<I>Chaetodon Aya</I> and Related Deep-Living Butterflyfishes

CHAETODON AYA AND RELATED DEEP-LIVING BUTTERFLYFISHES: THEIR "ARIA TION, DISTRIBUTION AND SYNONYMY CARL L. HUBBS Scripps Institution of Oceanography University of California, San Diego La Jolla, California

ABSTRACT The phyletic line of the Chaetodontidae here treated seems to comprise four species referable to Chaetodon (lalcifer. of the eastern Pacific, from islands off Baja California; aya, of the Gulf of Mexico and the South Atlantic coast of the United States; guyanensis, known from the Antilles and French Guiana; and marcellae, from tropical western Africa) plus the highly modified Prognathodes aculeatus, of the Antillean and Caribbean region, which parallels such Indo-Pacific genera as Forcipiger. Knowledge of this group has been increasing exponentially. All five species live at greater depths than are usually inhabited by chaetodontids. The known captures and observations of these species are tabulated. Their distribution, ecology, meristic variation, and other characters are treated. Seven of the 141 counts of principal caudal rays depart from the normal formula of 9 + 8 = 17: 4 specimens of C. aya and 3 of P. aculeatus have 8 + 8 = 16 rays, and 1 of the 3 counted specimens of C. guyanensis has 9 + 9 = 18 rays, which number is highly aberrant for any percomorph with a forked caudal fin. In having only 4 soft-rays in one pelvic fin, one specimen among 155 counted deviates from the almost invariable pelvic-ray formula, I, 5, that characterizes almost all percomorphs with well-developed pelvics. Direct observations are added on the life ways of Prognathodes aculeatus and of Chaetodon falcifer.

INTRODUCTION The group of deep-living chaetodont fishes herein discussed was first treated by Hubbs and Rechnitzer (1958), as an extension to their description of Chaetodon falcifer, a new species from Guadalupe Island, in the warm-temperate eastern Pacific off Baja California, Mexico. Since that paper was written I have continued to gather information on this previously little-known group of butterflyfishes, with the generous and extensive aid of many colleagues, listed below under Acknowledgments. Since the first species of this group was described in 1860, knowledge of the series has increased exponentially, with a particularly rapid rise during the past few years. This expansion in knowledge, which reflects the increase in exploratory effort, first slow and lately rapid, can be visualized by graph- ing, for the period of an approximate century (1860 through 1962), the cumulative numbers known in each year of valid species, of nominal species, and of records (collections and scientific observations) (Figure 1). The recent increase would be still more spectacular, if total numbers of

IContributions from the Scripps Institution of Oceanography, New Series. ]34 Bulletin of Marine Science of the Gulf and Caribbean [13(1)

9- r I I I 8- r I I I 80 7- r (/) I W I U , 70 ~ 6- ,---J (/) (/) PROGNATHOOES o I ACULEATUS a: I 60 o o"- 1 u a: ~- r' w w I a: 00 I I ~O :::;: I o"- ~ 4- C MARCELLAE a: Ir------~ w W 00 > I C GUYANENSIS :::;: I ::> ~ 3- r-----.J NOMINAL SPECIES z ...J I w ::> I :::;: I > ::> CHAETOOON AYA >= u ~ ...J ::> :::;: ::> u

C FALCIFER 1860 70 80 90 1900 10 20 30 40 ~O 1960

FIGURE 1. Increase in knowledge of the 5 species of chaetodontid fishes herein treated, from the discovery of 1 species in 1860 until 1962; expressed in terms of the cumulative number of valid species known, of nominal species described, and of number of published records. The disjunct column at the right shows the distribution by species of 87 records accumulated by 1962. The records are tallied in Table 6. One record each for C. aya (39a), C. guyanensis (41a) and two records (87 and 90) for P. aculeatus were received too late for inclusion on this figure. known specimens were graphed. A similar demonstration could be shown for many other groups, particularly for those representing species living at moderate to great depths, which are being so effectively collected by the OREGON and other exploratory fishing vessels. It is greatly to the credit of the officials concerned that the rich scientific harvests of the exploratory cruises are being exploited. The advancing knowledge of each of the five species of this group is documented by the chronologically tabulated list of collections (Table 6) and by an annotated synonymy, in which I have attempted to include all pertinent references. By oddly compounded coincidence each of the four Atlantic species was twice named, three of them within a one-year period. This species group was characterized, its relationships discussed, and the constituent species compared, by Hubbs and Rechnitzer (1958), by Durand (1960), and by Bauchot and Blanc (1961). The relationship of 1963] Hubbs: Chaetodon Aya and Related Butterflyfishes 135 Prognathodes aculeatus to this assemblage in Chaetodon rather than to the superficially similar Forcipiger was further noted by Randall (1961).

ACKNOWLEDGMENTS As noted above, many colleagues, of various institutions, have greatly and generously assisted me by providing specimens and data for this study. Among these, in alphabetic order, are W. W. Anderson, U. S. Bureau of Commercial Fisheries Biological Laboratory, Brunswick, Georgia; M.-L. Bauchot and Maurice Blanc, Museum National d'Histoire Naturelle Paris; M. Boeseman, Rijksmuseum van Naturlijke Historie, Leiden; James E. Bohlke and Charles C. G. Chaplin, Academy of Natural Sciences of Philadelphia; Harvey R. Bullis, Jr., Pascagoula Fishery Station, U. S. Bureau of Commercial Fisheries; Jean Cadenat, Institut Fran~ais d'Afrique Noire; David K. Caldwell, formerly of U. S. Bureau of Commercial Fishe- ries Biological Laboratory, Brunswick, Georgia, now of Los Angeles County Museum; Daniel M. Cohen, U. S. Bureau of Commercial Fisheries Laboratory, U. S. National Museum; C. E. Dawson, formerly of Bears Bluff Laboratories, now of Gulf Coast Research Laboratory; Myvanwy Dick, Museum of Comparative Zoology, Harvard College; Jean Durand. Museum National d'Histoire Naturelle, Paris; Donald S. Erdman, Puerto Rico Departamento de Agricultura y Comercio; Henry A. Feddem, Marine Laboratory, University of Miami; Charles Guthrie, U.S.N.; J. J. Hoede- man, formerly of the Zoblogisch Museum, Amsterdam; Paul Klihsbauer, Naturhistorisches Museum Fischsammlung, Wien; Robert H. Kanazawa, U. S. National Museum; Gerrit Klay and Ingvar Kristensen, Caraibisch Marien-Biologisch Instituut, Cura~ao; Francesca LaMonte, American Mu- seum of Natural History; John B. Lewis, the Bellairs Research Institute of McGill University, Barbados: the late Conrad Limbaugh, Scripps Institu- tion of OceanolZraphy; G. Robert Lunz, Bears Bluff Laboratories; N. B. Marshall, British Museum (Natural History); Giles W. Mead, formerly of U. S. Bureau of Commercial Fisheries Laboratory, U. S. National Mu- seum, now of Museum of Comparative Zoology, Harvard College; G. F. Mees, formerly of Rijksmuseum van Naturlijke Historie, Leiden, now of Western Australian Museum; George C. Miller, U. S. Bureau of Commer- cial Fisheries Biological Laboratory, Brunswick, Georgia; Robert Rush Miller, University of Michigan Museum of Zoology; Max Poll, Koninklijk Museum voor Midden-Afrika, Tervuren, Belgium; Keith W. Radford, Scripps Institution of OceanolZraphy; John E. Randall, formerlv of Marine Laboratory, University of Miami. now of University of Puerto Rico; An- dreas B. Rechnitzer, formerly of Scripps Institution of Oceanography, now of Autonetics, Anaheim, California; C. Richard Robins, Marine Labora- tory, University of Miami; Richard H. Rosenblatt, Scripps Institution of Oceanography; Leonard P. Schultz, U. S. National Museum; Walter A. 136 Bulletin of Marine Science of the Gulf and Caribbean [ 13(1) Starck, 11, Marine Laboratory, University of Miami; James R. Stewart, Scripps Institution of Oceanography; Ethelwynn Trewavas, British Museum (Natural History); Robert L. Wisner, Scripps Institution of Oceanography; F. G. Wood, Jr., Marineland, Florida; Loren P. Woods, Chicago Natural History Museum; Jacques S. Zaneveld, formerly of Caraibisch Marien- Biologisch Instituut, Curac;ao, now of College of William and Mary. Others have supplied information on the lack of material in other institutions and on the apparent absence of Chaetodon falcifer on the Mexican mainland. Though it is difficult aT'\d rather unfair to select from this long list of names for special acknowledgment, I must say that Caldwell, Dawson, Mees, G. C. Miller, Randall, Starck, and Zaneveld have all abstained in my favor from publishing on the western Atlantic species; Mees in particular had a manuscript on Prognathodes aculeatus in preparation and anticipated me in the study of critical material at Leiden and London; Boeseman supplied extensive notes from Mees's manuscript. Robins, Randall, Starck, and particularly Zaneveld, made available ecological and life-color pates on P. aculeatus. Stewart collected Chaetodon falcifer at a second locality and made further observations on this species. Limbaugh, Rosenblatt, and others searched for it along the Baja California coast. Radford and Wisner assisted by making radiographs and in other laboratory operations. Randall and Robins offered valued suggestions on reading the manuscript. In gathering data for this paper in many museums, in studying material at La Jolla, and in preparing the manuscript, I again owe much to my good wife, Laura. This report represents one phase of a program supported by a grant from the National Science Foundation.

MATERIAL During trips to various American and European institutions and through the kind intervention of colleagues I have endeavored to obtain data on the provenance and characters of all preserved specimens of the five species herein treated, and to obtain information on all the scientific observations that have been made on them in the field. The known collections and observations are listed in Table 6, chronologically for each species. With the kind permission and cooperation of the responsible officials or of associated scientists, listed above under Acknowledgments, I have examined the specimens that are deposited in the following collections, listed in the alphabetical sequence of the abbreviations herein adopted (the list includes a few collections, !!ive:l within square brackets, that are reported to contain specimens that I have not studied) : AMNH, American Museum of Natural History, New York. ANSP, Academy of Natural Sciences of Philadelphia. BBL, Bears Bluff Laboratory, Wadmalaw Island, South Carolina. 1963) Hubbs: Chaetodon Aya and Related Butterflyfishes 137 BLBG, Bureau of Commercial Fisheries Biological Laboratory, Brunswick, Georgia. BM (NH), British Museum (Natural History), London. CAS, California Academy of Sciences, San Francisco. [CMBI, Caraibisch Marien-Biologisch Instituut, Piscadera Baai, Curalfao.J CNHM, Chicago Natural History Museum. [COlEC. Centre d'Oceanographie de l'Institut d'£tudes Centrafricaines de Pointe-Noire.] [IF AT, Institut Franlfais d' Amerique Tropical a Cayenne.] IJ, Institute of Jamaica, Kingston. IMBLP, Institute of Marine Biology, La Parguera, University of Puerto Rico. ISNB, Institut Royal des Sciences Naturelles de Belgique. LACM, Los Angeles County Museum. [MNHN, Museum National d'Histoire Naturelle, Paris.) MCZ, Museum of Comparative Zoology, Harvard College. [NMFW, Naturhistorisches Museum Fischsammlung, Wien.J RMNHL, Rijksmuseum van Naturlijke Histoire, Leiden. SIO, Scripps Institution of Oceanography, University of California, San Diego. U Fla., University of Florida, Gainesville. UMML, University of Miami Marine Laboratory. UMMZ, University of Michigan Museum of Zoology, Ann Arbor. UPR, University of Puerto Rico, Mayaguez. USNM, United States National Museum, Washinl!ton. [UZMK, Universitetets Zoologiske Museum, K4>benhavn.] ZMA, ZoOiogisch Museum, Amsterdam.

An unsuccessful search for specimens of the species here treated was conducted at several other institutions, including the museums at Bergen, Stockholm, Lund, Goteborg, and Hamburg in Europe, and at Tulane Uni- versity, New Orleans, and University of Texas, Austin. Except for a few specimens of Chaetodon aya and of Prognathodes aculeatus from duplicated regions, the only known specimens that I have not studied are the single or few types of the following nominal species: Chaetoe/on eques (synonym of C. aya), C. guyanensis (on which Jean Durand has provided additional data), C. marcellae, and C. altipinnis (synonym of C. marcellae). Each entry in the synonymies and in the literature cited is accompanied by a number, or numbers, in parentheses, corresponding to the consecu- tively numbered items in the list of known collections and observations (Table 6), to indicate the collection(s) and observation(s) that were indicated or are assumed to have been utilized in the given treatise. t 38 Bulletin of Marine Science of the Gulf and Caribbean L 13(1)

DISTRIBUTION AND SPECIATION The recently augmented data strengthen the conclusion of Hubbs and Rechnitzer (1958: 307-308) that the distribution of this apparently natural group of relatively deep-living chaetodontids illustrates "two zoogeogra- phical tendencies that are being increasingly indicated by critical taxonomic studies; namely, (1) the tendency for New World fishes to possess a marked independence and integrity, indicative of lines of evolution essentially distinct from those of related Old World types, and (2) thc tendency for New World types to be sparingly but definitely represented in the fauna of tropical west Africa." The one Pacific representative of the group, Chaetodon faicifer, is sharply differentiated from the three closely related Atlantic species of the genus Chaetodon (aya, guyanensis, and marcellae). This circumstance is consonant with the evidence that the Panama portal was closed in Miocene or early Pliocene time. This limited representation in the Pacific of this essentially American group is consistent with the generally recognized de- pauperate character of the coral-reef fauna in the eastern Pacific: a circumstance that I have attributed (Hubbs, 1952) to the decimating effects of Pleistocene cold. Chaetodon falcifer seems to be holding out, as a relict, only on islands (Guadalupe and the San Benitos) off Baja California, on the northern fringe of the warm-water fauna (Figure 2). Intensive diving and collecting in relatively deep water has disclosed the presence of this species at West San Benito Island, and its apparent absence about Cabo San Lucas, as well as in Golfo de California and elsewhere in the eastern Pacific. No plausible explanation for the extremely restricted distribution of this species comes to mind. The group is chiefly characteristic of the warm waters of the western North Atlantic, where three of the five species occur (Figure 2). The three are essentially complementary in distribution. One, Prognathode.l' aeuleatus, a highly specialized derivative, is now shown to be widely distributed through the Greater and Lesser Antilles, including the Bahamas. It widely overlaps the range of the recently discovered Chaetodon guyanensis, but the two are probably segregated ecologically: P. aeuleatus lives chiefly on the coral-encrusted slopes, in shallower water, whereas C. guyanensis, to judge from the three known records and from its very close relationship with C. aya, probably lives chiefly in the outer part of the continental shelf (Figures 2 and 4). Prognathodes aeuleatus and Chaetodon aya seem to be largely allopatric, as they are now known to overlap in distribution only about the Florida Keys and in southern and southeastern Florida, and even here are presumably separated ecologically. Here as elsewhere the Prognathodes lives on the coral reefs, chiefly at considerable depths, whereas C. aya is definitely an inhabitant of the continental shelf, usually at even greater depths (Figure 4). 1963] Hubbs: Chaetodon Aya and Related Butterflyfishes 139

