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Osteology and Relationships of the Lanternfishes (Family Myctophidae) X I 03 on Icht Ny in I Notolychn I Ni Gym Nosc Omani

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Osteology and Relationships of the Lanternfishes (Family Myctophidae) X I 03 on Icht Ny in I Notolychn I Ni Gym Nosc Omani OSTEOLOGY AND RELATIONSHIPS OF THE LANTERNFISHES (FAMILY MYCTOPHIDAE) X I 03 ON ICHT NY IN I NOTOLYCHN I NI GYM NOSC OMANI 1 ./' WIC TOPHIN I DIAPHINI MYCTOPHINAE LAMPANYCTINAE Tribal relationships within the family Myctophidae, illustrating photophore patterns of repre- sentative species. OSTEOLOGY AND RELATIONSHIPS OF THE LANTERNFISHES (FAMILY MYCTOPHIDAE) By JOHN R. PAXTON BULLETIN OF THE NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY SCIENCE: NUMBER 13 JUNE 28, 1972 PROFESSIONAL PUBLICATIONS OF THE NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY The professional publications of the Natural History Museum of Los Angeles County include two series, Contributions and Bulletins. In the past, articles, monographs and cata- logs in the fields of history and science have appeared under various headings—Contribu- tions, Science Series, History Leaflet Series and unnumbered catalogs of exhibitions and collections. To simplify and to standardize matters, all professional publications of the History and Science Division of the Museum will now be issued at irregular intervals either as Contributions, or as Bulletins. The former will contain short, technical papers which may be occasionally gathered in volumes, octavo in size. The latter will contain longer, separate monographs and catalogs, usually quarto in size, although this will depend on the needs of the presentation. Papers in each series are to be numbered consecutively. These papers are original articles and studies based on the collections and work of the Museum, presenting newly acquired information and understanding in the fields of Anthro- pology, Botany, Geology, History, Mineralogy, Paleontology, Technology and Zoology. GILES W. MEAD, Director Natural History Museum of Los Angeles County VIRGINIA D. MILLER Editor All communications concerning science manuscripts, exchange of science publications, and the purchase of science publications should be sent to the Editor, Natural History Museum of Los Angeles County, 900 Exposition Boulevard, Los Angeles, California 90007. TABLE OF CONTENTS INTRODUCTION 1 Historical Review 3 Acknowledgments 4 MATERIALS AND METHODS 5 OSTEOLOGY 9 Superficial Dermal Bones 9 Mandibular Arch .................................................................................................................. 12 Palatine Arch ....................................................................................................................... 17 Hyoid Arch ............................................................................................................................. 19 Opercular Apparatus 23 Branchi al Arches .................................................................................................................... 24 Pectoral Girdle 28 Pelvic Girdle 31 Axial Skeleton ........................................................................................................................ 32 Caudal Skeleton 32 LUMINESCENT ORGANS ........................................................................................................................................... 37 Luminous Tissue ................................................................................................................... 37 Photophores ............................................................................................................................ 38 THE EVOLUTIONARY LINEAGES ........................................................................................................................... 40 Key to the Supraspecific Taxa .............................................................................................. 51 EVOLUTIONARY RELATIONSHIPS ........................................................................................................................... 54 Familial Affinities ................................................................................................................. 54 Subfamilial Relationships .................................................................................................... 56 Tribal Relationships 58 Generic Relationships .......................................................................................................... 63 EVOLUTIONARY SIGNIFICANCE OF PHOTOPHORE PATTERNS ............................................................... 73 LITERATURE CITED .................................................................................................................................................... 