BULLETIN OF MARINE SCIENCE, 65(3): 825–838, 1999

LARVAL AND JUVENILE DEVELOPMENT OF THE CUBBYU PAREQUES UMBROSUS WITH NOTES ON THE HIGH HAT LARVAE

G. Joan Holt and Cecilia M. Riley

ABSTRACT Larval and juvenile development of Pareques umbrosus is described from laboratory spawned and reared specimens. Eggs of P. umbrosus ranged from 1.08 to 1.38 mm diam- eter with a single, pigmented oil globule 0.23 mm diameter. Yolk-sac larvae averaged 2.82 mm NL and had heavy golden-green stellate chromatophores covering the trunk and tail in fresh specimens. Preserved larvae had scattered melanophores on the trunk, tail and yolk-sac. Transition to exogenous feeding occurred at 2.59 mm NL and melanistic pigmentation included a distinct band around the tail anterior to the tip of the notochord, another vertical swath from the dorsal finfold through the pectorals to the pelvic area, and a lateral stripe that extended from the snout through the eye to the cleithrum. Two preopercle spines appeared just prior to flexion at 4.67 mm SL. A continuous horizontal stripe the length of the trunk and tail first developed at about 8.16 mm SL. Metamorpho- sis to the juvenile stage occurred between 10.00 and 15.00 mm SL. Juveniles averaging 30.5 mm SL had a characteristic 3-stripe color pattern, and a darkly pigmented first dorsal with a white trailing edge. The first dorsal was 1.5 times longer than the body depth. The pigmented pelvic fin had white leading and trailing edges. All but the center longitudinal body stripe were narrower than the pupil in juveniles with 4 to 6-stripes. All stripes were narrower than the pupil when the adult 7-stripe pattern was developed by 41– 67 mm SL. Lab reared P. acuminatus larvae of known-age were distinguished from P. umbrosus by the presence of three distinct pre-opercular spines in flexion larvae, con- spicuous white leading and trailing edges on the first dorsal, and in juveniles by the extremely long first dorsal (~75% SL).

At least 21 of sciaenids occur along the U.S. Atlantic and Gulf coasts many of which are important in recreational and commercial fisheries. Growth and development of young are well documented in most species with the exception of the reef species (Ditty, 1989). These small sciaenids associated with natural and artificial reefs and rocky substrates, are commonly found in small groups from Bermuda and South Carolina to Rio de Janeiro (Randall, 1983). Bloch and Schneider, according to Chao (1976), described nine species of western Atlantic in 1801, three of which are included in the reef group: Eques americanus (now Equetus lanceolatus), Eques (now Equetus) punctatus and Grammistes (now Pareques) acuminatus. Jordan and Evermann (1898) divided Eques into two genera Eques and Pareques. McPhail (1961) justified division of the reef group into two genera Equetus and Pareques based on the presence or absence of free interneurals anterior to the first dorsal fin, and the relative development of the first dorsal fin. Using these char- acters, he concluded that E. punctatus and E. lanceolatus, which have no free interneurals and possess long and filamentous first dorsal fins belong in the Equetus. The other specimens which possessed free interneurals were placed in the genus Pareques. Chao (1976) retained the two genera and distinguished two species of Pareques by: 3 to 5 broad longitudinal stripes wider than the pupil, and a straight inter-orbital bar in P. acuminatus,

