Composition and distribution of larval fishes in New Jersey high marshes

The surface of the salt marsh is an important, but largely unrecognized, site for fish reproduction and larval growth. In an attempt to determine the composition and distribution of fishes utilizing these habitats, we sampled larval and juvenile fish with plankton nets, dip nets, and traps at a variety of microhabitats (tidal and nontidal ponds and ditches and the marsh surface) in three New Jersey high marshes. Two of the three marshes had been altered for mosquito control. During April to September 1980, we collected over 2,400 larvae and juveniles. All study sites were dominated by the larvae of the resident killifishes (Fundulus heteroclitus, Cyprinodon variegatus, F. luciae, andLucania parva) and less commonlyMenidia beryllina. However, the occurrence and abundance of each species varied with microhabitat. Larval production in all three marshes peaked during June–July, but extended from May until September. In most instances juveniles of the dominant fishes had microhabitat preferences similar to the larvae. High marshes may be more important for fish production than previously recognized because they serve as nursery areas for the resident killifishes.     

Benthic macrofaunal communities in partially impounded salt marshes in Delaware: Comparisons with natural marshes and responses to sediment exposure

This two-part study examined the benthic macrofaunal community in Delaware salt marsh impoundments having partial tidal restriction. The first part compared abundance, diversity, and taxonomic composition in three habitat types in impoundments—creeks, vegetated creek banks, and ponds—to those found in natural marshes. Impoundment effects were present but were habitat-specific. Abundances were higher in natural marsh creeks than in impoundment creeks, and diversities were higher in impoundment ponds than in natural marsh ponds. Vegetated bank communities in impoundments were about 50% insects and arachnids and 50% oligochaetes, while natural bank communities were primarily oligochaetes and the polychaeteManayunkia aestuarina. This is likely due to the decrease in flooding of the vegetated high marsh caused by partial impoundment. Pond and creek community composition also showed impoundment effects: in comparison with natural marshes, impoundments had higher proportions of the burrowing anemoneNematostella vectensis, nemerteans, andTubificoides sp. oligochaetes and lower proportions of the oligochaeteClitellio arenarius. The second part of the study compared benthic macrofauna in an impoundment before, during, and after the water level was lowered so that some bottom sediments were exposed and some covered with just a few centimeters of water for several weeks. During this event, macrofaunal abundances were reduced and the community shifted from being dominated by annelids, anemones, and nemerteans toward one dominated by annelids and insects. About 6 wk after reflooding, persistent effects of this disturbance were still suggested by greatly increased abundances and 96% dominance by one species of oligochaete,Paranais litoralis. Impoundment management plans calling for periods of sediment exposure or very low water may want to consider the potential for strong and persistent effects on the macrofaunal community.     

The ichthyofauna of the sundays estuary,South Africa,with particular reference to the juvenile marine component

The ichthyofauna of the Sundays Estuary was investigated by monthly seine netting over a period of a year. Forty-seven species were captured though 23 of these were represented by a total catch of less than 25 specimens each. The small clupeidGilchristella aestuarius was numerically dominant and constituted 80%, of the catch. The study confirms that in addition to this species, two species of goby,Caffrogobius multifasciatus andPsammogobius knysnaensis and the soleSolea bleekeri complete their life cycles in the estuary. Many other species such as mullet, utilize the estuary as a juvenile nursery area. First year juveniles ofRhabdosargus holubi, Lithognathus lithognathus, Pomadasys commersonni andMonodactylus falciformis, by virtue of their abundance in the Sundays Estuary and other South African estuaries, and their absence from other coastal environments, appear to be dependent on estuaries as juvenile nursery areas.     

Community structure of the demersal fish species of Laguna Joyuda,Puerto Rico

A year-long trawl survey of the mangrove-fringed Laguna Joyuda, Puerto Rico yielded 41 species of juvenile and adult fishes. Twelve percent of the species and 55% of the individuals were residents in the lagoon; 56% of the species and 44% of the individuals were cyclic visitors, mostly juveniles of species which spawn offshore. The lagoon yielded fewer species than tropical estuaries in continental regions of the Caribbean and temperate estuaries of North America. However, paralleling other estuarine habitats, Laguna Joyuda supported three primary types of fishes, a resident small flatfish (Achirus lineatus), a complex of transient juveniles (Gerreidae, paralleling the Sciaenidae of higher latitudes), and small planktivores (Anchoa spp.). Seasonal patterns in the abundance of individual species were not strong, but overall abundance showed a wet season maximum, particularly because of recruitment of anchovies and cyclic visitors in April, May, and June. Wet season abundance corresponded with highest water column productivity and detrital input to the benthos. Fishes associated with the water column appeared to show greater variation in abundance than those associated with more stable benthic food webs.     

