Habitat and Life History of Juvenile Hawaiian Pink Snapper, Pristipomoides Filamentosus 1

Habitat and Life History of Juvenile Hawaiian Pink Snapper, Pristipomoides filamentosus 1

ROBERT B. MOFFITT AND FRANK A. PARRISH 2

ABSTRACT: Eteline snappers are an important component of commercial demersal fisheries in the central and western Pacific, but there is a substantial gap in the knowledge of their life histories, specifically the larval and juvenile stages. Juvenile pink snapper, Pristipomoides filamentosus (Valenciennes), rang ing in size from 7 to 25 em fork length, inhabit a nearly featureless plain off shore of Kane'ohe Bay, O'ahu, at depths of 65-100 m. Bottom samples and underwater video footage showed the bottom to be uniformly composed offine, silty sand with little relief. Conductivity-ternperature-depth data indicate that an internal tide brings cold water over the bottom on a tidal basis. Telemetric studies show that juveniles undergo small-scale crepuscular migrations from deeper daytime locations to shallower nighttime locations but move relatively little during day and night periods. Analysis of length frequency distributions obtained over a 17-month period resulted in an estimate of the von Bertalanffy l growth constant (K) of 0.21 yr- .

ETELINE SNAPPERS SUPPORT existing or devel fishes (Leis 1987, Clarke 1991). Aside from a oping fisheries throughout the central and preliminary investigation by Parrish in 1989, western Pacific, where they are generally the information on the juvenile stage of these most valuable component of the demersal fishes is nonexistent. fisheries. These fishes are prized for their Parrish (1989) documented juvenile pink high-quality flesh and large size. Adults con snappers, Pristipomoides filamentosus (Valen centrate near high-relief features on the deep ciennes), in a flat, featureless habitat very slopes of Pacific islands and banks at depths different from high topographic relief areas of 100-400 m. Many aspects of the biology adults occupy. Juvenile eteline snappers have of the adults have been addressed: feeding not been observed during submersible sur (Kami 1973, J. D. Parrish 1987, Seki and veys of adult and subadult aggregations at Callahan 1988, Haight et ai. 1993), growth natural (Ralston et ai. 1986) and artificial (Ralston and Miyamoto 1983, Edwards 1985, (Moffitt et al. 1989) high-relief features. The Manooch 1987, Ralston and Williams 1988), use of a flat, featureless habitat as a nursery reproduction (Everson 1984, Kikkawa 1984, area is also very different from the situation Grimes 1987, Everson et ai. 1989), and ex reported for most other fishes, where the ploitable biomass (Polovina et ai. 1985, Po young utilize complex bottom habitats such lovina and Ralston 1986, Polovina and Sho as seagrass beds (Adams 1976, Heck and mura 1990). Information on the larval and Weinstein 1989, Sogard and Able 1991), algal early juvenile stages, however, is lacking. beds (Heck and Thoman 1981, Wilson et ai. Eteline snapper larvae are rarely encountered 1990), or natural and artificial reefs (Carlson in ichthyoplankton collections, providing lit and Straty 1981, Anderson et ai. 1989, Con tle biological information on presettlement nell and Jones 1991). Understanding the biology and the ecological requirements of juvenile eteline snap 'Manuscript accepted 26 January 1996. 2Honolulu Laboratory, Southwest Fisheries Science pers is necessary to appropriately manage Center, National Marine Fisheries Service, NOAA, 2570 both fish resources and their habitat through Dole Street, Honolulu, Hawai'i 96822-2396. out the Pacific region. This study expands on 371 372 PACIFIC SCIENCE, Volume SO, October 1996

