Leptocephalus Larvae of the Tribe Sphagebranchini (Pisces, Ophichthidae) in the Western North Atlantic
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BULLETIN OF MARINE SCIENCE, 32(1): 220-236, 1982 LEPTOCEPHALUS LARVAE OF THE TRIBE SPHAGEBRANCHINI (PISCES, OPHICHTHIDAE) IN THE WESTERN NORTH ATLANTIC Mark M. Leiby ABSTRACT Larvae of Apterichtus ansp, A, kendalli, Ichthyapus ophioneus, a problematical specimen tentatively identified as Microrhynchus (sensu Blache and Bauchot, 1972) sp. and Sticto- rhinus potamius are described. A. ansp, A. kendalli, I. ophioneus and Microrhynchus sp. are characterized by having the dorsal fin confined to the tail-tip and by reduction in gill arch development. S. potamius is characterized by its highly developed pigmentation and by its dorsal fin origin above the anus. An explanation for the difficulties encountered in deter- mining dorsal fin origin in ophichthid larvae is given. The ophichthid tribe Sphagebranchini (sensu McCosker, 1977) comprises ]2 nominal species in the Atlantic and Mediterranean. Blache (1977) described five series of larvae from the eastern Atlantic which he identified as Microrhynchus sp. aff. foresti, Apterichtus caecus, Verma kendaLli, V. monodi and Verma sp. Fahay and Obenchain (1978) identified two forms of larvae from the western Atlantic as Apterichtus (=Verma) kendalli and A. ansp. Recently, Leiby (in press) questioned these authors' identifications and descriptions of Verma mon- odi, Verma sp., Apterichtus (=Verma) kendalli, and A. ansp. This paper provides definitive larval descriptions of A. ansp, A. kendalli, Ichth- yapus ophioneus, a problematical specimen tentatively identified as Microrhyn- chus (sensu Blache and Bauchot, 1972) sp., and Stictorhinus potamius. It also provides an explanation for the difficulties occasionally encountered in determin- ing dorsal-fin origin in ophichthid larvae. METHODS Measurements were made to the nearest 0.1 mm using an optical micrometer in a dissecting mi- croscope. All counts, measurements, and growth stages are as defined in Leiby (1979a, b). At least one larva of each species was stained for bone and cartilage following Dingerkus and Uhler (1977) and dissected for osteological study. Drawings were made using a camera lucida on a dissecting microscope. Data on adult A. ansp, A. kendalli, and I. ophioneus were obtained from cleared and stained specimens, from the literature and from x-rays provided by Dr. James E. Bohlke, Academy of Natural Sciences, Philadelphia. Data on Microrhynchus foresti were obtained from the literature. Data on S. potamius were obtained from the literature and from x-rays provided by Eugenia B. Bohlke. Collection locations for the species examined are shown in Figure I. Institutional abbreviations used in this study are: Harvard University, Museum of Comparative Zoology, Cambridge, Massachusetts (MCZ); Marine Biomedical Institute, Galveston, Texas (MBI); Florida Department of Natural Resources, Marine Research Laboratory, St. Petersburg, Florida (FSBC); National Marine Fisheries Service, Northeast Fisheries Center, Highlands, New Jersey (NMFS-NFC); South Carolina Wildlife and Marine Resources Department, Charleston, South Car- olina (SCMRD); Rosenstiel School of Marine and Atmospheric Science, Miami, Florida (UMML); University of Rhode Island, Kingston, Rhode Island (URI). Apterichtus ansp (Bohlke) Figures 2, 4, 5; Table 3 ldentification.-An advanced metamorphic larva in a series of 28 specimens had two temporal pores (TP ]-2) and four preoperculomandibular pores (POP ]-4) 220 LEIBY: LARVAL SPHAGEBRANCHINI 221 •• :Apterichtus ansp o :Apterichtus kendalli 0: Ichlhyapus ophioneus .: ?Mlcrorhynchus sp. • : Stictorhinus potamlU5 C>-. 0 ,~, o •• o Figure I. Distribution of larval Microrhynchus sp., Stictorhinus potamius, Apterichtus ansp, A. kendalli and Ichthyapus ophioneus. in the cephalic lateralis system. This condition is found only in the tribe Sphag- ebranchini. The specimen was cleared and stained; the hydroid arch had ]6 bran- chiostegals; the gill arches (Fig. 2A) lacked basibranchials two and three (B 2-3), had basibranchial four (B 4) so weakly developed that it would be reduced or lost by the time metamorphosis was complete, and had a small ceratobranchial five (C 5) that would be reduced or absent in adults. Apterichtus is the only Atlantic sphagebranchin genus known to lack B 2-3, to have B 4 and C 5 either reduced or absent, and to have so few branchiostega]s. Comparison of total and nephric myomere counts of the larvae with total and precaudal counts of the six nomina] adult Apterichtus species in the Atlantic eliminated all but A. ansp (a western Atlantic species) and A. gracilis (an eastern Atlantic species). The presence of small (9.9 mm TL) larvae in the South Atlantic Bight indicates a recent spawning in the western North Atlantic. Consequently, the series of larvae can only be A. ansp. 1 mm 1m'" Figure 2. (Left) A, Gill arches of Apterichrus ansp from a 57.0-mm TL euryodontic specimen; B, Caudal skeleton of same specimen. Figure 3. (Right) A, Gill arches of Aptericlzrus kendalli from a 76.9-mm TL early metamorphic specimen; B, Caudal skeleton of same specimen. 