ANNOTATED LIST OF THE SPECIES 381 489. ORESTIAS ROSPIGLIOSIE Kigenmann and Allen, sp. nov. 12290,5,largest,type,53mm.,RioLanguf,Peru(above12,000feet)Eigenmann,December8,1918. Head 4.0-4.2; depth 5.33-5.5; D. 18-14; A. 14-15; eye 2.4—2.75; interorbital 4 in the head, snout very little more than half the eye in length; mouth small, very oblique; mandible vertical, forming a right angle with the lower surface of the head, equal to snout and half the eye.
Seales small, mostly lost, about 15 between the middle of the anal fin and dorsal; origin of the dorsal fin nearer to the base of the caudal than to the eye; origin of the anal fin about opposite to the origin of the dorsal; depth of the caudal peduncle 2.5-3.0 in its length.
For our friend Dr. Carlos Rospiglosi y Vigil, director of the Zoological Museum of the University of San Marcos, leader of the University’s expeditions into the Chanchomayo, member of the army medical corps, and of the army hospital staff. Having been unable to find the types, the description is incomplete, and I do not know where the species belongs.
Order SYNENTOGNATHI Family XIX: Belonidae Genus 196: POTAMORRHAPHIS Giinther Potamorrhaphis Giinther, 1866, Cat. Fish. Brit. Mus., VI, 234; Aigenmann and Kigenmann, 1891, Proc. U.S. Nat. Mus., XIV, 66; Eigenmann,1910,Rept.PrincetonUniv.Exped.Patagonia,IIT,463; Higenmann,1912,Mem.CarnegieMus.,V,461.
Type: Belone taeniata Giinther = B. guianensis Schomburgk Guiana to Peru and Paraguay Greatly elongated, needle-shaped fishes of small size, with snout greatly prolonged; dorsal rays subequal in length, anterior ones not forming a lobe. 490. PorAMORRHAPHIS GUIANENSIS (Schomburgk) BeloneguianensisSchomburgk,1843,Fish.Brit.Guiana,IT,131,pl.i. PotamorrhaphisguianensisKigenmannandHigenmann,1891,Proc.U.S.Nat.Mus.,XTV,66; Eigenmann,1910,Rept.PrincetonUniv.Exped.Patagonia,IIT,463; Higenmann,1912,Mem.CarnegieMus.,V,461; Fowler, 1919, Proc. Acad. Nat. Sci. Phila., LX-XI, 6, Nauta; Fowler, 1939 (1940), Proce. Acad. Nat. Sci. Phila., XCI, 278, two, 146 and 195 mm., Contamana.
BelonescolopacinaCuvierandValenciennes,1846,Hist.Nat.Poiss.,XVIII,428; Giinther, 1866, Cat. Fish. Brit. Mus., VI, 256. Belone taeniata Giinther, 1866, Cat. Fish. Brit. Mus., VI, 256; Cope, 1878, Proc. Amer. Phil. Soc., XVII, 695, Rio Maranon.
