and at Madrid, near the western margin of the plains
of Bogota. A return was made to Girardot, from which via Chicoral to Quatro
Esquinas, Ibague\ Toche, across the Quindio Pass of the central Cordilleras, at
an elevation of 11,200 feet, to Boquia, Piedra Moler, and Cartago near the
Cauca River. Up the Cauca Valley via Paila, Buga La Grande, Buga to Cali,
collections being made at Paila and at Cali, and in the Cauca near Cab.

After collecting at Caldas (elevation of 3722 feet), the valley of the Dagua
was descended, collections being made at Cisnero (1046 feet), at Cordova (120
feet), and in tide- water.

From Buenaventura, on the Pacific coast of Colombia, a steamer was
taken up the San Juan River to Puerto Negria; thence a dugout carried the
Expedition as far as Istmina; collections were made in both the latter places.
From Istmina, after a ride of two hours up a little stream, and across the low

14 THE AMERICAN CHARACIDAE.

continental divide (elevation 300 feet above sea-level) the valley of the Atrato
was entered near Tambo. By dugout to the settlement of Boca de Raspadura;
thence the Raspadura River was followed into the Quito River, then the Quito
River. Collections were made at Boca de Certegui and near the town of Quibdo,
at the junction of the Quito River with the Atrato. From Quibdo, a steamer
was taken to Rio Sucio, where additional collections were made. From Sucio,
a steamer carried the Expedition back to the starting point at Cartagena.

These collections have been supplemented since my return by material
collected by Manuel Gonzales, near Puerto Wilches, at Las Juntas on the Rio
Bogota, in the province of Santander, and along the way from Bogota to Villa-
vicencio. The first series of the specimens and the duplicates are in the Carnegie
Museum, the second series in Indiana University.

The Landon-Fisher Expedition to Colombia.

A second Expedition into Colombia was made possible by Mr. Hugh McK.
Landon and Mr. Carl G. Fisher, of Indianapolis.

Mr. Arthur W. Henn and Mr. Charles Wilson, undergraduates in Indiana
University, left in December, 1912. They landed at Tumaco, near the south-
western corner of Colombia. After devoting about a month's time to the
Telembi River, a tributary of the Patia, they separated. Mr. Wilson went to
the San Juan River, collecting in the Upper San Juan Basin, the Condoto River
at Condoto, and in the San Juan River at Istmina and Tado of the Pacific side,
and later on the Atlantic slope at Tambo, Raspadura, Boco de Raspadura,
Managru, Quibdo, in the Atrato between Quibdo and Rio Sucio, and especially
in the Truando River emptying into the Atrato near Rio Sucio.

The Landon Expedition to Colombia and Ecuador.

Through the continued liberality of Mr. Hugh McK. Landon of Indianapolis,
Arthur W. Henn was able to remain in South America and spend the time between
February 15, 1913 and March, 1914 in collecting fresh-water fishes in Colombia
and Ecuador.

He sailed from Tumaco, Colombia for Barbacoas on February 15th. From
Barbacoas he went by packtrain to Tuquerres (10,090 feet), Ancuya (5000 feet),
Los Llanos de Sandona (5000 feet), Tambo, Pefiol, and to Guayabillo on the
brink of the Canon of the upper Patia. Descending to the Patia River, about

SOURCES OF THE MATERIAL. 15

3000 feet collections were made above the mouth of the Guaitara. He returned
via Pasto to Tuquerres and Barbacoas. He next descended the Rio Telembi
to its mouth and ascended the Rio Patia and the Rio Magui, the first large
tributary of the Patia above the Telembi to the village Payan.

Returning to Tumaco on the coast he went via Buenaventura to Puerto
Negria, the head of steam navigation of the San Juan River. Drifting down
the San Juan to the Rio Calima, the last large tributary of the San Juan from
the east, he ascended the Calima, collecting in a small creek near its mouth,
and at Boca de Guineo about thirty miles from the mouth of the Calima. After
ascending the Rio El Guineo to a portage, he crossed over to the San Joaquin
and descended it to Buenaventura.

He next went to Guayaquil from where several shorter excursions were
made. The first of these lead to Naranjito. Collections were made south of
Naranjito in a small creek Estero Verdes, a tributary of the Rio Chan Chan,
and in a deep river, the Rio Barranca Alta. The second shorter trip from
Guayaquil was to the small rivers at Chone and Portoviejo. The third trip
took him to Daule, Santa Lucia, and Colimes on the Rio Daule. Returning
to Daule he crossed over along a winding cut-off from the Rio Palenque to Vinces.

Returning to Guayaquil from Vinces, he went by rail to Quito, collecting
en route at Huigra (4000 feet), Rio Bamba (9020 feet), Latacunga (9055 feet)
and Quito (9375 feet). A short trip was made from Quito to Mindo (4108 feet).
Later he went to El Angel (10,000 feet) and down the Rio Chota or Mira to
Maria Luisa, the property of Sn. C6sar Mena. Collections were also made in
the Rio Chota at Guallupi (5,000 feet). He returned to Quito and later went
via Angel by packtrain to Barbacoas and steamer to Tumaco.

By the various expeditions outlined above, collections have been obtained
from most of the Pacific slope streams between Panama and Peru.

The first series of the specimens collected in the Landon Expedition is in
Indiana University. Other series are in the Carnegie Museum, the Museum
of Comparative Zoology, the U. S. National Museum, the Field Museum, and
the Museum of Stanford University.

16 THE AMERICAN CHARACIDAE.

Central American Expeditions.

From time to time specimens have been secured by exchange with the Field
Museum of Chicago. These had been collected by the late Dr. S. E. Meek in
various parts of Mexico and Central America. Accounts of this material have
appeared in various publications of the Field Museum.

In collaboration with Mr. S. F. Hildebrand, Dr. Meek was engaged at the
time of his death on a report of the fishes of Panama. Mr. Hildebrand has com-
pleted the report in the Laboratory of Indiana University and has given me the
opportunity to examine many of the specimens collected in Panama. Dr.
Meek as the representative of the Field Museum, and Mr. Hildebrand, Director
of the Beaufort Laboratory of the Bureau of Fisheries, as the representative of
the Smithsonian Institution, spent two seasons in Panama to collect fishes.
A report on their itinerary and on their collections has been published by the
Field Museum (Field museum publication 191, zool. ser., 10, 1916, p. 217-374,
pi. 6-32). They collected in various localities of the east-slope Chagres Basin,
many of which are now covered by Gatun Lake, and in others of the Pacific
slope Grande basin of the Canal Zone. In addition they collected in the basins
of the Rio Chepo, and Tuyra of the Pacific slope, south of the Canal Zone, and
in a few of the smaller rivers both north and south of the Canal Zone on both
the east and west slopes.

The Expedition to Guatemala.

During January, February, and March of 1915, Mr. Newton Miller assisted
by Messrs. E. B. Williamson, C. C. Deam, and Prof. J. Hines collected fishes at
Tenedores, Los Amates, Algeria, Gualan, Zacapa, and El Rancho in the Motagua
Basin of Guatemala. Additional collections were secured at Puerto Barrios
and Santa Lucia. The first series of specimen is in Indiana University.

THE CHARACIDAE. 17

THE CHARACIDAE.

<Dermopteres Dumeril, Zool. analytique, 180(3, p. 146.

<Salmonidi Rafinesqite, Indice littiol. Siciliana, 1810, p. 32.

<Dcrmopteria Rafinesque, Analyse nature, 1815, p. 87.

= Charadni Muller, Archiv. nat., 1813, 1, p. 323.

= Characinidae Muller, Monatsb. Acad. wiss. Berlin 1842.

= Characins (Characidae) Agassiz, Rept. Brit, assos. adv. sci., 1814, p. 293.

X Characins Valenciennes, Hist. nat. poissons, 1848, 21, p. 159.

< Characina Vogt, Zool. briefe, 1851, 2, p. 150.

= Characinidae Richardson, Encycl. Brit., 1856, ed. 8, 12, p. 245.

X Characinoidei Bleeker, Enura. sp. piscium Arohip. Indico, 1859, p. 31.

= Characinidae Gunther, Cat. fishes Brit, mus., 1861, 5, p. 278.

= Characinidae Cope, Proc. Amer. assoc. adv. sci., 1872, p. 333.

= Characinidae Gill, Arrang. fam. fishes, 1872, p. 16.

= Citharini Fitzinger, Sitzungsb. Akad. wiss., Wien, 1878, 77, p. 37.

= Characinida Schmarda, Zool., 1878, 2, p. 377.

= Characinidae Jordan and Gilbert, Synop. fishes N. Amer., 1882, p. 254.

> Characidae Gill, Mem. Nat. acad. sci., 1893, 6, p. 131. Proc. 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 Eigenmann, Rept. Princeton univ. exped. Patagonia, 1909, 3, p. 252; 1910, p. 420.
= Characiformes Regan, Ann. mag. nat. hist., 1911, ser. 8, 8, p. 15.

> Characidae Regan, hoc. cit.

< Erythroiiles Valenciennes, Hist. nat. poissons, 1S46, 19, p. 480.
= Erythrinidae Richardson, Encycl. Brit., 1856, ed. 8, 12, p. 250.

> Erythrinoidei Bleeker, Enum. sp. piscium Archipel. Indico, 1859, p. 31.

> Erythrinidae Gill, 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 Cope, Proc. Amer. assoc. adv. sci., 1872, p. 333.

> Erylhrini Fitzinger, Sitzungsb. Akad. wiss. Wien, 1873, 67, abth., p. 37.

> Myletidae Adams, Man. nat. hist., 1854, p. 108.

> Gastropelecidae Regan, Ann. mag. nat. hist. 1911, ser. 8, 8, p. 19.

> Xiphostomatidae Regan, hoc. cit., p. 20.

> Anastomidae Regan, hoc. cit., p. 20.

> Hemiodontidae Regan, hoc. cit., p. 21.

> Citharinidae Regan, hoc. cit., p. 21 .

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 role 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 " inter calarium," representing the neural arch of the second
vertebra, is imbedded in the ligament extending from the tripus to the scaphium.
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 sympleetic; no
scales; supraoccipital and parietals coossified; 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 syrnplectie 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 Cyprinipormes).

66. 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.
f. Anus submedian; body variously shaped, never eel-shaped; dorsal and usually an adipose fin

present; ventrals abdominal (Heterogxathi or Characiformes).

cc. Anus at throat; body eel-shaped; dorsal absent, or reduced; ventrals absent.. . (Gymxoxoti).

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 jahrbuch, 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.

Radial 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 maxillaries 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 CHAR ACID AE. 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, 1 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 spateren wissenschaftlichen kenntnissnahme nach
geordnet, p. 213 of Abbildungen und beschreibungen der fische Syriens, 1843)
identifies Dislichodus nilolicus 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 firs.t brought to the notice of naturalists by
Marcgrav, who, in 1648 in his Historiae rerum naturalium Braziliae, 4, described
the following : —

Curimata, p. 156 = ProchUodus 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 dcntatus (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. 1 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 nilolicus Linn6, Syst. nat.,
ed. 12 is not the Salmo niloticus of Hasselquist. The date, 1757, of Hasselquist, however, being pre-
Linnean, the name Salmo nilolicus must be applied to the species of LinnS ed. 10. The S. nilolicus of
the Systema ed. 12, which is the same as that of ed. 10, is, according to Boulenger, identical with Mylelcs
baremose Joannis, and the latter = Alestcs baremose of Boulenger, p. 117, should stand as Mylelcs nilo-
licus. Further according to Boulenger, p. 141, the Salmo nilolicus of Hasselquist is Salmo aegyptiacus
of Gmelin. The name Salmo nilolicus being a synonym of Mylelcs niloticus cannot be used for any
other species and the oldest name after Hasselquist should be applied to his species. This oldest name
is Salmo aegyptiacus and since this is a Distichodus the species should be Dislichodus aegyptiacus instc ad
of Distichodus niloticus as given by Boulenger, p. 14 1 .

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, pi. 7, fig. 5.
Anostomus, 1756, p. 13, pi. 7, fig. 2.
Erythrinius, 1756, p. 6, pi. 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.

Linne 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 Linne 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) :

]

[758

176C

1788

Page

No.

Page

No.

Page

No.

Salmo argenbvwus 2

2

511

12

1372

12

Piabucus dentatus

gibbosus

311

19

513

20

1384

20

Charax gibbosus.

notatus

513

21

1385

21

Astyanax fasciatus

bimacvlatus

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 n iloticus

aegyptiacus

1386

49

Distichodus aegyptia-
cus & rostratus

pulverulcntus

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 3

319

6

524

7

?

?

sternicla

319

7

524

3

1384

48

Gasteropelecus stern tela

Cyprinus ccphalus part'

• 322

7

527

7

1417

(i

Erythrinus ccphalus

dentex

325

531

26

1383

47

Myletes dentex & hare-

1 Proc. TJ. S. N. M., 1895, 18, p. 225-227.

2 The generic names are those used in the tenth edition.

3 Clupa sima is credited to Asia by Linn6. It is placed in the synonymy of slei -nick by ISloch, p. 418.
I know nothing further about it.

THE CHARACIDAE. 25

The total number of species known to Linne 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. Linne" apparently knew none of the species described by Marcgrav except
Marcgrav's piabucu which is Salmo argentinus Linne\

In 1794 Bloch in his Auslandische 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: —

Synodw malabricus Bloch, p. 397. (tafel 392 of Bloch).

Synodvs erythrinus Bloch & Schneider, p. 397.

Synodus tareira Bloch & Schneider, p. 398, pi. 79.

Synod™ palustris Bloch & Schneider, p. 398.

Salmo frideriei Bloch, p. 403. (tafel 378 of Bloch).

Salmo fasciatus Bloch, p. 403. (tafel 379 of Bloch).

Salmo argentinus Linne, p. 403. (tafel 382, fig. 1 of Bloch).

Salmo rhombeus Linne, 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 pulverulcntm Linne, p. 406.

Salmo dentex Linne, p. 407.

Salmo edentulus Bloch, p. 412. (tafel 380 of Bloch).

Salmo melanurus Bloch, p. 412. (tafel 381, fig. 2 of Bloch).

Salmo unimacidatus Bloch, p. 412. (tafel 381, fig. 3 of Bloch).

Salmo bimaculatus Linne, p. 413. (tafel 382, fig. 2 of Bloch).

Salmo anastomus Linne, p. 414.

Salmo niloticus Linne, p. 414.

Salmo cyprinoides Linne, p. 414.

Salmo curimata Bloch & Schneider, p. 417.

Salmo aegyptius Gmelin, p. 418. {aegyptiacus of Gmelin).

Salmo gasteropelecu3 Gmelin, p. 418.

Salmo immaculatus Linne, p. 419.

Salmo gibbosus Linne, p. 419.

In 1802 Lacepede (Histoire naturelle des poissons) created for Salmo rhom-
beus Linne" 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 Regne animal, and a series of articles in the Memoires Museum
d'histoire naturelle. In the Regne 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. 1

In 1829, Memoires Museum d'histoire naturelle, 4, he further defined
Chalceus and in 1819, 6, 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 Ins new family Characinidae.

In 1844 Miiller 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
Miiller 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. Of these twenty-seven genera and eighty species were
American, the remainder African.

Miiller and Troschel's work was closely followed by the second revision of
the group. Cuvier and Valenciennes in the 19th (1846) and 22nd (1848) volumes
of their Histoire naturelle des poissons described many species and the genera
Lebiasina and Pyrrhulina, 19 and Parodon, Piabucina, Tometes, Mylesinus,
Chalcinus and Cynopotamus, 22. A retrograde step was taken in rejecting the
Characinidae and including the genera in the Salmonidae.

1 Gasteropelecus is attributed to Bloch.

THE CHARACIDAE. 27

In 1854 Girard (Proc. Acad. nat. sci. Phil., 6, p. 199) described the genus
Cheirodon, the first Characin reported from the Pacific slope of America. This
genus was afterward found to be widely distributed on the eastern slope.

In the same year Baird & Girard (Proc. Acad. nat. sci., Phil., 7, p. 27)
described Astyanax argenlatus, the only species that reaches the United States
and the first to be recorded of the overflow from South America northward.

The notable work by Castelnau, (1855) Exped. Amerique Sud. Poissons,
while containing many figures and description of species added nothing to our
understanding of the relations of the various members of the family.

In 1858 Gill published (Ann. Lye. nat. hist., N. Y., 6) a short paper on the
fresh-water fishes of Trinidad. In it he recognized Erythinus and Macrodon
as forming a distinct family and described a new subfamily, Stevardiinae. He
also established the genera Poecilurichthys and Hemigrammus.

In the following year (1859) appeared a very important contribution to
the knowledge of the Characins, Zur familie der Characinen, by Kner. He
defined the new genera Microdus, Rhytiodus, and Bryconops and described
and figured a large number of new species from various parts of South America.
Erythrinus and Macrodon are excluded from the Characidae on account of
the absence of an adipose fin. The same author (1863) defined the genera
Pseudochalceus, Chalcinopsis, and Saccodon, the latter in connection with
Steindachner who has since contributed so much to the knowledge of tropical
American fishes. In the same year Gunther defined the genus Crenuchus.

In 1864 we have the culmination of an epoch in the history of the Characins.
Before 1802, at least during the binomial period, naturalists concerned them-
selves altogether with the description of new species. Beginning with Lacepede,
in 1802, we have a series of descriptions of new genera by Cuvier, Spix, Agassiz,
and Muller. To this period belongs the work of Cuvier and Valenciennes
although it appeared later. In 1844 and 1845 we have the first attempts by
Muller and Troschel of a philosophic review of the material that had been
accumulated. All subsequent work was tinged by Muller and Troschel's Die
familie der Characinen. In 1864 was published the second revision of the
family. Cuvier and Valenciennes's work being an enumeration, or descriptive
catalogue, rather than a revision, and Kner's works being confined to American
species. Gunther, in the Catalogue of the fishes of the British Museum, 1864,
5, reunited the Erythrinidae with the Characinidae. He defined or used for
the first time the generic names Caenotropus, Brachyalestes, Creatochanes,
Hemibrycon, Scissor, Creagrutus, Anacyrtus, Roestes, Roeboides, Hystricodon,

28 THE AMERICAN CHARACIDAE.

Sarcodaces, Oligosargus, and Ichthyborus. Giinther recognized in all forty-
seven genera and two hundred and five species. Of these forty genera and one
hundred and eighty-one species are American.

Giinther groups the genera in a number of subfamilies whose characters
are largely the presence or absence of an adipose fin, perfection or imperfection
of dentition, the length of the dorsal fin, and the character of the gill-openings.

The following synopsis will indicate his subfamilies and the characters on
which they are based : —

a. Adipose fin absent Erythrinina, 1 American.

aa. Adipose fin present
6. Dentition imperfect

c. Dorsal fin short Curimatina, American.

cc. Dorsal fin rather long Cilharinina, African.

66. Dentition well developed
d. Dorsal fin short
e. Gill-openings narrow, the gill-membrane grown to the isthmus
/. Elements of the jaws separate

g. Nasal openings remote from each other Anostomatina, American.

(gg. Nasal openings close together Nannocharacina, African).

ee. Gill-openings wide, the gill-membrane not grown to the isthmus.
h. Teeth compressed, notched, or denticulated.

Telragonoplerina, American and African.

hh. Teeth all conical Uydrocyonina, American and African.

(jj. Both jaws very movable, their lateral halves being united into one piece.

Phagonina, African).
dd. Dorsal fin rather long

i. Gill-openings of moderate width, the gill-membrane being attached to the isthmus.

Distichodonlina, African.
ii. Gill-openings wide, the gill-membrane not being attached to the isthmus.

j. Belly rounded; jaws with conical teeth Ichthyborina, African.

jj. Belly rounded; canine teeth none. . . .Crenuchina, American and African.

jjj. Belly with a spinous serrature Serrasalmonia, American.

Giinther immediately (1865) added the Phagonina and shortly afterwards
(1867) the Nannocharacina from Africa, assigning the latter to a place
between the Anostomatina and the Tetragonopterina. Both are included
in parentheses in the foregoing synopsis.

Giinther's work stimulated exploration and description to a great extent,
and following the publication of his Catalogue we have contributions by Giinther
himself between 1864 and 1900; by Gill 1864-1903; Reinhard 1866; Cope
between 1870 and 1894; Lutken 1874-1890; Steindachner 1875 to the present;
Boulenger from 1887 to the present; Eigenmann and Eigenmann from 1889
to the present; Garman 1890-1895; Perugia 1891-1897; von Ihering 1893 to

1 Including the Stevardiinae of Gill.

THE CHARACIDAE. 29

the present; Berg 1895-1901; Lahille 1895 to the present; Ulrey 1895; Regan
1900 to the present; Meek 1885-1916; Fowler; Nichols; Hildebrand and Mrs.
Marion Durbin Ellis.

The discovery of the numerous species and genera since 1864 are too com-
plicated for detailed enumeration. Reinhard and Liitken considered chiefly
the species inhabiting the basin of the Rio das Velhas a tributary of the Rio
San Francisco, Central America and Trinidad. Cope, 1870-1878, dealt largely
with species of the Upper Amazons, and later with species from Rio Grande do
Sul. Steindachner has published descriptions of numerous new species, col-
lected by Agassiz and his associates during the Thayer Expedition and in part
by himself and correspondents. Boulenger has reported on numerous collec-
tions received by the British Museum from different parts of South America.
Eigenmann and Eigenmann have reviewed the Erythrininae and Curimatinae;
Carman has published critical revisions of a few genera; Ulrey has reviewed
parts of the Tetragonopterinae. Perugia reported on various collections re-
ceived by the Museum of Genoa. Regan is describing species from the collec-
tions of the British Museum and Holmberg, von Ihering, Senior and Junior,
Berg, Lahille, Ribeiro, and Goeldi were the first of a group of resident naturalists
who have made important observations on their own faunae, chiefly eastern
Brazil and Argentina. While many of the authors suggest modifications in
parts of the system proposed by Gunther, only Eigenmann and Regan con-
cerned themselves with the broader questions of the classification of the
Characins.

In 1884 Sagemehl demonstrated the close relationship of the catfishes,
electric eels (Gymnotidae), Cyprinidae, and Characidae, all of which he grouped,
on account of the common possession of the complicated Weberian apparatus, in
the superorder Ostariophysi. Further studies on the anatomy of the Characins
were published in 1903 by Rowntree.

Gill, in 1893, (Families and subfamilies of fishes) admitted the two families
of Heterognaths, Characinidae, and Erythrinidae. He had defined these in
1858 and redefined them in 1895 (Proc. U. S. N. M., 18) when he intimated the
existence of a third family, the Citharinidae. Together with Muller and Troschel
and Kner he considers that the Characinidae, even after the exclusion of the
Erythrinidae, "constitute a heterogenous group." Gill recognizes the sub-
families Erythrininae, Pyrrhulininae, Lebiasininae, Tetragonopterinae, Ser-
rasalmoninae, Hydrocyoninae, Myletinae, Distichodontinae, Anostominae,
Curimatinae, and Citharininae.

30 THE AMERICAN CHARACIDAE.

In a recent paper (Cambridge natural history. Fishes, 1904) Boulenger
while recognizing that "The classification of the family is still in an unsatis-
factory state "divides them into the following groups (hardly deserving the rank
of subfamilies) " : —

A. Erythrininae American.

B. Hydrocyoninae African and American.

C. Serrasalmoninae American

D. Ichthyoborinae African.

E. Xiphostominae American.

F. Anostominae American.

G. Hcmiodontinae American.

H. Distichodontinae African.

I. Citharininae African and American.

Regan (Ann. mag. nat. hist., 1911, ser. 8, 8) comes to quite different con-
clusions in regard to the "groups" of Boulenger. He divides his Characiformes,
i. e. the Heterognathi of authors, into six families (a) the Characidae which
equals the Erythrininae, Hydrocyoninae, and Serrasalmoninae of Boulenger,
(b) the Xiphostomatidae = Xiphostominae of Boulenger, (c) the Anostomidae
= Anostominae of Boulenger plus Curimatus and Prochilodus, (d) the Hemiodon-
tidae = Hemiodontinae of Boulenger, (e) the Citharinidae = Ichthyoborinae,
Distichodontinae, and Citharininae exclusive of Curimatus and Prochilodus
of Boulenger, and (f) the Gastropelecidae proposed for the flying Chara-
cins included in the Hydrocyoninae by Boulenger.

Regan's paper offers some criticism of my classification published in the
Reports of the Princeton University Expeditions to Patagonia, 1909, 3, p. 253-
256. In part I heartily agree with Regan, and long ago came to some of the
conclusions reached by him. There are, however, many points in Regan's
paper in which I think his conclusions are at variance with the facts. I reserve
my criticism of Regan's strictures and a discussion of the general classification
of the Characins until my study of all the subfamilies is completed. The keys
of the subfamilies given in the Patagonian reports do not agree entirely with my
present views, (cf. Rept. Princeton univ. exped. to Patagonia, 1909, 3, p.
253-256).

Geographical Distribution.

In the study of the Characins, as in that of all Tropical American fishes,
the question of the distribution of the genera and species must in the future be
among the first topics to be considered.

In 1891 and 1892 Eigenmann and Eigenmann enumerated all the known

THE CHARACIDAE. 31

Tropical American fishes. They (Proc. U. S. N. M., 14, 15) considered and
summarized the distribution of the genera and species so far as known. They
recognized sixty-five genera and about four hundred and sixty species, as com-
pared with the forty genera and one hundred and eighty-one species enumer-
ated by Gunther (1864). It is not necessary to give here the details of the
results of their inquiry into the geographical distribution of the species. It
was found that nineteen of the genera recognized were distributed over the
entire eastern slope of South America and that five of these had representatives
in the La Plata and the Amazon, but not in the small rivers emptying into
the Atlantic in southeastern Brazil. One genus, Saccodon, was confined to the
Pacific slope. Twenty-seven genera were limited to the Amazons, or to the
Amazons and the region north of it. The Guianas held two peculiar genera,
the Rio Magdalena one, the southeastern coast streams one, while four genera
had a wide but irregular distribution.

