<I>Rypticus</I> (Teleostei: Serranidae)

BULLETIN OF MARINE SCIENCE, 65(2): 337–379, 1999

REVISION, PHYLOGENY AND COMMENTS ON BIOGEOGRAPHY OF SOAPFISHES OF THE GENUS RYPTICUS (TELEOSTEI: SERRANIDAE)

Ricardo Zaluar Passos Guimarães

ABSTRACT Soapfishes of the Atlanto-East Pacific genus Rypticus are reviewed. Rypticus macrostigmus Courtenay and R. brachyrhinus Courtenay are junior synonyms of R. bornoi Beebe and Tee- Van and R. randalli Courtenay respectively. A phylogenetic analysis of the genus based on outgroup comparison with other members of the tribe Grammistini indicates Rypticus is a monophyletic lineage supported by six modifications in axial-skeleton structure and reduction in the number of infraorbital bones. A single, fully resolved tree illustrates the hypothesized relationships among species. Some of the cladogenetic events hypothesized based on morphology are corroborated by vicariant biogeography.

The tropical to warm-temperate marine percoid genus Rypticus Cuvier comprises the Atlanto-East Pacific serranids commonly referred to as soapfishes, alluding to their capacity to produce large amounts of skin mucous when stressed. This mucous contains a strong toxin, grammistin, which is repulsive for potential predators (Randall et al., 1971). Soapfishes are small to medium-sized cryptic inhabitants of reefs, where they feed primarily at night on small crustaceans, mollusks and fishes (Graham, 1972; Hobson, 1965; present study). They are protogynous hermaphrodites (Smith, 1965) and have planktonic eggs and larvae (Houde, 1982; Kendall, 1984). Rypticus was originally described more than 160 yrs ago and its position among other percoids varied considerably in proposed schemes (compare, for example, Smith and Atz, 1969: fig. 3; Kendall, 1976: fig. 1). Johnson (1983) cladistically defined the Serranidae and included Rypticus, some other soapfish genera (i.e., Grammistes, Pogonoperca and Grammistops) and the pseudogrammid genera (i.e., Pseudogramma, Suttonia and Aporops) in tribe Grammistini of the Epinephelinae. Subsequent studies (e.g., Kendall, 1984; Johnson, 1988) have continued to detail the phylogenetic structure of the Serranidae. Finally, Baldwin and Johnson (1993) considered Rypticus as the immediate sister-group to (Grammistops + (Jeboehlkia + (Aporops + (Pseudogramma + Suttonia)))), this whole group as the sister-group of (Grammistes + Pogonoperca), all in a monophyletic Grammistini. Intrageneric relationships of Rypticus were only previously hypothesized by Courtenay (1965, 1967) and McCarthy (1979), but neither author included all species in their analysis, and these were based on simple observations of shared characters. Eleven species of Rypticus are currently recognized, eight from the Atlantic (Courtenay, 1967) and three from the Eastern Pacific (McCarthy, 1979). Since reexamination of type specimens and/or topotypes indicated that some of these are invalid and because a significant amount of specimens and information have been collected and published since these last reviews, a complete taxonomic review of the group is warranted. Monophyly of the genus and relationships among species are hypothesized based on a cladistic analysis of characters. Possible vicariant scenarios are discussed for geographically isolated clades with a sister-group relationship.

337 338 BULLETIN OF MARINE SCIENCE, VOL. 65, NO. 2, 1999

METHODS

COUNTS AND MEASUREMENTS.—Following Courtenay (1967) and McCarthy (1979), counts and measurements were taken as described by Hubbs and Lagler (1949), with the tip of the upper jaw as the anterior landmark for snout, head, predorsal and standard lengths. Counts of vertebrae, supraneurals, total number of caudal rays and number of co-ossified proximal and middle radials in dorsal and anal fins are from radiographs or cleared and stained specimens. Counts of dorsal and anal-fin rays were taken from radiographs, cleared and stained or dissected alcohol specimens. In the descriptions, each meristic range is followed by the modal value in parentheses. Information on the number of gill rakers on the first arch and pectoral-fin rays are based on Courtenay (1967) and McCarthy (1979). In the descriptions and tables 1–2, 4–6, data for number of dorsal-fin spines, dorsal-fin rays, caudal vertebrae, anal-fin rays, predorsal distance, head length, snout length and upper jaw length of Atlantic species are a combination of the present data and those of Courtenay (1967). OSTEOLOGY.—Osteological terminology mostly follows that of Johnson (1983, 1984) and Baldwin and Johnson (1993). Osteological studies were performed primarily on specimens cleared and coun- terstained (cs) for bone and cartilage (Pothoff, 1984). Additionally, radiographs (r) and dried skel- etons (ds) were also examined. TYPES.—Among nominal species with extant type material, those of R. maculatus Holbrook, R. microps Castelnau, R. coriaceus Cope and R. bicolor Valenciennes were not examined in the present study, but examination of their original descriptions and/or topotypes indicates the synonymies of Courtenay (1967) and McCarthy (1979) should be followed. MAPS.—Locality records plotted on maps are those listed in the “Appendix: Material Examined” section and those cited in the following publications: Bullock and Smith (1991), Caldwell (1963), Courtenay (1967; 1981), Eskinazi and Lima (1968), Hetzel and Castro (1995), Hoese and Moore (1977), Koike and Guedes (1981), Mago-Leccia (1965), Martins-Juras et al. (1987), Maugé (1990), McCarthy (1979), Oliveira (1974; 1979), Rodriguez (1973), Rosa (1980), Roux (1973), Smith (1976) and Starks (1913). Records of R. saponaceus from Arraial do Cabo, state of Rio de Janeiro and Guarapari, state of Espírito Santo and that of R. subbifrenatus from the city of Rio de Janeiro are based on personal field observations. ABBREVIATIONS.—Institutional abbreviations are as listed in Leviton et al. (1985), except for UFRJ which refers to the fish collection at the Laboratório de Ictiologia Geral e Aplicada, Universidade Federal do Rio de Janeiro. PHYLOGENETIC ANALYSIS.—The character analysis conducted in this study follows the basic pre- mises of phylogenetic systematics (e.g., Hennig, 1966; Wiley, 1981), i.e., taxa are grouped based on shared apomorphies. Polarity of character states was determined by outgroup comparison (see Nixon and Carpenter, 1993). This was primarily accomplished via direct observation of other Grammistini available for examination. For those characters with more than one state in the outgroups (all Grammistini genera except Rypticus), its plesiomorphic condition in the ingroup (Rypticus) was determined with the aid of Maddison and co-workers’ (1984) algorithm applied upon Baldwin and Johnson’s (1993) phylogeny of the Epinephelinae. Because expansion of branches can alter topology of a previous tree, a simultaneous analysis of the entire Grammistini plus all species of Rypticus was conducted. This was done by adding to the matrix (Table 11) all those characters that contribute directly to the proposed Grammistini phylogeny, i.e., characters 16, 21, 32, 33 and 36 to 52 of Baldwin and Johnson (1993). Regarding these characters, the present work either supports or does not have any evidence to consider their states incorrectly assigned to the outgroups by those authors, except as noted in the text. A parsimony analysis in which reversals and convergences are permitted and counted equally as a single step was conducted by means of the “ie” algorithm of Hennig86, which certainly generates the tree with a minimum length (Farris, 1988). This was done with the aid of the Tree Gardener matrix and tree editor (Ramos, 1996). GUIMARÃES: REVISION OF RYPTICUS 339

Table 1. Frequency distribution of number of dorsal spines (A) and dorsal rays (B) in Rypticus.

AB III IVI I021222324252627282 R. bicolor -31 - ----2---- R. bistrispinus 57 ------12 320211 R. bornoi 22-5 ----18012- R. courtenayi 12 ------26--- R. maculatus 37 4- --1-8181- 7 R. nigripinnis 11-8 -----23-- R. randalli 977 - ---10 927--- R. saponaceus -79 - ---16 948--- R. subbifrenatus -1113 1141444461---

Table 2. Frequency distribution of total number of vertebrae (A), number of anal rays (B) and total number of caudal rays (C) in Rypticus.

AB C 254 23141516171425262 R. bicolor 3- ----2 -2- R. bistrispinus -63--71421 34- R. bornoi 1-5 -13216 32- R. courtenayi 8- - -36- 134 R. maculatus 318 114291 --1 R. nigripinnis 5- --121 -2- R. randalli 4-6 -2127- -53 R. saponaceus 7-6 -145442 -1- R. subbifrenatus 1-1206 158 7- 562

Table 3. Frequency distribution of number of pyloric caeca (A) and number of preopercular spines (B) in Rypticus.

AB 345 12345 R. bicolor -31 12 3 3-1 R. bistrispinus 4-- --321 7 R. bornoi 31- -912- 3 R. courtenayi -11 21 2 --- R. maculatus --1 771 --- R. nigripinnis 4-- 83 0 --- R. randalli 7-- 288-- 0 R. saponaceus -3- -44-- 4 R. subbifrenatus 3--19 237--

POLYMORPHISM.—Characters 13, 14, 15 and 18 have one or more taxa with more than one state. Because HENNIG86 does not support a “01” entry, each polymorphism was treated differently in order to avoid loss of information (e.g., Nixon and Davis, 1991). For example, three species presented polymorphism for character 15 (i.e., have two or three opercular spines), but each has a different frequency distribution of states. While R. randalli typically has two spines, R. nigripinnis and R. maculatus typically have three. Because reduction in number of opercular spines is a novelty within the Serranidae, it was assumed that presence of three spines (state 0, coded “0” in the ma- 340 BULLETIN OF MARINE SCIENCE, VOL. 65, NO. 2, 1999

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Table 5. Frequency distribution of head length in percent of standard length in Rypticus.

310 32333435363738393041424 R. bicolor -----12411--- R. bistrispinus --1-177121 42221 R. bornoi ------158432- R. courtenayi -----1111---1 R. maculatus - -11245195-1-- R. nigripinnis -----721--1-- R. randalli 2915 11 94------R. saponaceus - -12812 7261116--- R. subbifrenatus -----16992434232

Table 6. Frequency distribution of snout length (a) and upper jaw length (B) in percent of standard length in Rypticus.

AB 5678901213141516171819102 R. bicolor - - 81-- ---45- - -- R. bistrispinus -53 1 732 --218 61 6521 R. bornoi --1 5 71- ----2182- 0 R. courtenayi - - 311- ---13- -1- R. maculatus -120 611- -11412011-- R. nigripinnis - 33--- -2711---- R. randalli 143 6--- 1317282---- R. saponaceus -252 821- -65143028--- R. subbifrenatus -225 9561 6 ----734 35212

Table 7. Frequency distribution in Rypticus. A) eye length (in percent of SL); B) interorbital width (in percent of SL).

AB 678911 0121 234 R. bicolor -441--- -7- R. bistrispinus ---10121 -86 R. bornoi -----37 37- R. courtenayi -22---- -4- R. maculatus -14---- 32- R. nigripinnis 13331- - 29- R. randalli 8322- - - 21- 4 R. saponaceus 562---- 211- R. subbifrenatus -25521- 114 1 trix) was transformed into presence of two or three spines (state 01, coded “1” in the matrix), although this reduction is not fixed in any of the species. In regard to character 14, the parsimony analysis indicated a different transformation series. Presence of one or two supraneurals (state 01, coded “0” in the matrix) is plesiomorphic in regard to presence of only one supraneural (state 1, coded “1” in the matrix), since outgroups typically have two supraneurals (state 0, also coded “0” in the matrix). MULTISTATE CHARACTERS.—Characters 2, 5, 8, 10, 13 and 17 have more than two states within the group analyzed. In the absence of any evidence that would indicate a character transformation series, characters 2, 5, 8, 10 and 13 were treated as unordered (or nonadditive). In the case of 342 BULLETIN OF MARINE SCIENCE, VOL. 65, NO. 2, 1999

Table 8. Frequency distribution of body depth in percent of standard length in Rypticus.

265 27282920313233343536373839304 R. bicolor ---222-2------R. bistrispinus 113233------R. bornoi 122122------R. courtenayi ----1111------R. maculatus ------1-1-1-11 R. nigripinnis - -21216------R. randalli -131122212------R. saponaceus ------136111---- R. subbifrenatus - - -1131322- - -1- -

Table 9. Frequency distribution of number of supraneural bones (A), number of co-ossified proximal and middle radials in dorsal fin (B), number of co-ossified proximal and middle radials in anal fin (C) in Rypticus and outgroups.

ABC 123 1234 123 Grammistes -4- 3--- -2- Pogonoperca -12 3--- -3- Grammistops 12- 3--- -3- Jeboehlkia 11- 1--- -1- Aporops -3- 3--- -3- Suttonia 12- 3--- -2- Pseudogramma -5- 3--- 12- R. bicolor 3-- -2-- -12 R. bistrispinus 45- -5- - -31 R. bornoi 41- -211 121 R. courtenayi 8- - 152- 151 R. maculatus 1-- --1- -1- R. nigripinnis 4-- 1-2- -4- R. randalli 72- -321 -51 R. saponaceus 3-- -1-- -31 R. subbifrenatus 13 - - 352- 281

Table 10. Frequency distribution of number of localities (= stations) in which different species of Rypticus were collected simultaneously in the western Atlantic.

Rs. bornoi Ri. bistrispinu Rs. randall Rs. saponaceu R. subbifrenatu R. bornoi ---1 2 R. bistrispinus ---3 - R. randalli ---3 1 R. saponaceus 133-3 2 R. subbifrenatus 2-123- GUIMARÃES: REVISION OF RYPTICUS 343

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character 17, the states observed in adults of some taxa correspond to an early ontogenetic stage of other taxa. There has been much debate about the significance of ontogenetic information for character state coding (e.g., de Queiroz, 1985; Brooks and Wiley, 1985; Mabee, 1989, 1996; Nelson, 1985; de Pinna, 1994; Patterson, 1996). The present study follows Hauser and Presch (1991) by assuming that a character transformation series is to be determined by means of and not prior to a parsimony analysis. Therefore, multistate characters were not ordered. MISSING ENTRIES.—Two different types of missing entries (i.e., unknown data and inapplicable character) are represented in the matrix (Table 11). Although each is logically different with regard to binary characters (Platnick et al., 1991), both were coded as “-“ in the Hennig86 matrix, as this is the only character for missing entries supported by the program.

