BULLETIN OF THE FLORIDA STATE MUSEUM BIOLOGICAL SCIENCES Volume 4 Number 7 BOARFISHES OF THE GENUS ANTIGONIA OF THE WESTERN ATLANTIC Frederick H. Berry \·~CESI5'·/ UNIVERSITY OF FLORIDA Gainesville 1959 The numbers of THE BULLETIN OF THE FLORIDA STATE MUSEUM, BIOLOGICAL SCIENCES, will be published at irregular intervals. Volumes will contain about 800 pages and will not necessarily be completed in any one calendar year. OLIVER L, AUSTIN, JR., Editor The publication of this number of THE BUL- LETIN has been made possible by a grant from the Graduate School, University of Florida. All communications conceTning purchase or exchange of the publication should be addressed to the Curator of Biological Sciences, Florida State Museum, Seagle Building, Gainesville, Florida. Manuscripts should be sent to the Editor of the BULLETIN, Flint Hall, University of Florida, Gaine5ville, Florida. Published 20 May 1959 Price for this issue $.65 BOARFISHES OF THE GENUS ANTIGONIA OF THE WESTERN ATLANTIC 1 FREDERICK H. BERRY 2 SYNOPSIS: The recently named boarfish Antigonia combatia Berry and R~th- jen from the western Atiantic and the deeper-bodied Antigonia capros Lowe from the western Atlantic and the Indo-Pacific are described, illustrated, and compared with other nominal forms from the Indo-Patific. Appreciable individual and geographic variation characterizes Antigonia combatia and apparently also exists in Antigoma capros. The boarfishes range in the western Atlantic from off south- ern Brazil to off the northern Atlantic coast of the United States, in water ap- proximately 87 to 325 fathoms deep. INTRODUCHON The data developed on the biology and taxonomy of the genus Antigonia durijig an analysis of the relationships of Antigonia com- batia Berry and Rathjen (1959) comprise the basis of this studj>. Specimens of Antigonia previously were rare in most museum col- lections, but the recent work in the western North Atlantic by the U. S. Fish and Wildlife SerViCe VeSSelS OREGON, COMBAT, SILVER BA¥, and THEODORE N. GrnL has produced series of specimens that add appreciably to the knowledge of this group. The boarfish genus Antigonia is known from tropical and subtropi- cal waters of the Gulf of Mexico and the Caribbean Sea, the western and eastern Atlantic Ocean, the Indian Ocean, and the western Pacific eastward to Hawaii. It has not been reported from the east- ern Pacific off the American continents. Specimens are usually taken in waters of about 87 to 325 fathoms, but the larvae ap,parently are near-surface pelagic forms. Of the 13 nominal species that have been ascribed to this genus or its synonyms, 5 appear to be valid species: Antigonia capros Lowe, Hypsinotus rubescens Giinther, Anti- gonia eos Gilbert, Antigonia malayanus Weber, and Antigonia com- batia. Four others are synonyms, Caprophonus aurora Muller and Troschel (= A. ca-pros), Antigonia steindachneri Jordan,and Fowler (= A. capros), Antigonia fowleri Franz (= A. rubescens), and Anti- i Contribution No. 27 of the U. S. Fish and Wildlife Service Bureau of Com- mercial Fisheries Biological Laboratory, Brunswick, Georgia. ' The author is a Fishery Research Biologist of the U. S. Fish and Wildlife Service at the South Atlantic Fishery Investigations (Bureau of Commercial Fish- eries Biological Laboratory), Brunswick, Georgia. Manuscript submitted 14 July 1958. 206 BULLETIN FLORIDA STATE MUSEUM Vol. 4 gonia browni Fowler (= A. capros). Of the four remaining, the re- lationships of Antigonia rubicunda Ogilby and Antigonia rhomboidea McCulloch are uncertain, Hypsinotus benhatatate Bleeker is a nomen nudum, and Antigonia miilleri Klunzinger is of the family Zeidae. Two species of boarBsh occur in the western Atlantic: the deep- bodied Antigonia capros, with body depth generally greater than standard length, known also from eastern Atiantic and Indo-Pacific waters; and the recently described Antigonia combatia, a shallower- bodied species known only from the western Atlantic. ACKNOWLEDGMENTS I am indebted for suggestions concerning the manuscript to Wil- liam W. Anderson, Harvey R. Bullis, Jr., David K. Caldwell, Daniel M. Cohen, Hugh M. Fields, Jack W. Gehringer, and Donald Moore, all of the U. S. Fish and Wildlife Service, and to John C. Briggs, of the University of British Columbia. For making specimens and data available I am grateful to James E. B6hlke, Academy of Natural Sci- ences of Philadelphia, Edward H. Chandler, Lewis Crab Co., Bruns- widk, Georgia, Daniel M. Cohen, University of Florida, Herbert W. Graham and Theodore Gallagher, North Atlantic Fishery Investiga- tions, U. S. Fish and Wildlife Service, Gordon Gunter, Gulf Coast Research Laboratory, G. Robert Lunz, Bears Bluff Laboratories, G. Palmer, British Museum (Natural Hist6ry), Edward C. Raney, Cornell University, William C. Schroeder, Woods Hole Oceanographic Insti- tution and Museum of Comparative Zoology, Leonard P. Schultz, U. S. National Museum, Royal D. Suttkus, Tulane University, and Loren P. Woods, Chicago Natural History Museum; and for the use of X-ray facilities to Bert H. Maldne, Bruns#vick, Georgia. COLLECTION ABBREVIATIONS ANSP-Academy of Natural Sciences of Philadelphia BBL-Bears Bluff Laboratories BMNH-British Museum (Natural History) CNHM-Chicago Natural History Museum CU-Cornell University GCRL-Gulf Coast Research Laboratory MCZ-Museum of Comparative Zoology MMF-Museu Municipal do Funchal, Madeira MNRJ-Museu Nacional (Rio de Janeiro) NAFI-North Atlantic Fishery Investigations (U. S. Fish and Wildlife Service) 1959 BERRY: BOARFISHES OF THE GENUS ANTIGONIA 207 SAFI-South Atlantic Fishery Invesfigations (U. S. Fish and Wildlife Service) SU-Stanford Natural History Museum TU-Tulane University UF-University of Florida UMML-University of Miami Marine Laborat6ry USNM-U. S. National Museum DEFINITIONS AND METHODS STANDARD LENGTH (S.L.)-from tip of snout to middle of end of caudal base (hypural bones). All,body lengths given are standard length unless otherwise specified. TOTAL LENGTH (T.L.>-from tip of snout to. veftical of longest caudal , ray. BoDY DEPTH--from origin of pelvic spine t6 origih of 2nd dorsal spine. SNOUT TO lsT DORSAL ORIGIN-from tip of snout to origin of 1st 5pine of dorsal fin. SNOUT TO PECTORAL-from tip of snout to origin of: spine of pectoral En. SNOUT TO PELVIC:-from tip of snout to origin of spine bf pelvic fin. SNOUT TO 1ST ANAL SPINE-from tip Of snout to origin of 1st spine of anal En. PEDUNCLE DEPTH-Vertical depth at end of caudal base. ANAL BASE LENGTH-fr0m origin Of lst. anal spine to origin of last anal softray. THIRD DORSAL SPINE LENGTH-from orikin to tip of 8rd spine of dor- sal En. FIRST ANAL SPINE LENGTH-from origin to tip of 1st spine of anal fin. PECTORAL LENGTH-from origin of pectoral spine to tip of'longest pec- toral softray with fin in normal position against body. PEIX[C SPINE LENGTH-from origin to tip of pelvic spine. HEAD LENGTH-greatest distance frem tip of snout to fieshy margin of operculum. EYE DIAMETER-greateSt horizontal diameter between circumorbitals. SNOUT LENGTH-leaSt distance from tip of snout to inner margin of anterior circumorbitals. UPPER JAW LENGTH-from anteriomedian tip of upper lip (premax- illary junction) to afiterioventral. angle of upper jaw (maxillary). 