Stable isotope indicators of movement and residency for brown shrimp (<Emphasis Type="Italic">Farfantepenaeus aztecus</Emphasis>) in coastal Louisiana marshscapes

Brown shrimp (Farfantepenaeus aztecus) are an important commercial aquatic species experiencing loss of inshore marsh nursery habitat in coastal Louisiana. To study inshore brown shrimp movements and identify aspects of essential habitat important for sustaining brown shrimp populations, we collected juvenile brown shrimp in April and May 2000, the time of annual maximum brown shrimp abundance, in a small 1-km² marsh area on the central Louisiana coast. Drop sampling showed average shrimp densities of 1.6–2.4 m⁻² in shallow marsh ponds and seining indicated lower densities of 0.5–0.9 m⁻² in nearby shallow channel and open bay sites. Smaller shrimp (< 50 mm) fed disproportionately on benthic diatoms and small harpacticoid copepods, while large shrimp fed more frequently on larger-bodied amphipods and tanaids. We developed novel chemical approaches to estimate patterns of shrimp residency and movement using carbon and nitrogen stable isotopic determinations. Resident shrimp had isotopic values similar to average foods and showed consistent isotopic spacings between fast and slow turnover tissues. Residency was highest (47–55%) in ponds and shallow channel habitats and much less in open bays and deep channels (4–27%). There was sparse evidence for dietary specialization among individull shrimp. The results support the view that small 10–20 mm postlarval and juvenile brown shrimp arriving in estuaries from offshore waters continue movement through sub-optimal habitats (deep channels and open bays), but exhibit much less movement once an optimal habitat (marsh ponds or shallow channel margins) is reached. This study also indicated that combining estimates of shrimp densities, residency, growth rate, and mortality allows evaluation of the importance of different habitat types for shrimp production. Shallow ponds that in many ways resemble fertile aquaculture ponds appear to be hot spots for brown shrimp production, and coastal preservation and restoration efforts should focus on these areas as important for sustaining shrimp fisheries.     

Interactions between Northern-Shifting Tropical Species and Native Species in the Northern Gulf of Mexico

Niche models applied in the context of future climate change predict that as regional temperatures increase, the distribution of tropical species will shift poleward. While range expansions have been documented for a number of species, there is limited information on the ecological impacts of shifts on native species. Recently, abundances of tropically-associated gray snapper (Lutjanus griseus) and lane snapper (Lutjanus synagris) have increased in seagrass nurseries in the northern Gulf of Mexico (GOM), concurrent with regional increases in sea surface temperature. We investigated effects of increased abundances of these species on abundance and growth of pinfish (Lagodon rhomboides), the dominant native species. Because juvenile pinfish and snappers share common prey, predators, and habitat, the high degree of niche overlap suggests an equally high potential for competition. We used a multiple before–after control impact design to determine whether increased snapper abundances significantly affected abundance or growth of pinfish. Trawl surveys at six locations in the northern GOM in summer and fall 2010 were used to calculate pinfish and snapper abundances. We identified three locations with high snapper abundances and three locations with no snapper and compared pinfish abundance and otolith-determined growth rates in these locations before and after snapper recruitment. Paired t tests and two-way analysis of variance revealed no significant differences in pinfish abundance or growth in the presence of snappers compared to locations and seasons without snappers. We conclude that range expansions of tropically associated snappers have had no significant effect on abundance or growth of native pinfish in northern GOM seagrass habitats.     