382 FISHES OF WESTERN SOUTH AMERICA The Guianas to Peruvian Amazon; Rio Paraguay 15811, 1, 220 mm., Rio Morona, Allen, October, 1920. 15815, 1, 161 mm., Rio Pacaya, Allen, August, 1920. A marine relict reaching the extreme western margin of the Amazonian flood plain and the fringes of the Andean highlands. Genus 197: TYLOSURUS Cocco Tylosurus Coceo, 1833, Giorn. Sci. Lett. Sicilia, XLII, no. 124; Bigenmann and EHigenmann, 1891, Proc. U. 8. Nat. Mus., XIV, 65; Jordan and Gilbert, 1882, Bull. U. S. Nat. Mus., XVI, 372; Higenmann,1910,Rept.PrincetonUniv.Exped.Patagonia,IIT,462. Type: Tylosurus cantraini Coeco = T. acus Lacépéde and Hsox imperialis Rafinesque Both coasts of the Americas; West Indies; rivers Greatly elongate, subcylindrical; both jaws extended into a beak, the lower somewhat the longer; teeth of both jaws in bands, one band of small, sharp teeth, another of larger, unequal, conical ones; maxillaries and premaxillaries united; scales small, deciduous; lateral line along the belly, median on the tail; separated from Belone by lack of gill-rakers and vomerine teeth. 491. TyLOSURUS AMAZONICUS (Steindachner) Belone amazonicus Steindachner, 1875, Sitzb. Ik. Akad. Wiss. Wien, LX XII, 66. TylosurusamazonicusKigenmannandHigenmann,1891,Proc.U.S.Nat.Mus.,XTV,65; Jordan and Evermann, 1896, Bull. U.S. Nat. Mus., XLVIT, 1, 712; Eigenmann,1910,Rept.PrincetonUniv.Exped.Patagonia,ITT,462. Para to Peru 15812,2,180and212mm.,RioParanapura,Yurimaguas,Allen,November,1920. 15813,2,95and110mm.,Gosulimacocha,Allen,October,1920. 16161, 2, 40 and 52 mm., Rio Pichis, Pto. Bermudez, Allen, July, 1920. None of the familiars of the frontier post at Puerto Bermudez, so far as could be ascertained, had ever seen a specimen until they were shown my collections. At this point they are separated by more than three thousand miles of fresh water from their salt-water kindred.
Order ACANTHOPTERI Family XX: Nandidae Genus 198: MONOCIRRHUS Heckel Monocirrhus Heckel, 1840, Ann. Wien. Mus., IT, 439; Gunther, 1861, Cat. Fish. Brit. Mus., III, 371; Eigenmann and Eigenmann, 1891, Proc. U. 8. Nat. Mus., XIV, 66; Kigenmann, 1910, Rept. Princeton Univ. Exped. Patagonia, III, 466.
ANNOTATED LIST OF THE SPECIES 383 Type: Monocirrhus polyacanthus Heckel Rios Negro, Cupai; Guiana; Maranon Spiny rayed fishes of small size; body extremely compressed; dorsal and anal fins long with serrate margins, erect; soft dorsal and soft anal fins with vertical bases and hyaline, similar in both shape and color to the caudal; snout very long and sharply pointed; the mandible extending beyond snout, with a barbel; premaxillary with a long spine equal in length to the mandible; eye large, 4 in the head. 492. MonocrRRHUS MIMOPHYLLUS Eigenmann and Allen MonocirrhusmimophyllusEigenmannandAllen,1921,Biol.Bull.,XLI,5,301-305,figs.1-3. MonocirrhuspolyacanthusFowler,1939(1940),Proce.Acad.Nat.Sci.Phila.,XCI,279,one,71mm.. Contamana.
Thisspecimenmaybelonghere,asshouldbeexpected.
15715, 3, 44-51 mm., to base of caudal, brook near Rio Itaya, Iquitos, Allen, September, 1920. A leaf-mimicking species of spiny-rayed fishes was described by Heckel in 1840 (Johann Natterer’s Neue Flussfische Brasiliens). It was given the name Monocirrus polyacanthus, and later joined with Polycentrus to form the family Polycentridae, by Giinther. Scientifically it then went into complete desuetude for eighty years, although Dr. Myers informs me it was later rediscovered in Guiana and reported in certain aquarium journals, he himself having noted its occurrence at Rockstone and in the Essequibo.
On account of the imperfect original description, without any illustration, we were uncertain as to the exact identity of our specimens with Heckel’s species. The wide distance separating our locality from the original source on the Rio Negro, a thousand miles to the eastward, made reasonable the assumption that they were not identical, even though similar. Hence the paper of 1921 describing our three specimens as M. mimophyllus. As far as one could tell from Heckel’s account, the identity might still be uncertain.