222 BULLETIN OF MARINE SCIENCE, VOL. 32, NO. 1.1982 Figure 4. Apterichtus ansp, engyodontic stage. A, Leptocephalus 9.9 mm TL; B, Head of same specimen. General Morphology.-Twenty-eight larvae examined, 9.9-70.2 mm total length (TL). Body relatively elongate, much compressed, clear. Dorsal fin confined to tail-tip until late metamorphosis when all fins resorbed. Total myomeres 129-136; nephric myomeres 63-67; preanal myomeres 65-70, anus migrates anteriorly dur- ing metamorphosis; predorsal myomeres 114-125 in late euryodontic and early metamorphic specimens; predorsal myomeres cannot be determined in engyo- dontic or early euryodontic larvae. Branchiostegals 15-16. Generally nine much- reduced gut loops or swellings, often differentiable from preservation artifacts only by chromatophore patch beneath nephric duct on swelling (Table I); two usually distinct loops along esophagus, first at anterior end of liver, second at posterior end of liver; gut expands posterior to gall bladder and liver. Nephros terminates one to three myomeres anterior to anus. Relative preanal length (PAL) and head length (HL) decrease with growth of specimen due to differential growth in tail region (Table 2). Preanal myomere number remains constant. Pigmentation.-Pigment in formalin-preserved specimens inconspicuous (Figs. 4,5). Weak pigment on anterior half of upper and lower jaws near base of teeth; on lower jaw lateral to juncture of quadrate and Meckel's cartilage; on cheek lateral to hyomandibula. A variable number of chromatophores in body wall lateral to heart. One or more small chromatophores at midline on each myoseptum starting as far forward as myomere six, frequently uniting to form streak of pig- ment on myosepta. Weak pigment on dorsal surface of esophagus near pectoral- fin base. Five to nine irregular pigment patches along dorsal surface of gut, first two mark anterior and posterior lobes of liver, remaining three to seven patches mark gut swellings; number of pigment patches on gut increases from hatching to euryodontic stage. Small, scattered patches of pigment in body wall ventral to gut. Three to five faint, irregular subcutaneous pigment patches on tail just ventral to aorta near postanal myomeres 8-]2, 17-23, 25-35, 38-46, 49-57, number of patches increase from hatching, generally becoming five by early euryodontic stage. A single, weak chromatophore patch on ventral surface of each anal-fin pterygiophore. Material Examined.-Uncatalogued specimens from: NMFS-NFC (13 specimens, 20.2-70.7 mm TL); SCMRD (12, 9.9-29.5); UMML (2, 51.1-57.0); URI (I, 70.2). Engyodontic specimens collected in South Atlantic Bight; metamorphic specimens taken otT Grand Bahama Isle, and otT South Carolina. LEIBY: LARVAL SPHAGEBRANCHINI 223 Table I. Location of gut loops relative to myomere number in four species of sphagebranchin larvae. Values represent maximum range of occurrence observed for each gut loop. Loops are generally two to five myomeres long. (I = frequently low or absent; 2 = occasionally low but always present; 3 = occasionally low or absent) Gut Loops Species #1 #2 #3 #4 #5 #6 #7 #8 #9 A. ansp 10-16 18-23 24-28' 29-351 36-41' 41-46' 50-54' 56-62' 63-701 A. kenda/li 10-]4 18-25 27-30' 31-37 38-42t 44-5()2 52-583 60-66 67-74 I. ophioneus 9-12 17-21 26-30 35-41 47-54 S. potamius 10-13 18-23 26-29 33-38 42-45 49-53 57-62 67-73 71-76 Apterichtus kendalli (Gilbert) Figures 3, 6, 7; Table 3 ldentification.-Late euryodontic and early metamorphic larvae in a series of 24 specimens clearly had the dorsal fin confined to the tail-tip, indicating that the larvae were finless sphagebranchins. The gill arches (Fig. 3A) of a cleared and stained specimen lacked B 2-3 and had B 4 and C 5 so weakly developed that they would be reduced or absent in adults. The hyoid arch had 15branchiostegals. This combination of characters indicated that the series belonged to Apterichtus. The cephalic lateralis system was not fully developed in any of my specimens. Nevertheless, there were clearly four supraorbital pores, SO 1-4 (not including the ethmoidal pore or the median pore in the frontal commissure). The number and condition of the ossicles showed that TP 2 was absent, that POP 1-3 were present and that POP 4 was probably absent. Comparison of the cephalic lateralis system, total myomeres, and nephric myomeres of the larvae with the cephalic lateralis system, total vertebrae, and precaudal vertebrae of the known adult Atlantic Apterichtus spp. eliminated all but A. kendalli. General Morphology.-Twenty-four larvae examined, 5.8-90.3 mm TL. Body relatively elongate, much compressed, clear. Dorsal fin confined to tail-tip until late metamorphosis when all fins resorbed. Total myomeres 137-148; nephric myomeres 67-73; preanal myomeres 69-74, anus migrates anteriorly during meta- Table 2. Body proportions in five species of sphagebranchin larvae (TL = total length; PAL = preanal length; HL = head length) Developmental Size Range No.