A biogeographic assessment of the Samoan Archipelago

This report examines the marine biogeography of the Samoan Archipelago (~14º S latitude along the international date-line) with a focus on regional ocean climate, connectivity among islands due to larval transport, distributions of reef fish and coral communities, and the extent of existing marine protected areas. Management decisions and prior assessments in the archipelago have typically been split along the international political boundary between the islands of Samoa and those of American Samoa despite their close proximity and shared resources. A key goal in this assessment was to compile data from both jurisdictions and to conduct the characterization across the entire archipelago. The report builds upon earlier assessments by re-analyzing and interpreting many pre-existing datasets, adding more recent biogeographic data sources, and by combining earlier findings into a multidisciplinary summary of marine biogeography. The assessment is divided into 5 chapters and supporting appendices. Each chapter was written and reviewed in collaboration with subject matter specialists and local experts. In Chapter 1, a short introduction to the overall scope and approach of the report is provided. In Chapter 2, regional ocean climate is characterized using remote sensing datasets and discussed in the context of local observations. In Chapter 3, regional ocean currents and transport of coral and fish larvae are investigated among the islands of the archipelago and surrounding island nations. In Chapter 4, distinct reef fish and coral communities across the archipelago are quantified on the basis of overall biodiversity, abundance, and community structure. In Chapter 5, the existing network of MPAs in American Samoa is evaluated based on the habitats, reef fish, and coral communities that are encompassed. Appendices provide analytical details omitted from some chapters for brevity as well as supplemental datasets needed as inputs for the main chapters in the assessment. Appendices include an inventory of regional seamounts, a description of shore to shelf edge benthic maps produced for Tutuila, analytical details of reef fish and coral datasets, and supplemental information on the many marine protected areas in American Samoa.

Recruitment and spawning-stock biomass distribution of bay anchovy (Anchoa mitchilli) in Chesapeake Bay

Recruitment of bay anchovy (Anchoa mitchilli) in Chesapeake is related to variability in hydrological conditions and to abundance and spatial distribution of spawning stock biomass (SSB). Midwater-trawl surveys conducted for six years, over the entire 320-km length of the bay, provided information on anchovy SSB, annual spatial patterns of recruitment, and their relationships to variability in the estuarine environment. SSB of anchovy varied sixfold in 1995–2000; it alone explained little variability in young-of-the-year (YOY) recruitment level in October, which varied ninefold. Recruitments were low in 1995 and 1996 (47 and 31 Z 109) but higher in 1997–2000 (100 to 265 Z 109). During the recruitment process the YOY population migrated upbay before a subsequent fall-winter downbay migration. The extent of the downbay migration by maturing recruits was greatest in years of high freshwater input to the bay. Mean dissolved oxygen (DO) was more important than freshwater input in controlling distribution of SSB and shifts in SSB location between April– May (prespawning) and June–August (spawning) periods. Recruitments of bay anchovy were higher when mean DO was lowest in the downbay region during the spawning season. It is hypothesized that anchovy recruitment level is inversely related to mean DO concentration because low DO is associated with high plankton productivity in Chesapeake Bay. Additionally, low DO conditions may confine most bay anchovy spawners to the downbay region, where production of larvae and juveniles is enhanced. A modified Ricker stock-recruitment model indicated density-compensatory recruitment with respect to SSB and demonstrated the importance of spring-summer DO levels and spatial distribution of SSB as controllers of bay anchovy recruitment.

Larval development of the sidestriped shrimp (Pandalopsis dispar Rathbun) (Crustacea, Decapoda, Pandalidae) reared in the laboratory

Larval development of the sidestriped shrimp (Pandalopsis dispar) is described from larvae reared in the laboratory. The species has five zoeal stages and one postlarval stage. Complete larval morphological characteristics of the species are described and compared with those of related species of the genus. The number of setae on the margin of the telson in the first and second stages is variable: 11+12, 12+12, or 11+11. Of these, 11+12 pairs are most common. The present study confirms that what was termed the fifth stage in the original study done by Berkeley in 1930 was the sixth stage and that the fifth stage in the Berkeley’s study is comparable to the sixth stage that is described in the present study. The sixth stage has a segmented inner flagellum of the antennule and fully developed pleopods with setae. The ability to distinguish larval stages of P. dispar from larval stages of other plankton can be important for studies of the effect of climate change on marine communities in the Northeast Pacific and for marine resource management strategies.