825 826 BULLETIN OF MARINE SCIENCE, VOL. 65, NO. 3, 1999

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Ssseicep DlorsalAlnaPcectoraPlelviPrincipalCauda EquetuslanceolatusXIII-XIV,46−50Ι5Ι,61−615Ι8, 9+ EquetuspunctatusXII-XIII,45−47Ι−8Ι,71−8 715Ι8, 9+ ParequesacuminatusX-XI,36−41Ι−8Ι,71−7 615Ι8, 9+ ParequesiwamotoiXI,38Ι7Ι,715Ι9,9+8− ParequesumbrosusIX−XI,38−40Ι−8Ι,7∗ 51∗ Ι85, 9+ *thisstudy and longitudinal stripes narrower than the pupil, and a v-shaped interorbital bar in P. umbrosus. Taxonomic treatment by Darovec (1983) further separated Equetus from Pareques by differences in dorsal spine counts (13–16 in Equetus, 8–11 in Pareques) and by pigmen- tation differences. However, Darovec (1983) cautioned that the two species of Pareques (umbrosus and acuminatus) remain difficult to separate due to overlapping meristic ranges (Table 1) and because the primary distinguishing characteristics, juvenile color patterns and first dorsal fin lengths, change throughout ontogeny. , the black-bar drum, was a third recently described species (Miller and Woods, 1988). This new species was distinguished from P. acuminatus and P. umbrosus by the large wedge-shaped spot on the head above and posterior to the orbits, and by the broad oblique dark pigment band extending from the spinous dorsal fin to the pelvics. Although Robins et al. (1991) and Hoese and Moore (1998) retained the genus Equetus for all five species of reef sciaenids we follow Chao (1976) and separate Equetus and Pareques. We use P. umbrosus for the cubbyu and P. acuminatus for the high hat. Eggs and larvae of all species of reef sciaenids are undescribed. Herein we describe the growth and development of captive bred P. umbrosus larvae and juveniles, compare them to captive bred P. acuminatus larvae and juveniles, and provide new information for sepa- rating the larvae of these two species in field collections.

MATERIALS AND METHODS

Sub-adult cubbyu P. umbrosus were collected by SCUBA divers in September 1992 from oil platforms in the Gulf of Mexico off Port Aransas, Texas. The grew to approximately 150 cm SL during 1 yr in a 9 m3 rectangular tank with water recirculated through an external biofilter. Natural spawning first occurred in November 1993 and continued at a frequency of 3–5 times weekly through November 1996, with an average of 2000 eggs per spawning event. P. umbrosus spawned best under conditions of 25–26°C, salinity of 35‰, and 15:9 h light/dark photoperiod. The free- floating eggs were collected in mesh bags and placed into 150-L larval rearing tanks (Holt 1992). Larvae were raised under controlled conditions of 27–28°C, 12:12 h light/dark photoperiod, salini- ties of 34–35‰ and fed cultured rotifers (Branchionus plicatilis) (5 ml−1) and brine shrimp nauplii (Artemia franciscana) (1 ml−1). After metamorphosis (22–28 d) the juveniles were fed inert food. A few larvae were sampled daily for measurements and to monitor development. Larvae were measured live (NL = upper jaw to tip of notochord in preflexion larvae only and SL = upper jaw to posterior edge of hypurals) to the nearest 0.01 mm on a stereomicroscope equipped with a drawing tube and digitizing pad using Jandel Scientific SIGMASCAN 3.90 software. Juveniles were mea- sured live to the nearest millimeter on a standard metric rule. Illustrations of individuals were made HOLT AND RILEY: DEVELOPMENT OF PAREQUES UMBROSUS AND NOTES ON PAREQUES ACUMINATUS 827

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Seegat AegerangLnengthrangmhtgneealnnm/LeaDnaemLP cas-kloY 093− 2.24−3.024.825dn Preflexion44−92.47−3.221.931dn Preflexion170−123.42−4.4462.19 0.8 Flexion193−154.11−5.040.58221.1 Postflexion166−254.93−8.1761.10 1.5 Juvenile206−3010.07−15.184.44111.9 Juvenile3-stripe301−4813.70−27.2490.93 1.7 Juvenile4-stripe419−10027−4300.529 7 1.7

using a dissecting microscope and camera lucida attachment from live, anesthetized larvae before they were preserved and stored in 80% ethyl alcohol. Fifteen specimens of high hat P. acuminatus larvae were obtained from the Baltimore Aquarium. Mature high hats had been subjected to hormonal manipulation and spawned in 280 gallon rectan- gular tanks and the larvae reared on rotifers and brine shrimp (Harris, et al., 1994). Drawings and measurements of P. acuminatus were made on preserved specimens.