Many modifications in these summaries of distribution are necessary,
both on account of the changes in the boundaries of the genera, and owing to
the increase in our knowledge of the distribution of the several species. The
papers published since the enumeration of 1891 and 1892 have dealt largely
with the fauna of Paraguay, Rio Grande do Sul, Guiana, Colombia, and Mexico
at nearly opposite ends of the range of the family and with the fauna of the
Pacific slope of Ecuador. 1 A reconsideration of the entire problem of the dis-
tribution of the fresh-water fishes of South America may be found in The fresh
water fishes of Patagonia and an examination of the Archiplata-Archhelenis
theory. (Reports of the Princeton university expeditions to Patagonia, 1909,
3, p. 225-374. Catalogue of the fresh-water fishes of tropical and south tem-
perate America. Ibid., 1910, 3, p. 375-512.

1 While no attempt has been made to trace the details of the evolution of our knowledge of the
African Characins the present account would be most inadequate and incomplete without reference to
Boulenger's work on the Characins of the Congo and Nile Basins. The new genera and species from
the Congo were described for the most part in volume 1 and 2 of the Annales Musee du Congo. A general
account in which all the species were considered, formed part of his Les poissons du basin du Congo,
1901. The Nile representatives are described and figured in his superb volume, The fishes of the Nile,
1907. Among other recent authors on the African Characins is J. Pellegrin, who is describing the
material of the Paris Museum. Finally Boulenger (Catalogue of the fresh-water fishes of Africa, 1909,
1, p. 174-298) redefines the subfamilies and genera and redescribes all of the African species. He recog-
nizes twenty genera and one hundred species.

32

THE AMERICAN CHARACIDAE.

Chronological List of Generic Names.

Original name

Date

Current Name

Charax Gnonow

1754 & 1777

Erythrinus Gronow

1756 & 1777

Gasteropelecus Gronow

1756 & 1777

Anostomus Gronow

1756 & 1777

Tetragonopterus Artedi

1758

Characinus Lacepede

1S02

Serrasalmo Lacepede

1S02

Curimates Cuvier

1815

Tetragonoptere Cuvier

1815

Les Curimates Cuvier

1817

Curimatus

Les Anostomes Cuvier

1817

Anostomus

Les Piabuques Cuvier

1817

Piabucus.

Tetragonopterus Cuvier

1817

Myletes Cuvier

1817

Hydrocynus Cuvier

1817

Citharinus Cuvier

1817

Gasteropelecus Cuvier

1817

Erythrinus Cuvier

1817

Curimatus Oken

1817

Piabucus Oken

1817

Chalceus Cuvier

1818

Curimata Cloquet

1818

Hydrocyon Cuvier

1819

Hydrocynus

Anodus Spix

1829

Prochilodus Agassiz

1829

Leporinus Spix

1829

Schizodon Agassiz

1829

Anostomus

Rhaphiodon Agassiz

1829

Xiphostoma Spix

1829

Pacu Spix

1829

Prochilodus

Cynodon Spix

1S29

Xiphorhynchus Agassiz

1829

Acestrorhynchus

Salminus Agassiz

1829

Macrodon Miiller

1842

Hoplias

Hemiodus Miiller

1842

Chilodus Miiller & Troschel

1844

Distichodus Miiller & Troschel

1844

Alestes Miiller & Troschel

1S44

Brycon Miiller & Troschel

1844

Exodon Miiller & Troschel

1S44

Epicyrtus Miiller & Troschel

1844

Charax

Hydrolycus Miiller & Troschel

1844

Pygocentrus Miiller & Troschel

1844

Pygopristis Miiller & Troschel

1S44

Catoprion Miiller & Troschel

1844

THE CHARACIDAE.

33

Original name

Date

Current Name

Myleus Muller & Troschel

1844

Xiphorhamphus Muller & Troschel

1845

Acestrorhamphus

Agoniates Muller & Troschel

1845

Grundulus Cuv. & Valenciennes

1846

Lebiasina Cuv. & Valenciennes

1846

Pyrrhulina Cuv. & Valenciennes

1846

Parodon Cuv. & Valenciennes

1848

Brycinus Cuv. & Valenciennes

1848

Alestes?

Piabucina Cuv. & Valenciennes

1848

Tometes Cuvier & Valenciennes

1848

Mylesinus Cuv. & Valenciennes

1848

Chalcinus Cuv. & Valenciennes

1S4S

Cynopotamus Cuv. & Valenciennes

1848

Hydropardus Reinhardt

1849

Raphiodon

Cheirodon Girard

1854

Astyanax Baird & Girard

1854

Poecilurichthys Gill

1858

Astyanax

Hemigrammus Gill

1858

Stevardia Gill

1858

Corynopoma Gill

1858

Stevardia

Nematopoma Gill

1858

Stevardia

Microdus Kner

1859

Caenotropus

Rhytiodus Kner

1859

Bryconops Kner

1859

Ctenolucius Gill

1861

Luciocharax

Hydrocyonoides Castelnau

1861

Sarcodaces

Crenuchus Giinther

1863

Pseudochalceus Kner

1863

Chalcinopsis Kner

1863

Brycon

Saccodon Kner & Steindachner

1863

Caenotropus Giinther

1884

Brachyalestes Giinther

1864

Creatochanes Giinther

1864

Hemibrycon Giinther

1864

Scissor Giinther

1864

Creagrutus Giinther

1864

Anacyrtus Giinther

1864

Charax

Roestes Giinther

1864

Roeboides Giinther

1864

Hystricodon Giinther

1864

Exodon

Sarcodaces Giinther

1864

Oligosargus Giinther

1864

Ichthyborus Giinther

1865

Phago Giinther

1865

Piabina Reinhardt

1866

Characidium Reinhardt

1866

Nannocharax Giinther

1867

Xenocharax Giinther

1867

34

THE AMERICAN CHARACIDAE.

Original name

Date

Current name

Aphyocharax Giinther

1868

Megalobrycon Giinther

1869

Brycon

Stethaprion Cope

1870

Holotaxis Cope

1870

Plethodeetes Cope

1870

Chalceus

Odontostilbe Cope

1870

Laemolyta Cope

1871

Nannaethiops Giinther

1871

Iguanodectes Cope

1871

Nannostomus Giinther

1872

Bryconaethiops Giinther

1873

Leporellus Liitken

1874

Curimatopsis Steindachner

1S76

Liitkenia Steindachner

1876

Stichanodon

Paragoniates Steindachner

1876

Bramocharax Gill

1877

Luciocharax Steindachner

1878

Elopomorphus Gill

1878

Potamorhina Cope

1878

Metynnis Cope

1878

Leptagoniates Boulenger

1887

Psectrogaster Eigenmann & Eigenmann

1889

Curimatella Eigenmann & Eigenmann

1889

Semitapicis Eigenmann & Eigenmann

1889

Henochilus Garman

1890

Schizodontopsis Garman

1S90

Laemolyta

Pseudocorynopoma Perugia

1891

Bergia Steindachner

1891

Pseudocorynopor

Chalcinopsis Holmberg

1891

Pseudocorynopor

Neolebias Steindachner

1S94

Asiphonichthys Cope

1894

Chorimycterus Cope

1894

Diapoma Cope

1894

Petersius Hilgendorf

1894

Nanognathus Boulenger

1895

Hoplerythrinus Gill

1895

Eugnathichthys Boulenger

1898

Paraphago Boulenger

1899

Neoborus Boulenger

1899

Micralestes Boulenger

1899

Neolebias Boulenger

1899

Catabasis Eigenmann & Norris

1900

Mesoborus Pellegrin

1900

Hemistichodus Vaillant & Pellegrin

1900

Citharidium Boulenger

1902

Gymnocharacinus Steindachner

1903

Hoplias Gill

1903

Anisitsia Eigenmann & Kennedy

1903

. THE CHARACIDAE.

35

Original name

Date

Current Name

Lahilliella Eigenmann & Kennedy

1903

Holoshesthes Eigenmann

1903

Holoesthes

Holoprion Eigenmann

1903

Holopristis Eigenmann

1903

Pristella

Markiana Eigenmann

1903

Moenkhausia Eigenmann

1903

Othonophanes Eigenmann

1903

Bryconodon Eigenmann

1903

Stichanodon Eigenmann

1903

Evermannella Eigenmann

1903

Eucinopotamus

Acestrorhynchus Eigenmann

1903

Acestrorhamphus Eigenmann

1903

Acestrocephalus Eigenmann

1903

Boulengerella Eigenmann

1903

Gilbertella Eigenmann

1903

Gibertolus

Aenodon Eigenmann

1903

Myleocollops Eigenmann

1903

Piaractus Eigenmann

1903

Orthomyleus Eigenmann

1903

Colosoma Eigenmann

1903

Mylosoma Eigenmann

1903

Eucinopotamus Fowler

1900

Ophiocephalops Fowler

190G

Hoplerythrinus

Copeina Fowler

1906

Cyphocharax Fowler

1906

Steindachnerina Fowler

1906

Peltapleura Fowler

1906

Eigenmannina Fowler

1906

Chilomyzon Fowler

1900

Prochilodus

Hemiodopsis Fowler

1906

Pithecocharax Fowler

1906

Anostomus

Poecilosomatops Fowler

1906

Characidium

Garmanina Fowler

1906

Abramites Fowler

1906

Leporinus

Pellegrinina Fowler

1906

( haleeus

Coscinoxyron Fowler

1906

Thoracocharax Fowler

1906

Cyrtocharax Fowler

1906

Cynopotamus

Cynocharax Fowler

1906

Sphyraenocharax Fowler

1906

Acestrorhamphus

Belonocharax Fowler

1906

Lucioeharax

Waiteina Fowler

1900

Colosoma?

Reganina Fowler

1906

Colosoma?

Starksina Fowler

1906

Mylosoma

Sealeina Fowler

1900

E\'ermannolns Eigenmann

1907

Eucinopotamus

Gilbertolus Eigenmann

1907

Phenacogrammus Eigenmann

1907

36

THE AMERICAN CHARACIDAE.

Original Name

Date

Current Name

Pogonocharax Regan x

1907

Phoxinopsis Regan

1907

Mimagoniates Regan

1907

Ctenocharax Regan

1907

Grundulus

Eobrycon Jordan

1907

Bryconamericus Eigenmann

1907

Deuterodon Eigenmann

1907

Phenaeogaster Eigenmann

1907

Astyanacinus Eigenmann

1907

Fowlerina Eigenmann

1907

Ephippicharax

Joinvillea Steindachner

1908

Deuterodon

Coelurichthys Ribeiro

1908

Gymnocorymbus Eigenmann

1908

Thayeria Eigenmann

1908

Ctenobrycon Eigenmann

1908

Pristella Eigenmann

1908

Psellogrammus Eigenmann

1908

Hyphessobrycon Durbin

1908

Brycochandus Eigenmann

1908

Poptella Eigenmann

1908

Champsoborus Boulenger

1909

Anostomoides Pellegrin

1909

Jobertina Pellegrin

1909

Pterodiseus Eigenmann

1909

Carnegiella Eigenmann

1909

Holobrycon Eigenmann

1909

Triurobrycon Eigenmann

1909

Brycon

Poecilocharax Eigenmann

1909

Microcharax Eigenmann

1909

Poecilobrycon Eigenmann

1909

Archicheir Eigenmann

1909

Hollandichthys

1910

Nematobrycon Eigenmann

1911

Knodus Eigenmann

1911

Bivibranchia Eigenmann

1911

Hasemania Ellis

1911

Probolodus Eigenmann

1911

Psalidodon Eigenmann

1911

Spintherobolus Eigenmann

1911

Glandulocauda Eigenmann

1911

Hysteronotus Eigenmann

1911

Vesicatrus Eigenmann

1911

Apodastyanax Fowler

1911

Ctenobrycon.

Rhodsia Fowler

1911

Parastremma Eigenmann

1912

Genycharax Eigenmann

1912

1 This is probably a Cyprinoid from Ceylon not a Characin from South America.

THE CHARACIDAE.

37

Original name
Gephyrocharax Eigenmann
Pterobrycon Eigenmann
Argopleura Eigenmann
Microgenys Eigenmann
Zygogaster Eigenmann
Ephippicharax Fowler
Prionobrama Fowler
Gnathocharax Fowler
Tyttocharax Fowler
Xenurocharax Regan
Landonia Eigenmann
Phenagoniates Eigenmann & Wilson
Microbrycon Eigenmann & Wilson
Ceratobranchia Eigenmann
Bleptonema Eigenmann
Pareebasis Eigenmann
Myocharax Fowler
Xiphocharax Fowler
Leptobrycon Eigenmann
Macropsobrycon Eigenmann
Megalamphodus Eigenmann
Microschemobrycon Eigenmann
Oligobrycon Eigenmann
Aphyocheirodon Eigenmann
Compsoura Eigenmann
Mixobrycon Eigenmann

Date

Current name

1912

1913

1913

Bryconamericus

1913

1913

Astyanax

1913

1913

1913

1913

1913

1914

1914

Phanagoniates

1914

1914

1914

Prionobrama

1914

1914

1914

1915

1915

1915

1915

1915

1915

1915

1915

38 THE AMERICAN CHARACIDAE.

THE AMERICAN TETRAGONOPTERINAE.

Whether we regard the Heterognaths as a single family, or as several fami-
lies, the fact remains that there are few groups of fishes within which the lines
of evolution are so clearly portrayed by existing forms as in the Characidae.
In order more satisfactorily to discuss their evolution I have divided the
family into a large number of small groups of genera with undoubted affinity.

Several facts, aside from the general structure point to the Tetragonop-
terinae or perhaps the closely allied Cheirodontinae as the group nearest
to the ancestral Characins.

1 . Tetragonopterid fishes have been found fossil in the Tertiary fresh-water
deposits at Taubate.

2. They are found both in Africa and South America, the most nearly
allied genera on the two sides, Astyanax and Petersius, are scarcely generi-
cally distinct. No other subfamily has representatives on both sides of the
Atlantic.

3. They are the dominant groups both in Africa, where they form more
than 36% of the Characins, and in America, where they form about 40%.

4. They are found over the entire area of distribution both in America
and Africa. In America members of this subfamily form everywhere the
vanguard in the distribution.

5. There are several lines of evolution diverging from the two subfamilies,
Cheirodontinae and Tetragonopterinae.

Some of the lines of evolution radiating from different sections of the Tetra-
gonopterinae, or the closely allied Cheirodontinae are minor lines that have
not diverged greatly. For instance Diapoma and Stevardia are Tetragonop-
terids with modified opercles. The Gymnocharacinae are naked Tetragonop-
terinae. The Crenuchinae are apparently an offshoot from the Cheirodontinae.
The Stethaprioninae have developed a predorsal spine but are otherwise very
close to Tetragonopterus and especially to Moenkhausia. The Mylinae and
Serrasalmoninae have possibly diverged from the Stethaprioninae, increasing
the number of the dorsal rays, increasing the depth, compressing the ventral
surface, and adding spines and emphasizing the dentition without much alter-
ing it. Another offshoot from the Cheirodontinae has given rise to a series of
fishes with a decrease in the size of the mouth and the effectiveness of the

THE AMERICAN TETRAGONOPTERINAE. 39

dentition. This series includes the Anastomatinae, Chilodinae, Prochilodinae,
Hemiodontinae, Elopomorphinae, and finally, the toothless Curimatinae.

Another fine diverging from the Cheirodontinae has given rise to the Sal-
mininae, Characinae, Acestrorhamphinae, Cynodontinae, and ultimately the
Hydrocyninae.

Another line of divergence from the Cheirodon or Tetragonopterus group
led through the Bryconinae, Iguanodectinae, Pyrrhulininae, Piabuscininae,
to the Lebiasininae. A side branch from this leads to the Chalcininae which
points the way to the flying Gasteropelicinae. For the above reasons it seems
best to begin the detailed examination of the species with the Tetragonopterinae.

It may be left an open question whether the African and American genera
owe their similarity to convergence or to community of origin. They are for
the most part small or minute fishes ranging from 50-200 mm. in length.
Myletes in Africa reaches 460 mm. The Tetragonopterinae are closely related
to the Cheirodontinae, Diapominae, Glandulocaudinae, Stethaprioninae, Bry-
coninae and to other subfamilies. In shape they vary from the fusiform
Creatochanes to the deeply compressed Tetragonopterus. In the majority
of the genera the mouth is small. The lower jaw is heavy, and on account of
the obliquity of the mouth, when the mouth is opened it is thrown forward,
so as to project beyond the snout; when the mouth is closed the teeth of the
lower jaw usually fit in behind the innermost series of the premaxillary. In
some genera from Africa there is an inner pair of conical teeth in the lower jaw.
These appear in lieu of an inner series of teeth which in the American Brycon
are still present on the sides of the lower jaw. Conical teeth like these appear
in isolated (not closely related) genera of other subfamilies both in Africa and
America, and, since they cannot be genetically connected, appear to offer an
example of homoplastic development. The skull is smooth in cross-section or
slightly grooved in the smaller specimens; two fontanels are well developed in
all the genera but Brycinus of Africa. In this genus there is no frontal fontanel.
The bridge between the fontanels is either flush with the surface or sunk beneath
it; the occipital process varies directly with the depth of the species, and serves
to bridge the space between the skull and first interneurals, i. e. the space over
the coalesced vertebrae which lack interneurals. In the deeper species the
process is curved or bent upward and is long, reaching as much as one third of
the distance of its base from the dorsal. In the slenderest species it is short and
insignificant. It is always grooved to its tip, the groove leading to the parietal
fontanel.

40 THE AMERICAN CHARACIDAE.

The eyes are always large, the species depending largely on living and
moving food. The cheeks may be narrow or deep and are one third or entirely
covered by the second suborbitals.

The premaxillary teeth are always in at least two series. The inner series
consists of from 4-12 graduated; three to many-pointed (a few may be conical)
teeth arranged in a regular series. The outer row is very variable both in the
group and in individual species and ranges from one or more teeth near the
middle of the premaxillary, to a complete, compact series of teeth, narrower
than those of the inner series in Tetragonopterus. If the outer row consists of
four or more teeth the third tooth usually drops out of line and tends to form a
third series of teeth. In Moenkhausia melanogramma the third tooth is entirely
withdrawn from the line of the others. In the related Brycon in which the
teeth are more numerous than in the genera of this subfamily, the fourth tooth
and a few others also drop out of line and join the third tooth of the lower inner
series which has moved forward. Other teeth of the inner series have also
moved forward giving rise to a third series of teeth. A third series of teeth has
independently arisen in Creagrutus and Bryconops, and in the Bryconinae.

The maxillary may have no teeth, or a few may be crowded along the part
of the margin nearest to the premaxillary, or it may have conical or tricuspid
teeth along its entire margin. Usually the variation in any one species is very
limited, but in Astyanax fasciatus nicaraguensis there is a variation of from one
to nine teeth in the maxillary.

The teeth in the lower jaw may all be alike and graduate, or the lateral
teeth may be more or less abruptly smaller, the more abruptly the more proba-
bly the lateral teeth will be conical. The larger teeth, 3-6 in number, may form
a nearly transverse series or be arranged in a curve.

The denticles of the individual teeth of the inner series of the premaxillary
may be in a straight line, i. e., the teeth may be strictly incisors or, the line
joining the cusps may be more and more curved so that it will be U-shaped,
the open part of the U forward. The teeth in the lower jaw are usually the
reverse of those of the inner series of the upper jaw.

Gill-rakers are usually slender and not very long, they are all but absent
in one genus, Scissor. The gill-membranes are free from each other, the nares
close together. The breast is flat or rounded, never keeled. The scales are
usually cycloid, rarely crenate or even ctenoid. They vary from 26 to about
60 in the lateral line, which is variously developed.

The caudal may be naked, the scales of the sides passing on to its base and

THE AMERICAN TETRAGONOPTERINAE. 41

ending in slightly enlarged scales, or the scales may become minute on the
caudal adhering and covering the lobes to a greater or less extent. The anal
may similarly have simply a basal sheath or may be covered to near its tip with
minute scales.

The scales may be regularly imbricate or in certain regions there may be
interpolated rows. In some species there are but a few interpolated scales,
i. e., a single series becomes divided into two above the anal. The point of
division becomes more and more removed from the anal and the number of
divided series increases. In long slender species, or larger-scaled, deep species,
there are no interpolated rows. In deep, many-scaled species they sometimes
become numerous. Closely allied species or even varieties may differ in this
respect. In Astyanax fasciatus individuals with interpolated series are rarely
found except in the Rio Parahyba where the usual variety has been entirely
replaced by one with interpolated series.

The dorsal is short, of between 9 and 12 rays, counting everything, and
its origin is usually in the middle of the body. The adipose dorsal is small
but almost always well developed. The caudal is always forked, the lobes
equal or subequal, rarely markedly different. The anal is another variable
element. Its origin usually below or behind the last dorsal ray may (Psello-
grammus and Phenacogaster) fall below its origin. Other things being equal
the deeper species have the larger number of anal rays. The number of rays
varies from 10 to 48.

The reach of the pectorals and ventrals varies with the shape of the fish.
In deep species with long anal they overlap, in slender species they ordinarily
do not reach each other. The notable exception to this is found in Gephyro-
charax.

The alimentary canal varies but little from the entire length of the fish.
There are a few pyloric coeca.

The air-bladders are large, the posterior about twice the length of the
anterior, curved down behind in the deeper species.

The range of color is limited. In life the caudal of the male is frequently
cherry-red, the dorsals and anal (and caudal in females) are frequently yellow.
A shoulder-spot of varying shape is usually present. There is usually a silvery
lateral band overlying a black band which becomes evident in formalin prepara-
tions. There is frequently a dark spot on the base of the caudal which is often
continued on the middle caudal rays. In Moenkhausia dichrourus and Brycona-
mericus exodon the tips or bands across the caudal lobes are dark. In Moenk-

42

THE AMERICAN CHARACIDAE.

hausia lepidurus and the species of Creatochanes the middle rays and upper
caudal lobes are black. In Astyanax lineatus and A. steindachneri, in Moenk-
hausia latissimus, and in Hollandichthys and Pseudochalceus dark lines follow
the spaces between successive rows of scales. The dorsal and anal are marked
with black in a few species.

The points of greatest variability within a genus are: —

1. The depth and all that this carried with it, length of occipital process, rows of scales,
number of rays of anal. 2. The size of the mouth and the dentition. 3. The degree of
armature of the cheek. 4. The scaling of the caudal, anal, predorsal, and preventral areas.
5. The degree of development of the lateral line.

Contrasted Generic Characters.

The following mutually exclusive characters are found variously combined
in different genera. The characters appearing in the largest number of genera
are given in the first column.

A. Caudal fin naked. a.

B. Lateral line complete. b.

C. Maxillary with few teeth or none. c.
V. Premaxillary teeth in two series. d.

E. Cheeks partly naked. e.

F. Anterior edge of maxillary a simple /.

curve.

G. Predorsal line scaled. g.
H. Teeth of the sides of the dentary h.

abruptly smaller.
I. Premaxillary meeting the maxillary of an i.

angle.

J. Scales entire. j-

3-

K. Anal naked except at the base. k.

L. Lateral line nearly straight. /.

M. Preventral area with normal scales. m.

N. Gill-rakers setiform. n.

0. Adipose fin present. o.

P. Origin of anal behind origin of dorsal. p.

Q. Caudal without glandular scales. q.

Caudal scaled.
Lateral line incomplete.
Maxillary with teeth along its entire edge.
Premaxillary teeth in three series.
Cheeks entirely covered by the third sub-
orbital.
Anterior edge of maxillary sigmoid.

Predorsal line naked.

Teeth of the sides of the dentary gradu-
ated.

Maxillary-premaxillary border a simple
curve.

Scales ctenoid.

Scales crenate.

Anal scaled to near its tip.

Lateral line sharply decurred in front.

Preventral area with paired scales.

Gill-rakers lanceolate.

Adipose fin wanting.

Origin of anal under or in front of origin
of dorsal.

Caudal with glandular scales.

The characters appearing in the larger number of genera and given in the
first column are all found in Astyanax, which may for that reason, be the central,

THE AMERICAN TETRAGONOPTERINAE. 43

possibly the most primitive member, of the subfamily. Poecilurichthys, which
differs from it only in having the character I instead of L, is scarcely distin-
guishable. 1

The different characters of the first column are found in various combina-
tions with most of the characters of the second column. Thus A is found in
combination with all (not counting its contrasted character, a) except k and I.
B is found in combination with all but o and p. C is found in combination with
all the characters of the second column and the same is true of all the rest of
the characters of the first column.

POLYPHYLETIC CHARACTERS.

It is quite certain that the characters of the right-hand column of the
contrasted generic characters have sometimes, at least, been independently
derived from the characters of the left-hand column, not only by the different
genera possessing the character, but sometimes by different species of the same
genus. That is, some of the genera possessing characters given in the second
column are of polyphyletic origin.

Beginning at the bottom of the list, the character o is found in Hasemania
from southeastern Brazil, and in the very different genus, Nematobrycon,
from the west of the western Cordilleras of Colombia. The two genera are not
related, and are widely separated geographically. There can be no doubt but
that they have independently lost the adipose fin.

The next undoubted case is that indicated by g. This character is found
in Poecilurichthys, scarcely distinct from Astyanax, and in Gymnocorymbus,
a very different fish. Here again the character has very probably been inde-
pendently acquired by the two genera.

1 In Indiana Univ. Studies, 1914, No. 20, I have tried to carry the inference a step further.

"The common possessions of all members of the Tetragonopterina; enumerated above, permit us
to picture the ancestral type of the subfamily. In brief, it must have been a fish similar in most char-
acters to Astyanax fasciatus Cuvier. This species, besides possessing all the characters common to all
members of the subfamily, possesses also many of those positive (as contrasted with absent) characters
enumerated for the family, and lacks some characters, like the highly specialized scaling of the ventral
surface, ctenoid scales, extreme length of anal, extreme development of second suborbital, which are
evidently highly specialized characters in a few of the genera. It is more widely distributed than any
other species and has given rise to numerous variations.