Growth, mortality, and hatchdate distributions of larval and juvenile spotted seatrout (Cynoscion nebulosus) in Florida Bay, Everglades National Park

Life history aspects of larval and, mainly, juvenile spotted seatrout (Cynoscion nebulosus) were studied in Florida Bay, Everglades National Park, Florida. Collections were made in 1994−97, although the majority of juveniles were collected in 1995. The main objective was to obtain life history data to eventually develop a spatially explicit model and provide baseline data to understand how Everglades restoration plans (i.e. increased freshwater flows) could influence spotted seatrout vital rates. Growth of larvae and juveniles (<80 mm SL) was best described by the equation loge standard length = –1.31 + 1.2162 (loge age). Growth in length of juveniles (12–80 mm SL) was best described by the equation standard length = –7.50 + 0.8417 (age). Growth in wet weight of juveniles (15–69 mm SL) was best described by the equation loge wet-weight = –4.44 + 0.0748 (age). There were no significant differences in juvenile growth in length of spotted seatrout in 1995 between three geographical subdivisions of Florida Bay: central, western, and waters adjacent to the Gulf of Mexico. We found a significant difference in wet-weight for one of six cohorts categorized by month of hatchdate in 1995, and a significant difference in length for another cohort. Juveniles (i.e. survivors) used to calculate weekly hatchdate distributions during 1995 had estimated spawning times that were cyclical and protracted, and there was no correlation between spawning and moon phase. Temperature influenced otolith increment widths during certain growth periods in 1995. There was no evidence of a relationship between otolith growth rate and temperature for the first 21 increments. For increments 22–60, otolith growth rates decreased with increasing age and the extent of the decrease depended strongly in a quadratic fashion on the temperature to which the fish was exposed. For temperatures at the lower and higher range, increment growth rates were highest. We suggest that this quadratic relationship might be influenced by an environmental factor other than temperature. There was insufficient information to obtain reliable inferences on the relationship of increment growth rate to salinity.

Maturity, ovarian cycle, fecundity, and age-specific parturition of black rockfish (Sebastes melanops)

From 1995 to 1998, we collected female black rockfish (Sebastes melanops) off Oregon in order to describe their basic reproductive life history and determine age-specific fecundity and temporal patterns in parturition. Female black rockfish had a 50% probability of being mature at 394 mm fork length and 7.5 years-of-age. The proportion of mature fish age 10 or older significantly decreased each year of this study, from 0.511 in 1996 to 0.145 in 1998. Parturition occurred between mid-January and mid-March, and peaked in February. We observed a trend of older females extruding larvae earlier in the spawning season and of younger fish primarily responsible for larval production during the later part of the season. There were differences in absolute fecundity at age between female black rockfish with prefertilization oocytes and female black rockfish with fertilized eggs; fertilized-egg fecundity estimates were considered superior. The likelihood of yolked oocytes reaching the developing embryo stage increased with maternal age. Absolute fecundity estimates (based on fertilized eggs) ranged from 299,302 embryos for a 6-year-old female to 948,152 embryos for a 16-year-old female. Relative fecundity (based on fertilized eggs) increased with age from 374 eggs/g for fish age 6 to 549 eggs/g for fish age 16.

Description and growth of larval and pelagic juvenile pygmy rockfish (Sebastes wilsoni) (family Sebastidae)

A developmental series of larval and pelagic juvenile pygmy rockfish (Sebastes wilsoni) from central California is illustrated and described. Sebastes wilsoni is a non- commercially, but ecologically, important rockfish, and the ability to differentiate its young stages will aid researchers in population abundance studies. Pigment patterns, meristic characters, morphometric measurements, and head spination were recorded from specimens that ranged from 8.1 to 34.4 mm in standard length. Larvae were identified initially by meristic characters and the absence of ventral and lateral midline pigment. Pelagic juveniles developed a prominent pigment pattern of three body bars that did not extend to the ventral surface. Species identification was confirmed subsequently by using mitochondrial sequence data of four representative specimens of various sizes. As determined from the examination of otoliths, the growth rate of larval and pelagic juvenile pygmy rockfish was 0.28 mm/day, which is relatively slow in comparison to the growth rate of other species of Sebastes. These data will aid researchers in determining species abundance.

