BULLETIN OF MARINE SCIENCE. 29(3): 312-319. 1979
REDISCOVERY AND REDESCRIPTION OF THE CARIBBEAN ANOMALOPID FISH KR YPTOPHANARON ALFREDl SILVESTER AND FOWLER (PISCES: ANOMALOPIDAE)
Patrick L. Colin, Deborah W. Arneson, and William F. Smith- Vaniz
ABSTRACT The Caribbean anomalopid fish Kryptophanaron alfred; is redescribed from one specimen collected off western Puerto Rico at 200-m depth, and six specimens from Grand Cayman Island taken in 30-36 m. These specimens differ from the original description in lacking vomerine teeth and in having only two anal spines. Live specimens are now being maintained in aquaria. White scales at the bases of the second dorsal and anal fins may serve as "re- f1ectors." The species is easily distinguished from its eastern Pacific relative K. harvey; Rosenblatt and Montgomery, in having more ventral scutes (7-9 versus 13) and smaller scales (ca. 120-140 scale rows along the back vs. ca. 80).
The description of Kryptophanaron alfredi by Silvester and Fowler (1926) was based on a single specimen found floating on the surface south of Kingston, Jamaica. This specimen, deposited in the Yale University fish collection, was subsequently lost and presumed destroyed during the 1930's. Interest in the bi- ology of the Anomalopidae has increased greatly during the past decade and the discovery of a second species of the genus Kryptophanaron, K. harveyi, in the Gulf of California in 1972 (Rosenblatt and Montgomery, 1976) has stimulated considerable interest in the enigmatic Caribbean member of the family. Impetus for this paper was the collection of a single specimen of K. alfredi taken in a deep fish pot west of Puerto Rico. After the original manuscript had been accepted for publication, the junior author called to our attention six spec- imens that he and several colleagues had recently collected at Grand Cayman Island. This new material together with the Puerto Rican specimen forms the basis for the redescription of the species. Subsequently a number of live specimens have been collected both in Puerto Rico and the Cayman Islands allowing behavioral observations.
Kryptophanaron Silvester and Fowler Some statements given in the original description of Kryptophanaron do not agree with our specimens and are considered to be erroneous. Silvester and Fowler (1926: 246) cited the presence of villiform teeth on the vomer in the generic description, yet on the same page in the species description they stated that there were teeth on "jaws and palatine; vomer and tongue toothless." This discrepency was noted by Rosenblatt and Montgomery (1976: 512). The reference given in the generic description is an obvious error, as all our specimens lack vomerine teeth. Silvester and Fowler (1926: 246) reported 3 anal spines in both the generic and species descriptions. Our specimens possess only 2 anal spines as does Krypto- phanaron harveyi (Rosenblatt and Montgomery, 1976). The number of dorsal spines (6) in our specimens agrees with that reported by Silvester and Fowler (1926), but differs from the 5 spines reported for K. harveyi. The last spine in K. alfredi is segmented distally in some of our specimens and is very difficult to distinguish from a "soft" ray. The discrepancy in number of dorsal-fin spines
312 COLIN ET AL.: CARIBBEAN ANOMALOPID FISH 313
A
B
Figure I. Kryptophanaron alfredi Silvester and Fowler. A) Puerto Rican specimen (ANSP 138144, 105.6 mm SL) from 200-m depth. Negative reversed to bring specimen to normal head left orientation. Photographed shortly after collection, and white color of the scales at the base of the second dorsal fin is still apparent. B) Grand Cayman Island specimen (ANSP 136510, 64.4 mm SL) showing white scales at base of dorsal and anal fins. reported for the two species of Kryptophanaron is more apparent than real and results from differences of interpretation. A variant spelling of the generic name (Kryptophaneron) was introduced by Fowler (1944: 461) and followed by Woods and Sonoda (1973) and Rosenblatt and Montgomery (1976); in accordance with the recommendation of the Inter- national Code of Zoological Nomenclature, we here adopt the original spelling.