~ ~ g 140 Bulletin of Marine Science of the Gulf and Caribbean 113(1) The cognate species C. aya and C. guyanensis seem to be completely allopatric (Figure 2). Further exploration in the area intervening between their known ranges (between Puerto Rico and the Straits of Florida) may throw light on their distributional and systematic interrelations. The Puerto Rico locality for C. guyanensis is deeper than any reported for C. aya, and it was associated there with a typically bathybial fauna (Woods, 1960), but the French Guiana locality lies well within the deeper part of the bathymetric range of C. aya (Figure 4). The numerous recent collections of Chaetodon aya seem to delimit its geographic and bathymetric ran

20· 20·

AFRICA 20' 20'

o o

20' 20· o 20·

FIGURE 3. Distribution of Chaetodon marcel/ae, from all published records. pyge argi Woods and Kanazawa (1951: 636-638, fig. 135; see also Phillips, 1958). The usual depth range seems to be 20 to 40 meters for C. faldfer, 35 to 120 meters for C. aya (with no indication of regional differences), apparently about the same for C. marcellae, likely somewhat greater for C. guyanensis. Prognathodes aculeatus seems to occur most commonly in 15 to 55 meters, but occasionally ventures onto the very shallow part of the reef, and at times descends to greater depths, with records at 69, 77, and 9 J meters. Over this depth range from the surface meter to more than 142 Bulletin of Marine Science of the Gulf and Caribbean 113(1) 90 meters the species must vary its associates from typical shallow-reef species to more or less bathybial types. In general, even where occurring in rather deep water, the species ot this group remain in warm to moderate temperatures (see Table 6). The distribution of the species may be more or less limited to areas where the upper mixed zone is rather deep. This is true of Guadalupe Island, where Chaetodon falcifer has been taken at depths of about 20 meters where essentially surface temperatures prevailed (16.7°C in January, 18.3°e in

CFAL- C C.MAR- PROGNA rHODES CHAErODON AYA CIFER CELLAE ACULEArus 0 = • I•• •• 25 '--1'1 °0 ""ST • e. • II!AB~DANT 50 ~ if) • a: • •••• w ° • f- "" W ·=00•• • :::;: 75 • ~ ·~~o~ • :I: • • f- • Q.l 00 w o .. • °1 0. 25

° FOR C. AYA ONLY 50 ° SOUTH ATLANTIC COAST () STRAITS OF FLORIDA • NORTHEAST COAST OF MEXICO CAMPECHE BANK 175 " • •

2 FIGURE 4. Bathymetric distribution of the 5 species of chaetodontids herein treated; by specified regions for Chaetodon aya. For this species, trawling records with limiting depths are indicated by a hollow vertical bar, with a circle at the median depth. The dark vertical bars for Prognathodes aculeatus represent the range over which the species was collected or observed. Some of the depth records are approximated. The observations showing differential abundance are listed as 85 and 86 in Table 6 and are discussed on page 174. Data are plotted for all collections and observations except 5-7, 39a, 41a, 45, 48-53, 57, 58, 61, 66, 68, 81, 87, and 90. 1963] Hubbs: Chaetodon Aya and Related B utterfly[ishes 143 November). Available temperature records for C. aya vary from 12.8 to 27.2°C, usually from 15 to 22°. The one record for C. marcellae is about 16.5°C at 85 meters. Prognathodes aculeatus must inhabit a wide range of temperature: in one 5-meter habitat at Dominica, Charles C. G. Chaplin recorded temperatures between 24 and 27°C; at depths as great as 91 meters the water must be colder (though no definite record seems to be available) .

MERISTIC VARIA nON The relatively abundant new material has made possible an analysis ot meristic variation (Tables 1 and 2) in the Chaetodon aya group (including Prognathodes aculeatus). Throughout the group the dorsal spines are usually constant in number (13) as well as in their strength, height, and excision. The number occasionally varies to 12 or 14 in Chaetodon aya and in Prognathodes aculeatus, and will presumably be found to vary similarly in the three other species, for which few counts are yet available. The number of dorsal soft-rays is more variable but seems to be essentially similar in range and in mean in four of the five species. The number is higher in C. falcifer, with some overlap, at least with the counts for C. aya. The anal spines consistently number 3 in the 158 specimens counted for the five species-as might be expected, since this is the normal number for most acanthopterygian fishes. The anal soft-rays number usually 15, occasionally 14 or 16, in C. aya and P. aculeatus, and are not certainly different in the other species. The formula of principal caudal rays is normally 9 + 8 = 17, which is characteristic of most acanthopterygians with a forked caudal fin. This number holds for 133 of the 141 specimens counted. In 4 specimens of C. aya and in 3 of P. aculeatus, with apparently uninjured fins, the number is reduced to 8 + 8 = 16. The type of C. goniodes (Woods, 1961) has the much more unusual, all but unique, increase in number to 9 + 9 = ]8, which led to the speculation that this species might be characterized by a primitively higher number of caudal rays. The two available type specimens of C. guyanensis (of which C. goniodes is a synonym), however, have the normal formula of 9 + 8, according to Jean Durand (personal communication, 1961), and we may assume that the high number in the type of goniodes was a rare coincidence. The pectoral rays vary from 13 to 15 and are usually 14 in C. aya and in P. aculeatus. The average number is apparently higher, usually 14 or 15, in the three other species, but the number of counts is inadequate. Normally the first (upper) principal ray, as well as the short ray above it, is unbranched, and the lowermost ray is either unbranched or narrowly branched. 144 Bulletin of Marine Science of the Gulf and Caribbean 1/3(1 ) In respect to bilateral variation in number of pectoral rays (Table 3) the two species with an adequate number of counts agree with the rules that we have found to hold for fishes in general (Hubbs and Hubbs, 1945). The percentage of asymmetry-of specimens with the counts different on the two sides-is 16 for C. aya and 7 for P. aculeatus; 12.5 for the combined counts of the two species. There is a bare suggestion that in C. aya the higher count occurs more frequently on the right than on the left side, as it clearly does in Leptocottus armatus (Hubbs and Hubbs, 1945: 264- 272). Ordinarily, as I find it to be a virtually universal rule, the lowermost ray is the shorter on the side with the higher count. As is usual and to be expected, the chance of asymmetry is negatively correlated with the frequency of the count; that is, the more deviant the count, the greater the chance for it being different from the count of the fin on the other side. Positive correlation is indicated between the counts of the two sides, but the deviates are too few to yield a highly significant coefficient of correlation. Among the 310 counts of pelvic rays available for the five species there is one variant: one specimen of Chaetodon aya violates the almost universal rule that percoid fishes with unreduced pelvics have I, 5 pelvic rays. This is one more example of the generalization that variation is sufficiently universal that we may expect, even though very rarely, to encounter deviation in the counts of even the most meristically stable characters. This specimen has, significally on one side only, in an apparently uninjured fin, only I, 4 rays. The number of pored scales in the lateral line (Table 2) varies rather widely and differs in the several species. The number is low (22 to 28) in Prognathodes aculeatus and high (42 to 46) in Chaetodon falcifer. The three very closely related Atlantic species have an intermediate number, overlapping on the range of counts for the Prognathodes. There is a suggestion of an increase in the average number from C. aya through C. guyanensis to C. marcellae. The vertebral count is probably uniform in the five species. though we have only a few counts, 3 for C. falcifer, 7 for C. aya and 13 for P. aculeatus. The number in all 23 specimens, as also in 4 of C. humeralis Gunther, is 10 + ] 4 = 24, which has long been interpreted as standard for pcr- comorph fishes, through the deviation from this number, even in the central ~TOUp of Serranidae, is greater than is generally realized (Hubbs, ] 96'0: 382-383). That the vertebral formula (though not the total number of vertebrae) diverges within the Chaetodontidae has been discovered by Randall (1961: 55), who found that Forcipiger has 11 + 13 rather than ]0 + 14 vertebrae. Since Forcipiger seems to be related to Heniochus, forming with it and certain other genera a seemingly natural group having a complete lateral line (Hubbs and Rechnitzer, 1958: 296-300), radio- 1963] Hubbs: Chaetodon Aya and Related Butterflyfishes 145 graphs were prepared of Heniochus nigrirostris to see if it too might have II + 13 vertebrae, but it proved, on the basis of counts of 15 specimens from Bahia San Lucas, Baja California Sur, to have the "standard" formula of 10 + 14. Since neither Chaelodon aya nor Prognathodes aculealus shows any ob- vious geographical variation, all counts of each species were combined in an analysis of correlation between the number of rays in different fins (Table 4). In line with the virtually universal trend (Hubbs and Hubbs, 1954: 192-193), there is an indication of negative correlation between the anterior and posterior elements in a linear series (dorsal spines and dorsal soft-rays) and of positive correlation between opposed elements (anal soft-rays and dorsal soft-rays).

(f) W 10 (f) (f)

•o C FALCIFER

20 40 60 80 100 120 140

STANDARD LENGTHS, mm. FIGURE5. Size-frequency distribution (standard length) by 5-mm classes of the known specimens of the five species herein treated, based on the published records for Chaelodon guyanensis and C. marcellae and on original data for the other species. Included, by courtesy of George C. Miller, are measurements of 40 specimens of C. aya that were taken with collection 38 (see Table 6), but were discarded after being measured. Five specimens of C. aya of collection 39a (65, 73, 76, 78, and 87 mm long) were measured too late for inclusion. Specimens of Prognathodes aculeatus 53 and 58 mm long, from collections 87 and 90, respectively, are not plotted. 146 Bulletin of Marine Science of the Gulf and Caribbean [ 13(1)

SIZE AND LIFE SPAN In comparison with chaetodontids in general, Chaetodon falcifer is relatively large, Prognathodes aculeatus definitely small, and the three At- lantic species in this group of Chaetodon about medium (Figure 5). The complex form of the size-frequency graph for C. aya suggests a life span of several years, whereas the sharply unimodal size distribution of the adults of the small species, P. aculeatus, suggests definite size at maturity and, possibly, an annual life cycle.

DESCRIPTIONS OF SPECIES Chaetodon falcifer Hubbs and Rechnitzer Figures 1, 2, 4-7

Chaetodon falcifer Hubbs and Rechnitzer, 1958 (I) 2: 273-294, 297, 298, 300, 303-305, 307, 308, map 1, pI. I (original description; camparisons and relationships; endemism; associated species; Guadalupe Island. Baja California, Mexico).-Myers, 1959 (1): 158 (reference; relationships).- Woods, 1961 (1): 420 (comparisons; relationships) .-Bauchot and Blanc, 1961 (1): 89-91 (compared with related species, C. marcellae and C. aya). Material and distribution.-Two additional records arc now available for this very distinctive eastern Pacific relative of Chaetodon aya. For both I am indebted to the keen eye and diving skill of James R. Stewart of Scripps Institution of Oceanography. On January 26, 1960 he saw one specimen, which he estimated to be about 40 mm in standard length, at a depth of 21 meters, on the northeast side of Guadalupe Island, at 28° 18' 30" N. Lat., 1150 36' 05" W. Long., 21 kilometers north of the type locality; field number SIO 60-1'O-32(A). On February 28, 1959, he saw and collected by spear 2 fine specimens, 102 and 137 mm long, off the west side of West San Benito Island, at a depth of about 38 meters; Cat. No. SIO 59-46-32(A). From an examination of smears of gonad tissue, the larger specimen appears to be a male; the smaller one, with higher spines, to be a female. Thus, Chaetodon falcifer is not, as was originally thought probable, one of the Guadalupe Island endemics. Its occurrence on Guadelupe and the San Benitos, with no indication of occurrence elsewhere, parallels the distribution of Gibbonsia nome Hubbs (1952: 126-130). Next to Guada- lupe and the little explored, more tropical Alijos Rocks, the Benitos are the most offshore of the islands off the west coast of Baja California, and they lie approximately opposite Guadalupe. Since C. falcifer was described, much additional collecting and observing in relatively deep water around the Cape region of Baja California and on the Golfo de California coast

2Figures in parenthesis in the synonymies, entered after the date in each item, refer to the chronological list of collections (Table 6), and indicate the collection stated, or assumed, to have been used in the given reference. 1963] Hubbs: Chaetodon Aya and Related Butterflyfishes 147

FIGURE 6. The specimens of Chaetodon falcifer from West San Benito Island, Baja California Norte, 102 and 13? mm in standard length. See Table 6, collection 2. Photography by Scripps Institution of Oceanography. 148 Bulletin of Marine Science of the Gulf and Caribbean 113(1) of the peninsula, by Richard H. Rosenblatt and others, has failed to locate the species on the mainland coast, though a close watch has been main- tained for it by him and others. It does indeed seem to be endemic to Guadalupe and the San Benitos. The San Benito specimens were taken together about 2 or 3 meters off the bottom, in very clear (probably oceanic) water. The bottom was of rubble rock, with blue coral, at the base of a rock pinnacle, about 2 km offshore, that rises to within about 6 meters of the surface. The new Guadalupe locality was among lava boulders close to the boulder-strewn beach at the base of high lava cliffs; there were growths of brown algae (Eisenia and Sargassum). On the same dive the collectors, Jay C. Quast and James R. Stewart, secured other tropical types (Holacanthus passer, Apogon guadalupensis, and Pseudojulis sp.).