Identification of formalin-preserved eggs of red sea bream (Pagrus major) (Pisces: Sparidae) using monoclonal antibodies

Catches of important commercial fish such as red sea bream, flat fish, and yellowtail are decreasing in Japan. In order to sustain these species it is especially important that their distribution and biomass at all life stages are known. However, information on the early life stages of these species is limited because identifying the eggs and larvae of such fish is sometimes extremely difficult.

Elemental signatures in otoliths of larval walleye pollock (Theragra chalcogramma) from the northeast Pacific Ocean

The population structure of walleye pollock (Theragra chalcogramma) in the northeastern Pacific Ocean remains unknown. We examined elemental signatures in the otoliths of larval and juvenile pollock from locations in the Bering Sea and Gulf of Alaska to determine if there were significant geographic variations in otolith composition that may be used as natural tags of population affinities. Otoliths were assayed by using both electron probe microanalysis (EPMA) and laser ablation inductively coupled plasma mass spectrometry (ICP-MS). Elements measured at the nucleus of otoliths by EPMA and laser ablation ICP-MS differed significantly among locations. However, geographic groupings identified by a multivariate statistical approach from EPMA and ICP-MS were dissimilar, indicating that the elements assayed by each technique were controlled by separate depositional processes within the endolymph. Elemental profiles across the pollock otoliths were generally consistent at distances up to 100 μm from the nucleus. At distances beyond 100 μm, profiles varied significantly but were remarkably consistent among individuals collected at each location. These data may indicate that larvae from various spawning locations are encountering water masses with differing physicochemical properties through their larval lives, and at approximately the same time. Although our results are promising, we require a better understanding of the mechanisms controlling otolith chemistry before it will be possible to reconstruct dispersal pathways of larval pollock based on probe-based analyses of otolith geochemistry. Elemental signatures in otoliths of pollock may allow for the delineation of fine-scale population structure in pollock that has yet to be consistently revealed by using population genetic approaches.

Seasonal distribution, abundance, and growth of young-of-the-year Atlantic croaker (Micropogonias undulatus) in Delaware Bay and adjacent marshes

We examined the spatial and temporal distribution, abundance, and growth of young-of-the-year (YOY) Atlantic croaker (Micropogonias undulatus) in Delaware Bay, one of the northernmost estuaries in which they consistently occur along the east coast of the United States. Sampling in Delaware Bay and in tidal creeks in salt marshes adjacent to the bay with otter trawls, plankton nets and weirs, between April and November 1996–99, collected approximately 85,000 YOY. Ingress of each year class into the bay and tidal creeks consistently occurred in the fall, and the first few YOY appeared in August. Larvae as small as 2–3 mm TL were collected in September and October 1996. Epibenthic individuals <25 mm TL were present each fall and again during spring of each year, but not in 1996 when low water temperatures in January and February apparently caused widespread mortality, resulting in their absence the following spring and summer. In 1998 and 1999, a second size class of smaller YOY entered the bay and tidal creeks in June. When YOY survived the winter, there was no evidence of growth until after April. Then the YOY grew rapidly through the summer in all habitats (0.8–1.4 mm/d from May through August). In the bay, they were most abundant from June to August over mud sediments in oligohaline waters. They were present in both subtidal and intertidal creeks in the marshes where they were most abundant from April to June in the mesohaline portion of the lower bay. The larger YOY began egressing out of the marshes in late summer, and the entire year class left the tidal creeks at lengths of 100–200 mm TL by October or November when the next year class was ingressing. These patterns of seasonal distribution and abundance in Delaware Bay and the adjacent marshes are similar to those observed in more southern estuaries along the east coast; however, growth is faster—in keeping with that in other northern estuaries.