RESULTS

LARVAL MORPHOLOGY AND PIGMENTATION.—A comparison of age, and mean length at each developmental stage is presented in Table 2. The pelagic eggs of P. umbrosus ranged from 1.08 to 1.38 mm diameter and averaged 1.24 mm with a single, pigmented oil glob- ule measuring 0.23 mm diameter (Fig. 1A). At late egg stages, the larva, finfold and oil globule were heavily pigmented with golden-green chromatophores. Larvae hatched in approximately 12 h (at 27°C and 35‰) and averaged 2.82 mm NL (Fig. 1B). Newly hatched larvae had heavy golden-green stellate chromatophores on the trunk and tail, anterior yolk-sac and the oil globule, that reduced to scattered melanophores on the trunk, tail and yolk-sac after preservation. Additional, web-like pigment was present on the ex- ternal surface of the brain, and in the dorsal and ventral finfold (Fig. 1B). No significant changes in pigmentation occurred throughout the first 2 d of development. Transition from the yolk-sac stage to exogenous feeding occurred at 2.59 mm NL on day 3 posthatch when eyes and mouth were fully developed (Fig. 1C). Distinct pigmenta- tion changes occurred with this transition. The previously scattered melanophores of the tail had coalesced to form a distinct band of pigment on the tail about halfway between the anus and tip of the notochord. There was a broad oblique bar from the nape, through the pectoral fins to the abdomen and a dark, lateral stripe extended from the snout through the eye to the dorsal surface of the gut. Pigment was also present on the lower jaw and snout. The base of the pectoral fin-bud was deeply pigmented. When larvae reached about 3.25 mm NL, the contraction of the melanophores in the tail band resulted in the forma- tion of a single pigment spot on the ventral margin of the tail in the area of the latent anal fin (Fig. 2A). Just prior to flexion, 4.47 mm NL larvae develop two preopercle spines, an 828 BULLETIN OF MARINE SCIENCE, VOL. 65, NO. 3, 1999

Figure 1. Early developmental stages of the cubbyu Pareques umbrosus. A) late embryo egg, 1.24 mm diameter, n = 76 with a single oil globule at 0.23 mm diameter, n = 9; B) 2.87 mm NL newly hatched larva; C) 2.59 mm NL, 3 d posthatch. elongated spine located at the preopercular angle and another short spine just anterior to the preopercular angle (Fig. 2B). Notochord flexion occurred at 4.67 mm (~14 d posthatch) (Fig. 2C). At this time changes in pigmentation included a deeper concentration of black and gold chromatophores in the bar through the trunk, and the development of large stellate melanophores over the mid- brain, behind the eye and on the operculum. Soft tissues of the first dorsal were heavily pigmented with fine stellate melanophores. The two preopercle spines in flexion larvae appeared reduced in length but remained visible just opposite the pectoral base. A row of HOLT AND RILEY: DEVELOPMENT OF PAREQUES UMBROSUS AND NOTES ON PAREQUES ACUMINATUS 829

Figure 2. Preflexion developmental stages of the cubbyu Pareques umbrosus. A) 3.25 mm NL, 6 d posthatch; B) 4.47 mm NL, 10 d posthatch; C) 4.67 mm NL, 14-d old larva at flexion. stellate melanophores began to develop along the lateral midline of the tail and caudal fin in post flexion larvae (Fig. 3A). These melanophores formed a continuous row or hori- zontal stripe the entire length of the tail at about 8.16 mm SL (Fig. 3B). Melanophores also began to form vertical bars on the trunk and over the brain. Metamorphosis to the juvenile stage occurred between 10.00 and 15.00 mm and 26 to 30 d posthatch (Fig. 3C). The horizontal body stripes characteristic of juveniles and adults 830 BULLETIN OF MARINE SCIENCE, VOL. 65, NO. 3, 1999