the work of Parrish (1989), providing greater Temporal Sampling detail of the physical features of the habitat and its spatial and temporal utilization by Length frequency samples were collected juvenile P. filamentosus in Hawai'i. monthly between October 1989 and Febru ary 1991. Intervals between samples were usually 4-6 weeks, with one gap of 9 weeks caused by poor weather conditions. Obtain MATERIALS AND METHODS ing each sample required between one and five fishing days and ended after 1 week or Fishing Operations when about 100 fish had been caught. A drifting 6-m boat equipped with a fish An estimate of the von Bertalanffy growth finder was used as the fishing platform for parameter, K, was obtained using the ELE most of this study. All fish used in length FAN computer program (Gayanilo et al. frequency analysis were caught by rod and 1989), with L oo restricted to 78 cm as esti reel, fishing with light tackle. Terminal gear mated by Ralston and Miyamoto (1983). The consisted of about 2 m of 7-kg monofilament length-weight relationship was calculated line with a 0.1- to 0.2-kg lead weight on the using the SAS nonlinear regression program bottom and four evenly spaced 30-cm-Iong 2 (NUN). to 3-kg monofilament branch lines ending in circle hooks with a 5.5-mm gap. Strips of Habitat Description squid, Loligo opalescens Berry, were used as bait. Lines were fished with leads on or near Environmental conditions of the juvenile the substrate. Depth of capture and fork snapper habitat off Kane'ohe Bay were length (FL) to the nearest 1 mm were re studied on a Townsend Cromwell cruise (TC corded for all pink snapper taken. Prey items 90-02) in February 1990. Three transect lines regurgitated from pink snapper upon landing were established perpendicular to the depth were noted, but no gut analysis was con contours off the southern, central, and north ducted. Whenever possible, fish were re ern portions of the bay (Figure 1). Bottom turned to the ocean after we vented excess grabs and conductivity-ternperature-depth gas from their swim bladder with a hypo (CTD) casts were taken at five depths (45, 60, dermic needle and repositioned the stomach 75, 90, and 105 m) along each transect. when everted. Moribund fish were iced and Sediment samples were frozen when col returned to the laboratory for other use (e.g., lected, then oven-dried at 50-60°C, stored in weighing and otolith studies). We fished only a desiccator until cooled to room temper during daylight hours, between 0900 and 1600 ature, and weighed to the nearest 0.1 g. The hours. samples were washed through a series of three sieves with mesh sizes of 2, 0.42, and 0.06 mm. The material remaining on each of Survey ofSites the screens was redried and weighed. Based Thirteen exploratory fishing trips were on the classifications established by Went conducted around O'ahu between 2 October worth (1922), the materials collected on the and 9 November 1989 to identify a site that 2-mm sieve are termed "granules or larger," would yield enough juvenile pink snapper for those on the 0.42-mm sieve are "medium to a temporal analysis. Survey effort focused coarse sand," those on the 0.06-mm sieve are on leeward O'ahu, where better weather con "fine to medium sand," and the material lost ditions permitted more regular sampling. through the last sieve (calculated by sub Depths between 50 and ISO m were fished tracting the weight of sediment collected on over 40% of the island's coastline (Figure 1). the three sieves from the original weight) is Windward sites where fish had previously termed "mud." A sand-mud ratio was calcu been found were also included (Parrish 1989). lated by dividing the combined weight of the Juvenile Hawaiian Pink Snapper-MoFFm AND PARRISH 373

& 0.

/ / / / / / / / / / / / / / / CO::> Kapapa I.

KANEOHE BAY lJ Coconut I. OAHU, HAWAII OAHU

Kilometers o 2 J

FIGURE 1. O'ahu and Kiine'ohe Bay (shaded areas on the O'ahu map represent coastline fished in the initial survey).

