BULLETIN OF MARINE SCIENCE. 33(4): 846-854,1983

SYNOPSIS OF THE SUBFAMILY (PISCES: CARAPIDAE)

Jeffrey T. Williams

ABSTRACT The pearl fish subfamily Pyramodontinae is characterized by the following synapomorphies: 24-30 pectoral fin rays, distinctive upward curvature of the posterior portion of the lateral canal above the opercle, visceral cradle, elaborate predorsal bone and relatively deep-bodied vexillifer larvae with short predorsal length, Pyramodon differs from Snyderidia in having pelvic fins present, pterygiophores 3-22 to 24 forming visceral cradle (3-13 in Snyderidia) and total lamellae in olfactory rosettc 13 (15-16 in Snyderidia), The Atlantic Ocean S. bothrops differs from the Indo-Pacific S. canina in having one row of dentary teeth (2-3 rows in the latter). The geographic range of Pyramodon is extended to the eastern Pacific, off Chile, and that of Snyderidia bothrops from the Caribbean off Panama north to the Gulf of Mexico.

Additional material belonging to the genera Pyramodon and Snyderidia ob- tained from the National Museum of Natural History, Smithsonian Institution, represents significant geographic range extensions and allows a comparison ofthe genera. Analysis of this material revealed several characters possessed by these specimens that had not been noted previously. These and other diagnostic char- acters are presented herein along with geographic range extensions. The pearl fish family Carapidae contains the subfamilies Pyramodontinae and Carapinae, which Trott (1981) recognized as distinct families, although he gave no reason for this action. I follow Robins and Nielsen (1970) and Cohen and Nielsen (1978), who treated the two groups as subfamilies of Carapidae. The family Pyramodontidae was originally erected by Smith (1955) to include Pyramodon, and possibly Snyderidia. The major distinguishing characters given by him for Pyramodontidae were as follows: small parietals, separated by supra- occipital; maxilla expanded posteriorly; dorsal origin opposite anal origin and vent, and no scales. Gosline (1960) modified this definition to include Snyderidia. which necessitated the deletion of two characters: separation of the parietals by the supraoccipital, and position of the dorsal fin origin. On the basis of the emended definition, the family Pyramodontidae differed from Carapidae in the number of pectoral rays, and, although not specifically stated, the lack of a known vexillifer larval stage. Strasburg (1965) described a vexillifer larva of Snyderidia canina and, based on this finding, placed Snyderidia in the family Carapidae. Robins and Nielsen (1970) supported this action and suggested that Snyderidia and Pyramodon be placed in the subfamily Pyramodontinae within the Carapidae. The recent description of the vexillifer larva of Pyramodon ventralis (Markle and Olney, 1981) verified the existence of this unique larval stage in both genera of the Pyramodontinae and reinforced the placement of this group in the family Carapidae. The vexillifer larval stage is now known for all carapid genera (Markle and Olney, 1981), and is apparently a specialization unique to the family. Subfa- milial designation of the pyramodontines has been supported by Cohen and Niel- sen (1978) on the basis of meristic characters, and by Markle and Olney (1981) on the basis of larval characteristics.

846 WILLIAMS: SYNOPSIS OF PYRAMODONTINAE 847

Figure I. General appearance ofa Pyrarnodon ventralis female (103 mm SL) from off Isla San Felix, Chile.

METHODS

Counts and measurements and cephalic sensory pore series follow Williams and Shipp (1982) with the following exceptions: the last precaudal vertebra is the last trunk centrum without a distinct hemal spine, and the total number oflamellae in the olfactory rosette are recorded instead of pairs oflamellae.

D,o and A30 refer to the number of fin rays whose bases lie anteriad of the 31st vertebra. Cephalic sensory pores were located by directing a jet of air through the sensory canals. Cleared and stained specimens were prepared following the method of Dingerkus and Uhler (1977). Cranial sutures are extremely difficult to discern in pyramodontines and discussions of suture patterns are tentative. Osteological and meristic characters were examined from X-rays, cleared and stained specimens, and partially dissected specimens. The following abbreviations are used to denote locations of specimens studied: ANSP-Academy of Natural Sciences of Philadelphia, BMNH-British Museum (Natural History), CAS-California Academy of Sciences, UF - Florida State Museum at the University of Florida, USAIC- University of South Alabama Ichthyological Collection, and USNM-National Museum of Natural History of the Smithsonian Institution.