It represents an average in length of head (4.3), depth, (2.6-3); length of anal (about 30); scales
(about 38); size of eye (2.5-3); general shape (compressed subfusiform) ; position of dorsal (its base
being in the space above the origins of the ventral and anal) ; size of mouth ; and the characters of the
teeth. The fossil fishes found at Taubate — south of Rio de Janeiro — are similar to it in most char-
acters. They are a little larger and may be members of the genus Brycon. In all but the teeth, they
are very similar to Astyanax fasciatus."

44 THE AMERICAN CHARACIDAE.

The next case is that of e, the enlarged third suborbital. This character is
found in Knodus with a scaled caudal and in Creagrutus, Piabina, and Brycon-
americus with a naked caudal. The last three genera are undoubtedly closely
related and for the present purpose count as one. Knodus is a Bryconamericus
in all but its scaled caudal. We are, therefore, compelled to assume either that
if the ancestors of Knodus had the caudal scaled, that it has paralleled Brycon-
americus in the character of its cheek or that if its ancestors were identical with
those of the latter genus that its caudal has independently acquired scales.

The case presented by the character d, three rows of premaxillary teeth, is
much simpler and clearer. This character is found in Microgenys, Creagratus,
Piabina, and Bryconops. It is quite certain that the latter genus has been
derived from an ancestor like Creatochanes and that the others have been de-
rived from an Astyanax-like ancestor. Microgenys, Creagratus, and Piabina,
on the one hand, and Bryconops on the other, have independently acquired
three series of teeth. In this case the steps by which this has been accomplished
are indicated in a variety of species of the subfamily. In many species alternate
teeth of the front series of the premaxillary are withdrawn from the line of the
rest, thus forming an incipient third series. In Moenkhausia melogramma and
in Bryconamericus exodon this process has almost yielded additional genera
with three series of teeth. The modification from one to the other condition
is a perfectly progressive one, without notable breaks or saltations.

The next character c, teeth along the entire maxillary, has again been inde-
pendently derived from C several times. Here we have not only the evidence
of several distinct, not closely related, genera which have the character but
also the evidence from changing species. I have elsewhere called attention to
some specimens referred to as Aslyanas aeneus nicaraguensis. Of thirty-five
specimens from Lake Nicaragua, there are nine with two teeth, two with three
teeth, five with four teeth, five with five teeth, five with six teeth, five with seven
teeth, three with eight teeth and one with nine teeth in the maxillary. The
normal number is two. Phenacogaster is in a similar state of transition. Pris-
tella, Hemibrycon, Nematobrycon, Hollandichthys, and Pseudochalceus have
acquired complete dentition for their maxillaries in at least three independent
groups.

The most interesting and conclusive evidence of the independent origin
of the same character in different genera is presented by the character b, the
incompleteness of the lateral line. Not only have we the evidence from widely
divergent genera with this character, but we again have species in a state of true
mutation. Most remarkable of all is one species of which I have been able

THE AMERICAN TETRAGONOPTERINAE.

45

to examine hundreds of specimens from the Amazon, not one of which showed
signs of mutation, while the specimens coming from another region are in an
evident state of mutation.

Genera with a complete lateral line and the genera with an incomplete
lateral line to which they have given rise are : —

Genera with

lateral line complete.

Genera with lateral line incomplete

Tetragonopterus

none.

Entomolepis

none.

Moenkhausia

Hemigrammus (further changed into Pristella and Thayeria)

Astyanax

Hyphessobrycon (into Hasemania).

Knodus

none.

Markiana

none.

Gymnocorymbus

none.

Ctenobrycon

Psellogrammus.

Creatochanes

Brycochandus.

Bryconops

none.

Creagrutus

none.

Piabina

none.

Microgenys

none.

Bryconamericus

none.

Zygogaster

none.

Ceratobranchia

none.

Landonia

none.

Deuterodon

none.

Hemibrycon

Hollandichthys, Pseudochalceus, and Nematobrycon.

Phenacogaster

Vesicatrus.

Scissor

none.

Henochilus

none.

Psalidodon

none.

The deviation is so evident in a number of cases that the polyphyletic origin
of the character 6, an incomplete lateral line, is beyond the faintest shadow of a
doubt.

Without considering sporadic individuals of otherwise constant species,
the species which are undoubtedly mutuating at the present time are Hemi-
grammus inconstans, Moenkhausia cotinho, Phenacogaster beni, Astyanax mutator,
and Hyphessobrycon poecilioides. The details for these species are given under
their descriptions. Another species which has crossed the line but has not
reached a state of equilibrium is Psellogrammus kennedyi. Many of the details
of this species are again given under the proper caption.

Of Ctenobrycon hauxwellianus I have been able to examine over fourteen
hundred specimens from various places on the Amazon. In all of these the
lateral line is complete. In at least six specimens out of nineteen from the Lagoa

46 THE AMERICAN CHARACIDAE.

Parnagua, Paranahyba basin, the lateral line "stutters." This species, in other
words, is mutuating at Parnagua. Moenkhausia cotinho is similarly locally
mutating and the same seems to be true of Hyphessobrycon poecilioides and
Astyanax fasciatus.

The scaling of the caudal I am not able to cope with satisfactorily. It is
certain that it has several times been acquired independently by different members
of the family, if not by different members of the subfamily under consideration.

Selective Grouping of Characters.

Another line of inquiry leads us to consider whether the contrasted unit char-
acters are entirely combined as if by chance or whether there is a selected com-
bination. The characters from g-q are found in but one or two genera and these
may be omitted from the discussion since they would needlessly complicate it.

Taking only the first six pairs of contrasted characters, there are 62 or 64
possible combinations. Considering each combination a distinct genus we
should have 64 genera, without considering the characters between g and q.
In reality we have but about half as many.

Taking only the first three characters we should have eight possible com-
binations. An examination of the genera shows that six of these combinations
are actually found, but not by any means in the same proportions. If we take
the first four characters, each of which has probably several times, and inde-
pendently, been modified into its contrasted character, these four pairs give us
sixteen possible combinations. Of these only eight actually occur.

A certain amount of selective grouping is thus found, if we take only three
contrasted pairs of characters, a greater amount of selective omission if we take
four pairs and this increases rapidly as we increase the number of contrasted
forms. It is evident that either many of the possible combinations have never
arisen, or, having appeared, they have not been preserved.

POLYPHYLETIC GENERA.

In my paper, Indiana University Studies, 1914, no. 20, I wrote —

"This independent origin of characters is responsible for the fact that some of the
accepted genera of the Tetragonopterinse are of polyphyletic origin, i. e. our definitions of
genera are in many cases enumerations of characters frequently independently acquired,
not enumerations of the characters of the ancestral type of the genus from which the species
have diverged. A result of this independent divergence is that frequently in a restricted,
isolated area the species of different genera represented in the area are more nearly related
to each other than to members of their own genera in remote regions. For instance Astyanax

THE AMERICAN TETRAGONOPTERINAE. 47

festce and Bryconamericus peruanus of the Pacific slope of Ecuador are more intimately related
than festa is to Astyanax anterior of the upper Amazon. And in this case, Astyanax brevi-
rostris or Bryconamericus brevirostris whichever it may be, is intermediate between the two.
I am not competent to say whether brevirostris is moving from Bryconamericus to become
an Astyanax, or whether it has just completed the reverse process. Certainly festa and
brevirostris are more intimately related, have had a common ancestor at a less remote time,
than either of them with an Astyanax or Bryconamericus of southeastern Brazil.

We recognize two types of genera, one a group of closely related species, descended
from a common ancestor and having certain distinguishing characters in common. Phena-
cogaster is such a genus. The peculiar scaling of the ventral surface has been developed
but once; and the species are all closely allied, differing from each other in but a few char-
acters. The other, a polyphyletic type, consists of species having a certain combination of
definite characters in common which easily distinguish members of the genus, but which,
instead of indicating a single ancestral line from which the species have diverged, are acquired
possibly one at a time along distinct lines converging to a common definition. Sometimes
the polyphyletic origin can be detected, sometimes not. Bryconamericus seems to me to be
such a genus; Hemibrycon, Deuterodon, and the larger genera are probably also polyphyletic.

" Since it is difficult, or impossible, to say in any case which of the given characters has
appeared first, it is extremely difficult to point out lines of evolution leading to different gen-
era or species. We can only insist that certain innate possibilities may become actualities
anywhere along the line, possibly wherever they may prove advantageous, though the advan-
tage, to say the least, is not always obvious.

" We may be permitted to assume that the more frequent character is the primitive one,
although this is certainly not always a safe assumption."

Deuterodon with the character h is a genus of polyphyletic origin. Deu-
terodon iquape is found in southeastern Brazil, Deuterodon nasatus in Central
America. These two species technically belong to the same genus, genetically
they are most certainly not derived from an immediate common ancestor and
it is very probable that Deuterodon pinnatus and Deuterodon potaroensis from
Guiana, and Deuterodon acanthogaster from the Paraguay, are also independent
derivatives from the genus Astyanax.

A somewhat similar case is presented by a character not mentioned in the
list, because I hesitate to propose a generic designation for it. It is this: —
In the vast majority of the species of Characins, the innominate bone is feeble
and entirely concealed. In Deuterodon acanthogaster and in Astijanax mucrona-
tus, this bone has become firm and the anterior end projects out of the body as
a distinct spine. There is no doubt whatever that this modification is arising
independently in the two species.

It must be quite evident from the foregoing that the subfamily is a paradise

for the student of divergent evolution. But the very conditions that make it

of interest to the student of evolution make it the despair of the systematist

whose object is to express relationship by grouping the species in an orderly

■ array of genera and the individuals in an orderly array of species, always, if

48 THE AMERICAN CHARACIDAE.

possible, in the form of the conventional phylogenic tree. In order to better
express the relationship of smaller groups of species the genera have been made
as small as consistent with facts. The Tetragonopterinae seem to form an
interlacing fabric rather than a branching tree. An illustration of the interest
and difficulty is given by the series of genera Poecilurichthys, Ctenobrycon,
Psellogrammus. The genus Poecilurichthys as here understood, grades in-
sensibly into the genus Astyanax, and perhaps should have been retained
united with the latter. Be that as it may, P. spilurus is much more closely
related to Ctenobrycon hauxwellianus than it is to any species of either Poe-
cilurichthys or Astyanax, and yet there is scarcely a technical character by
which it can be separated from the remaining species of Poecilurichthys and
united with Ctenobrycon. The latter is sharply distinguished by ctenoid scales
from all other members of the subfamily. In an another direction, the speci-
mens of Ctenobrycon hauxwellianus from Paranagua form a complete bridge so
far as the technical characters are concerned, between the genera Ctenobrycon
and Psellogrammus. From the parental form, P. spilurus, the Amazonian
Ctenobrycon hauxwellianus has become generically distinct. C. hauxwellianus
in its turn, in Paranagua, is becoming and, in Paraguay, has developed the char-
acter which distinguishes Psellogrammus.

The difficulties of the systematist could of course be obviated by retaining
all of these things in the single old genus Tetragonopterus, but this procedure
would neither remove the facts nor explain them.

I have attempted to express the relationship of the genera by the accom-
panying diagram, in which Astyanax occupies the center, and in which it is
made to appear that the scaled caudal has appeared once, of which I am not at
all certain, and in which the line of the outer ellipse indicates the boundary
within which the lateral line is complete. This boundary has been crossed
many times and by different radiating lines.

Fig. 1. — The genera Astyanax, and Moenkhausia, within the central oval, are closely allied and
the difference between them is partly bridged by the species M. intermedia. They are considered typical
and central for the subfamily. From Moenkhausia have radiated directly Markiana, Gymnocorymbus,
Tetragonopterus, and Entomolepis. It is left in doubt whether the genera Hemigrammus and Hyphesso-
brycon have diverged independently from Moenkhausia and Astyanax, and then converged till now only
the scaled and naked condition of the caudal separate them, or whether they have been derived from
one of the above and then developed or lost the caudal scales. The same applies to the genera Brycon-
americus and Knodus. The species Astyanax mutator and Hyphessobrycon proteus furnish bridges between
Hyphessobrycon and Astyanax. Moenkhausia colinho and Hemigrammus inconslans furnish a similar
bridge between Hemigrammus and Moenkhausia. From Astyanax many lines have radiated, some of
which have been continued beyond the line of the outer oval, which indicates that their lateral line has
become broken. Where a bridge exists between the so-called genera this has been indicated by extending
the name of the genus beyond its boundary or by supplying the name of the species which constitutes
the bridge in brackets. The origin of two of the radial lines is uncertain, a fact indicated by an interro-
gation point.

Cheir °aom, m

Rho&dsins

I^uanodecLinee.

Figure 1.

50 THE AMERICAN CHARACIDAE.

Key to the Genera.

a. Upper lip covering the teeth of the premaxillary ; dentary with a single series of teeth.
b. Some or all of the teeth serrate or notched.
c. Gill-rakers setiform or branched.
d. Preventral area with a median series of scales or with normal scales irregularly arranged, or
more or less trenchant with the scales of the two sides bent over the middle.
e. Maxillary border a simple more or less pronounced curve, or with a break in the symi letry
of the curve at the end of the tooth bearing portion.
/. Caudal fin at least partly covered with small, normal scales.

g. Lateral line complete (see also under Hemigrammus inconstans, Moenkhausia cotinho).
h. Predorsal area with a median series of scales which are not notably smaller than the
other scales; D. 10-11.
i. Anal with a basal sheath of scales, its margin straight or emarginate.

j. Lateral line much decurved in front, frequently several odd scales in front
between it and the next regular series; depth at least half the length; pre-
ventral area flat with sharply bent scales on the sides; profile depressed over
the eyes; occipital crest 5 of the distance from its base to the dorsal; outer
series of premaxillary teeth complete; anal long . . 1. Tetragonoptcrus Cuvier.
jj. Lateral line little decurved, parallel with the row of scales below it.

k. Scales crenulate; an enlarged scale on each side of the base of the occipital

crest 2. Entomolepis Eigenmann.

kk. Scales entire; no enlarged scale on sides of the occipital crest.

{I. Preventral area trenchant Slichonodonlinae) .

II. Preventral area rounded.

(m. A procumbent predorsal spine Slethaprioninae).

mm. No procumbent predorsal spine.

n. Second suborbital leaving a naked area between it and the lower
limb of the preopercle; at least five teeth in the inner row of the
premaxillary; outer series of premaxillary teeth in a line parallel
with the inner series except that one tooth frequently retreats

from the line of the others 3. Moenkhausia Eigenmann.

nn. Second suborbital in contact with the subopereulum below; four
teeth in the inner row of the premaxillary. Similar to Brycon-

amerieus 4. Knodus Eigenmann.

ii. Anal scaled to near its tip; caudal unequally lobed; anal margin convex.

5. Markiana Eigenmann.

hh. Predorsal line largely naked, a few scales of the sides near the dorsal with their

margins bent over the back; D. 11 or 12; ventrals in front of the dorsal; anal

emarginate or rounded 6. Gymnocorymbus ' Eigenmann.

gg. Lateral line incomplete.

o. Lower caudal lobe much the longer; maxillary without teeth.

7. Thayeria Eigenmann.
on. Caudal lobes equal or subequal.

p. Maxillary with teeth along its entire length.

8. Prislella Eigenmann.
pp. Maxillary teeth, if present, crowded on the upper anterior angle.

9. Hemigrammus Gill.
ff. Caudal fin naked except at its base.

1 Poptella (Slethaprioninae) is almost identical with this genus but contains a hidden predorsal spine.

THE AMERICAN TETRAGONOPTERINAE. 51

q. Premaxillary teeth in two series.

r. Second suborbital not in contact with the preopercle below,
or with 5 teeth in the inner series of the premaxillary.
8. Lateral line incomplete. See Psellogrammus.

/. Few teeth if any near the upper angle of the premaxillary.

u. An adipose fin 10. Hyphessobrycon Durbin.

uu. No adipose fin 11. Ease-mania Ellis.

It. Maxillary with teeth along its entire edge.

v. Anterior teeth of the lower jaw all alike, tricuspid;

anterior pair of premaxillary teeth little larger

than the rest ... 12. Hollandichthys Eigenmann.

w>. Lateral teeth of lower jaw enlarged, the fourth

usually largest; middle teeth of premaxillary

much enlarged 13. Pseudochalceus Kner.

(vvv. Anterior teeth of the lower jaw all alike, multi-
cuspid ; caudal scaled . . 8. Pristella Eigenmann) .
ss. Lateral line complete.

w. Teeth of the sides of the lower jaw more or less
abruptly minute,
x. Maxillary-premaxillary border angulated
where they meet.
y. Scales cycloid.

14. Astyanax Baird & Girard.

(z. Predorsal line scaled Astyanax.

zz. Predorsal line at least partly naked.

Poecilurichthys Gill).
yy. Scales ctenoid at least on breast; anal long,
39-47, predorsal line scaled.
A. Lateral line complete.

15. Ctenobrycon Eigenmann.
A A. Lateral line stuttering.

16. Psellogrammus Eigenmann.
xx. Maxillary-premaxillary border a continuous
curve, half as long as the head.

17. Astyanacinus Eigenmann.
ww. Teeth of the lower jaw all alike in character
and regularly graduate from in front to the
last tooth on the sides; two teeth in the
front row of the premaxillary on each side;
teeth of the second row multicuspid incisors
with a contracted base, their anterior and
posterior surfaces alike, convex, without dis-
tinct ridges. ... 18. Deuterodon Eigenmann.
ivww. Teeth of the lower jaw similar, the lateral ones
much wider than the anterior ones; maxil-
lary with two very broad incisors whose
combined width is about half the length of
the maxillary .... 19. Landonia Eigenmann.
rr. Second suborbital in contact with the lower limb of the pre-
opercle; four teeth in the inner series of the premaxillary.

52 THE AMERICAN CHARACIDAE.

B. No adipose fin ; lateral line incomplete ;
outer and middle caudal rays of the
male filiform.

20. Nematobrycon Eigenmann.
BB. Adipose fin present.

C. Anal very short, with but ten rays.
21. Microgenys Eigenmann.
CC. Anal moderate or long.

D. Outer row or premaxillary teeth
the dominant one. Gill-rakers
branched.
22. Ceralobranchia Eigenmann.
DD. Inner row of premaxillary teeth
at least as well developed as
the outer. Gill-rakers simple.
E. Few teeth along the upper por-
tion of the maxillary.
23. Bryconamericus Eigenmann.
(F. Males with a pouch covered
with scales at the base of
the caudal.

Argopkura Eigenmann.
FF. Males without a pouch on
the caudal.

Bryconamericus.)
EE. Teeth along the greater part or
along the entire edge of the
maxillary.

24. Hemibrycon Gunther.
qq. Premaxillary teeth in three series.

G. Anal short of not more
than 14 rays.
25. Crcagrutus Gunther.
GG. Anal of 20 or more rays.

26. Piabina Reinhardt.

ee. Maxillary border meeting premaxillary border at a right angle, its upper anterior margin

then describing a quarter circle and continued in a direction nearly parallel with that of

the premaxillary border; maxillary slender, partially slipping under the preorbital and

first suborbital for nearly its entire length. Plate 2, fig. 5.

H. Lateral line incomplete.
27. Brycochandus Eigenmann.
HH. Lateral line complete.
/. Premaxillary teeth

five-pointed, in an
outer row of 6-10
teeth; anterior man-
dibular teeth strong-
er than those of the
premaxillary, 10-12
in number; sides of

THE AMERICAN TETRAGONOPTERINAE. 53

the lower jaw with a

long row of very small

single-pointed teeth.

28. Creatochanes Giinther.

//. Premaxillary teeth in

three series.

29. Bryconops Kner.

(dd. Preventral area compressed, the scales of the two sides separated by a series of narrow

median scales, the marginal scales of the two sides with their lower margins straight; outer

ventral rays filiform Zygogasler Eigenmann ').

ddd. Preventral area with two series of large overlapping scales; origin of anal in advance of
the origin of the dorsal or but little behind it; premaxillary with an inner series com-
posed of several tricuspid and several conical teeth; the outer series of the premaxillary
of two or three tricuspid and usually one or more conical teeth.

/. Lateral line com-
plete; caudal
naked.
30. Phenocogasler Eigenmann.
JJ. Incomplete lateral
line.
31. Vesicatrus Eigenmann.
66. Teeth all conical; mouth very large, the lower jaw entering the profile.

32. Genycharax Eigenmann.
cc. Gill-rakers short, lanceolate; anal long (29); lateral line complete; maxillary with conical teeth
along half of its length; premaxillary and mandibular teeth as in Tetragonopterus.

33. Scissor Giinther.
aa. Upper lip not covering the premaxillary teeth.

K. Premaxillary

teeth in a main
outer series
and a minor
inner series of
two or more
teeth; a pair
of conical

teeth behind
the main series
of the man-
dibulary teeth
in front.
34. Henochilus Garman.
KK. Premaxillary

with one series
of teeth; no
conical teeth
in the lower
jaw.
35. Psalidodon Eigenmann.

1 This has been made a subgenus of Astyanax.

54 THE AMERICAN CHARACIDAE.

1. Tetragonopterus Cuvier.

rerpas four, yuwla angle, TTTepoe, to, wing = square winged.

Tetragonopterus Artedi, Seba, Locupl. rerum, 1758, 3, pi. 34, fig. 3, (argenteus).

Tetragonopterus Cuvier, Regne animal, 1817, 2, p. 166, (argenteus). Eigenmann, Rept. Princeton univ.
exped. Patagonia, 1910, 3, p. 438.

Type. — Tetragonopterus argenteus Cuvier.

Small fishes, pug-nosed, much compressed and very deep, the depth at least
half the length; humped at the occiput, concave over the eyes; interorbital
broad, rounded; snout very short, the maxillary nearly vertical; nostrils sepa-
rated by a valve; a long frontal fontanel extending beyond middle of eye;
parietal fontanel continued as a groove to the tip of the occipital crest, which
reaches £ to the dorsal; cheek largely covered by the suborbital; opercle very
short (nearly four times as high as long, in the type) ; premaxillary teeth in two
rows, the teeth of the outer series small, of nearly uniform size, the row more
or less regular, the teeth of the inner series larger, graduated, multicuspid, the
cusps of each tooth arranged in a curve, the middle cusp much the longest;
several large, graduated, several pointed teeth in the front of the lower jaw,
abruptly minute teeth on the sides; maxillary with or without teeth on its upper
anterior edge; gill-membranes entirely free from the isthmus; gill-rakers long,
slender; fins all well developed, the anal long, 32-37; scales entire, large on
the middle of the sides, becoming smaller in all directions, notably toward front
of anal 1 ; lateral fine complete, much decurved, several scales between its
origin and that of the regular series below it, caudal scaled; preventral area flat,
bounded by sharp angles, a median series of scales on the breast; seven or more
series of scales between the lateral line and the dorsal; postventral surface
trenchant; tongue thick, but little free.

Vertebrae 10 + 19; alimentary canal about 1.4 the length over all.

This genus, most nearly allied to Moenkhausia, is readily distinguished
from it by the greatly decurved lateral fine which is not parallel with the row
of scales' below it in front.

Habitat. — Orinoco and Guianas, Amazons and south to Rio de Janeiro
and La Plata.

1 In a specimen of T. argenteus the exposed edge of the 9th scale of the lateral line is about twice
as high as the exposed edge of the 4th, equal to two scales just below the beginning of the dorsal, greater
than the width of the flat ventral surface, equal to the scales of the lateral line plus the series above and
below it on the caudal peduncle, or equal to | the depth of the caudal peduncle.

THE AMERICAN TETRAGONOPTERINAE. 55

Key to the Species.

A. 36 or 37; depth 1.6-1.8 rarely 2; eye in adult less than interorbital; scales toward occiput
decreasing in size; occipital process bordered by 5-7 scales on each side; caudal lobes scaled
for about half their length; distance of dorsal from tip of snout greater than the distance of
the ventrals from tip of snout; ventrals equidistant from tip of snout and last third or last
fourth of anal; pectorals reaching beyond origin of ventrals; 12 to 16 azygous, predorsal scales.

1. argenteus Cuvier.

A. 32 or 33; depth 1.66-2; eye in adult at least equal to the interorbital; occipital process bor-
dered by 3 or 4 scales on each side; caudal lobes densely scaled to near their tip; dorsal and
ventrals about equidistant from tip of snout; ventrals equidistant from tip of snout and
end of anal; pectorals usually not reaching origin of ventrals; 8 or 9 azygous predorsal
scales. Scales 7-29 to 34-3.5; a caudal spot 2. chalceus Agassiz.

A. 32; depth 1.66; eye 2.7 in the head, a little less than the interorbital, which is 2.33 in the
head; pectorals reaching a little beyond origin of the ventrals; scales 8.5-31-4.5; caudal
scaled to its tip 3. huberi Steindachner.

A. 31-33; depth 1.66-1.8; eye 2.2-2.4 in the head, greater than interorbital; ventrals nearer
the snout than the dorsal, equidistant from snout and last fourth of the anal; pectorals reach-
ing past origin of ventrals; scales 7-28 to 29-3.5 to 4 4. gibbosus Steindachner.

1. Tetragonopterus argenteus Cuvier.
Plate 2, fig. 1; Plate 4, fig. 2.

Tetragonoplerus argenteus Cuvier, Mem. Mus. hist, nat., 1848, 4, p. 455 (Bahia?) ; Muller & Troschel,

Horae ichthyol., 1845, 1, p. 13 (Brazil & Guiana); Fische British Guiana, 1848, p. 634 (Amucu);

Cuvier & Valenciennes Hist. nat. poissons, 1848, 22, p. 132 (Bahia?) ; Kner, Characinen, 1S59,

p. 38 (Cujaba; Guiana); Gunther, Cat. fishes Brit, mus., 1864, 6, p. 318; Steindachner, Ichthyol.

beitr., 1876, 6, p. 46 (Santarem); Flussf. Sudamer., 1879, 1, p. 7 (Orinoco, near Ciudad Bolivar);

1S82, 4, p. 13 (Amazons, Iquitos); Eigenmann & Eigenmann, Proc. U. S. N. M., 1891, 14, p. 52;

Ulrey, Ann. N. Y. acad. sci., 1895, 8, p. 273; Boulenger, Trans. Zool. soc. London, 1896, 14, p.