two sand categories by the weight of the mud Telemetry for each sample. Visual documentation of the bottom type and associated fauna was ob Acoustic tags were implanted in two tained from eight baited video camera drops hooked pink snappers using a subsurface conducted between October 1990 and Jan tagging method developed in this study and uary 1991. reported in Parrish and Moffitt (1992). Temperature profiles were generated from Within 25 min, each fish was captured, the CTD data collected at the same times and tagged, and released near the point of cap sites as the bottom grabs. An additional ture. After release, each fish was inter series of CTD data was taken for a 24-hr mittently tracked over a 5-day period (the period along the northern transect line. In approximate life of the tag's battery) by a this series, CTD data were collected for surface vessel with a directional hydrophone. each depth on the transect every 1-2 hr. Time, bottom depth, and location were re Temperature profiles were also obtained in corded every !-I hr during tracking. The December 1989 (n = 3), November 1990 tags were ca. 8 by 30 mm cylinders with (n = 16), and December 1990 (n = 6). a specific acoustic frequency between 65 and 374 PACIFIC SCIENCE, Volume 50, October 1996

75 kHz depending on the tag. On the first during those months. The ELEFAN estima tracking, positions were calculated from vis tion of the von Bertalanffy growth constant ual and radar sightings of shoreline features. (K), with a preselected L oo of 78 cm, was 1 On the second tracking, positions were ob K = 0.21 yc • tained from a portable Global Positioning Weight (W) in grams was recorded for 125 System (GPS) unit. individuals ranging in FL from 8.0 to 20.8 cm, and the data were used to determine the b length-weight relationship (VV = aFL ). Val ues obtained for the parameters (with the RESULTS ranges of the 95% confidence intervals) were Sites Surveyed a=0.018 (0.012-0.025) and b=2.99 (2.87 3.12). A total of 66 juvenile pink snapper was captured at only two sites during this survey. Habitat Description Four specimens were caught on leeward O'ahu, off Honolulu Harbor, at depths of The sieve analysis ofbottom grabs (n = 15) 63-100 m during 5 days of fishing. The re showed that sediments at 45 m consisted maining 62 specimens were taken at similar mostly of coarse sand and larger materials depths during 3 days of windward coast fish (Table 1). At greater depths, where the snap ing off Kane'ohe Bay. The temporal and pers inhabit, the sediments were dominated habitat evaluation focused on the high snap by fine sand and mud in almost all samples. per yield of the Kane'ohe site. Review of the video camera tapes, and observations from the depth finder during Temporal Sampling fishing operations, confirmed the lack of ver tical relief in the area. The bottom appeared A total of 1047 juvenile pink snapper to be entirely soft, with many invertebrate ranging from 8 to 25 cm FL was caught in burrows and occasional shallow depressions the northern and central portions of the that may have been made by either dasyatid Kane'ohe Bay site at depths of 60-100 m rays or portunid crabs. Species recorded by during the monthly sampling. No pink snap the camera over this habitat include juvenile per were caught in the southern portion of pink snapper; white crab, Portunus sanguino the study site. Fishing frequently extended lentus (Herbst); dasyatid ray; subadult am beyond the 60-100 m targeted depth range, berjack, Seriola dumerili (Risso); spiny puffer, but no juvenile pink snapper were captured Torquigener sp.; sharp-nosed puffer, Can outside the target depths. Catches ofjuvenile thigaster rivulata (Schlegal); and carcharhinid pink snapper on the "highest" of the fishing sharks. On two occasions, when a strong lines' four hooks (about 2 m off the bottom) current was present, the camera frame was were rare, suggesting that these fish have a dragged across the bottom, passing derelict close association with the bottom. Large, crab traps. In the vicinity of the traps, species obvious regurgitated prey included postlarval composition changed dramatically, to one fishes (gobies, synodontids, and Lutjanus kas of Chaetodon miliaris Quoy & Gaimard, mira [Forsskal]), small cephalopods, and por Heniochus diphreutes Jordan, Lutjanus kas tunid crabs. mira, Priacanthus sp., Parupeneus chrysone Fish length was not correlated with depth mus (Jordan & Evermann), and Canthigas of capture (R2 = 0.035), indicating no obvi ter coronata (Vaillant & Sauvage). No pink ous stratification of size by depth. ELE snapper were observed near the traps. FAN analysis of the 13 length frequency dis Analysis ofCTD temperature data showed tributions (Figure 2) identified two modes in evidence of an internal tide with a semi the October to February data for both 1989 diurnal periodicity that provides an influx of 1990 and 1990-1991, indicating the presence cold water over the bottom on the juvenile of two year classes on the nursery grounds pink snapper nursery grounds during high Juvenile Hawaiian Pink Snapper-MOFFITT AND PARRISH 375