Subfamily Pyramodontinae Smith, 1955 Diagnosis. - Pectoral fin rays 24-30. Posterior portion of lateral temporal canal angling upward above the opercle at an angle of about 45 degrees to longitudinal axis of body, terminating near dorsum at about a vertical through pectoral fin base. Anal fin pterygiophores relatively longer than carapines and modified into a visceral cradle (Markle and Olney, 1981) consisting of alternate bending of pterygiophores left or right between at least the third to thirteenth pterygiophores. Predorsal bone elongate, with a wavy ventral surface, its midpoint at dorsal fin origin. Premaxillae each with 2 small conical teeth anterior to and mesial to the large symphyseal canines. In addition to the above, Markle and Olney (1981) reported pyramodontine larvae to be unique in having a deep, compressed head and trunk (depth at first anal ray 0.77-0.92 head length-HL), and a short predorsallength (1.12-1.2 HL). Comments. -An additional character not previously reported in pyramodontines is the presence of an unnamed bone located between the ethmoid and the dorsally projecting premaxillary processes. The rostral cartilage is covered dorsally and laterally by this bone, which appears to be the same as that described by Tyler (1970) for Onuxodon parvibrachium. Tyler (1970), who did not use a cartilage counterstain in preparing his material, called this an "X bone" and, understand- ably, did not mention the fact that the rostral cartilage, while reduced in size compared to other carapines, is present and fits into a hollow on the ventral side of the X bone in Onuxodon. This is the same formation found in pyramodontines, but in no other carapines. 848 BULLETIN OF MARINE SCIENCE. VOL. 33. NO.4. 1983

Table I. Summary of differences between pyramodontines

Snyderidia Pvramodon ·~·(!f1(ralis borhrops canina Pelvic fins present absent absent Pterygiophores forming visceral cradle 3-22 to 24* 3-13t ? Total lamellae in olfactory rosette 13 15-16 15 Rows of dentary teeth 1 1 2-3 • From Markle and Olney (1981). t Based on onc cleared and stained specimen.

Pyramodontines appear to be free-living carapids. Snyderidia bothrops has been collected from areas with large holothurians (Robins and Nielsen, 1970), but has not been reported from a host and may be free-living. Ifpyramodontines are free- living, then they share this free-living life style with the carapine Echiodon, which also inhabits relatively deep water on the shelf and/or upper slope.

Genus Pyramodon Smith and Radcliffe, 1913 Figure I

Pyramodon Smith and Radcliffe in Radcliffe (1913: 175; type species by original designation Pyr- amodon ventralis Smith and Radcliffe). Cynophidium Regan (1914: 16; type species by monotypy Cynophidium punctatum Regan, 1914). Diagnosis. -See Table I. Comments.-Smith (1955) stated that the parietal bones were separated dorsally by the supraoccipital bone. I have found it extremely difficult to discern cranial suture lines, but, in the specimen I examined, the parietals seem to meet for at least a few millimeters in front of the supraoccipital. This observation is tentative because the cranial bones are adorned with numerous ridges and other ornamen- tation that obscure the sutures. If my interpretation is correct, then this character is shared by all genera ofCarapidae (personal observation). Gosline (1960) stated that the meeting of or separation of the parietals is correlated with the size of the occipital crest. He found that the parietals meet in many flatheaded species and are separated by the supraoccipital in narrow, high headed species. A recent de- scription of the cranium of the flatheaded ophidiid Enchelybrotula supports this idea as it shows the parietals meeting in front of the supraoccipital (Cohen, 1982). Although it is likely that the character state of parietals meeting dorsally has evolved independently in several different ophidiiform lines, I believe it is infor- mative in the Carapidae when used in conjunction with other derived characters.

pyramodon ventralis Smith and Radcliffe, 1913

Pyramodon ventralis Smith and Radcliffe in Radcliffe, 1913: 175-176 (Type locality: Doworra Island; holotype USNM 74155). Cynophidium punctatum Regan, 1914: 16 (Type locality: Cape North, New Zealand; holotype BMNH 1913.12.4.37). Diagnosis. -See Table 1.. Description. - The following account is based on a mature female specimen col- lected off Isla San Felix, Chile, and is the first record of the genus from the eastern Pacific Ocean.