Characters.-The original diagnosis holds, except that there are apparently only average differences between this species and its closer relatives in the number of dorsal and pectoral soft-rays and little if any difference in the number of anal soft-rays. The indicated increased number of teeth, as is indicated below, apparently merely reflects the larger size attained. The ray and pore counts on the San Benito specimens are included in Tables 1 and 2. The vertebrae number 10 + 14 = 24 in each. The scales that impinge above on the lateral line number 62-64 and 60-60; on each side of each specimen there are 16 scales in the row from the edge of the sheath at the base of the first dorsal spine downward and backward to (but not including) the lateral line; 25 to 27 in the row from the edge of the scaly sheath at the side of the first anal spine to the lateral line; about 28 and 29 around the caudal peduncle. The lateral line ends close to the end of the soft dorsal fin. The rows of fine inner teeth in the upper jaw number 8 in the smaller specimen and about 12 in the larger specimen; in the lower jaw, 8 and about 15, respectively. The new specimens agree closely with the description of the type. Minor differences are indicated in the following account, in which the measurements, when entered, are given first for the larger specimen, then, in parenthesis, for the smaller one. The extreme base of the soft dorsal measures 1.7 (1.6) in the spinous dorsal base (the statement was reversed in the original description). The height of the dorsal spines varies greatly, and is much greater in the smaller specimen than in the larger. This is possibly a sexual difference, with the spines longer in the female. In the larger specimen, the third and fourth dorsal spines are highest and subequal, whereas in the smaller one, the third is definitely higher than any of the others. In the larger specimen, the third and fourth spines, as measured from the scaly sheath, each enter the soft-dorsal base about 1.4 times, whereas in the smaller specimen the fourth spine from the sheath is about as long as the soft-dorsal base and 1963] Hubbs: Chaetodon Aya and Related Butterflyfishes 149 the third spine is about 0.1 longer. The least width of the membrane in front of the two longest spines in the larger specimen is little less than one-fourth the height of the spine, whereas in the smaller specimen the height of the membrane enters the height of the spine about 7 times for the third spine and about 4.5 times for the fourth spine. The margin of the soft dorsal starts to curve downward from the first ray. The second anal spine when depressed falls somewhat short of the tip of the third spine in the larger specimen, but passes beyond the third spine for about O. 1 this length in the smaller specimen. The free front edge of the second spine is about as long as the soft-anal base in the larger specimen and about 0.1 longer in the smaller one. The membrane in front of the second spine is attached on one side of the spine very close to the edge of the scaly sheath. The pectoral fin enters the head length 1.3 (1.2) times. The free length of the pelvic filament is somewhat less than half as long as the orbit in the larger specimen, but is somewhat longer than the orbit in the smaller one. There is a slip in the original description, wherein the length of the pelvic is said to be equal to that of the snout plus orbit; this statement applies to the length of the pelvic spine. In the specimens at hand, the length of the pelvic spine in the larger specimen is a little less than the snout plus orbit; in the smaller specimen, a little more than the snout plus orbit. The direction of scale rows shows some variation in the two new specimens, rendering a little less impressive the indications of the value of this character in the separation of species groups. On the left side of the smaller specimen, above the axis of the body, the scale rows in the light area below the scythe-like mark slope upward considerably; below the axis, the rows over the body cavity are curved weakly downward. On the right side the rows are more nearly horizontal throughout this area. There is a tendency for the rows to arch downward a little above the anal spines. In the smaller specimen, the scale rows behind the pectoral fin are arched slightly upward on the left side, but are nearly horizontal on the right side. In both specimens small rough scales extend across the premaxillaries in front of the rostral groove. There is a narrow scaleless area both above and below each nostril on each side. The scaleless groove in front of the anterior nostril is weak in the smaller specimen and almost obliterated in the larger one, verifying the prediction in the original description that this character will prove to be variable. In each specimen scales are obsolete in the denticulate bony area above and behind the upper bony orbital rim, and on the upper fleshy orbital rim. In the smaller specimen the squamation of the caudal fin is essentially as described for the type, but in the larger one small ctenoid scales cover the broad caudal rays almost to the tip (the rows of scales branch to correspond with the branching of the rays). The anterodorsal profile from the dorsal origin to the snout forms an angle 150 Bulletin of Marine Science of the Gulf and Caribbean [13(1) of 56° (58°) with the body axis, and an angle of 87° (85°) with the chord of the anteroventral contour behind the muzzle. The angle between the dorsal profile of the snout and the profile between the snout and the dorsal fin is 145° (149°); the angle of the muzzle is 5'0° in lateral view. The greatest width of the muzzle in front of the mouth steps 1.6 (1.7) times in the length of the snout. The length of the head steps 2.7 (2.85) in standard length. The lower border of the upper jaw forms an angle of 29° (27°) with the body axis. The length of the upper jaw steps 3.35 (3.65) times in the head. In the larger specimen the midline length of the upper lip is more than half the length of the orbit, but in the smaller one it is just half the length of the orbit. The lower fissure in the upper lip is about two-thirds as long as the orbit. The midline length of the tooth patch in each jaw varies from 0.25 to about 0.2 the length of the orbit. The gill-rakers, including rudiments, number 4 + lIon the outer arch on each side of each specimen. The margins of the facial bones are variously denticulate. The preorbital border in the larger specimen is weakly denticulate on one side, near the middle, and obsolescently denticulate toward either end; on the other side, the margin is nearly smooth near the posterior end, but elsewhere is rather strongly denticulate, with the denticulations definitely increasing in strength toward the squarish anterior corner. In the smaller specimen the preorbital denticulations are obsolescent, except near the posterior end. The preoper- cular denticulations are rather uniformly developed in the smaller specimen, but are obsolescent over part of either the upper or lower limb in the larger one. There is a slight tendency for the denticulations to become somewhat enlarged just above the angle, in the region where they are directed backward. The exposed borders of the scapular processes are also variable in denticulation, from obsolescent to fairly well developed. The upper border of the orbit is roughened by several series of minute denticulations in the larger specimen, but in the smaller one these denticulations are fully developed, except in one outwardly directed series along the extreme margin. In each specimen, but more strikingly so in the larger one, the region above and behind the orbit is tuberculate and denticulate. The orbital rim is scarcely expanded in the prefrontal region. Anteriorly, the bony rim departs slightly from the margin of the orbit and extends toward but not to the posterior end of the posterior nostril. The bony rim continues around the back and bottom of the eye, and somewhat up the anterior border. All along this region the extreme edge is denticulate, and there are some minute bony points outside the marginal row. Coloration.-In general pattern and color the new specimens are strikingly like the type (their color was described after nearly five weeks of preserva- 1963] Hubbs: Chaetodon Aya and Related Butterfiyfishes 151 tion in formalin, but had changed little). The scythe-like mark, and the other black markings, are almost identical in position. The main markings are deep brown-purple, almost jet-black. The course of the main mark across the opercles is essentially as described for the type. The larger specimen, like the type, has dusky dapplings on the adjoining branchiostegal membrane, but the smaller one has blackish streaks along the branchioste- 0 0 gals. The axis of the mark anteriorly forms an angle of about 65 (61 ) with the axis of the body. The angle of the axis near the change in course 0 0 is about 87 , rather than about 80 as in the type. The upper angle of the mark is approximately under the fourth rather than the fifth dorsal spine, but the separation, as in the type, is approximately two-thirds the length of the orbit. The apex of the lower border of the mark lies below the posterior exposed base of the fifth dorsal spine, in either the second or the third scale row below the lateral line. The posterior course of the mark is approximately as in the type, except that the mark ends on the fourth rather than the sixth anal soft-ray. The lower border of the mark crosses the lateral line at the extension of the course of the tenth or eleventh (eleventh or twelfth) dorsal spine, as the fin is normally expanded (a little farther forward than in the type). The posterior course of the mark is very much as in the type. The separations from the end of the dorsal and anal fins, and from the base of the caudal, are as described for the type. The width of the band, measured as in the type, steps 2.6 (2.2) times in the head length. The least width, not far behind the apex, measures 3.6 (2.7) times in the head. The anterior extension of the anterior bar extends from the eye to the rostral fold, but here is weakly dusky, less conspicuous than in the type. The lower border, above the mouth, is somewhat oblique. The narrow light area in front of the band, from above the eye nearly to the dorsal fin, is pale yellowish cream in the smaller specimen, but purplish gray in the larger one. There is a purplish area behind the upper end of the anterior band, under the second and third dorsal spines. On the body a yellow streak follows the anterior black band ventrally, then runs below the purplish area just mentioned, to reach a point below the fourth dorsal spine (as exposed). This yellow streak meets another one, less distinct, along the front of the upper anterior portion of the scythe-like mark. The anterior bar is not very noticeably blacker at the edges, and remains almost jet-black to the dorsal fin. Near the junction of the yellow stripes described above, under the fourth dorsal spine, as exposed, there are, particularly in the larger specimen, rather black marks on the center of the scales, on a lighter base with purplish reticulations. The lighter marks represent the silvery specks seen in the holotype after preservation. The greatest width of the anterior band, where well defined, is about 0.75 that of the orbit. The blue glints, in a fine pattern, still remain on the sides of the head 152 Bulletin of Marine Science of the Gulf and Caribbean [/3(1) in front of the scythe, especially on the opercles. A trace remains of the very narrow yellow edge of the orbit posteroventrally. The posterior arm of the scythe in the larger specimen is bordered by yellow-gilt, and the lower border of the scythe on the body has some concentration of blue, but these marks are not very evident on the smaller specimen. Above and behind the scythe, and above the caudal peduncle, the color is yellowish, with a purplish gray reticulum around the scales. In the larger specimen the yellowish color predominates. but in the smaller one. except for some mottlings, purplish gray predominates. Small silvery specks remain in both specimens along the lateral line, where it is exposed behind the scythe. The caudal peduncle remains yellow-brown, becoming almost pure yellow ventrally. Below the scythe the sides are extensively brightened by silvery blue reflections, and there is a broad purplish streak along the middle of each scale row, particularly in the larger specimen. In the two new specimens. this pattern remains distinct anteriorly. The narrow ventral surface is lemon-yellow, and this color extends onto the pelvic and anal fins, most conspicuously behind the spines. The outer part of the pelvic fin becomes almost blackish. The anal is pale purplish gray near the scaly base, but outward, becomes blackish brown. particularly along the interradial membranes. The pelvic filament is clear whitish. The anal spines are pale yellowish. The outer edge of anal, except on the first, or first and second soft-ray, is abruptly clear. The pectoral is very pale purplish, with very fine streaks along the edges of the rays and their branches. The upper margin is not very conspicuously darkened. The caudal fin has some purplish on the rays, particularly on the larger specimen, and the rays and branches are finely marked by blackish lines, obscurely so anteriorly. The upper and lower borders are black. The spinous dorsal fin is almost entirely black, backward to and including most of the third spine. Most of the fourth spine is blackened, but the membranes behind the third and behind the fourth spine are pale yellowish. with some purplish. The posterior part of the spinous dorsal above the scaly base is almost entirely blackish, but there is a light purplish base anteriorly, from the fifth spine backward, in an area narrowing posteriorly to the eighth spine. This marking is conspicuous only in the smaller specimen, and is largely obscured by dapplings of the dark color in the larger one. The submedian blackish mark on the exposed part of the dorsal fin is almost solid in color throughout in the smaller specimen, and not very conspicuously dappled with light dorsally in the larger one. The rays are lighter than the membranes in this mark, which is slightly scalloped by conspicuous clear membranes along its edge. The main body of the light border on this fin, more conspicuously than on the anal, is slightly purplish, due to the fine bordering of the branches of the rays. 1963] Hubbs: Chaetodon Aya and Related Butterflyfishes 153

FIGURE 7. Radiographs of the specimens of Chaetodon falcifer from West San Benito Island, Baja California Norte, shown in Figure 6. Photography by Keith W. Radford. . 154 Bulletin of Marine Science of the Gulf and Caribbean [J3(1) Comparative study of skeletal features.- The skeletal features of all 3 specimens are in essential agreement. The vertebrae number 10 + 14 = 24 in all. The first vertebra is wedge-shaped, narrow ventrally, and the second is narrower than those following. In one of the San Benito specimens the upper 15 principal caudal rays originate on the 6 hypurals according to the same formula as in the holotype, namely 3 + 4 + 2 = 9, 1 + 4 + 1 = 6; in the other specimen, according to the formula 1 + 6 + 2 = 9, 1 + 31/2 + 1'h = 6. The 2 enlarged hemal elements that lie just in front of the hypural plate and with which the lowermost principle caudal ray and the 3 lower procurrent rays are associated, arise separately from the last 2 normal centra, rather than, as abnormally in the type, from the last centrum (in the type the antipenultimate centrum bears a hemal spine like those that precede it; none of the other specimens X-rayed (see Fig. 7) shows a comparable peculiarity; all have the two enlarged hemal elements on separate centra). The interneurals of the last 2 dorsal spines arise on either side of the neural spine of the twelfth vertebra, as in all of the other specimens X-rayed excepting 1 of Chaetodon aya and all 15 of Heniochus nigrirostris (this may therefore be a character of some value). The neural spine of the twelfth vertebra is the last one that is sagittally expanded basally, rather than the eleventh as in Chaetodon aya or the tenth as in C. humeralis, Prognathodes aculeatus, and Heniochus nigrirostris. The interneural of the second dorsal spine is deflected forward to fit between the first 2 neural spines close to the vertebrae, as in all of the specimens of the 5 species X-rayed. The interneural of the first dorsal spine extends down along the front of the succeeding interneural almost to its base, as also in Chaetodon aya and Prognathodes aculeatus; in C. humeralis and in H. nigrirostris the interneural of the first spine extends only about half-way to the base of the following spine. The 2 mOre anterior interneurals arise on either side of the first, short, suberect neural spine and remain separate to their distal tips, where they are sharply deflected forward to fill the short gap between the anterior process of the interneural of the first dorsal spine and the apex of the supraoccipital spine of the skull. This arrangement is characteristic also of C. aya, C. humeralis, and H. nigrirostris. In the last-named species these interneurals are more nearly parallel than in the others, because they are forced apart ventrally by the somewhat clavate tip of the first neural spine. A contrasting pattern is exhibited by P. aculeatus, in which these 2 interne urals fuse distally. In that species, furthermore, the more elevated supraoccipital spine instead of ending about opposite the tip of the first interneural continues backward to, or almost to, below the first dorsal spine. The nearly straight first hemal spine is broadly expanded ventrally to an oblique tip, which lies against the upper-rear edge of the very strong second interhemal and just in advance of the tip of the third interhemal. The same general arrangement holds for C. aya and P. aculeatus, but in those species 1963] Hubbs: Chaetodon Aya and Related Butterflyfishes 155 this hemal spine is typically more clavate and more arched caudad to accommodate a posterior expansion of the visceral organs. In C. humeralis and H. nigrirostris this neural spine is slenderer and is otherwise less modified. The more or less conjoined first and second interhemals extend only about half-way to the vertebral column, as also in C. aya and P. aculeatus; in C. humeralis and H. nigrirostris this compound interhemal is slenderer and extends somewhat more than half-way to the vertebral column. Obviously, there are osteological features that will be helpful in classifying the chaetodontids.