New data on diatoms from the marine Paleogene of Western Kamchatka

Diatom assemblages from the marine Paleogene of Western Kamchatka (the Kovachina, Viventek, and Kuluven formations) are studied. The shallow-water neritic assemblage from the Kovachina Formation of the Pyatibratskii Cape section is tentatively attributed to the middle Eocene, as it is lacking zonal index species. A high abundance and great taxonomic diversity are characteristic of diatoms from the upper part of the Viventek Formation in the Kovran River section. The zonal index species Cavitatus rectus identified here among the diatoms suggests that the respective assemblage is macerated from the synonymous diatom zone of the middle Oligocene (29.6?28.2 Ma). According to its taxonomic composition and structure, this assemblage is indicative of transgression environments and the presumable influence of the middle Oligocene climatic optimum. The Rocella gelida (upper part) and Thalassiosira praefraga diatom zones of the Oligocene-Miocene transition are established in the Kuluven Formation sediments of the Mt. Uvuch section in Kovachina Bay.     

The fish community of a shallow tropical lagoon in Belize,Central America

Trawl collections indicate that the fish community of the Belize barrier reef lagoon is dominated numerically and in biomass by grunts (Haemulidae), especiallyHaemulon sciurus andHaemulon flavolineatum. Although the gear selected for small sizes, length frequency analysis indicated seasonality in recruitment of the dominant species of grunts. Apogonids and tetraodontiform fishes were also dominant components of the community. Most fishes collected were juveniles of species that occur as adults on the main reef, or were small species that are resident in the lagoon. Of three habitats sampled, the mangrove creek had the greatest relative abundance and biomass of fishes, followed by the seagrass bed and the sand-rubble zone. There were no significant seasonal differences in fish relative abundance or biomass. Community structure analysis indicated a uniqueness in the mangrove fish community. Diversity (H′) was high, and was due to high species richness and evenness of distribution of individuals among species. The Belize barrier reef lagoon serves as an important nursery habitat for juvenile fishes.     

Miocene freshwater diatoms from the eastern slope of the submarine Ulleung Plateau (Krishtofovich Rise) in the Sea of Japan

The freshwater diatom flora from the tuffogenous sedimentary rocks of the submarine Ulleung Plateau of the southern Sea of Japan is studied and its complete taxonomic composition (96 species and varieties belonging to 33 genera) is revealed. It is characterized by species diversity and abundance of representatives of the freshwater genus Aulacoseira. The predominance of Aulacoseira is characteristic of the Miocene lacustrine deposits of the Sea of Japan and other regions, but the presence of ellipsoid species of narrow stratigraphic distribution allows the age of the recognized diatom flora to be limited to the Early Miocene. The species diversity and abundance of diatoms in the deposits of the Ulleung Plateau indicate the presence in the Early Miocene of a large freshwater lake in this region. Such lakes also existed on other large submarine rises of the Sea of Japan and its periphery before the onset of the tectonic activation which led to the descent of vast fragments of the continent and expansion of the Sea of Japan.     

The potential role of roots and rhizomes in structuring salt-marsh benthic communities

The density of the Carolina marsh clam,Polymesoda caroliniana (Bosc), was determined in three adjacent tidal marsh communities which differed only in plant species composition. Clam density was inversely related to the density (biomass) of plant roots and rhizomes in sediments and directly related to density of plant stems (numbers). Clam abundance was not related to the basal area of plant stems. Each plant community contained clams of various ages from juvenile to adult indicating continued recruitment and survival. These data suggest thatP. caroliniana is most abundant inJuncus roemerianus marshes because there are fewer roots and rhizomes (mean of 2.5 kg m^?2) to hamper burrwing as compared toSpartina alterniflora andcynosuroides (5.1 and 6.3 kg m^?2, respectively) dominated marshes. Salinity, floding frequency, distance from flooding water, and sediment type were essentially constant among the three plant communities. Root/rhizome density should be collected along with other environmental parameters during studies of benthic organisms on marshes because it potentially limits the occurrence or abundance of some species.     

Seasonal occurrence of larval and juvenile fishes in a Virginia Atlantic coast estuary with emphasis on drums (Family sciaenidae)

The seasonal occurrence and relative abundance of larval and juvenile fishes, particularly members of the family Sciaenidae, from a Virginia Atlantic coast estuary were determined from ichthyoplankton and otter trawl collections made from March 1979 to March 1980. The larvae of 19 species in 14 families were identified in the ichthyoplankton. Larvae of the engraulid, Anchoa mitchilli (bay anchovy), and the atherinid, Menidia menidia (Atlantic silverside), dominated the samples and made up 13 and 22%, respectively, of the 9,440 larvae collected. Peak occurrence of all larvae was from May to August. The juveniles of 28 species in 19 families were identified from otter trawl collections. Juvenile sciaenids numerically dominated the trawl collecions and made up 68% of the trawl catch. Juvenile density peaked during September through December.     