D3047. A3044. Pectoral fin rays 30-30. Pelvic fin rays 1-1. Caudal fin rays 8. Branchiostegal rays 7 (5 on ceratohyal, 2 on epihyal). Total nasal lamellae in right WilLIAMS: SYNOPSIS OF PYRAMODONTINAE 849

B

A c

MD LT

MAX--1

POP Figure 2. Semidiagrammatic illustration of cephalic sensory canals and pores of Pyramodon ventralis: A, left lateral view; B, dorsal view; C, ventral view oflower jaw. Lines indicate first and last pores of each series. AN, anterior nostril; 10, infraorbital series; LT, lateral series; MAX, maxillary bone; MD, mandibular series; POP, preopercular series; PN, posterior nostril; ST, supratemporal series. olfactory rosette 13. Pseudobranchiae 2-2. Gill rakers (3 rudiments) 0 + I + 2 (II rudiments). Precaudal vertebrae 15. Upper lip with 2 small fleshy tabs located above lip just anterior to anteriormost pore of supraorbital sensory pore series on each side of snout. No pyloric caecae present. The following measurements are given in mm: total length (TL) 107.8, standard length (SL) 103.0, head length (HL) 43.0, predorsal length 52.0, preanal length 58.9, preanus length 62.6, upper jaw length 23.0, snout length 7.8, eye diameter 6.7, bony orbit diameter 8.3, least bony interorbit length 8.5, pectoral fin length 42.8, pelvic fin length 13.7, caudal fin length 4.8. Peritoneum black. Swim bladder white, thick walled and located outside perito- neal lining. Stomach and intestines creamy tan colored. Cephalic sensory canals and pore positions illustrated in Figure 2. Supraorbital and infraorbital canals not connected anteriorly. In Snyderidia these canals appear to be connected. No lateral lines discernible (Smith, 1955, reported three for his South African specimen). Each premaxilla has one strong, recurved fang (left fang broken off at base), head of each premaxilla bears two small conical teeth located mesial to and in front of each fang. Slight untoothed gap behind fangs, followed by band of conical teeth extending to posterior edge of premaxilla and consisting of the following: two rows anteriorly with inner row teeth depressible and about three times size of outer, immovable, conical teeth; inner teeth decrease in size posteriorly, be- coming equal in size to other teeth at about Ih to '12 length of band; band widens to three irregular rows at midpoint, and tapers to 2 irregular rows at posterior end of premaxilla. Vomer with round patch of small conical teeth on anterior face, about three or four rows of small conical teeth forming arch across posterior face of vomer, arch begins at base of vomer, in front of palatine, and arches toward peak of vomer, then back to its base at point in front of other palatine. Peak of vomer with large fang bending to right of midline and second, smaller fang bending to left of midline from its base slightly anterior to base of other fang (smaller fang on vomer may be a replacement for one lost earlier). Palatines each with a single 850 BULLET1N OF MARINE SCIENCE, VOL. 33, NO.4. 1983