SYSTEMATICS

Rypticus Cuvier

Rypticus is characterized by the combined presence of two to four dorsal spines; 20 to 28 dorsal rays; no anal spines; 13 to 18 anal rays; pelvic fin with one spine and five rays; 24 to 26 total caudal rays; 13 to 17 pectoral rays; two or three opercular spines; one to five preopercular spines; small embedded scales with concentric rings and ctenii absent; numerous subcutaneous mucous glands; dorsal and anal-fin bases thick and fleshy; dorsal fin low at spiny portion, increasing in height posteriorly; vertical and pectoral fins rounded; pelvic fins reduced, their bases slightly anterior to pectoral-fin bases, innermost rays attached to abdomen by a membrane; mouth large and oblique, with projecting lower jaw; supramaxillary bone present; villiform teeth in bands on pre-maxillary, dentary, vomer and palate; presence of a light brown stripe from the tip of lower jaw to first dorsal spine; nasal rosette with a unique series of longitudinally oriented lamellae; three to five pyloric caeca; five infraorbitals; five to twelve total gill rakers on first arch; seven branchiostegals; one or two supraneurals; 24 (10 + 14) or 25 (10 + 15) vertebrae; seventh interneural space not vacant; first proximal-middle anal radial supporting two elements; three epurals; one uroneural; five hypurals; hypurapophysis present and procurrent spur absent.

Rypticus bistrispinus (Mitchill) (Fig. 1A)

Bodianus bis-trispinus Mitchill, 1818 (original description, Straits of Bahama). Rypticus bistrispinus, Courtenay, 1967 (synonymy, redescription, figure); Eskinazi and Lima, 1968 (range extension); Roux, 1973 (range extension; description); Smith, 1976 (range extension); Guitart, 1985 (diagnosis, figure); Bullock and Smith, 1991 (range extension, description, figures); Carvalho, 1992 (diagnosis); Guimarães, 1997 (range extension).

Diagnosis.—The following combination of characters separates R. bistrispinus from its congeners: dorsal-fin spines two, opercular spines three; preopercular spines three, body covered with small closely-set reddish brown spots, spots concentrating dorsoanteriorly; pores along ventral surface of lower jaw and posterior margin of preopercle large and simple. Description.—Dorsal-fin spines two; dorsal-fin rays 24–28 (25–26); anal-fin rays 15– 17 (16); pectoral-fin rays 13–16 (16); total caudal-fin rays 24–25; gill rakers on first arch GUIMARÃES: REVISION OF RYPTICUS 345

Figure 1. A) Rypticus bistrispinus, live adult, 82 mm SL, UFRJ 2204, Arraial do Cabo, Brazil. B) Rypticus bornoi, preserved adult , 45 mm SL, UFRJ 3061, Port-au-Prince Bay, Haiti. C) Rypticus subbifrenatus, preserved subadult, 61 mm SL, MZUSP 48014, Arembepe, Brazil. 346 BULLETIN OF MARINE SCIENCE, VOL. 65, NO. 2, 1999

Figure 2. Head, ventral view. A) Rypticus bistrispinus, MCZ 46644, 80 mm SL; B) Rypticus nigripinnis, MCZ 45625, 90 mm SL; C) Rypticus saponaceus, MCZ 42637, 141 mm SL. SMP: sub-mandibular pore; POP: preopercular pore. Scale bars equal 5 mm.

seven to nine (eight); opercular spines three; pre-opercular spines three to five (three); pyloric caeca three; supraneural bones one or two; vertebrae 25. Body cylindrical, with body depth typically 27 to 30% SL and head width modally 18 to 19% SL; dorsal profile of head convex; eyes large (usually 10 to 11% SL in adult specimens). Four large, simple pores along ventral surface of lower jaw and four on posterior margin of preopercle, last two submandibular pores rarely broken into two smaller ones (Fig. 2A). Largest examined specimen 117 mm SL; reaches 121 mm SL (Courtenay, 1967). Frequency distributions of selected characters are presented in Tables 1–9. GUIMARÃES: REVISION OF RYPTICUS 347

Figure 3. Map with distributions of Rypticus bistrispinus (circles) and R. bornoi (stars). Scale bar equals 1000 km.

Coloration.—Body cream to pale brown, covered with small closely-set reddish brown spots; spots concentrated on dorsal half of body, especially on head; spots extending to proximal portion of soft dorsal-fin and caudal peduncle; mosaic of spots sometimes broken by pale brown gaps; distal portion of soft dorsal-fin, caudal fin, pectoral fins and anal fin pale brown; spinous dorsal-fin whitish cream. Distribution.—Known from South Carolina south to Arraial do Cabo, southeastern Brazil (Fig. 3). Habitat.—Rypticus bistrispinus is more frequently collected from insular habitats (sensu Robins, 1971), i.e., those with warm, clear waters. It is a relatively common species along the rocky shorelines of Arraial do Cabo, where it greatly outnumbers the sympatric R. saponaceus and R. subbifrenatus. Behavior.—In Arraial do Cabo, R. bistrispinus is more often seen during day-time hours, especially at dawn, whereas it is rare at night. It swims closely to the bottom, usually above the sand at the edge of the reef. Remarks.—Records of R. bistrispinus from the eastern Gulf of Mexico and eastern coast of the United States are from offshore deeper waters (up to 430 m), where there is influence of warm Caribbean currents (Bullock and Smith, 1991; Smith, 1976; present study). In the coastal portion of these regions, R. bistrispinus is replaced in abundance by R. maculatus (Bullock and Smith, 1991). Specimens from southeastern Brazil presented modal displacement in number of anal rays in comparison to those from the Caribbean area (15 in Brazil vs 16 in Caribbean) (Guimarães, 1997). 348 BULLETIN OF MARINE SCIENCE, VOL. 65, NO. 2, 1999

Rypticus bornoi Beebe and Tee-Van (Fig. 1B)

Rypticus bornoi Beebe and Tee-Van, 1928 (original description, Lamentin Reef, Port-au-Prince Bay, Haiti; figure); Courtenay, 1967 (redescription; figures). Rypticus macrostigmus Courtenay, 1967 (original description, Grand Bahama, Bahamas; figures).

Diagnosis.—Differs from other soapfishes by possessing typically two dorsal spines, usually three preopercular spines, large dark-brown rounded spots on anterior portion of body, pores along ventral surface of lower jaw and posterior margin of preopercle large and simple, and in reaching a maximum size of about 60 mm SL. Description.—Dorsal-fin spines two or three (two); dorsal-fin rays 24–27 (25–26); anal-fin rays 14–17 (16); pectoral-fin rays 13–15 (13); total caudal-fin rays 24–25; gill rakers on first arch 8–12 (10); opercular spines three; pre-opercular spines two to four (three); pyloric caeca three or four (three); supraneural bones one or two (one); vertebrae 24 (10+14). Eyes large (11 to 12% SL in adults); pectoral fins elongate (modally 22–23% SL). Body moderately compressed (body depth 26–30% SL). Head convex on its dorsal profile. Juveniles and adults with four large undivided pores along ventral surface of lower jaw and four on posterior margin of preopercle. The smallest species of the genus, reaching 61 mm SL. See also Tables 1–9 for frequency distributions of selected characters. Coloration.—Body, including base of vertical fins, tan to pale brown, with rounded dark spots, these typically larger than the eye’s pupil dorsoanteriorly, smaller ventrally and absent posteriorly; spots sometimes coalesced immediately behind orbit; distal portion of pectoral and vertical fins clear; a cream stripe runs from the tip of lower jaw to base of dorsal fin, sometimes weakly delineated in specimens >40 mm SL. Courtenay (1967) illustrated a 23 mm SL specimen, which has fewer and smaller spots than do adults. Distribution.—Known from the Bahamas, Belize, Haiti, Honduras and Panama (Fig. 3). Habitat.—Among examined lots with available ecological data, two bear “coral reef ” in their labels (USNM 170572, FMNH 95453), and two bear “drop-off ” (AMNH 34471, ANSP 114563). This information, together with other listed localities, indicates R. bornoi is also exclusively found in stable, clear water habitats. There is no evidence supporting Courtenay’s (1967) statement that “the areas in which this species occurs is typically characterized by a substrate of silty marl interspersed with dead coral formations”. Remarks.—The original description of R. bornoi was based on a single specimen that agrees with the types of R. macrostigmus in the following aspects: two dorsal spines; 24 vertebrae; submandibular and preopercular pores large and simple; dark spots in anterodorsal portion of body. Furthermore, this specimen has 26 dorsal rays (24 to 26 in R. macrostigmus types), 16 anal rays (14 to 17 in macrostigmus), 13 pectoral rays (13 to 14 in macrostigmus), predorsal distance 45% SL (42 to 47 in macrostigmus) and head length 41% SL (36 to 41 in macrostigmus). The supposed peculiarities of R. bornoi’s type are: three narial openings on the same side of the body (an aberration already noticed in some R. macrostigmus specimens, e.g., ANSP 114563, some specimens with posterior nare divided in two, and some R. bistrispinus, e.g., one specimen from the lot UFRJ 2204 with a bifid tube on first nare), presence of three preopercular spines on one side of the body and two on the other (different counts of preopercular spines on each side are not so rare GUIMARÃES: REVISION OF RYPTICUS 349

in Rypticus), upper jaw length 19% SL (17 to 18 in R. macrostigmus, one specimen examined in this study with 19), snout length 9% SL (7 to 8 in R. macrostigmus, this higher value being probably due to the shape in which R. bornoi’s holotype was fixed, that is with its mouth opened and premaxillary projected anteriorly).

Rypticus subbifrenatus Gill (Fig. 1C)

Rhypticus subbifrenatus Gill, 1861 (original description, St. Thomas, Virgin Islands). Rypticus subbifrenatus, Courtenay, 1967 (synonymy, redescription, figures); Courtenay, 1970 (range extension, brief description, figure); Randall et al., 1971 (figure); Almeida, 1973 (range extension); Hoese and Moore, 1977 (diagnosis, range extension); Koike and Guedes, 1981 (range extension); Guitart, 1985 (diagnosis, figure); Maugé, 1990 (synonymy; range in eastern Atlantic); Humann, 1996 (diagnosis, picture); Guimarães, 1997 (range extension).

Diagnosis.—Dorsal spines three or four (three), opercular spines three, body pale brown with dark brown spots, these rounded, usually the size of eye’s pupil and restricted to the head in adults; pores along ventral surface of lower jaw and posterior margin of preopercle small and numerous in adults. Description.—Dorsal-fin spines three or four (three); dorsal-fin rays 20–25 (23); anal- fin rays 13–16 (15); pectoral-fin rays 14–17 (15); total caudal-fin rays 24–26 (25); gill rakers on first arch seven to ten (nine); opercular spines three; pre-opercular spines one to three (two); pyloric caeca three; one supraneural bone; vertebrae 24 (10 + 14). Anterior region of body with a distinct positive allometry: predorsal distance modally 42% SL, head length 36% SL, snout length 8% SL and upper jaw length 18% SL. Dorsal profile of head convex in adults. Juveniles (to ca. 25 mm SL) with four large undivided pores along ventral surface of lower jaw and four on posterior margin of preopercle. Larger specimens with successive subdivisions in these pores, resulting in patches of very small pores in adults (with 60 mm SL or more). Reaches 150 mm SL. Frequency distributions of selected characters are presented in Tables 1–9. Coloration.—Most collections are of small to moderate-sized specimens (20–90 mm SL), and these have the body pale brown to olivaceus, with dark rounded spots, these usually weakly ocellated and smaller than the eye’s pupil. Larger specimens (100 to 150 mm SL) have spots restricted to the anterior portion of the body and are posteriorly dark brown and covered with pale faint spots. Courtenay (1967) fully described the ontogeny of pigmentation in R. subbifrenatus. Distribution.—In the western Atlantic, R. subbifrenatus occurs from Florida south to Rio de Janeiro city (Fig. 4). Hoese and Moore (1977) indicated it also occurs in the Gulf of Mexico. In the eastern Atlantic, R. subbifrenatus occurs from Senegal south to Angola; also reported from Cape Verde and Annobon (Courtenay, 1970; Maugé, 1990). Habitat.—Most of the material examined is from tide-pools and shallow water coral reefs. In Florida, Courtenay (1967) observed R. subbifrenatus preferably occupies deep holes in the reef and suggested it’s possibly a nocturnal species. In the state of Rio de Janeiro, adult specimens are rarely seen around rocky reefs, always at night. According to Maugé (1990), adults favor clearer water habitats than juveniles. Humann (1996), reported R. subbifrenatus from depths of up to 43 m. 350 BULLETIN OF MARINE SCIENCE, VOL. 65, NO. 2, 1999

Figure 4. Map with the distribution of R. subbifrenatus. Scale bar equals 1000 km.

Rypticus maculatus Holbrook (Fig. 5A)

Rypticus maculatus Holbrook, 1855 (non-Gill, 1862; original description, Cape Romaine, South Carolina); Courtenay, 1967 (synonymy, redescription, figures); Smith, 1976 (range extension); Hoese and Moore, 1977 (diagnosis; anal-fin counts in error; habitat); Bullock and Smith, 1991 (range extension, figures); Humann, 1996 (diagnosis, picture).