Chaetodon aya Jordan Figures 1, 2, 4, 5, 8, 9 Chaelodon aya Jordan, 1886 (6)3: 225 (original description; distinguished by coloration; Snapper Banks, near Pensacola, Florida; " 'spewed up' by a Red Snapper (Lutjanus aya)").-Eigenmann and Horning, 1887 (6): 5,8,18 (comparisons; diagnosis, from holotype; northern Gulf coast of Florida, Pensacola).-Jordan, 1887 (6): 593 (West Indies).-Jordan and Evermann, 1898 (6): 1673, 1676-1677 (comparisons; description; rather deep water, Gulf of Mexico).-Ahl, 1923 (6): 122-123 (description, after Jordan and Evermann; in section Chaetodontops, misspelled Chaedontops on p. lll).-Breder, 1929 (6): 215,217 (comparisons; Gulf of Mexico; chiefly in rather deep water; "may represent the young of other species").-Nicholas and Firth, 1939 (11): 87-88 (off North Carolina; faunal relationships; description) .-Hildebrand, in Longley and Hildebrand, 1941 (9, 10): 150-151 (Tortugas, Florida; depth distribution; color; cOunts and proportions; except smallest specimen, which is C. acel/atus, as was shown by Hubbs and Rechnitzer, 1958: 302) .-Springer and Bullis, 1956 (16-18, 20-22): 89 (records, with detailed data, for "Oregon" stations 33, 727, 728, 944, 1022, and 1024-see Table 6).- Briggs, 1958: 282 (vernacular, "bank butterfly fish"; North Carolina to eastern Gulf of Mexico; shore).-Hubbs and Rechnitzer, 1958 (6-10, 13, 14, 16, 17, 19, 26): 278, 280-282, 284, 285, 287, 288, 292, 300-308, map 1, pI. 2, fig. 1, pI. 3, fig. 1 (comparisons; depth distribution; annotated synonymy; description; compilation of published and original records-see serial numbers above and Table 6; relationships) .-Myers, 1959 (5, 6): 158 (G. eques as synonym; history of name eques; relationships) .-Bailey, el al., 1960 (6, etc.): 32 (listed as of the United States; vernacular, "bank butterflyfish") .-Durand, 1960 (6, etc.): 209-212, fig. 1 (characters; comparisons; relationships; Gulf of Mexico to North Carolina and South Carolina).-Bauchot and Blanc, 1961 (6, etc.): 89-92, pI. 1, fig. B (compared with related species, C. marcellae and C. falcifer) .-Randall, 1961 (6, etc.): 54 (Prognathodes probably derived from species like C. aya).-Woods, 1961 (6, etc.): 417, 419-420 (compared with C. goniodes [= C. guyanensis]; depth distribution). Chaelodon eques Steindachner, 1903 (5): 18-20, fig. 1 (original description; similar in coloration to C. aya). Ahl, 1923 (5): 123 (description, abridged from Steindachner).-Myers, 1959 (5): ]58 (as synonym of C. aya). --- lFigures in parenthesis in the synonymies, entered after the date in each item refer to the chronological list of collections (Table 6), and indicate the collection ;tated or ""umed, to ha' e been used in the given reierence. ' 156 Bulletin of Marine Science of the Gulf and Caribbean [/3(1 ) "Common butterfly fish."-Zim and Shoemaker, 1956: 131, col. fig. on p. 131 and on cover ("well-known reef dweller"; color; "5 to 8 in."; "fast and aggressive, feeds on small invertebrates") .-Ray, 1960: 32, 1 fig. on col. pI. (fig. after Zim and Shoemaker). Synonymy.-Myers (1959: 158) correctly pointed out that Chaewdon eques Steindachner is a synonym of C. aya. [ have not reexamined the type of C. eques, but Steindachner's excellent description and magnificent figure leave no room for doubt. Furthermore, topotypes of C. eques have been studied.

Distribution.-Thanks to the aid of numerous colleagues, I am able to expand still further our knowledge regarding this species, which until very recently was inadequately known (Figure 1). The 37 known records, summarized in Table 6, show that the half-grown and adults are rather common at moderate depths (Figure 4) on the continental shelf on Campeche Bank in the southern part of the Gulf of Mexico, off the peninsula of Yucatan; on the outer part of the broad shelf of the northeastern part of the Gulf, east of the Mississippi Delta; around the tip of Florida; and on the outer shelf waters in the course of the Gulf Stream, along the South Atlantic coast of the United States north to the vicinity of Cape Hatteras (Figure 2). No tholichthys prejuveniles of this species, however, have as yet been found. There is no record for the species from the Antilles, from the Caribbean area, or from elsewhere off South America. Those regions are indicated, by the three known locations (off Puerto Rico, off Barbados, and off French Guiana), to be inhabited by an apparently allopatric close relative, Chaetodon guyanensis. Recent exploratory fishing in the Bahamas confirms the indications that C. aya is absent there (Harvey R. Bullis, Jr., personal communication, 1962). Nor are there any records from the western or northwestern shores of the Gulf of Mexico. That there are no extant unpublished records from this region seems evident from the information supplied to me by several ichthyologists who have been studying the fish fauna of the Texas coast. Included among these are Gordon Gunter, Henry Hildebrand, Hinton D. Haese, Clark Hubbs, and William N. McFarland. Loren P. Woods and C. Richard Robins have pointed out to me (personal communications, 1962) that the recent exploratory fishing operations of the Fish and Wildlife Service off Texas and in the northern Gulf west of the Mississippi Delta have been so extensive at appropriate depths as to leave little doubt regarding the absence of C. aya in that area. A consideration of various environmental factors has yielded no plausible explanation for the apparent absence of Chaetodon aya from the western shores of the Gulf of Mexico, between the Campeche Bank and the Mississippi Delta. In the southern part of this area the continental shelf 1963] Hubbs: Chaetodon Aya and Related Butterflyfishes 157 is narrow, but in the northern part it is wide (Figure 2). Since C. aya almost certainly passes through a pelagic prejuvenile stage, the recognized pattern of surface currents (Leipper, 1954: 119-125, figs. 34-35) would seem to be conducive to the dispersal of the species from the Campeche Bank to the coast of Texas and northeastern Mexico, where it appears to be absent, as well as to the broad shelf off western Florida, where it does occur. The winter and early spring temperatures in the northwestern part of the Gulf from the surface down through the usual depths inhabited by the species (Phleger, 1951 : 7-25, figs. 2-19) are cold enough presumably to exclude a tropical fish, but equally cold surface temperatures during the winter characterize the northeastern Gulf (Fuglister, 1947; data repeated by Hedgpeth, 1953: fig. 15, and by Leipper, 1954: figs. 39-40) and such temperatures prevail there (Adams and Sorgnit, 1951) to the bottom at the moderate depths inhabited by C. aya. A number of other species of fishes (and of invertebrates) seem to be limited in the Gulf of Mexico waters either to the east or to the west of the Mississippi Delta, but except for vague statements about ecological differences no plausible explanations for such a limitation seem to have been made. No clue was found in the publications just cited nor in the paper on the oceanography of the Gulf by Parr (1935). The system of currents and the distribution of temperatures does nicely explain the occurrence of C. aya along the course of the Gulf Stream north to Cape Hatteras. Whatever the oceanographic, historical, ecological or other basis may be, the common occurrence of C. aya about the peninsulas of Yucatan and Florida, with more or less complete absence in adjacent areas, seems to fit a pattern (Table 5), which was mentioned by me (Hubbs, 1936: 159-16'0). A number of hypotheses may be erected to explain the more or less specifically common occurrence of the same or related fishes on the two peninsulas. Anyone hypothesis is not necessarily exclusive of the others, for two or more factors may be involved in any joint occurrence, and different factors may be involved in different joint occurrences, as indeed one would expect, considering the ecology of the species noted in Table 5. One hypothesis would invoke regular or occasional transport by current; the pelagic stage of C. aya may be dispersed at present, or may have been dispersed in the past, from a Yucatan origin to Florida by the Yucatan and Florida currents, or, possibly, in view of the indicated bathypelagic habitat of the tholichthys of Prognathodes aculeatus (p. 173), from a Florida base to Yucatan by way of a counter-flowing undercurrent (if such exists here, as it seemingly does in the Gulf Stream). Or, the joint occurrences may have a basis in past land connections, along such lines as were discussed earlier by me (Hubbs, 1936: 160). Another hypothesis, perhaps more plausible, is that ecolo~ical factors, as yet not determined, may be preserving the populations, as relicts, on the two peninsulas. 158 Bulletin of Marine Science of the Gulf and Caribbean [13(1) Variation, size, and skeletal features.-Meristic variation in Chaetodon aya and related species is discussed on page 143, for each fin, for the lateral-line pores, and for the vertebrae (Tables 1 and 2), along with an analysis of bilateral asymmetry in the pectoral-ray numbers (Table 3) and of correlations between the number of rays in different fins (Table 4). No significant geographical variation is indicated when the data for C. aya are analyzed for the 4 main regions of Occurrence (Tables 1 and 2). The size frequencies for this medium-sized chaetodontid (Figure 5) suggest a life span of several years (see also p. 146). Comparative data on skeletal features are shown for C. aya and 2 related species on Figures 7, 9, and 14, and are discussed in the account of C. falcifer (p. 154). Coloration.-Through a study of color descriptions and color photographs made in the field, and of freshly caught specimens preserved in formalin- all kindly made available by W. W. Anderson and George C. Miller, of the Bureau of Commercial Fisheries Biological Laboratory, Brunswick, Georgia, I am able to present the following information on the life colors as well as the color pattern of Chaetodon aya. The conspicuous color markings of this species are the black and blackish bars (Figure 8). The first bar, which extends unpaired along the midline from the tip of the snout to midway between the orbits, is brown- black, with a f1ded appearance even in fresh specimens; it further fades, considerably, on preservation. This bar may be of uniform width throughout, or may widen somewhat near the maxillary. The second bar extends on each side from near the rear end of the mouth to the first 2 dorsal spines. It is inky black above (and behind) the eye. but is often somewhat faded, as well as narrower, below (and before) the eye. The two segments of this bar are connected through the middle of the silvery iris by a yellow-brown band. The pupil is dark green in life. The second jet-black bar extends from the unconcealed parts of the fifth to eighth dorsal spines in a broad band, slightly arched backward, to above the posterior half of the anal fin, onto which it is extended downward, or downward and somewhat forward, as a narrowing and rather pointed process, which fades to dusky near its tip. Both black body bars are bordered on either side by a narrow white stripe, most conspicuously along the lower front border of the posterior bar. These white margins tend to fade soon if the fish has been preserved in formalin. Between the black bars the body varies from silvery to dusky gray, depending on the amount of melanin; on preservation the black pigment generally fades to a dark gray. In life there is a trace of gilt toward the dorsal fin ami of bluish on the breast and cheeks: early in preservation the body above the axis becomes more distinctly yellowish, ani the lower parts 1963] Hubbs: Chaetodon Aya and Related Butterflyfishes 159

FIGURE 8. Six specimens of Chaetodon aya, 82 to 121 mm in standard length, from the South Atlantic coast of the United States at 31°53'N. Lat., 79°3O'W. Long., at a depth of 66-7 [ meters. Note the unique black opercular spot on the left side of one specimen. The dusky appearance of the outer part of the vertical fins is due to the dark background showing through the translucent part of the fins. Photography by C. E. Dawson, reproduced with his permission. See Table 6, collection 23. show purplish reflections. Between the black bars a cream or yellowish brown streak may show, above an irregularly edged dusky longitudinal band. These marks occasionally remain on preservation. Another yellowish brown streak extends along the base of the second dorsal fin, and on the 160 Bulletin of Marine Science of the Gulf and Caribbean [13(1)

FIGURE 9. Radiograph of a specimen of Chaetodon aya 103 mm in standard length, from the South Atlantic coast of the United States at 32°55'N. Lat.. 78°31'W. Long., at a depth of 38 meters. See Table 6, collection 36. Photog- raphy by Keith W. Radford. scaly sheath. Still another yellowish brown band runs along the basal sheath of the anal fin. The dorsal spines are largely silvery gray in life, but the first 2 may be bright yellow; the next two are largely white. The posterior dorsal spines and the soft-dorsal fin may also show considerable yellow. The caudal fin is largely translucent, with a distinctly yellowish tinge basally. The exposed parts of the anal may be deep orange-yellow; the spines are largely silvery. The pectoral fin is mostly translucent, but may be dusky on the base and may show some yellow near the base. The pelvic fin in life varies from silvery to dusky gray; the front edge is silvery, but the rest of the fin may be yellow distally, and in some fish also basally, on each soft-ray. The lips may be yellow, and there may be some yellow on the branchiostegal region, in front of the anterior black bar, and some about the opercle. 1963] Hubbs: Chaetodon Aya and Related Butterflyfishes 161 The yellow color on the face, as on the fins, fades on preservation. One specimen (Figure 8) is unique in displaying a black blotch on the opercIe, on the left side. Chaetodon guyanensis Durand Figures 1, 2, 4, 5, 10, 11 Chaetodon guyanensis Durand, August 5, 1960 (40)4: 209-213, fig. 2 (original description; comparisons; relationships; subgenus; associated species; Guyane [French Guiana], at 06°35'N. Lat., 52°34'W. Long.). Chaetodon goniodes Woods, "1960" [January 19, 1961] (41): 417-420, fig. 1 (original description; relationships; deep habitat, at 125 fathoms, with deep-sea associated species (listed); off west coast, Puerto Rico, at 18°26'N. Lat., 67°11'W. Long. [=18°24'N. Lat., 67°15'W. Long., in 120 fathoms = 183 meters]). Synonymy.-It is a surprising circumstance that this species, like Chaeto- don marcellae and Prognathodes aculeatus, was described twice under different names within the span of one year. As in the other species, the priority seems clear: Jean Durand has informed me that the description of C. guyanensis appeared on August 5, 1960, whereas C. Richard Robins has told me that the account of C. goniodes was published on January 19, 1961 (not in 1960, as dated on the printed copies). There seems to be no reason to doubt the identity of the two nominal species. The diagnostic features show well on the type figure of each nominal species (Figures 10 and 11). Characters and comparisons.-The known differences of consequence between this species and its nearest relatives (c. aya and C. marcellae) involve details of the color pattern (compare Figures 8, 10-12). In C. guyanensis the anterior band below the eye is more nearly horizontal than vertical, rather than the reverse, and extends to the maxillary in advance of, instead of to, the rear end of that bone (when extrapolated, the axis of the bar below the eye strikes near the junction between the two parts of the dorsal fin, rather than in advance of the middle of the spinous part); the two body bars approach each other along the edge of the dorsal-fin sheath to within less than the diameter of the eye, rather than nearly twice that dimension; instead of being diverted downward or even slightly forward in a narrowing extension on the anal fin, the posterior bar extends backward and downward across the fin in a broad band, so as to leave only a narrow clear border on the posterior (actually upper) part of the fin, rather than a broad and complete light margin; a black band, at most very faintly suggested in C. aya, extends along the entire length of the anal fin; a dark bar, unrepresented in C. aya, extends across the caudal