Figure 3. General appearance of Snyderidia. Based on a S. bothrops male (USNM 213505, HL 44.4 mm). row of strong conical teeth. Each dentary with two large recurved, symphyseal fangs, posteriormost fang of each pair flexible basally. Short untoothed gap sep- arates single row of stout, conical teeth from dentary fangs. This single row of dentary teeth interdigitates with palatine teeth when mouth is closed. In alcohol, the body has a grey sheen and is densely covered with small me- lanophores. Dorsal, anal, pectoral, pelvic, and caudal fins with small scattered melanophores in the skin covering these fins. Comments.-As there is some confusion regarding the number of species of Pyr- amodon, I tentatively identify the Chilean specimen as P. ventralis. A definitive identification of this specimen will have to await the completion of Douglas F. Markle's study of the relationships within this group. Distribution. -pyramodon ventralis is known to occur on the shelf or upper slope off South Africa, Kenya, Australia, New Zealand, Sumatra, Doworra Island (In- donesia), Japan, Hawaii and Chile (based on Markle and Olney, 1981; D. F. Markle, personal communication; present work).

Material examined. - Pyramodon ventralis, USNM 231353, female-l 03 mm SL, 25°40'S, 85°23'W, 185-230 m, R/V IKHTlANDR trawl 41. BMNH 1913.12.4.37, holotype of Cynophidium punctatum. New Zealand, "Terra Nova."

Genus Snyderidia Gilbert, 1905 Figure 3

Snyderidia Gilbert (1905: 654; type species by monotypy Snyderidia canina). Diagnosis. -See Table 1. Comments. -An account of the osteology of Snyderidia canina was provided by Gosline (1960). Vexillifer larvae of S. canina were described by Strasburg (1965) and by Markle and Olney (1981). Markle and Olney (1981) did not observe a visceral cradle in cleared and stained vexillifer larvae of Snyderidia canina, but in a cleared and stained adult S. bothrops this structure is clearly visible. It is possible that this cradle is found in S. bothrops and not in S. canina, but it may develop only in adult Snyderidia. This would explain its absence in S. canina larvae. Gosline (1960) did not discuss the anal WILLIAMS: SYNOPSIS OF PYRAMODONTINAE 851

B

A c

MAX

10 POP

Figure 4. Semidiagrammatic iJJustration of cephalic sensory canals and pores of Snyderidia. A, left lateral view; B, dorsal view; C, ventral view of lower jaw. Some specimens have one more pore in posterior part of LT. Abbreviations as in Figure 2. fin pterygiophore arrangement for S. canina, but the visceral cradle would be difficult to discern in a skeletonized specimen, and may have been overlooked. Cephalic sensory canals and pores are illustrated in Figure 4. Two small, fleshy tabs are present on each side of the snout above the upper lip. These tabs are present in specimens of both nominal species of Snyderidia, in Pyramodon, in Echiodon (overlooked by Williams and Shipp, 1982), and in a modified condition in a new species of sub-Antarctic pearlfish (D. F. Markle, J. T. Williams and J. E. Olney, in preparation), but in no other carapid. This structure is not unique to these pearl fishes, as I have found it in many other ophidiiform genera. I interpret the possession of this structure (except in the modified state) as a plesiomorphic character state with the modification or loss of the structure representing apomorphic (derived) conditions. The phylogenetic implications of this character will be discussed further in another paper (in preparation). There are two nominal species in Snyderia, S. canina in the Indian and Pacific Oceans, and S. bothrops in the Atlantic Ocean (Robins and Nielsen, 1970). Anal- ysis of additional specimens of both nominal species reveals a high degree of variability in the characters initially used to separate the two species. In view of the fact that I have not examined material from the Gulf of Guinea and that the two nominal species occur in different oceans, I recognize both forms as distinct. Douglas F. Markle and John E. Olney are revising this genus and related taxa.

Snyderidia bothrops Robins and Nielsen, 1970

Snyderidia bothrops Robins and Nielsen, 1970: 287 (Type locality: Gulf of Guinea; holotype USNM 201420). Diagnosis. -See Table 1. Comments. -Counts made on available material are presented in Table 2. None of the specimens had a caudal fin or the posterior vertebrae due to breakage. This condition is not uncommon in pearl fish specimens because of their long attenuate tails. Robins and Nielsen (1970) stated that the major difference betweeen Snyderidia 852 BULLETIN OF MARINE SCIENCE. VOL. 33. NO.4. 1983

Table 2. Frequency distribution of selected counts for specimens of Snyderidia examined (D30 and