Diagnosis.—Presence of few scattered and well defined white spots on sides of body is an unique character among other Rypticus. Dorsal spines typically two (sometimes three); pores along ventral surface of lower jaw and posterior margin of preopercle large and simple; body deep (depth 33 to 40% SL). Description.—Dorsal-fin spines two or three (two); dorsal-fin rays 22–27 (25); anal- fin rays 13–17 (15); pectoral-fin rays 13–17 (15); total caudal-fin rays 26; gill rakers on first arch seven to nine (nine); opercular spines three; pre-opercular spines one or two (two); pyloric caeca three to five; one supraneural bone; vertebrae 24 (10 + 14). Body markedly deep (depth 33–40% SL). Dorsal profile of head straight. Four large simple pores along ventral surface of lower jaw and four on posterior margin of preopercle. Largest examined specimen 201 mm SL. Frequency distributions of selected characters are presented in Tables 1–9. GUIMARÃES: REVISION OF RYPTICUS 351

Figure 5. A) Rypticus maculatus, preserved, 93.7 mm SL, USNM 315518, off Georgia, USA. B) Rypticus nigripinnis, preserved subadult, 72 mm SL, USNM 321625, Panama City, Panama. C) Rypticus randalli, live subadult, 76 mm SL, UFRJ 3040, Ilha Grande Bay, Brazil. 352 BULLETIN OF MARINE SCIENCE, VOL. 65, NO. 2, 1999

Figure 6. Map with distributions of Rypticus maculatus (squares), R. nigripinnis (circles) and Rypticus randalli (stars). Scale bar equals 1000 km.

Coloration.—Body, including bases of dorsal and caudal fins dark brown (except for belly and lower half of head which are pale brown) with a few, scattered and well defined white spots, these smaller than the eye’s pupil, usually absent in fins and sometimes coalesced. Distal portions of dorsal and caudal fins, anal fin and pectoral fin pale brown. Bullock and Smith (1991) described the juvenile coloration of R. maculatus. Distribution.—Rypticus maculatus is the only species in the genus with a warm-temperate pattern of distribution (or Carolinian sensu Smith, 1976). It is known from Cape Hatteras, North Carolina, to the western Gulf of Mexico (Fig. 6). Habitat.—According to Courtenay (1967), R. maculatus prefers “cooler deeper waters and a sandy rather than calcareous marl or mud.” However, other authors indicate that this species preferably inhabits rocky bottomed areas, such as rocky reefs in the eastern Gulf of Mexico (Bullock and Smith, 1991), hard bottoms and reefs or oil platforms and jetties in the inshore Gulf of Mexico (Hoese and Moore, 1977). It has been reported from 91 m (Courtenay, 1967), although in the deeper and warmer offshore waters of the eastern Gulf of Mexico and east coast of the United States, R. maculatus is replaced in abundance by its congener R. bistrispinus (Bullock and Smith, 1991). GUIMARÃES: REVISION OF RYPTICUS 353

Rypticus nigripinnis Gill (Fig. 5B)

Rhypticus nigripinnis Gill, 1861 (original description, Panama). Rypticus nigripinnis, McCarthy, 1979 (synonymy, redescription, figures); Thomson et al., 1987 (diagnosis, figure).

Diagnosis.—The combined presence of two (sometimes three) dorsal spines; three (sometimes two) opercular spines; four undivided pores along ventral surface of lower jaw and four on posterior margin of preopercle in juveniles (to 50 mm SL), each pore broken into two to four smaller pores in larger specimens; non-ocellated pale spots over sides of a dark brown body in juveniles, spots ocellated in adults and a short snout (snout length 6% SL) separates this species from its congeners. Description.—Dorsal-fin spines two or three (two); dorsal-fin rays 24–28 (26); anal- fin rays 14–18 (16); pectoral-fin rays 15–17 (15); total caudal-fin rays 25; gill rakers on first arch seven to ten (eight); opercular spines with typically three spines well developed and externally visible, lowermost spine sometimes weak or absent; pre-opercular spines one or two (two); pyloric caeca three; a single supraneural bone; vertebrae 24 (10 + 14). Head, snout and upper jaw shorter than in all other species except R. randalli. Dorsal profile of head moderately concave in adults. Juveniles (to 50 mm SL) with the typical set of four large undivided pores along ventral surface of lower jaw and four on posterior margin of preopercle. Adults with all submandibular pores except anteriormost divided into two to four smaller ones (Fig. 2B). Reaches 180 mm SL. Frequency distributions of selected characters are presented in Tables 1–9. Coloration.—Adults have a dark brown body covered with ocellated pale spots, these extending onto vertical fins and head and sometimes diffuse; distal margins of vertical fins darker. McCarthy (1979) described and illustrated the ontogeny of pigmentation in R. nigripinnis. Distribution.—Tropical eastern Pacific, from upper Sea of Cortez to northern Peru, including the Galápagos (Fig. 6). Habitat.—Information from examined lots indicate R. nigripinnis preferably inhabits coastal areas, such as sheltered bays and mangroves. One lot (USNM 321626, from Toboguilla Island, Panama) exceptionally bears “coral reef next to shore” on its label. It is sometimes collected together with R. bicolor from the same station (Table 10). According to Graham (1972), R. nigripinnis lives in depths ranging from shallow waters to 15 m. Remarks.—Based on experiments with physiology, Graham (1972) concluded that R. nigripinnis is not capable of performing seasonal acclimatation, i.e., it doesn’t tolerate large seasonal variations in temperature. This fact, together with the significant seasonal variation in water temperature from the northern Sea of Cortez (Graham, 1972), could explain the rarity of R. nigripinnis from that area (see Fig. 6).

Rypticus randalli Courtenay (Fig. 5C)

Rypticus randalli Courtenay, 1967 (original description, off Río Añasco, north of Mayagüez, Puerto Rico, figures); Randall et al., 1971 (toxin; figure), Rodriguez, 1973 (range extension); Oliveira, 1974 (first record for the Parnaiba estuary); Oliveira, 1979 (notes on habitat); Menezes and 354 BULLETIN OF MARINE SCIENCE, VOL. 65, NO. 2, 1999

Figueiredo, 1980 (range extended to São Paulo state; brief description; illustration); Rosa, 1980 (range extension); Guitart, 1985 (diagnosis); Martins-Juras et al., 1987 (range extension); Carvalho, 1992 (description; figure). Rypticus brachyrhinus Courtenay, 1967 (original description, Mindi Cut, Panama Canal, figures).

Diagnosis.—Dorsal spines two or three (two); opercular spines two or three (two); body dark brown sometimes with pale blotches on its posterodorsal portion; pores along ventral surface of lower jaw and posterior margin of preopercle typically undivided, except posteriormost submandibular one, which is typically subdivided into two or more pores in adults; snout length six percent of SL. Description.—Dorsal-fin spines two or three (three); dorsal-fin rays 23–25 (24); anal- fin rays 14–16 (15); pectoral-fin rays 14–17 (15–16); total caudal-fin rays 25–26 (25); gill rakers on first arch seven to eleven (nine); two dorsalmost opercular spines always well developed, lowermost variably developed, usually absent; pre-opercular spines one to three (two); pyloric caeca three; supraneural bones one or two (one); vertebrae 24 (10 + 14). Anterior region of body with a marked negative allometry; adults with a small head (head length modally 33% SL), short snout (its length modally 6% SL), short upper jaw (length 14% SL) and small eyes (6% SL). Specimens up to 90 mm SL with four large, undivided pores along ventral surface of lower jaw and four on posterior margin of preopercle. Larger specimens with the last submandibular pore typically broken into two, rarely three smaller ones. Largest examined specimen 180 mm SL. Frequency distributions of selected characters are presented in Tables 1–9. Coloration.—Body brown, sometimes with distal portions of vertical fins darker and usually with small pale blotches on posterodorsal region; venter and lower half of head pale in juveniles to pale brown in adults; a cream stripe runs from the tip of lower jaw to posteriormost dorsal-fin spine, stripe tending to fade with growth; live specimens in situ with a grayish-blue cast, notably on distal portions of vertical fins. Distribution.—Known from Cuba and southern Bahamas south to Santa Catarina, southern Brazil (Fig. 6). Habitat.—Rypticus randalli typically lives in rocky mainland areas with hyposaline continental influence. It has been collected in silty bottom habitats in the Caribbean from as deep as 15 m (Courtenay, 1967; Rodriguez, 1973), estuaries in the north, northeast and southeastern Brazil (Martins-Juras et al., 1987; Menezes and Figueiredo, 1980; Oliveira, 1979: in salinities varying from 13.5 to 30.6‰; Rosa, 1980), or even in lower parts of rivers (Courtenay, 1967; present study). In the rocky patch reefs under the influence of the Guanabara and Ilha Grande bays, southeastern coast of Brazil, this species can be seen at night swimming just above the rocky bottom, sometimes sympatricly with R. saponaceus. Behavior.—A cryptic species, difficult to approach, even at night. Remarks.—Among differences used by Courtenay (1967) to separate R. randalli from R. brachyrhinus, none appeared consistent. He stated the most evident difference to be number of opercular spines, i.e., two in randalli vs three in brachyrhinus. All kinds of intermediate stages for this character have been recognized in the present study, ranging from a sharply pointed third (lowermost) spine clearly visible externally to its total absence. Courtenay (1967) also suggested that brachyrhinus has the posterior margins of anal and dorsal fins moderately rounded, while in randalli these are sharply rounded, and that brachyrhinus has a darker coloration, with its vertical and pectoral fins almost black GUIMARÃES: REVISION OF RYPTICUS 355

vs “brown-gray body with paler-colored belly, dorsal and caudal fins dark brown, anal fin gray with dark gray to black distal border” in randalli. Such differences are consistent if only the holotypes were considered, but inclusion of remaining specimens indicate these are unrealistic.

Rypticus saponaceus (Bloch and Schneider) (Fig. 7A)

Anthias saponaceus Bloch and Schneider, 1801 (original description). Rypticus saponaceus, Carvalho, 1950 (listed); Courtenay, 1967 (synonymy, redescription, figures); Courtenay, 1970 (brief description, figure); Randall et al., 1971 (toxin, figure); Koike and Guedes, 1981 (range extension); Randall, 1983 (diagnosis, picture); Guitart, 1985 (diagnosis, figure); Lubbock, 1980 (range extension); Maugé, 1990 (synonymy, eastern Atlantic range); Lubbock and Edwards, 1981 (range extension); Carvalho, 1992 (brief description; range extension); Hetzel and Castro, 1995 (figure); Guimarães, 1997 (range extension).

Diagnosis.—The following combination of characters separates R. saponaceus from its congeners: dorsal spines always three, opercular spines always three, preopercular spines typically two; lower jaw projected and with a fleshy protuberance at anterior tip in adults; adults with numerous, small, patched pores along ventral surface of mandible and preopercular margin; body dark brown, except anteroventrally, covered with pale roundish spots, spots sometimes coalesced anterodorsally. Description.—Dorsal-fin spines three; dorsal-fin rays 23–25 (24); anal-fin rays 14–17 (16); pectoral-fin rays 14–17 (15–16); total caudal-fin rays 25; gill rakers on first arch five to eleven (eight); opercular spines three; pre-opercular spines two or three (two); pyloric caeca three; one supraneural bone; vertebrae 24 (10 + 14). Body moderately deep (depth modally 33% SL), compressed. Dorsal profile of head concave, especially in adults. Lower jaw projected, with a small fleshy protuberance at tip sometimes visible in adults. Juveniles (to 55 mm SL) with four large, simple pores along ventral surface of lower jaw and four on posterior margin of preopercle. In larger specimens, these pores break into smaller ones, starting with the last submandibular pore. In adults (ca 100 mm SL or more), each juvenile pore is represented by patches of very small pores (Fig. 2C). Largest examined specimen 285 mm SL. Frequency distributions of selected characters are presented in Tables 1–9. Coloration.—Body brown, darker posteriorly, with numerous pale brown spots, these typically roundish, about the size of the eye’s pupil and sometimes coalesced anterodorsally; pectoral and vertical fins darker than body. Live specimens have a marked bluish gray cast over darker areas of body. Courtenay (1967) described and illustrated development of color patterns in R. saponaceus. Distribution.—The most widely distributed species in the genus, R. saponaceus occurs from southern Florida south to Rio de Janeiro in the western Atlantic, Atlantic mid-ridge islands (Lubbock and Edwards, 1981; present study), Cabo Verde Islands and in the eastern Atlantic from Mauritania south to Angola (Maugé, 1990; Courtenay, 1970, 1981) (Fig. 8). Habitat.—Available ecological data on material examined and personal field observations in southeastern Brazil agree with Courtenay’s (1967) statement that “R. saponaceus is an ubiquitous species, occurring from the shallow silty waters near shore to clear water 356 BULLETIN OF MARINE SCIENCE, VOL. 65, NO. 2, 1999

Figure 7. A) Rypticus saponaceus, preserved subadult, 70 mm SL, MZUSP 47993, Arembepe, Brazil. B) Rypticus bicolor, preserved subadult, 64 mm SL, USNM 330424, Utria Bay, Colombia. C) Rypticus courtenayi, preserved, 81.9 mm SL, USNM 218386, Revillagigedo Islands. GUIMARÃES: REVISION OF RYPTICUS 357

around coral reefs.” Nevertheless, there is no evidence supporting Courtenay’s (1967) statement that R. saponaceus or any other soapfish are shallow burrowers in muddy areas. Rypticus saponaceus is collected syntopically with R. bistrispinus, R. randalli and very oftenly from the same tide-pool as R. subbifrenatus (Table 10). Already recorded from 60 m (Maugé, 1990). Behavior.—In southeastern Brazil, R. saponaceus is typically found either in caves during the day or swimming openly at night, usually close to bottom, sometimes in the water column. In this area as well as in others, it sometimes lies motionless or flushes its body against rocks (Lubbock, 1989; Lubbock and Edwards, 1981; present study).