A new lizard (Reptilia: Squamata) from the Lower Cretaceous Yixian Formation of China,with a taxonomic revision of Yabeinosaurus

Yabeinosaurus was the first lizard genus described from the Early Cretaceous Jehol Biota of China. The holotype of the type species, Y. tenuis, is an immature skeleton but it has been lost for decades. A second species, Y. youngi, was erected based on another immature skeleton and is distinguished by its longer, more gracile limbs. In 2001, a juvenile skeleton from the Jingangshan locality of Liaoning was designated as the neotype for Y. tenuis. Subsequently, several further specimens from other localities, juvenile and adult, have been attributed to this species. Here we describe a second lizard specimen from the neotype locality. In anterior tooth shape, finger-like mandibular angular process, cranial sculpture, maxilla shape, and short, robust humerus, the new specimen resembles adult and subadult material currently attributed to Y. tenuis, but it differs in having bicuspid posterior teeth and a straight rather than a hooked angular process of the mandible. We propose a new species (Y. bicuspidens sp. nov.) for this specimen which represents the first occurrence of bicuspid teeth in an Early Cretaceous lizard from China. However, the recognition of a second species at the neotype locality raises a taxonomic problem. Due to the immaturity of the designated holotype and neotype of Y. tenuis, they cannot be coded for the jaw and dental characters that distinguish the two robust-limbed species. This renders Yabeinosaurus tenuis a nomen dubium. Here we propose that Yabeinosaurus tenuis should be treated as a historic taxon, permitting retention of the generic name under ICZN rules, and erect a second new species (Y. robustus sp. nov.) for the principal robust-limbed morphotype in the Jehol Biota with monocuspid posterior teeth and a hooked angular process.     

The distribution of shallow water juvenile fishes in an urban estuary: The effects of manmade structures in the lower Hudson River

The objective of this study was to determine what effect, if any, large pile-supported platforms (piers) have on the habitat distribution and abundance of juvenile fishes. Trapping techniques were used in 1993 and 1994 under piers, in pile fields, and in open-water habitat types in shallow areas (<5 m) in the lower Hudson River estuary (40°44′N, 70°01′W). Nearly 1500 fishes, mostly juveniles, representing 24 species were collected in 1865 trap-days from May through October in the 2-yr study. The presence of relatively large numbers of young-of-the-year (YOY) fish during both years lends support to the idea that shallow areas in the lower Hudson River estuary currently function as nursery habitats for a variety of fishes. Two seasonal assemblages were apparent, but their composition varied somewhat between years.Microgadus tomcod andPseudopleuronectes americanus YOY dominated an early summer assemblage (May–July) while large numbers of YOYMorone saxatilis were collected as part of a late summer assemblage (August–September). The effects of habitat type on fish assemblage structure were significant during both years. Fish abundance and species richness were typically low under piers; YOY fishes were rare andAnguilla rostrata accounted for a large proportion of the total catch. In contrast, YOY fishes dominated collections at pile field and open-water stations, where abundance and species richness were high. These results indicate that habitat quality under the platforms of large piers (>20,000 m²) is probably poor for YOY fishes when compared with nearby pile field and open-water habitat types.     

Small sized charophyte gyrogonites in the Maastrichtian of Coll de Nargó, Eastern Pyrenees: An adaptation to temporary floodplain ponds

Charophytes bearing small sized fructifications dominated in fluviatile floodplain facies (red beds) from the Maastrichtian of Coll de Nargó and neighbouring basins in the southern Pyrenees (Catalonia, Spain). These charophytes mainly belong to the genus Microchara and are often disregarded in biostratigraphic studies, which focus their attention instead on facies from permanent lakes that are richer in species which usually bear fructifications with a larger size range. However, small sized gyrogonites are also significant for biostratigraphic purposes and even include some of the index species for Maastrichtian biozones in Europe. Indeed, the charophyte assemblages from the Maastrichtian of Coll de Nargó belong to the Microchara punctata biozone, recently calibrated to the middle-upper Maastrichtian.Floodplain ponds from the Maastrichtian red beds of Coll de Nargó (Lower Red Unit) were extremely shallow, received considerable terrigenous influx and were frequently exposed, probably resulting in turbid, warm waters with high light availability. These conditions could explain the almost exclusive occurrence of charophytes with small fructifications in the Lower Red Unit. The available data, mainly based on oospores from extant species, indicate that the small size observed in gyrogonites from temporary ponds may represent an adaptation to environmental stress. Fossil species with small gyrogonites of Microchara cristata, M. punctata and Microchara nana would thus develop massively in stressed shallow ponds on fluvial floodplains. To contrast these hypotheses, we compared our results to those of four well-known case studies with similar sedimentological contexts, ranging from the Lower Cretaceous to the upper Eocene–lower Oligocene. Gyrogonite size patterns were similar in all cases, possibly suggesting that characeans display a long history of adaptation to shallow, temporary and turbid floodplain ponds by means of producing a high number of small gyrogonites, probably representing short life cycles and opportunistic strategies.     