208 BULLETIN FLORIDA STATE MUSEUM ' Vol. 4 DoRSAL FIN couNTs-the En consists of spines continuous with soft- rays. ANAL FIN couNTs-the fin consists of spines continuous with softrays. BRANCHING OF DORSAL AND ANAL SOFTRAYs-frequently the terminal softray in the dorsal and anal fins is branched several times be- low the scaly sheath, and the next to last softray may be in close proximity to the last, making an accurate count difficult. PECTORAL FIN COUNTS-the fin consists of a spine at the fin origin and softrays. Counts were made on both pectoral fins of each fish be- cause slight bilateral variations exist. PELVIC FIN COUNTS-the fin consists of a spine and branched softrays. CAUDAL FIN COUNTS-the fin ConSiSts of 4 dorsal secondary rays, one unbranched and 5 branched dorsal principal rays, 5 bfanched and one unbranched ventral principal rays, and 3 ventral sec- ondary rays. The principal rays originate on the hypural bones; the secondary rays do not. Counts of secondary rays were made on only a few specimens that were cleared and stained with alizarine. BRANCHIOSTEGAL'COUNTs-branchiostegal rays were counted on each side of only a few cleared and stained. specimens. GILLEAKER COUNTS-upper limb and lower limb gillrakers were counted on the Brst arch. SCALE couNTs-the number of diagonal scale rows from the junction of the operculum and shoulder girdle to the caudal base were counted. This count was begun at the operculum-shoulder girdle junction, progressing posteriorly to about the tip of the pectoral fin (in normal position against body); the last row as counted above was traced ventrally to a level with the straight part of the lateral line on the caudal peduncle, and the count was con- tinued to the caudal base. Because of incomplete diagonal rows and intervening rows, the counts made are considered to have an accuracy of + 1 scale. Counts of scales along the lateral line were not recorded because most of the scales along the curved portion of the lateral line do not have pores and do not cross the lateral line, but group aboutit dorsally and ventrally. In sam- ple comparisons, from about 8 to 10 more scales were counted along the lateral line of a ~ fish than were recorded for the scale row count. 1959 BERRY: BOARFISHES OF THE GENUS ANTIGONIA 209 KEY To THE S*ECIES OF Antigonia OF THE ATLANTIC OCEAN 3 A. Dorsal spines 9 (rarely 10); dorsal shftrays 26 to 80; anal softrays 23 to 28; pectoral rays I-12 (rarely I-11·or I-18);.body depth.from ,about 15 to 70 mm. S.L. 64 20 100% .S.L., ab6ve 70·fnm. S.L. 62 to 85% S.L.; pelvic spine length 12 to 28% S.L. -_-- - .--_..«e.««--Antigonia combatia Berry and Rathjen AA. Dorsal spines 8 (rarely 7 or 9); dorsal softfays 81 to 87; anal softrays 29 to 34; pectoral rays. I-18 (rarely I-12 of I-14); body· depth from about 15 to 7.0 mm. S.L. 126 to :186% S.L., above 70 mm: S.L. 98 to 127% S.L.; pelvic spine length 24 to 88% .S.L.· _ . 1--____.._._.....Antigonia' capros Lowe Aftigonia combatia Berry and Rathjen (Figurds 1, 86 10) Antigonia capros (non Lowe), L6n*ley and Hildebrand, 1941, p. 147 (in part; Tortugas, Flofid~).-Springer and Bullis, 1956, p. 89 (in part; OREGON Sta. 32, 29'10' N., 85'55' W., Sta. 1026, 25'08' N., 84'19' W., and Sta. 1843, 22'59' N., 79'17' W.). Antigonia browni (hon. Fowler), Fowler, 1952, .p, 4(off Boynton Beach, south of· Sombrero Key Light; ·and southeast of Sand Key Light, Florida). -Briggs, 1958, p. 271 (in part; based on Fowler, 1952, p. 4, southern Florida). SYNONYMY: It is pr6bable that in their description of A. caprosi from Tortugas, Longley· and Hildebrand (1941: 147) considered two specimens of A. · combatia, USNM 171784 and USNM 116755, col- lected by W. H. Longley from south, of Tortugas, "Florida, and bearing the identity label of A. capros.. The three paratypes of A. combatia that Fowler (1952: 4) identified as A. browni from off southern Florida are cataloged as .ANSP 74840, ANSP 75409, and ANSP 75410. DIAGNOSIS: 4 Ali Antigonia with dorsal spines 9 (rarely 10), dorsal softrays '29 (26 to 86), anal softrays 27 (23 to 28), pectoral I-12 (rarely I-11 or I-13), scale rows 58 (about 49 to 57), 'body depth 66.7 (100.8 to 62.4), third (16ngest) dorsal spine length- 16.2 (30.2 to 16.2), first (longest) ·anal spine length 7.7 (16.2· to 7..7) 4 pelvic spine length 12.4 (22.6 to 12.4), upper jaw length 9.0 (8.1 to 11.1). DESCRIPS[foN: Counts and; measurements are giv. en in tables 1, 2, and S. Body compressed and elevated. Greatest vertical body depth (from base of 2nd spine of' dorsal fin to pelvic spine origin) 3 Nominal species 6f Indo-Pacific distribution are not, kexed because of their uncertain relationships. They are distinguished in the sections on relationships. 4 The number aft6f each charatter is the value recorded for the holotype. Nuinberb in parentheses ~ following £his Te,r~nges in yalues 6f all specimens. Where the proportion decreases with growth, the larger extreme of the range is placed .first. Body proportions· are'in percent of standard length (% S,L.). This procedure is followed in the Description, 210 BULLETIN FLORIDA STATE MUSEUM Vol. 4 66.7 (100.8-62.4), decreasing in proportion to S.L. with growth (fig. 4). Greatest body width (near lateral midline of body at cleithrum) 19.7 (value of holotype only). Snout to ist dorsal spine 50.0 (69.4-50.0), de- creasing iii proportion to S.L, with growth. Snout to pectoral origin 88.0 (46.8-86.4), decreasing in proportion to S.L. with growth. Snout to pelvic origin 57.7 (78.5-54.9), decreasing in proportion to S.L. with growth. Snout to 1st anal spine 68.8 (84.5-66.7), decreasing in pro- portion to S.L. with growth. Catidal peduncle deeper than long, its depth at caudal base 12.0 (16.2-11.9), decreasing in proportion to S.L. with growth. Lateral line curving upward to below about 4th dorsal spine, then curving downward approximating the dorsal body con- tour posteriad to midline of body below the dorsal softrays, extending past the caudal base, and ending immediately anterior to the teri-nina- tion of the scaly covering of the caudal ray bases. 