78 ••■ OSTEOLOGY AND RELATIONSHIPS OF THE 1 LANTERNFISHES (FAMILY MYCTOPHIDAE) By JOHN R. PAXTON2 ABSTRACT: Analyses of the osteological features, except the neurocra- nium and axial skeleton, and the luminescent organs of representatives of all currently recognized genera in the family Myctophidae revealed a number of evolutionary trends and groups within the family. The subfamilies Mycto- phinae and Lampanyctinae differ in the number of extrascapular elements, the presence or absence of a shelf on the cleithrum, the number and position of the Pic photophores, and the shape of the larval eyes. Two tribes, the Myc- tophini and the Gonichthyini, within the Myctophinae differ in the shape of the antorbital bone, the length of the jaw, the fusion of the procurrent caudal rays, and the position of the mouth and PLO photophore. The Myctophini includes the genera Protomyctophum, Electrona, Diogenichthys, Benthosema, Hygophum, Myctophum, and Symbolophorus; the Gonichthyini includes Loweina, Tarletonbeania, Gonichthys, and Centrobranchus. Four tribes, the Notolychnini, Lampanyctini, Diaphini, and Gymnoscopelini, comprise the subfamily Lampanyctinae. The monotypic Notolychnini displays some char- acters of each subfamily. The remaining three tribes differ in the number, shape, or position of the procurrent caudal rays, distal pectoral radials, supra- maxillary, dentary teeth, and Dn, PO, and PVO photophores. The Lam- panyctini includes the genera Lam padena, Taaningichthys, Bolinichthys, Lepidophanes, Ceratoscopelus, Stenobrachius, Lampanyctus, and Triphoturus. Diaphus and Lobianchia comprise the Diaphini and Lampanyctodes, Gymno- scopelus, Notoscopelus, Scopelopsis, Lampichthys, and Hintonia the Gymnoscopelini. Osteological evidence indicates that myctophids are most closely related to and probably derived from the family Neoscopelidae. A supramaxillary is present in some species, as is a foramen in the ceratohyal, features not pre- viously reported for the family. Osteological trends within the family include streamlining of the body in the Gonichthyini, lengthening of the jaws, sculp- turing of the circumorbitals, and fusion of the extrascapulars in the Lam- panyctinae, increase in the number of procurrent caudal rays and develop- ment of a supramaxillary in the Gymnoscopelini, and a fusion of these rays in a number of lines. Changes in photophore pattern and development of secondary luminous structures closely parallel the osteological trends apparent in the various lineages and the phylogeny presented is basically similar to those previously proposed on the basis of photophore patterns alone. INTRODUCTION Myctophids undertake extensive vertical migra- tions; in one study they were found to range diur- Lantemfishes of the family Myctophidae are nally from 400 to 750 meters and nocturnally to world-wide in distribution and are found in the the upper 100 meters (Paxton, 1967a). Many midwater regions of all major oceans. Myctophids species are taken at the surface under a night-light, are the most speciose of midwater fish families with more than 300 nominal species and probably about 200 valid species. In a long-term trawling 'EDITORIAL COMMITTEE FOR THIS BULLETIN Robert J. Lavenberg project off Bermuda, these representatives of the H. Goeffrey Moser order Myctophiformes (Iniomi, Scopeliformes) Basil G. Nafpaktitis were second only to the bristlemouth Cyclot hone of Camm C. Swift the family Gonostomatidae in the number of speci- 'The Australian Museum, Sydney, New South Wales, mens captured (Beebe, 1937) ; in many areas, Australia; and Research Associate in Ichthyology, lantemfishes make up the largest fish biomass of a Natural History Museum of Los Angeles County, Los midwater trawl catch. Angeles, Calif. 90007. 1 2 BULLETIN OF THE NATURAL HISTORY MUSEUM OF LOS ANGELES COUNTY No.13 while a very few forms apparently undergo no phology and position of body photophores do not vertical migration. Studies on the migration pat- suggest their adaptive significance, and several terns and swimbladder structure have indicated that hypotheses have been advanced to explain the lanternfishes are important contributors to the relatively common occurrence of luminous organs sonic scattering layers; bathyscaphe observations or tissue on the bodies of deep sea fishes. Harvey have confirmed that their migrations correspond (1952) and McAllister (1967) have made the to the scattering layer migrations (Barham, 1966; most extensive, recent reviews on the subject. Backus et al., 1968). Recent captures indicate None of the proposed hypotheses have been some species are associated with the bottom and tested experimentally in midwater fishes, because not restricted to the midwaters. none can presently be maintained alive for any Myctophids are utilized as a food source by length of time. In the shallow water midshipman many larger organisms, including a number of Porichthys, photophore display during courtship commercially important fishes. Lanternfishes them- has been reported (Crane, 1965). Most species of
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