Spatio-temporal fluctuations in the distribution and abundance of demersal fish and epibenthic crustaceans in Yaquina Bay,Oregon

A total of over 32,000 demersal fish and epibenthic crustaceans belonging to 62 species were caught in 42 biweekly trawls from 10 stations in Yaquina Bay, Oregon, during 1967 and 1968. English sole,Parophrys vetulus, was the most abundant species. Seventeen species (13 fishes and 4 crustaceans) constituted 95% of the catch. Total numerical abundances of both individuals (mainly juvenile fishes) and species were greatest in the lower 12 km of the estuary during summer and early fall, a period of water mass stability and increased water temperature and salinity. This section of the estuary is used by many immature fishes and crustaceans as a “nursery area”. These fishes generally emigrate from the estuary as subadults in the fall around the onset of the rainy season. The fewest species were taken in January 1968 from the central, upper-estuarine, and riverine areas of the bay, this being a time when high rainfall and river discharge result in low salinity and temperature. Crustaceans (shrimp and subadult crabs) were generally most abundant in late winter and early spring throughout the estuary. Changes in diversity indices reflected variations in community structure, the influence of migratory species and juvenile fishes, and seasonal changes in dominance. Year-to-year fluctuations in abundance may be due, in part, to local hydrographic and meteorological conditions along the central Oregon coast.     

The abundance and diversity of larval and juvenile fish in a tropical estuary

The larval and juvenile fish of the Cayenne river estuary (French Guiana, South America) were sampled at two stations from June 1989 until October 1990. A total of 52,989 individuals from 59 species, some still incompletely identified, were collected. Three families, Engraulidae, Gobiidae and Sciaenidae, accounted for over 97% of the total number of juveniles. The analysis of data over this period showed low diversity, and a difference in diversity between the two sampling locations (H′ = 1.24 and 1.68). The results conform to some theoretical models of abundance that suggest a relative equilibrium of juvenile assemblages. In contrast, the seasonal variations in diversity and abundance and the results of a correspondence analysis showed significant differences in species distribution and in their relative abundance at the two sampling locations at certain periods, mainly in the rainy season. Our study indicates that, in spite of an apparent stability, the year to year variation in salinity and freshwater inputs could affect juvenile recruitment of some species and induces modifications in the composition of larval and juvenile estuarine fish assemblages.     

Lentic and lotic assemblages of zooplankton in a tropical reservoir,and their association with water quality conditions

Zooplankton assemblages in relation to water quality parameters at lentic and lotic habitats of Air Itam Reservoir, Penang, Malaysia, were analysed. Five sampling stations were designated, including three stations located in the reservoir (Stations A, B, and C) and two stations in the upstream inflow and downstream outflow (Stations D and E, respectively). The most common and dominant zooplankton found in the lake were Polyarthra vulgaris, Ceriodaphnia cornuta, Mesocyclops leuckarti, and Thermocyclops crassus. Water Quality Index, calculated as recommended by the Malaysian Department of Environment, ranged between 88.46 and 93.6, showed that the reservoir was clean. Low numbers of species and value of the Shannon–Wiener Diversity Index were recorded at stations located upstream and downstream of the reservoir. Cluster analysis based on the abundance of zooplankton species distinguished the sampling stations into two groups (lentic and lotic groups, comprised of Stations A, B, and C; and Stations D and E, respectively). This study showed that zooplankton occurrence and abundance were associated with the quality of their environment, and zooplankton community provides information about the reservoir ecosystem, thus reflecting the importance of biomonitoring in lake assessment and management.     