Kryptophanaron alfredi Silvester and Fowler Figures 1-2
Anomalops sp., Dahlgren, 1908: 454-455. Kryptophanaron alfredi Silvester and Fowler, 1926; Jordan, Evermann and Clark, 1930: 234 (listed); Caldwell, 1966: 37 (listed); Marshall, 1966: 179; McCosker, 1977: 106-114, Kryptuphaneron alfredi , Fowler, 1944: 461 (listed); Woods and Sonoda, 1973: 328-330; Rosenblatt and Montgomery, 1976: 510-515, 314 BULLETIN OF MARINE SCIENCE, VOL. 29, NO.3, ]979
Material Examined.-ANSP 138144(1, 105.6 mm SL), 8.5 km west of buoy "8" (Tourmaline Reef) off western Puerto Rico (approximately 18°10'N; 67°27'W) in 2oo-m depth, fish trap, 10 June 1977. Evengelista Silva and Victor Padilla. ANSP 136508(1, 65.2 mm), Grand Cayman Island, southwest coast opposite Spot Bay at edge of drop-off, 30 m, 27 January 1978, 0100-0120 h, W. F. SmiIh- Vaniz and Paul Humann. ANSP 136509(I, 70.2 mm), Grand Cayman Is., West Bay, ca. 0.2 km south of Northwest Point, "Orange Canyon" at drop-off, 30.5 m, 27 January 1978, 1940-2000 h, W. F. Smith-Vaniz. ANSP 136510 (2,58.0-64.4 mm), Grand Cayman Is., West Bay, ca. 0.2 km south of Northwest Point, "Orange Canyon" at drop-off, 30-36 m, 27 January 1978, 1800-1820 h, Paul Humann and Don Kincaid. ANSP 136511 (2, 60.0--84.4 mm), Grand Cayman Is., southwest coast opposite Spot Bay at edge of drop-off, 30 m, 26 January 1978, 1800-1820 h, W. F. Smith-Vaniz and Paul Humann. Description.-Dorsal-fin rays IV-II, 14; anal-fin rays II, 10; pelvic-fin rays I, 6; pectoral-fin rays 16 or 17 (typically 16); caudal fin rays 9-11 procurrent, 10 + 9, 8-10 procurrent; branchiostegals 8, gill rakers 6 or 7 + 18-20; vertebrae 12 + 17. Morphometric and meristic characters of the recent specimens are compared with those of the lost holotype in Table 1. The Puerto Rican specimen is shown in Figure la and one Cayman Island specimen in Figure lb. Body compressed, width 2.0 in depth. Snout blunt. An- terior nostril about 4.0 in posterior. Mouth oblique, the posterior end of the maxilla reaching to the midline of the eyeball. Maxilla with large posterior su- pramaxilla covering most of its posterior one-third with a narrow process pro- jecting anteriorly along the dorsal surface of the maxilla. A small anterior supra- maxilla located on the outer surface of the maxillary ahead of the posterior supramaxilla. Premaxilla somewhat protrusible with a deep notch at the sym- physis. A projection of the lower jaw matches this notch so that a gap does not exist when the mouth is closed. Surface of preopercle slightly sculptured. Margin entire except for a few ser- rations at the angle. Opercular series sculptured with low ridges. Cleithrum sculp- tured and expanded above the pectoral fin base. Supracleithrum and posttemporal exposed. Circumorbital series and cheek covered by roughened patches, but scales may not be present. Cephalic canals of lateral line system well developed, often resembling ramose cavities covered by skin rather than tube-like canals. Subocular canal with 3 openings anteriorly through bone followed by 5 skin-covered cavities with small pores. Postorbital canal with opening on rim of orbit near tubercle. Supraorbital canal with 2 openings anteriorly, one below the nares and the second at the anterior end of a large skin-covered fossa above the nares. The preopercular canal is dorsally narrow, with 3 wide skin-covered channels on its lower one-half, particularly near the angle. The mandibular canal has 3 pores, the anterior two opening through bone and the posterior at the end of another skin-covered depres- sion. The canals of the dorsal surface of the head have no pores and are not roofed by bone. The enlarged pored lateral-line scales number 32-34. Eye large, about one-third head length. Tubercle inside posterior rim of orbit 1.5 mm in height. Prominent luminous organ below eye in orbit, attached by its anterior end. As in some other anomalopids, this organ is partially rotatable (Dahlgren, 1908). Black membrane in the lower portion of the orbit is capable of being raised to occlude the light organ when it is ventrally rotated (Fig. 2). This membrane assists in complete occlusion of the light. G. David Johnson and Rich- ard H. Rosenblatt (pers. comm.) will present a detailed description of the mech- anism of light organ occlusion in a subsequent paper now in preparation. On the premaxilla, the outer, inner and lower surfaces are paved with tiny, knoblike teeth which are expanded on the edges of the symphyseal notch. There is a band directly at the symphysis which lacks teeth. Lower teeth are similar, COLIN ET AL.: CARIBBEAN ANOMALOPID FISH 315