35 (Descalvados, Northern Paraguay); Pellegrin, Bull. Mus. hist, nat., 1899, 6, p. 157 (Apurc;

Manaos) ; Boulenger, Boll. Mus. univ. Torino, 1900, 15, no. 370, p. 2 (Urucum) ; Eigenmann, Ann.

Carnegie mus., 1907, 4, p. 126, fig. 10 (Puerto Murtinho; Bahia Negra); Rept. Princeton univ. exped.

Patagonia, 1900, 3, p. 439; Mem. Carnegie mus., 1912, 6, p. 319, fig. 37 (Tumatumari) ; Fowler,

Proc. Acad. nat. sci., Phil., 1914, p. 242 (Rupununi). 1
Tetragonoplerus rufipes Valenciennes, d'Orbigny, Voy. Amer. Merid. Poissons, 1847, pi. 11, fig. 1

(Buenos Aires); Cuvier & Valenciennes, Hist. nat. poissons, 1848, 22, p. 136 (Buenos Aires);

Gunther, Cat. fishes Brit, mus., 1864, 6, p. 318 (Buenos Aires); Eigenmann & Eigenmann, Proc.

U. S. N. M., 1891, 14, p. 52; Perugia, Ann. Mus. civ. storia nat. Genova, 1891, ser. 2, 10, p. 42

(Candclaria, Rio Parana; Rio Paraguay at Asuncion); Lahille, Rev. Mus. de la Plata, 1895, 6,

p. 7 (Punta Lara, Isla Santiago); Ulrey, Ann. N. Y. acad. sci., 1895, 8, p. 272. 2

1 Tetragonopterus argenteus about 100 mm. Cuvier's type (said to contain thirty-four anal rays by
Cuvier and forty by Valenciennes) has thirty-seven and a half anal rays; dorsal broken. Lateral line
30; a dark vertical bar at origin of caudal; two vertical bars behind the head.

2 The types, three specimens, of T. rufipes in the Jardin des Plantes from Buenos Aires are in bad
condition; they are 100, 110, and 118 mm. long. Those are evidently the ones figured and described
by Valenciennes who wrote the label. They have A. 37; interorbital slightly convex. It is impossible
in their state of disintegration to distinguish them from T. argenteus, with which they are very probably
identical. I have not seen the type of T. sawa.

56

THE AMERICAN CHARACIDAE.

Tetragonopterus sawa Castelnau, Exped. Amer. Sud. Poissons, 1855, p. 65, pi. 33, fig. 1 (Rio Coxas).
Telragonopterus chalceus Eigenmann & Kennedy (non Agassiz), Proc. Acad. nat. sci. Phil., 1903, p. 523
(Rio Paraguay; Arroyo Trementina).

Habitat. — From the Orinoco and Guiana to the Paranahyba basin, Ama-
zons and Paraguay, Parana to Buenos Aires. Not in the coastwise streams
between Buenos Aires and the Paranahyba. It is rare in Guiana and the
only species in the La Plata basin.

Specimens examined.

Catalogue Number of

number

specimens

Size in mm.

Locality

Collector

18982 2

3

80-125

Tabatinga

Bourget

20793

20794
20797

15

55-125

Hyavary

Bourget

20799

820

1

119

Fonte Boa

Fletcher

20748

1

118

Fonte Boa

Agassiz

20819

7

70-130

19a

James

20745

14

65-115

Tocantins

Agassiz
f James,

20859

3

100-110

Jutahy

Thayer,
I Talisman

20822

2

100-115

San Paolo

James

20726

4

87-104

Agassiz

20728

6

69- 85

Lake Hyanuary

Agassiz

20976 1
29982 J

> 6

75-about 130

Lago Alexo

Thayer

20741

5

68- 84

Tajapurii

Agassiz

2071S

4

71- 7S

Villa Bella

Agassiz

20847

1

60

Obidos

Bentos

20776 '

20777

20786

13

49- 85

Santarem

Bourget.

20791

27729

?

9

63- 85

Monte Alegre

Agassiz

20953

1

83

Jatuarana

Navez

2978*

2. 2

69- 72

San Joaquin, Guapore basin

Haseman

2976 <

D. 4

67- 86

Santarem

Haseman

21007

1

about 90

Jose Fernandez

Coutinho

21052 1

21053 J

64-108

Rio Puty, into Rio Paranahyba St. John.

1 1 have also examined the types of T. argenteus and T. rufipes.

8 Numbers without designation indicate the Museum of Comparative Zoology; C. = Carnegie
Museum, Pittsburgh; I. = Indiana University; U. = U. S. National Museum.

THE AMERICAN TETRAGONOPTERINAE.

57

Catalogue
number

Number of
specimens

Size in mm.

21051

75»

66-107

21056

12

bad

10022 I.

10023 I.
10253 I.

1
2
3

about 90
about 56-about 85
about 82-about 95

10255 I.

S 2

about 64-100

1489 C.

1

67

3317 C.

3

24-104

3318 C.

1

33

3319 C.

35

38- 96

2977 C.
2979 C.

3

1

28- 80

55 to base of

caudal

2980 C.

4

58- 67

Locality Collector

Rio Puty St. John.

San Goncallo, Rio Paranahyba St. John.

Arroyo Trementina, Paraguay Anisits.

Asuncion, Paraguay Anisits.

Bahia Negra, Paraguay Anisits.

Puerto Murtinho, Paraguay Anisits.

Tumatumari, Brit. Guiana Eigenmann.

Asuncion, Paraguay Haseman.

Sapucay, Paraguay Haseman.

Villa Hays, Paraguay Haseman.

Corumba, Rio Paraguay Haseman.

Rio Jauru, into Rio Paraguay Haseman.

Caceres, Rio Paraguay Haseman.

This species can always be readily distinguished by its great depth and
the two vertical humeral bars. The latter, though in some cases shadowy, are
evident in all of the specimens examined.

Head 3-3.66; depth 1.6-1.8, rarely 2; D.ll; A. usually 36 or 37 3 ; scales
7 to 9-32 to 35-3 £-5- 4 Eye 2.25 in the young, 3 in the adult; interorbital less
than the eye in the young, greater than the eye in the adult.

Deep, compressed, ventral profile evenly rounded from the tip of the lower
jaw to the origin of the anal; anal basis nearly straight, very steep. Dorsal
profile depressed over the eyes, humped to the dorsal; dorsal basis straight,
oblique; postdorsal to caudal nearly straight, oblique. Preventral area with a
median series of flat or slightly keeled scales, bordered on the sides by series
of angularly bent scales; postventral area with a median series of small, narrow

1 Of these, twenty-five cf, judging by the anal armature, range from 60-90 mm., only three reach-
ing the latter size; the smallest female is SO mm. long, while thirty-six of them exceed the length of the
largest male. Of twelve taken at random six have thirty-six anal rays, four have thirty-seven, two
have thirty-eight.

2 Of these, three have thirty-five anal rays, two have thirty-six, one has thirty-eight, and two have
thirty-nine; three have eight scales between the dorsal and lateral line, four have nine; all but one have
four and one half scales between the lateral line and the ventrals, one has five. The scales in the lateral
line are thirty-two in two, thirty-three in two, thirty-four in two, and thirty-five in two.

3 Counting all rudimentary rays at the beginning and the double one at the end as one, out of fifty-
six specimens taken at random, four have thirty-four, nine thirty-five, eighteen thirty-six, fourteen thirty-
seven, seven thirty-eight, and four thirty-nine.

4 In Amazonian specimens fifteen out of eighteen had seven scales between the origin of the dorsal
and the lateral line, the other three had eight. Out of twenty-four Amazonian specimens one has thirty-
one, one thirty-two, twelve have thirty-three, nine thirty-four, one has thirty-five. Thirteen out of
eighteen have three and one half scales between the ventrals and the lateral line, five have four. In the
specimens from Puerto Murtinho there is a larger number of scales above and below the lateral line.
Three out of seven have eight, the other four have nine scales between the dorsal and lateral line; six
out of seven have four scales between the lateral line and the anal and one has five. The scales in the
lateral line do not differ,

58

THE AMERICAN CHARACIDAE.

bent scales, bordered on the sides with small asymmetrical scales, the area com-
pressed; predorsal region keeled; predorsal scales small, crowded, the median
series of scales (12-16) reaching to occipital process.

Occipital process long, one third of the distance from its base to the dorsal,
bordered by from five to seven scales on each side, the groove of the occipital
fontanel reaching to its tip; interorbital convex; second suborbital not covering
the entire cheek; maxillary equal to the distance from tip of snout to pupil.
From five to seven (rarely four to eight) teeth forming a continuous front series
of the premaxillary, the third (in one case the fourth) usually withdrawn some-
what from a straight line. Five (rarely four or six) teeth in the inner row;
maxillary with three (rarely one, two, or four) small teeth. Lower jaw with
four, or very rarely five, large teeth; the second largest, graduated to the fourth
or fifth; many small teeth on the side. 1 Gill-rakers about 9 + 13, the longest
one third the diameter of the eye.

Scales deeply imbricate, an occasional line on the scales of the sides, lines
more numerous on those near the caudal; scales of the sides continuous with
the anal sheath consisting of three or four rows of scales in front and tapering
to a single series on the last ray. Lateral line obliquely descending on the
first seven scales, then nearly straight; a well-developed axillary scale; caudal
lobes scaled for about half their length. Origin of dorsal about equidistant from
tip of snout and base of middle caudal rays, its first divided ray two and one half
times as long as the last, three times in the length. 2 Anal in the adult low, its
margin nearly straight; in the young the anterior rays are higher than the rest.
Origin of ventrals much nearer tip of snout than the dorsal, equidistant from

1 An examination of a variety of specimens gives the number of teeth in the:

first series of pre-

second series of

mandible

maxillary

premaxillary

maxillary

4 5

4 5 6 7 S

4 5 6

12 3 4

Localities

males

10

1 5 4

10

7 2

Rio Puty

females

7 3

7 3

10

115 3

Rio Puty

6

2 4

6

5 1

I^a

13

13 6 3

1 10 2

3 10

Santarem

6

1 3 2

4 2

2 4

In this table the first line of figures gives the different numbers of teeth found on the structures
mentioned above and the following lines represent the number of individuals having the given number.

2 In the specimen from Tumatumari the first divided ray is three and one half times as high as the
last ray which is two thirds the length.

THE AMERICAN TETRAGONOPTERINAE. 59

tip of snout and last fourth of anal. Ventrals reaching to anus or anal; pectorals
beyond the origin of the ventrals.

Two oblique dark bars, one from in front of the dorsal to the pectoral, the
other parallel to it from behind the tip of the occipital crest; a dusky predorsal
line continued on the first dorsal ray; a dark spot at end of caudal peduncle,
inconspicuous in the adult, sometimes extending across the entire peduncle
and base of caudal in the young. Anal sometimes margined with dusky. Dorsal
membrane thickly punctate. Otherwise bright silvery.

About twelve of the anterior anal rays of the male with hooklets turned
toward the base of the fin, about twelve of the hooklets on the middle half of
the first rays, the number decreasing backward.

Air-bladder very large, each section a cone, their bases contiguous, the
posterior section nearly twice as long as the anterior, bent downward and ex-
tending to the origin of the anal, the diameter of its large end equaling two thirds
the length of the head. Alimentary canal about one and two fifths times the
length over all.

Vertebrae 10 + 19 counting the coalesced as one. Tip of occipital process
extending much beyond the posterior face of the skull.

2. Tetragonopterus chalceus Agassiz.
Plate 4, fig. 1; Plate 98, fig. 4.

Coregonus amboinensis Artedi, Species, 1738, p. 44.

Tetragonopterus argenteus Artedi, Seba, Lecupl. rerum, 1758, 3, tab. 34, fig. 3, p. 174 (Rio Negro; Suri-
nam).

Tetragonopterus chalceus Agassiz, Spix Selecta gen. et spec. Pise. Bras., 1829, p. 70, tab. 33, fig. 1 (Brazil) ;
Cuvier & Valenciennes, Hist. nat. poissons, 1848, 22, p. 140; Kner, Characinen, 1859, p. 38
(Rio Negro; Surinam); Gunther, Cat. fishes Brit, mus., 1864, 6, p. 320 (British Guiana, Essequibo);
Cope, Proc. Acad. nat. sci. Phil., 1871, p. 260 (Ambyiacu); Steindachner, Ichthyol. beitr., 1876
6, p. 47 (Xingu, near Porto do Moz); Eigenmann & Eigenmann, Proc. N. S. N. M., 1891, 14,
p. 52; Ulrey, Ann. N. Y. acad. sci., 1895, 8, p. 227; Vaillant, Bull. Mus. hist, nat., 1899, 5,
p. 154 (Carsevenne) ; Pellegrin, Bull. Mus. hist, nat., 1899, 6, p. 157 (Apure); Fowler, Proc.
Acad. nat. sci. Phil., 1906, p. 440 (Ambyiacu; Maraiion; Pebas); Eigenmann, Rept. Princeton
univ. exped. Patagonia, 1910, 3, p. 438; Mem. Carnegie mus., 1912, 6, p. 320, fig. 38 (Wismar;
Bartica; Tumatumari; Crab Falls; Rockstone); Fowler, Proc. Acad. nat. sci. Phil., 1914, p. 242
(Rupununi).

t Tetragonopterus artedii Cuvier & Valenciennes, Hist. nat. poissons, 1848, 22, p. 128 (Surinam);
Gunther, Cat. fishes Brit, mus., 1864, 6, p. 319; Eigenmann & Eigenmann, Proc. U. S. N. M.,
1891, 14, p. 52; Ulrey, Ann. N. Y. acad. sci., 1895, 8, p. 272. 1

1 The type of this species was given to the Paris museum by the Ley den museum. It measures
68 mm. to base of caudal. Lat. line 31; A. 32 (Valenciennes says 40).

60

THE AMERICAN CHARACIDAE.

Tetragorwpterus schomburgkii Cuvier & Val., Hist. nat. poissons, 1848, 22, p. 137 (Essequibo) 1 .
Tetrugonopterus ortonii Gill, Proc. Acad. nat. sci. Phil., 1870, p. 92 (Brazil ?) ; Cope, Proc. Amer. philos.
soc, 1869, 11, p. 566 (Pebas).

Habitat. — Guianas and Amazons, Rio San Francisco.

Specimens examined. 2

Catalogue

Number of Size in

number

specimens

mm.

Locality

Collector

20747

8 .

76-118

Maues, Rio Madeira

Agassiz.

20994

1

88

Serpa

Thayer.

20714

1

65

Villa Bella

Agassiz.

20828

1

about 68

Obidos

James.

20781

8

81-92

Santarem

Bourget.

21009

1

135

?

Justa.

20923

2

62-88

San Francisco below the falls

Hartt.

20852

2

about 68

Jutahy

James, Thayer,
and Talisman.

20746

7

Porto do Moz

Agassiz.

1377 C.

118521.

4

58-68

Wismar, Demerara River

Eigenmann.

1378 C.

118531.

6

65-71

Bartica, Essquibo River

Shideler.

1379 C.

118541.

15

47-76

Turaatumari, Potaro River

Eigenmann.

1380 C.

11855 I.

18

50-91

Crab Falls, Essequibo River

Eigenmann.

1381 C.

11856 1.

4S

41-49

Rockstone, Essequibo River

Eigenmann.

2989 C.

2

65-80

Maciel, Rio Guapore

Haseman.

2990 C.

21

largest 61

Santa Rita, San Francisco Basin

Haseman.

2991 C.

13

60-76

Joazeiro, Rio San Francisco

Haseman.

2992 C.

3

54-80

Penedo, Rio San Francisco

Haseman.

2993 C.

3

52-63

(to base of

caudal)

Penedo, Rio San Francisco

Haseman.

2994 C.

6

48-81

Pirapora, San Francisco

Haseman.

2995 C.

5

35-75

Barreiras, Lagoas of Rio Grande,
Rio San Francisco

Haseman.

2996 C.

19

largest 42

Boqueirao, near mouth of Rio
Preto

Haseman.

2997 C.

31

71 '

Lagoa Pereira, Rio San Francisco

Haseman.

Head 3.2-3.66; depth l|-2; D. 11; A. usually 32 or 33. 3 Scales 7-29 to

1 The type, one specimen in bad condition, from the Essequibo, 160 mm. A. 32; scales large,
about 30; depth a little less than 2; head 3.66; eye very large, about 2.33 in the head, about equal to
the convex interorbital; maxillary with three small teeth; pectorals extending to ventrals; anal not
falcate, dorsal behind the vertical from the ventrals, not reaching the adipose. No occipital process.
The nape of the specimen seems to have met with an accident and it is difficult to determine how much
of the difference between it and the types of T. rufipes is due to bad preservation and to the mutilation.
The eye is much larger than in the type of T. rufipes. Caudal spot not continued on the middle rays.

2 1 have also examined the types of T. artedii and T. schomburgkii.

3 Out of twenty-one one has twenty-eight, one has thirty, four have thirty-one, eight have thirty-
two, five have thirty-three, and two have thirty-four anal rays.

THE AMERICAN TETRAGONOPTERINAE. 61

34 '-3^. Eye 2.2-2.3; interorbital 2.2-2.66, equal to the eye or less than the
eye in the adult.

Deep, compressed, ventral profile evenly curved to the anal, not as greatly
arched as in T. argenteus; anal basis nearly straight. Dorsal profile depressed
over eye; arched to the dorsal; dorsal basis oblique, straight; postdorsal part
of profile nearly straight. Preventral area with a median series of flat or slightly
keeled scales, bordered on the side by a series of angularly bent scales; post-
ventral area with a series of small, narrow bent scales, the area compressed;
predorsal region bluntly keeled, the scales not notably small, the median series
of eight or nine scales reaching the occipital process.

Occipital process long, one third of the distance from its base to the dorsal,
bordered by three or four scales on each side; groove of the occipital fontanel
reaching to its tip; interorbital convex; second suborbital not covering the
entire cheek; maxillary equal to the distance from tip of snout to pupil; five
or six teeth in the front series of the premaxillary (five in thirteen cases, six in
fourteen), the third tooth from the middle usually withdrawn from the line;
five teeth in the inner series of the premaxillary; usually three teeth (two in
five cases, three in eighteen, four in four) on the maxillary; lower jaw with
four large graduated teeth and many small ones on the side.

Gill-rakers about 4 + 13, the longest about one third the diameter of the
eye.

Scales deeply imbricate, striae lacking or an occasional line on the scales,
the striae fairly numerous near the caudal; scales of the sides continuous with
the anal sheath which is composed of three series of scales near its anterior end,
becoming reduced to a single series on the last rays; caudal lobes densely scaled
to near their tips in the adult, the scales much more readily caducous than those
of the sides; lateral line obliquely descending on the first seven scales then
nearly straight. A well-developed axillary scale.

Origin of dorsal equidistant from tip of snout and tip of adipose or caudal,
its highest ray three to four times as high as its lowest, two to three in the length;
anal in the adult with its margin nearly straight; the anterior rays being but
slightly prolonged; origin of ventrals equidistant from the tip of snout with
the dorsal and equidistant between the tip of snout and end of anal, reaching
to anus; pectorals usually falling short of the ventral, except in Guiana speci-
mens.

1 Of ten specimens from Lagoa Pereira one has twenty-nine, three have thirty, four thirty-one, and
two thirty-two. Of ten specimens from Roekstone one lias thirty-one, three have thirty-two, five thirty-
three and one has thirty-four.

62 THE AMERICAN CHARACIDAE.

Fresh specimens from the Essequibo River have the fins more or less suf-
fused with red. The first prolonged anal ray milk-white, the first fully formed
dorsal ray either dark or milk-white. In these specimens there is a humeral
spot prolonged above and below with a bar and a second bar fainter but evident
extends down from the dorsal parallel with the first. There is a well-marked
dark band-like spot at the base of the caudal. These approach T. argenteus
in color. The first three developed rays of the dorsal sometimes reach the
caudal, the first three anal rays are also sometimes prolonged to a length about
two thirds that of the anal base.

No humeral bars in old alcoholic specimens in which the caudal spot is
more or less obscure with age; dorsal membranes thickly peppered; general
color bright and iridescent, silvery.

Air-bladder very large, the sections conical, the posterior about twice as
long as the anterior, sharply pointed behind, its largest diameter, at its base,
equals about half the length of the head.

Alimentary canal somewhat longer than the entire fish.

Vertebrae 10 + 19; occipital process extending much beyond the posterior
face of the skull.

3. Tetragonopterus huberi Steindachner.

Tetragonopterus huberi Steindachner, Anz. K. akad. wiss. Wien, 1909, no. 12, p. 172 (Rio Pums);
Eigenmann, Rept. Princeton univ. exped. Patagonia, 1910, 3, p. 438.

Habitat. — Rio Purus, Upper Amazon Basin.

This species known only from the brief description of two specimens is said
to differ from T. argenteus chiefly in having its caudal completely scaled. It is
evidently closely allied if not identical with T. chalceus.

Head 2.7; depth 1.66; D. 11; A. 32; scales 8.5-31-4.5. Eye 2.7 in the
head; interorbital 2.33.

Belly with lateral keels; profile of head concave, that of nape very convex;
maxillary with 3-4 teeth; height of prolonged dorsal rays 3 in the length; pec-
torals little shorter than head, reaching beyond origin of ventrals; ventrals
H in the head; origin of ventrals in front of the vertical from the dorsal; pre-
dorsal area keeled; distance between dorsal and adipose equal to the head.

A row of scales along the base of the anal; traces of two dark vertical
humeral bands; caudal spot wanting in a small specimen, faint in a large one.

THE AMERICAN TETRAGONOPTERINAE. 63

4. Tetragonopterus gibbosus Steindachner.

Tetragonoptcrus gibbosus Steindachner Siisswf. sudostl. Bras., 1876, 3, p. 4, pi. 1, fig. 1 (Rio Parahyba);
Eigenmann & Eigenmann, Proc. U. S. N. M., 1891, 14, p. 52; Ulhey, Ann. N. Y. acad. sci., 1895,
8, p. 277; Eigenmann, Rept. Princeton univ. exped. Patagonia, 1910, 3, p. 438.

Habitat. — Parahyba.

This species is known only from the types in the Vienna Museum. If the
caudal is naked as the artist has drawn it and if as Dr. Steindachner says the
"Oberkiefer, unter der Loupe gesehen, am vorderen Rande dicht mit Ziihnen

besetzt " then this species does not belong to Tetragonopterus as here

understood. In the characters other than the above and those mentioned in
the key this species closely resembles the Tetragonopterus chalceus which geo-
graphically is its closest neighbor.

Head 3.6; depth 1.66-1.8; D. 10-11; A. 31-33; scales 6.5 or 7-28 or
29-3.5 to 4. Eye 2.2-2.4 in the head; interorbital 3 in the head.

Much compressed, the profile little concave over eye, rising rapidly; ven-
tral profile regular to the anal; depth of caudal peduncle 5 in the greatest depth;
mouth oblique, terminal; 10-11 mandibular teeth, with the exception of the
outer pair more than twice as long and thick as the premaxillary teeth; second
suborbital nearly covering the cheek; origin of the dorsal a little in advance
of the middle; pectorals reaching a little beyond origin of the ventrals, equal
to the length of the head in the male, considerably shorter in the female, ventral
considerably in advance of the vertical from the dorsal, not reaching anal;
basis of anal scaled; its base half an orbital diameter longer than the head.

Caudal only moderately forked and about as long as the head. Lateral
line rapidly descending on the first six scales; scales of the middle of the sides
very deep, radiae variable, 1-7; the three preventral series of scales very bluntly
keeled; the lateral keels forming the edge of the belly. Scales of the nape
apparently not decreasing in size.

A light yellow lateral band; humeral and caudal spots very faint, the former
sometimes absent.

2. Entomolepis, gen. nov.

ivToitos, cut; \ewh, i}, scale. In allusion to the crenate scales.

Type. — Tetragonopterus steindachneri Eigenmann.

Caudal scaled; lateral line but little decurved, complete; the scales cre-
nate; an enlarged scale on either side of the occipital process; maxillary with few

64 THE AMERICAN CHARACIDAE.

teeth, second suborbital leaving a narrow naked area on the cheek. This genus
differs from Moenkhausia in having crenate instead of entire scales.
Habitat. — Middle and Upper Amazon.

1. Entomolepis steindachneri (Eigenmann).
Plate 3, fig. 1-3, Plate 6, fig. 3.

Tetragonoplerus lineatus Steindachner (non Perugia), Ichthyol. beitr. 1891, 15, p. 26, pi. 2, fig. 1 (Iqui-

tos).
Tetragonoplerus steindachneri Eigenmann, Proc. U. S. N. M., 1893, 16, p. 53.

Moenkhausia steindachneri Eigenmann, Rept. Princeton univ. exped. Patagonia, 1910, 3, p. 437
fAslyanax oligolepis Fowler (non Gtinther), Proc. Acad. nat. sci. Phil., 1906, p. 439, fig. 37 (Peruvian

Amazon).

Habitat. — Middle and Upper Amazon.

Aside from the types in Vienna I have examined the following, from which
the present description is drawn:

Two specimens, 20967, 71 and 74 mm. to base of caudal. Cudajas. Thayer
& Bourget.

Head 3.6; depth 2.26; D. 10 or 11; A. 25 to 26; scales 5-32 to 33-4 or 4.5.
Eye 3-3.2; interorbital 2.4 in head.

Deep oval, without distinct humps or depressions; pre ventral area somewhat
compressed, keeled; postventral area more narrowly compressed; predorsal
area with a blunt median keel; a median series of nine scales reaching from
dorsal to occipital process.

Occipital process about one fifth of the distance of its base from the dorsal,
bordered by one or two scales, the first of which is unusually large; skull heavy
and broad; frontal fontanel nearly circular, not half as long as the occipital
fontanel; interorbital convex; second suborbital corrugate, leaving but a very
narrow naked border; interopercle distinctly visible from the side; maxillary
3.33 in head; four teeth in the outer series of the premaxillary, the middle two
close together, the third withdrawn from the line of the others; five teeth in
the inner series; one or two teeth on the maxillary; mandible with four large
graduated teeth. Gill-rakers 8 and 10, very slender, the longest two thirds of
the pupil.