6 Figure 1. Antigonia combatia, holotype, USNM 159597, 117 mm. in stand- .ir d length, COMBAT Sta. 259, 24'29' N., 88"28' W., southwest of Tortugas, Flcirida. Head length 34.2 (48.4-84.2). decreasing in proportion to S.L. with growth. Profile changing with growth (fig. 8); an apex produced by upper end of supraoccipital crest with concavities between apex 1959 BERRY: BOARFISHES OF THE GENUS ANTIGONIA 211 and dorsaI origin (fig. 3: F, K, Pi·Q) and between apex and snout (fig. 3: E, T, and others); these concavities varying with growth -stage and individhals (fig. 8: B, D. Snout length 10.3 (i2.5-9.9), decreasing 6nly slightly in proportion to S.L. with growth. Cleft of mouth angu- lar. Lower jaw slightly projecting. Upper jaw length (fig. 7) 9.0 (8.1-11.1), growing neady in direct proportion to S.L. Eye diameter 13.2 (20.4-18.2), decreasing in proportion to S.L.'with growth. Bony interorbital width 10.4 (value of holotype only). Denticulated ridges, about 12 to 15 in a deBnite pattern on head abdve eye, curving anteriad to posteriad. Denticulated fidges oh snout below and in front of eye, On preopercle and subopercle. Teeth small and conical in a narrow batid in .front and partially around the side of each jaw. No teeth on vomer, palatines, or tongue. Gillrakers moderate, 5 (8-5) + 16 (14-16) Branchiostegals 6. Dorsal spines 9 (9-10) (table 1). One specimen with 10 spines, cleared and stained, 65.5 mm. S.L., has 10th spine and Ist softray articulating with same intemeural, spine, while 6ther stained speci- mens with 9 spines have separate interneural spines for the 9th spine and the 1st softray. One specimen with 9 spines has the 7th spine split to about three-fourths the distance from the tip, forming two -. points. Third spine the longest 16.2 (80.2-16.2), generally decreasing in proportion to S.L. with growth; 1st spine the shortest; 2nd spine the next to the shortest· in length; and the Brd through the last spine in a graduated series decreasing in length posteriad. First and second dorsal fins continuous, shallowly notched at connection. Anal spines 8 (8). One aberrant specimen, 66 mm. S.L., has ele- ments of all 8 spines fused into one spine mass. First spine the longest 7.7 (16.2-7.7), generally decreasing in proportion to S.L. with growth; the 2nd and 8rd spines successively shorter. Third spine and 1st softray connected by a shallow ncstched membrane. Dorsal softrays 29 (26 to 30). Anal softrays 27 (28 to 28) (table 2). In both fins the 2nd softray is the longest; the 1st softray is unbranched iIi smaller speeimens and is slightly branched in some larger speci- mens, the last softray is frequentiy branched 6 or 7 times to the scaly sheath in specimens of all sizes, and the Other softrays are moderately branched m spedimens of all 5izes (softrays broken.in many specimens). Proximal portion of the softrays covered by a Reshy, scale-covered sheath. Anal fin base length 44.4 (55.2-40.7), decreasing in proportion to S.L. with growth. Caudal. fin subtruncate. Caudal principal rays 6 + 6 (in all·speci- mens); the most dorsal ray and the most ventral ray are unbranched, 212 B U LLE TIN FLO R ID A STATE M U SEU M Vol. 4 TABLE 1 RELATION, OF NUMBER OF DORSAL ,SPINES TO NUMBERS OF PECTORAL SOFTRAYS FOR Antigonia combatia (179 SPECIMENS) AND Anti- gonia Capros (108 SPECIMENS). PECTORAL SOFTRAY COUNTS ARE FROM BOTH SIDES OF EACH FISH, AND THE,SINGLE SPINE ON EACH SIDE IS NOT INCLUDED IN THIS COUNT. NUMBER OF SPECIMENS WITH EACH COMBINATION IN PARENTHESES. PECTORAL SOFTRAYS (Spine of each fin excluded from count) 11 - 12 12-12 12 - 18 18 - 18 18 - 14 VII capros (1) D O R S A L SP IN ES VIII capros (8) capros (93) capros (8) IX combatia (1) combatia (161) combatia (11) capros (8) combatia (8) X combatia (1) combatia (1) combatia (1) TABLE 2 RELATION OF' NUMBERS OF DORSAL SOFTRAYS 'rO NUMBERS OF ANAL, SOFTRAYS FOR 177 SPECIMENS OF Antigonia combatia,(com) AND 97 SPECIMENS OF Antigonia CaprOS (Cap) NUMBERS ARE THE COUNTS OBTAINED FOR EACH COMBINATION. 1959 B E R R Y: B O A R FIS H E S O F T H E G E N U S A N T IG O N IA 218 DORSAL SOFTRAYS 26 27 28 29 80 31 82 83 84 85 36 37 28 com 1 24 com com 11 25 com Col-11 corn com 2 11 12 2 26 coin corn corn com coin 1 14 57 25 2 27 com com com 10 82 5 A N A L S OF TR AY S 28 com 1 29 capcap eklp 1 1 2 80 capcap cap 4 64 81 cap cap cap 2 15 18 32 cap Icap cap cap cap 2 9 18 5 . 1 38 cap cap cap 611 84 cap 1 TABLE 8 RANGES OF MEASUREMENTS OF BODY AND FIN PARTS ~IN PERCENT OF STANDARD LENGTH) FOR DESCRIBED SIZES OR SIZE RANGES OF Antigonia combatia AND A. capros 214 B U LLE T IN F LO R ID A S TATE M U S E U M Vol. 4 Standard Body Snout to Snout to Snout to Snout to Peduncle Anal A nt ig on ia c ap ro s co m ba tia Species Locality No. Length Depth 1st Dorsal Pectoral Pelvic 1st Anal Depth Base 18 100-118 68.6- 78.5 50.0-54.0 87.7-39.7 56.1-62.8 68.2-77.9 11.9-18.2 40.7-46.6 Atlantic 11. 79-99 66.7- 79.7 50.8-56.0 36.4-41.1 54,9-64,1 68.2-77.8 12.0-18,6 42.5-50.0 & Gulf 1 62 87.1 57.3 40.8 66.1 75.8 51.6 , U. S. 9 81-45.5 89.6-100.6 61.3-69,4 40,6-45.1 62.7-78.5 75.2-84.5 14.0-15.5 50.6-55.2 2 23.5-26 92.8-100.8 68.5-68.5 45.0-46,8 69.2-70.2 82.7-83.0 15.8-16.2 49.6-52.3 Cuba 1 100 85.0 57.0 42.0 66.0 76.5 14.0 48.0 H6nduras & 2 60.5-65 86.0- 90.8 56.7-60.0 41.8-41.5 68:6-66.9 75.2-78.5 18.8-14.0 51.2-54.8 Nicaragua 4 40.5-42 84.5- 91.6 60:0-62.7 40.5-41.6 61.9-66.7 72.6-76.5 18.6-15.4 49.4=50.6 ·9 British 6 69-84.5 75.4- 85.1 51.4-56.8 89.7-41.5 61.9-68.8 71.0-75.2 12.6--18.5 46.8-51.1 5 Guiana 2 57-60 85.1- 89.2 55.8-56,7 40.8-41.2 68.2-66.7 74.2-75.8 18.5-13.7 50.9-53.8 0 Surinam 32 61-88 . 62.4- 77.2 50.8-55.9 87.0-48.6 56.4-69.7 66.7-75.5 11.7-18.3 41.3-47.9 9 51-59.5 78.2- 81.7 53.6-58.5 40.2-44,2 62.6-69.2 72.2-78.8 12.6-14.2 47.0-50.9 2 39.2-44.5 87.6- 94.4 58.4-60,7 42.7-48.6 67.4-68.9 76.4-79.1 18.5-14.8 51.7-53.6 8 69-81 64.8- 78.8 50.6-52.3 87'.7-89.7 55.6-57.6 67.9-70.9 11.9-12.5 42.1-48.5 Brazil 6 56-67.5 64.9- 75.2 51.1-54.6 88.8-42.7 57.8-65.0 67..2-74.4 12,1-13.0 42.0-44.7 8 82-51.5 72.8- 92.2 51.5-62.8 S8.8-44.4 59.2-68,8 68.9-75.6 13.6-14.7 44.7-51.3 Madeira 1 188 105.8 67.3 89.7 67.8 79.0 14.2 56.2 8 109-127 105.5-122.2 60:9-67.6 37.4-89.8 71.5-78.7 79.7-88.4 13.8-15.3 55.9-68.9 4 68.5-85 111.7-122.4 64.8-70.1 88.2-41.2 76.4-78.2 83.9-85.9 14.2-15.8 58.2-68.1 Atlantic 5 50-65.5 118.7-184.3 66.7-77.2 40.4-42.0 78.1-82.9 85.1-91.1 18.6-15.8 68.6-66.7 U. S. 8 44.5-48 116.7-182.6 64.6-71.7 89.6-48,5 79.2-88.7 85.4-90.0 15.2-15.2 60.4-67.4 1 26 129.2 75.0 44.2 81.9 86:5 16.9 69.2 19 110-184 98.8-108.7 59.6-65.0 37.4-40.9 69.3-74.8 76.4-88.5 18.9-15.8 52.5-61.0 11 100-109 100,9-118.6 60.2-66.5 87.740.8 70.2-76.1 78.6-85.8 18.9-16.5 52.8-60.8 16 69.5-97 108,5-126.6 62.7-71.2 86.2-41,8 71.2-79.1 78.0-87.1 14.0-16.2 57.1-65.5 Gulf 3 52-58.5 117.1-127.9 68.4-70.8 40.5-45.2 77.8-88.7 86.8-91.8 15.0-16,2 66.7-65.4 - U. S. 8 89-47.5 125,8-186.0 70.5-76.7 40.9-48.0 80.9-85.4 88.2-92.7 12.8-16.1 65.8-69.8 8 25:5-29 125.9-132.1 72.4-74.7 48.1-48.9 81,0-86.0 87.9-92,5 16.2-16.9 64.5-65.1 1 I81 108,8 62.2 88.2 72.1 - 76.6 14.9 57.8 Honduras & 4 76.88.5 110.2-121.7 65.5-68.5 87.8-41.8 72.9-78.8 81.4-86.2 18,4-15.8 61.0-64.5 Nicaragua 6 88-59 120.8-181.2 69.5-74.1 42.4-46.1 79.7-86.0 85.9-92.0 14.1-14.6 Surinam 2 45.5-57.5 120.0-124.2 70.3-70.4 42.4-48.5 78.8-80.9 85.2-86.8 13.6-14.6 61.2-68.7 2 88.5-89 106.8-110.1 62.7-68.7 89.8-39.8 72.0-75.8 79.1-82.2 14,6-14.7 58.4-58.8 · TABLE 3-Continued Standard 8-rd Dorsal 1st Anal Pectoral Pelvid Head Eye Snout Upper JawSpticies Length Spine Spine Length Spine Length Diameter Length Length 1959 B E R R Y : B O A R FIS H E S O F TH E G E N U S ANTIGONI 215 100-118 16.2-22.1 7.7-11:1 28.5-38.8 12.4-15.5 34.2-88.2 18.2-16.0 9.9-11.6 8.8- 9.879-99 