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' Figure 2. Specimen of Kryptophanaron alfredi with light organ occluded resting in diver's hand after being blinded by a diver's light. The reflective nature of the margins of much of the fins, the soft rays, the scutes and basal scales is apparent. Grand Cayman Island, depth about 30 m, photo by F. McConnaughey. but laterally restricted to the upper surface of the jaw. Teeth on the knoblike projection on the anterior end of the lower jaw expanded to cover all the enlarged tip and extend ventrally on the lower one half of this bone. There is a narrow band lacking teeth at the symphysis of the dentaries. The teeth on the palatines are similar, but slightly smaller, covering the entire exposed surface of the bone. The vomer is toothless. Body covered with tiny ctenoid scales, extremely difficult to enumerate. There are about 140 scale rows in lateral series. Between the lateral line and the dorsal- fin origin there are about 15 rows; about 30 rows occur between the lateral line and the anal-fin origin. There are about 10 rows of larger cycloid scales behind the pectoral-fin base. Weakly developed pelvic-fin sheath present. The chest and belly are densely scaled and dorsally scales reach to the rear of the belly between the pelvic fin and the anus. The second dorsal fin and the anal fin have a sheath of enlarged, plate-like ctenoid scales at their bases extending outward almost forming a groove in which the fin can be folded. Several hours after capture of the Puerto Rican specimen, 9 of the right and 6 of the left enlarged scales were white while other adjacent scales were as dark as the remainder of the body. A few of the scales at the anal- fin base were also white. The Cayman Island specimens had the enlarged scales similar in color. Underwater photographs taken of these before collection show that the basal scales, lateral-line scutes and belly scutes were light reflective. The tips of the first 5 dorsal spines together with much of each soft ray of the dorsal, anal and pelvic fins were also reflective. The entire margin of the caudal fin and the rear margins of the soft dorsal, anal and pelvic fins were reflective (Fig. 2). No portion of the pectoral fin was reflective. The only indication of this property in preserved specimens was the pale color of the enlarged scales and scutes with all trace of the reflective margins of the fins and their rays being lost. While these reflective areas have no light-producing function, they may serve to reveal the position of an individual with its light organs occluded to another which is emitting COLIN ET AL.: CARIBBEAN ANOMALOPlD FISH ·317 light. There is variation in the numbers and position of the "reflectors" so that individual recognition is potentially possible. The first dorsal fin has 4 spines, the 2nd is the longest and the 4th shortest. The second dorsal fin has 2 spines, the first short and about one-half the length of the second. The 4th soft ray is the longest and rays 2-14 have branched tips. The anal fin has 2 spines, the second nearly twice the length of the first. The first soft ray is the longest with rays 3 through 10 bifurcate. The rear margin of the fin is concave. The pectoral fins are short, about 0.6 to 0.7 in head, fourth and fifth rays longest and subequal. The pelvic fins have the 2nd soft ray longest and the last ray is slightly longer than the spine. The caudal fin is deeply forked. The second ray is longest on the upper lobe while the 18th is longest on the lower lobe. The shortest caudal ray, the 10th, goes into the longest over 4 times. The color is predominantly black with the head and fins darkest. The sides are slightly lighter. The lateral-line scutes and ventral scutes are white, but become less conspicuous after preservation. The light organ is pale with visible blood vessels. The membranes of the upper jaw and the inside of the mouth are pale. The light emitted by the light organ is blue-green in color. Discussion.-We have a report of 3 other specimens of K. alfredi collected at one time under circumstances similar to our original collection in Puerto Rico. The trap in which the original K. alfredi was collected had been baited with tuna roe and set for 3 days. The nature of the bottom where the trap was set is unknown. The Mona Channel, however, has many areas of high relief at 100-400 m depths and although the general area around the collection area does not have great slope, the bottom could be quite rugged. The Cayman Island specimens were all collected at night with lights and hand nets in 30-36 m on the crest of a nearly vertical drop-off into deep water. The bottom was characterized by deep erosion channels, undercut ledges and nu- merous caves. The gut contents of three of the Grand Cayman specimens were examined. One fish (ANSP 136509) collected about 4 h after sunset had the stomach jammed full with at least 45 shrimps of 5-10 mm rostral length (RL) and a few copepods. The other two individuals (ANSP 136510) collected shortly after sunset had less material in the stomach with one having only three shrimps of 8-16 mm RL and the other with over 120 copepods averaging 1.5 mm in length and a single shrimp of 10 mm RL. The intestines of the fish were nearly empty and the items in the stomach poorly digested. Puerto Rican specimens maintained in aquaria accepted a variety of small shrimps and fishes as food. In the field K. alfredi actively swam close to the bottom apparently inspecting the reef face for prey. Since it does not remain stationary for more than a few seconds, it seems unlikely that it attracts organisms with its luminescent organs. Rather it uses these organs to search out prey on the reef. The gut of K. alfredi is short, being nearly the same or slightly less than the standard length of the fish. It is possible that the fish rise into shallow water and feed only long enough to fill the stomach and then return to depth below diving range, even though it is still dark. There were indications of such behavior during the dives to collect specimens. The Puerto Rican specimens maintained alive were similarly collected at the edge of a steeply sloping drop-off at 27-40 m depth south of Margarita Reef, La Parguera. The fish were present in this area about I h after sunset. They moved 318 BULLETIN OF MARINE SCIENCE. VOL. 29. NO.3. 