Scales large, crenate, very regularly imbricate, the exposed margin of those
on the caudal peduncle three fourths as high as the highest on the middle of the
sides; the scale at the base of the occipital crest much larger than usual; each

THE AMERICAN TETRAGONOPTERINAE. 65

scale with a few widely diverging striae and numerous shorter ones along the
margin; anal sheath composed of two series of scales in front, of a single series
behind; caudal lobes scaled for about half their length; lateral line not greatly
descending, the rows of scales above and below it parallel with it; a well-devel-
oped axillary scale.

Dorsal about equidistant from snout and caudal, short and high, its height
3.66 in the length; anal margin nearly straight, but the anterior rays nearly
twice as high as the posterior; pectorals reaching ventrals, the latter not to the
anal, equidistant from tip of snout and base of last anal ray.

Brownish above, silvery on the sides; a series of dark lines between the
series of scales; a faint humeral spot just above the fourth scale of the lateral
line; a large caudal blotch not extending forward or to the end of the middle
rays.

This species can easily be distinguished by its crenate scales, compressed
breast, and peculiar coloration.

It is very probable that Fowler's Astyanax oligolepis is this species. If
the rows of scales are as Fowler figures then his specimens are distinct, differ-
ing from all other species of the group.

3. Moenkhausia Eigenmann.
For William J. Moenkhaus.

Moenkhausia Eigenmann, Smithsonian misc. coll. quart., 1903, 45, p. 145.

Type. — Tetragonopterus xinguensis Steindachner.

Small fishes differing from Tetragonopterus in the course of the lateral
line. The line is straight or but little decurved. In other characters some
of the species of this genus agrees with Tetragonopterus. The species differ
greatly in shape. The extreme in one direction are very compressed and very
deep, the depth being more than one half the length. The extreme in the other
direction are quite slender, subspindle shaped, the depth being only one fourth
of the length. The preventral area may be either narrowly rounded, as in M.
profunda, latissima, comma, and oligolepis, more broadly rounded as in M.
chysargyrea, or flat with lateral angles as in M. jamesi, megalops, et al. There
is a regular median series of scales in front of the ventrals. The anal rays range
from 18-37, in number; the scales from 22-39; the lateral line is complete except
as noted under M. australe, cotinho, and sanctae filomenae. In these species the

66 THE AMERICAN CHARACIDAE.

lateral line is complete in the specimens from some localities and incomplete
in those from other localities, or both types may be found in the same place.
The genus differs from Astyanax in having its caudal partly covered with minute
scales. It merges directly into the genus Hemigrammus which has an incom-
plete lateral line.

Habitat. — Very abundant in Guiana and the Amazons, extending south
to the Paraguay and Parana Rivers and to the Rio Doce. It has not been
recorded from the Magdalena Basin from Central America, the western slope of
South America, or the lower reaches of the La Plata.

Key to the Species.

a. Depth considerably more than half of the length; much compressed. A. 33; scales 8-34-7.

1. bondi Fowler.
to. Depth 2-2.66 in the length, sometimes 3.25 in M. costae. See also aaa. A. 23-36; Occipital pro-
cess one third to one fifth as long as the distance from its base to the dorsal.
6. Five or more scales between the lateral line and ventrals.

c. Horizontal dark lines between the rows of scales; no caudal spot; humeral spot very long,
pointed in front. Median line behind occipital process sometimes covered by the bent over
margins of the scales along the side of the compressed back. Head 3.75; depth 2; A. 31-
36; scales 5| or 6-33 to 36-6; preventral area narrowly rounded; predorsal area sharply
keeled; occipital process two sevenths of the length from its base to the dorsal; origin of
ventrals equidistant from tip of snout and origin of last third of anal.

2. latissima Eigenmann.

cc. No horizontal dark lines; seven scales between dorsal and lateral line; ventrals equidistant

from tip of snout and last anal ray, or nearer the latter; a median series of scales from the

dorsal to the occipital process.

d. A caudal spot; a large faint vertically crescentic humeral spot; mouth minute; preventral

area flat; head 4; depth 2-2.2.

e. A. 33-35; scales 7-35 to 37-5; anal margin nearly straight; maxillary short, its length

equals length of snout; premaxillary teeth three- or five-pointed; maxillary without

teeth; five graduated teeth on each side of the lower jaw; second suborbital leaving

but a narrow naked area; maxillary 4 in head; mandible 3 3. jamesi Eigenmann.

ee. A. 31; scales 7-36-6; maxillary with a single tooth; premaxillary teeth four or five
pointed; four graduated teeth on each side of the lower jaw; maxillary less than snout
in length, 5 in head; mandible 3 in head; second suborbital but two thirds the width

of the cheek 4. justae Eigenmann.

dd. No caudal spot; a well-defined longitudinally ovate or round humeral spot; preventral area

rounded; mouth large; fins large.

/. Maxillary with many teeth; eye large, 2-2.5 in the head, the profile much depressed

over the eye; interorbital less than diameter of eye; one fourth of the cheek naked;

humeral spot elongate, beginning over the second scale of the lateral line; depth

2.1-2.6; head about 4; A. 34-37; scales 7 or 8-34 to 37-6 or 7; highest dorsal ray

not reaching adipose 5. doceana (Steindachner).

ff. Maxillary with 1 or 2 teeth, equal to the distance from tip of snout to pupil; eye 2.5

THE AMERICAN TETRAGONOPTERINAE. 67

or more in the head, equals interorbital ; profile not much depressed over eye; hum-
eral spot short, nearly equidistant from opercle and dorsal; depth 2.1-2.4; A. 26-30;
scales 7-33 or 35-5 or 6; highest dorsal ray usually not reaching adipose.

6. chrysargyrea (Giinther).
///. Maxillary with two teeth; maxillary longer than the distance from tip of snout to pupil;
eye 2.5 or more in the head, equals interorbital; the profile scarcely depressed; hum-
eral spot prolonged forward to a point; depth 2-f-; A. 26; scales 6-32-5; highest

dorsal rays reaching the adipose 7. comma Eigenmann.

66. Three and a half or four scales between the lateral line and ventrals.

g. Six scales between the dorsal and the lateral line. Depth 2.3; no caudal spot, no
humeral spot; A. 26, lateral line 34; maxillary length equals distance from tip
of snout to pupil; third tooth of the anterior premaxillary series entirely withdrawn
from the line of the rest, forming a separate series; maxillary with 2 teeth; a black
line along base of anal; occipital process one fifth the distance from its base to the

dorsal 8. melogramma Eigenmann.

gg. Five scales between the dorsal and lateral line.

h. A broad black band across the anterior part of caudal. A. 23-28; preventral area

narrowly rounded (See also M. cotinho).

i. Scales 5-29 to 32-4 9. oligolepis (Giinther) .

ii. Scales 22-26, rarely 28 10. sanclae filomenae (Steindachner).

hh. Caudal without black band.

j. Some striae of the scales diverging from the middle line of each scale in nearly
opposite directions, up and down; eye 2.33-2.5 in the head, equal to the
interorbital; origin of dorsal directly over the origin of the ventrals; no
caudal spot in the adult; a round humeral spot. Anal usually 26-28,
emarginate; depth usually 2 (sometimes 2.4) scales 5-24 to 32-4.

11. grandisquamis (Mttller & Troschel).
jj. Striae of the scales not as under j.

k. Depth 2 in the length; anal not emarginate; dorsal rounded, the anterior
rays not much higher than the posterior, its origin behind that of the
ventrals. Eye 3 in the head, 1.33 in interorbital; scales 5-31—4 (to
ventrals); A. 27, its margin straight; head 3.66; depth 2; profile scarcely
depressed; maxillary with two teeth, extending somewhat beyond the
anterior margin of the eye; preventral area flat; a large caudal spot, not
extending to the end of the middle rays. 1 . . . . 12. ovalis (Giinther).
kk. Depth 2.2-2.6 in the length. Upper caudal lobe not black.

I. Eye 2.4 or more in the head.

to.. A. 30-33; scales 5.5-37-4.5; head 4.2-4.3; depth 2.4; interorbital
smaller than eye; a faint vertical humeral spot; tips of caudal

and middle rays faintly dusky 13. barhouri Eigenmann.

toto. A. 26; scales 5-32-4; head 3.33; depth 2.33; maxillary reaching to
near middle of eye; no caudal spot; humeral spot large, faint.

14. xinguensis (Steindachner).
mmm. A. 23-24; scales 5-30 to 34-3; head 3.75-^; depth 2.3-2.6. Scales
with dark margins; a well-developed humeral spot.

15. browni Eigenmann.

II. Eye 2-2.2 in head; interorbital 2.4-3; scales 5-35^; head 3.6-3.7; depth

2.5-2.66;

1 Based on type in the British museum.

68 THE AMERICAN CHARACIDAE.

n. No caudal spot; a vertical humeral spot. A. 28-30.

16. megalops Eigenmann.
nn. A median caudal spot; no humeral spot. A. 26.

17. shideleri Eigenmann.
kkk. Upper caudal lobe with an oblique black band, continued downward

across the caudal peduncle and along the basal portion of the anal.
A. 27-28, scales 5.5-32 or 33-3.5; head 3.4-3.75; depth 2.25-3.25.
Longitudinal extent of premaxillary insignificant.

18. costae (Steindachner).
aaa Depth usually more than 2.75 in the length (2.6 in M. dichroura, M. lata, and gravid M. colletli),

anal less than 28 in all but exceptional specimens of M . dichroura and M. lata,
o. Base of caudal without definite spot or band.

p. Premaxillary very short; maxillary with its anterior margin
very convex. Caudal lobes black at the tips, or with a black
band across them, their tips white; base of caudal pale;
depth usually 3(2.75); A. 25-27 rarely 28; lat. line 36 or 37
(rarely 34, 35, or 39); gill-rakers one third to one half the

length of the eye 19. dichroura (Kner).

pp. Anterior margin of maxillary not sharply and evenly curved
premaxillary longer than under p, the mouth strong.
q. Tips of caudal lobes black or a black band across the lobes.
r. A. 25; scales 5-35-3.5; depth 3.1-3.75.

20. intermedia Eigenmann.
qq. Lobes of caudal without well-defined cross-band,
s. Base of anal without a black line.
t. Base of upper caudal lobe yellow or orange, this fol-
lowed by black which fades toward the tip with water
markings.
u. Depth 3-3.5; A. 24 (21-27).

21. lepidura lepidura (Kner) .
uu. Depth 2.6; A. 26 (25-28); lateral line 33 or 34,
middle caudal rays faintly colored.

22. lepidura lata Eigenmann.
It. Upper caudal lobe not black.

v. Depth 3; A. 23 or 24; lateral line 31-33; pectorals
not reaching ventrals; upper caudal lobe and
sometimes the distal part of the other rays

dusky 23. lepidura icae Eigenmann.

w. Depth 3.4; A. 24; lateral line 37; pectorals not
reaching ventrals; caudal lobes plain.

24. lepidura hasemani Eigenmann.
vw. JDepth 4; A. 21-23; lateral line 35-36; pectorals

to ventrals; usually the middle caudal rays
and distal part of all the other rays dusky.

25. lepidura gracilirna Eigenmann.
ss. Anal with a dark (zigzag) line along its base; silvery

lateral line very narrow, bordered with dark above,
no caudal spot; upper part of cheek and opercle with
numerous chromatophores.

THE AMERICAN TETRAGONOPTERINAE. 69

w. Anal rays 23-24; depth 2.6-3.3; lateral line
33-35; a well-defined humeral spot.

26. collelii (Steindachner).
ww. Anal rays 18-20; depth 3.33-3.66; lateral line

32-34 27. copei (Steindachner).

oo. A large, conspicuous black spot on the base of the caudal; origin
of ventrals equidistant from tip of snout and base of caudal.
x. Second suborbital leaving a wide naked area;
five or six teeth in the inner series of the
premaxillary; gill-rakers about 9 and 15,
one half as long as eye; 2.5 scales between
lateral line and front of anal; middle caudal
rays jet black, the color spreading over the
base of the fin. Depth 3.25; A. 19; eye
greater than interorbital.

28. ceroa Eigenmann.
xx. Second suborbital leaving a very narrow naked
area; five teeth in the inner series of the
premaxillary; gill-rakers 7 and 9, one fifth
as long as eye; 2.5-3.5 scales between
lateral line and front of anal; a very large
and conspicuous vertically oval spot occupy-
ing nearly all the base of the caudal to near
the tip of the middle rays, bordered by milk-
white; depth 3 + ; A. 20 or 21; eye about
equal or less than interorbital.

29. cotinho Eigenmann.

1. Moenkhausia bondi (Fowler).
Plate 14, fig. 3; Plate 100, fig. 7.

Phenacogastcr bondi Fowler, Proc. Acad. nat. sci. Phil., 1911, p. 419 (Corisal, Venezuela).
Moenkhausia profunda Eigenmann, Mem. Carnegie mus., 1912, 5, p. 322, pi. 46, fig. 1 (Issorora Rubber
Plantation).

Habitat. — Northern Guiana and Venezuela.

I have been able to examine the type of M. bondi in the collections of the
Academy of Natural Science in Philadelphia and 2207 C. Type of M. pro-
funda, 51 mm. Issorora Rubber Plantation. Shideler.

Head 4; depth 1.66-1.8; D. 11; A. 33; scales 8-34-7 (9 to the anal).
Eye 2.6; interorbital 2.5.

Very much compressed; the ventral profile nearly a regular section of a
circle; dorsal profile not so greatly and less regularly arched; preventral area
very narrowly rounded, without a median series of scales, the lateral scales
small and their margins bent over the median ridge; predorsal area narrow,

70 THE AMERICAN CHARACIDAE.

scaled, but without a distinct median series, the scales along the ridge much
smaller than those of the sides.

Occipital process reaching one third of the distance to the dorsal, bordered
by about five scales on each side; interorbital slightly convex; mouth small;
maxillary with the anterior margin convex, equal to snout and anterior width
of iris or snout and half the eye. Second suborbital leaving a considerable
naked area; maxillary with two teeth, premaxillary with four teeth in the outer
series, five in the inner; mandible with four strong teeth, five-pointed, abruptly
minute teeth on the sides.

Gill-rakers about a third as long as the eye.

Scales thin, with a few feeble diverging striae, margins convex; anal with
a sheath of two rows of scales; lateral line almost straight; no interpolated
rows of scales.

Origin of dorsal considerably behind the ventrals, three in the length;
anal long, but little emarginate, its origin and base of last dorsal ray equidistant
from the snout; pectorals reaching about to middle of ventrals; ventrals not
to anal.

A diffuse humeral band followed after a light band by a fainter dark band.

This species is technically a Moenkhausia. In reality its relationship is
with Ephippicharax of another subfamily. It differs from Ephippicharax in
not having a movable predorsal spine.

2. Moenkhausia latissima Eigenmann.
Plate 6, fig. 2.

Moenkhausia latissimus Eigenmann, Bull. M. C. Z., 190S, 62, p. 101 (Tabatinga); Rept. Princeton univ.
exped. Patagonia, 1910, 3, p. 437.

Habitat. — Tabatinga.

Specimens examined.

Number of
Catalogue number specimens Size in mm. Locality Collector

207G2

22 about 55-92 Tabatinga Bourget

•iU/oy J

This species, resembling Entomolepis steindachneri in its coloration, com-
pressed preventral region, and keeled predorsal area, is easily distinguished by
its entire scales, small nuchal scale, and much longer anal fin.

THE AMERICAN TETRAGONOPTERINAE. 71

Head 3.75; depth 2 on an average; D. 11; A 33-35; ' scales 5.5 or 6-33-
36-6. 2 Eye 2.75-4; interorbital 2.5.

Very deep, compressed, profile slightly depressed over eye, ventral and
dorsal profiles about equally arched; preventral area narrowly rounded or
keeled; postventral area compressed; predorsal area sharply compressed and
keeled, with a few notched median scales, further forward the scales of the sides
are bent over the sharply compressed area; postdorsal area narrowly rounded.

Occipital process narrow and long, two sevenths in the length to the dorsal,
bordered by four scales; interorbital broad, rounded; fontanels narrow, the
anterior less than half as long as the posterior which is continued as a groove
to the tip of the occipital process; second suborbital large, striate, leaving a
narrow naked area between it and the lower limb of the preopercle; maxillary
three in head, mandible 2.33. Usually four, rarely three or five teeth in the
outer series of the premaxillary, the second and third close together, the third
removed from the line of the others, five teeth in the inner series; two, rarely
three, in one case one and in one four teeth in the maxillary; lower jaw with
four graduated teeth and numerous minute ones.

Gill-rakers 8 + 11, the longest two thirds as long as the pupil.

Scales regularly imbricate, the exposed margin of those on the caudal
peduncle but little lower than that of those on middle of the sides, the width
of the exposed part a little more than half of the height; a few divergent striae,
the margins not crenate; nuchal scales not enlarged; anal sheath composed of
two series of scales in front, of a single one behind; lateral line but little decurved,
caudal lobes scaled for at least half their length.

Dorsal about equidistant from tip of snout and base of caudal; its height
three and a half in the length, its tip when laid back removed by five scales
from the adipose. Lower caudal lobe somewhat the longer, three and a fourth
in the length; anal emarginate, origin of anal and last dorsal ray equidistant
from tip of snout ; origin of ventrals below the third scale in front of the dorsal,
equidistant from tip of snout and origin of last third of anal, reaching to or nearly
to the anal; pectorals reaching the tip of the axillary scale.

No caudal spot; a well-defined, narrow, but very long, humeral spot reaching
from above the third to above the ninth scale of the lateral fine, on the upper
part of the series of scales above the lateral fine and the lower part of the series

1 Of seventeen one has thirty-one, six have thirty-three, five have thirty-four, four have thirty-five
and one has thirty-six.

2 Qf sixteen two have thirty-three, two thirty-four, seven thirty-five, five thirty-six.

72

THE AMERICAN CHARACIDAE.

next above that. The spot is rounded behind, pointed in front, and bordered
by a lighter area; horizontal dark lines between successive series of scales.

Vertebrae 11 +19.

Posterior air-bladder bent conical, its diameter greater than that of the
eye, two and a half in its length; anterior air-bladder one and a half in the
length of the posterior.

Alimentary canal about equal to the length; containing insects and small
fishes.

3. Moenkhausia jamesi Eigenmann.

Plate 6, fig. 1, Plate 100, fig. 8.

Moenkhausia jamesii Eigenmann, Bull. M. C. Z., 1908, 52, p. 102 (I<;a; Obidos; Lago do Maximo; Taja-
puru); Eigenmann, Rept. Princeton univ. exped. Patagonia, 1910, 3, p. 437.

Habitat. — Amazon Basin.

Specimens examined.

Catalogue

Number of

Size

number

specimens

in mm.

Locality

Collector

20816 Cotypes

2

62,68

I<?a

James.

20827 Cotypes

1

53

Obidos

James.

3734 C.

12

50-68

Santarem

Haseman

3735 C.

1

65

Rio Mamore

Haseman

Head 4; depth 2.2; D. 11; A. 32-35; scales 7-35 to 37-5 (to ventrals).
Eye 2.4-2.66; interorbital equals eye.

Deep, compressed, ventral profile slightly more arched than dorsal, sym-
metrically rounded; dorsal profile slightly humped at the end of the occipital
process; preventral area flattish, with a median series of scales and a lateral,
angularly bent series; postventral area compressed, very narrowly rounded;
predorsal area narrowly rounded, with a median series of ten scales reaching
from the dorsal to the occipital process.

Occipital process moderate, bordered by four scales on each side; inter-
orbital distinctly and evenly convex; second suborbital leaving a narrow naked
area; mouth very small, maxillary short and anteriorly convex, not nearly
reaching to end of first suborbital, its length equal to that of the snout. Three
or four tricuspid teeth in the outer series of the premaxillary, the second or
third very slightly removed from the fine; five tricuspid or quinquecuspid grad-
uated teeth in the second row; five rather small, graduated teeth in each side of

THE AMERICAN TETRAGONOPTERINAE. 73

the lower jaw, followed by a few minute ones on the side; maxillary without
teeth.

About 8 + 12 gill-rakers, about one third as long as the eye.

Scales deeply imbricated, with a few divergent striae; anal sheath of one
or two series of scales which are well demarked from the lateral scales; caudal
lobes scaled to near their tip; lateral line scarcely decurved; the rows of scales
above it and below it parallel with it.

Origin of dorsal nearer the tip of the snout than the base of the caudal;
origin of ventrals a little nearer to the tip of snout than the dorsal, equidistant
from tip of snout and about end of anal; highest dorsal ray a little more than
three in length, the shortest more than two and a half times in the longest. Anal
distinctly emarginate in front, the fourth ray two and three tenths times as
long as the fourteenth; ventrals scarcely reaching anal, pectorals to ventrals.

A dark vertical caudal spot on the base of the caudal, sometimes on base
of all but the outermost rays; the spot not continued on the middle rays; a
silvery lateral band half as wide as the eye; an ill-defined vertical humeral
spot crossing the space between the fourth and seventh scales of the lateral line;
no other dark markings; iridescent, silvery, and brassy.

4. Moenkhausia justae Eigenmann.

Moenkhausia justae Eigenmann, Bull. M. C. Z., 1908, 52, p. 102 ( ); Rept. Princeton univ. exped.

Patagonia, 1910, 3, p. 437.

One specimen 21014 Type 60 mm.

This species is very similar to M. jamesi from which it differs in having a
tooth on the maxillary; the teeth four- or five-pointed, four teeth on each side
of the lower jaw. A. 31: scales 7-36-6; the second preorbital much narrower
than in M. jamesi.

The exact locality is unknown. The specimen came with others from
Dr. Justa through Major Coutinho and was probably found in the neighborhood
of Manaos.

5. Moenkhausia doceana (Steindachner).

Plate 14, fig. 1.

Tetragonopterus doceanus Steindachner, Susswf. sudostl. Bras., 1876, 3, p.. 14 (Rio Doce); Eigenmann

& Eigenmann, Proc. U. S. N. M., 1895, 14, p. 52; Ulkey, Ann. N. Y. acad. sci. 1895, 8, p. 277.
Moenkhausia doceana Eigenmann, Rept. Princeton univ. exped. Patagonia, 1910, 3, p. 437.

Habitat. — Rio Doce, Rio Mucuri.

74

THE AMERICAN CHARACIDAE.

Specimens examined.

Catalogue

Number of

Size

number

specimens

in mm.

Locality

Collector

20909

38

G9-106

Rio Doce

Hartt & Copeland,

20914

4

01-70

Porto Alegre

Hartt & Copeland

20879 \

20880 J

12

04-106

Sao Matheos

Hartt & Copeland.

This species has hitherto been known from the types only. It is the only
species of Moenkhausia in the coastwise streams of eastern Brazil, south of the
Rio San Francisco.

There can be no doubt about the identification, although Steindachner says
that the entire maxillary border is toothed.

Head 3.7; depth 2 + -2.4; D.ll (rarely 12); A. 34-38 l ; scales mostly
removed, 7 or 8-34 to 37-6 or 7. Eye 2.14-2.5; interorbital 2.7-3.25.

Much compressed, the ventral profile regularly arched from the chin to
the end of the anal; dorsal outline arched as much as the ventral, but less
regularly; the occipital process rising rapidly, the profile of the head corre-
spondingly concave; preventral area rounded, with a median series of scales;
predorsal area trenchant, with a median series of about eleven scales.

Occipital process a little less than one third of the distance from its base to
the dorsal, bordered by about five scales on each side; interorbital convex;
frontal fontanel not much shorter than the parietal exclusive of the groove on
the occipital process; second suborbital leaving a naked area about one fourth
as wide as the covered portion ; length of maxillary equals the distance from tip
of snout to pupil; mandible equals distance from tip of snout to end of maxil-
lary; usually five teeth in the front row of the premaxillary, the third tooth
removed from the line, more rarely three or four teeth; five, more rarely six,
teeth in the second series; maxillary most frequently with five teeth; of the
maxillaries examined two have three, four have four, twelve have five, six have
six, six have seven, and two have eight teeth; mandible with four large teeth,
frequently a smaller one on the side, and then a series of minute ones.

Gill-rakers about 8 + 13, one third the diameter of the eye.

Scales caducous, entire, with a few diverging striae, regularly imbricate;
anal sheath of a single series of scales; caudal lobes scaled for a third of their
distance at least; lateral line very little decurved, the scales below it parallel
with it, a few auxiliary scales over the muscles of the anal.

1 Of seventeen specimens examined two have thirty-four, one thirty-five, six thirty-six, seven thirty-
seven, and one thirty-eight rays.

THE AMERICAN TETRAGONOPTERINAE. 75

Origin of dorsal a little in advance of the middle, its height four in the length;
caudal large, the lower lobe slightly longer, about one third of the length; anal
basis equals distance from dorsal to caudal, its origin equidistant from tip of
snout and base of middle caudal ray; anal emarginate, the highest anterior ray
twice the height of the fifteenth ray; ventrals reaching beyond origin of anal,
equidistant from tip of snout and last third of the anal; pectorals reaching to
middle or the second third of the ventrals.

A large, horizontally oval, humeral spot, nearly two thirds as long as the
head; a silvery lateral stripe not much more than half the width of the pupil;
general color, including all the fins dark, the fins with numerous pigment cells.

Vertebrae 12 +20.

Alimentary canal not equal to the total length. Contains ants in the one
dissected.

Measurements of eleven specimens M. C. Z. 20909.

Premax-

Maxil-

Inter-

illury

lary

orbital

teeth

teeth

Specimen Dorsal Anal Length Depth Head Eye

mm. mm. mm. mm. mm.