Figure 3. Development of postflexion and juvenile stages of the cubbyu Pareques umbrosus. A) 5.39 mm SL, age 20 d; B) 8.16 mm SL, age 16 d; C) 12.16 mm SL, age 28 d. in all of the reef Sciaenids, began forming with the appearance of small melanophores at the base of the dorsal fin and the increasing thickness of the lateral mid-line pigment. Additional pigment was scattered in small patches of melanophores anterior to the orbit. Preopercle spines were still visible in early metamorphosis but were not detectable on any of the older age classes due to epidermal thickening prior to scale formation. HOLT AND RILEY: DEVELOPMENT OF PAREQUES UMBROSUS AND NOTES ON PAREQUES ACUMINATUS 831

Figure 4. Color pattern development in the juvenile stages of the cubbyu Pareques umbrosus. Arrows indicate white margins on fins. A) 3-stripe phase juvenile 13.70 mm SL; B) 4-stripe phase juvenile 30.0 mm SL; C) color pattern of adult individual at 110 mm SL. 832 BULLETIN OF MARINE SCIENCE, VOL. 65, NO. 3, 1999

JUVENILE MORPHOLOGY AND COLOR PATTERNS.—Juveniles older than 30 d posthatch and averaging 20.9 mm SL had a characteristic 3-stripe color pattern (Fig. 4A). Pigmentation anterior to the orbits was highly variable among individuals but body stripes and the mid- body bar were consistently present. The heavily pigmented first dorsal fin had a white trailing edge and was about 1.5 times longer than the body depth, i.e., the vertical dis- tance between the dorsal margin of body and the ventral symphysis of the cleithra. The heavily pigmented pelvic fin had white leading and trailing edges. Scale development began at mid-body, progressed over the head and finally over the trunk and tail, and was complete in the larger individuals (27 mm) of this age class. Juveniles 27 to 41 mm SL and 49–100 d posthatch were characterized by the development of a fourth lateral stripe (Fig. 4B). The pelvic and dorsal fins remained heavily pigmented but were reduced in length relative to earlier development. Pigment between the eyes formed a u-shaped pat- tern that in some individuals was closed by the presence of a narrow horizontal band. The number of longitudinal stripes increased gradually until all individuals greater than 50 mm SL had the full 7-primary stripe pattern observed in the adult (Fig. 4C). All seven primary stripes were narrower than the pupil width. Between the seven stripes were vari- able numbers of very fine dark stripes. The first dorsal fin now less than 1/3 of the body depth, retained some pigmentation and a white trailing edge. All other fins were without pigment as in the adult. FIN FORMATION.—Fin development in P. umbrosus occurred in the following sequence: first dorsal and pelvics, second dorsal, anal, and pectorals, resulting in a full meristic complement of IX-XI,38-40 dorsal, II+7-8 anal, I + 5 pelvic and 15 pectoral rays. On day 3 post hatch, a pectoral fin bud was present. Development of the spinous dorsal and pelvic fins began simultaneously at 4.67 mm NL (Fig. 2C). By 5.39 mm SL segmentation of the caudal rays was visible as were early developing rays of the second dorsal (Fig. 3A). First dorsal fin length was approximately one-half the body depth and the longest pelvic ray was about 60% of the length of the longest dorsal spine (Table 2). The full complement of 10 first dorsal spines was present at 8.16 mm SL and the longest spines had elongated to a length estimated to be equal to the body depth; the pelvics were 60% of the body depth (Fig. 3B). Segmentation, though not complete, was visible on all fin rays. Fin formation was complete by 12.16 mm SL (Fig. 3C) with the first dorsal fin length approximately one and one half times the body depth. DESCRIPTIVE NOTES ON P. ACUMINATUS.—Since only 15 specimens were available for study, a complete larval description of P. acuminatus is not presented here. Yolk-sac and early larvae (Fig. 5) were similar to P. umbrosus larvae. Pigment patterns were nearly identical when preserved specimens of both species were compared, but with only four specimens of P. acuminatus yolk-sac and preflexion larvae to examine there may be subtle differ- ences that we missed. Three pre-opercular spines developed by flexion (Fig. 5D) and persisted throughout larval development. The pelvics and first dorsal also began to form at this time. By 5.5 mm (day 14), postflexion larvae had five dorsal spines and developing rays in all other fins (Fig. 6A). The first dorsal had a white leading edge and the length was less than the length of the longest pelvic rays (Table 3). A series of stellate chromatophores were present along the midline of the body and extended onto the developing caudal fin. An oblique band of pigment ran from the spinous dorsal to the pelvic fin, and stellate chromato- phores were scattered on the head above, behind, and below the eye (Fig. 6A). HOLT AND RILEY: DEVELOPMENT OF PAREQUES UMBROSUS AND NOTES ON PAREQUES ACUMINATUS 833