1979 steadily over the bottom but seemed undisturbed by a diver with light extin- guished. In some instances the fish approached the stationary observer closely. The light organs are normally occluded for less than ] sec when the fish changes direction. One group of 9 fish was observed at 36 m close to the substrate in a volume of water less than] m3• They were "blinking" the light organs rapidly and swimming rapidly around one another in an action which may have been courtship or spawning behavior. Specimens collected alive and returned to aquaria survived well being brought directly to the surface. The females in the aquaria commonly shed infertile eggs into the water for at least 3 days after collection during April. The eggs were 1.6 mm in diameter with numerous small oil droplets. The potential for obtaining fertile eggs in the laboratory is good. In addition to the Puerto Rican specimens being ripe, three of the 6 Cayman Island specimens collected in January were ripe females. We believe that K. alfredi will prove to be broadly distributed throughout the Caribbean where suitable habitat exists. John E. McCosker (in litt.) has informed us of the recent collection of two additional specimens taken in 60-76 m at Cu- ra\;ao. The rarity of specimens in museum collections simply reflects the difficulty of collecting the species. During daylight hours K. alfredi occurs in habitats unsuitable for trawling and at depths too deep for SCUBA diving. Although the species is now known to ascend to relatively shallow depths at night, the timing and duration of the upward movement may be strongly influenced by the moon; there was a noticeable decrease in the number of individuals observed at Grand Cayman Island when the moon had risen and was unobstructed by clouds. Kryp- tophanaron alfredi is very sensitive to strong illumination and rapidly seeks cover when dive lights are turned on. Thus, successful capture of specimens requires initiation of nocturnal dives, closely approaching an individual and only then blinding it with a strong light so that it can be dip-netted. Although the blue-green light organs of K. aifredi can be detected underwater from a distance of ap- proximately 10-]5 m, initially it is often difficult to correctly identify the struc- ture, especially in areas where bioluminescent invertebrates are relatively com- mon. ACKNOWLEDGMENTS We thank the two fishermen, E. Silva and V. PadiUa, who called to our attention the first Puerto Rican specimen of K. alfredi. The junior author is indebted to Fred McConnaughey who organized and provided financial support for the Grand Cayman Island trip. The successful capture of the Grand Cayman Island specimens was greatly facilitated by fellow divers Capt. P. Humann, D. Kincaid and F. McConnaughey. Dr. R. H. Rosenblatt read and made valuable comments on the manuscript. C. Arneson assisted in photographing the Puerto Rican specimen and collecting live individuals. LITERATURE CITED Caldwell. D. K. 1966. Marine and freshwater fishes of Jamaica. Bull. Inst. Jamaica. Sci. Ser. 17: 1- 120. Dahlgren, U. ]908. The luminous organ of a new species of Anomalops. Science 27: 454-455. Fowler, H. W. 1944. Results of fifth George Vanderbilt expedition (1941). The fishes. Acad. Nat. Sci. Philadelphia, Monograph 6: 57-529. Jordan, D. S., B. W. Evermann, and H. W. Clark. 1930. A checklist of the fishes and fishlike vertebrates of North and Middle America north of Venezuela and Colombia. Rpt. U.S. Comm. Fish. for 1928, part 2: 1-670. Marshall, N. B. 1966. The life of fishes. World Pub!. Co., Cleveland. 402 pp. McCosker. J. E. 1977. Flashlight fishes. Sci. Amer. 236: 106-114. Rosenblatt, R. H., and W. L. Montgomery. ]976. Kryptophaneron harveyi, a new anomalopid fish from the eastern tropical Pacific and the evolution of the Anomalopidae. Copeia ]976: 510-515. COLIN ET AL.: CARIBBEAN ANOMALOPID FISH 319 Silvester. C. F., and H. W. Fowler. 1926. A new genus and species of phosphorescent fish, Kryp- topllllna/"Onalfredi. Proc. Acad. Nat. Sci. Philadelphia 78: 245-247. Woods, L. P., and P. Sonoda. 1973. Order Berycomorphi (Beryciformes). Pages 263-396 in Fishes of the western North Atlantic, part 6. Mem. Sears Fndtn. Mar. Res. No. I. DATEACCEPTED: January 9, 1978. ADDRESSES: (PLC) Departme/11 of Marine Sciences. University of Puerto Rico, Mayagaez, Puerto Rico 00708; (DWA) Commercial Fisheries Laboratory, Department of Agriculture, Commonwealth of Puerto Rico. P.O. Box 3665, Mayaguez, Puerto Rico. 00708; (WFS-V) Academy of Natural Sci- {'IICCS of Philadelphia, 19th and the Parkway, Philadelphia, Pennsylvania, /9103.