A30 refer to the number of dorsal and anal rays, respectively, whose bases are anteriad of a vertical through the junction between the 30th and 31st vertebral centra)

Olfoctary Lomellae in Eaeh Precaudal Nosal Verl. D" Pectoral Roselle 25- 25- 26- 26- 27- 13 14 15 47 48 49 50 51 43 44 45 46 25 26 26 27 27 15 16 bothrops Campeche 1 2 2 I Surinam I I? I I? French Guiana 4 2 3 2 I? canina Philippines I E. of Marianas I 2 Hawaii I? 2 3 bothrops and S. canina was that of dentition; i.e., there are more and stronger teeth in S. bothrops. They gave the following as the major differences: "S. bothrops has 2 fangs anteriorly on each dentary instead of one and the remaining dentary teeth are much larger than in canina and in a single row rather than in two. Occasionally there are a few teeth posteriorly in a second row. The vomerine canine is much longer and more curved than in canina and it has teeth in front as well as behind (S. canina apparently has fewer teeth behind the vomerine canine and may lack the symphyseal teeth on the premaxilla)." The material that has been collected since the original description of S. bothrops makes it necessary to disallow all but one of these differences. In my material, specimens of S. canina usually have: two fangs anteriorly on each dentary; one or two vomerine canines of varying length and curvature, but within range ob- served for bothrops; numerous conical teeth in front of and behind vomerine canine(s), also variable but in range of bothrops; two or three small conical sym- physeal teeth on each premaxilla like those of bothrops. The only significant difference observed was in the number of rows of conical teeth behind the dentary, two or three rows for S. canina vs. one row for S. bothrops. Robins and Nielsen (1970) reported 24-25 pectoral fin rays for specimens of Snyderidia bothrops from the Gulf of Guinea, 26 for the specimen from Caribbean Panama, and 25-26 for S. canina. Specimens of S. bothrops from the Gulf of Mexico and western Atlantic have 26-27 pectoral fin rays (Table 2). This difference between eastern and western Atlantic populations would be a significant difference were it not for the 27 pectoral fin rays reported for S. canina from Hawaii (Gosline, 1960). It seems likely that this is a variable character, and that sampling bias has resulted in apparent diflerences that may not reflect the actual situation. The cephalic sensory pore system is illustrated diagrammatically in Figure 4. This diagram of the pore pattern differs from that of Robins and Nielsen (1970) in the following: lateral temporal canal (LT; their lateral canal) with 4 or 5 pores beyond its junction with preopercular cana~ (POP) (they indicated 3 pores along this portion of the canal in Gulf of Guinea specimens), and POP with 4 pores (they did not observe a pore behind the posterior edge of the maxilla). In the material examined, the anterior portions of the supraorbital (SO) and infraorbital (10) canals are continuous. Robins and Nielsen (1970) reported 15 precaudal vertebrae for their material WILLIAMS:SYNOPSISOF PYRAMODONTINAE 853 of Snyderidia bothrops, including a specimen from Caribbean Panama. My Gulf of Mexico and western Atlantic specimens have 13 or 14 precaudal vertebrae (Table 1). The significance of this difference is being investigated by Douglas F. Markle. The Gulf of Mexico and western Atlantic specimens agree (with the above exceptions) with the original description of Snyderidia bothrops, and represent a significant range expansion north to the Gulf of Mexico, as well as documenting its occurrence off French Guiana and Surinam. Osteological characters agree with Gosline's (1960) account of the osteology of Snyderidia canina. Distribution. -Snyderidia bothrops is known from shelf and upper slope regions in the Gulf of Guinea and Caribbean Panama (Robins and Nielsen, 1970), the Bahia de Campeche in the Gulf of Mexico, and off Surinam and French Guiana in the western Atlantic. With the exception of one specimen from St. Helena Island reported from 4-9 m by Robins and Nielsen (1970), adults have been collected at depths of 260-1,500 m, with most specimens collected at depths of 600-900 m. Robins and Nielsen (1970) did not comment on the specimen cap- tured at 4-9 m, but if their data are correct, I can only hypothesize that this specimen may have been collected just before abandoning its planktonic life stage to begin its adult life in the benthos.