4 Figures in parenthesis in the synonymies, entered after the date in each item, refer to the chronological list of collections (Table 6). and indicate the collection stated, or assumed, to have been used in the given reference. 162 Bulletin of Marine Science of the Gulf and Caribbean 113(1 )

FIGURE10. One of the two syntypes of Chaetodon gll)'anensis, 86 or 90 rnm in standard length, from off French Guiana (Guyane). Photograph received from and reproduced by permission of Jean Durand, Laboratorie d'Ichthyologie du Museum National d'Histoire Naturelle, Paris. peduncle; the inner part of the pelvic fin is more deeply pigmented. The light margins along the black bars appear to be broader and bolder than in C. aya. Three other differences pointed out by Woods or indicated by his data do not seem to hold. The supposedly shorter and stronger dorsal spines are about matched in some specimens of C. aya and do not seem to hold for the figured syntypes of C. guyanensis (see Figures 8-11). Woods correctly gave the principal caudal-ray formula for the type of C. goniodes as i, 8-8, i (= 18), which is not represented in any of the 83 counts for C. aya, but since Jean Durand has informed me that the two syntypes of C. guyanensis have the normal number of i, 8-7, i (= 17), characteristic of Chaetodon and of most typical spiny-rayed fishes, I conclude that the count of 18 principal caudal rays in the type of C. goniodes is an unexp~ct- ed coincidence. The pectoral rays in the type of C. goniodes number 15, which is unusual for C. aya, but Durand gave counts of 13 to 15 for the syntypes of C. guyanensis. Possibly C. guyanensis has, on the average, slightly more pectoral rays 1963] Hubbs: Chaetodon Aya and Related Butterflyfishes 163 and slightly more pored scales than C. aya, and in these respects it may correspond more closely with C. marcellae. However, the data for C. guyanensis and C. marcellae (Tables 1 and 2) are inadequate to secure this point. Durand indicated higher counts for dorsal and anal soft-rays than is usual for C. aya, but it is thought likely that he enumerated separately what I would treat as the two elements together comprising the last ray. For this reason his counts for these fins are not included in computing the averages for C. guyanensis in Table 1. Relationship, distribution, and status.- Very probably, C. guyanensis and C. aya are allopatric geminates. No specimen confusable with C. guyanensis has appeared in the large sample of C. aya now available, and the ranges appear to be distinct. It may be assumed that C. guyanensis will be found at various localities around and between the three known locations, off Puerto Rico, off Barbados, and off northern South America. It is to be looked for also along the coasts of other Antillean islands. Perhaps C. guyanensis and C. aya occur together, or intergrade, along the northwestern coast of Cuba, but until or unless intergradation is demonstrated, full specific status will be indicated. Subjective considerations of closeness of relationship or of hypothetical chances of interbreeding if the two forms could be kept together would not, in my opinion (Hubbs,

FIGURE II. Holotype of Chaetodon goniodes, synonym of C. guyanensis, 95 mm in standard length, from off Puerto Rico. Photograph received from and reproduced by permission of Loren P. Woods, Chicago Natural History Museum. 164 Bulletin of Marine Science of the Gulf and Caribbean [13(1) 1943), justify according these forms subspecific status. I hold to this view despite the contrary statements and actions of certain ornithologists and other vertebrate zoologists. Chaetodon guyanensis is obviously a relatively deep-water species, and probably lives deeper than Prognathodes aculeatus and, at least on the average, deeper than C. aya. The Puerto Rico specimen was trawled at a depth of 183 meters, where, as Woods indicated, it was associated with typically bathybial fishes. The specimens from off French Guiana were taken at a depth of 105 meters. The data for the type specimen of Chaetodon goniodes as given in the original description are somewhat in error, as they were taken from an incorrect Station List. The corrected list gives the location as 180 24' N. Lat., 67° 15' W. Long., rather than as 18° 26' N. Lat., 670 11' W. Long., and the depth as 100 instead of 125 fathoms. A third record almost surely pertaining to C. guyanensis has been furnished by John B. Lewis (personal communication, 1962), of the Bellairs Research Institute of McGill University, at St. James, Barbados, West Indies. Some specimens that had been caught recently in a deep-set fish-pot off the west coast of Barbados agreed well with the original figure of C. goniodes. Unfortunately the specimens have not been found. The record must remain clouded with some degree of doubt. Chaetodon marcellae Poll Figures 1, 3-5, 12 Chaetodon (Oxychaetodon) marcellae Poll, November, 1950 (44)': 2-7, fig. 1 (original description; 25 miles WSW. of Pointe de Banda, west Africa, at 03°57.5'S. Lat., 10°36.5'£. Long.). Chaetodon marcellae: Cadenat, 1953 (44-47): 1057 (C. altipinnis a synonym, west Africa; 11 kilometers SW. of Goree, in 38-40 meters, and at 14°30'N. Lat., in 50 meters).-Cadenat, 1954 (44-47): 574 (feeds on polychetes).-Roux, 1957 (44-48): 201,278 (synonymy; description; comparisons; "vers 70 metres").-Hubbs and Rechnitzer, 1958 (44,45): 275, 278-281, 284, 304, 305, 307, 308, map 1, pI. 2, fig. 2 (c. altipinnis as junior synonym; comparisons and relationships; west African records cited).-Myers, 1959 (44, 45): 158 (relationships).-Cadenat, 1960 (44-48): 1403 (west Africa).-Durand, 1960 (44-48): 210-212, fig. 3 (characters; comparisons; relationships; C. altipinnis a synonym; tropical Africa).-Bauchot and Blanc, 1961 (42, 43, 48): 87-94, fig. 3, pI. 1, fig. A (off Liberia, at 04°57'N., 09°26'W., in 95 meters; off Ivory Coast, at 04°38'N., 06°18'W., in 90 meters; "region de Pointe-Noire"; review of records; C. altipinnis as synonym; description; compared with C. aya, C. fabfer, and other eastern Atlantic species; referred to subgenus Chaetodon).-Cadenat, 1961 (44-48): 242 (C. altipinnis a synonym).- Woods, 1961 (44,45): 420 (comparisons; relationships). --- 5Figures in parenthesis in the synonymies, entered after the date in each item, refer tu the chronological list of collections (Table 6), and indicate the collection, stated or assumed, to have been used in the given reference. 1963] Hubbs: Chaetodon Aya and Related Butterflyfishes 165 Chaetodon altipinnis Cadenat, 1951 r not 1950; see Hubbs and Rechnitzer, 1958: 304] '(45): 239,307,315,318, fig. 174 (original diagnosis: Senegal). Relationships and distribution.-The synonymy of Chaetodon marcellae, the west African cognate of C. aya and C. guyanensis, has been authenti- cated. The published figures show the differences in coloration, which, though highly consistent and therefore distinctive, are mere modifications of the same basic pattern. Other differences are not apparent. There may be slight, but at most only average, differences in ray and pore counts (Tables 1 and 2). The indicated average differences between C. marcellae and C. aya in numbers of dorsal and anal soft-rays may merely reflect, as for C. guyanensis, a difference in mode of counting. As indicated earlier (p. 140), C. marcellae was probably derived from

FIGURE 12. Holotype of Chaetodon marcellae, reproduced, by permission, from Poll (1950: fig. 1). 166 Bulletin of Marine Science of the Gulf and Caribbean [13(1) a vagrant from the American side of the Atlantic. It is now known (Bauchot and Blanc, 1961; Figure 3) to be widespread along the coast of tropical west Africa. Like its relatives, it inhabits rather deep water, apparently at about the same depth as C. aya (Figure 4). Prognathodes aculeatus (Poey) Figures 1, 2, 4, 5, 13, 14 Chelmon aculeatus Poey, July, 1860 (50) 6: 202-203 (original description; rare; [vic. Habana,] Cuba).-Metzelaar, 1922 (52): 137 (characters; Caracasbay, Cura9ao). Prognathodes aculeatus: Poey, 1868 (50): 354 (reference; characters).- Eigenmann and Horning, 1887 (49, 50): 2-3 (diagnosis: relationships; Chelmo pelta a synonym; after GUnther and Poey).-Jordan, 1887 (49, 50): 593 (references; West Indies; Chelmo pelta a synonym) .-Jordan and Evermann, 1898 (49, 50): 167] (description; synonymy; after GUnther and Poey) .-Ahl, 1923 (49, 50): 11 (synonymy; relationships: description; Havana, Cuba; West Indies) .-Erdman, 1956 (58): 339 (possible identification-Capros aper also suggested--of young specimen [tholichthys prejuvenile] from stomach of blue marlin caught at La Par- guera, Puerto Rico, March 26, 1956).-Hubbs and Rechnitzer, 1958 (49, 50): 281, 304-308, map 1, pI. 3, fig. 2 (synonymy; Chelmo pelta as junior synonym; description; relationships) .-Bailey, et al., 1960 (65): 32 (listed for United States; vernacular, "longsnout butterflyfish") .-Erdman, 196] (58?, 66, 68, 81): 39 (addition to Puerto Rico list). Randall, 1961 (50, etc.): 54 (probably a long-snouted derivative from a species like C. aya; origin apparently independent of Forcipiger) .-Caldwell, 1962 (54,55,82): 276 (Jamaica). Prognathodus [emendation] aculeatus: Bleeker, 1876a: 303; 1876b: 3] 5; and 1871 (49,50): 32,34 (characters; genus; synonymy). Chelmo pelta GUnther, September, 1860 (49): 38 (original description; location?) . Chelmo peltae [emendation]: Gill, 1862 (49): 238 (cited under "Progna- thodes, Gill"). Chaetadan sp. CoIlins, 1956 (56, etc.): 15th p., col. fig. in pI. on 11th p. [fish part prepared by J. S. Zaneveld (fide Zaneveld, 1956: 5, and personal communication, 1958)]. Material.-Many new collections and much new information from collectors and observers show that this twice-named species is widely distributed from the islands off the north coast of South America through the Lesser and Greater Antilles to the Bahamas and around the southern tip of Florida (Figure 2). Despite this wide distribution and wide bathymetric range (Fi~ure 4), this species appears to exhibit no pronounced regional variations. Knowledge regarding this species has advanced rapidly during the past few years. Only two specimens, the respective types of Chelmon aculeatus Poey and Chelmo pelta Gunther, were known to Hubbs and Rechnitzer

6Figures in parenthesis in the synonymies, entered after the date in each item, refer to th~ chronological list of collections (Table 6), and indicate the collection stated, or assumed, to have been used in the given reference. 1963] Hubbs: Chaetodon Aya and Related Butterflyfishes 167 (1958: 305), who, however, overlooked Metzelaar's (1922: 137) record of Chelmon aculeatus and Collins's (1956) record of "Chaetodon sp.," both from Cura~ao. Numerous additional specimens came to light in 1958, during my trip to European museums, and throughout the five-year period, 1958-1962, during which specimens have been examined in American institutions. Both in Europe and America numerous colleagues (see acknowledgments) have made old and new material freely available to me. A short time previously, G. F. Mees, then of Rijksmuseum van Naturlijke Historie, Leiden, had studied the European material (see p. 136). The third-collected specimen (see Table 6, collection 51), not previously published on, is listed in the Records of 1866 in the British Museum (Natural History) as having been obtained at "Dominicia" by Me. Cutter; this fish was doubtless taken at Dominica in the Lesser Antilles, for adjacent entries are for fishes taken by the same collector in Barbados and Trinidad (furthermore, we now report, as collection 83 in Table 6, a recent observation of the species at this, its southeastern most area of known occurrence). The 10 specimens reported by Metzelaar, which were collected by C. J. van der Horst in 1920, were studied in the Zoological Museum at Amsterdam. Topotypes of Chelmon aculeatus, collected at Habana, Cuba, by Luis Howell Rivero in 1934, were found in the Museum of Comparative Zoology at Harvard University and in the Academy of Natural Sciences at Philadelphia. Specimens collected by Victoria Smith in Jamaica in 1950 and 1951 were located by David K. Caldwell in the Institute of Jamaica, at Kingston. From 1955 to 1962 specimens have been taken each year through the range of the species. So far as known to me, all specimens and scientific observations are tallied in Table 6, and, except as notcd in this table, were utilized in preparing Figures 1, 2, 4, and 5.

Relationships and status.-As previously indicated (Hubbs and Rechnitzer, 1958: 305; Randall, 1961: 54), Prognathodes aculeatus appears to be a close relative of Chaetodon aya and similar species (C. marcellae and the more recently discovered C. guyanensis). A comparative study of radiographs shows essential agreement with C. aya in most respects, but discloses a rather striking distinction from C. aya and other species of Chaetodon examined (see p. 154-155): the two anteriormost, free interneurals, which arc attached on either side of the first neural spine, converge above thcir base and fuse, rather than diverging throughout and remaining separate; furthermore the supraoccipital crest of the cranium extends to below the front of the dorsal fin or even farther, rather than only to the first interneural. These features may help to define the genus Prognathodes as a supplement to the previously used specialized structure of the forceps-like jaw (see Hubbs and Rechnitzer, 1958: 305-306). I see no particular advantage in synonymizing Prognathodes with Chaetodon, even though it be recognized as a specialized offshoot of the aya group of Chaetodon, with 168 Bulletin of Marine Science of the Gulf and Caribbean [13(1)

______.R"R ••• _.H"

FIGURE 13. Two specimens of Prognathodes aculeatus from Jamaica, respectively 62 and 67 mm in standard length. See Table 6, collections 54 and 55. Photograph received from and reproduced by permission of David K. Caldwell. 1963] Hubbs: Chaetodon Aya and Related Butterfiyfishes 16Y no direct relationship with other chaetodontid genera, such as Chelmon and Forcipiger, having analogously modified beaks. The independently derived beak structure, however, provides a new example of the low phyletic importance of modifications related to feeding.