Rypticus bicolor Valenciennes (Fig. 7B)

Rypticus bicolor Valenciennes, 1846 (illustration only, issued before description, Galápagos). Smecticus bicolor Valenciennes, 1855 (description, Galápagos). Rypticus bicolor, Hobson, 1965 (behavior); McCarthy, 1979 (synonymy, redescription, figures); Randall et al., 1971 (figure); Thomson et al., 1987 (diagnosis, figure); Humann, 1993 (diagnosis, picture). Rypticus nigripinnis, Humann, 1993 (misidentification, picture).

Diagnosis.—Differs from other soapfishes in having the following combination of characters: dorsal-fin with three, rarely two spines; specimens larger than 60 m SL with submandibular pores small, numerous and patched; body brown covered with numerous small cream spots, these faint and sometimes coalesced. Description.—Dorsal-fin spines two or three (three); dorsal-fin rays 23–26 (24); anal- fin rays 16–18 (17); pectoral-fin rays 14–17 (16); total caudal-fin rays 25; gill rakers on first arch seven to nine (eight); opercular spines three; preopercular spines one to five (two); pyloric caeca four or five (four); one supraneural bone; vertebrae 24 (10 + 14). Body moderately compressed (head width modally 14% SL) and deep (depth around 30 % SL). Lower jaw projected, with fleshy protuberance at tip sometimes visible in adults. Dorsal profile of head convex in juveniles and subadults and slightly concave in adults. Juveniles (to ca. 35 mm SL) with four simple pores along ventral surface of lower jaw and posterior margin of preopercle (Fig. 2A). Pores break into a series of small ones with growth, starting with posteriormost submandibular positions (Fig. 2B,C). Frequency distributions of selected meristic and morphometric characters are presented in Tables 1–9. Coloration.—Body, including vertical fins and excluding opercular area, of preserved adult specimens (larger than about 60 mm SL) is brown (darker at tips of vertical fins) and has numerous pale cream spots, these usually roundish and faint, sometimes coalesced. Anteroventral portion of body uniformly pale brown. McCarthy (1979) described the ontogeny of pigmentation in R. bicolor. Distribution.—Tropical eastern Pacific, from southern Sea of Cortez to northern Peru, also Cocos and Galápagos islands, absent from the Revillagigedo Islands, where it is replaced by R. courtenayi (Fig. 8). Habitat.—Rypticus bicolor preferably inhabits oceanic islands or unprotected coastal reefs within a depth range of 0 to 50 m. It has already been reported at depths of 69 m in the Sea of Cortez (Thomson et al., 1979) and 75 m in the Galápagos (Humann, 1993). 358 BULLETIN OF MARINE SCIENCE, VOL. 65, NO. 2, 1999

Figure 8. Map with distributions of Rypticus bicolor (stars), R. courtenayi (squares) and R. saponaceus (triangles). Scale bar equals 1000 km.

Behavior.—During daytime, R. bicolor mostly hides among crevices, sometimes lies motionless on the bottom or flushed against rocks (Hobson, 1965; Thomson et al., 1979). It is more active at night, when solitary specimens swim usually close to the bottom, occasionally in water column (Hobson, 1965).

Rypticus courtenayi McCarthy (Fig. 7C)

Rypticus courtenayi McCarthy, 1979 (original description, Revillagigedo Islands, figures).

Diagnosis.—Differs from its congeners in possessing typically two dorsal-fin spines, adults with pores along preopercular margin and anterior portion of ventral surface of mandible numerous, small and evenly spread; body, except anteroventral region, dark brown with irregular pale spots, these sometimes coalesced and usually concentrated anterodorsally and absent in vertical fins. Description.—Dorsal-fin spines two or three (two); dorsal-fin rays 23–26 (25); anal- fin rays 15–17 (16); pectoral-fin rays 14–16 (16); total caudal-fin rays 24–26 (26); gill rakers on first arch seven to nine (eight); opercular spines three; pre-opercular spines one or two (two); pyloric caeca four or five; one single supraneural bone; vertebrae 24 (10 + 14). Dorsal profile of head convex. Juveniles (to ca 60 mm SL) with four large undivided GUIMARÃES: REVISION OF RYPTICUS 359

pores along ventral surface of lower jaw and four on posterior margin of preopercle. Pores breaking with growth. Adults with distinct patches of small pores on two posteriormost submandibular pore positions; other pore positions indistinct, i.e., broken into small pores but these not displayed in patches. Reaching 200 mm SL (McCarthy, 1979). Frequency distributions of selected characters are presented in Tables 1–9. Coloration.—Adults have a brown body covered with pale cream spots, these usually roundish, the size of the eye’s pupil and concentrated in anterodorsal portion of body, spots sometimes coalesced and typically absent in vertical fins. McCarthy (1979) described the ontogeny of coloration in R. courtenayi. Distribution and Habitat.— This species is restricted to the Revillagigedo Islands (Fig. 8), where it probably lives in association with the rocky substrate

PHYLOGENY

HISTORICAL BACKGROUND.—New World soapfishes were initially treated in three different genera by Gill (1861), but Jordan and Evermann (1896) and Schultz and Reid (1939) indicated such division was unrealistic. Since then, Rypticus has been treated as a natural assemblage either by pre-cladistic authors (e.g., Gosline, 1960; 1966; Kendall, 1976; 1979; Randall et al., 1971; Smith, 1965) or cladistic ones (Baldwin and Johnson, 1993; Johnson, 1983; 1988), although none clearly tested monophyly of the genus. From the initial work of Jordan and Eigenmann (1890), who reserved an exclusive sub-family to accommodate Rypticus in the Serranidae to the detailed phylogeny of the Epinephelinae of Baldwin and Johnson (1993), others have contributed significantly to the understanding of the phylogenetic relationships of Rypticus. Katayama (1959) was the first to recognize affinities of soapfishes (sensu lato) with groupers and allies. Gosline (1960) suggested that pseudogrammids (genera Aporops, Suttonia and Pseudogramma) are closely related to soapfishes based on morphology of the olfactory lamellae and other characters and included both in the Grammistidae. In contrast, Smith and Atz (1969), argued that Rypticus and Pseudogramma belong to distant lineages, based on evidence from their gonadal morphology. Later, Gosline (1966) removed the Latiinae and Percichthyinae from Serranidae, and recognized the presence of three opercular spines as an unique character uniting the Serraninae, Anthiinae, Epinephelinae and Grammistidae. Kendall (1976), corroborating Katayama’s early hypotheses, recognized within the serranoids one Epinephelinae-Grammistidae lineage, based on predorsal bone pattern. This lineage also included Liopropoma and the pseudogrammids (sensu Gosline). Later, Kendall (1977) found within the Epinephelinae-Grammistidae lineage a close relationship among Liopropoma, Pseudogramma and Rypticus based on larval morphology. He then included the three genera in Grammistinae (this time again recognized as a Serranidae sub-group). Johnson (1983) cladistically defined monophyly of Serranidae based on four derived features (namely the loss of the small posterior uroneural pair or its fusion to the larger anterior pair, absence of a procurrent spur and third preural radial cartilages, and presence of three opercular spines) and kept Gosline’s subdivisions of the group. Johnson (1983) also agreed with Kendall’s epinepheline-grammistid lineage (and treated it as the serranid subfamily Epinephelinae), but suggested it can only be cladistically supported by absence of the autogenous distal element of the first pterygiophore, rather than by the predorsal pattern (although later he would use this character as a synapomorphy for a 360 BULLETIN OF MARINE SCIENCE, VOL. 65, NO. 2, 1999

Figure 9. Cladogram showing hypothesized relationships among Rypticus species. Numbers followed by ‘ indicate a third state of a single character, numbers followed by the symbol ® indicate reversal to the primitive state. See Baldwin and Johnson (1993) for outgroup characters. group formed by Epinephelinae tribes Epinephelini, Diploprionini, Liopropomini and Grammistini). He further added that the relative position of genera within Kendall’s lineage is inconsistent with other findings, especially concerning the placement of Pogonoperca. Finally, Johnson suggested that the Epinephelinae should be divided in five monophyletic tribes (see Johnson 1983, Table 1 for genera placement). Although he listed some autapomorphies for the tribes, he omitted evidence for relationships among them. Kendall (1984) suggested monophyly of a group consisting of Johnson’s Epinephelinae tribes Diploprionini, Liopropomini and Grammistini based on distinct morphology of dorsal spines of larvae. Johnson (1988) corroborated Kendall’s (1984) hypotheses and suggested that tribe Epinephelini is the sister group of the clade described above. He also listed three synapomorphies for a clade consisting of the four tribes and recognized tribe GUIMARÃES: REVISION OF RYPTICUS 361

Figure 10. Anterior portion of dorsal fin. A) Rypticus bistrispinus, UFRJ 2204, 58 mm SL; B) Rypticus randalli, MCZ 44415, 61 mm SL; C) Rypticus saponaceus, AMNH 29970, 86 mm SL. SN: supraneural, NS: neural spine, PR: proximal radial, MR: medial radial, DR: distal radial, DS: dorsal spine. Scale bars equals 5 mm.

Niphonini as its sister group. Baldwin and Johnson (1993) in a broader survey of characters corroborated monophyly of Johnson’s epinepheline tribes, except for placement of Jeboehlkia in Grammistini rather than in Liopropomini. To Johnson’s (1983) list of autapomorphies for Epinephelinae tribes, they added one more for Diploprionini and ignored one and added two others for the Grammistini. They used all these characters to build a phylogeny of the Epinephelinae, corroborating Johnson’s (1988) clades. A sister group relationship between Liopropomini and Grammistini was suggested based on four characters, two of them being larval features first suggested as unique to Pseudogramma, Rypticus and Liopropoma by Kendall (1977). Baldwin and Johnson (1993) indicated a monophyletic Grammistini is supported by ten synapomorphies. Within the tribe, Grammistes + Pogonoperca are considered the sister group of Rypticus + (Grammistops + (Jeboehlkia + (Aporops + (Pseudogramma + Suttonia)))) (Fig. 9). 362 BULLETIN OF MARINE SCIENCE, VOL. 65, NO. 2, 1999

MONOPHYLY OF RYPTICUS

(1) REDUCTION IN NUMBER OF DORSAL-FIN SPINES.—The presence of two to four spines in the dorsal fin is unique for Rypticus in the Serranidae. Other Grammistini have from six to eight dorsal spines. (2) INCREASE IN TOTAL NUMBER OF DORSAL-FIN ELEMENTS.—Rypticus has from 26 to 28 total dorsal elements, vs 20 to 21 in Grammistes and Pogonoperca, 19 in Grammistops, 16 in Jeboehlkia and 28 to 31 in pseudogrammids. In the most parsimonious scenario, an increase in number of elements occurred independently in Rypticus and pseudogrammids. (3) ABSENCE OF ANAL-FIN SPINES.—Absence of anal spines is unique for Rypticus among other serranids. In remaining Grammistini there are two or three spines. (4) FIRST PROXIMAL-MEDIAL RADIAL OF ANAL FIN ELONGATE, NOT EXPANDED DISTALLY AND BEARING TWO ELEMENTS.—Except for Rypticus, in all Grammistini the first anal proximal- middle element is thick and distally expanded in order to support three elements, these either three spines (e.g., Jeboehlkia; Fig. 11A) or two spines and a soft ray (e.g., Grammistes; Fig. 11B). In Rypticus, the first proximal-middle element is elongated, not expanded distally and gives support to only two elements, the first two soft rays (Fig. 11C). (5) INCREASE IN TOTAL NUMBER OF ANAL ELEMENTS.—Rypticus is the only Grammistini genus with 15 to 16 total anal elements (vs 11 in Grammistes and Pogonoperca, 12 in Grammistops, 10 in Jeboehlkia or 18 to 24 in pseudogrammids. Again, the most parsimonious scenario is that in which an increase in number of anal-fin elements occurred independently in Rypticus and pseudogrammids. (6) REDUCTION IN NUMBER OF INFRAORBITALS.—With the exception of Jeboehlkia, which has only two ossified infraorbital elements (probably caused by truncation in ontogeny, see Baldwin and Johnson, 1993), the remaining Grammistini non-Rypticus (e.g., Grammistes; Fig. 12A), the Liopropomini and the Diploprionini have six infraorbitals. The presence of five infraorbitals is synapomorphic for Rypticus and is probably caused by the co-ossification of infraorbitals 5 and 6, since three pores are sometimes identified in infraorbital five (Fig. 12B). (7) SEVENTH INTERNEURAL SPACE NOT VACANT.—As mentioned by Baldwin and Johnson (1993: Fig. 14), all Grammistini except for Rypticus have the seventh interneural space vacant, i.e., there are no proximal radials between the seventh and eighth neural spines. In Diploprion, Liopropoma and Rypticus, the seventh interneural space is occupied by the seventh proximal radial. It is most parsimonious to assume that a vacant space is synapomorphic for the Grammistini and a reversion synapomorphic for Rypticus. (8?) INCREASE IN NUMBER OF CO-OSSIFIED PROXIMAL AND MIDDLE RADIALS IN THE SOFT DORSAL FIN.—Except for Rypticus, in all grammistins only the first proximal radial serially associated with the soft dorsal fin is co-ossified with its correspondent medial radial (Table 9). In Rypticus, usually the two or three first proximal radials serially associated with the soft dorsal fin are co-ossified with their correspondent medial elements (Fig. 10; Table 9). As the remaining serranids have at least the first two proximal radials fused to the middle elements (Baldwin and Johnson, 1993), it is equally parsimonious to suppose that the condition observed in the first (Grammistes + Pogonoperca) and second (Grammistops + Jeboehlkia + pseudogrammids) outgroups of Rypticus is autapomorphic for each as it is to suppose only one co-ossification was the state in the ancestor of the Grammistini and GUIMARÃES: REVISION OF RYPTICUS 363

Figure 11. Anterior portion of anal fin. A) Jeboehlkia gladifer, USNM 330430, 52 mm SL; B) Grammistes sexlineatus, USNM 218886, 68 mm SL; C) Rypticus nigripinnis, MCZ 42608, 75 mm SL. PM: proximal-middle radial S: spines R: soft ray. Scale bars equal 5 mm.

a reversion occurred in Rypticus. In this last hypothesis, an increase in the number of co- ossified elements would be synapomorphic for Rypticus. (9?) MUCOUS CELLS TYPE II PRESENT ONLY IN BASAL LAYER OF EPIDERMIS.—Randall et al. (1971) examined histology of some soapfish species and found two mucous-cell types: type I, generally found in fish and type II, exclusive of soapfishes. Randall et al. (1971) and Aida et al. (1973) observed that in Grammistes and Pogonoperca, cells of type II are present both in the distal and basal layer of epidermis, while in R. bicolor, these are restricted to the basal layer. Type II cells are also present in Aulacocephalus, Diploprion and Grammistops, but absent in pseudogrammids (Baldwin and Johnson, 1993; Randall et al., 1971). If further studies indicate all soapfishes except Rypticus have type-II cells in both layers, then absence of those cells from the distal layer would be synapomorphic for Rypticus. 364 BULLETIN OF MARINE SCIENCE, VOL. 65, NO. 2, 1999

Figure 12. Infraorbital series, left side. A) Grammistes sexlineatus, USNM 218886, 68 mm SL; B) Rypticus bicolor, MCZ 45607, 81 mm SL. I: infraorbital. Scale bars equal 1 mm.

