<I>Pareques Umbrosus</I>
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BULLETIN OF MARINE SCIENCE, 65(3): 825–838, 1999 LARVAL AND JUVENILE DEVELOPMENT OF THE CUBBYU PAREQUES UMBROSUS WITH NOTES ON THE HIGH HAT PAREQUES ACUMINATUS 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 species 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 Sciaenidae 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 genus 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 arenegehtnisdineaicsfeerrofscitsireM.1elbaT suteuqE dna seuqeraP yttiDmorfdeifidom, .)4991(wahSdna 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. Pareques iwamotoi, 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 fish 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 uybbucderbevitpacfohtgneldnaegA.2elbaT susorbmuseuqeraP ,hctahtsopsyadniegA. nifcivleptsegnolehtsiLP,)mm(htgnellasrodtsriftsegnolehtsiLD,LSroLNmmnishtgnel .depolevedtonerewsnifehtsnaemdn,)mm(htgnel 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.