Fork Length (em)

FIGURE 2. Length frequency distribution ofjuvenile pink snapper by month. tide. The presence of this tide is best shown 90 m at low tide. During high tide, temper with the 24-hr CTD casts taken along the ature in the upper 70 m remained unchanged, north transect line (Figure 3). Surface tem while temperature from 70 m to the bottom perature at all sites was about 23.0-23.5°C, declined rapidly to a low of about 20.5 gradually declining to about 22.0-22SC at 21.0°C on the bottom. This internal tidal 376 PACIFIC SCIENCE, Volume 50, October 1996

TABLE I

SEDIMENT SIZE BY DEPTH ON TRANSECTS TAKEN ON JUVENILE PINK SNAPPER NURSERY GROUNDS

PARTICLE SIZE (% of total)

DEPTH COARSE FINE SAND/MUD TRANSECT (m) ~GRANULE SAND SAND MUD RATIO

Southern 45 55.4 37.0 6.6 1.0 41.9 60 3.1 16.6 74.8 5.5 16.5 75 6.0 23.9 43.3 26.9 2.5 90 3.2 14.3 36.7 45.8 1.1 105 10.3 35.3 39.1 15.2 4.9 Central 45 46.7 43.4 7.3 2.6 19.4 60 30.6 48.7 16.6 4.0 16.2 75 0.2 1.6 44.3 54.0 0.8 90 1.4 5.3 41.1 52.2 0.9 105 7.4 13.4 44.3 34.9 1.7 Northern 45 29.8 18.8 46.5 4.9 13.3 60 0.7 6.1 78.0 15.1 5.6 75 1.0 2.0 50.1 46.9 1.1 90 0.03 0.5 41.7 57.8 0.7 105 0.4 1.1 52.0 46.5 1.1 feature with a temperature inflection at 60 we were able to observe its return to capture 80 m was observed to some extent at all position. It was caught at a depth of 72 m, sample depths except the shallowest (45 m) the GPS position was recorded, and then it and was observed on all sampling days, sug was released (after processing during a drift) gesting that it is a persistent phenomenon. where the depth was 61 m (about 300 m from the capture location). The fish gradually Telemetry moved deeper over the next 2 hr, arriving at its capture location, where it remained until Two juvenile pink snappers (each 19 cm dusk. Video records of the day and night FL) were successfully tagged and tracked positions of the second fish revealed no over a period of 5-6 days (Table 2). Both fish discernable difference in bottom features, moved similarly from deeper daytime posi though the fish are quite obviously able to tions to shallower nighttime locations. The recognize particular locations on the bottom. migration was not linked to the temperature fluctuation caused by the internal tide de scribed above but was crepuscular in peri DISCUSSION odicity. Each fish exhibited discrete day and night depths and positions, migrating be The results of this study can now be used tween the two positions rapidly (within 30 as a framework in which the early life history 40 min) at dawn and dusk. Day ranges were of juvenile pink snapper off Kane'ohe Bay double or more than double the area of the can be described. Juveniles first arrive at a night ranges for both fish, indicating greater size of about 7-10 cm FL (the smallest cap daytime activity. Recorded day and night tured was 7.3 cm FL) in October or No positions for the first fish contained a small vember. Although it is not known whether degree of overlap; however, the lack of over juveniles are just settling after a long pelagic lap in the depths recorded for these positions stage or immigrating from another benthic indicate that this was an artifact of the poor habitat, juveniles of this size or smaller have positioning method used. With the better po not been reported from other benthic hab sitions obtained for the second tagged fish, itats, either deeper or shallower. Leis (1987) LOW TIDE HIGH TIDE