1 11 34 SO 3S 21 S.5 8 5'-5 2 5M3 4

2 11 36 81 38 20 8 7 5-5 5-1

3

11

37

80

35

19

8

7

5-5

3-7

4

11

36

78

33

19

S

6.5

5-5

6-7

5

12

37

52

22

14

6.5

4.5

5-5

5-6

6

11

36

56

22

15

6

5

5-6

5-5

7

11

37

49

20

13

6

4.5

[4-5
I 3-6

7-5

8

11

36

48

20

13

6

4

4-5

7-7

9

11

37

69

29

IS

8

6

5-5

6-5

10

11

39

64

26

10

7

5

4-5

4-4

11 11 36 59 25 15 7 5 5-5 8-7

6. MOENKHAUSIA CHRYSARGYREA (Giillther).

Plate 6, fig. 3; Plate 100, fig. 9, 10.

Tetragonopterus chrysargyreus Gunther, Cat. fishes Brit, mus., 1864, 5, p. 328 (Essequibo); Eigenmann
& Eigenmann, Proc. U. S. N. M., 1891, 14, p. 53; Ulrey, Ann. N. Y. acad. sci., 1895, 8, p. 281.

Moenkhausia chrysargyrea Eigenmann, Rept. Princeton univ. exped. Patagonia, 1910, 3, p. 45. Mem.
Carnegie mus., 1912, 5, p. 323.

Moenkhausia chrysargyrea leucopornis Fowler, Proc. Acad. nat. sci. Phil., 1914, p. 245 (Rupununi).

Habitat. — Essequibo, Amazons.

1 Front row. ! Second row. 3 On one maxillary. * On the other.

76

THE AMERICAN CHARACIDAE.

Specimens examined.

Catalogue

Number of Size

number

specimens in mm.

Locality

Collector

3070 C. 118411.

43

51-61

Rockstone, Essequibo River

Eigenmann.

1371 C. 118421.

7

51-91

Crab Falls, Essequibo River

Eigenmann.

1372 C. 118481.

15

53-69

Konawaruk, Essequibo River

Eigenmann.

1366 C.

1

58

Warraputa Cataract, Esse-
quibo River

Eigenmann.

1373 C. 118441.

25

52-101

Packeoo Falls, Essequibo River

Eigenmann.

1374 C. 118451.

10

60-87

Tumatumari, Potaro River

Eigenmann.

1375 C.

1

60

Wismar, Demerara River

Eigenmann.

1376 C. 118461.

2

58-60

Rupununi River

Grant.

20754

3

About 45-

70
About 60

Tabatinga

Bourget.

20705

1

Teffe

Agassiz.

20851

2

About 68
and 75

Jutahy

James, Thayer,
and Talisman.

21003 1

5

About 56-

Jose-Fernandez

Coutinho.

21005 J

54

Head 3.5-4; depth 2.4 in young, 2 in old; D. 11; A. 27-30; scales 7-33
to 34-5. Eye 2.5-2.75 in head; interorbital equals eye.

Deep, compressed, subrhomboidal, the anal basis straight, nearly parallel
with the upper anterior profile in the young, more oblique in the oldest; profile
depressed slightly over the eyes and rounded above the occipital process. Pre-
ventral area rounded, with a median series of scales, the lateral series not
sharply angulated except just in front of the ventrals; predorsal area keeled,
with a median series of about 10 scales from the dorsal to the occipital crest.

Occipital process long, equal to one third of the distance from its base to
the dorsal, bordered on the sides by three scales; second suborbital leaving a
narrow naked area which is widest below; maxillary not nearly reaching end of
first suborbital, but little beyond anterior margin of eye, its length equal to
distance from tip of snout to pupil. Four or five teeth in the outer series of
the premaxillary; five or six five-pointed, graduated teeth in the inner series;
maxillary with one or two minute teeth; lower jaw with four large graduated
teeth on each side and many minute ones.

Gill-rakers about 6 + 14, one third as long as eye.

Scales deeply imbricate, with several divergent striae; anal sheath of two
scales in front; caudal lobes scaled for more than half their length. Lateral
fine somewhat descending, parallel with the row of scales below it.

Origin of dorsal equidistant from tip of snout and base of caudal, its highest

THE AMERICAN TETRAGONOPTERINAE. 77

ray three in the length; ventrals a little nearer to tip of snout than the dorsal,
about equidistant from tip of snout and end of anal; ventrals reaching to anal
somewhat shorter in old specimens; pectorals reaching two scales beyond
ventrals; anal emarginate, the highest ray about two thirds the length of the
base. Iridescent silvery. A deep lying, well-defined, horizontally oval, circular
or rhomboidal, black, humeral spot over the space between the fifth to the eighth
scale of the lateral line and at least the width of a scale removed from it; a few
scattered cells nearer the surface extend from the front of the spot across
the lateral line; humeral spot surrounded by a light court behind which there is
a faint vertical bar; a narrow silvery lateral fine, its width equal to about one
fourth the length of the maxillary; no caudal spot; sides variously dotted with
chromatophores. In the Tumatumari specimens the scales of the abdomen are
margined with dark, there being faint streaks following the rows of scales. Scales
of the second row below the dorsal each with an oblique dark band noticeable
only with a lens; outer margin of pectoral and ventral dark; middle of anal
lobe and bases of the remaining rays yellow; dorsal rays each tinged with yellow
at its middle, pectoral tinged with yellow; upper part of iris red. Vertebrae
11 + 19.

Posterior section of air-bladder about twice as long as anterior; its width
about equal to the eye; bent down behind and reaching to near the origin of the
anal.

7. Moenkhausia comma Eigenmann.

Plate 6, fig. 2.

Moenkhausia comma Eigenmann, Bull. M. C. Z., 1908, 62, p. 102 (Cudajas); Rept. Princeton univ.
Exped. Patagonia, 1910, 3, p. 437.

One specimen 20972 part Type 77 mm. Cudajas Thayer and Bourget.

A second specimen in very bad condition of the same length and from the
same place.

This species can readily be distinguished by its elevated dorsal, and the
peculiar humeral spot.

Head 3.4; depth 2 + ; D. 11; A. 26; scales 6-32(?)-5. Eye 2.5; equal to
interorbital.

Deep, compressed, subrhomboidal, the anal basis much steeper than the
predorsal profile. Profile very slightly depressed over the eye. Preventral
area narrowly rounded, the lateral series of scales without a distinct bend.
Postventral area compressed; predorsal area narrowly compressed and keeled,
with eight median scales between occipital process and dorsal.

78 THE AMERICAN CHARACIDAE.

Occipital process long, equal to one third of the distance between its base
and the dorsal, bordered on the side by three scales. Second suborbital leaving a
naked border distinctly wider than in M . chrysargyreus; maxillary rather longer
than the distance from tip of snout to pupil, a little more than three in the head.

Three to six teeth 1 in the front row of the premaxillary, five or six in the
second row; maxillary with two small teeth; mandible with four large teeth
and numerous small ones on the side.

Gill-rakers 7 + 15, a little more than half the length of the pupil.

Scales regularly imbricate, each with a few divergent striae, the exposed
portion of those on the caudal peduncle about two thirds of the depth of the
exposed portions of those on the middle of the sides; anal sheath composed of
two scales in front ; (all but one of the caudal scales missing) ; lateral line but
little decurved.

Origin of dorsal a little nearer tip of snout than to caudal, the anterior rays
elevated, three in the length, reaching adipose when depressed; caudal lobes
equal to height of dorsal; origin of anal equidistant from tip of snout with a
point midway between the dorsals, the margin damaged but evidently emargi-
nate, the highest anterior ray reaching at least to the base of the seventh from
the last when depressed; ventrals reaching anal, one and a half in the head,
their origin equidistant from tip of snout and caudal; pectorals reaching beyond
origin of ventrals.

A deep lying, well-defined humeral spot, beginning in a point above the
origin of the lateral line increasing in width backward and ending above the
seventh scale of the lateral line; a very narrow, silvery lateral line; upper poste-
rior part of the interradial membranes of the dorsal dark; remaining fins hyaline.

Vertebrae 11 +19.

8. Moenkhausia melogramma Eigenmann.
Plate 6, fig. 1; Plate 100, fig. 4.

Moenkhausia melogrammus Eigenmann, Bull. M. C. Z., 1908, 52, p. 102 (Tabatinga); Rept. Princeton
univ. exped. Patagonia, 1910, 3, pt. 4, p. 437.

One specimen 20825 Type 40 mm. to base of caudal Tabatinga Bourget

Readily distinguished by the depth and by the black line along the base
of the anal.

1 In the type there are four on the left side and five on the right, hut one has evidently been lost
from the latter; in the second specimen there are five on the left and three on the right side.

THE AMERICAN TETRAGONOPTERINAE. 79

Head 4; depth 2.5; D. 11, counting the divided ray as 2; A. 26; scales
6-34-4. Eye equals twice the length of the snout, 2.4 in the head; interorbital
3 in the head.

Compressed, dorsal and ventral profiles nearly equally curved, the ventral
more regularly; profile slightly depressed over eye; preventral area rounded,
postventral area compressed; predorsal area slightly keeled, with a median
series of ten scales from dorsal to occipital crest.

Occipital process a little more than one fifth of the distance between its
base and the dorsal, bordered by three scales on each side; interorbital very
slightly convex; second suborbital leaving a naked area on cheek which is
widest below; maxillary not reaching to the end of the first suborbital, equal
to the distance from tip of snout to pupil. Five teeth in the front row of the
premaxillary, the third entirely withdrawn from the line of the rest; five gradu-
ated teeth in the inner series; two teeth on the maxillary; four graduated, large
teeth on each ramus of- the lower jaw followed by smaller ones on the side.

Scales deeply imbricate, with a few not greatly divergent striae; anal
without a sheath (in this specimen only ?) ; caudal lobes covered with caducous
scales for at least half their length; lateral line scarcely descending in front.
Origin of dorsal equidistant from tip of snout and base of caudal, distinctly
further from tip of snout than the ventrals; ventrals equidistant from tip of
snout and base of penultimate anal ray, reaching not quite to anal, pectorals
to ventrals. Anal probably emarginate.

A black line along the base of the anal; a faint dark line along the sides,
otherwise faintly silvery without spots.

9. Moenkhausia oligolepis (Giinther).
Capaule of the Potaro Indians.
Plate 7, fig. 4; Plate 95, fig. 4; Plate 100, fig. 5.

Tetragonoplerus oligolepis Gunther, Cat. fishes Brit, mus., 1S64, 5, p. 327 (British Guiana), Eigen-
mann & Eigenmann, Proc. U. S. N. M., 1891, 14, p. 53; Ulrey, Ann. N. Y. acad. sci., 1S95, 8,
p. 282; Vaillant, Bull. Mus. hist, nat, 1899, 5, p. 155 (Carnot).

Moenkhausia oligolepis Eigenmann, Rept. Princeton univ. exped. Patagonia, 1910, 3, p. 437; Mem.
Carnegie mus., 1912, 6, p. 321, pi. 46, fig. 3.

Tetragonoplerus agassizil Steindachner, Ichthyol. beitr., 1876, 6, p. 41, pi. 8, fig. 2 (Tabatinga; Cuda-
jas; Hyavary); Cope, Proc. Amer. philos. soc, 1878, 17, p. 691 (Peruvian Amazon); Eigenmann &
Eigenmann, Proc. U. S. N. M., 1891, 14, p. 53; Ulrey, Ann. N. Y. acad. sci., 1895, 8, p. 281.

fAslyanax atahualpuanus Fowler, Proc. Acad. nat. sci., Phil., 1906, p. 436, fig. 36 (Pebas). 1

f Moenkhausia alahualpiana Eigenmann, Rept. Princeton univ. Exped. Patagonia, 1910, 3, p. 437.

•This species is said to have: Head 3.3; depth 2.75; D. 12; A. 24; lat. 1. 26-27.

80

THE AMERICAN CHARACIDAE.

Habitat. — Amazons and north to Guiana.

I have examined the types of T. oligolepis and there is no doubt but that
they are identical with T. agassizii.

Specimens examined.

Catalogue

Number of

Length

number

specimens

in mm.

Locality

Collector

2075G, 20759,

106

41-95

Tabatinga

Bourget

20761, 20775

20854

1

59

Jutahy

James, Thayer,
and Talisman

20S08

1

85

Manacapuru

James

20729

8

about 50-66

Lake Hyanuary

Agassiz

20G93

8

54-65
(to base of caudal)

Goyaz

20966, 20972, 20963,

118

45-85

Cudajas

Thayer and

20974, 20975

Bourget

21018

81

59-90

Curupira

Coutinho

1361 C,

11835 1.

30

35-96

Holmia, Upper Potaro

Eigenmann.

1362 C,

118361.

11

39-118

Aruataima Cateract,
Upper Potaro

Eigenmann and
Grant

1363 C,

118371.

11

33-53

Potaro River, 2 hrs. be-
low Holmia

Eigenmann

1364 C,

118381.

7

28-44

Savannah Landing, Upper Eigenmann

Potaro

1365 C,

11339 I.

3

37-61

Erukin, a tributary of the Eigenmann

Potaro below Amatuk.

1367 C.

1

88

Potaro Landing

Eigenmann

1368 C,

118401.

8

40-47

Tumatumari

Eigenmann

1369 C.

1

78

Maripicru, branch of
Ireng.

Grant.

Head 3.64; depth 2.25 on an average; D. 11; anal most frequently 25
or 26 '; scales 5-30 to 31^4 2 . Eye 2.5-3 in the head; interorbital wider than
eye.

Rather deep, compressed, especially above the anal; ventral profile some-
what more arched than the dorsal, no distinct humps or depressions, the out-
line subrhomboidal. Preventral area narrowly rounded or with an indistinct
median keel, no lateral keels except in the very largest; postventral area com-
pressed, with a median series of bent scales, the scales bordering it not much
smaller than those above it ; five scales from the median scale to the lateral line ;

1 Of fifty-two specimens, two have twenty-three, six have twenty-four, nineteen have twenty-five,
fourteen have twenty-six, ten have twenty-seven, and one has twenty-eight.

2 Of fifty specimens among those in the M. C. Z., six have twenty-nine, twenty-five have thirty,
sixteen have thirty-one and three have thirty-two scales in the lateral line.

THE AMERICAN TETRAGONOPTERINAE. 81

predorsal area rounded, with an obscure median keel; eight scales between the
dorsal and the occipital process; postdorsal area rounded.

Occipital process equal to about one fifth the distance from its base to
the dorsal, bordered by two or three scales on the sides; interorbital broad,
but little convex, the frontal fontanel narrower and about half as long as the
parietal; second suborbital very broad, covering the entire cheek or leaving a
very narrow naked border behind; maxillary but little longer than snout,
more than three in the length of the head ; four or five teeth in the front row of
the premaxillary, the third withdrawn from the line, the second and third and
sometimes the fourth close together; five teeth in the inner series; usually two
teeth on the maxillary; mandible with four large teeth and a number of minute
ones on the sides.

Gill-rakers about 8 + 11, a little more than half the length of the pupil.

Scales very regularly imbricate, the exposed edge of those on the caudal
peduncle about two thirds as high as that of the largest scale on the middle
of the sides; scales above the lateral line in specimens 50 mm. long with about
four nearly parallel striae, in the largest with 6-10 striae, frequently a notch
in the margin of the scale at the end of a line; anal sheath composed of two
or three series of small scales in front of a single series behind; caudal lobes
scaled to near their tip; lateral fine nearly straight from its sixth scale; axil-
lary scale well developed.

Dorsal equidistant from tip of snout and base of caudal, its highest ray
nearly four in the length, its eighth ray about two thirds of the height of the
second. Anal emarginate in the young, the highest ray reaching base of fifth
from the last; not emarginate in the adult but the anterior rays two and a
half times as long as the one next to the last; origin of the ventrals equidistant
from tip of snout and tip of last anal ray, nearly or quite reaching the anal
in the young; pectorals beyond origin of ventrals for one or two scales.

Dorsal, adipose, ventrals, and anal pink to orange; upper part of iris red,
the rest golden. A broad black band across the end of the caudal peduncle
and base of caudal, distal part of all the caudal rays light pink; in life a faint
vertically oval humeral spot; no silvery lateral band; color otherwise variable
in intensity with the locality, the margins of all the scales dark, the centers light ;
in life the younger ones have the adipose and caudal peduncle bright yellow.

Vertebrae 12 + 16.

Posterior air-bladder as wide as and three times as long as eye, twice as
long as the anterior, curved down behind to near the origin of the anal.

Alimentary canal not quite equal to the length, without the caudal.

82

THE AMERICAN CHARACIDAE.

10. Moenkhausia sanctae filomenae (Steindachner) .
Plate 16, fig. 1, 2.

Telragonopterus agassizii Boulenger (non Steindachner), Bull. Mus. univ. Torino, 1895, 10, no. 196,

p. 3 (Colonia Risso) ; Trans. Zool. soe. London, 1896, 14, p. 35 (Descalvados and Monte Sociedad) ;

Boll. Mus. univ. Torino, 1897, 12, no. 279, p. 4 (San Lorenzo); 1900, 15, no. 370, p. 2 (Urueum).
Poecilurichlhys agassizii Eigenmann, Proc. Acad. nat. sci., Phil., 1903, p. 522 (Arroyo Trementina;

Arroyo Chagalalina).
Moenkhausia agassizii Eigenmann, Ann. Carnegie mus., 1907, 4, p. 138, pi. 12, fig. 2 (Corumba).
Tetragonoplerus sanctae filomenae Steindachner, Anz. K. akad. wiss. Wien, 1907, p. 82 (Sancta Filomena,

Parnahyba). 1
Moenkhausia australe Eigenmann, Bull. M. C. Z., 1908, 62, p. 102.

Habitat. — Paraguay Basin, Rio Parnahyba, and southern Madeira.

Specimens examined.

Catalogue

Number of

number

specimens

Size in mm.

Locality

Collector

9991 I.

1

41
(to base of caudal)

Arroyo Trementina

Anisits

9992 I.

1

32
(to base of caudal)

Arroyo Chagalalina

Anisits

3315 C.

2

41-68
(to base of caudal)

Jauru

Haseman

2998 C.

1

55

Barreiras, Lagoas of Rio Grande

Haseman

3307 C.

2

39^9

Cacequy

Haseman

2999 C.

1

49

San Francisco

Haseman

3303 C.

10

33-66

Rio Sapon, Prazer, Bahia

Haseman

3304 C.

4

32-69

Above Cachoeira da Velha, Rio
Novo, Goyaz

Haseman

5305 C.

1

47

Rio Tiete above the fall

Haseman

3306 C.

1

34

Salto das Cruzes, Rio Tiete

Haseman

3308 C.

10

36-43

Arequa, Lake Ipacarary, Paraguay

Haseman

3310 C.

2

42-43

Corumba

Haseman

3309 C.

4

40^5

Puerto Suarez

Haseman

3311 C.

22

36-65

San Luiz de Caceres

Haseman

3312 C.

8

32-67

San Luiz de Caceres

Haseman

1 The following is an abstract of Steindachner's description of the types: —

Head 3.6-3.5; depth 2.25-2.2. A. 25-27; scales 4.5-22 to 24-3.5; eye nearly 3 in head; inter-
orbital 2.33-2.

Interorbital slightly convex; snout equals eye; maxillary not reaching center of eye; mandibular
teeth brown, darkest at tip; two small teeth on maxillary.

Origin of dorsal equidistant from caudal and front of eye; ventral in front of vertical from dorsal;
pectoral reaching ventral, ventral not to anal.

Lateral line frequently interrupted, skipping several scales in the posterior part of the body.

Silvery dark above; scales margined with darker; humeral spot not sharply demarked, rounded
or oval; caudal spot deep, dark brown, very large, forming a crossband in base of caudal.

THE AMERICAN TETRAGONOPTERINAE.

83

Catalogue

Number of

number

specimens

Size in mm.

Locality

3313 C.

2

42-43

Rio Santa Rita

3314 C.

6

40-50

Rio Boa Ventura

3315 C.

5

39^7

San Joaquin

Collector
Haseman
Haseman
Haseman

This species differs from the Amazonian M. oligolepis in having usually
but twenty-five scales in the lateral line instead of thirty or thirty-one.

In many of the smaller specimens the lateral line is either interrupted, i. e.
it skips a scale or more, or it is incomplete. The incompleteness is positively
correlated with the shorter lateral line. This is especially well brought out in
the specimens from San Luiz de Caceres. These specimens with a complete
lateral line have usually thirty scales in the lateral line, while those with an
incomplete lateral line have usually only twenty-five though of the former
one has but twenty-five and of the latter one has as many as twenty-eight scales
in the lateral line. Out of eighty-four specimens examined twenty-nine have
imperfect lateral lines.

The count of the anal rays and lateral lines is given in detail for differ-
ent localities in the first part of the following table, and summarized in the
second part of the table for those specimens of both M. oligolepis and sanctae
filomenae in which the character of the entire line could be determined.

Anal rays

Scales in the lateral line

Locality

Species.

22

23

24

25

26

27

28

22

23

24

25

26

27

28

29
6

30

25

5
30

31
16

1
17

32

oligolepis
sanctae filomenae
sanctae filomenae
sanctae filomenae
sanctae filomenae
sanctae filomenae
australe
australe
australe
australe
australe
oligolepis?
australe

3

2
6
1

6

19

14

10

1

?

Rio Sapon
Rio Tiet6

1

1
1

4

3
1

1
1

1
3

2

1

1
1
4
1
2

1

1

4

35

Cacequy

1

1

2
1
1
1
3

1
1

1

2

2

2
2
1
1
1
5
18

Arequa

1
1

1

Rio Boa Ventura

1

1

4

3

1
14

3

2

21

1

1
7

1

1

6

Totals

15

12

1

1

5

8

3

Correlation between completeness of the lateral line and anal and number of scales in the lateral line.

Lateral line complete
Lateral interrupted
Lateral line incomplete

2

1
2

9
2

4

8
2

6

9

1

11

15

12

1
1

2
2
2

3

2

2

7
2

12

2
1
4

6
1

29

17

1

84 THE AMERICAN CHARACIDAE.

In the specimens with the lateral line imperfect the scales along the lateral
line series are as follows, the scales with pores being in italics: — '

Corumba 10 + 15; 8 + 16; Puerto Suarez 8 + 14; Caceres 22 + 1 + 49;
22 + 3+3; 8 + 17; 10 + 15; 9 + 16; 10 + 16; 9 + 16; 8 + 17; Rio
Boa Ventura 8 + 16; 10 + 16; 10 + 1Q; 3 + 1 + 8 + 16; 9+ 17; Rio
Sapon 6+18; 19+2+2; 12 + 1+3; Cachoeira 12+2+9 + 1; 14 +
2 or 3 + 9; 15 + 1+8 + 1; Rio Tiete" 13 + 11 and two small ones with
short lateral line; Caceguy 15 + 3+7 + 1; Arequa 8 + 16; 10 + 15; 10 +
15; Rio Jauru 11 + 14; San Joaquin 8 + 17; 10 + 17 (and three others with
an incomplete line). In the other specimens the lateral line is complete.

Summarizing the data we find that there are two groups of specimens.
The first consist of one with six scales with pores, seven with seven scales, three
with nine scales, eight with ten scales, one with eleven and one with thirteen
scales with pores. In each of these cases the remaining scales of the series are
without pores.

To this series also belongs one specimen in which the lateral line series con-
sists of three pores, one scale without a pore, three with pores and sixteen with-
out pores. This is evidently a modification of seven scales with pores and
sixteen without pores. Most of these come from the Paraguay and Parana
Basins and represent M. australe.

In the remainder of those enumerated the modifications are of another sort.
The lateral fine reaches the caudal or within one scale of the caudal, but is
interrupted, skipping one, two, or three scales. All of these with the exception
of two from Caceres came from southeastern Brazil and represent M. sanctae
filomenae. It is scarcely possible that they indicate the road from the com-
plete lateral line to the incomplete, for in those with the incomplete lateral line
the number of scales at the beginning of the line with pores is usually eight to
ten, and specimens having this character occur at the same locality with others
with complete lateral lines. In those with the interrupted lateral line there
are from fourteen to twenty-two scales at the beginning of the line with pores,
and while these also occur along with others with complete lateral line, they
in general are found in other river basins in an area geographically widely sepa-
rated from the area in which those with incomplete lateral lines occur.

There seem to be variations (constantly occurring or mutations) in two
different directions, one of these directions leads to Hemigrammus which is a

1 1 am indebted to Mrs. Marion Durbin Ellis for the examination of these specimens in connection
with her study of Hemigrammus and Hypheasobrycon.

THE AMERICAN TETRAGONOPTERINAE.

85

Moenkhausia with an incomplete lateral line, the other leads towards the condi-
tion found in Psellogrammus in which the lateral line is always interrupted but
continued to the caudal.

11. Moenkhausia grandisquamis (Miiller and Troschel).
Plate 12, fig. 3; Plate 100, fig. 6.

Tetragonoplerus grandisquamis Muller & Troschel, Horae ichthyol, 1845, 1, p. 27, tab. 8, fig. 2 (Suri-
nam) ; GtJNTHER, Cat. fishes Brit, mus., 1864, 6, p. 324 (British Guiana) ; Eigenmann & Eigen-
mann, Proc. U. S. N. M., 1891, 14, p. 53.

Moenkhausia grandisquamis Eigenmann, Rept. Princeton univ. exped. Patagonia, 1910, 3, p. 437;
Mem. Carnegie mus., 1912, 6, p. 323; pi. 77, fig. 2.

Habitat. — Guianas, Amazons.

Catalogue Number of
number specimens

20694 19

20811, 20815 7

20981, 20979 3

20977

20875 1

20988 2

20707

20879 (part) 2

20830, 20879 (part) 4

20844 6

20739 3

20014 2

20725 14 '

3736 C. 6

1348 C, 118571. 51

1349 C, 118581. 66

1350 C, 118591. 35

1351 C, 118601. 8

1352 C. 1

1353 C. 1
3000 C. 1

Specimens examined.