Figure 5. Early developmental stages of the high hat Pareques acuminatus. A) 2.29 mm NL, newly hatched larva; B) 2.60 mm NL, age 3 d posthatch; C) 2.83 mm NL, age 5 d posthatch; D) 5.18 mm SL, age 10 d posthatch. 834 BULLETIN OF MARINE SCIENCE, VOL. 65, NO. 3, 1999

Figure 6. Development of flexion and postflexion larvae of the high hat Pareques acuminatus. A) 5.85 mm SL, age 14 d; B) 7.70 mm SL, age 19 d.

A 7.7 mm (19 d) postflexion larva ( Fig. 6B) had nearly complete fin development and a pigment pattern little changed from the previous stage. Melanophores now formed a vertical band from the first dorsal through the pelvics, and another band immediately behind the eye. Pigment also formed a horizontal stripe along the mid body from behind the eye through the caudal fin. The first dorsal had leading and trailing white edges and was longer than the pelvic fins. Fin formation was complete by 19.25 mm and scales covered the mid-body and head regions (Fig. 7). The spinous dorsal on this three stripe phase juvenile was extremely long HOLT AND RILEY: DEVELOPMENT OF PAREQUES UMBROSUS AND NOTES ON PAREQUES ACUMINATUS 835

Figure 7. A juvenile high hat Pareques acuminatus aged 31 d, 19.25 mm SL. (75% of SL), extending beyond the tip of the caudal, with a very clear white leading edge. Pelvic fins were also long (40% of SL), reaching the middle of the anal fin base and had white leading edges.

DISCUSSION

Larvae of P. umbrosus are characterized during ontogeny by specific pigment patterns, the presence of two preopercular spines, and elongate, precocious spinous dorsal and pelvic fins. There is very little published data on larvae of other Pareques or Equetus for comparison (Ditty, 1989). Preflexion P. acuminatus described here are very similar to P.umbrosus but can be distinguished by development of three spines on the preopercle compared to two spines in P. umbrosus. In flexion larvae, the length of the longest pelvic ray in P. acuminatus is 1.3 to 3 times the length of the longest dorsal spine while both fins are about equal in P. umbrosus (Tables 2,3). Three larvae described as Pareques by Powles and Burgess (1978) differ from P. umbrosus by the number of spines on the preopercle. The two that were collected from Florida (figs. 1 and 2 in Powles and Burgess, 1978) are probably P. acuminatus based on relative length of the longest dorsal spine to length of the first pelvic ray and the presence of three preopercular spines. The third specimen, collected off Santa Marta, Colombia was tentatively identified by Miller and Woods (1988) as P. iwamotoi based on pigment patterns and the longer spinous dorsal fin relative to the pelvic fins. Thus it may be pos- sible to distinguish the three species of Atlantic Pareques from one another at flexion based on number of spines on the preopercle (3 in P. acuminatus and P. iwamotoi, and 2 in P. umbrosus) and by the greater length of the pelvic fins to spinous dorsal in P. acuminatus. 836 BULLETIN OF MARINE SCIENCE, VOL. 65, NO. 3, 1999

derbevitpacfohtgneldevreserpdnaegA.3elbaT sutanimucaseuqeraP ,hctahtsopsyadniegA. nifcivleptsegnolehtsiLP,)mm(htgnellasrodtsriftsegnolehtsiLD,LSroLNmmnishtgnel .depolevedtonerewsnifehtsnaemdn,)mm(htgnel