45m 40

120

Or-~----,---.-----,--,.----r--....--r--...., GOm 40 GOm 40

80 80

120 120

r-.....,.....---,,--...... ----,----r-...,.__,.-----, 0 -E 40 75 m 75 m 40 .r::... Q, 80 80 CD C 120 120

...-..,..-----,---.-----,--...,....-----r-----,.,.----., 0

40 90 m 90 m 40

80 80

120 120

O...-.....,.....---,,--...... ----.,.-...,....-----r-----.,.,....---., r-~---,---..,.--...,.__-r-----, 0

40 105 m 105 m 40

80 80

120 120

20 21 22 23 24 20 21 22 23 24 Temperature (OC) Temperature (OC)

FIGURE 3. Temperature profiles at high and low tides. 378 PACIFIC SCIENCE, Volume 50, October 1996

TABLE 2

RESULTS OF TELEMETRY

PARAMETERS FISH 1 FISH 2

Dates tracked 4-8 December 1989 13-18 September 1990 Daytime depth range (m) 84-87 71-73 Nighttime depth range (m) 74-77 65-67 2 Area of daytime positions (km ) 0.40 0.06 2 Area of nighttime positions (km ) 0.20 0.02

reported a pelagic juvenile of the congeneric particularly strong or the 1990 (and possibly Pristipomoides sieboldii (Bleeker) of 5.0 cm 1988) year class was particularly weak. length, suggesting that our smallest juveniles The nursery habitat used by the juvenile off Kane'ohe Bay may indeed have recently pink snapper appears to be very different settled from their pelagic stage. Juvenile pink from that described for other juvenile fishes. snapper remain on the nursery grounds for Most authors suggest that highly complex a period of 6-7 months or until they reach substrates (e.g., coral reefs, artificial reefs, 18-20 cm FL (although some individuals as seagrass, and algal beds) reduce predation by large as 25 cm FL remain in the area). After providing protective shelter (Adams 1976, leaving the flat, featureless nursery grounds, Heck and Thoman 1981, Wilson et al. 1990, the juveniles appear to concentrate around Connell and Jones 1991) for juveniles ofvar high-relief features in deeper water. A docu ious species. Juvenile pink snapper at the mented example is an artificial reef at 150 m Kane'ohe Bay nursery may achieve com off leeward O'ahu, where pink snappers parable protection from predation by utiliz (~20 cm FL) are routinely caught by State ing flat, featureless areas where predators are of Hawai'i Division of Aquatic Resources absent. At the depths involved here, high personnel (Okamoto 1993). level predators (adult snappers, amberjacks The estimate of growth based on length [Okamoto 1993], and other jacks) concen l frequency distributions (K=0.21 ye ) falls trate around high-relief structures and do not within the range of values (0.10-0.25) re occur in large numbers over featureless bot ported for species of Lutjanidae (Manooch toms (Ralston et al. 1986, Moffitt et al. 1987). It is supported by preliminary results 1989). of otolith examination for these juvenile pink It may not be merely coincidental that snapper, indicating ages of about 180 days 60 m is the shallowest depth at which juve for a 10.5 cm FL fish and about 350 days for nile pink snapper have been observed off an 18.5 cm FL fish (DeMartini et al. 1994). Kane'ohe Bay and is also the shallowest Recruitment to this nursery ground ap depth where effects of the internal tide were pears to vary considerably between years. In recorded. Studies of internal waves near the February 1990, the 1989 year class (FL ~ 16 surface of the ocean indicate that these cm) was much more abundant than that from phenomena assist in shoreward transport 1988 (FL ~ 17 cm); in February 1991, the of larval fish and crustaceans (Kingsford 1990 year class was much less abundant than and Choat 1986, Shanks and Wright 1987, that from 1989. Differences in catch per unit Shanks 1988). Although similar studies have effort obtained during the two sampling pe not been conducted on subsurface internal riods (103 fish on 1 day in February 1990 waves, such as the internal tide described versus 55 fish on 2 days in February 1991) here, it is possible that a similar process may furnish crude supporting evidence for this occur. The internal tides may assist in shore variability. Currently, we have no way of ward transport of postlarval pink snapper. determining whether the 1989 year class was Examination of the sand/mud ratios (Table Juvenile Hawaiian Pink Snapper-MoFFm AND PARRISH 379