Meanwhile, since 1921, the leaf-mimic has again come to light in the collections of dealers in tropical fishes, imported into Germany and thence to the United States. Here it is reported as having been successfully propagated in aquaria, and its habits much more fully described. I have unsuccessfully attempted to procure specimens of these importations for comparison with our types. Mr. Innes’s halftone of polyacanthus appears very similar (Innes, 1933; ibid, 1935). However, the source of his material is not made known. The source of commercial collections is usually very indefinite. Whether the commercial species of unknown source is the same as MW. polyacanthus or mimophyllus, or both, or neither, is not yet positively demonstrated.
Dr. Myers has examined a good series of Monocirrhus from the dealers. He assures me that there is but one species, polyacanthus. He does not state where the specimens came from originally. He is sure our specimens are not different in any respect.
However, the Innes photographs (1933, p. 7, and 1935, p. 335) both clearly
384 FISHES OF WESTERN SOUTH AMERICA show about 16-17 spinous rays in the anal fin, where our specimens have but 12 (at most 13). His description does not allude to the ashen markings seen in mimophyllus, nor do they appear in the photograph from life. Mr. Innes’s illustrations show a specimen with mouth distended, clarifying better than our fig. 2 (1921 paper) the significance of the unusual visceral skeleton. It is of interest to note here that the Agassiz in their book, A Journey in Brazil, pages 161 and 238, make allusion to certain fishes of the Amazon which are mimics of dead leaves. These they assign to a family Folhidae, a Portuguese word latinized. The derivation of the name is quite appropriate, but I have found no evidence that it ever attained proper standing.
THE PRESENT STATUS OF THE THEORY OF MIMICRY McAtee and others have recently held up for much adverse criticism the entire matter of mimicry in animals. The ground for their antagonism seems to be the fact that animals possessing it do not appear to enjoy a measurably higher survival rate in Nature than others. We are constrained, they say, to reject the doctrine entirely. If other animals not so protected live as long on the average, the species has not been subjected to the operation of Natural Selection, and no mechanism other than random selection has brought about the infringement of copyright.
So far as it reaches, the argument mentioned is very convincing, and doubtless completely satisfying to many good zoologists. However, should the term mimicry fall into complete disuse and be forgotten, we still have a phenomenon needing a name, one of the most vivid and interesting in all the realm of Nature. Perhaps a better name will be forthcoming, but likely not if we are looking for a term which will explain the phenomenon to the general satisfaction of the zoological world. We do have a well-known phenomenon. Certain animal forms do show a clear-cut resemblance to other objects in their environment. We do have a masquerading act whether we like it or not, and whether we can explain it or not. This masquerade goes on in the immediate vicinity of the object which is imitated, and not at remote, unrelated points. Perhaps the greatest objection to the word mimicry is the fact that it seems to imply psychic factors, that the imitator exercises a voluntary control of its own evolution. We are unable to understand the intermediate steps by which the mimic has arrived at perfection, except as a long line of consciously-directed mutations or gradations.
This phylogeny cannot be shown to have had psychic elements; it has not been demonstrated that mimicry has effective selection value; its processes are not known, nor are the respective cases of mimicry known to have arisen in the same manner. My thesis here is to defend the use of the name until its true name shall have been revealed.
At this juncture it would appear that what we need is a new definition rather than a new provisional name, since no new explanation is available. We might raise some inquiry as to the occurrence in Nature of intermediate stages in the evolution of this at least quasi-mimicry. For such mimicry-in-the-
ANNOTATED LIST OF THE SPECIES 385 making we might consider the numerous classical examples of protective form, such as that of the walking-stick, and protective coloration, applied in some measure to most animals. In none of these cases would we speak of them as mimicry. These factors exist in greater or less perfection, either protective form alone, protective coloration alone, or in varying degrees in combination. Species with colors blending with those of the background, and more or less unusual body form also resembling the animal’s background, are not at all uncommon. We may still be unable to consider them as good examples of mimicry. They may be only good cases of protective resemblance.
