Munroe: Systematics of western Atlantic Symphurus 
157 
piger, S. billy krietei, and S. stigmosus, all of which 
share the 1-3-2 ID pattern. Symphurus marginatus 
is the largest western Atlantic species possessing the 
1-3-2 ID pattern (ca. 146 mm, matures at ca. 79-90 
mm). The only western Atlantic species character- 
ized by a 1-3-3 ID pattern, S. trewavasae, is also in- 
cluded in this size range, as is S. kyaropterygium, 
the largest member of the 1-4-2 species group. Both 
species occur at similar depths on the inner conti- 
nental shelf off Brazil. In this size range are S. 
plagusia and S. caribbeanus, the smallest of five At- 
lantic species in the S. plagusia species complex 
(Munroe, 1991), featuring a 1-4-3 ID pattern. Un- 
like other tonguefishes in this size category, these 
two inhabit shallow estuaries and nearshore coastal 
habitats throughout the tropical western Atlantic. 
All large-size western Atlantic tonguefishes pos- 
sess the same ID pattern ( 1-4-3). Interestingly, these 
species, together with S. chabanaudi and S. 
elongatus, eastern Pacific species that have a 1-5-3 
ID pattern, are the largest species of symphurine 
tonguefishes. These are the predominant tongue- 
fishes collected in abundance in shallow-water envi- 
ronments located in both north and south warm-tem- 
perate and tropical regions throughout the western 
Atlantic and eastern Pacific oceans, the only areas 
worldwide where such diversity and abundance of 
shallow-water species of Symphurus is found. Of spe- 
cial note is the fact that only in the New World do 
large-size Symphurus occur, and this interestingly 
is the only tropical, shallow-water, region of conti- 
nental seas where large-size tonguefishes of other 
cynoglossid genera (Cynoglossus and Paraplagusia) 
are absent. 
More complete analysis of geographical distribu- 
tions and patterns of historical ecology for western 
Atlantic tonguefishes must await further resolution 
of monophyletic groups within the genus. Such reso- 
lution and testing of hypotheses put forth in this work 
can only be made with a more detailed osteological 
study of the entire genus and suitable outgroups, an 
analysis beyond the scope of this investigation. 
Acknowledgments 
Initially representing a portion of a dissertation sub- 
mitted as partial requirement for the Ph.D. degree 
at the College of William and Mary, this work was 
enhanced during my tenure as a postdoctoral re- 
search fellow in the Division of Fishes, National 
Museum of Natural History, Smithsonian Institution. 
Final data collection and drafting of the manuscript 
occurred after my arrival at the National Systemat- 
ics Laboratory. This study would not have been pos- 
sible without assistance provided by many people 
whose generosity is greatly appreciated. The project 
benefited from efforts, criticism, support, and encour- 
agement of a large number of people including mem- 
bers of my dissertation committee, especially J. 
Musick, J. Merriner, B. Collette, and M. Weinstein. 
This study required and benefited from financial 
support arranged by J. Merriner, J. Musick, H. Aus- 
tin, G. Grant, W. Hargis, and F. Perkins of the De- 
partment and Division of Fisheries and Director’s 
Office at the Virginia Institute of Marine Science 
(VIMS). Additional financial support was provided 
by Sigma Xi, the Raney Fund of the American Soci- 
ety of Ichthyologists and Herpetologists, William and 
Mary Minor Research Grants, and Student Confer- 
ence Fund of William and Mary College. Fellow stu- 
dents and colleagues at VIMS who provided relevant 
criticisms and suggestions included G. Sedberry, M. 
E. Anderson, M. Armstrong, R. Rosa, B. Raschi, J. 
Govoni, R. Crabtree, J. Carter, J. Desfosse, T. 
Sminkey, J. Colvocoreses, D. Hata, and D. Wyanski. 
Special thanks to K. Sulak and R. Vari for their en- 
couragement, support, and insights; this study ben- 
efited substantially through their advice and con- 
structive criticism. Donna Munroe provided logisti- 
cal and moral support and assisted with innumer- 
able different tasks during the dissertation portion 
of the study. Thanks to I. B. Ratnose for maintenance 
of a proper perspective on reality through countless 
hours of counting fin rays. Statistical advice was of- 
fered by J. Loesch, J. Desfosse, and M. Nizinski 
(VIMS). A special thanks to C. R. Robins who made 
the extensive collections of tonguefishes from UMML 
available to me, which permitted me to greatly ex- 
pand the scope of this study. The following people 
(presently or formerly associated with respective in- 
stitutions following their names) assisted with loan 
of specimens, supplied specimens and information, 
or use of facilities: L. Munroe, H. Boschung and D. 
Nieland (ALA); G. Nelson and B. Brown ( AMNH); M. 
McGrouther (AMS); W. Smith -Vaniz and W. Saul 
(ANSP); D. Markle, L. Van Guelpen and K. Sulak 
(ARC); A. Wheeler, M. Holloway, B. Brewster, N. 
Merrett, R Campbell, J. Chambers and O. Crimmen 
(BMNH); W. Eschmeyer, M. E. Anderson, D. Catania 
(CAS); J. Gourley, Corpus Christi State University, 
TX; S. Michels, Delaware Division of Fish and Wild- 
life; J. Darovec, M. Leiby, and J. Gartner (FDNR); C. 
Spurrier, M. Rogers, R. Johnson, and D. Stewart 
(FMNH); C. E. Dawson, S. Poss and M. O’Connell 
(GCRL); W. Anderson Jr., A. Harold and J. McKinney 
(GMBL); M. T. Gaspar-Dillanes and H. Espinosa- 
Perez (IBUNAM); T. Farr, Institute of Jamaica; A. 
Acero (INVEMAR), Colombia; R. Claro and J. P. 
Arteaga, Instituto de Oceanologia, Havana; R. 