Characters and variation.-In meristics (Tables 1 and 2) P. aculeatus approaches or corresponds with its presumed nearest relatives (in the genus Chaetodon). The dorsal spines are normally 13, rarely 12 or 14; dorsal soft-rays, 18 or 19, rarely 16 or 17; anal spines, consistently 3; anal soft-rays, usually 15, occasionally 14 or 16; pectoral rays, usually 14, occasionally 13, rarely 15, consistently with the uppermost short ray and the upper principal ray unbranched, and often with the lowermost one or two rays unbranched; the principal caudal rays 9 + 8 = 17, with a deviation to 8 + 8 = 16 in 3 of 52 specimens; pelvic rays consistently I, 5; all essentially as in C. aya. The pored scales of the lateral line (Table 2) average fewer than in C. aya: 22 to 28, usually 24 to 26, instead of 25 to 34, usually 27 to 30. The vertebrae are consistently 10 + 14 = 24 (in the] 5 specimens counted). Features of bilateral asymmetry in number

FIGURE ] 4. Radiograph of a specimen of Prognathodes aculeatus from Puerto Rico, 65 mm in standard length. See Table 6, collection 66. Photography by Keith W. Radford. 170 Bulletin of Marine Science of the Gulf and Caribbean [13(1) of pectoral rays and of correlation between members of rays in different fins are indicated on page 144 and in Tables 3 and 4. The general features of body, head, and fin form, indicated in Figures 13 and 14, are remarkably close to those of Chaetodon aya--except for the much more exaggerated development of the beak. Various other characteristics of P. aculeatus are very similar to those of C. aya, as Hubbs and Rechnitzer (1958: 305) noted. In general, the different lots of specimens, including the types of Chelmon aculeatus and Chelmo pelto, agree well in the diagnostic features of the species. The lateral line ends close to the extreme structural base of the soft-dorsal rays; usually about one scale row short. The bones around the orbit are finely denticulate. The muzzle is well scaled forward to the rostral fold, but there are no scales on the top of the premaxillary; the chin is well scaled almost to the front. A more or less scaleless groove extends from the nostril to the rostral fold. In the type of Chelmo pelta, in the specimen from "Dominicia," and in some others, a thin layer of flesh seems to have covered the upper surface of the premaxillaries; the upper lip is very little developed, along the very edge of the beak; the cross groove is weak or absent; laterally, the lip is narrow, and runs out posteriorly. There is considerable variation in the length of the beak. It is particularly short in the specimen from "Dominicia," in which its length steps into the head 2.35 times, contrasted with 2.15 times in the type of Chelmo pelta; the length of the production of the upper jaw beyond the rostral fold is only about 1.3 rather than 1.7 times the basal width and steps 3.3 instead of 2.8 times into the length of the snout. Despite the magnitude of these differences, no specific differentiation seems involved. There is considerable variation in the denticulation of the head bones, which I examined chiefly in the series of 10 specimens that Metzelaar (1922: 137) reported from Cura~ao. The scapular edges are finely denticulate. Occasionally the interopercular edge is weakly denticulate. A rounded notch is more or less developed above the angle of the preopercle, and a minor notch is sometimes developed farther up. On the upper part of the upper limb the edge may be wholly smooth, or vary to rather finely denticulate, with the points directed backward or backward and upward. On the rounded angle the denticulations are commonly strengthened, but may be no stronger than elsewhere or even very weak; the points may be directed backward or slightly upward or slightly downward, and commonly tend to radiate. The denticulations on the lower limb of the preopercle also vary greatly in strength and regularity, and may be entirely lacking: they also vary considerably in their deviation, either backward or forward, from a vertical direction. Obviously no great reliance can be placed on these characters. The caudal peduncle is typically slender in P. aculeatus, but varies 1963] Hubbs: Chaetodon Aya and Related Butterflyfishes 171 considerably. The least depth steps into head 4.2 times in the type of C. pelta, 4.0 times in the "Dominicia" specimen, and 3.7 to 4.3 times (average 4.0) in the 1'0 specimens reported from Cura9ao by Metzelaar.

Coloration.- The most striking external differences between P. aculeatus and its nearest relatives (in the genus Chaetodon) lie in the coloration. The anterodorsal midline bar of C. aya and cognate species is moderately developed, but the bar that extends from near the origin of the dorsal fin through the eye to the snout is dusky orange instead of jet-black and the posterior jet-black bar of the other species is lacking. There are striking differences also in the ground color in life. The life colors of P. aculeatus are purportedly shown for "Chaetodon sp." in a pamphlet listed as written by Collins (1956), though the fish section, I understand, was prepared by Jacques S. Zaneveld, who has informed me that the upper parts of the body and the caudal fin are shown too brown and the bars on the opercle and on the caudal peduncle too red. These indicated inaccuracies are not due to the reproduction of the plate for they appear also on the original painting, which I saw in Amsterdam. With the aid of color descriptions and color transparencies kindly furnished me by David K. Caldwell, John E. Randall, and Loren P. Woods, supplemented by the examination of specimens preserved for short to long periods in formalin and in alcohol, I am able to compose the following description of the color and color pattern of this interesting species. A dark-edged dusky band, which may be more or less disrupted, at least in preserved specimens, extends from the front of the beak along the snout middorsally to the upper posterior part of the interorbital region. The most conspicuous dark marking is the dusky orange to bright orange band, margined narrowly with brown, that extends from the eye to the front of the dorsal fin, sometimes forming a commissure around the origin of the fin. The anterior part of the band extends from the rostral fold near the midline horizontally backward, with a slight upward curve posteriorly, to the front of the eye. In preserved specimens the two sections of this band are more or less prominently light-centered. Between the converging upper ends of the pair of dark bands and behind the middorsal stripe a triangular light area is more or less boldly developed. These dark bands fade markedly with age and with preservation. The opercular margin from the upper end of the gill opening to opposite the upper edge of the pectoral base is bright orange. The upper limb of the preopercle is narrowly yellow. The side of the snout and in some fish the lower part of the cheek are yellowish to light yellow. A usually narrow but occasionally broad, rarely doubled, bar of bright orange, dull red, or red- dish brown extends vertically across the caudal peduncle. This colorful bar is usually set off, more or less prominently, by a following cream or yellow streak. 172 Bulletin of Marine Science of the Gulf and Caribbean [13(1) The ground color of the body and head grades from dark brown dorsally through dusky orange, more or less obscured by dusky olive, and through yellowish to white with blue and violet reflections ventrally, and often becomes somewhat orange on the back in front of the fourth dorsal spine and on the caudal peduncle. In some individuals the upper anterior parts are washed with pale purplish, so that the orange-brown color is largely on the upper posterior parts. Lengthwise bluish lines are just visible on the lower half of the body. In preserved specimens the scales are more or less definitely marked off by individual dusky blotches. The spinous part of the dorsal fin and the basal part of the soft-rayed portion are very dusky orange-brown to blackish with an orange cast. The outer part of the soft dorsal is hyaline beyond a dusky orange to rather clear orange stripe that runs from near the tip of the first soft-ray in an almost straight and nearly vertical line to near the middle of the last rays. This stripe is narrowly bordered on its outer (posterior) edge by brown or blackish, and tends to be separated by a light band from the dark basal portion, which may be intensified toward its outer margin. In some preserved specimens a submarginal dusky streak is evident on the soft dorsal. The interradial membranes may be almost black between the spincs, especially behind the fourth one, and on the dark part of the soft fin. The first 4 dorsal spines and the brief membranes betwecn them are more or less conspicuously lighter. The sooty color on the dorsal fin is diffused to a varying distance and in varying intensity onto the adjacent part of the body, but there is no tendency toward the formation of an oblique bar. The markings on the dorsal fin, like the bars on the body, fade with age. The other fins are less distinctively colored. In life, the caudal fin is usually hyaline, with dusky rays; after preservation, it varies from pale gray to almost blackish; a broad darker bar commonly crosses the base of the fin. The anal and pelvic fins are typically clear, bright yellow in life. In preserved specimens the anal is either clear or more or less darkened, either basally or distally, with the pigment more intensified on the membranes than on the rays; on the posterior (upper) half, there is variously developed a narrow clear edge, just within which the dark pigment is sometimes con- centrated. In preserved material, the pelvic fin grades from sooty blackish in the smaller specimens to whitish in the largest ones. The pectoral in life is hyaline, with the basal part slightly yellowish; on preservation, the fin tends to be somewhat darkened toward the base. The color description has been compounded from notes made on specimens from the Bahamas, Florida, Jamaica, and Curac;ao. No regional differences are indicated.

Size and life history.-As compared with Chaetodon aya, C. guyanensis, and C. marcellae, and more notably as contrasted with C. faicifer, Prog- nathodes acuieatus is a small species (Figure 5). The sharply dominant 1963] Hubbs: Chaetodon Aya and Related Butterflyfishes 173 5-mm size-class is 60 mm (standard length). The two specimens in the largest (70-mm) size-class measure 7 I mm. The apparently unimodal size- frequency of the adults suggests a definitive size at maturity and very likely an annual life cycle. Immature specimens, in the 35- to 50-mm size-classes, were taken in June (2 collections), July (3), and September (3). Accord- ing to Jacques S. Zane veld (personal communication, 1958), who had extensive experience with the species in 1955-1958 while he was director of the Caraibisch Marien-Biologisch Instituut at Cura<;ao, specimens "are mostly collected during the months of November-March; however, they are also present outside the rainy season." What proportion are of adults at the different seasons was not mentioned. Ingvar Kristensen, present director of the marine institute at Cura<;ao, has added (personal communication, 1962) that: "We do not know, any- thing about spawning but small specimens [the size] of a dime or so we see in February-March. They seem to grow quite rapidly." The single specimen in tholichthys prejuvenile (Hubbs, 1958) stage, 23 mm in standard length (see Table 6, collection 58), was collected on March 26 (a date compatible with that for the appearance of young on the reefs at Cura<;ao). It was found in the stomach of a bathypelagic fish, Pseudoscopelus sp., itself from the stomach of a blue marlin, Makaira nigricans Lacepede, caught at La Parguera, Puerto Rico. The specimen was referred to by Erdman (1956: 339) as "the young of Prognathodes aculeatus if it is not the young of Capros aper." On reexamination, this specimen (USNM 114652) proves to be the tholichthys of a chaetodontid. Its radial formula (D XIII, 19; A III, 15; C 9 + 8 = 17; Pt 14-14, P2 1,5-1,5) and number of lateral-line pores (26-26) and the position of the lateral line, as well as the peculiar pre juvenile-stage characters, fixes it as a species of Chaetodontidae, definitely excluding Capros. The anterior parts are heavily armored in the tholichthys style. There is a pair of large plates on each side of the body, respectively above and below the lateral line, and the usual plates on the head, covering the postorbital, suborbital, and preorbital regions forward to the narial fossa. The muzzle is sharp, already produced about as much as in C. aya, and about one-third as long as the head. The caudal peduncle is slender and the coloration is plain. These meristic, morphological, morphometric, and colorational characters all suggest P. aculeatus. The pore count would appear to exclude C. guyanensis (see Table 2), which is also known from Puerto Rico, as well as C. aya, which appears to be excluded on distributional evidence. The later capture (collection 68, Table 6) of a full-sized adult of P. aculeatus at La Parguera, Puerto Rico confirms the identification.

Habitat.-On the basis of the information in the preceding paragraph, it appears that P. aculeatus is bathypelagic in the tholichthys stage. The half- grown to adult specimens have been taken at depths from less than 1 to 91 174 Bulletin of Marine Science of the Gulf and Caribbean 1/3(1) meters, with the center of abundance at 15 to 55 meters (Figure 4). All collections and observations came from above the three depth records (105 and 183 meters) for the sympatric species C. guyanensis. The depths usually inhabited are apparently similar to those known for C. falcifer and average less than those for the other Atlantic species, C. aya and C. marcellae. Concerning the depths and ecological conditions usually inhabited we have the testimony of collectors who have obtained and observed this species in various parts of its range. At Cura\fao (see collections 52,56,57, and 59-61, Table 6), accoding to Jacques S. Zaneveld (personal communication, 1958), who was director of the Caraibisch Marien-Biologisch Instituut there in 1955-1958, the species is usually taken, in fish-pots, at the rather considerable depth of about 20 to 40 meters (somewhat deeper than the depth inhabited by most other colorful reef fishes). The first specimens came from the fish-pots set at 52 meters. "Diving with snorkel and glasses I never saw the species between the coral reefs close to shore, where the Chaetodon species are always to be observed ... I have the impression that it occurs only in somewhat deeper water." Occasionally, however, the species comes into shallow water on the reefs about Cura\fao (see collection 85, Table 6). John E. Randall reported (personal communication, 1962): " ... I observed Prognathodes in Cura- \fao in as little as 3 feet of water right at a rocky shore. I saw it several times in shallow water, so it was not a case of one displaced fish. It is, however, more common in the deeper water at Cura\fao ... I observed it down to about 200 feet. My deepest dive at the island was 210 feet." Still later in 1962, referring to collection 89, Randall reported (personal communication, 1962) as follows: "Prognathodes aculeatus does occur in shallow water off Cura\fao, not always in or near caves. I saw 3 in one day in 15 feet with no more than the usual cover afforded reef fishes around small coral heads. The water was very clear. Still, there is no question about its 'preference' for deeper water." More details regarding the habitat of the species at Cura\fao were provided under date of June 6, 1962 by Ingvar Kristensen, present Director of Caraibisch Marien-Biologisch Instituut on that island. He noted that Prog- nathodes aculeatus is not very common at Cura\fao, less so than the species of Chaetodon. It is seldom found at a depth of less than 20 meters; most commonly, at about 40 meters, where patches of coral grow, at about the depth limit for corals. However, depth per se is apparently not the con- trolling factor. Gerrit Klay has caught specimens at a depth of only 2 to 3 meters near Lagoen, on the south coast of Cura\fao in an underwater cave, wherein light measurements indicate the same light intensity as at 40 meters outside the cave. The same cave is known for its beautiful specimens of 1963] Hubbs: Chaetodon Aya and Related Butterflyfishes 175 Stygnobrotula latebricola and for the deepwater habitus of some coral species. Concerning his observations about the Virgin Islands, Randall has written (personal communication, 1959): "I think there is no question about Prognathodes being a fish of deeper water (deep for reef fishes, that is). 1 have dived a great deal around St. John since last November, but never in a cora]iferous area deeper than about 60 feet. During this time I never sighted Prognathodes. Then I made one 25-minute dive off French- cap Cay in from 100 to 110 feet (collection 67, Tab]e 6) and saw five ... The bottom was about three-fourths covered with small heads of coral, very few over two feet high; the rest of the bottom was sand. Very few Diadema were seen. From the depth of ] 10 to ]20 feet off Frenchcap Cay the bottom tended to level off and consisted mostly of sand, with a very few, scattered coral heads; I sighted very few reef fishes in this sandy zone and no Prognathodes." Loren P. Woods informed me (personal communication, 1959) that 3 specimens of P. aculeatus trawled by the OREGON about the Virgin Islands (collections 77 and 78, Table 6, from 69 and 53 meters respectively) were from sponge areas. The specimen from Dade County, F]orida (collection 87, Tab]e 6) was captured alive at a depth of 24 meters along a rocky ledge (Henry A. Feddern, personal communication, 1962). In the Bahamas, according to James E. Boh]ke (personal communication, ] 961 ), this species was taken by diving at depths of 12 to 23 meters about cora] reefs, coral and rock, and cora] and sand (collections 71-75, Table 6) ; and at a depth of 55 meters, at the edge of the dropoff to deep water (col- ]ection 84). John E. Randall and associates have seen or collected other specimens from the Bahamas (collections 62, 63, 69, and 70), at depths of 15 to 55 meters, and he has indicated (persona] communication, 1959) that in the Bahamas "this species appears to be the most common butterflyfish at depths greater than about ] 30 feet." At the southeastern tip of Florida John E. Randall and Walter A. Starck, II (personal communication, ] 958) took Prognathodes aculeatus for the first time in the United States (collection 65, Tab]e 6). Following is the substance of Starck's field data, with additions from an accompanying sketch. Here the species occurs in a depth zone between 23 and 29 meters on the outer 6 meters of flat bottom inside a vertical dropoff of about 3 or 4 meters, more or less undercut above the gently sloping sand bottom. It never strays far from the dropoff. The habitat is primarily one of large head-forming corals, sponges, and a1cyonarians. Live growths of these organisms occur behind the lip, and some coral grows where this fish occurs on the upper part of the vertical cliff, where this fish ceases to occur, near the middle of the vertical cliff, the coral is dead and pock-marked. In general terms John E. Randall has written (personal communica- 176 Bulletin of Marine Science of the Gulf and Caribbean [13(1) tion, 1962): "this butterflyfish is more common in the deeper waters (100 to 200 feet) where I have seen it (Bahamas, Virgin Islands, Puerto Rico, Cura<;ao) than the common species of Chaetodon, which are in shallower water (C. capistratus is usually the commonest). It did not seem so common on the Florida reefs."