An approach to estimating rockfish biomass based on larval production, with application to Sebastes jordani

An assessment of the total biomass of shortbelly rockfish (Sebastes jordani) off the central California coast is presented that is based on a spatially extensive but temporally restricted ichthyoplankton survey conducted during the 1991 spawning season. Contemporaneous samples of adults were obtained by trawl sampling in the study region. Daily larval production (7.56 × 1010 larvae/d) and the larval mortality rate (Z=0.11/d) during the cruise were estimated from a larval “catch curve,” wherein the logarithm of total age-specific larval abundance was regressed against larval age. For this analysis, larval age compositions at each of the 150 sample sites were determined by examination of otolith microstructure from subsampled larvae (n=2203), which were weighted by the polygonal Sette-Ahlstrom area surrounding each station. Female population weight-specific fecundity was estimated through a life table analysis that incorporated sex-specific differences in adult growth rate, female maturity, fecundity, and natural mortality (M). The resulting statistic (102.17 larvae/g) was insensitive to errors in estimating M and to the pattern of recruitment. Together, the two analyses indicated that a total biomass equal to 1366 metric tons (t)/d of age-1+ shortbelly rockfish (sexes combined) was needed to account for the observed level of spawning output during the cruise. Given the long-term seasonal distribution of spawning activity in the study area, as elucidated from a retrospective examination of California Cooperative Oceanic Fisheries Investigation (CalCOFI) ichthyoplankton samples from 1952 to 1984, the “daily” total biomass was expanded to an annual total of 67,392 t. An attempt to account for all sources of error in the derivation of this estimate was made by application of the delta-method, which yielded a coefficient of variation of 19%. The relatively high precision of this larval production method, and the rapidity with which an absolute biomass estimate can be obtained, establishes that, for some species of rockfish (Sebastes spp.), it is an attractive alternative to traditional age-structured stock assessments.

Larval development of the southern sea garfish (Hyporhamphus melanochir) and the river garfish (H. regularis) (Beloniformes: Hemiramphidae) from South Australian waters

Larval development of the southern sea garfish (Hyporhamphus melanochir) and the river garfish (H. regularis) is described from specimens from South Australian waters. Larvae of H. melanochir and H. regularis have completed notochord flexion at hatching and are characterized by an elongate body with distinct rows of melanophores along the dorsal, lateral, and ventral surfaces; a small to moderate head; a heavily pigmented and long straight gut; a persistent pre-anal finfold; and an extended lower jaw. Fin formation occurs in the following sequence: caudal, dorsal and anal (almost simultaneously), pectoral, and pelvic. Despite the similarities between both species and among hemiramphid larvae in general, H. melanochir larvae are distinguishable from H. regularis by 1) having 58–61 vertebrae (vs. 51–54 for H. regularis); 2) having 12–15 melanophore pairs in longitudinal rows along the dorsal margin between the head and origin of the dorsal fin (vs. 19–22 for H. regularis); and 3) the absence of a large ventral pigment blotch anteriorly on the gut and isthmus (present in H. regularis). Both species can be distinguished from similar larvae of southern Australia (other hemiramphids and a scomberosocid) by differences in meristic counts and pigmentation.

Larval abundance, distribution, and spawning habits of spotted seatrout (Cynoscion nebulosus) in Florida Bay, Everglades National Park, Florida

The spotted seatrout (Cynoscion nebulosus) is one of the most sought after recreational fish in Florida Bay, and it spends its entire life history within the bay (Rutherford et al.,1989b). The biology of adult spotted seatrout in Florida Bay is well known (Rutherford et al., 1982, 1989b) as is the distribution and abundance of juveniles within the bay. The habitats and diets of juveniles are well documented (Hettler, 1989; Chester and Thayer, 1990; Thayer et al., 1999; Florida Department of Environmental Protection1). Nevertheless, the spatial and temporal spawning habits of spotted seatrout and the distribution of larvae have only been partially described (Powell et al., 1989; Rutherford et al., 1989a).