18.4-23,8 8.2-f2.7 30.3-83.3 18.2-17.8 35.4-38.8 14.3-16.0 10.0-11.8 9.0-9.762 28.4 13.7 34.2 19.2 87.9 15.5 11.0 8.931-45.5 25.7-28.1 12.7-16.2 88.7-38.7 15.9-22.6 88.4=41.9 14.9-17.8 10.5-12.5 8.4- 9.1 ia c ap ro s tg on ia c om ba tia 28.4-26 25.0-29.4 11.9-12.8 88.8-40.4 16.5-19.6 41.9-48.4 18.5-20.4 11.5-12.8 8.1-10.2 100 24.5 12.5 32.3 17.5 89.0 16.4 11.7 9.9 60.5-65 27.8-30.2 14.0-16.2 38.8-84.0 18.2-20.6 87.147.4 16.2-16.9 10.8-11.7 9.8-10.240.5-42 28.8-28.9 18.6-14.8 35.7-88.3 18.8-20.5 37.8-89.8 15.9-19.8 10.8-11.9 8.4-10.5 69-84.5 21.9-28:9 18.0 32.3--84.5 16.8-18.4 37.649.8 15.5-17.8 10.6-11.9 9.6-10.6- 57-60 24.6 12.3-18.5 86.3 17.7-20.0 39.5-40.0 15.0-16.0 11.0-11.4 9.1- 9.861-88 17.2-23.1 8.9-11.8 30.146.2 14.1-18.2 86.8-40,9 15.4-18.9 9.9-12.8 9.2-11.151-59.5 22.8-26.5 11.5-18.8 81.9-85.8 17.3-20.2 87.4-40.4 15.9-16.5 10.1-11.8 9.2-10.039.2-44.5 25.8-28.1 18.5-18,8 34.2-86.7 18.0-20.4 40,4-41.6 15.7-16.6 11.5-12.1 8.9- 9'.969-81 17.8-19.9 8.3-10.8 81.5-83.1 14.5-14.9 36.7-38.0 14.9-15.9 10.9-11.4 9.7-10.356-67.5 18.9-28.1 9.2-12.0 88.0-36.8 14.0-16.4 37.1-40.2 15.9-17.9 11.1-12.4 10.0-11.132-51.5 22.2-26.1 12.6-14.7 85.046.0 17.1-18.9 36.9-40.9 16.5-17.5 10.5-12.0 9.8- 9.5 138 26.4 81.4 28.0 37.8 16:9 11.0 8.7 109-127 27.8-80.0 15.2-18.5 81.8-84.7 26.0-27.0 83.9-88.0 18.4-14.8 9.4-11.7 7.4- 9.168.5-85 29:6-38:2 14.6-20.4 38.3-85.0 24.1-31.2 85.8-88.2 15.8-16.1 8.9-11.1 8.9- 9.550-65:5 38.8-44.5 17.4-21.6 , 86.040.2 29.8-85.8 86.6-40.2 15.8-18.8 8.9-11.6 8.2- 9.644.5-48 88.8 16.8-19,8 85.4-88.0 27.1-81.2 '85.4-40.4 15.4-17.4 9.6-1-0.1 8.1-8.926 15.4 41.5 27.7 42.8 16.5 8.8 7.7110-184 28.8-30.0 14.1-16.7 '31.1-36.6 24.8-28.7 88.9-39.2 13.4-16.4 8.8-11.7 7.4-9.7100-109 25.7-29.8 14.2-16.6 81.5«37.5 26.5-28.4 85.8-87.6 14.4-15.6 9.4-10.7 8.1-10.069.5-97 28.9-37.1 15,4-18.5 33.8-85.9 26.0-38.1 35.1-37.4 14.0-16.8 9.0-41.2 8.6- 9.952-58.5 35.0 18.9-19.8 36.2-40.4 30.8-35.1 87.8-88.5 16.2-18.8 10.5-10.9 8.1- 9.239-47.5 85.5-48.0 16.8-20.9, '85.8=40.0 82.6-37.2 86.4-40.7 15.9-17.9 9.8-11.6 7.8- 9.625.5-29 88.2 15.5-17.6 88.6-89.2 80.2-80.6 87.9-41.2 17.0-17.8 9.8-16.7 7.5- 7.9 A131 16.0 84.4 26.0 35.1 18.7 10.5 8.876-88.5 27.7-36.2 16.6-20.5 85.6-38:0 28.2-34,9 86.6-88.2 14.9-16.6 9.7-IO.8 9.0- 9.9 -88-59 35.6-36.5 17,5-19.1 88.6-48.5 29.7-32.9 89.0-42.1 17.1-20.2 9.5111.1 9.1-10.245.5-57.5 38.8 18.8-18.7 87.4-89.6 86.0 87.9-44.0 17.6-19.1 9.9-10.8 8.8-10.488.5-89 29.5-81.5 84.5-85.4 27.1-29.9 34.5-85.6 15.4-15,5 10.1-16.4 8.5- 9.0 216 BULLETIN FLORIDA STATE MUSEUM Vol. 4 others branched. CaudaI secondary rays 4+8 (in seven cleared and stained specimens). Continuation of fleshy part of body extending posteriad over bases of caudal rays covered by about 5 vertical rows of scales. Pectoral fin bluntly pointed. Pectoral rays I-12 (rarely I-11 or I-13) (table 1). Length of pectotal spine decreasing in proportion to length of pectoral softrays with growth. First softray unbranched, others branched. Pectoral length 29.1 (40.4-28.5), decreasing in pro- porti6n to S.L. with growth. Third or fourth softray the longest. In- serted in advance of a vertical to 1st dorsal fin 6rigin. Pelvic fin pointed. Pelvic rays I-5 (in all specimens), Sth softray rudimentary in one small specimen, rays connected by a membrane, membrane connects 5th softray to body. Pelvic spine length 12.4 (22.6-12.4), decreasing in proportion to S.L. with growth (fig. 5). First softray the longest element of the fin. Inserted below 1st dorsal An. Scale rows 58 (about 49 to 57). Body completely scaled except for small bony area immediately above pectoral origin. Scales cover mandibles (exposed portion), cheeks, throat, opercle; few scales on interopercle and subopercle and on and above snout. Scale rows, usually single, extend along each (lateral) side of the spines of the dorsal, anal, pectoral, and pelvic fins, and On the softrays of the pelvie fin. Several scale rows extend distally on the membranes between ' the softrays of the pectoral and caudal fins. Most body scales have on exposed portion an elevated ridge that is bent posteriorly, aligned dorsoventrally, and denticulated on the distal margin. Most body scales also have, arising from scale base and with same alignment as the longer ridge, about 2 to 26 fiattened ridgelike structures bearing from 1' to 4 points or denticles (fig. 2). Scales immediately below the dorsal softrays and on lower portion of the peduncle· tend to lack the smaller ridgelike structures, and the number.of these structures apparently increases with growth. Scales on bones of head have ridges aligned dorsoventrally along central part of scale, those on body with ridges on posterior portion of scale. Group of scales posterior to horizontally curvihg denticulated ridges above the eye have a centrally located crescentic ridge. On most body scales circuli originate at base of the dorsoventral ridge and align with contour of anterior field of the scale. Radii occur on imbedded portion (fig. ·2). Pigmentation essentially faded out on preserved specimens. A light orange area at the dorsal and anal fin bases over the position of the interne-ult*E -Ed-ifitdfli€fnals, and a narow orange stripe sloping from the operculum-shoulder girdle junction to the midline of the 1959 BERRY : BOARFISHES OF THE GENUS ANTIGONiA 217 body at the caudal peduncle are visible on many of the specimens in formalin. A Kodachrome slide (35 mm.) taken of several specimens immediately after capture at OREcoN Sta. 2013 shows the dorsaI aspects of the head and body to be reddish pink blending into pink on the sides and then into silver on the ventrolateral aspects. f-- 1.0 mm. Figure 2. Top and side views of stale of holotype of Antigonia combatia, taken from below lateral iine under second dorsal fin. Line of dashes on top view represents the posterior extension of the scale base, lying beneath the ele- vated dorsoventral ridge. Gonads were developed in specimens from Rve collections. (12 Off Tortugas, Florida, COMBAT Sta. 259, 7 March 1957: 1 male (107 mm., with flacid testes that may have been partially spent); 2 females (111 and 117 mm., with well developed ovaries containing enlarg6d eggs). (2) West of Tortugas, Florida, OREGON Sta. 1026,19 April 1954, 29 specimens: 2 with gonads not developed (88.5 and 89.5 mm.); 10 males (62 to 111 mm., mean size 91.2 mm.; smallest with slightly de- veloped testes, others from 80 to 111 mm. with well developed testes); 17 females (79 to 118 mm; mean size 102.4 mm.; all with enlarged eggs in ovaries). (3) Off northern Cuba, OREGeN Sta, 1348, '16 July 1955: 1 female (100 mm., with ovaries smaller than previously listed females and contaiding moderately developed eggs). (4) Off Surinam, OREGON Sta. 2014 8 November 1957, of 107 specimens: 2 with gonads not developed (89.2 and 44.5 mm.); 58 males (51 to 75.5 mm., mean size ~ 65.9 mm.; testes very enlarged in specimens larger than 65 mm.); 47 - females (52 to 88 mm.,-mean size 68.8 mm.; specimens 55 mm. and smaller with small ovaries containing small eggs, specimens 55.5 