Length in mm. Locality Collector

about 64-91 Goyaz Sn. Honario

40-85 lea James

70-97 Lago Alexo Thayer

66 Manacapuru James

45-52 Serpa Thayer

47-53 Villa Bella Agassiz

44-51 Obidos James

44-53 Obidos James

16-84 Obidos. Col. Bentos

60-75 Monte Alegre Agassiz

74 Justa

poor Gurupa Agassiz

48-52 Santarem Haseman

45-79 Bartica, Essequibo River Shideler

51-86 Rockstone, Essequibo River Eigenmann

50-115 Crab Falls, Essequibo River Eigenmann

64-110 Tumatumari, Lower Potaro Eigenmann

122 Cangaruma, Lower Potaro Eigenmann

55 Wismar, Demerara R. Eigenmann

54 ' Maciel, Rio Guapore Haseman

This species can very readily be distinguished from all others of the genus
by the very widely diverging striae of the scales. The striae are very irregular

• To base of caudal July 23, 1909.

86 THE AMERICAN CHARACIDAE.

in the type forming a network in the median scales. In others they are more
regular, rarely anastomosing but those from above and below meeting along
the median line.

Head 3.5^1; depth usually 2 in the adult (2.4 in some of the specimens
from Obidos and in the young). D. 10 or 11; A. usually 26-28 1 ; scales 5-31
to 34 2 -4. Eye 2.33-2.5 in the head; interorbital equals eye, or but a trace larger.

Compressed, oval, without notable breaks in the nearly symmetrical curves;
profile scarcely depressed over the eyes, ventral profile slightly more arched than
the dorsal; pre ventral area with a median series of about twelve flat scales
bordered on the sides by series of angularly bent scales; postventral area with
a median series of moderate sized thin scales, bent and bordered on the sides
with a series of symmetrical or nearly symmetrical scales, the area compressed;
predorsal region narrowly rounded, the median series of about nine scales extend-
ing from dorsal to occipital crest.

Occipital process one fourth to one fifth in the distance from its base to
the dorsal, bordered on the sides by three scales; interorbital very slightly con-
vex; second suborbital leaving but very narrow naked border; maxillary not
reaching to end of second suborbital, equal to the distance from tip of snout to
pupil ; four or five, rarely six, in one case seven, teeth in the front series of the
premaxillary forming a continuous series on each side; the third tooth slightly
behind the line of the others; five graduated teeth in the second row; four large
teeth in the mandible, slightly graduated; minute teeth on the sides of the
lower jaw, maxillary with one or two teeth.

Gill-rakers about 9 + 12.

Scales deeply imbricate with one to two pairs of widely divergent striae
exposed and several pairs concealed; anal sheath consisting of one to three
rows of minute scales in front and a single row behind, more or less sharply
marked off from the scales of the sides; lateral line scarcely bent up in front,
a well-developed axillary scale; caudal lobes densely scaled to near their tip.

Origin of dorsal a little in advance of middle. Its first divided ray two and
a half times as long as the last, three and a half in the length; anal emargi-
nate, the first rays at least twice the length of some of the posterior ones; origin
of ventrals about equidistant from tip of snout and tip of last anal ray, equi-
distant from tip of snout with the dorsal; ventrals scarcely reaching anal in

1 In thirty specimens from the Amazon Basin one has twenty-four, one twenty-five, seven twenty-six,
ten twenty-seven, five twenty-eight, two twenty-nine, two thirty, and two thirty-two anal rays.

2 Of nineteen specimens from the Amazon Basin, three have thirty-one, five thirty-two, eight thirty-
three, and three thirty-four scales with pores,

THE AMERICAN TETRAGONOPTERINAE. 87

the young, shorter in adult, pectorals to ventrals or the width of one scale farther
in young, not to ventrals in the largest.

A round humeral spot over the third to fifth scales of the lateral line, dis-
appearing with age; no caudal spot in adult, sometimes a dusky area at end
of caudal peduncle in young; a distinct silvery lateral band, the width of the
free margin of a row of scales; fins all plain, mostly hyaline; sides silvery, highly
iridescent.

Air-bladder slender, not bent down behind to the anal, posterior section
not twice as long as the anterior, abruptly narrowed behind; diameter at its
widest part about equal to the eye. Alimentary canal about one and one third
times the entire length with the caudal; intestines containing insects and vege-
table fibers.

Vertebrae 11 + 18. Tip of occipital process not extending beyond the
posterior face of the skull.

As stated above, specimens from Obidos are distinctly more elongate than
others, whereas the largest specimen, the one from Cangaruma, is more than
half as deep as long.

12. Moenkhausia ovalis (Gunther).
Plate 7, fig. 3.

Tetragonoplerus ovalis Gunther, Proc. Zool. soc. Lond., 1868, p. 245 (Xeberos) ; Eigenmann & Eigen-
mann, Proc. U. S. N. M., 1891, 14, p. 53; Ulbey, Ann. N. Y. acad. sci., 1895, 8, p. 282; Fowler,
Proc. Acad. nat. sci., Phil., 1906, p. 441, fig. 38 (Maraflon); Regan, Ann. mag. nat. hist., 1913, ser.
8, 12, 281 (Ucayali).

Tetragonoplerus clialceus Cope, Proc. Amer. philos. soc, 1878, 17, p. 691 (Maraflon).

Moenkhausia ovalis Eigenmann, Rept. Princeton univ. exped. Patagonia, 1910, 3, p. 437.

Habitat. — Maraflon.

I have examined the type in the British Museum, represented in Plate 7,
figure 2. It is evidently closely related to M. grandisquamis.

Head 3.66; depth 2; D. 11; A. 27; scales 5-31-4 (6 to anal). Eye 3 in
the head, 1.33 in interorbital.

Upper profile rather more convex than the lower, scarcely concave at the
nape; preventral area flat; maxillary with two teeth, extending somewhat
beyond front margin of the eye; origin of dorsal just behind origin of ventrals;
pectoral extending beyond the ventrals, ventrals to the vent. Humeral spot
indistinct, caudal spot diffuse, extending over the base of the fin, not to the end
of the middle rays.

88 THE AMERICAN CHARACIDAE.

13. Moenkhatjsia barbouri Eigenmann.
Plate 7, fig. 1; Plate 100, fig. 1.

Moenkhausia barbouri Eigenmann, Bull. M. C. Z., 1908, 62, p. 103 (Villa Bella); Rept. Princeton
univ. exped. Patagonia, 1910, 3, p. 437.

Habitat. — Amazon Basin.

Specimens examined.

Catalogue

Number of

Size

number

specimens

in mm.

Locality

Collector

20708 Cotypes

2

62, 65

Villa Bella

Agassiz

3723 C. 1 64 San Antonio, Rio Madeira Haseman

Very similar to M . grandisquamis, the striae of the scales are different and
the caudal lobes dark.

Head 4.2-4.3; depth 2.4; D. 11; A. 30-33; scales 5.5-37-4.5. Eye 2.5;
interorbital slightly smaller than eye.

Compressed, moderately deep, the dorsal and ventral profiles symmetri-
cally curved, without humps or depressions; preventral area rounded, with
obscure lateral keels; postventral area narrowly rounded; predorsal area
obscurely keeled, with a median series of nine scales from the dorsal to the
occipital process.

Occipital process equals one fourth the distance from its base to the dorsal,
bordered by three scales on the sides; the occipital fontanel continued as a
groove to its tip; interorbital distinctly convex; second suborbital leaving but
a very narrow naked border; maxillary 2.66 in head, equal to the distance from
the tip of the snout nearly to pupil; four teeth in the front series of the pre-
maxillary, five teeth in the second series; maxillary with a single tooth. Lower
jaw with four large teeth in front and several minute ones on the sides. Gill-
rakers about 7 + 13, the longest about one fourth of the eye.

Scales of the sides with four or more diverging striae; anal sheath consist-
ing of a single series of scales; caudal lobes scaled for at least half their length;
lateral fine nearly straight.

Origin of dorsal equidistant from tip of snout and base of upper caudal
lobe; ventrals equidistant from tip of snout and base of last but four rays of

THE AMERICAN TETRAGONOPTERINAE. 89

anal, their distance from tip of snout slightly less than that of the dorsal; high-
est dorsal ray three and one half in the length, equal to the caudal lobes; anal
but slightly emarginate, its highest ray reaching about to the base of the six-
teenth ray, its base about one third of the length ; ventrals not reaching to anal,
and the pectorals not quite to the ventrals.

A faint vertical humeral spot; tips of caudal lobes and middle rays faintly
dusky; a well-defined, silvery lateral band, two thirds as wide as the eye; sides
iridescent silvery.

14. Moenkhausia xinguensis (Steindachner) .

Tetragonopterus xinguensis Steindachner, Flussf. Siidamer., 1882, 4, p. 32 (Xingu); Eigenmann &

Eigenmann, Proc. U. S. N. M., 1891, 14, p. 53; Ulret, Ann. N. Y. acad. sci., 1895, 8, p. 281.
Moenkhausia xingue?isis Eigenmann, Rept. Princeton univ. exped. Patagonia, 1910, 3, p. 438.

This species is known only from the types.

"Head 3.33; depth 2.33; D. 11; A. 26; scales 5-32-4; interorbital 3 in the
head, snout 3.6.

Dorsal profile a little more arched than ventral; interorbital nearly flat;
maxillary narrow, reaching nearly to the vertical from the middle of the eye.
Height of dorsal equals length of head; pectoral equals length of head without
the snout, reaching ventrals; ventrals shorter than pectorals, reaching to anal;
caudal slightly longer than head. Golden brown, lower parts of head silvery;
humeral spot large, ill defined. No caudal spot. Differs from lepidurus in
having fewer scales in the lateral line and in the greater depth." Steindachner.

15. Moenkhausia browni Eigenmann.

Conia of the Wacusi Indians.

Plate 12, fig. 2.

Moenkhausia browni Eigenmann, Ann. Carnegie mus., 1909, 6, p. 13 (Aruataima Falls, Potaro River);
Rept. Princeton univ. exped. Patagonia, 1910, 3, p. 438; Mem. Carnegie mus., 1912, 6, p. 324, pi.
47, fig. 3.

Habitat. — Potaro River both above and below the Kaieteur Fall.

Number

of

Size

Catalogue number

specimens

in mm.

1004 C. Type

1

66

Paratypes

1005 C, 11711 I.

25

46-82

1006 C, 11712 I.

12

28-68

1007 C, 11713 I.

69

23-80

100S C.

1

31

1009 C, 11714 I.

9

30-62

1010 C, 11715 I.

2

48-50

1011 C.

1

65

90 THE AMERICAN CHARACIDAE.

Specimens examined.

Locality Collector

Aruataima, Upper Potaro River Eigenmann

Holmia, Upper Potaro River Eigenmann

Two hours below Holmia Eigenmann

Savannah Landing above Kaieteur Eigenmann

Creek below Savannah Landing Eigenmann

above Kaieteur

Tukeit, Lower Potaro River Eigenmann

Amatuk, Lower Potaro River Eigenmann

Tumatumari, Lower Potaro River Eigenmann

Very similar to M. oligolepis but without trace of caudal spot and with
the anal falcate.

Head 3.75-4; depth 2.3-2.6; D. 11; A. 23 or 24; scales 5-30 to 34-3.
Eye 2.4-2.5; interorbital 2.8-3.

Compressed, elevate; the dorsal profile high, angulated at the origin of
the dorsal; profile depressed over the eye; ventral profile regularly arched
from the snout to the end of the anal. Predorsal area narrowly rounded, with
a median series of eight to ten scales; preventral area bluntly keeled, with a
median series of scales; postventral area narrowly rounded, with a series of
saddle-shaped median scales.

Occipital process four in the distance from its base to the dorsal, bordered
by three scales on the side; head narrow, interorbital convex, smooth; fontanels
of equal width, the posterior considerably longer, continued as a groove to the
tip of the process. Second suborbital, striate, leaving a considerable naked area.
Maxillary 2.6 in the head. Usually five teeth in the front row of the premaxil-
lary, the third tooth withdrawn from the line of the rest; five graduated teeth
in the inner series; the mandible with four large graduate teeth in the front
and small ones on the sides. Three small teeth in the maxillary.

Scales regularly and deeply imbricate, without interpolated rows; each
scale with numerous radiating striae; lateral line sagging to below the middle
of the dorsal; anal sheath of a single series of scales along the first twelve rays;
caudal lobes scaled for half their length.

Origin of dorsal in advance of the middle of the body, its longest ray two and
a half as long as the penultimate, three and a third in the length; caudal lobes
equal, a little longer than the longest dorsal ray; anal emarginate, its longest

THE AMERICAN TETRAGONOPTERINAE. 91

ray when depressed reaching the base of the last but four anal ray; ventrals
reaching anal, pectorals about one scale beyond origin of ventrals.

No caudal spot, a large horizontally oval humeral spot continued below
to the origin of the pectoral; a dark band from origin of dorsal obliquely down-
ward and forward to the lateral line; a dark median lateral line; white below,
dark along back, each scale of the side with a conspicuous dark crescent along
its middle.

In life all fins but the adipose strongly tinged with red; middle of adipose
yellow.

16. Moenkhausia megalops Eigenmann.
Plate 7, fig. 2.

Tclragonopterus grandisquamis Ulbey (non Miiller & Troschel), Ann. N. Y. acad. sci., 1895, 8, p. 281

(Itaituba).
Astyanax megalops Eigenmann, Proc. U. S. N. M., 1907, 33, p. 29 (Itaituba, Brazil).
Moenkhausia megalops Eigenmann, Rept. Princeton univ. exped. Patagonia, 1910, 3, p. 438; Mem.

Carnegie mus. 1912, 5, p. 325 (Rockstone).

Habitat. — Amazon and Guiana.

Specimens examined.

Catalogue

Number

Size

number

of specimens

in mm.

Locality

Collector

5192 Type

1

42 »

Itaituba

Agassiz

20723

1

50 »

Gurupa

Agassiz

2488 C.

1

56

Rockstone, Essequibo River

Eigenmann

3724 C.

1

65

Santarem, in Rio Tapajos

Haseraan

Allied to M. grandisquamis with different proportions and different striae,
with larger eye.

Head 3.6-3.7; depth 2.5-2.66; D. 9-11; A. 28-30; scales 5-34-3 or 4.
Eye 2-2.2 in head, twice as long as snout ; interorbital 2.8-3 in the head.

Elongate, compressed, dorsal and ventral profile evenly curved, a slight
depression over eye; preventral region with a median series of flat scales bor-
dered by a series of scales angularly bent ; postventral region compressed, with a
median series of large, thin scales, bordered on each side by slightly asymmetri-
cal scales; predorsal region compressed, the median series of eight scales extend-
ing from the dorsal to the occipital crest which is contained three to three and a

1 To base of caudal.

92 THE AMERICAN CHARACIDAE.

half times in the distance of its base from the dorsal, and bordered by three
scales on each side.

Groove of the occipital frontanel reaching to tip of occipital process; inter-
orbital slightly convex; second suborbital leaving a narrow naked margin of
the cheeks; maxillary not reaching to the end of the first suborbital two and
three fourths in the head, equal to the distance of the tip of the snout from the
pupil. Four teeth in the front row of the premaxillary, the third slightly with-
drawn from the line of the others, five graduated teeth in the inner series; maxil-
lary with three teeth; mandible with four teeth and numerous small ones on the
side.

Gill-rakers 9 + 14.

Scales deeply imbricate, each with several divergent striae. Anal sheath
of a single series of scales. Lateral line but little descending. A well-developed
axillary scale; caudal scales caducous.

Origin of dorsal in front of middle of body; ventrals nearer tip of snout
than the dorsal, equidistant from tip of snout and base of last anal ray or a little
nearer the former; pectorals extending little beyond origin of ventrals; ventrals
not quite to anal; anal slightly emarginate.

A vertical humeral spot above the space between the third to the fifth scale
of the lateral line, faint; no caudal spot; a broad silvery lateral line. Some
metallic reflections.

The specimen from the Essequibo 2488C. is deeper and has a narrower
second suborbital than the type.

17. Moenkhausia shideleri, Eigenmann.
Plate 12, fig. 1.

Moenkhausia shideleri Eigenmann, Annals Carnegie mug., 1909, 6, p. 15 (Bartica); Rept. Princeton univ.
exped. Patagonia, 1910, 3, p. 438; Mem. Carnegie mus., 1912, 6, p. 325, pi. 47, fig. 4.

Habitat. — Essequibo River.

Specimens examined.

Catalogue

Number of

Size

number

specimens

in mm.

Locality

Collector

1012 C. Type

1

65

Bartica

Shideler

11716 I. Cotype

1

73

Bartica

Shideler

THE AMERICAN TETRAGONOPTERINAE. 93

Head, 3.7-3.8; depth 2.5-2.7; D. 10; A. 26; scales 5-34-3 or 4. Eye 2.1;
interorbital 2.4 or 2.5.

Elongate, subrhomboidal, the ventral surface in front of the anal distinctly
arched; a very slight depression over the eye; preventral area rounded, a series
of median scales; postventral region narrowly rounded, with a median series of
large scales, bordered by slightly asymmetrical scales. Predorsal area keeled
to near the occipital crest. A median series of scales between the dorsal and
occipital crest whose length is about one fifth of the distance from its base to the
dorsal.

Interorbital slightly convex; second suborbital leaving a narrow naked
border on the cheek. Maxillary three in the head; four or five teeth in the front
row of the premaxillary, the third somewhat removed from the rest; five grad-
uated teeth on the inner row; two small teeth on the maxillary; mandible with
four large teeth and numerous small ones on the side.

Gill-rakers about 9 + 12.

Scales regularly and deeply imbricate, each with several divergent striae.
Anal sheath of a few scales in a single row near front of anal; lateral line but little
decurved; caudal scaled for about one fourth of its length.

Origin of dorsal in front of middle of body, but little more remote from snout
than dorsal; highest dorsal ray about two and two thirds in the length; caudal
deeply forked, the lobes longer than the dorsal; anal deeply emarginate; ven-
trals not reaching anal, pectorals just to ventrals.

No humeral spot; caudal with a small diffuse dark spot at the base of the
middle rays. Scales of sides margined with dark, the marginal spots tending
to form lines along the sides; pigment more profuse toward the back; a series
of dark spots on the median series of scales of the back.

18. Moenkhausia costae (Steindachner).
Plate 14, fig. 2; Plate 100, fig. 2.

Telragonoplerus costae Steindachner, Anz. K. akad. wiss. Wien, 1907, p. 83 (Rio San Francisco; Rio
Grande do Norte; Rio Preto).

Habitat. — Rio San Francisco; Rio Itapicuru.

94 THE AMERICAN CHARACIDAE.

Specimens examined.

Catalogue Number of Size

number

specimens

in mm.

Local]

ity

Collector

3713 C.

19

57-64

Joazeiro

Haseman

3714 C.

51

largest 59

Pirapora

Haseman

3715 C.

76

largest 61

Lagoa Pereira

Haseman

3716 C.

1

69

Lagoa de Porto

Haseman

3717 C.

26

48-60

Barreiras, Lagoas

of Rio Grande

Haseman

3718 C.

18

36-57

Boqueirao

Haseman

3719 C.

31

35-58

Santa Rita

Haseman

3720 C.

37

35-56

Queimadas, Rio Itapicurfi

Haseman

3721 C.

44

40-70

Penedo

Haseman

3722 C.

16

11M8

Penedo

Haseman

Head 4; depth 2.25-3.25; D. 11; A. 26-28; scales 5-32 to 34-3.5. Eye
2.6 in the head; interorbital equal to the eye or a little narrower.

Much compressed, the depth varying greatly; dorsal and ventral outline
regularly and nearly equally arched; area immediately in front of ventrals flat,
region between the pectorals bluntly keeled, about fourteen scales in the median
series in front of the ventrals; predorsal area bluntly keeled, with a median
series of ten scales.

Occipital process about one fifth of the distance from its base to the dorsal,
bordered by three scales on each side; interorbital convex; frontal fontanel dis-
tinctly shorter than the parietal without its groove.

Third suborbital covering the entire cheek, leaving but a small naked tri-
angle under its anterior corner; mouth small, the antero-posterior extent of the
premaxillary very small, maxillary large, about as long as the eye, its anterior
margin convex; the mouth similar to that of M. dichrourus. Teeth all feeble,
each ramus of the mandible with five or six graduate teeth, of which the second
is more forward, out of line with the rest, forming an incipient second series,
sides of the ramus with about seven to ten minute conical teeth; premaxillary
with three to five teeth in the outer row, five in the inner; maxillary with or
without a microscopic conical tooth.

Gill-rakers slender, about equal to the snout in length, 11 + 14. Scales thin,
entire, with widely diverging radiae; no interpolated scales; anal with a feeble
sheath of one series of scales in its front half; caudal lobes densely scaled to near
the tip, the central rays nearly naked.

Dorsal pointed, the first rays about three and a half in the length; anal

1 Specimens only fifteen mm. long have the characteristic coloration.

THE AMERICAN TETRAGONOPTERINAE. 95

emarginate, its origin behind the vertical from the base of the last dorsal ray;
ventrals reaching anal, pectorals about one scale beyond base of ventrals.

Straw-color in alcohol, a narrow silvery, lateral band; sometimes a faint,
always minute, humeral spot; a black stripe along base of anal, along ventral
surface of caudal peduncle, then upward and backward along the posterior
margin of the upper caudal lobe. This black stripe is diagnostic for the species,
although it is at times (Penedo specimens) scarcely perceptible. It is especially
conspicuous in the specimens from Queimadas.

19. Moenkhausia dichroura (Kner).
Plate 8, fig. 3; Plate 15, fig. 1; Plate 95, fig. 3; Plate 100, fig. 3.

Telragonopterus dichrourus Kner, Characinen, 1859, p. 41, tab. 9, fig. 21 (Rio GuaporS; Caigara; Para-
guay); Guntheb, Cat. fishes Brit, mus., 1864, 5, p. 324; Eigenmann & Eigenmann, Proo. U. S.
N. M., 1891, 14, p. 53; Ulrey, Ann. N. Y. acad. sci., 1895, 8, p. 279; Perugia, Ann. Mus. civ. storia
nat. Genova, 1891, ser. 24, 10, p. 45 (Chaco Centrale); Boulenger, Trans. Zool. soc. Lond., 1896,
14, p. 35 (San Luis and Descalvados) ; Boulenger, Boll. Mus. univ. Torino, 1897, 12, no. 279, p. 4
(San Lorenzo); 1900, 15, no. 370, p. 2 (Urucum); Eigenmann & Kennedy, (in part), Proc.
Aead. nat. sci., Phil., 1903, p. 522 (Ascuncion; Arroyo Trementina).

Moenkhausia dichrourus Eigenmann, Ann. Carnegie mus., 1907, 4, p. 138, pi. 41, fig. 1 (Tuyuyu;
Corumba; Asuncion); Rept. Princeton univ. Exped. Patagonia, 1910, 3, p. 438.

Moenkhausia dichroura Eigenmann, Mem. Carnegie mus., 1912, 5, p. 326.

Habitat. — Guiana south to Bolivia, Paraguay and the Paranagua.

Catalogue Number of Length

number specimens in mm. Locality Collector

3698 C. • 5 28-62 Rio Jauru Haseman

3699 C. 3 44-91 Puerto Suarez Haseman
3701 C. 5 53-75 San Joaquin, Bolivia Haseman

3703 C. 2 43-52 San Joaquin Haseman

3704 C. 2 51-60 Corumba Haseman

3705 C. 2 56-60 Bastos Haseman

3706 C. 8 52-66 Asuncion Haseman

3707 C. 2 62-65 Arequa Haseman

3708 C. 26 40-06 Villa Hays Haseman

3709 C. 16 52-67 Lagoa de Paranagua Haseman
3738 C. 2 58-62 Santarem Haseman

1345 C, 11861 I. 2 59-62 Warraputa Cataract, Essequibo River Eigenmann

1346 C. 2 28-65 Crab Falls, Essequibo River Eigenmann

1347 C, 11S62 I. 12 58-63 Konawaruk, Essequibo River Eigenmann

These specimens are without exception more slender than the types. In
shape the types resemble M. barbouri, from which they differ in the number of

96 THE AMERICAN CHARACIDAE.

anal rays. Distinguished by the oblique mouth, feeble dentition, rounded
anterior margin of the maxillary, and color of the caudal.

Head about 4; depth usually 3, ranging from 2.75 to 3.5; D. 11; A. 25-28 ';
scales 5 or 5.5-34 to 39 2 -3 or 3.5 (to ventrals). Eye 2.4-2.6 in the head; inter-
orbital slightly less than eye.

Elongate, moderately compressed; dorsal and ventral profiles nearly sym-
metrical, the ventral profile slightly more convex than the dorsal which is slightly
depressed over the head; preventral area rounded with obscure keels on the sides,
postventral area very narrowly rounded or keeled; predorsal area keeled, with a
median series of about ten scales from the occiput to the dorsal.

Occipital process one fifth the distance from its base to the dorsal, bordered
by three scales on the sides; interorbital distinctly and evenly convex; second
suborbital variable, leaving a narrow naked area on the cheek; maxillary about
three in the head, equal to the distance from the tip of the snout to the pupil,
its anterior margin regularly arched; mandible two and a half in the head, very
oblique; mouth small, dentition feeble, usually four (3-5) teeth in the front
row of the premaxillary, the third removed slightly from the line of the others;
five, rarely six teeth in the inner series; maxillary without teeth or with one,
rarely two, feeble ones; lower jaw with four larger teeth and several minute
ones on the sides, the mouth is much smaller (narrower) and the dentition much
feebler than in M. lepidura of the same size.

Gill-rakers long and slender, much longer than M. lepidura, about 11 + 16,
those of the upper arch similar to but shorter than those of the lower, the longest
one third to half the length of eye.

Scales nearly semicircular, deeply imbricate, with several diverging striae 3 ;
anal sheath of a single series of scales; caudal lobes scaled for two thirds of their
length; a well-developed axillary scale.

Dorsal about equidistant from tip of snout and base of caudal, its height
three and a half in the length; ventrals nearer tip of snout than the dorsal,
about equidistant from tip of snout and end of anal; origin of anal behind

1 In twenty specimens from the Amazon Basin, six have twenty-five, seven twenty-six, five twenty-
seven, and two twenty-eight rays.

2 In sixteen specimens from the Amazon Basin one has thirty-four, two have thirty-five, seven thirty-
six, and six thirty-seven pores in the lateral line. A specimen from Paraguay has thirty-nine; of fifteen
specimens from the Essequibo, six have thirty-six, eight thirty-seven, and one has thirty-nine.