(10?) TESTIS CONFINED TO A REGION INTERMINGLED WITH OVARIAN TISSUE.—Smith (1965) discussed patterns of sexuality in serranids and found three basic types: simultaneous hermaphrodites (typically found in the Serraninae), protogynous hermaphrodites with mixed testicular and ovarian tissues (typically found in the Epinephelinae) and protogynous hermaphrodites with separate male and female tissue (typically found in the Anthiinae). In Rypticus, Smith (1965) found a state intermediate between the last two, in which there is also protogyny, and both testicular and ovarian tissues are histologically separate, but morphologically entwined. Smith and Atz (1969) found a fifth state in Pseudogramma, in which the testis is confined to a median lobe placed dorsally in the oviduct. An unequivo- cal phylogenetic interpretation of the state in Rypticus depends on a more detailed comparison of gonadal morphology within the Grammistini. GUIMARÃES: REVISION OF RYPTICUS 365

Figure 13. Last precaudal vertebra and anterior portion of first proximal-medial radial of anal fin, ventral view. A) Rypticus bicolor, MCZ 52656, 80 mm SL; B) Rypticus bistrispinus, MCZ 90071, 85 mm SL. C: centrum, HA: hemal arch, P: parapophyses, PM: first proximal-middle element of anal fin.

PHYLOGENY OF RYPTICUS

Rypticus bornoi + R. bistrispinus (11) PARAPOPHYSES OF LAST PRECAUDAL VERTEBRA FUSE POSTERIORLY BUT DO NOT BIFURCATE DISTALLY.—In all remaining Grammistini, the parapophyses of the last precaudal vertebrae fuse posteriorly and bifurcate ventrolateraly (Fig. 13A). In Rypticus bistrispinus and bornoi, these parapophyses fuse posteriorly to support the posterior tip of the first proximal-middle element of anal fin (Fig. 13B). Rypticus bornoi (12) PRESENCE OF DARK ROUNDED SPOTS.—Both Rypticus bornoi and R. subbifrenatus show the presence of dark-brown, rounded spots on sides of body at least during part of their ontogeny (Fig. 1B,C). Although such a color feature is unique in the Grammistini, its appearance is more parsimoniously interpreted as independent for each species. Rypticus maculatus + R. nigripinnis + R. randalli + R. subbifrenatus + R. saponaceus + R. bicolor + R. courtenayi (13) REDUCTION IN NUMBER OF PREOPERCULAR SPINES.—Known larvae of all grammistins genera have five or six preopercular spines (Baldwin and Johnson, 1993). Except for Jeboehlkia, which retain the larval condition in adults, in all genera these spines disap- pear at the juvenile stage and are replaced by new ones (Baldwin and Johnson, 1993). Adults of Grammistops and the pseudogrammids have only one preopercular spine. Adults of Grammistes, Pogonoperca, R. bornoi and R. bistrispinus typically have three spines on the preopercle while the remaining species of Rypticus have two (Table 3). In the 366 BULLETIN OF MARINE SCIENCE, VOL. 65, NO. 2, 1999

Liopropomini, the preopercle is either devoid of spines or weakly serrated (Baldwin and Johnson, 1993). The ancestral state for Rypticus cannot be unequivocally calculated based on the outgroups (is either one or three spines), but since one state of the ingroup is also found in the outgroup (three spines), this state is assumed plesiomorphic. Hence, presence of two spines is synapomorphic for maculatus, nigripinnis, randalli, subbifrenatus, saponaceus, bicolor and courtenayi. (14) REDUCTION IN NUMBER OF SUPRANEURAL BONES.—Presence of two supraneural bones is a primitive condition in the Epinephelinae (Baldwin and Johnson, 1993) that is present (partially at least) in all Grammistini except Rypticus (Table 9). Rypticus bistrispinus, bornoi and randalli have one or two supraneurals, while in maculatus, nigripinnis, subbifrenatus, saponaceus, bicolor and courtenayi, the first supraneural is always absent (Fig. 10; Table 9). The condition observed in R. randalli is best interpreted as a reversion. Rypticus maculatus + R. nigripinnis + R. randalli (15) REDUCTION IN NUMBER OF OPERCULAR SPINES.—Rypticus coriaceus Cope (1871), correctly recognized as a junior synonym of R. saponaceus by Courtenay (1967), has histori- cally been referred to a form with only two opercular spines (e.g., Jordan and Evermann, 1896; Beebe and Tee-Van, 1928). Courtenay (1967) correctly created a new name to accommodate this form (Rypticus randalli), but failed to recognize the polymorphic condition of that character and erected another name to include those specimens with three spines (see also ‘Remarks’ under R. randalli above). Later, Bullock and Smith (1991) pointed out that some specimens of R. maculatus also have the unusual two opercular spines condition, and in the present study it was observed that such condition is also found in some R. nigripinnis individuals. Presence of three opercular spines was recognized by Gosline (1966) as a distinct feature that grouped the serranids and grammistids known at that time. Johnson (1983) cladistically redefined the Serranidae based on four synapomorphies, one of them being the presence of three opercular spines. Absence or reduction of the lowermost opercular spine is unique for these three soapfish species among all other serranids. Rypticus nigripinnis + R. randalli (16) CAPACITY TO INVADE HYPOSALINE HABITATS.—Information based on personal field observations, habitat characterization in labels of examined material and published ecological data indicate that R. nigripinnis and R. randalli show a marked preference for inhabiting protected coastal areas, both often invading estuaries. Although ecological information is incomplete for the outgroups, tolerance for hyposaline habitats appears to be a derived condition within the Grammistini. (17) POSTERIORMOST SUBMANDIBULAR PORES AND UPPERMOST PREOPERCULAR PORES SPLIT INTO TWO TO FOUR SMALLER ONES DURING ONTOGENY.—Juveniles of all Grammistini have four pores on the ventral surface of the mandible and other four in the distal margin of the preopercle. Adults of Grammistops, Jeboehlkia, Suttonia, Pseudogramma, R. bistrispinus, bornoi and maculatus typically retain the juvenile condition (Fig. 2A). In adults of all other Rypticus there are at least a few subdivisions of some of these pores. In Rypticus randalli and nigripinnis, the third and fourth submandibular pores are transformed into a series of two to four smaller ones with growth (Fig. 2B). In Rypticus subbifrenatus, saponaceus, bicolor and courtenayi, all original pores are transformed into numerous tiny ones (Fig. 2C). Although the condition in outgroups indicates that the state in the ancestor of Rypticus is a retention of the juvenile state, and since other states of the ingroup are equivalent to consecutive ontogenetic steps, this character was treated unorderly in GUIMARÃES: REVISION OF RYPTICUS 367

the parsimony analysis (see justification in “Phylogenetic Analysis” section above). This analysis indicated division of pores either appeared independently in randalli + nigripinnis and subbifrenatus + saponaceus + bicolor + courtenayi or appeared in the ancestor of maculatus + randalli + nigripinnis + subbifrenatus + saponaceus + bicolor + courtenayi and suffered reversion in maculatus. Rypticus subbifrenatus + R. saponaceus + R. bicolor + R. courtenayi (17') SUBMANDIBULAR AND PREOPERCULAR PORES SPLIT INTO NUMEROUS SMALLER ONES IN ADULTS.—As described by Courtenay (1967) and McCarthy (1979), the four submandibular and four preopercular pores of the lateral-line system of R. subbifrenatus, saponaceus, bicolor and courtenayi split into numerous tiny terminals with growth (Fig. 2C). Within other Grammistini, the presence of numerous small pores was observed only in Aporops. Rypticus subbifrenatus (12) Presence of dark rounded spots.—See discussion above. Rypticus saponaceus + R. bicolor + R. courtenayi (18) INCREASE IN NUMBER OF PYLORIC CAECA.—In Grammistes and in all Rypticus except maculatus, saponaceus, bicolor and courtenayi there are typically three pyloric caeca. In maculatus there are five, in saponaceus four and in bicolor and courtenayi either four or five (Table 3). In the most parsimonious scenario, an increase in the number of pyloric caeca occurred independently in maculatus and in saponaceus + bicolor + courtenayi. Rypticus bicolor + R. courtenayi (19) PRESENCE OF THREE DARK STRIPES IN THE JUVENILE STAGE.—McCarthy (1979) reported that juveniles of R. bicolor with about 20 mm SL have three dark longitudinal stripes beginning just posterior to the orbit and reaching posteriorly to the caudal peduncle. He also noticed that juveniles of R. courtenayi (with about 35–40 mm SL) sometimes show vestiges of darkened bands immediately posterior to the eye, and suggested these color features are homologous for both species. Such pattern of juvenile pigmentation has not been described in any other grammistin.

DISCUSSION

Although there has been no previous attempt to cladistically test monophyly of Rypticus, the group has long being recognized as a distinct lineage by systematists. All previous proposed subdivisions of the group into more than one genus were proved inconsistent (see Courtenay, 1967: 243). In the present study, ten synapomorphies (three putative) are hypothesized to support monophyly of Rypticus. Among these, seven (one putative) are modifications in axial skeleton structure which are probably closely related. Partially because it includes a different set of terminal taxa and is based on the cladistic methodology, the analysis presented herein differs largely from previous hypotheses regarding intrageneric relationships of Rypticus, i.e., those of Courtenay (1965: fig. 27; also 1967—relationships discussed in the text) and McCarthy (1979). Nevertheless, it roughly agrees with those studies in recognizing R. bornoi and bistrispinus as sister- group taxa, randalli and maculatus as somewhat closely related and nigripinnis as closer to other Atlantic congeners rather than eastern Pacific ones. The tree (Fig. 9) resulting from the cladistic analysis of a matrix of polarized character states (Table 11) is the single most parsimonious one when characters are treated as unordered (or non-additive). For instance, if it is assumed that ordering multistate characters 368 BULLETIN OF MARINE SCIENCE, VOL. 65, NO. 2, 1999

(e.g., based on a priori observation of similarity or ontogenetic adjacency) is a sounder procedure, then two equally parsimonious trees would result: one which has the same topology of that illustrated in Figure 9 and one in which R. maculatus is placed as an immediate outgroup to (randalli + nigripinnis) + (subbifrenatus + saponaceus + bicolor + courtenayi). Because maculatus, randalli and nigripinnis share is a reversion of a character that defines the whole Serranidae among other percoids (i.e., loss of the lowermost opercular spine, see Gosline, 1966; Johnson, 1983), while all other characters that define monophyletic groups within Rypticus are not unique within the family, placement of maculatus as an outgroup to (randalli + nigripinnis) seems more informative. That indicates ordering multistate characters would decrease the accuracy of this analysis. It is also worth noting that some of the monophyletic groups indicated in Figure 9 also agree in some other aspects, which cannot be clearly polarized based on available information, but are thought as extra indications of close relationships. For example, both Rypticus bistrispinus and bornoi have the distal extremities of impaired fins weakly pig- mented, show a preference for insular habitats (sensu Robins, 1971), have reduced sizes (61 and 121 mm SL the largest known specimens of each vs 150 to about 300 in the other Rypticus), and present a typical configuration of two dorsal-fin spines and 25 to 26 dorsal-fin rays. One single character does not support the idea that both are sister-group taxa and that is the presence of dark spots against a light background (character 12), a derived character within the Grammistini which is shared by R. bornoi and subbifrenatus. On another group, both Rypticus maculatus, nigripinnis and randalli show a preference for coastal habitats and do not overlap geographically (Fig. 6), a fact that could indicate recent vicariance. Rypticus nigripinnis and randalli also share an elongate body with the shortest modes for snout length (modally 6% SL vs 7 in others) and upper jaw length (modally 14 % SL vs 15 to 18 in congeners). Aside from being almost indistinct meristi- cally, Rypticus saponaceus, bicolor and courtenayi also share similar body shape and coloration and show a preference for open ocean habitats.