The early life history of swordfish (Xiphias gladius) in the western North Atlantic

Lengths and ages of sword-fish (Xiphias gladius) estimated from increments on otoliths of larvae collected in the Caribbean Sea, Florida Straits, and off the southeastern United States, indicated two growth phases. Larvae complete yolk and oil globule absorption 5 to 6 days after hatching (DAH). Larvae <13 mm preserved standard length (PSL) grow slowly (~0.3 mm/d); larvae from 13 to 115 mm PSL grow rapidly (~6 mm/d). The acceleration in growth rate at 13 days follows an abrupt (within 3 days) change in diet, and in jaw and alimentary canal structure. The diet of swordfish larvae is limited. Larvae <8 mm PSL from the Caribbean, Gulf of Mexico, and off the southeastern United States eat exclusively copepods, primarily of one genus, Corycaeus. Larvae 9 to 11 mm eat copepods and chaetognaths; larvae >11 mm eat exclusively neustonic fish larvae. This diet indicates that young larvae <11 mm occupy the near-surface pelagia, whereas, older and longer larvae are neustonic. Spawning dates for larvae collected in various regions of the western North Atlantic, along with the abundance and spatial distribution of the youngest larvae, indicate that spawning peaks in three seasons and in five regions. Swordfish spawn in the Caribbean Sea, or possibly to the east, in winter, and in the western Gulf of Mexico in spring. Elsewhere swordfish spawn year-round, but spawning peaks in the spring in the north-central Gulf of Mexico, in the summer off southern Florida, and in the spring and early summer off the southeastern United States. The western Gulf Stream frontal zone is the focus of spawning off the southeastern coast of the United States, whereas spawning in the Gulf of Mexico seems to be focused in the vicinity of the Gulf Loop Current. Larvae may use the Gulf of Mexico and the outer continental shelf off the east coast of the United States as nursery areas. Some larvae may be transported northward, but trans-Atlantic transport of larvae is unlikely.

Reproduction and recruitment of white mullet (Mugil curema) to a tropical lagoon (Margarita Island, Venezuela) as revealed by otolith microstructure

The reproductive activity and recruitment of white mullet (Mugil curema) was determined by observations of gonad development and coastal juvenile abundance from March 1992 to July 1993. Adults were collected from commercial catches at three sites in northeastern Venezuelan waters. Spawning time was determined from the observation of macroscopic gonadal stages. Coastal recruitment was determined from fish samples collected biweekly by seining in La Restinga Lagoon, Margarita Island, Venezuela. The examination of daily growth rings on the otoliths of coastal recruits was used to determine their birth date and estimate the period of successful spawning. Fish with mature gonads were present throughout the year but were less frequent between September and January when spawning individuals migrated offshore. In both years, juvenile recruitment to the lagoon was highest between March and June when high densities of 25–35 mm juveniles were observed. Back-calculated hatching-date frequency distributions revealed maximum levels of successful spawning in December–January that were significantly correlated with periods of enhanced upwelling. The relation between the timing of successful spawning and the intensity of coastal recruitment in white mullet was likely due to variations in food availability for first-feeding larvae as well as to variations in the duration of the transport of larvae shoreward as a result of varying current conditions associated with upwelling.

Larval and settlement periods of the northern searobin (Prionotus carolinus) and the striped searobin (P. evolans)

This study reports new information about searobin (Prionotus spp.) early life history from samples collected with a Tucker trawl (for planktonic stages) and a beam trawl (for newly settled fish) from the coastal waters of New Jersey. Northern searobin, Prionotus carolinus, were much more numerous than striped searobin, P. evolans, often by an order of magnitude. Larval Prionotus were collected during the period July–October and their densities peaked during September. For both species, notochord flexion was complete at 6–7 mm standard length (SL) and individuals settled at 8–9 mm SL. Flexion occurred as early as 13 days after hatching and settlement occurred as late as 25 days after hatching, according to ages estimated from sagittal microincrements. Both species settled directly in continental shelf habitats without evidence of delayed metamorphosis. Spawning, larval dispersal, or settlement may have occurred within certain estuaries, particularly for P. evolans; thus collections from shelf areas alone do not permit estimates of total larval production or settlement rates. Reproductive seasonality of P. carolinus and P. evolans may vary with respect to latitude and coastal depth. In this study, hatching dates and sizes of age-0 P. carolinus varied with respect to depth or distance from the New Jersey shore. Older and larger age-0 individuals were found in deeper waters. These variations in searobin age and size appear to be the combined result of intraspecific variations in searobin reproductive seasonality and the limited capability of searobin eggs and larvae to disperse.