Behavior and Food.-Jacques S. Zaneveld, John E. Randall, Walter A. Starck, II, Ingvar Kristensen, and other observers have commented (personal communications, 1958, 1959, 1962) on the distinctive behavior of this species, as compared with the chaetodontids of shallower water (but there have been no observations on Chaetodon aya or on C. guyanensis). Zaneveld has interpreted its actions as more pleasing and attractive, and its present favor as a home-aquarium fish, mentioned below, probably reflects such behavior. Starck has written that on the Florida reef it swims about on the face of the underwater cliff and on top of the reef, but never very far from the dropoff; he regards it as a solitary species, which he has never observed in pairs or schools such as he has recorded for other chaetodontids that he has observed; it "flits about quite rapidly, always keeping close to the coral and not allowing itself to be approached as closely as other chaetodontids do." At Frenchcap Cay in the Virgin Islands, at a depth of 30 to 34 meters, 2 of the 5 specimens seen by Randall were swimming as a pair. The food of Prognathodes aculeatus is probably similar to that of the other chaetodontids that have developed a beaklike muzzle, along one or more other phyletic lines, presumably as an adaptation for food probing in small recesses of the intensively foraged tropical reefs. Conrad Limbaugh (personal communication, 1958) found Forcipiger feeding at Clipperton Island on the tube-feet of the venomous-spined sea urchin Diadema, though stomach contents of the Forcipiger contained a variety of small invertebrates (Randall, ]961) . Jacques S. Zaneveld has reported (personal communication, 1959) that Prognathodes aculeatus feeds on the tube-feet of echinoids at Cura<;ao: "we observe that Prognathodes and all other chaetodontids very often look for food between the spines of the sea urchins; they pluck the seaweeds growing on the spines of Eucidaris tribuloides, they often pull on the spines of Tripneustis esculentus and Lytechinus variegatus, and likewise they try to clean Diadema, but the long movable spines make it impossible to approach; so the fish tries it from the oral side, where it has the least trouble." In the aquarium of the Caraibisch Biologisch lnstituut in Cura<;ao "it lives on small animals occurring on and in corals, especially Acropora palmata and A. cervicornis." The stomach contents of two of the specimens from the Virgin Islands (in collection 67, Table 6), examined by Randall and me, include one nemertean, one mysid, one very small caprellid, at least 1963] Hubbs: Chaetodon Aya and Related Butterflyfishes 177 one copepod, and several other small crustaceans, and, apparently, a number of echinoid pedicellariae. More information on the behavior and food of Prognathodes aculeatus has just been received from Ingvar Kristensen, who on June 6, 1962 transmitted the following observations by himself and Gerrit Klay on Cura~ao. The adults are solitary, but the young often keep together. At times as many as 8 are swimming around an isolated piece of coral. Klay has seen this species feeding on small crustaceans that live between the spines of sea urchins. In the aquarium (at Piscadera Baai) it feeds on Mysis and often browses. Unlike some other butterflyfishes, it never picks parasites from the bodies of other fishes. Toward one another, but not toward other fishes, specimens of P. aculeatus can be rather intolerant. When attacked they do not flee, but rather elevate their strong, high dorsal spines and orient themselves in a more or less horizontal position with the spines directed toward the attacker. This seems to be effective behavior, as most fishes quickly cease the attack. Kristensen provided another interesting bit of information which suggests that thigmotaxis may play an important role in the activities of this species: the fish usually swim with belly upward along the roof of the underwater cave mentioned above. Vernacular names.-Collins (1956: 15th p., "Check-list of the Fishes of the Netherlands Antilles") cites "forceps-fish" as the English name of "Chaetodon sp." (=Prognathodes aculeatus), and this, I was informed at the Amsterdam Museum, is merely the English version of the Dutch name, "pincet vis." Collins assigns a native (Papiamento) name, "chamba," to the species. The American Fisheries Society approved list of names (Bailey, et al., 1960: 32) assigns "Iongsnout butterflyfish" to the species. In the newly developed aquarium trade, it is going under the name of "Ross butterflyfish." It is odd that a species so poorly known a few years ago should already have developed a vernacular synonymy. Prognathodes as an aquarium fish.-For some years Prognathodes aculeatus has been displayed in the aquarium of the Caraibisch Marien-Biologisch Instituut at Cura~ao, where, as mentioned above, it feeds on small animals living on and in corals. More recently, John E. Randall (personal communication, 1962), wrote that this very attractive species has entered the aquarium trade, as the "Ross butterflyfish." Still later, Director Ingvar Kristensen of the Caraibisch Marien-Biologisch Instituut informed me that this is a very hardy fish much easier to keep than the species of Chaetodon. They do not suffer from Oodinium or other diseases and are among "the most pleasant aquarium fishes you can imagine." Gerrit Klay, a diver at Cura~ao, catches specimens, which he sells at a reported price of $5.00 each to New York aquarium dealers. So, what not long ago was a scientific curiosity of great rarity has now become an article of commerce! 178 Bulletin of Marine Science of the Gulf and Caribbean 113(1)

TABLE 1

FIN-RAY VARIATION IN DEEP-LIVING SPECIES OF Chaelodon AND Prognathodes Counts are original for all specimens except the types of C. eques. C. guyanensis, C. marcellae, and (from figure) C. allipinnis, and except for ten specimens of C. marcellae reported by Bauchot and Blanc (1961); those counts are in parentheses; those for dorsal and anal soft-rays are probably I too high (see p. 163) and were disregarded in computing means. Regions under C. aya are: I, South Atlantic coast, U. S. A.; II, Straits of Florida; Ill, NE. Gulf of Mexico; IV, Campeche Bank. The last 2 dorsal and anal soft-rays that are separate through the concealed base are counted as 1.

IC. fal- Chaelodon aya (by Regions) ,c. gUy-I c. mar- P. awl- I eifer II 1I1 IV Total ,_al1ensis c~~e eallis Dorsal spines 12 I I I 2 I 13 3 45 7 17 10 (+1) 79 (+1) 1 (+2) ( 12) 51 14 3 I 4 I •. 1 No. 3 49 8 18 10 (+1) 85 (+1) I 1(+2) ( 12) 53 Mean 13.0 13.0 12.9 13.1 13.0 13.0 13.0 ( 13.0) 13.0 Dorsal soft-rays , 16 I 2 17 5 1 6 18 25 3 11 3 42 i : ; I (~: +?) 19 19 5 5 6 35 ~g 20 2 ] I (+1) 2 (+1) (2) i(Usual) I 21 1 (I, +?)I No. 3 49 8 18 10(+1) 85 (+1) 1(+2) (11) 53 Mean 20.3 18.3 18.6 18.3 ]8.8 18.4 19.0 18.5

Anal spines-3 3 49 8 18 10 (+1) 85 (+2) I 1 (+2) (12) 53 Anal soft-rays 14 8 I 1 I 11 4 15 39 7 16 9 71 I 41

16 2 1 (1) 3 (+1) (2) (Usual) 1 8 17 No. 3 49 8 18 10 (+1) 85 (+1) ' . i (+2) ~ht)1 53 Mean 14.9 14.9 15.0 14.9 14.9 I 15.1 15.0 15.0 I" , Caudal principal rays 8 8 = 16 3 I 4 1 3 + .. I 9 + 8 = 17 3 46 6 18 9 79 2 49 9 + 9 = 18 1 No. 3 49 6 18 10 83 3 I •. 52 Mean 17.0 16.9 17.0 17.0 16.9 17.0 17.3 I .. 16.9 Pectoral (both sides) 13 3 3 6 1 Il 14 2 89 14 27 18 148 , 2 (l, +?), 92 ] IS 2 6 4 2 13 2 (+1) (Usual) I' 3 16 1 .. 1 No. 4 98 16 34 20 168 5(+])\(11) 106 Mean 14.5 14.0 14.0 14.0 14.] 14.0 , 14.3 13.9 Pelvic rays 1,5-1,4 I .. I 1,5-1,5 3 48 8 18 11 85 I (12) 53 1963] Hubbs: Chaetodon Aya and Related Butterflyfishes 179

TABLE 2

VARIATION IN NUMBER OF PORES IN LATERAL LINE IN DEEP-LIVING SPECIES OF Chaetodon AND Prognathodes Both sides were counted. Poreless scales within course of lateral line are included. Regions under C. aya are: !, South Atlantic coast, U. S. A.; II, Straits of Florida; III, NE. Gulf of Mexico; IV, Gulf of Campeche. Counts from the literature (in parentheses) are included In the computed means, except for C. marcellae.

------~ ---- I C. lal-\ Chaelodon ara I c. guy- C. mar- P. acul- I ciler II III IV Total anensis cellae ea/us - ----i-- Pores in I lateral line 22 2 23 11 24 22 25 3 3 19 26 9 2 (1) 11 (+1) 25 27 12 2 2 2 18 13 28 23 6 3 32 1 2 29 ]4 3 4 (+1) 21 (+1) (1) 30 14 4 3 6 27 1 (+1) (1) 31 11 2 14 X

32 6 8 X 33 2 2 X

34 (I) X

35 X 36 X

42 43 44 45 46 3 No. 6 95 14 10 18 (+2) 137 (+2) 2(+3) (II) 94 Mean 44.8 28.8 28.1 29.4 29.2 28.8 30.2 25.1 180 Bulletin ot Marine Science of the Gulf and Caribbean [13(1) •... OJ) " ....• ~ N "< ..s:: C. l>:: i.J 0 f-o J!J ..• .•..• PJ l>:: c 0 0 ::l "U ! ~ p,. 0 "5 >. ~ u 0•.... a- a- •... ;... E" ". '" >. -'"0 u c « [/) p,. E "::l ';;J 0' 0 ';;J "~ ~ .g 't .Q ••... Vi'" .•..• OIl >< "0 ,..., ,..., .>- E 0 ..J •... ,...,'0 » E ;;- .~ " 00 V) '" " "E ..c

TABLE 4

FIN-RAY CORRELATIONS IN Chaetodon aya AND Prognathodes aculeatus

------Dors'al soft-rays - Dorsal spines Cilaetodoll a)'a Progllathodes aculea/us 17 18 19 20 16 17 18 19 12 ...... 2 ...... 1 13 ••••••••• '0. 5 39 33 2 1 1 20 29 14 ...... I 3 .. . . 1 ...... r = -0.26 ± 0.10 .! = -0.41 ± 0.11

Anal soft-rays Dorsal soft-rays ------Chaetodoll aya Prognathodes aculeatus - 17 18 19 20 16 17 18 19 14 ...... 2 7 2 .. 1 .. 1 2 ]5 . , ...... 4 35 30 2 1 1 17 22 16 ...... 3 ...... 2 6 ---- ~= +0.28 ± 0.10 r - +0.65 ± 0.08 ------,-

TABLE 5

SPEC]FICALLY COMMON OCCURRENCES OF FISH TYPES ABOUT THE PEN]NSULAS OF YUCATAN AND FLORIDA

Genus or Genera (and Family) Repres'entative in Yucatan F]orida Authority Floridichthys F. carpio polyommus F. carpio carpio Hubbs, 1936 (Cyprinodontidae) F. carpio barbouri Garmam;lIa and. Jordanella G. pulchra J. floridae Hubbs, 1936 (Cyprmodonl1dae) . Chriodorus (Hemiramphidae) *. C. atherinoides C. a/herinoides Hubbs, 1936 Chaetodoll (Chaetodontidae) .. Chae/odoll aya Chae/odoll aya Hubbs and Rechnitzer. 1958; present paper Menidiat (Atherinidae) M. colei M. concho rum Hubbs, 1936