Material Examined.-Snyderidia bothrops: USNM 213505 (4), GulfofMexico, off Mexico, 18°56'N, 94°36'W, 735 m, OREGONII Cruise 18 stat. 10961. USNM 210397 (I), western Atlantic off French Guiana, 7°41'N, 53°37'W, 735 m, OREGONII stat. i0621. USNM 214034 (i), western Atlantic off French Guiana, 7°41'N, 53°35'W, 881.5 m, OREGONII stat. 10798. USNM 214035 (I spec. cleared and stained), western Atlantic off French Guiana, r44'N, 53°59'W, 594 m, OREGONII stat. 10807. USNM 2i4037 (I), western Atlantic off French Guiana, r20'N, 53°02'W, 731.5 m, OREGONII stat. 10610. USNM 214039 (i), western Atlantic off French Guiana, 7°32'N, 53°i9'W, 658 m, OREGONII stat. i 0814. USNM 214038 (I), western Atlantic off Surinam, 7°49'N, 54°22'W, 731.5 m, OREGONII stat. 10604. USNM 211297 (I), western Atlantic off Surinam, 7°46'N, 54°06'W, 585-640 m, OREGON II stat. 10817. Snyderidia canina Gilbert, 1905

Snyderidia canina Gilbert, i905: 654 (Type iocaiity: Kaui, Hawaiian Islands; holotype USNM 51646). Diagnosis. -See Table I. Comments. -Counts made on available material are presented in Table 2. The material examined lacked caudal fins (all had been broken off), but Robins and Nielsen (1970) gave 7 caudal fin rays for the holotype. A discussion of the relationship of Snyderidia canina to S. bothrops is provided in the account of the latter. The specimens examined agree in external characteristics with the original description. All previous accounts of Snyderidia canina have given a count of 15 precaudal vertebrae. My material has a range of 13-15 precaudal vertebrae (i.e., those without a distinct hemal spine). Distribution. -Adults are known from the shelf and the upper slope at depths of 150 to 662 m from the Indian Ocean, the Philippine Islands and the Hawaiian Islands (Robins and Nielsen, 1970; this study). Pelagic vexillifer larvae are known from the equatorial Central Pacific and the Hawaiian Islands (Markle and Olney, 1981).

Material Examined.-Snyderidia canina: USNM 99230 (I), Philippine Islands, Sialat Point Light, 13°40'57"N, 123°57'45"E, ALBATROSS.USNM 210602 (3), western Tropical Pacific, 21°09'N, 157°42'W, 662 m, CROMWELLstat. TC 36-31. USNM 215467 (I), Hawaiian Islands, Oahu, 2l020'N, I 58°20'W 854 BULLETINOFMARINESCIENCE.VOL33. NO.4. 1983 to 21°30'W, 0-350 m, CROMWELLstat. TC 73-8-28. USNM 215700 (I) spec., 21°20'N, I58°20'W to 21°30'N, 158°30'W, 150 m, TOWNSENDCROMWELLstat. TC 52-52. USNM 215715 (1), Hawaiian Islands, Oahu, 12°15'N, 158°15'W to 21°30'N, 158°30'W, 0-530 m, TEUTHISCruise 6 stat. 43. Comparative Material Examined. --Brotula barbata. USAIC 4073 (I )-cleared and stained, Gulf of Mexico. Echiodon dawsoni, UF 30886 (I )-cleared and stained, Gulf of Mexico. Lepophidiumjeannae. UF 26817 (4)-1 cleared and stained, Gulf of Mexico. Onuxodon margaritiferae, CAS 48975 (3)-1 cleared and stained, Maldive Islands. Onuxodon parvibrachium. ANSP 109210 (1)-cleared and stained, Seychelles Islands. Ophidion ho/brooki. UF 948 (4)-1 cleared and stained, Florida. Otophi- dium omostigmum, USAIC 2248 (2)-cleared and stained, GulfofMexico. Parophidion schmidti, UF 25086 (8)-1 cleared and stained, Caribbean Sea, Providencia Island.