2) indicates a strong discontinuity in the a more intensive study of the Kane'ohe Bay transition from larger to smaller particle sizes nursery ground. After more research, we at the 60-m depth. A trend of higher sandi should be able to determine the relevance of mud ratios in the southern versus northern nursery grounds for eteline snappers through and central portions of the study area for out the Indo-Pacific region. The low-relief each depth sampled may be associated with shelf areas used by juveniles of this and per the lack of juvenile pink snapper in the haps other eteline species could be a limited southern area. Interrelationships between the resource because of the steep slopes of most internal tide, sediment particle size, and oc oceanic islands. Consequently, management currence of juvenile pink snapper currently and protection of these habitats may be im are not understood, but the transition in all portant. Such protection appears contrary three factors at 60 m on the nursery grounds to conventional wisdom that expounds the raises intriguing questions about the processes benefits in terms of productivity of creation involved. of artificial structures on just such low-relief Telemetry suggests that juvenile pink areas. snapper, like the adults (Moffitt 1980), mi grate crepuscularly. Our tagged fish moved as if they were able to recognize bottom ACKNOWLEDGMENTS locations and return to their habitual sites, both on a routine daily basis and when arti We appreciate the work of all those who ficially displaced from their initial location contributed to our field study, particularly (i.e., our second tagged fish moved between Karl Bromwell, Wayne Haight, Theresa capture and release). The benefit obtained Martinelli, and Leslie Timme for the long, from these short-range daily migrations is wet days and nights spent in a small, rocking not known. Day and night positions for boat during our telemetry operations. We are both tagged fish occurred within the range also grateful for the comments of those who of depths where pink snapper are routinely reviewed the manuscript at various stages: captured during the day. Thus, night posi Edward DeMartini, Denise Ellis, Wayne tions of our tagged fish were probably uti Haight, Gene Huntsman, Sharon Kramer, lized by other juvenile pink snapper during James Parrish, James West, and two anony the day and day positions utilized by yet mous reVIewers. others at night. This study has examined a relatively con centrated, isolated, and persistent population LITERATURE CITED ofjuvenile pink snapper occupying a uniform featureless habitat in northern Kane'ohe Bay. ADAMS, S. M. 1976. The ecology of eelgrass, Successive year classes were observed to use Zostera marina (L.), fish communities. I. the same habitat, with individual fish appar Structural analysis. J. Exp. Mar. BioI. ently making only small diel movements. Ecol. 22: 269-291. Observations, measurements, and collections ANDERSON, T. W., E. E. DEMARTINI, and were made on some characteristics of the D. A. ROBERTS. 1989. The relationship habitat that may affect its use by juvenile between habitat structure, body size and pink snapper. However, this work is descrip distribution of fishes at a temperate artifi tive and is not compared with that on other cial reef. Bull. Mar. Sci. 44(2): 681-697. pink snapper nursery grounds. Thus, we are CARLSON, R, and RR STRATY. 1981. Hab unable to attach the importance of any of itat and nursery grounds of Pacific rock these characteristics to juvenile snapper with fish, Sebastes spp., in rocky coastal areas any confidence. Future research should be of southeastern Alaska. Mar. Fish. Rev. directed toward locating other pink snapper 43(7): 13-19. nursery grounds and comparing those hab CLARKE, T. A. 1991. Larvae of nearshore itats with that described above as well as to fishes in oceanic waters near Oahu. U.S. 380 PACIFIC SCIENCE, Volume 50, October 1996

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