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LITERATURE CITED The numbers in parentheses following certain citations indicate the collections (corres- pondingly numbered in Table 6) that were stated to have been utilized, or may be assumed to have been reported, in the given citation. ADAMS, RICHARD M. AND ERNEST F. SORGNIT 1951. Comparison of summer and winter sea temperatures (Gulf of Mexico). A.&M. ColI. Texas, Dept. Oceanogr., Project NR 083 036, Rep. 3: 8 pp., figs. 1-3. AHL, ERNST 1923. Zur Kenntnis der Knochenfischfamilie Chaetodontidae, inbesondere der Unterfamilie Chaetodontinae. Arch. Naturgesch. (Abt. A) 89 (5): 1-205, pis. 1-2. (5, 6) BAILEY, REEVE M., ERNEST A. LACHNER, C. C. LINDSEY, C. RICHARD ROBINS, PHIL M. ROEDEL, W. B. SCOTT, AND LOREN P. WOODS 1960. A list of common and scientific names of fishes for the United States and Canada (2nd ed.). Amer. Fish. Soc. Spec. Pub I. 2: 1-102. (6,65, etc. ) BAUCHOT, M. L. AND M. BLANC 1961. Poissons marins de l'Est Atlantique tropical. II. Percoidei (Teh~osteens Perciformes) 1 ere partie. Atlantide Rep., 6: 65-100, figs. 1-4, pI. 1. (42,43,48) BLEEKER, PIETER 1876a. System a percarum revisum. Pars II. Arch. neerl. Sci. (Ser. B, Sci. Nature!.), 11: 289-340. (49,50) 1876b. Notice sur les genres et sur les especes des Chetodontoides de la sousfamille des Taurichthyiformes. Vers!. Meded. Kon. Akad. Wet- ensch. Amsterdam, Afd. Natuurk., (2) 10 (3): 308-320. (49, 50) 1877. Revision des especes insulindiennes de la famille des Chetodontoides. Verh. Akad. Wet. Anst. (Afd. Natuur.), 17: 1-174. (49, 50) BREDER, CHARLES M., JR. 1929. Field book of marine fishes of the Atlantic coast from Labrador to Texas. G. P. Putnam's Sons, New York and London: i-xxxvii, 1-332, many figs. (6) BRIGGS, JOHN C. 1958. A list of Florida fishes and their distribution. Bull. Fla. St. Mus. bioI. Sci., 2 (8): 223-318. (4,9-11) CADENAT, J. 1951. Poissons de mer du Senegal. Init. afr., 3: 1-345, figs. 1-241. [Not "1950"-sec Hubbs and Rechnitzer, 1958: 304.1. (45) 1953. Notes d'lchtyologie ouest-africaine. VI.-Poissons. Bull. Inst. Franc;. Afr. noire (A), 15 (3): 1051-1102, figs. 1-43. (44-47) 1954. Note d'lchtyologie ouest-africaine. VII.-Biologie. Regime alimen- taire. Bull. Inst. Franc;. Afr. noire (A), 16 (2): 564-583. (44-47) 1960. Notes d'lchtyologie ouest-africaine. XXX.-Poissons de mer ouest- africa ins observes du Senegal au Cameroun et plus specialement au large des Cotes de Sierra Leone et du Ghana. Bull. Inst. Franc;. Afr. noire (A), 22 (4): 1358-1420, pIs. 1-3. (44-48) 1961. Notes d'Ichtyologie ouest-africaine. XXXIV.-Liste complementaire des especes de Poissons de mer (provenant des cotes de I'Afrique occidentale) en collection a la Section de Biologie Marine de 1'1. F. A. N. a Goree. Bull. Inst. Franc;. Afr. noire (A), 23 (1): 231-245. (44-48) CALDWELL, DAVID K. 1962. The ecology and systematics of the shore fishes of Jamaica [report on grantl Yearb. Amer. phil. Soc., 1961: 275-277. 190 Bulletin of Marine Science of the Gulf and Caribbean [13(1)

COLLINS, HENRY H. 1956. Birds and fish of the Netherlands Antilles (Netherlands West Indies). Caribou Press, Bronxville, New York: 18 pp., 25 figs. [fish part prepared by J. S. Zaneveld (fide Zaneveld, 1956: 5, and personal communication, 1958)]. (56, etc.) DURAND, J. 1960. Chaetodontidae (Poissons teleosteens Percoidei) recoltes au large de la Guyane. Description d'une espece nouvelle. Bull. Mus. Hist. nat. Paris, (2),32 (3): 209-213, figs. 1-3. (Pub. Aug. 5,1960.) (40) EIGENMANN, CARL H. AND JENNIE E. HORNING 1887. A review of the Chaetodontidae of North America. Ann. N.Y. Acad. Sci., 4 (7-8): 1-18. (6,49,50) ERDMAN, DONALD S. 1956. Recent fish records from Puerto Rico. Bull. Mar. Sci. Gulf & Carib., 6 (4): 3]5-340. (58) 1961. New fish records from Puerto Rico. Caribbean J. Sci., 1 (2): 39-40. (58?, 66, 68, 81) FUGLISTER, FREDERICK C. 1947. Average monthly sea surface temperatures of the western North Atlantic Ocean. Pap. phys. Oceanogr., 10 (2): 1-25, 2 figs. 16 pIs. GILL, THEODORE 1862. Remarks on the relations of the genera and other groups of Cuban fishes. Proc. Acad. nat. Sci. Philad., 14 (5): 235-242. (49) GUNTHER, ALBERT 1860. Catalogue of the acanthopterygian fishes in the collection of the British Museum. Squamipinnes, Cirrhitidae, Triglidae, Trachinidae, Sciaenidae, Polynemidae, Sphyraenidae, Trichiuridae, Scombridae, Carangidae, Xiphiidae. London, 2: i-xxi, ]-548, 1 fig. (49) HEDGPETH, JOEL W. 1953. An introduction to the zoogeography of the northwestern Gulf ot Mexico with reference to the invertebrate fauna. Pub!. Inst. Mar. Sci. Univ. Texas, 3 (1): 107-224, figs. ]-46. HUBBS, CARL L. 1936. Fishes of the Yucatan Peninsula. Publ. Carneg. Instn., 457: 157-287, fig. 1, pis. 1-15. 1943. Criteria for subspecies, species and genera, as determined by re- searches on fishes. Ann N.Y. Acad. Sci., 44 (2): 109-121. 1952. Antitropical distribution of fishes and other organisms. Proc. Seventh Pacif. Sci. Congr., 3: 324-329. 1958. Dikellorhynchus and Kanazawaichthys: nominal fish genera interpreted as based on prejuveniles of Malacanthus and A ntennarius, respectively. Copeia, 1958 (4): 282-285. 1960. Fauna Japonica [review]. Copeia, 1960 (4): 381-383. HUBBS, CARL L. AND LAURA C. HUBBS 1945. Bilateral asymmetry and bilateral variation in fishes. Pap. Mich. Acad. Sci., 30: 229-310, figs. 1-2, pI. 1. 1954. Data on the life history, variation, ecology, and relationships of the kelp perch, Brachyistius frenatus, an embiotocid fish of the Cali- fornias. Calif. Fish Game, 40 (2): 183-]98, figs. 1-3. HUBBS, CARL L. AND ANDREAS B. RECHNITZER 1958. A new fish, Chaetodon falcifer, from Guadalupe Island, Baja Cali- fornia, with notes on related species. Proc. Calif. Acad. Sci. (4th Ser.), 29 (8): 273-313, pis. 1-3, 1 map. (1, 6-10, 13, 14, 16, 17, 19, 26, 44, 45, 49, 50) 1963] Hubbs: Chaetodon Aya and ReLated Butterflyfishes 191

HUBBS, CLARK 1952. A contribution to the classification of the blennoid fishes of the family Clinidae, with a partial revision of the Eastern Pacific forms. Stanford ichthyol. Bul!., 4 (2): 41-165, figs. ]-64, graphs 1-79. JORDAN, DAVID S. 1886. Notes on some fishes collected at Pensacola by Mr. Silas Stearns, with descriptions of one new species (Chaetodon aya). Proc. U.S. nat. Mus., 9: 225-229. (6) ]887. A preliminary list of the fishes of the West Indies. Proc. U.S. nat. Mus., 9, ]886: 554-608. (50) JORDAN, DAVID STARR AND BARTON WARREN EVERMANN 1898. The fishes of North and Middle America: a descriptive catalogue of the species of fishlike vertebrates found in the waters of North America, north of the Isthmus of Panama. Bull. U.S. nat. Mus., 47 (2): i-xxx, 1241-2183, 1 fig. (6, 50) LEIPPER, DALE F. ]954. Physical oceanography of the Gulf of Mexico. In: Gulf of Mexico / Its origin, waters, and marine life. Fish. Bull., U.S., 55 (89): ] ]9-137, figs. 34-43. LONGLEY, WILLIAM H. AND SAMUEL F. HILDEBRAND 1941. Systematic catalogue of the fishes of Tortugas, Florida, with observations on color, habits, and local distribution. Publ. Carneg. Instn. 535, Pap. Tortugas Lab., 34: i-xiii, 1-331, pis. 1-34. (9, 10) METZELAAR, J. 1922. On a collection of marine fishes from the Lesser Antilles. Bijdr. Dierk., 22: 133-141. (52) MYERS, GEORGE S. ] 959. A Caribbean chaetodont fish, Chaetodon eques Steindachner, now referred to Chaetodon aya Jordan. Copeia, 1959 (2): 158. (5, 6) NICHOLS, J. T. AND F. E. FIRTH ]939. Rare fishes off the Atlantic coast, including a new grammicolepid. Proc. bioI. Soc. Wash., 52: 85-88, fig. 1. (11) PARR, ALBERT EIDE 1935. Report on hydrographic observations in the Gu]f of Mexico and the adjacent straits made during the Yale Oceanographic Expedition on the "Mabel Taylor" in 1932. Bull. Bingham oceanogr. Col!., 5 (1): 1-93, figs. 1-33. PHILLIPS, CRAIG 1958. A "lost" fish rediscovered. Aquarium J., 29 (11): 410-412,3 figs. PHLEGER, FRED B 1951. Foraminifera distribution. In: Ecology of Foraminifera, northwest Gulf of Mexico, by Fred B Phleger and Francis L. Parker. Part 1. Mem. geo!. Soc. Amer., 46: v-ix, 1-88, figs. 1-33. POEY, FELIPE 1860-1861. Poissons de Cuba. Memorias sobre la historia natural de ]a isla de Cuba, acompafiadas de sumarios latinos y extractos en frances, 2 (49): 115-356, pis. 12-19 (pp. 115-336 published 1860). (50) 1868. Synopsis piscium cubensium. Repertorio fisico-natural de ]a isla de Cuba, 2.' 279-484 (reprinted, in same year, with same pagination and with added title, Catalogo razonado de los peces de la isla de Cuba). (50) POLL, MAX 1950. Description de deux Poissons percomorphes nouveaux des eaux 192 Bulletm of Marine Science of the Gulf and Caribbean [13(1) cotieres de I'Atlantique Sud (1948-1949). Bull. Inst. Sci. nat. Belg., 26 (49): 1-14, figs. 1-2. (44) RANDALL, JOHN E. 1961. Two new butterflyfishes (family Chaetodontidae) of the Indo-Pacific genus Forcipiger. Copeia, 1961 (1): 53-62, figs. 1-6. (1, 6, 50, etc.) RANDALL, JOHN E. AND CARLETON RAY 1958. Bahamian land-and-sea park. Sea Frontiers, 4(2): 72-80, figs. 63,66) RAY, CARLETON 1960. The adventure book of underwater life. Capitol Pub!. Co., New York: 1-95, many figs. Roux, CH. 1957. Poissons marins. In: Collignon, J., M. Rossignol, and Ch. Raux, Mollusques, Crustaces et Poissons marins des cotes d'A. E. F. en collection au Centre d'Oceanographie de l'Institut d'e.tudes Cen- trafricaines de Pointe-Noire. Office de la Recherche Scientifique et Technique Outre-Mer, Paris: 137-368, figs. 1-121, 1 pI. (44-48) SCHULTZ, LEONARD P. 1948. A revision of six subfamilies of atherine fishes, with descriptions of new genera and species. Proc. U.S. nat. Mus., 98 (3220): 1-48, figs. 1-9, pIs. 1-2. SMITH, SANDERSON 1889. Lists of the dredging stations of the U.S. Fish Commission, the U.S. Coast Survey, and the British Steamer Challenger, in North American waters, from 1867 to 1887, together with those of the principal government expeditions in the Atlantic and Arctic oceans. Rep. U.S. Comm. Fish., 14: 871-1017, charts 1-5, figs. 1-4. (1886.) (5) SPRINGER, STEWART AND HARVEY R. BULLIS, JR. 1956. Collections by the Oregon in the Gulf of Mexico. Spec. Sci. Rep. U.S. Fish Wild. Serv., Fish., (196): 1-134 (16-18, 20-22) STEARNS, SILAS 1887. The fishing-grounds of the Gulf of Mexico belonging to the United States. In: George Brown Goode and associates, Fisheries and Fishery Industries of the United States, United States Commission of Fish and Fisheries, Sect. 3: 55-58, charts 16-17. (6) STEINDACHNER, FRANZ 1903. Ober einige neue Reptilien- und Fischarten des Hofmuseums in Wien. S.B. Akad. Wiss. Wien, 112: 15-21, figs. 1-2a. (5) TOWNSEND, C. H. 1901. Dredging and other records of the United States Fish Commission Steamer Albatross, with bibliography relative to the work of the vessel. Rep. U.S. Fish Comm., Pt. 26 (1900): 387-562, pIs. 1-7 (7) WOODS, LOREN P. 1961. Chaetodon goniodes, a new butterfly fish from Puerto Rico. Bull. Mar. Sci. Gulf & Carib., 10 (4): 417-420, fig. 1. (41) WOODS, LOREN P. AND ROBERT H. KANAZAWA 1951. New species and new records of fishes from Bermuda. Fieldiana, Zool. 31 (53): 629-644, figs. 134-137. ZAN EVELD, J. S. 1956. Eenige algemeen voorkomende zeevissen in de Nederlandse Antillen. De West-Indische Gids, 37 (1): 5-17, pIs. 1-2. (See also Collins, 1956). (56, 57) ZIM, HERBERT S. AND HURST H. SHOEMAKER 1956. Fishes-A guide to fresh- and salt-water species. Golden Nature Guide, Simon and Schuster, New York: 1-160, many figs.

&lt;I&gt;Chaetodon Aya&lt;/I&gt; and Related Deep-Living Butterflyfishes

<I>Chaetodon Aya</I> and Related Deep-Living Butterflyfishes