Protective resemblance may be thought of as a good name for a phenomenon consisting of elements in varying degrees of perfection called protective coloration and protective form, which are doubtless associated with discrete genes. In the most perfect cases of protective resemblance it might be argued that they have reached the threshold of actual mimicry, but for the sake of a definition let us say that they fall just short of it, that they lack the psychic element. Yet Nature affords examples in which the psychic factor is as evident as in the small boy in the policeman’s costume. Many animals assume attitudes and go through movements similar to the models after which they are formed. A third factor climaxes the whole drama, that of protective behavior. An animal species resembling some other object in its environment in form, color, and behavior is a very good mimic, whether or not we have a sufficiently broad view of itsevolution to understand the steps by which it came into being. Whether or not we may say that the species as a whole has responded to the model after which it is patterned, or that its evolution has been directive, still the individual member of the species does consciously imitate the model in its behavior. The psychic factor is the “priceless ingredient” which finishes our definition of mimicry. We might put the meaning of the above paragraphs as follows: OLE CVC TOUNUD aA aes Lea) - Protective resemblance) Protective coloration.......... -Mimicry Protective behavior. . .
Monocirrhus is a form which admirably exemplifies all the factors of mimicry. It is true that a photograph, a dead specimen, or a drawing may not be especially convincing, even to an open mind. The scholar in the museum, the laboratory, or the library might well remain skeptical as to this performance. We may concede that seeing the living fish in an aquartum might also fail to convince. But in the natural background of the forest pool, with its bottom strewn with dead leaves, and the fish in the midst of them, there is no room for doubt as to the actuality of the phenomenon.
Dr. Julian Huxley recently wrote (Cott; Adaptive Coloration in Animals, Preface): ‘““Among a certain section of experimental biologists, any time in these last thirty years, it has been fashionable, and indeed almost a matter of professional conscience to display a radical skepticism on the subject of adaptations, especially
386 FISHES OF WESTERN SOUTH AMERICA colour adaptations, and most particularly mimetic adaptations. Upholders of the theories of protective and warning coloration and of mimicry have often been attacked as ‘armchair theorists’ (whereas they have in fact almost invariably been first and foremost field naturalists) insufficiently acquainted with modern work in genetics, which for some unexplained reason is held to do away with adaptive interpretations. Dr. Cott has .. . shown that it is they who are the armchair GHIUUGS are ae It is not my purpose here to enter the debate among worthier contenders, but rather to bear witness as to the validity of a natural phenomenon which few men have been privileged to see at its best, a phenomenon which even the unclad savage has observed, and given his testimony in calling the fish ‘“‘pira-cad-a,’’ or leaf-fish. Family X XI: Sciaenidae Genus 199: PLAGIOSCION Gill Plagioscion Gill, 1861, Proc. Acad. Nat. Sei. Phila., 82; Jordan and Eigenmann, 1886 (1889), Rept. U.S. Fish Comm., 380; EKigenmann and Eigenmann, 1891, Proc. U. 8. Nat. Mus., XIV, 67; EKigenmann,1910,Rept.PrincetonUniv.Exped.Patagonia,IIT,467; Kigenmann, 1912, Mem. Carnegie Mus., V, 471.