mm. and larger with very large ovaries contaming well developed eggs 218 BULLETIN FLORIDA STATE MUSEUM Vol. 4 and one of these,appeared to be running-ripe). (5) Off Amazon River, OREGON Sta, 2066, 15 November 1957, lI specimens: 2 with gonads not developed (87.5 and 51.5 mm.); 8 males with slightly developed testes (56, 58.5, and 64 mm.); 2 males with well developed testes (65.5 and 67.5 mm.); 4 females with well developed ovaries (66,69,75.5, and 81 mm.). Although Weber and de Beaufort (1929: 267) stated that males of A. malayanus had longer pelvic fins than the females, I found no such discrete differences attributable to sexual dimorphism in A. combatia. Of the specimens from OREGON Sta. 1026, the males tended to have shorter pelvic spines and a lesser body depth than the females, but the values were decidedly overlapping. The smallest paratype, estimated at about 19 mm. S.L., from the stomach of a Thunnus atlanticus (Lesson), has the tail and peduncle and most of the fin rays missing. Its body depth (19.1 mm.), 1st anal spine length (2.4 mm.), pelvic spine length (8.4 mm.), and other body measurements identify it as continuous with the A. combatia series and as distinct from small A. capros. This specimen was identified as A. capros in Anderson and Gehringer (1957: 58). DISTRIBUTION: Antigonia combatia is recorded from off Brazil (Amazon River), Surinam, British Guiana, Caribbean Nicaragua and Honduras, the Leeward Islands, northern Cuba, Florida, North Caro- lina, and Massachusetts (fig. 8). DEVELOPMENT .AND GEOGRAPHIC VARIATION: The possibility that the specimens designated as paratypes of A. combatia may comprise two or more species· was considered. The variations in body propor- tions and estimated relative growth of specimens from different locali- ties suggested this, and the futute acquisition of an adequate size series of specimens from other localized areas of the western Atlantic may demonstrate the description of A. combatia to be based in part on a yet undescribed species. However, the differences encountered in this series of specimens more probably are attributable to individual and geographic intraspecific variation. The more conspicuous changes during growth of A. combatia are a depression of the body outline (decrease in body depth), and a pro- portional decrease in the lengths of the dorsal spines (principally the , 8rd), the anal spines (principally the lst), and the spine of the pelvic fins (compare F through J of fig. 8). Table 3 shows that all of the body parts measured increase in size or length mere slewly than the standard length. Therefore these body parts are proportionally smaller on large fish than on small fish. The proportional decrease of body depth to standard length with growth is apparent in figure 4. 1959 BERRY: BOARFISHES OF THE GENUS ANTIGONIA 219 Although the data available and the size range of specimens are incompletes comparisons between collections from off Brazil (Amazon River), Surinam, British Guiana, Nicaragua, Honduras, Cuba, and the United States coast in the Gulf of Mexico and the Atlantic-each of progressively more northern latitude-indicate that the southern pop- ulations reach comparable stages in their allometric or heterogonic growth at smaller body sizes than the northern populations. Specimens from off British Guiana, from the Caribbean, and. from the Gulf and Atlantic off the United States seem to be closely related in this differential development, for specimens of similar size have generally similar body proportions. This group differs most from the specimens from off Surinam and Brazil, with the Brazilian specimens showing the most extreme divergence. Geographic variation is mani- fest in several body proportions, as may be seen in the relation of body depth to standard length in Bgure 4. At smaller sizes, estimated be- tween 15 and 80 mm. S.L., specimens from the different localities apparently are directly comparable, so that measurements of both depth and sta4dard length of a small specimen from off Brazil might be expected to equal both these measurements for a specimen of the same size from off the United States. However, at sizes larger than 30 or 40 mm. S.L., geographic variation is apparent in comparison of specimens of similar length. The more southern, Brazilian specimens average the least body depth. The Surinam specimens avefage a lesser depth than those from more northern localities. Specimens from British Guiana to the United States apparently average a more closely comparable depth-length relation (an incomplete series of specimens limits this interpretation), and have greater depths than specimens from Surinam and Brazil. The 100-mm. Cuban specimen (fig. 8: 0) has a greater depth than specimens of similar length from the United States, but is generally similar in depth and other body proportions and in body outline to some smaller specimens fr6m more southern localities, as to the 65-mm. Honduran specimen (fig. 3: N). The 82.5- mm. specimen from off Surinam has a slightly different head and body profile and less body depth (62.4% S.L.) than all other specimens, and it may represent the ultimate stage ·in the development of body out- line (fig. 8: D. The northern populations probably attain greater lengths than those to the south. Assessment of the relative positions of the coordinates of the groups of specimens in figure 4 suggests that the relative increase of body depth to standard length is heterogonic, and that a complete series , of specimens from any locality might be described adequately by a curvilinear regression or by rectilinear regressions interrupted by at 220 BULLETIN FLORIDA STATE MUSEUM Vol. 4 least two growth inflections. Other body parts or measurements- snout to 1st dorsal spine, snout to 1st anal spine, snout to pelvic, 1st anal spine length, and 8rd dorsal spine length-are similar to body depth, both in sh6wing a geographic difference in relative growth and in having a heterogonic growth pattern. . 1 *'~JO (©f]/7«3-dj/U j , K %-r . P _/*« 39,2 m/i, '.38 mm, . ,- - 4.4 'L , -0% -1--- 41V }4. 44In- 6~*0 5< i- - ril' R - 56= 1,60:Sm/, 39/m '954//.3 / 1 -N ,, s 45.Smm. - Uzzr---1 f*)/1 »41_1 91 A ~L«' ' \*JU to \42 .J j f 65 5 m. - -a////2/A *...lill./.- BRAZIL SURINAM HONDURAS UNITED NICARAGUA STATES CUBA Figure 8. Body outlines of specimens of Antigonia combgtia from five differ- ent areas, illustrating intraspecific changes occurring with growth and variation in outline, and interpopulation differences in degrees of development. Reference line under each outline represents 10 mm. Body length for eadh specimen is given in millimeters of standard length. 1959 BERRY : BOARFISHES OF THE GENUS ANTICONIA 221 A\° O\0 90125 1 ' ' ' 1/1 1 71 ' r .1 1 7- x ATLANTIC & GULF U.S. / / / --1.9 CUBA 51- - o HONDURAS'B NICARAGUA If100 - 1-• SUR IN AM a BRAZIL 1 . 1 1 - ®·BRITISH GUIANA iii 010 , 1 /0./ 9 7.FE) B O D Y D E P T H (M M ) - It /- . 75 - -1 1 . - 1 1. - =t - 1 0 0 /0% / / e Zt·1< 2 , - 50- - / - . .6 / - _ 25 - - Antigonjo combolia ji'f,0,1 11'1111,1111,11'111'11'11 0 25. 