3 The striae usually start from about the same vertical line and diverge backward, one specimen
from Villa Bella (20711) and two from Jose-Fernandez (21006) differ from the rest. The condition
approaches that of M. grandisquamis. Successive pairs of striae are joined at the base and follow each
other like a series of stacked v's placed horizontally. The specimen from Jos6-Fernandez have the
depth 2.75 in the length.

THE AMERICAN TETRAGONOPTERINAE.

97

the vertical from the last dorsal ray; ventrals not reaching anal, pectorals to
ventrals.

A well-defined, silvery, lateral band, half as wide as the eye along the middle
of the sides, tapering from below the dorsal forward; middle caudal rays black,
caudal lobes beyond the tips of the middle rays black, the tips milk-white. The
intensity of the caudal color differs very much with the condition of the speci-
mens, the method of preservation, and the locality; an insignificant humeral
spot not evident in alcoholic specimens.

The silvery lateral band of the sides is underlaid with a black band of the
same size and shape. In formaline specimens in which the silvery band is
dissolved a black band takes its place. Base of caudal lobes in life bright yellow,
dorsal and adipose yellowish.

Vertebrae 13 + 18.

Posterior air-bladder sausage-shaped, its diameter about equal to that of
eye, its length twice that of the anterior section, bent down somewhat to near
the origin of the anal.

Alimentary canal less than the total length with the caudal.

20. Moenkhausia intermedia Eigenmann.
Plate 16, fig. 2; Plate 101, fig. 8, 9.

Moenkhausia dichrourus intermedins Eigenmann, Bull. M. C. Z., 1908, 62, p. 103 (Tabatinga); Rept.

Princeton univ. exped. Patagonia, 1910, 3, p. 438.
Moenkhausia lepidura madeirae Fowler, Proc. Acad. nat. sci. Phila., 1913, p. 540, fig. 11 (Tributary of

Rio Madeira near Porto Velho).

Habitat. — Amazon, Paraguay, and Parana Basins.

Specimens examined.

Catalogue Number of Length
number specimens in mm.

20762 Cotypes 2

3709 C. 4

3700 C. 1

3702 C. 2

Locality Collector

42-64 Tabatinga Bourget

25-43 San Louiz de Caceres Haseman

47 Rio Santa Rita, near frontier of Bolivia Haseman

60-67 Salto Avanhandava Haseman

Closely resembling M. dichroura from which it differs mostly in the shape
of the open mouth.

98 THE AMERICAN CHARACIDAE.

Head 4 + ; depth 3 + ; D .11 ; A. 25; scales 5-35-3.5. Eye 2.25-2.75; inter-
orbital 3 in the head.

Slender, dorsal and ventral profiles regular; preventral area flat, with a
median series of twelve scales; predorsal area rounded, with a median series of
ten scales; occipital process about one fifth of the distance from its base to the
dorsal; interorbital very little arched; second suborbital nearly covering the
entire cheek in the largest, with a wider naked border in the smaller; maxillary
about one third of the length of the head; its anterior margin not as greatly
arched as in M. dichroura. Premaxillary with three or four teeth in the anterior
series, five in the inner series; dentary with four large teeth, the anterior two
not in a continuous series with the posterior two in the larger specimens, the third
tooth being farther in. Gill-rakers one fourth to four tenths the length of the eye.

Origin of dorsal in middle of the body; origin of ventrals further forward;
pectorals reaching the ventrals, ventrals to anal. Scales with up to eight
divergent striae.

A silvery lateral band, middle caudal rays and tips of the lobes or a submargi-
nal band black.

The specimens described by Fowler are more slender than the others.

21. MOENKHAUSIA LEPIDUBA LEPIDURA (Kner).

Plate 8, fig. 2.

Tetragonopterus lepidurus Kner, Characinen, 1859, p. 40, tab. 8, fig. 20 (Rio Guapor6) ; Giinther, Cat.
fishes Brit, mus., 1864, 6, 328; Steindachner, Flussf. Sudam., 1882, 4, p. 32 (Tabatinga; Cudajas;
Obidos; Villa Bella); Eigenmann & Eigenmann, Proc. U. S. Nat. Mus., 1891, 14, p. 53; Ulrey,
Ann. N. Y. acad. sei., 1895, 8, p. 278; Vaillant, Bull. Mus. d'hist. nat, 1899, p. 155 (Carnot).

Moenkhausia lepidurus Eigenmann, Rept. Princeton univ. exped. Patagonia, 3, 1910, p. 438.

Moenkhausia lepidura Eigenmann, Mem. Carnegie mus., 1912, 6, p. 326.

Habitat. — Amazons Basin and north to Guiana.

The types in the Vienna Museum have the depth three and one half in the
length. Aside from the types I have examined the following:

Catalogue Number of Length

number specimens in mm. Locality Collector

20954 5 63 to about 90 Jatuarana Navez

20743 4 70-85 Tajapurii Agassiz

21062 12 68-80 Arary Thayer

20722 1 90 Gurupa Agassiz

20790,20788,20778 34 63-84 Santarem Bourget

THE AMERICAN TETRAGONOPTERINAE.

99

Catalogue

Number of Length

number

specimens in mm.

Locality

Collector

20965, 20863, 20864 7

45-81

Rio Tapajos

Dexter, James,

and Talisman

20849

2

88

Obidos

Bentos

Part of 20829

2

about 40 to

Obidos

James

about 50

20712

3

about 70

Villa Bella

Agassiz

21019

1

about 75

Ueranduba

Coutinho

20992

7

about 60 to
about 80

Serpa

Thayer

21010

1

85

Justa

20730, 20733

6

54-72

Lake Hyanuary

Agassiz

20735

12

about 55-73

Lago Maximo

Agassiz

20701, 20702, 20703 l 33

49-85

Teffe

Agassiz

20744

1

about 76

Tocantins

Agassiz

20853, 20857

3

45-60

Jutaby

James, Thayer,

and Talisman

3733 C.

1

68

Cachoele de Riberao, Rio Madeira Haseman

3731 C.

7

44 to about 89

Maciel, Rio Guapore"

Haseman

3732 C.

3

67-77

Bastos

Haseman

1335 C,

11825 I

203

54-75

Bartica, Essequibo River

Shideler

1336 C,

11826 I

12

47-71

Rockstone, Essequibo River

Eigenmann

1337 C,

11827 I

8

42-56

Gluck Isl., Essequibo River

Eigenmann

133S C,

11828 I

51

44-80

Crab Falls, Essequibo River

Eigenmann

1339 C,

11829 I

22

45-58

Konawaruk, Essequibo River

Eigenmann

1340 C,

11830 I

7

45-82

Warraputa, Essequibo River

Eigenmann

1341 C,

11831 I

215

42-95

Tumatumari, Potaro River

Eigenmann

1342 C,

11832 I

20

45-94

Creek below Potaro Landing

Shideler

1343 C,

11833 I

4

51-61

Erukin, Potaro below Amatuk

Eigenmann

1344 C,

11834 I

4

62-108

Malali, Demerara River

Shideler

The species is very variable in shape, number of anal rays and scales. It
is distinguished by the peculiar coloration of the upper half of the caudal.

Head 3.75-L4; depth 3 2 -3.5; D. 11; A. most often 24, ranging from 22-
27 3 ; scales 5-35-4 4 . Eye 2.5-3, about equal to the interorbital.

Elongate, moderately compressed, greatest depth under origin of dorsal,
the dorsal and ventral profiles evenly curved, or the ventral profile more arched,
or preventral area broadly rounded; postventral area very narrowly rounded.

1 In part.

2 In some of the specimens (20788) from Santarem, and some of those from Guiana, the depth is
much greater, 2.75 in the length.

3 Of fifty-five specimens six have twenty-two anal rays, sixteen twenty-three, twenty-one, twenty-
four, seven twenty-five, two twenty-six, and three twenty-seven.

4 In one case there were found six scales above the lateral line, in two cases five and one half. The
scales along the lateral line average thirty-five, but range from thirty-one to thirty-seven. In twenty-
four one has thirty-one scales, five thirty-four, eleven thirty-five, five thirty-six, and two thirty-seven.

100 THE AMERICAN CHARACIDAE.

Predorsal area obscurely keeled, a median series of scales may extend from the
dorsal to the occipital process or a variable number of the median scales near
the occipital process may be divided into two halves, leaving a naked line con-
tinued from the occipital process; the median series of scales thus varies from
six to nine.

Occipital process moderate, four or five times in the distance from its base
to the dorsal, bordered by three scales on the side. Interorbital convex; second
suborbital covering a variable amount of the cheeks but always leaving a com-
paratively broad naked border even in the large specimens (20722) ; maxillary
not reaching to the end of the first suborbital, equal to the length of the snout
or a little more. Premaxillary usually with four teeth in the front series, the
second tooth withdrawn from the line; rarely three or five teeth (in one case
six) in the front row, second row with five graduated teeth; maxillary usually
with one, rarely with two teeth; lower jaw with four, large graduated teeth on
each side and a number of smaller ones.

Gill-rakers very small, about 6 + 9, longest about four in the diameter of
the eye.

Scales regularly imbricate, thin, those on the sides with two to ten striae.
Anal naked, or its sheath composed of a single series of scales; caudal lobes
scaled for at least half their length; lateral fine very slightly decurved; four
scales between the origin of the anal and the lateral line; axillary scale well
developed.

Origin of dorsal little nearer tip of snout than base of caudal; origin of ven-
trals scarcely nearer tip of snout than the dorsal; highest dorsal ray three and
a half in the length; anal emarginate, its origin below the first or second scale
behind the dorsal; ventrals about reaching anal or but little beyond anus, pecto-
rals nearly or quite to ventrals.

Upper half of caudal, except a semicircular spot at the upper part of its
base, black of varying intensity, sometimes shading into light at the upper
margin and with the tip light; lower half of the caudal hyaline or dusky toward
the tip (20730) shading into the black of the upper lobe at the middle; a small,
horizontally oval, or round humeral spot over the second and third scales of
the lateral line; a silvery lateral band one fourth the diameter of the eye in
width, bordered above by dusky; iridescent steel-blue above, brassy below.
In the young the caudal markings are very faint.

In living specimens from Guiana the base of the upper caudal lobe is con-
spicuous!}' yellow or orange or cherry, base of lower caudal lobe, adipose, dorsal

THE AMERICAN TETRAGONOPTERINAE. 101

rays and anal lobe less intensely yellow or orange. The black of the caudal is
most variable, darkest near the orange spot or nearer the tip, the part of one
color shading gradually into that of the other or with an abrupt broken divid-
ing line or "water marked."

Vertebrate 14+17.

Posterior air-bladder of nearly equal width throughout, its width equal to
the diameter of the eye, its length nearly twice that of the anterior section; ali-
mentary canal not quite equal to the entire length.

22. Moenkhausia lepidura lata Eigenmann.
Plate 101, fig. 10.

Moenkhausia lepidurus lotus Eigenmann, Bull. M. C. Z., 1908, 52, p. 103 (Rio Tapajos); Rept. Prince-
ton univ. exped. Patagonia, 1910, 3, p. 438.

20860 Cotypes 27 55-75 mm. Rio Tapajos. Dexter, James, and Talisman.

These specimens differ from others from the Tapajos and from various other
localities. The anal rays average 26 + ; five specimens have twenty-five,
eleven twenty-six, eight twenty-seven, and four twenty-eight rays. The body
is deeper, the depth averaging 2.6 of the length; the scales in the lateral line
vary from 32 to 34, being most frequently 33.

The middle caudal rays are but faintly colored if at all, but the upper caudal
lobe is black.

23. Moenkhausia lepidura icae Eigenmann.
Plate 101, fig. 6.

Moenkhausia lepidurus icae Eigenmann, Bull. M. C. Z., 1908, 62, p. 103 (Iga); Rept. Princeton univ.
exped. Patagonia, 1910, 3, p. 438.

Habitat. — lea.

20810, 20812 Cotypes 46 30-50 mm. Ica James

Deep, compressed fishes, depth three in the length; anal rays usually
twenty-three or twenty-four; two have twenty-one, six twenty-two, ten twenty-
three, thirteen twenty-four, and one has twenty-five rays; the scales in the

102 THE AMERICAN CHARACIDAE.

lateral line number with about equal frequency, thirty-one, thirty-two, or thirty-
three.

Pectorals just reaching ventrals.

Upper caudal lobes and sometimes the distal part of the other rays dusky,
a very small but well-defined humeral spot of about ten chromatophores over
the fourth or fifth scale of the lateral line; tip of anal lobe sometimes milk-
white.

24. MOENKHATJSIA LEPIDTJRA HASEMANI, Subsp. nOV.

Plate 15, fig. 3; Plate 101, fig. 5.
3746 Type 45 mm. to base of caudal Santarem, Dec. 12, 1909 Haseman

Head4 + ; depth 3.4; D .11; A .24 ; scales 6-37-3.5. Eye 2.66 in the head,
snout 3.33; interorbital 2.66.

Slender, dorsal and ventral profiles gently and evenly curved, preventral
area flattish with a median series of thirteen scales; predorsal area rounded with
a median series of ten scales. Occipital process one fifth as long as the distance
from its base to its dorsal, bordered by three scales. Interorbital rounded,
maxillary a little less than one third of the length of the head, its anterior mar-
gin greatly arched forward, less so posteriorly. Second suborbital leaving a
naked border all around. Three or four teeth in the anterior row of the pre-
maxillary, a small tooth on the maxillary; dentary with four large teeth and
abruptly small ones on the sides.

Origin of dorsal midway between tip of snout and base of middle caudal
rays; pectorals not reaching the ventrals whose origin is in advance of the
vertical from the origin of the dorsal; anal slightly falcate.

Scales firm with up to ten prominent radiae. Anal sheath very feeble,
axillary scale well developed.

A faint humeral spot on and above the third and fourth pores of the lateral
fine. A silvery lateral band, most prominent between the dorsals and ending
a few scales in advance of the lateral line. No caudal spot.

EXPLANATION OF THE PLATES.

PLATE 1.

PLATE 1.

Map of a part of South America, showing the routes Spix and Martius (1817-1820), of Castelnau
(1844-1847), Natterer (1817-1835), Schomburgk (1840-1847), the Thayer Expedition (1865-1866),
and the common route along the Amazon.

Mem Mu

PLATE 2.

PLATE 2.

Fig. 1. Tetragonopterus argenteus Cuvier.

20718 M. C. Z. Length of head 17.5 mm. Villa Bella, Brazil.

Fig. 2. Deuterodon iguape Eigenmann.

9265 I. U. M. Type. 110 mm. Iguap6, Southeastern Brazil.

Fig. 3. Hemibrycon guppii Regan.
11326 I. U. M. Trinidad.

Fig. 4. Hollandichtys multifasciatus Eigenmann & Norris.

9288 I. U. M. Type. 70 mm. Cubatao, Southeastern Brazil.

Fig. 5. Creatochanes affinis (Gunther).

21076 M. C. Z. Head 24 mm. Porto do Moz, Brazil.

Fig. 6. Astyanacinus moorii Fowler.

A. N. S. P. Cotype. 102 mm. Chapala Plateau, Brazil.

MEM. MUS. COMP. ZOOL

CHARACIDAE PLATE 2

COCKAYNE. BOSTON

PLATE 3.

PLATE 3.

Fig. 1. Entomolepsis steindachneri Eigenmann.
20967 M. C. Z. 86 mm. Iquitos, Peru.

Fig. 2. Scale from near the anal.

Fig. 3. Last scale above the lateral line in Fig. 1.

Fig. 4. Henochilus wheatlandi Garman.

21109 M. C. Z. 415 mm. Type. Southeastern Brazil.

Fig. 5. Thayeria obliqua Eigenmann.
M. C. Z. Type. Obidos, Brazil.

MEM. MUS. COMP. ZOOL

CHARACIDAE PLATE 3

HEL10TYPE CO. BOSTON

PLATE 4.

PLATE 4.

Fig. 1. Tetragonopterus chalceus Agassiz.

11852 I. U. M. 68.5 mm. Wismar, British Guiana.

Fig. 2. Tetragonopterus argenteus Cuvier.

20718 M. C. Z. 76 mm. Villa Bella, Brazil.

MEM. MUS. COMP. ZOOL.

CHARACIDAE PLATE 4

HELIOTYPE CO. BOSTON

PLATE 6.

PLATE 5.

Fig. I. Moenkhausia jamesi Eigenmann.

20742 M. C. Z. 72 mm. Type. Amazon.

Fig. 2. Moenkhausia latissima Eigenmann.

20767M.C.Z. 71.5 mm. Type. Tabatinga, Brazil.

Fig.S. Entomolepis steindachneri (Eigenmann).
20967 M. C. Z. 86 mm. Iquitos, Peru.

MEM. MUS. COMP. ZOOL

CHARACIDAE PLATE 5

;

COCKAYNE BOSTON

PLATE 6.

PLATE 6.

Fig. 1. Moenkhausia melogramma Eigenmann.

20825 M. C. Z. About 47 mm. (caudal broken). Type. Tabatinga, Brazil.

Fig. 2. Moenkhausia comma Eigenmann.

20972 part, M. C. Z. About 85 mm. (caudal broken). Type. Cudajas, Brazil.

Fig. 3. Moenkhausia chrysargyrea (Gunther).

11842 I. U. M. 78.5 m. Crab Falls, British Guiana.

MEM. MUS. COMP. ZOOL.

CHARACIDAE PLATE 6

HEUOTvPE CO. BOSTON

PLATE 7.

PLATE 7.

Fig. 1. Moenkhausia barbouri Eigenmann.

20108 M. C. Z. 65 mm. Type. Villa Bella, Brazil.

Fig. 2. Moenkhausia megalops Eigenmann.

20723 M. C. Z. 56 mm. (caudal broken). Gurupa, Brazil.

Fig. 3. Moenkhausia ovalis (Giinther). From Type in British Museum. Xeberos, Brazil.

Fig. 4. Moenkhausis oligolepis (Giinther).

118351. U. M. 58 mm. Holmia, British Guiana.

MEM. MUS. COMP. ZOOL.

CHARACIDAE PLATE 7

HELIOTYPE CO. BOSTON

PLATE 8.

PLATE 8.

Fig. 1. Moenkhausia colletti (Steindachner).

11809 I. U. M. 66 mm. Tumatumari, British Guiana.

Fig. 2. Moenkhausia lepidura lepidura (Kner).
20788M.C.Z. 80 mm. Santarem, Brazil.

Fig. 3. Moenkhausia dichroura (Kner).

11862 I. U. M. 63 mm. Konawaruk, British Guiana.

MEM. MUS. COMP. ZOOL.

CHARACIDAE PLATE 8

COCKAYNE. BOSTON

PLATE 12.

PLATE 12.

(Mem. Carnegie Mus., 6, Plate 47.)

Fig. 1. Moenkhausia shideleri Eigenmann.

1012 C. M. 65 mm. Type. Bartica, British Guiana.

Fig. 2. Moenkhausia browni Eigenmann.

1004 C. M. 66 mm. Type. Aruataima Falls, British Guiana.

Fig. 3. Moenkhausia grandisquamis (Miiller and Troschel).
1350 C. M. 100 mm. Crab Falls, British Guiana.

MEM. MUS. COMP. ZOOL.

CHARACIDAE PLATE 12

HEL10TYPE CO., BOSTON

PLATE 14.

PLATE 14.

Fig. 1. Moenkhausia doceana (Steindachner).

20909 M. C. Z. 102 mm. Rio Doce, Brazil. (The scales are mostly lost from the caudal
and are not indicated by the artist).

Fig. 2. Moenkhausia costae (Steindachner).
3722 C. M. 48 mm. Penedo, Brazil.

Fig. 3. Moenkhausia bondi (Fowler). (Mem. Carnegie mus., 6, Plate 46, fig. 1).

2207 C. M. 51 mm. Type of M. ■profunda Eigenmann. Issorora Rubber Plantation, British
Guiana.

MEM. MUS. COMP. ZOOL

CHARACIDAE PLATE 14

\

■P

MEHOTvPE CO., BOSTON.

PLATE 16.

PLATE 16.

Fig. 1. Moenkhausia diehroura (Kner).

3699 C. M. 90 mm. Puerto Suarez, Bolivia.

Fig. 2. Moenkhausia intermedia Eigenmann.

3702 C. M. 60 mm. Salto de Avanhandava, Southeastern Brazil.

Fig. 3. Moenkhausia lepidura hasemani Eigenmann.
3746 CM. 58 mm. Type. Santarem, Brazil.

Fig. 4. Moenkhausia collettii (Steindachner).
3742 C. M. 55 mm. Maciel, Brazil.

MEM. MUS. COMP. ZOOL.

CHARACIDAE PLATE 15

-'.■' : -'^*?

MELIOTYPE CO., BOSTON

PLATE 16.

PLATE 16.

Fig. 1. Moenkhausia sanctae filomenae (Steindachner).
3312 CM. 45 mm. Caceres, Brazil.

Fig. 2. Moenjchausia sanctae filomenae (Steindachner).
3311 C. M. 50 mm. Caceres, Brazil.

Fig. 3. Knodus victoriae (Steindachner).

3751 C. M. 53 mm. Lagoa Paranagua, Brazil.

MEM. MUS. COMP. ZOOL.

CHARACIDAE PLATE 16

•■■iOBfell

1

HEUOTYPE CO., BOSTON

PLATE 96.

PLATE 96.

Fig. 1. Astyanax fasciatus (Cuvier).

12739 I. U. M. 162 mm. Quibdo, Colombia.

Fig. 2. Phenacogaster megalostictus Eigenmann.

11746 I. U. M. 63 mm. Tumatumari, British Guiana.

Fig. 3. Moenkhausis dichroura (Kner).

11862 I. U. M. 60 mm. Konawaruk, British Guiana.

Fig. 4. Moenkhausia oligolepis (Giinther).

118361. U. M. 114 mm. Aruataima, British Guiana.

Fig. 5. Astyanax festae (Boulenger).

13119 I. U. M. Largest, 67 mm. Puerto viejo, Ecuador.

Fig. 6. Astyanax bimaculatus (Linnd).
11308 I. U. M. 134 mm. Trinidad.

Fig. 7. Deuterodon iguape Eigenmann.

11636 1. U. M. 112 mm. Raiz da Serra, Southeastern Brazil.

Fig. 8. HoUandichthys multifasciatus (Eigenmann & Norris).
13297 1. U. M. 105 mm. Raiz da Serra, Southeastern Brazil.

MEM. MUS. COMP. ZOOL.

CHARACIDAE PLATE 95

>~te^

»^Y i >^frw

riti^^^^

8

HELIOTVPE CO., BOSTON

PLATE 98.

PLATE 98.

Fig. 1. Creatochanes melanurus (Bloch).

11887 I. U. M. 136 inni. Wismar, British Guiana.

Fig. 2. Genycharax tarpon Eigenmann.

12674 1. U. M. 135 mm. Cali, Colombia.

Fig. 3. Stichanodon insignis (Steindachner).

20803 M. C. Z. 65 mm. Manacapuru, Brazil.

Fig. 4. Tetragonopterus chaloeus Agassiz.

118551. U. M. 85 mm. Crabs Falls, British Guiana.

Fig. 5. Ephippicharax orbicularis (Cuvier & Valenciennes).
12212 I. U. M. 82 mm. Rockstone, British Guiana.

Fig. 6. Stethaprion erythrops Cope.

19268 M. C. Z. 94 mm. Jutahy, Brazil.

Fig. 7. Ctenobrycon hauxwellianus (Cope).
20814 M. C. Z. 84 mm. Ica, Brazil.

MEM. MUS. COMP. ZOOL.

CHARAC1DAE PLATE 98

HEUOTYPE CO., BOSTON

PLATE 100.

PLATE 100.

Camera outlines of the heads.

Fig. 1. Moenkhausia barbouri Eigenmann.

Fig. 2. Moenkhausia costae (Steindachner).

Fig. 3. Moenkhausia diehroura (Kner).

Fig. 4. Moenkhausia melogramma Eigenmann.

Fig. 5. Moenkhausia oligolepis (Gunther).

Fig. 6. Moenkhausia grandisquamis (Muller and Troschel).

Fig. 7. Moenkhausia bondi Fowler.

Fig. 8. Moenkhausia jamesi Eigenmann.

Fig. 9, 10. Moenkhausia chrysargyrea (Gunther).

MEM. MUS. COMP. ZOOL.

CHARACIDAE PLATE 100

HEUCnPE CO., BOSTON

PLATE 101.

PLATE 101.
Camera outlines of the heads.

Fig. 1. Moenkhausia ceros Eigenmann. Type.

Fig. 2. Moenkhausia cotinho Eigenmann. Type.

Fig. 3. Moenkhausia collettii (Steindachner).

Fig. 4. Moenkhausia copei (Steindachner).

Fig. 5. Moenkhausia lepidura hasemani Eigenmann.

Fig. 6. Moenkhausia lepidura icae Eigenmann.

Fig. 7. Moenkhausia lepidura gracilima Eigenmann.

Figs. 8, 9. Moenkhausia intermedia Eigenmann.
Fig. 10. Moenkhausia lepidura lata Eigenmann.

MEM. MUS. COMP. ZOOL.

CHARACIDAE PLATE 101

10

HEUOTYPE CO., BOSTON

PUBLICATIONS

MUSEUM OF COMPARATIVE ZOOLOGY

AT HARVARD COLLEGE.

There have been published of the Bulletin Vols. I. to LIV., and
Vols. LVI., LVIII. to LX.; of the Memoirs, Vols. I. to XXXIV.,
and also Vols. XXXVI. to XXXVIII., XL. to XLII., XLIV., and
XLVI.

Vols. LV., LVII., LXL, and LXII. of the Bulletin, and Vols.
XXXV., XXXIX., XLIII., XLV., XLVII. toXLTX. of the Memoirs,
are now in course of publication.

.1 price lint of the publication* of the Museum will In sent on appli-
cation to the Director of the Museum of Comparative Zoology, Cambridge,
Mass.

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