BIOGEOGRAPHY

According to Brown and Gibson (1983), biogeography’s last frontier is to enable reconstruction of sequences of events that generate endemisms in marine organisms. In Rypticus, as well as in other groups with a planktonic larval stage, overlapping patterns of distribution caused by their strong capacity of dispersion constrain such reconstruction. Never- theless, four of the cladogenetic events hypothesized above relate geographic isolated taxa, and can be associated with three known “Earth’s history-events” (sensu Rosen, 1990; i.e., the combination of paleogeography, tectonic, eustactic, climatic and oceanographic events): CLOSING OF THE TETHIAN SEAWAY.—Rypticus is a monophyletic group distributed along the continental shelves and adjacent oceanic islands of the tropical Atlanto-East Pacific (Figs. 3,4,6,8). Since other monophyletic groups of organisms are limited to the same area, it has been recognized as a generalized biogeographic track, the east Pacific–east Atlantic track (Rosen, 1976). With the exception of Jeboehlkia, which occurs only in the Caribbean area (Baldwin and Johnson, 1991), and Pseudogramma, absent only in the tropical eastern Atlantic (Myers, 1989; Randall, 1983), the remaining Grammistini are restricted to the Indo-West Pacific (Myers, 1989; Randall, 1986; Randall et al., 1990). GUIMARÃES: REVISION OF RYPTICUS 369

Other examples of monophyletic Atlanto-East Pacific percoid groups with their immediate sister-groups in the Indo-Pacific are the jacobus group of the holocentrid genus Myripristis (Greenfield, 1968) and the brasiliensis group of Spanish mackerels (Collette and Russo, 1985). The most parsimonious explanation to such congruent patterns is that in which the ancestor of each Atlanto-East Pacific group became isolated in the end of the tertiary, when the faunas of the Indo-West Pacific and Atlanto-East Pacific were sepa- rated by the closing of the Tethian seaway (Ekman, 1967). If this phylogeny of Grammistini genera (Fig. 9) is accepted, then the presence of Jeboehlkia in the New World must be attributed to secondary dispersion. If a different topology in the phylogeny of the Grammistini is assumed (i.e., that cited by Baldwin and Johnson (1993) in which soapfishes and pseudogrammids are each regarded as monophyletic), then secondary dispersion is avoided as an explanation for any current distribution of grammistin genera. RISE OF THE PANAMA ISTHMUS.—This event was completed around 2.4 mya in the beginning of the Pleistocene (Cox and Moore, 1993) and has been recognized as a major cause of divergence in lineages of marine organisms in the tropical New World (Ekman, 1967; Rosen, 1976; Vermeij, 1978). In Rypticus, such event is intuitively associated with two of the hypothesized cladogenetic events, namely the differentiation between R. randalli and R. nigripinnis, two sister species isolated by the isthmus (Fig. 6), and between R. saponaceus and R. bicolor + R. courtenayi, two monophyletic sister-groups also isolated by the isthmus (Fig. 8). DISPLACEMENT OF THE PACIFIC TECTONIC PLATE.—Rypticus bicolor is distributed over the coastal portion of the tropical eastern Pacific, also present in the Cocos and Galápagos Islands, while its sister species, R. courtenayi, is restricted to the Revillagigedo Islands (Fig. 8). Unlike the Cocos and Galápagos Islands, the Revillagigedo archipelago is an- chored on the Pacific tectonic plate, which is drifting northwestward (Springer, 1982), thus separating from the American coast. It seems likely that such displacement caused the ancestor of both species to become isolated at some point in time. Conversely, a dispersalist explanation would suggest that larvae of R. bicolor eventually reached the Revillagigedo archipelago and differentiated into R. courtenayi according to the “Founder- effect” model of allopatric speciation (Bush, 1975). Among the 16 species of the circumtropical genus Myripristis (squirrelfish), Greenfield (1968) recognized a lineage in which M. jacobus (Atlantic) is the sister group of M. leiognathus (eastern Pacific) + M. clarionensis (Clipperton and Revillagigedo Islands). Although Greenfield (1968) provided no basis for such hypothesis (there are no shared characters listed), the resulting area cladogram is congruent with that observed for Rypticus saponaceus + bicolor + courtenayi (see Figs. 8,9), except for the supposed absence of Rypticus from the Clipperton Islands. It is also noteworthy that both Rypticus maculatus and Scomberomorus maculatus (a mackerel of the regalis group; see Collette and Russo, 1985) have a Carolinian distribution and a sister-group relationship with a tropical eastern Pacific-western Atlantic monophyletic clade. Such area cladogram cannot be explained by means of those traditionally known events, and supports the idea that detailed biogeographic analysis based upon groups of reef fishes await a refinement in the knowledge about the “Earth’s history” as well as in the knowledge about phylogenetic relationships of such groups, especially at the species level. 370 BULLETIN OF MARINE SCIENCE, VOL. 65, NO. 2, 1999

ACKNOWLEDGMENTS

I wish to especially thank my advisor, W. J. E. M da Costa, who was a major source of inspiring discussions during this research. For also easing this endeavor I thank my colleagues at the Laboratório de Ictiologia Geral e Aplicada: F. Autran, F. Bockmann, R. C. da Paz, A. Sarraf, M. Britto, C. A. Rangel and A. C. Bacellar. A significant part of this study was conducted at the Museum of Com- parative Zoology, and I wish to thank K. F. Liem, J. Craddock, L. Kaufmann, E. Drucker, Kristen, John, Miriam and especially Karsten E. Hartel for their kindness and extensive help. The following people provided all the necessary assistance and materials during my visits to their institutions and/ or relevant information to this study: G. W. Nunan, D. F. de Morais Jr., J. L. de Figueiredo, O. Oyakawa, R. L. de Moura, S. L. Jewett, K. Murphy, S. Raredon, D. Smith, J. Finan, J. Williams, M. Stiassny, N. Feinberg, R. Arrindel, B. Chernoff, S. Schaefer, W. R. Courtenay Jr., W. G. Saul, E. Böhlke, K. M. Grosser, Z. M. S. de Lucena and M. A. Bemvenuti. I also thank G. W. Nunan, M. P. Paiva, R. Lavenberg, R. L. de Moura and W. R. Courtenay, Jr., for the loan of additional relevant material. T. C. Ramos kindly provided a copy of his Tree Gardener software and assisted in some technical details. The manuscript benefited from the criticisms of W. R. Courtenay. Jr., and C. R. Robins. This study is a condensed version of a thesis, completed in partial fulfillment of the re- quirements for a Masters degree at the Universidade Federal do Rio de Janeiro, Museu Nacional. Research was financed by CNPq (Conselho Nacional de Desenvolvimento Científico e Tecnológico — Brazilian Federal Government).

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Wiley, E. O. 1981. Phylogenetics—The theory and practice of phylogenetic systematics. John Wiley and Sons, New York.

DATE SUBMITTED: February 4, 1998. DATE ACCEPTED: April 4, 1998.

ADDRESS: Laboratório de Biodiversidade de Recursos Pesqueiros, Núcleo de Inovação em Gerenciamento Pesqueiro (NIGP), UFRJ, Cidade Universitária, CEP 21941-569, Rio de Janeiro, RJ, Brazil. Phone: (021) 280-2394 ext. 31, e-mail: .

APPENDIX

MATERIAL EXAMINED

OUTGROUPS: Aporops bilinearis: AMNH 30905 (3), AMNH 51743 (1), AMNH 51762 (2), AMNH 72652 (2), MCZ 37301 (1), USNM 218920 (3, cs, 39, 43, 58). Jeboehlkia gladifer: USNM 330430 (1, cs, 52), USNM 201422 (1, r, 40, holotype). Grammistes sexlineatus: USNM 218886 (1, cs, 68), MCZ 09758 (5), AMNH 58205 (5), AMNH 31435 (1), AMNH 37873 (1), AMNH 18439 (2), MCZ 09759 (2), AMNH 27018 (1), MCZ 24486 (1), MCZ 05985 (2), MCZ 03652 (3, r, 81, 85, 127), AMNH 18041 (2), MCZ 09760 (1), MCZ 09761 (4), AMNH 89993 (1), AMNH 90023 (10), AMNH 89839 (3), MCZ 01073 (1), MCZ 65224 (2). Grammistops ocellatus: USNM 218873 (3 [1 cs, 86], [2, r, 56, 93] ), USNM 260562 (2). Pogonoperca punctata: USNM 205491 (1, r, 163), AMNH 38198 (1), MCZ 05933 (1, r, 195), MCZ 25771 (1, r, 158). Pseudogramma gregoryi: AMNH 31219 (12), AMNH 34111 (46), AMNH 27294 (5), AMNH 25922SW (1 cs, 34), AMNH 33016 (2), MCZ 47402 (3), MCZ 45688 (4). Pseudogramma polyacantha: USNM 209575 (3, cs, 24, 31, 41), MCZ 46976 (1), MCZ 12838 (7), AMNH 72481 (1), AMNH 51733SW (1, cs, 50), AMNH 71694 (1), AMNH 72664 (3), AMNH 72245 (5), AMNH 72649 (13), AMNH 18038 (1), MCZ 99270 (1), AMNH 58183 (2), MCZ 65240 (1), MCZ 51455 (3), AMNH 18132 (1), AMNH 15563 (1), AMNH 50557 (2), AMNH 50537 (1), AMNH 50551 (1). Pseudogramma thaumasium: AMNH 16035 (1), AMNH 16025 (2), MCZ 29409 (1), MCZ 52734 (1), MCZ 44588 (15), MCZ 44589 (2), MCZ 65155 (2), MCZ 45681 (4), MCZ 44587 (12), AMNH 73356 (5), MCZ 101611 (1), MCZ 45685 (6), MCZ 45682 (1), AMNH 17160 (3). Suttonia lineata: USNM 209705 (2 [1 r, 36], [1 cs, 38] ). Suttonia sp: MCZ 101609 (1). Suttonia suttoni: USNM 285959 (2, r, 52, 60). Rypticus bicolor (138 specimens, 18–195 mm SL): COLOMBIA: USNM 330424 (8), USNM 330403 (1), Bahía Utría. USNM 330420 (1), USNM 330387 (3), Isla Gorgona. COSTA RICA: FMNH 61492 (1), Golfo de Nicoya. Equador: MCZ 40463 (5), MCZ 40466 (1), Isla Baltra. GALÁPAGOS: FMNH 41295 (1), FMNH 41698 (1). FMNH 41787 (1), FMNH 41788 (1), ANSP 85987 (9), USNM 321619 (1), USNM 94010 (1). COCOS ISLAND: AMNH 08533 (1). MEXICO: Baja California Sur: USNM 300740 (1, holotype of Rhypticus xanti), Cabo San Lucas. AMNH 05489 (1), Isla Carmen. USNM 321624 (1), Los Frailes. USNM 321621 (1), Pichilingue. USNM 321622 (2), Puerto Chileno. Guerrero: MCZ 2842 (1), FMNH 88034 (2), ANSP 87880 (1), Acapulco. FMNH 88274 (3), Bahía de Puerto Marquez. Sinaloa: USNM 321623 (13), Isla San Ignacio de Farallon. Sonora: USNM 330382 (1), Bahía Choya. PANAMA: ANSP 13231 (1), no data. AMNH 73349 (1), MCZ 42616 (1), MCZ 52651 (3), MCZ 45692 (1), MCZ 45544 (1), MCZ 45842 (1), ANSP 84997 (1), Arquipelago de Las Perlas. ANSP 120891 (5), Bahía Pinas. MCZ 52656 (26, [2 cs, 66, 80] ), MCZ 52657 (5), MCZ 52658 (3), UFRJ 3062 (2), Isla Coiba. MCZ 45609 (1), Isla Taboga. MCZ 58188 (1), MCZ 82764 (1), USNM 321618 (2), USNM 321616 (2), Isla Toboguilla. USNM 321617 (1), Isla Urava. MCZ 45607 (9, [1 cs, 81] ), Ilhote Valladolid. ANSP 79450 (1), Monte Sapo. USNM 321620 (1), Peninsula Azueiro. USNM 321318 (1), Punta Mariato. FMNH 35618 (3), Recife de San Francisco. PERU: USNM 127971 (1), Bahía de Lobos de Afuera. Rypticus bistripinus (180 specimens, 14–117 mm SL): BAHAMAS: ANSP 100962 (1), Andros. GUIMARÃES: REVISION OF RYPTICUS 375