ACKNOWLEDGMENTS

The discovery of the eastern Pacific specimen of Pyramodon was made possible by a 10-week fellowship awarded by the Smithsonian Institution. I thank the curators of the fish collection at the National Museum of Natural History for providing free access to the collection, to V. G. Springer for providing work space, and to D. M. Cohen for discussing ophidiiform classification with me during my residence at the museum. I thank W. F. Smith-Vaniz for his assistance and hospitality during a visit to the Academy of Natural Sciences of Philadelphia and W. N. Eschmeyer for his assistance during a visit to the California Academy of Sciences. A. Wheeler supplied information on and radio- graphs of the holotype of Cynophidium punctatum. I. Echavarria assisted in X-raying specimens. D. M. Cohen, C. R. Robins, and D. F. Markle read the manuscript and provided helpful suggestions and comments.

LITERATURE CITED

Cohen, D. M. 1982. The deepsea fish genus Enchelybrotula (Ophidiidae): description of new species, notes on distribution and osteology. Bull. Mar. Sci. 32: 99-111. --- and J. G. Nielsen. 1978. Guide to the identification of genera of the fish order with a tentative classification of the order. NOAA Tech. Rept. NMFS Circ. 417: 1-72. Dingerkus, G. and L. D. Uhler. 1977. Enzyme clearing ofalcian blue stained whole small vertebrates for demonstration of cartilage. Stain Tech. 52: 229-232. Gilbert, C. H. 1905. The aquatic resources of the Hawaiian Islands, part II, section II. The deep- sea fishes. U.S. Fish Comm. Bull. 23: 577-713, pis. 66-101, figs. 230-276. Gosline, W. A. 1960. Hawaiian lava-flow fishes, Part IV. Snyderidia canina Gilbert, with notes on the osteology of ophidioid fishes. Pacif. Sci. 14: 373-381. Markle, D. F. and J. E. Olney. 1981. A description of the vexillifer larvae of Pyramodon ventralis and Snyderidia canina (Pisces, Carapidae) with comments on classification. Pacif. Sci. 34: 173- 180. Radcliffe, L. 1913. Descriptions of seven new genera and thirty-one new species of fishes of the families Brotulidae and Carapidae from the Philippine Islands and the Dutch East Indies. Proc. U.S. Nat. Mus. 44: 135-176. Regan, C. T. 1914. Diagnoses of new marine fishes collected by the British Antarctic ('Terra Nova') Expedition. Ann. Mag. Nat. Hist., Ser. 8,13: 11-17. Robins, C. R. and J. G. Nielsen. 1970. Snyderidia bothrops, a new tropical, amphi-Atlantic species (Pisces: Carapidae). Stud. in Trop. Oceanogr. 4: 285-293. Smith, J. L. B. 1955. The genus Pyramodon Smith & Radcliffe, 1913. Ann. Mag. Nat. Hist., Ser. 12,8: 545-550. Strasburg, D. W. 1965. Description of the larval and familial relationships of the fish Snyderidia canina. Copeia 1965: 20-24. Trott, L. B. 1981. A general review of the pearl fishes (Pisces, Carapidae). Bull. Mar. Sci. 31: 623- 629. Tyler, 1. C. 1970. A redescription of the inquiline carapid fish Onuxodon parvibrachium. with a discussion of the skull structure and the host. Bull. Mar. Sci. 20: 148-164. Williams, J. T. and R. L. Shipp. 1982. A new species of the genus Echiodon (Pisces: Carapidae) from the eastern Gulf of Mexico. Copeia 1982: 845-85 I.

DATEACCEPTED: September 13, 1982.

ADDRESS: Florida State Museum and Department oj Zoology. University oj Florida. Gainesville. Florida 32611. PRESENTADDRESS:Division oj Fishes, National Museum of Natural History, Smith- sonian Institution. Washington, D.C. 20560.