S egat Aeeg LhengthrangmNLeanlengtDL/P Y1olk-sac 2−.29 1dn Y3olk-sac 29.58−2.602.52dn Preflexion52−.83 1dn Flexion 1042.28−5.184.5440.3 Flexion1459.52−6.295.8350.7 Postflexion** 197−.70 141.8+ J31uvenile3-stripe 154.2−19.217.3312.2 **firstdorsalbroken Although little information is available for larvae, several authors have examined juve- niles and described differences among the species of reef sciaenids. Juvenile E. lanceolatus (jackknife-fish) and E. punctatus (spotted drum) are separable from Pareques spp. by having more than 10 dorsal spines, greater than 45 dorsal rays (Table 1), and by pigmen- tation pattern. Young P. iwamotoi (blackbar drum) have a large dark wedge-shaped spot on the head between the orbits and a single lateral body stripe (Miller and Woods 1988). Juvenile P. umbrosus (cubbyu) have been separated from P. acuminatus (high-hat) by the difference in width of the longitudinal body stripes, and the pigment pattern between the orbits (Miller and Woods, 1988). Juvenile P. acuminatus have 3 to 5 broad longitudinal stripes wider than the pupil and narrow stripes in between them; a straight bar connecting the orbits; and 8 to 9 dorsal spines (Chao, 1978). Juvenile P. umbrosus described here have 3 to 7 longitudinal stripes with all but the center stripe narrower than the pupil and all stripes becoming more narrow with age. Young P. umbrosus often have a u-shaped band low (closer to the snout) between the orbits, that may be closed by a straight bar connecting the orbits and 10 dorsal spines. A conspicuous white leading edge on the spinous dorsal of young P. acuminatus (Bohlke and Chaplin, 1993) is a good additional character for separating the post-flexion larvae and three to five stripe juveniles from the young of P. umbrosus. In our study of lab reared juvenile P. acuminatus, we confirmed the presence of the conspicuous white leading and trailing edges on the first dorsal, and the extremely long and flexible first dorsal (~2× head depth; >50% body length). Conversely, P. umbrosus juveniles have only a white trailing edge on the first dorsal, white leading and trailing edges on the pelvic fins, and a short, stiff first dorsal (~1–1.5× head depth; <50% body length).

SUMMARY

The availability of laboratory reared specimens enabled us to describe ontogenetic de- velopment in P. umbrosus and a closely related species P. acuminatus. This information allows preflexion larvae as small as 4.5 mm (live) SL of these two species to be distin- guished from each other. Larvae of P. umbrosus have two spines on the preopercle that gradually become incon- spicuous prior to the juvenile stage. The length of the longest first dorsal element in HOLT AND RILEY: DEVELOPMENT OF PAREQUES UMBROSUS AND NOTES ON PAREQUES ACUMINATUS 837 flexion larvae is equal to or longer than the length of the pelvic fin. The first dorsal fin in fish with three body stripes has only a white trailing edge while the pelvics have both white leading and trailing edges; the spinous dorsal fin is relatively short, reaching a length <50% body length; the pelvic fins do not reach the base of the anal fins. Pigment between the eyes forms a horseshoe or u-shape pattern that is sometimes closed by a bar between the orbits. This pigment pattern constricts to a dark spot on the snout and a dark line between the eyes on preserved specimens. Larvae of P. acuminatus have three spines on the preopercle that remain conspicuous until the juvenile stage. The length of the longest dorsal element in flexion larvae is only 34–75% of the length of the pelvic fin. The first dorsal fin in larvae with three body stripes has both white leading and trailing edges and is long and flowing reaching a length 50% or greater of the standard length; pelvic fins reach to the middle of the anal base. The pigment pattern between the eyes consists of a bar between the orbits and a dark mark on the snout, not different than some of the P. umbrosus specimens.