50. 75 100 125 150 S'TANDA'RD LENGTH (MM) Figure 4. Relation Of body depth to standard length for Antigonia combatia from five different areas. Reference lines of dashes represent proportional values; at "90%" body depth is 90% or nine-tenths of standard length. The use of the term population in the ab6Ve discussion refers to samples collected at designated separate localities. The specimens examined may be representatives of a continuous clinal population, but the sparsity of samples over the broad area considered fails to show any geographic centers of abundande or gaps in the species range. Hence, assigning subspecific designations to the collections from these discontinuous localities is not advisable. RELATIONSHIPSE Antigonia combatia with dorsal softrays 30 or less and anal softrays 28 or less differs from A. capros Lowe (- A. browni Fowler = Caprophonus aurora Mfiller and Troschel = A. steindachneri Jordan and Fowler) and A. eos Gilbert in having fewer dorsal and anal softrays. A. combatia also differs from A. capros and A. eos in having a shallower depth of body and shorter 8rd dorsal, 1st anal, and pelvic spines. Antigonia combatia is similar in numbers of dorsal and anal soft- rays to A. rubescens (Ciinther), A. rubicunda Ogilby, A. malayanus 222 BULLETIN FLORIDA STATE MUSEUM Vol. 4 Weber, A. fowleri Franz, and A. rhomboidea McCulloch. The five species are nominal forms of Indo-Pacific distribution and of uncertain relationships. Fraser-Brunner (1950) synonymized A. mulleri (ques- tionablk possibly unidentifiable) and A. malayanus with A. rubicunda; and he synonymized A. fowleri and A. rhomboidea with A. rubescens. My examination of a larger series of specimens suggests modification5 of his interpretations (discussed below). As the relationships of these - O\0 35 - -* 0 4 2- I. f 30 - - , , .... / 25 - 0 • P E LV IC SP IN E LE N G TH ( MM ) /- . / . 1 20- ,. , . /.....15- F , . .: ./e.. ......... ' 10 - :.f V ... Antigonia~ combatio // .A.5- , O,1 1 lilli 1 1 1 1 1 1. , 1 1 1 1 1 , 1 1 1 1 1 0 25 50 75 100 125 150 STANDARD LENGTH (MM) Figure 5. Relation of pelvic spine length to standard length for Antigonia combatia and A. capros. Reference line of dashes indicates values of pelvic spine length equivalent to 28% of standard, length. 1959 BERRY: BOARFISHES OF THE GENUS ANTICONIA 223 Indo-Pacific forms are not definite, it is b.est to distinguish A. combatia from the nominal species individually. Antigonia rubescens was named Hypsinotus' sp. by Temminck and Schlegel (1844: 84, pl. XLII, fig. 2) and described from a specimen about 4 inches in total length (which I estimate -as about 80 mm. S.L.) from Japan, and the figure of the type is a natural size (1:1) reproduction. Compared to Temminck and Schlegel's figure bf A. rubescens, A. combatia has a shallower body depth (107% S.L. in A. rubescens, 62,4 to 100.8% S.L. in all specimens of .all sizes of A. combatia); a shorter 8rd dorsal spine (84% S.L. in A. rubescens, 16.2 to 30.2% S.L. in A. combatia); a shorter ist anal spine (15.6% S.L. in A. rubescens, 7.7 to 14.8% S.L. in A. combatia); and a shorter pelvic spine (28% S.L. in A. rubescens, 12.4 to 22.6% S.L. in A. combatia). The earliest redescription of A. rubescens was by Jordan and Fowler (1902: 523), and the larger specimen they illustrated appears to be conspecific (their fig. 2, USNM 50803, 119 mm. S.L.). How- ever, their illustrated specimen has an open and extended mouth, and this probably prompted Fraser-Brunner (1950: 50) to describe A. rubescens as haying, "Cleft of mouth small, nearly horizont#l." The mouth is relatively sm~11 compared to certain specimens discussed below, but the cleft of the mouth is Oblique and not nearly horizontal. Jordan and Fowler gave a scale count of. "14-60-40" (the count of 60 apparently was made along the lateral line, as I counted approxi- mately 58 and 61 scales along the lateral line for the two sides of the USNM specimen). This specimen has 44 and 45 scale rows, right and left sides, respectively, and the other 17 specimens I have identified as this species have about 38 to 47 scale rows. A. combatia differs from A. rubescens in having 49 to 57 scale rows. A. combatia has a shorter pelvic spine than A. rubescens at com- parable body sizes (from about 20 to 50 mm. S.L., 24 to 80% S.L. in A. rubescens, 14 to 28% S.L. in A. combatia, above 50 mm. S.L., 21 to 26% S.L. in A. rubescens, 12 to 20.9% S.L. in A. combatia). The 3rd dorsal spine of A. combatia averages a shorter length than that of A. rubescens (27 to· 42% S.L. in A. ritbescens, 16 to 30% S.L. in - A. combatia); the 1st anal spine averages shorter (12.8 to.17.6% S.L. in A. .rubescens, 7.7 to 16.2% S.L. in A. combatia); the pectoral fin above 50 mm. S.L. averages slightly longer (about 30 to 82% S.L. in A. rubescens, 28 to 36% S.L. in A. combatia). A. combatia has a shallower body depth at all 00mparable body lengths (84 to 106% S.L. in A. rubescens, 62 to 100.8%·S.L. in A. combatia), a slightly shorter snout to -lst dorsal spine distanee above'50 mm. S.L. (57 to 68% ' 224 BULLETIN FLORIDA STATE MUSEUM Vol. 4 S.L. in A. rubescens, 50 to 60% S.L. in A. combatia), and a longer upper jaw (8.0 to 9.5% S.L. in A. mbescens, 8.1 to ·11.1% S.L. in A. combatia). The two species also show slight differences in profile (see figure 6). The figure of Temminck and Schlegel, or its outline, has been erroneously reproduced by Goode and Bean (1895, fig. 285), by Fowler (1986. fig. 881), and probably by Thompson (1943, fig. 524), to represent A. capros. Obviously the small first dorsal spine was omitted from Temminck and Schlegel's figure and its copies, and Boseman (1947: 84) stated that Burger's manuscript gave a count of 9 spines. [Specimens of A. rubescens examined: USNM 50808 (1 specimen) 119 mm. S.L., Suruga Bayi Japan, ALBATRoss Sta. 8720, 18 May 1900, 84 fathoms. USNM 50804 (6) 40 to 79 mm. S.L., Totmi Bay, Japan, ALBATROSS. USNM 57588 (2) 77.5 and 79 mm. S.L., Yokohama, Japan. USNM 75465 (8) 26 to 60.5 mm. S.L., Mishai, Japan. USNM 185698 (2) 22 and 80 mm. S.L., Gulf Of Tokyo, Japan, ALBATROSS, 26 October 1906.] Antigonia fowleri was described by Franz (1910: 58) apparently because he believed specimens he examined from Japan and the illustrafion and description by Jordan and Fowler (1902: 528, fig. 2) of A. rubescens were specifically distinct from Temminck and Schlegers (1884: 84, pl. XLII, fig. 2) figure and description. He did not indicate adequate separations of the two forms, and because I consider Jordan and Fowler's speeimens to be conspecific with A. rubescens, I also iiiclude A. fowleri as a synonym. As A. fowleri was based in part on Jordan and Fowler's description, A. combatia is considered to differ from A. fowleri in the same respects as it does from A. rubescens. Antigonia rhomboidea, described and illustrated by McCulloch (1915: 111, pl. XVIII, fig. 1) from Victoria, Australia, may be identical to A. rubescens. McCulloch's Bgure, apparently