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Pisces: Carapidae) BULLETIN OF MARINE SCIENCE. 33(4): 846-854,1983 SYNOPSIS OF THE PEARLFISH SUBFAMILY PYRAMODONTINAE (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 genus 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
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