Type: Sciaena squamosissima Heckel Atlantic drainage from R. Magdalena to La Plata Skull not excessively cavernous nor lacking in firmness; preorbital not turgid, interorbital space moderate; teeth not villiform, more or less unequal, upper teeth in two series, the inner somewhat the larger teeth; pseudobranchiae small; lateral line scales the largest and overlapped by the smaller ones. 493. PLAGIOSCION SQUAMOSISSIMUM (Heckel) SciaenasquamosissimaHeckel,1840,Ann.WienerMus.,I,432. Diplolepis squamosissima Steindachner, 1863, Sitzb. KK. Akad. Wiss. Wien, XLVII, 2; Steindachner, 1882 (1883), Denksch. KIx. Akad. Wiss. Wien, XLVI, 1. PachyurussquamosissimusGiinther,1860,Cat.Fish.Brit.Mus.,IT,526. PlagioscionsquamosissimaGill,1861,Proe.Acad.Nat.Sci.Phila.,82; Jordan and Eigenmann, 1886 (1889), Rept. U.S. Fish Comm., 381; Jordan and Evermann, 1898, Bull. U.S. Nat. Mus., no. 47, IT, 1418; Eigenmann, 1910, Rept. Princeton Univ. Exped. Patagonia, III, 468; Fowler, 1939 (1940), Proce. Acad. Nat. Sei. Phila., XCI, 280, five, 140-210 mm., Contamana. PlagioscionsquamosissimumiquitensisNakashima,1941,Bol.Mus.Hist.Nat.Lima,V,68. JohniuscrouvinaCastelnau,1855,Anim.Amér.Sud,Poiss.,11,pl.v,fig.1. JohniusamazonicusCastelnau,1855,Anim.Amér.Sud,Poiss.,12,pl.iv,fig.1. Peruvian Amazons to Guiana 16146,1,125mm.,Gosulimacocha,Allen,October,1920. 1OL47. 3; , creek near Yurimaguas, Allen, November, 1920. 16148, 1, 180 mm., Manaos, Allen, December, 1920. 16149, 1, , Iquitos, R. Nanay, Allen, September, 1920. 16150, 5, . Iquitos, Amazon, Allen, September, 1920.
ANNOTATED LIST OF THE SPECIES 387 The largest of the above specimens was 250 mm. The species is known throughout the Amazons, the Orinoco basin, and the Guianas. — It is another of the relict marine ichthyfauna of the Amazon.
494. PLAGIOSCION AURATUM (Castelnau) Johnius auratus Castelnau, 1855, Anim. Amér. Sud, Poiss., 12, pl. iv, fig. 2, Rio Ucayali. Sciaena aurata Giinther, 1860, Cat. Fish. Brit. Mus., II, 287. PlagioscionauratusJordanandEigenmann,1886(1889),Rept.U.S.FishComm.,383; Eigenmann and Eigenmann, 1891, Proc. U.S. Nat. Mus., XIV, 67; Higenmann, 1910, Rept. Princeton Univ. Exped. Patagonia, IIT, 468; Higenmann,1912,Mem.CarnegieMus.,V,472.
PlagioscionauratusiquitensisNakashima,1941,Bol.Mus.Hist.Nat.Lima,V,67. Guianas, Rios Apuré, Sao Francisco, Peruvian Amazons Family XXII: Cichlidae Genus 200: CHAETOBRANCHUS Heckel ChaetobranchusHeckel,1840,Ann.WienerMus.,IT,401; Giinther, 1862, Cat. Fish. Brit. Mus., IV, 309; Eigenmann and Higenmann, 1891, Proc. U.S. Nat. Mus., XIV, 70; Pellegrin, 1904, Bull. Soc. Zool. France, XXTX, 200; Higenmann, 1910, Rept. Princeton Univ. Exped. Patagonia, III, 469; Higenmann,1912,Mem.CarnegieMus.,V,483.
Type: Chaetobranchus flavescens Heckel The Guianas to the Rio Guaporé Most readily to be distinguished from other Cichlidae by the exceptional length of the numerous, setiform gill-rakers. 495. CHAETOBRANCHUS FLAVESCENS Heckel ChaetobranchusflavescensHeckel,1840,Ann.WienerMus.,II,401; Miller and Troschel in Schomburgk, 1848, Reisen, IIT, 625; Gunther, 1862, Cat. Fish. Brit. Mus., IV, 310; Eigenmann and Higenmann, 1891, Proce. U. 8. Nat. Mus., XIV, 70; Pellegrin, 1902, Bull. Mus. Hist. Nat., VIII, 182; Pellegrin,1903(1904),Mém.Soc.Zool.France,XVI,190,200; Bigenmann, 1910, Rept. Princeton Univ. Exped. Patagonia, IIT, 469; Eigenmann, 1912, Mem. Carnegie Mus., V, 485; Fowler, 1939 (1940), Proc. Acad. Nat. Sei. Phila., XCI, 280, one, 118 mm., Contamana. Aquidensflavescens?Haseman,1911e,Ann.CarnegieMus.,VII,338. ChromysucayalensisCastelnau,1855,Anim.Amér.Sud,Poiss.,15,pl.vi,fig.2,one,Sarayacu. Geophagus badiipinnis Cope, 1871, Proc. Acad. Nat. Sci. Phila., XXIII, 251, pl. x, fig. 1, R. Ambyiacu.