ACKNOWLEDGMENTS

We would like to thank J. Lyczkowski-Shultz and D. Hoese for comments and suggestions on an early draft of this manuscript. Special thanks to M. Harris and the Baltimore Aquarium for the specimens of P. acuminatus larvae. Partial funding for this work was provided by grants from DOC\NOAA’s Saltonstall-Kenedy, and the Sid W. Richardson Foundation. This is contribution num- ber 1113 of The University of Texas at Austin Marine Science Institute.

LITERATURE CITED

Böhlke, J. E. and C. C. G. Chaplin. 1993. of the Bahamas and adjacent tropical waters. Univ. Texas Press, Austin. 771 p. Chao, L. N. 1976. Aspects of systematics, morphology, life history and feeding of western Atlantic Sciaenidae (Pisces: ). Ph.D. Thesis, College of William and Mary, Williamsburg, Virginia. 342 p. ______. 1978. A basis for classifying western Atlantic Sciaenidae (Teleostei: Perciformes). NOAA Tech. Rpt. Circ. 415. Darovec, J. E. 1983. Sciaenid fishes (Osteichthyes: Perciformes) of western peninsular Florida. Memoirs of the Hourglass Cruises. 6: 1–73. Ditty, J. G. 1989. Separating early larvae of Sciaenids from the western North Atlantic: A review and comparison of larvae off Louisiana and Atlantic coast of the U.S. Bull. Mar. Sci. 44: 1083–1105. ______and R. F. Shaw. 1994. Preliminary guide to the identification of the early life history stages of sciaenid fishes from the western central Atlantic. NOAA Tech. Mem. NMFS-SEFSC- 349. 118 p. Harris, M. C., A. Henningsen, B. Hecker, C. Andrews, S. Hassin, J. Stubblefield and Y. Zohar. 1994. Controlled reproduction of the high hat (Equetus acuminatus) at the National Aquarium in Baltimore. AZA Ann. Conf. Proc., Atlanta. 1994: 256–260. Holt, G. J. 1992. Experimental studies of feeding in larval red drum. J World Aquacult. Soc. 23: 265–270. Hoese, H. D. and R. H. Moore. 1998. Fishes of the Gulf of Mexico, Texas, Louisiana, and adjacent waters. Texas A&M Univ. Press, College Station. 422 p. Jordan, D. S. and B. W. Evermann. 1898. The fishes of North and Middle America. Bull U.S. Nat’l. Mus. 47:1485–1490. 838 BULLETIN OF MARINE SCIENCE, VOL. 65, NO. 3, 1999

McPhail, J. D. 1961. A review of the tropical eastern Pacific species of Pareques (Sciaenidae). Copeia 1: 27–32. Miller, G. C. and L. P. Woods. 1988. A new species of sciaenid fish, Pareques iwamotoi, from the western Atlantic, with color descriptions of prejuvenile and juvenile Pareques acuminatus and Pareques umbrosus. Bull. Mar. Sci. 43: 88–92. Powles, H. And W. E. Burgess. 1978. Observations on benthic larvae of Pareques (Pisces: Sciaenidae) from Florida and Columbia. Copeia 1: 169–172. Randall. J. E. 1983. Caribbean reef fishes. 2nd ed. T.F.H. Publications, Neptune, New Jersey, 350 p. Robins, C. R., R. M. Bailey, C. E. Bond, J. R. Brooker, E. A. Lachner, R. N. Lea and W. B. Scott. 1991. Common and scientific names of fishes from the United States and Canada. 5th ed. Amer. Fish. Soc. Spec. Publ. 20. Bethesda, Maryland. 183 p.

DATE SUBMITTED:January 22, 1999. DATE ACCEPTED: July 23, 1999.

ADDRESS: The University of Texas at Austin, Marine Science Institute, 750 Channel View Drive, Port Aransas, Texas 78373. CURRENT ADDRESS: (C.M.R.) Gulf Coast Bird Observatory, 9800 Richmond Ave., Suite 150, Houston, Texas 77042. CONTACT NUMBERS: (G.J.H.) (361) 749-6749 (Tel); (361) 749-6749 (Fax); e-mail: [email protected]. (C.M.R.) (713) 789-4226 (Tel); e-mail: .