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2003


A preliminary survey of the dorsal gill arches of flatfishes (Pleuronectiformes) and an examination of potential phylogenetic consequences

Amanda Cass
Research Training Program, 2003



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A preliminary survey of the dorsal gill arches of flatfishes (Pleuronectiformes) and an examination of potential phylogenetic consequences.


ABSTRACT

The flatfishes (Pleuronectiformes) are a diverse and widespread order of fishes united by their ocular asymmetry. Due to their reduced skeletal morphology, much of their classification is based on loss and fusion characters, especially those of the caudal skeleton. Though these characters are used in other classifications and may be phylogenetically informative, no studies of pleuronectiform ontogeny have been conducted to determine homologies of the remaining elements, thus making them a weak tool for phylogenetic analysis. Another character system used to determine phylogenies is the gill arches, specifically the dorsal complex. To date, no systematic study of the Pleuronectiformes has included a detailed analysis of gill arch variation. The aim of the current study was to document variation within this structure across taxa and determine whether it warranted further study. Representatives from each of 10 families were prepared for dissection via clearing and counter-staining. The gill arches were removed from these specimens, observed and documented. Pleuronectiform gill arches were found to vary widely, with several characters delineating previously identified taxa and others drawing connections between groups thought to be only distantly related. Further studies including more taxa are planned and may elucidate the underlying patterns present in this important character system. It is hoped that these studies will shed some light on the obscure interrelationships of these fishes.
 
The Pleuronectiformes

The flatfishes (Pleuronectiformes) are a diverse order of fishes, comprising over 570 species in 11 families. For many years they were united by only one shared derived character, their unique metamorphosis from pelagic symmetrical larvae to benthic asymmetrical adults. During this metamorphosis many changes occur, the most dramatic of which is the migration of one eye over the middorsal line. As Pleuronectiformes are popular food fishes (including flounder, sole and halibut) and live worldwide, they are of great importance to many economies. Due to their atypical development and adult morphology, they are also of particular interest to science. Yet, much about them is still unknown. Their unusual asymmetry has minimized challenges to their monophyly, but both super- and sub-ordinal relationships are difficult to determine and have been revised many times. Some difficulty in determining their relationships arises from skeletal reduction—many bones, distinct in other fishes, are absent or fused. Furthermore, their tendency to lose rather than gain characters means that more often one finds shared losses, rather than shared structures. Few synapomorphies have been identified that unite families or groups of genera. To date, the caudal skeleton has provided most of the characters that define subgroups.


Dorsal Gill Arch Characters

Posterior to the mouth and ventral to the braincase lies a complex structure with a framework of bones and cartilage called the gill arches. This apparatus serves two important functions; it supports the gill filaments used in respiration and assists with the ingestion and mastication of food. This structure is composed of a medial and ventral series of bones, basibranchials, with bilaterally paired elements extending vertically. These paired elements are composed of many smaller bones, each described as lying either in the dorsal (pharyngobranchials and epibranchials) or ventral (ceratobranchials and hypobranchials) portion of the arch. Although the ventral complex is phylogenetically informative, more characters are usually found in the arrangements of tiny bones and cartilages in the dorsal gill arches, which vary considerably between groups. To date, pleuronectid gill arch characters have not been studied closely.


Caudal Skeleton Characters

The caudal (tail) fin of fishes is composed of many fan-like bones associated with the terminal centrum. Bones that form above the notochord of a developing embryo are referred to as epurals (EP) and those that develop below are referred to as hypurals (HP). Other, smaller bones referred to as uroneurals (UN) may be present in the dorsal portion of the fin. Primitively, fish have many bones in the tail, but these may be lost or fused. Similarities and differences in loss and fusion patterns are often used in systematic studies, as there are many possible patterns of fusion and shared patterns can indicate relationships. However, adult fins that appear similar do not necessarily result from the same developmental pathways and thus may not be homologous. There is no way to determine definitively or refute homology, but careful study of larval series can identify ontogenetic fusions. Currently a great deal of pleuronectiform classification rests on caudal skeleton characters; however, ontogeny has been studied in few taxa. Though a great deal of change has occurred in these fishes from a caudal skeleton like that of Psettodes (long presumed to be the basal taxon) to the forms below, because the changes are reductions, shared ontogenetic trajectories would give us more confidence when using similar adult endpoints as characters.


ACKNOWLEDGMENTS

The National Science Foundation Research Experiences for Undergraduates program for funding this research (Award: #DBI-0243512 ). The Smithsonian Museum of Natural History Research Training Program for providing the circumstances for it to occur. Dr Lynne Parenti and Dr G. David Johnson for their advice and guidance. Sandra Reardon for her help with digital photography and x-ray.


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The information presented here represents preliminary research as the result of ten-weeks of investigation in-residence at the National Museum of Natural History. This is not an official publication of the information.

As preliminary information, results and/or findings should not be cited as part of conclusive work. Please contact the authors first if you wish to utilize the information presented here.

 

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