a natural size (1:1) reproduction of a fish 115 mm. in total length (about 90 mm. S.L.), is similar in outline and proportions of body parts to Jordan and Fowler's , figure and specimen of A. rubescens (1902: 528, fig. 2, USNM 50808), and is intermediate in body depth between this figure and Temminck and Schlegel's figure. One objection to this proposed synonymy is in scale counts. McCulloch . stated that his specimen had "About fifty- eight scales between the operculum and the hypural." This descrip- tion seems to imply that McCulloch counted scale rows rather than scales along the lateral line, and as specimens I have identified as A., rubescens have about 38 to 47 scale rows, a difference exists. Using values given for A. rhomboidea by McCulloch, A. combatia has a lesser body depth (1.08 in S.L. in A. rhomboidea, .99 to 1.60 in S.L. in A. combatia) and a shorter pelvic spine (1.4 in head length in 1959 BERRY : BOARFISHES OF THE GENUS ANTIGONIA 225 A. rhomboidea, 1.75 to 2.78 in head length in A. combatia). Based on my measurements of McCulloch's figure of A. rhomboidea, A. combatia has a lesser body depth (105.6% S. L. in A. rhomboidea, 62.4 to 100.8% S. L. in A. combatia), a shorter 1st anal spine (17.8% S.L. in A. rhomboidea, 7.7 to 14.8% S.L. in A. combatia), a shorter pelvic . spine (26.4% S.L. in A. rhomboidea, 12.4 to 22.6% S.L. in A. combatia), a lesser snout to 1st dorsal spine distance (72.8% S.L. in A. Thorn- boidea, 50.0 to 68.5% S.L. in A. combatia). Antigonia rubicunda was described by Ogilby (1910: 108) from Queensland, Australia, and was redescribed and illustrated by Mc- Culloch (1915: 118, pl.,XVIII, fig. 2). I have not seen Ogilby's de- scription, but McCulloch (loc. cit.) stated that in Ogilby's (1910) description of his two type specimens, 62 and 65 mm. T.L. (about 48.5 and 52 mm. S.L.), "Ogilby has wrongly counted the number of spines ahd rays of the fins, while some of the proportions given by him are also incorrect." Using values given for A. rubicunda by McCulloch, A. combatia has more scales (45 rows in A. fubicunda, * 49 to 57 rows in A. combatia), a shorter 8rd dorsal spine (1.06 into head length in A. rubicunda, 1.28 to 2.12 into head length in A. com- batia), and a shorter pelvic spine (1.2 into head length in A. rubicunda, 1.75 to 2.78 into head length ib A. combatia). Based on my measure- ments taken from McCulloch's 8gure of A. rubicunda, A. combatia has a shorter 3rd dorsal spine (84.0% S.L. in A. rubicunda, 16.2 to 30.2% S.L. in A., combatia), and a shorter pelvic spine (27.8% S.L. in A. rubicunda, 12.4 to 22.6% S.L. in A. combatia). IA. rubicunda and A. rhombmdea may be synonymous. . The difference in scale counts given by McCulloch (45 for A. mbicunda, 58 for A. rhomboidea) suggests a specific difference, however, and McCulloch's figure of A. fubicunda 5hows the upper jaw to be longer than that in his figure of A. rhomboidea. Fraser- Brunner's (1950: 721) interpretation of this last character is _ que5tioned because he synonymized A. mahlt/ant* Weber with A. rubicunda-and I regard. A. malat/- ant,8, with a long upper jaw and a nearly vertical cleft of mouth, to be distinct from any other nominal species. I have been unable to identify two small speci- mens (BMNH 1912.11.28.68) froib the Kai Islands which Fraser-Brunner desig- nated as A. rubicunda. I have examined one of the specimens (33.8 mm. S.L.), and data on the other (88 mm. S.L.) have been kindly furnished by G. Palmer of the British Museum. The 38.3-min. specimen has a longer upper jaw (12.8% S.L. in the Kai I51ands specimen, 8.1 to 11.1% S.L. in A. combatia) than was f6und in 5pecimens of A. combatia.1 Antigonia malayanus was described and figured by Weber (1918: 299, fig. 69) from the Arafura Sea, and was redescribed by Weber and de Beaufort (1929: 267, fig. 71). I have examined a 64.5-mm. specimen of A. malayanus (CNHM · 52482) labeled "Cotype, Siboga 226 BULLETIN FLORIl)A STATE MUSEUM Vol. 4 Figure 6. Top: Antigonia rubescens, 65.5 mm. S.L., USNM 5()8()4. Middle: Antigonia malayanus. 67.5 min. S.L.. USNM 150996. Bottom: Antigonia combatia, 67.5 min. S.L.. SAFI. OREGON Sta. 2.018. 1959 BERRY: BOARFISHES OF THE GENUS ANTIGONIA 227 258, Max Weber," and a 67.5-mm. specimen (USNM 150996) from Talogo Light, Philippilie Islands, that is identical to the catype and to Weber's description and figure. These spedimens have upper jaw lengths of 12.5% S.L. (CNHM) and 18.9% S.L. (USNM), which values are greater than those of any other described species of Antigonia (figs. 6.and 7). Both specimens have 8 dorsal spines, which is the count Weber (1918: 299) gave for,14 specimens of 76 to 90 mm. T.L. (about 59 to 70 mm. S.L.). A. malayanus is similar to A. combatia in having a relatively short pelvic spine. A. combatia is distinct from A. malauanus in having normally 9 aorsal spines, a shorter upper jaw, and in alignment of the 16wer jaw (less protruded and less vertically cleft in A. combatia) (fig. 7). Herre and Herald (1951: 832) recorded this species from Luzon, Philippines, but gave no confirming descrip- Mon. Antigonia midleri was described by Klunzinger (1880: 380, pl. V, Bg. 3) from two specimens of about 85 mm, T.L. from New Zedland. In appearance and known characters, A. mulleri belougs to the family Zeidae rather than Caproidae. Fraser-Brunner (1950: 722) placed it in questionable synbnymy with A. ·rubiounda, but noted that this was uncertain because the sha116w body and large mouth of A. mulleri Were "more like that. of a Cyttid." Antigonia benhatatate (Bleeker) is apparently a nomen nudum. The earliest record I have been able to find for the specific name is in- Bleeker's 1876 Systema Percarum Rexisum (p..810), where, with no description- or Bgure, he gives, Hypsinotus Sch. (1842 ?) = Antigonia Lowe (1843) - Caproph 20 15 5 I0 95 90 85 80 75 70 65 49. 55 50 Figure 8. Distribution of Antigonia combatia and A. capros in the western Atlantic. Records of A. capros from Rio de Janeiro, Brazil (22'55' S., 43'19' W.). are not included. 1959 BERRY : BOARFISHES OF THE GENUS ANTICONIA 231 OREGON 'Sta. 2088, 01°49'N., 46°48'W., about 200 miles northeast of mouth of Amazon River, Brazil, 17 November 1958, 225 fathoms. SPECIMEN EXAMINED: SAFI. (1) 68.5 mm, S,L., OREGON Sta. 2291, 07°27'N., 54°27'W., about 80 miles northward of Paramaribo, Suri- nam, 9 September 1958, 120-185 fathoms. Antigonia capros Lowe, 1848 (Figukes. 10 and 11) Antigonia capros Lowe, 1848,' p. 85 (Madeira).-Gunther, 1887, p. 44 (in part; Madeira; Barbados).-Jordan, 1887, p. 577 (baged on records of LOwe.1843 and Muller and Troschel 1849):-Goode and Bean, 1895, p.,229 (in part; based , on records of Lowe 1843 from Madeira, Miiller and Troschel 1849 from Barbad65, and Gunther 1887 from Barbados and Madeira;, excluding fig. 285 which is A. rubescens).-Jordan and Evermann, 1896, p. 419 (in part;' based on records of Lowe 1848 and Miiller and Troschel 1849; excluding Pacific rec- , ords).