Widely dispersed Guianas to the Guaporé 17750, 6, 115-195 mm., Lago Cashiboya, Allen, August, 1920. 17751, 2, 102 and 105 mm., pond, Iquitcs, Allen, September, 1920.
388 FISHES OF WESTERN SOUTH AMERICA 17752, 1, 132 mm., Rio Itaya, Allen, September, 1920. 17753, 5, 126-215 mm., Contamana, Allen, August, 1920. 17754, 3, 132-188 mm., Yarinacocha, Allen, August, 1920. 17755, 1, 240 mm., Iquitos, Morris, 1922.
17756, 1, 220 mm., Manaos, Allen, December, 1920. 17757,1,180mm.,Maranon,Allen,October,1920. In general form Chaetobranchus may be thought of as bearing the same relationship to Cichla as does the croppie to the black basses of North America. Genus 201: ACARONIA Myers Acaropsis Steindachner, 1875, Sitzb. KIX. Akad. Wiss. Wien, LX-XT, 281; HKigenmann and Eigenmann, 1891, Proc. U.S. Nat. Mus., XIV, 68; Pellegrin, 1903 (1904), Mém. Soc. Zool. France, XVI, 144; FEigenmann,1910,Rept.PrincetonUniv.Exped.Patagonia,III,470; Kigenmann,1912,Mem.CarnegieMus.,V,485.
Acaronia Myers, 1940, Stanford Ichth. Bull., I, 170. Type: Acaropsis nassa Heckel Amazonia, Orinoco drainage, Guianas The characters are largely those of 4quidens; the maxillary exposed to the extremity, the premaxillary highly protractile, mouth very large. 496. ACARONIA NASSA (Heckel) Acara nassa Heckel, 1840, Ann. Wiener Mus., IT, 353, R. Guaporé; Miller and Troschel, in Schomburgk, 1848, Reisen, ITT, 624; Gunther, 1862, Cat. Fish. Brit. Mus., IV, 281. Acara (Acaropsis) nassa Steindachner, 1875, Sitzb. KIX. Akad. Wiss. Wien, LXXT, 81; Pellegrin, 1902, Bull. Mus. Hist. Nat., VITI, 182. Acaropsis nassa Kigenmann and Bray, 1894, Ann. N. Y. Acad. Sei., VII, 613; Pellegrin, 1903 (1904), Mém. Soc. Zool. France, XVI, 180; Regan, 1905, Ann. Mag. Nat. Hist., (7), XV, 345; Eigenmann, 1910, Rept. Princeton Univ. Exped. Patagonia, III, 470; Higenmann,1912,Mem.CarnegieMus.,V,485.
AcaronianassaMyers,1940,StanfordIchth.Bull.,I,170. Guianas, Orinoco, Amazons 17794, 1, 240 mm., Manaos, Allen, December, 1920. This specimen, larger than any described from the Guianas, shows some departures from type.
Scales deeply embedded; spines of dorsal and anal fins very stout, and arranged alternately with one another; the membranes of these fins and of the caudal much thickened, dark, and opaque; markings obscured, except for a caudal ocellus; posterior border of operculum with about twenty small, irregular, dark spots; vertical bands faintly perceptible; the lateral line decurved just before its interruption, and separated from the dorsal fin by 5-73 rows of scales.