AMNH 30331 (3), AMNH 30334 (1), AMNH 30336 (3), AMNH 30339 (1), AMNH 30382 (1), AMNH 21242 (4), AMNH 26174 (4, 2 r), AMNH 26186 (2), AMNH 14892 (1), AMNH 14896 (1), AMNH 14890 (4, [1, cs, 87] ), Bimini Islands. ANSP 100950 (5), Exuma Cays. ANSP 100969 (1), ANSP 111469 (1), Grand Bahama. USNM 53137 (1), Green Cay. ANSP 100813 (2), ANSP 100945 (2), Hog Island. ANSP 100949 (2), New Providence Island. ANSP 113422 (2), Rose Island. ANSP 100944 (12), ANSP 100961 (1), Silver Cay. BELIZE: FMNH 95451 (1), Salt Creek. BERMUDA: FMNH 48759 (2). BRAZIL: off Amapá: FMNH 69424 (1). Bahia: MZUSP 46382 (1), Arembepe. Rio de Janeiro: UFRJ 2204 (2, cs, 57, 82), UFRJ 3041 (13), MNRJ 12822 (3), MZUSP 47483 (2), Arraial do Cabo. CUBA: USNM 37493 (1), no data. USNM 330390 (1), Cayo Coco. USNM 188217 (1), Havana. GULF OF MEXICO: AMNH 85440 (1), AMNH 85149 (1), AMNH 85605 (1), AMNH 85796 (1), AMNH 85771 (1), no data. FMNH 46440 (1), Campeche Bay. USNM 131580 (1), Port Charlotte. USNM 23555 (1, holotype de Rhypticus pituitosus), AMNH 82920 (1), AMNH 82800 (1), AMNH 85370 (1), AMNH 82035 (2), AMNH 82316 (3), AMNH 82172 (1), AMNH 81053 (1), AMNH 84997 (1), MCZ 28111 (1), FMNH 61289 (1), FMNH 61300 (2), FMNH 91484 (1), FMNH 45483 (1), FMNH 45484 (4), USNM 330410 (1), USNM 130807 (1), USNM 330405 (2), USNM 133907 (1), USNM 131609 (1), USNM 330411 (1), USNM 330406 (2), USNM 158241 (1), USNM 43559 (1), USNM 330414 (2), USNM 188228 (1), USNM 148860 (1), off Florida. FMNH 59890 (8), Florida Keys. AMNH 84076 (1), Florida Middle Ground. USNM 117194 (2), Tortugas. ISLANDS VIRGIN: USNM 183601 (1), Eustatia Island. PANAMA: ANSP 45275 (1), Colon. PUERTO RICO: FMNH 3261 (1), FMNH 52050 (2), FMNH 69425 (1), USNM 50208 (11), USNM 126189 (2), Isla Culebra. UNITED STATES ATLANTIC COAST: MCZ 101612 (1), MCZ 46644 (1, r, 78), AMNH 76941 (4), MCZ 90071 (3, [1, cs, 85] ), MCZ 39527 (1), FMNH 65572 (3), FMNH 69423 (1), USNM 330402 (1), off Florida. AMNH 76998 (1), MCZ 59538 (1, r, 105), USNM 315676 (1), off Georgia. USNM 315535 (1), off North Carolina. AMNH 77197 (1), AMNH 76987 (1), AMNH 76985 (1), MCZ 90072 (1), off South Carolina. Rypticus bornoi (75 specimens, 34–61 mm SL): BAHAMAS: AMNH 34471 (1), Cat Island. AMNH 24053 (1), USNM 201408 (1, r, 45, paratype of R. macrostigmus), Eleuthera Island. ANSP 100968 (1, holotype of R. macrostigmus), Wood Cay. BELIZE: FMNH 95453 (1), Barrier Reef. HAITI: ANSP 111385 (5), ANSP 114415 (10), ANSP 117941 (6), ANSP 119043 (4), ANSP 120546 (4), Gulf of Gonave. ANSP 114563 (28 [2 cs, 45, 48] ), USNM 170572 (1, r, 49, holotype), UFRJ 3061 (10), Port-au-Prince Bay. HONDURAS: FMNH 95668 (1), Northwest Key. PANAMA: MCZ 43620 (1, r, 53, paratype of R. macrostigmus), Shepherd Island. Rypticus courtenayi (20 specimens, 62 - 160 mm SL): REVILLAGIGEDO ISLANDS: LACM 32097-19 (10, [2 cs, 70, 85], [6 r, 80, 93, 109, 120, 147], all paratypes), Isla Clarion. ANSP 136546 (3, paratypes), USNM 218386 (7, paratypes), Isla Socorro. Rypticus maculatus (44 specimens, 65–201 mm SL): BAHAMAS: FMNH 50345 (1), no data. GULF OF MEXICO: AMNH 87024 (1), AMNH 86976 (1), AMNH 85606 (1), AMNH 79558SD (1, ds, 155), AMNH 92557SD (1, ds, 145), AMNH 94639SD (1, ds, 150), AMNH 94829SD (1, ds, 105), no data. AMNH 55824SD (1, ds, 125), Alabama. USNM 158229 (2), USNM 330404 (1), off Alabama. USNM 118949 (1), Cedar Keys. MCZ 39527 (1), off Florida. FMNH 45485 (1), off Laguna Madre, Mexico. USNM 124302 (1), North Key. USNM 188229 (2), off Sarasota. AMNH 85206 (1), FMNH 64196 (1), USNM 158236 (1), off Texas. UNITED STATES ATLANTIC COAST: USNM 21544 (1), Charleston. AMNH 76943 (1), USNM 106514 (2), USNM 142916 (2), off Florida. AMNH 77120 (1), USNM 190337 (1), USNM 315766 (2), USNM 315518 (3), off Georgia. USNM 155190 (1), off North Carolina. USNM 62695 (1), Palm Beach. AMNH 77185 (1), AMNH 76903 (1), AMNH 77089 (1), AMNH 77122 (1), AMNH 77225 (2, [1, cs, 85] ), AMNH 76169 (2), off South Carolina. Rypticus nigripinnis (350 specimens, 10–180 mm SL): COLOMBIA: USNM 330409 (7), Bahía Utría. USNM 218870 (1, cs, 60), USNM 330418 (1), USNM 218870 (2), Bocana. USNM 330429 (22), N of Buenaventura. USNM 330400 (1), USNM 330412 (1), off Cabo Manglares. COSTA RICA: FMNH 61494 (1). Puntarenas Estero. EL SALVADOR: FMNH 93636 (9), FMNH 93637 (8), FMNH 93635 (11), USNM 220801 (10), USNM 220782 (5), Bahía de Jiquilisco. MEXICO: 376 BULLETIN OF MARINE SCIENCE, VOL. 65, NO. 2, 1999

Baja California Sur: USNM 3689 (1, holotype de R. maculatus Gill), Cabo San Lucas. Guerrero: FMNH 72300 (4), Acapulco. FMNH 72301 (1), Bahía de Acapulco. FMNH 88020 (2), Bahía de Puerto Marquez. PANAMA: USNM 270278 (1, cs, 35), MCZ 28784 (1), MCZ 2843 (1, r, 158), FMNH 20575 (1), FMNH 20577 (3), FMNH 20578 (1), FMNH 20579 (1), FMNH 20581 (3), FMNH 20582 (1), FMNH 20583 (1), USNM 30961 (1, holotype of Promicropterus decoratus), USNM 3700 (1, holotype), no data. MCZ 45674 (2), MCZ 45661 (3), MCZ 52653 (1), MCZ 52652 (1), Arquipelago Las Perlas. USNM 321629 (2), Bahía Honda. ANSP 70203 (1), Bahía de San Miguel. FMNH 20576 (1), FMNH 20580 (1), ANSP 77179 (2), Balboa. MCZ 45625 (17, [1, cs, 63]), USNM 321625 (8), USNM 321630 (10), Panama City. USNM 80232 (1), Corozal. USNM 321631 (4), El Rompio. MCZ 42614 (1), MCZ 42615 (1), Fort Amador. MCZ 45541 (5) MCZ 45838 (2), MCZ 45700 (1), Isla Culebra. USNM 321628 (4), USNM 321626 (2), Isla Toboguilla. USNM 321627 (11), Isla Urava. ANSP 84996 (2), Playa Muerto. MCZ 52654 (3), Playa Venado. MCZ 42618 (6), Punta Bruja. AMNH 73384 (68), MCZ 42605 (2), MCZ 42606 (10), MCZ 42607 (6), MCZ 42608 (17, [1 cs, 75] ), MCZ 42609 (11), MCZ 42610 (4), MCZ 42611 (8), MCZ 42612 (3), MCZ 42613 (2), MCZ 42617 (2), FMNH 72323 (11), UFRJ 3063 (5), Punta Paitilla. MCZ 52655 (2), Punta Vique. FMNH 35618 (1), 35619 (1), 35620 (1), San Francisco Reef. PERU: USNM uncat. (2), Bahía de Lobos de Afuera. USNM 127970 (1), Gulf of Guayaquil. Rypticus randalli (174 specimens, 25–180 mm SL): BAHAMAS: AMNH 20624 (1), Little Inagua. BELIZE: USNM 300519 (2), Dangrica. FMNH 95439 (1), Placentia Town. FMNH 95455 (3), Punta Gorda. BRAZIL: MZUSP 48033 (2), USNM 43278 (1, paratype of R. brachyrhinus), no data. MZUSP 48016 (1), Bahia and Espírito Santo coasts. Bahia: MZUSP 47994 (1), Maragogipe. MZUSP 47995 (2), Ilha de Itaparica MZUSP 48031 (3) Monte Serrat. Maranhão: MZUSP 48036 (2), São Luís. Rio de Janeiro: MZUSP 46311 (1), UFRJ 3040 (2), UFRJ 3039 (1), UFRJ 3635 (4), Baía da Ilha Grande. MZUSP 48034 (3), Cabo Frio. MNRJ 12821 (1), Baía de Guanabara. ANSP 121163 (1, r, 30), Manguinhos. MNRJ 8744 (1), MNRJ 8745 (1), MNRJ 8746 (1), MNRJ 8747 (1), MNRJ 8748 (1), MNRJ 8749 (1), MNRJ 8750 (1), MNRJ 8751 (1), MNRJ 8752 (1), MNRJ 8753 (1), MNRJ 8754 (1), Praia de Sepetiba. UFRJ 3037 (1), UFRJ 3038 (2), MNRJ 12823 (1), Praia Vermelha. Santa Catarina: MZUSP 46648 (2), Porto Belo. São Paulo: MZUSP 48028 (3), MZUSP 48029 (1), MZUSP 48030 (1), Cananéia. MZUSP 48035 (1), Ubatuba. Sergipe: MZUSP 48032 (5), Aracajú. MZUSP 48027 (1), Chica Chaves. CENTRAL AMERICA: USNM 44716 (1, paratype of R. brachyrhinus). COLOMBIA: USNM 330423 (3), Isla Baru. USNM 330422 (2), Isla Cabruna. DOMINICAN REPUBLIC: USNM 314453 (1 r, 93), Santo Domingo. GUIANA FRANCESA: FMNH 69426 (1). HAITI: USNM 330316 (2), USNM 330317 (4), no data. AMNH 18931 (1), AMNH 18976 (1), AMNH 18995 (2), ANSP 87100 (1, paratype), ANSP 77203 (2), ANSP 81865 (1), ANSP 83057 (1), ANSP 83090 (2), ANSP 83668 (1), ANSP 83886 (6), ANSP 83893 (2), USNM 132544 (1, paratype), Port-au-Prince. HONDURAS: USNM 44468 (1, paratype of R. brachyrhinus), Patuca River. JAMAICA: ANSP 31707 (2, paratypes), FMNH 2824 (2), USNM 30130 (1, paratype). PANAMA: AMNH 73280 (1), Bocas del Toro. Canal Zone: USNM 80234 (1, r, 122, holotype of R. brachyrhinus), Mindi Cut. USNM 106625 (6, paratypes of R. brachyrhinus). SE of Almirante: MCZ 44413 (2, r, 85, 86), Ambrosia Bight. MCZ 44414 (2), Jungle Point. MCZ 44415 (4, [1 cs, 61] ). PUERTO RICO: MCZ 106929 (1, cs, 103), no data. FMNH 61495 (1), USNM 197917 (1, r, 127, holotype), USNM 126136 (1, paratype), Bahía de Mayagüez. USNM 50188 (1, paratype), Hucares. FMNH 61489 (1), Punta Arena. ANSP 100970 (2, paratypes), Tres Hermanos. TRINIDAD & TOBAGO: ANSP 44980 (1, paratype), Gulf of Paria. VENEZUELA: USNM 121809 (1, r, 115), Caño de Sagua. ANSP 101603 (1, paratype), Península de Araya. LOCALITY UNKNOWN: FMNH 69427 (1). Rypticus saponaceus (406 specimens, 12–285 mm SL): ANTIGUA & BARBUDA: ANSP 114404 (2), Antigua. AMNH 47865 (1), Leeward Island. ASCENSION ISLAND: USNM 330383 (1), USNM 330384 (3), USNM 330417 (7), USNM 330419 (7), USNM 330421 (2), USNM 330426 (1). BAHAMAS: FMNH 50345 (1), no data. ANSP 100939 (1), ANSP 106350 (1), ANSP 106352 (6), Salt Cay Bank. AMNH 24176 (2 r, 63, 78), Fortune Island. AMNH 28767 (3), ANSP 100940 (3), ANSP 106353 (2), ANSP 100941 (2), Grand Bahama. ANSP 72419 (1), ANSP 72420 (1), ANSP GUIMARÃES: REVISION OF RYPTICUS 377