-Jordan and Evermann, 1898, p. 1665 (in part; based on records of Lowe -1843 from Madeira and Muller and Tr6schel 1849 from Barbados; ex- cluding A. mullmi and A. mbescens from synonymy).-Miranda Ribeiro, 1908, p. 175 (Brasil).-? Osorio, 1909, p. 67 (Ilha de Santo Antao, Cape Verde Is- lands).-? Clark, 1918, p. 885 (Ascension Island).-Miranda Ribeir6, 1915 (Brasil).-Miranda Ribeiro, 1918, p. 76 (in part; synonymy, excluding H. rube- scens and A. mutteri).-? Route, 1919, p. 52 (southeast of Flores,.Azares, 1860 mdters; no description).-Jordan Evermann, and Clark; 1980, p. 358 (based.on records of Lowe 1843 and Muller and Trosehel 1849).-? Firth, 1981,9. 162 (75 to 90 miles southeast of Cape Henry, Virkinia, 28 to 50 fathoms; no de- scription).-Fowler, 1986, p. 894 (in part; based on records of Lowe 1848, Clark 1918, and Roule 1919 from Madeira, Ascension, and Azores; excluding fig. 881 which is A. rubescens), ? p. 1306 (based on record of Osorio 1909 from Cape Verde Islands)..Cadenat, 1987, p. 491, fig. 86 (from seven stations olf, West Africa at Cape Verde Islands, Senegal, and French Guinea, between depths of 120 and 265 meters).-Longley and Hildebrand, 1941, p. 146 (in part, off Tortuga5, Florida, 40 to 140 fathoms; probably als6 included speci- mens of A. combatia).- ? Hildebrand, 1941, p. 228 (based on. record of Firth 1931).-Howell Rivero, 1941, p. 844 (Matanzas, Cuba).-Fowler, 194la, p. 169 (based on record of Miranda Ribeiro 1903 from Rio de Janeiro, Brazil).- Raney and Ross, 1947, p. 68 (off Block Island, Rhode Island, 53 fathoms).-7 Maul in Noronha aud Sarmento, 1948, p. 78 and 149 (Madeira; no descrip- tion).-Fraser-Brunner, 1950, p. 722 (Madeira; Barbados).-Bigelow and Sthroeder, 1958, p. 488, fig. 225 (south of Nantucket Lightship, off Massa- chusetts, 55 to 80 fathoms).-Springer and Bullis, 1956, p. 89 (in part; speci- mens examined and records verified for OREGON ·Stations 156,278,277,278, 826,602,782,864,895, afid 945; excluding rec6rds from OREGON Stations 32, 1026, and 1848 which are A. combatia).-? Salzen, 1957, p. 77 (Gold Coast, West Africa; no description).-Briggs, 1958, p. 271 (froin Rhode Island to Rio de Janeiro and the northern Gulf of Mexico; compiled). Caprophonus aurora Muller and. Troschel, 1849, p. ·28, DI. V, fig, 1 (Barbados). Antigonia brotoni Fowler, 1934, p. 856, :fig. 8 (about 50 miles southeast of Five- fathom Bank Light Ship, New Jersey); 1935, p. 9 (reference to Fowler 1984); 232 BULLETIN FLORIDA STATE MUSEUM Vol. 4 1987, p. 802, (Cape Henry, New Jersey); 1952, p. 119 (New Jersey; listed).-= Briggs, 1958, p. 271 (in part; New Jersey; compiled), ? Antigonia otwavi (nomen nudum) Fowler, 194lb, p. 59 (lapsus calami ?.; "Deep Boar Fish", off New Jersey). SYNONYMS AND REFERENCES: References are included only for the Atlantic Ocean. Identifications by name only, with no descrip- tion or illustration that' can be identified, are question-ed because they may have included or have been composed of the previously undis- tinguished A. combatia. DIAGNOSIS: 6 An Antigonia with d6rsal spines 8 (rarely 7 or 9), dorsal softrays 81 to 87, anal softrays 29 to 84, pectoral I-18 (rarely I-12 or I-14), scale rows about 46 to 54, body depth 186.0 to 98.8, third (longest) dorsal spine length 44.5 to 23.8, Erst (longest) anal spine length 21.6 to 18.9, pelvic spine length 87.2 to 24.1, upper jaw length 10.2 to 7.3. DESCRIPTION: Counts and measurements are given in tables 1, 2, and 8. Body compressed and elevated. Greatest vertical body depth 136.0 to 98.3, apparently increasing in proportion to S.L. with growth to about 50 or 60 mm. S.L. and decreasing in proportion to - S.L. above this size (fig. 9; compare specimens in figs. 10 and 11). Snout to 1st dorsal spine 77.2 to 59.6, decreasing in proportion to S.L. with growth. Snout to pectoral origin 46.1 to 86.2, decreasing in proportion to S.L. with growth. Snout to pelvic origin 86.0 to 67.8, decreasing in proportion to S.L. with growth. Snout to 1st anal spine 92.7 to 76.4, decreasing in proportion to S.L. with growth. Caudal peduncle deeper than long, its depth at caudal base 16.9 to 12.8, tending to decrease in proportion to S.L. with growth. Lateral line as in A. combatia. Head length 42.8 to 83.9, decreasing in proportion to S.L. with gr6wth. Body 6utline becoming shallower with growth, but anterio- dorsal profile remaining characteristic with slight concavities between snout and upper end of supraoccipital crest and between upper end of supraoccipital crest and dorsal fin origin. Snout length 8.8 to 11.7, increasing nearly in direct proportion to S.L. with growth. Cleft of mouth angular, and lower jaw slightly projecting. Upper jaw length 7.8 to 10.2, increasing nearly in direct proportion to S.L. with 5 Only values f6r specimens from the Atlantic Ocean are given. The numbers after each character are the ranges in values for these specimens. Where the pro- portion decreases with growth, the larger extreme of the range is placed first. Body proportions are in percent of standard length. This procedure is followed in the Description. 1959 BERRY: BOARFISHES OF THE GENUS ANTIGONIA 288 growth. Eye diameter 20.2 to 13.4, decreasing in proportion to S.L. with growth. Denticulated ridges on head similar to A. combatia. Teeth similar to A. combatia. Gillrakers 5 to 6 + 13 to 16. Branchiostegals 6. Dorsal spines 8 (rarely 7 or 9) (table 1). One specimen with 9 dorsal spines has the 5th spine split to about one-third 6f the distance from the tip forming two points and with the 9th spine appearing normal. X-ray of another specimen with 9 spines shows that the 9th spine abnormally has no interneural spine associated with it. Third spine the longest 44.5 to 28.8, decreasing in proportion to S.L. with growth. Spines in graduated ~ length series and 1st and 2nd dQrsal fins connected as in A. combatia. Anal spines 8. First spine the longest 21.6 to 13.9, decreasing in proportion to S.L. with growth. Graduation of spines and connection to 1st anal softray as in A. combatia. Dorsal softrays 81 to 37. Anal softrays 29 to 84 (table 2). A 128-mm. specimen has an abnormal anal fin consisting of only two spines and 26 softrays, with the first two softrays hardened and short- ened like spines but possessing segment marks (the dorsal fin of this Bsh has 9 spines and 33 softrays). Approximate length of softrays, branching, and basal sheath as in A. combatia. Anal fin base length 69.8 to 52.5, decreasing in proportion to S.L. with growth. Caudal fin subtruncate. Caudal principal rays 6+6. (in·all speci- mens examined). Caudal secondary rays 4+8 (in two stained speci- mens). Branching of rays and scales on caudal as in A. combatia. Pectoral fin bluntly pointed. Pectoral rays I-18 (rarely I-12 or I-14) (table 1). Pectoral length 43.5 to 31.1, decreasing in proportion to S.L. with growth. Relative growth of spine, branching and com- parative length of rays, and insertion of pectoral as in A. combatia. Pelvic fin pointed. Pelvic rays I-5 (in all specimens). Pelvic spine length 37.2 to 24.1, decreasing in proportion to S.L. with growth