72421 (1), ANSP 72422 (1), ANSP 83582 (1), ANSP 83584 (3), ANSP 83586 (1), ANSP 83587 (1), ANSP 83591 (1), ANSP 83592, ANSP 83594 (4), ANSP 83599 (1), ANSP 115041 (7), ANSP 116282 (3), ANSP 126851 (2), Hog Island. ANSP 106351 (2), Hogsty Reef. ANSP 83596 (2), Balmoral Island. AMNH 22930 (2), AMNH 29970 (6, [1, cs, 86] ), AMNH 30013 (1), AMNH 34668 (2), Berry Island. AMNH 74706 (1), Bimini Islands. ANSP 83590 (2), Eleuthera Island. MCZ 2840 (1), ANSP 72420 (1), ANSP 83583 (3), ANSP 83585, (1), ANSP 83588 (2), ANSP 83593 (2), ANSP 83595 (4), ANSP 83597 (2), ANSP 83598 (7), New Providence Island. AMNH 21275 (2), Little Inagua. AMNH 22997 (2), Little San Salvador. AMNH 33239 (3), Mayaguana. ANSP 126851 (2), USNM 53136 (1), Nassau. AMNH 24992 (1), Orange Cay. ANSP 83582 (1), ANSP 113398 (3), Rose Island. ANSP 83589 (1), ANSP 115041 (7), Sandy Cay. AMNH 28695 (1), ANSP 106349 (6), Wood Cay. BARBADOS: ANSP 83647 (5), ANSP 83648 (5), USNM 330407 (1). BELIZE: FMNH 95443 (1), FMNH 95446 (2), FMNH 95467 (2), Glover’s Reef. FMNH 95450 (1), Northeast Cay. FMNH 95473 (1), Turneffe. BERMUDA: FMNH 5295 (1), FMNH 48169 (1), FMNH 48671 (1), AMNH 48993 (1), AMNH 39294 (3), ANSP 133304 (1), ANSP 133571 (1), ANSP 133619 (1) ANSP 133620 (2), ANSP 133703 (1), ANSP 139096 (1), ANSP 148243 (1), ANSP 148445 (1), ANSP 168548 (1), ANSP 168549 (2), ANSP 133572 (1), USNM 170057 (1), USNM 175760 (1), USNM 330427 (3), USNM 178604 (1). BRAZIL: Bahia: MZUSP 48026 (3), MZUSP 48019 (1), MZUSP 47993 (3), Arembepe. MZUSP 48018 (2), MZUSP 48017 (1), MZUSP 48025 (1), Ilha de Itaparica. MZUSP 48021 (1), MZUSP 48024 (1), MZUSP 48022 (2), MZUSP 48023 (3), Itapoã. MZUSP 48020 (1), Parque Interlagos. Espírito Santo: MZUSP 44672 (1), Guarapari. Rio de Janeiro: MNRJ 12820 (1), Baía de Guanabara. UFRJ 1289 (1), Ponta do Marisco. BRITISH VIRGIN ISLANDS: ANSP 83646 (1), Guana. CARIBBEAN SEA: FMNH 65585 (1), FMNH 74815 (1), USNM 314551 (12), Serrana Bank. CAYMAN ISLANDS: ANSP 102334 (1), ANSP 105104 (1), ANSP 121932 (2), Grand Cayman. COLOMBIA: MCZ 47446 (1), Bahía Gairaca, Santa Marta. CUBA: MCZ 21755 (2), no data. MCZ 2841 (1), Santa Cruz del Sur. DOMINICA: ANSP 120554 (1), ANSP 121459 (2), ANSP 126713 (10), USNM 330391 (8), USNM 330401 (1), USNM 330413 (1), USNM 330408 (3), USNM 330428 (2), USNM 330389 (5). GRENADA: ANSP 113326 (1), ANSP 113398 (3), ANSP 113911 (2), ANSP 52471 (2), ANSP 120548 (2), ANSP 121678 (2). HAITI: USNM 89654 (1), Bahía Petit Baraderes. ANSP 120857 (1), ANSP 120967 (1), Canal de Saint Marc. ANSP 91761 (1), Port-au-Prince. HONDURAS: FMNH 95480 (1), Half Moon Bay. FMNH 101105 (1), Roatan. JAMAICA: USNM 160725 (1), no data. ANSP 95636 (1), Lucae. LIBERIA: USNM 193658 (1), Mesurado R beach. NIGERIA: USNM 188458 (1, r, 215), off Lagos. PANAMA: FMNH 61493 (1), Isla Cuili. MCZ 42637 (1), MCZ 42638 (1), MCZ 42639 (1), MCZ 42640 (1), MCZ 42641 (1), MCZ 42642 (2), MCZ 52733 (1), Isla Galeta. MCZ 44416 (1), Playa Blanca. MCZ 45668 (1), MCZ 45673 (1), San Lorenzo. PUERTO RICO: USNM 330416 (1), USNM 330394 (3), Boca de Cangrejos. FMNH 62296 (4), FMNH 61490 (3), FMNH 61541 (13), Guanica. ANSP 70987 (1), ANSP 145833 (10), Isla Mona. ANSP 118430 (2), ANSP 115556 (2), ANSP 115646 (1), ANSP 118714 (1), Puerto Yabucoa. SAINT LUCIA: ANSP 124701 (1), Anse des Pitons. ANSP 112812 (6), ANSP 117986 (4), ANSP 121663 (3), ANSP 122702 (1), Pigeon Island. SAINT VINCENT: ANSP 113223 (1), ANSP 121460 (1), Anse Mahaut. ANSP 121946 (1), ANSP 121952 (1), Bequia Island. ANSP 120550 (2), Canouan Island. ANSP 121498 (3), Little St. Vincent Island. SANTA HELENA ISLAND: USNM 267872 (1, r, 150). SOMBRERO ISLAND: MCZ 2849 (3), MCZ 2915 (3). TRINDADE ISLAND: MNRJ 1772 (3). TRINIDAD & TOBAGO: ANSP 97959 (1), USNM 318542 (7), USNM 318522 (1), USNM 318536 (3), Tobago. UNITED STATES: Florida: USNM 167604 (1), Biscayne Bay. USNM 198139 (1, neotype of Anthias saponaceus), Key Biscayne. ANSP 32676 (1), ANSP 74875 (1), Key West. U. S. VIRGIN ISLANDS: ANSP 17111 (1), Saint Croix. VENEZUELA: USNM 194112 (1), USNM 179260 (4), Los Roques. LOCALITY UNKNOWN: MCZ 106931 (1, cs, 75). Rypticus subbifrenatus (652 specimens, 12–150 mm SL): BAHAMAS: ANSP 149010 (1), Acklins Island. ANSP 100952 (1), Andros. ANSP 100942 (1), Athol Island. AMNH 35302 (1), Great Stirrup Bay. ANSP 106355 (2), Conception Island. AMNH 29258 (2), ANSP 106354 (1), Crooked Island. ANSP 100943 (3), ANSP 100948 (5), ANSP 111917 (4), Eleuthera. AMNH 28766 (1), AMNH 378 BULLETIN OF MARINE SCIENCE, VOL. 65, NO. 2, 1999

28768 (2), ANSP 100946 (2), ANSP 100955 (2), ANSP 106357 (1), Grand Bahama. AMNH 21347 (3), AMNH 27167 (1), Great Inagua. ANSP 100951 (1), ANSP 100966 (1), ANSP 106356 (1), ANSP 100964 (1), ANSP 147170 (1), ANSP 121746 (2), Green Cay. ANSP 72423 (1), ANSP 72626 (1), Hog Island. AMNH 27352 (6), ANSP 106358 (2), Hogsty Atoll. AMNH 33101 (1), Little Inagua. ANSP 121759 (1), ANSP 121802 (3), Mayaguana. ANSP 143266 (1), Nassau. ANSP 72627 (1), ANSP 72628 (3), ANSP 100947 (1), ANSP 100954 (2), ANSP 100956 (1), ANSP 100957 (1), ANSP 100959 (1), ANSP 100960 (1), ANSP 100963 (1), ANSP 100965 (1), New Providence. ANSP 100953 (2), North Cay. ANSP 121741 (2), ANSP 121817 (3), ANSP 121753 (1), AMNH 28488 (2), AMNH 28434 (1), AMNH 29224 (1), Plana Cays. ANSP 115029 (1), Sandy Cay. ANSP 100967 (1), Wood Cay. BELIZE: FMNH 95470 (2), FMNH 95471 (1), FMNH 95810 (1), Ambergris Cay. FMNH 95475 (3), FMNH 95463 (2), FMNH 90188 (1), FMNH 90186 (3), FMNH 90187 (1), FMNH 90176 (1), FMNH 90177 (1), FMNH 90182 (1), FMNH 90184 (1), FMNH 90185 (2), FMNH 95442 (1), USNM 276233 (1), USNM 276231 (1), USNM 321041 (1), Carrie Bow Cay. FMNH 95465 (1), FMNH 95466 (3), Curlew Cay. FMNH 90174 (1), FMNH 90175 (1), Gallow’s Point. FMNH 95445 (2), FMNH 95446 (1), FMNH 95447 (2), FMNH 95448 (3), FMNH 95444 (2), FMNH 95456 (1), FMNH 95457 (2), FMNH 95458 (2), FMNH 95459 (1), FMNH 95460 (1), FMNH 95461 (3), 95462 (1), FMNH 95468 (1), FMNH 95469 (1), FMNH 90180 (1), FMNH 90183 (3), FMNH 90178 (2), FMNH 90181 (1), FMNH 90179 (1), Glover’s Reef. FMNH 95452 (1), Goff’s Cay. FMNH 95454 (1), Punta Gorda. FMNH 95464 (1), FMNH 95487 (2), FMNH 95488 (1), FMNH 95489 (3), FMNH 95490 (1), South Water Cay. FMNH 95665 (1), FMNH 95474 (1), FMNH 95472 (1), Turneffe. BERMUDA: ANSP 148246 (2). BONAIRE: MCZ 57195 (1). BRAZIL: Bahia: MZUSP 48012 (1), MZUSP 48014 (1), MZUSP 47996 (4), Arembepe. MZUSP 47997 (4), MZUSP 48013 (2), MZUSP 48015 (1), MZUSP 9110 (1), Itapoã. Rio de Janeiro: UFRJ 5038 (1), Arraial do Cabo. CARIBBEAN SEA: FMNH 65586 (2), Serrana Bank. CAYMAN ISLANDS: ANSP 102290 (2), ANSP 102349 (1), ANSP 102350 (1), ANSP 104997 (1), ANSP 105053 (1), ANSP 105075 (1), Grand Cayman. COLOMBIA: MCZ 47441 (1), Bahía Concha. MCZ 47408 (2, [1, cs, 67] ), MCZ 47347 (2), Bahía Gairaca. USNM 330397 (17), Ceycen. USNM 330398 (4), Isla Grande. ANSP 143145 (3), ANSP 146553 (2), Old Providence. USNM 330425 (2), USNM 330386 (1), USNM 330385 (7), Santa Marta. CUBA: USNM 82432 (1), San Antonio. DOMINICA: ANSP 112826 (7), ANSP 121845 (3), USNM 330392 (5), USNM 330393 (4), USNM 330381 (7), USNM 330396 (6). DOMINICAN REPUBLIC: USNM 314418 (1), Santo Domingo. GRENADA: ANSP 112856 (10), ANSP 112862 (4), ANSP 114554 (7), ANSP 121483 (13), ANSP 121483 (13). GUADELOUPE: ANSP 121415 (1), ANSP 121422 (1). GULF OF GUINEA: MCZ 45286 (1, r, 49), ANSP 109203 (1, r, 58), Fernando Poo Island. HAITI: ANSP 120560 (6), Port-au- Prince Bay. ANSP 112523 (2), ANSP 117953 (7), ANSP 119061 (9), ANSP 119155 (6), Gulf of Gonave. ANSP 120526 (1), Isla de Sable. FMNH 61488 (1), Petit Gonave. HONDURAS: FMNH 95481 (5), FMNH 95486 (7), Half Moon Bay. FMNH 95479 (1), ‘Calvin’s Crack’. FMNH 95440 (7), Caribbean Point Bite Bay. FMNH 95485 (1), Concha Cay. FMNH 95482 (1), Crab Wall. FMNH 95667 (2), French Harbor. FMNH 95664 (1), Big Hog Island. FMNH 95666 (1), Isla Guanaja. FMNH 95441 (1), Little Hog Island. FMNH 95476 (1), FMNH 95477 (1), Punta Royale. FMNH 95483 (1), FMNH 95484 (1), Reef House. FMNH 95478 (2), ‘Tortuga Reef’. JAMAICA: AMNH 74084 (1), AMNH 36452 (1), Discovery Bay. FMNH 72522 (1), Drunkenman Cay. MARTINIQUE: ANSP 112954 (2). MEXICO: ANSP 114912 (4), Quintana Roo. NIGERIA: USNM 330395 (1), Lagos. PANAMA: AMNH 73542 (1), Las Mulatas. FMNH 35621 (1), Bahía Limon. MCZ 44425 (1), Boca del Drago. AMNH 73250 (2), Bocas del Toro. MCZ 42629 (1), MCZ 42636 (6), MCZ 44417 (2), MCZ 44419 (3), MCZ 44420 (2), MCZ 44424 (13, [3 cs, 12, 14, 21] ), MCZ 45635 (9), MCZ 45653 (21, [3 cs, 28, 45, 66] ), MCZ 45660 (10), MCZ 45668 (1), MCZ 45678 (2), Fort Sherman. AMNH 36685 (1). FMNH 61429 (1), Isla Cuili. MCZ 42619 (3), MCZ 42620 (5), MCZ 42621 (2), MCZ 42622 (1), MCZ 42623 (5), MCZ 42625 (3), MCZ 42626 (1), MCZ 42627 (5), MCZ 42630 (1), MCZ 42631 (1), MCZ 42632 (2), MCZ 42633 (1), MCZ 42634 (1), MCZ 42635 (2), MCZ 44418 (1), MCZ 44421 (2), Isla Galeta. FMNH 61430 (3), FMNH 61491 (1), Islas San Blas. MCZ 44423 (1), Juan Chiquita. MCZ 44422 (3), Playa Blanca. MCZ 42624 (1), Porto Bello. GUIMARÃES: REVISION OF RYPTICUS 379

MCZ 42628 (1), Río Piedras. MCZ 45695 (4, r, 115, 117, 120, 125), San Lorenzo. PUERTO RICO: ANSP 145702 (1), ANSP 145852 (4). Isla Mona. ANSP 143823 (2), La Parguera. ANSP 118462 (2), Porto Yabucoa. SAINT LUCIA: ANSP 121664 (1), Anse des Pitons. ANSP 112462 (1), ANSP 113233 (5), ANSP 119019 (1), Pigeon Island. ANSP 114545 (2), Port Castries. SAINT THOMAS: USNM 106516 (1). SAINT VINCENT: ANSP 119025: (3), ANSP 121421 (2), Anse Mahaut. ANSP 119014 (3), ANSP 121679 (1), Bequia Island. ANSP 117996 (2), Little St. Vincent. TRINIDAD & TOBAGO: ANSP 97943 (1), ANSP 97947 (1), ANSP 97949 (1), USNM 318533 (5), USNM 318546 (3), USNM 318527 (4), USNM 318539 (6), USNM 318530 (1), USNM 318524 (1), USNM 318532 (1), USNM 318535 (2), USNM 318529 (1), USNM 318523 (2), Tobago. UNITED STATES, Florida: MCZ 106930 (1, cs, 80), no data. FMNH 46901 (1), Old Rhodes Key. VENEZUELA: USNM 194094 (1), USNM 179259 (2), Los Roques.

&lt;I&gt;Rypticus&lt;/I&gt; (Teleostei: Serranidae)

<I>Rypticus</I> (Teleostei: Serranidae)