The Effects of Integrated Marsh Management (IMM) on Salt Marsh Vegetation,Nekton, and Birds

An integrated marsh management (IMM) project in an urbanized watershed on Long Island, New York, USA, aimed to mitigate salt marsh degradation and to reduce mosquito production by an innovative combination of restoration and open marsh water management methods. The grid ditch network at two treatment marshes was replaced with naturalized tidal channels and ponds. Effects of the hydrologic alterations were monitored utilizing a before–after–control–impact approach. The treatment marshes experienced a number of beneficial outcomes including a fourfold reduction in the invasive Phragmites australis and increased native vegetation cover in the most degraded portions of the marsh, increased abundance and diversity of marsh killifish and estuarine nekton species, higher shorebird and waterfowl densities, and increased avian species diversity. The successful implementation of IMM concept led to improved marsh health and diminished mosquito production. Therefore, this study may serve as a template for similar large-scale integrated salt marsh restoration projects.     

Posthurricane Recovery of Riverine Fauna Reflected in the Diet of an Apex Predator

Hurricane Charley, a category 4 storm on the Safford–Simpson scale, passed over the Peace River, Florida, and its watershed on 13 August 2004 causing widespread hypoxia. An electrofishing study investigating the abundance of the euryhaline Centropomus undecimalis (common snook) began in the freshwater portions of the Peace River 3 months after the storm (November 2004) and included diet sampling. Samples were taken seasonally up to August 2006. Changes in the prey assemblages of C. undecimalis determined through multivariate analyses (e.g., nonmetric multidimensional scaling) were used as a proxy to estimate the recovery of riverine fauna. The initial posthurricane prey assemblages were dominated by species well adapted to low dissolved oxygen, namely Gambusia holbrooki and Hoplosternum littorale. A significant long-term serial change in assemblage structure occurred as the abundance of G. holbrooki and H. littorale decreased and other river fauna such as Procambarus spp. and ictalurid catfish increased. Assemblages were presumed as recovering 1 year after hurricane landfall, a posthurricane recovery period that is an order of magnitude longer than those reported for estuarine fish assemblages. The approach of using stomachs as sampling tools detected changes at the assemblage level that included an armored catfish species (e.g., H. littorale) and cryptic invertebrate species (e.g., Procambarus spp.) that went unnoticed in a study using standard electrofishing methods. Although sampling the diet of C. undecimalis was useful in detecting assemblage-level changes, other metrics—such as changes in health, sex ratio, relative abundance, and juvenile year-class strength of fishes and invertebrates collected by traditional fishing methods—are needed to fully evaluate the effects of the hurricane. Nonetheless, this study demonstrates that diet studies can supplement standard fishery gear to create a more comprehensive sampling strategy for aquatic fauna.     

Abundance and seasonality of infauna and epifauna inhabiting a Halodule wrightii meadow in Apalachicola Bay,Florida

The fauna inhabiting a Halodule wrightii meadow in Apalachicola Bay, Florida, was studied from March 1975 through Feburary 1976. The infaunal community was sampled by monthly coring. Fifty-eight species were recorded, averaging 35 species per month. A maximum faunal abundance of 104,338 organisms per m² in April was one of the highest infaunal densities recorded in the literature. Sixteen species accounted for 84% of the total numbers and 80% of the total biomass over the study period. Numerical dominants were Hargeria rapax, Heteromastus filiformis, Ampelisca vadorum, Aricidea fragilis, and oligochaetes. Biomass dominants were Tagelus plebeius, Neritina reclivata, Ensis minor, and Haploscoloplos fragilis. Life history notes are given for several dominant species. Epibenthic fishes and macroinvertebrates were sampled by monthly trawling. Twenty-three species of fishes (mostly juveniles) were collected near the coring site, with most species and individuals recorded during the months May through September. Bairdiella chrysoura, Orthopristis chrysoptera, and Lagodon rhomboides comprised 76% of the total fish numbers. Eleven species of macroinvertebrates were collected mainly in June and July. Callinectes sapidus comprised 61% of the total invertebrate numbers. It is postulated that the influx of juvenile fishes and crabs into the Halodule meadow in summer months leads to a coincident decline in infaunal population densities (number per m²) through predation. Infaunal biomasses are largely unaffected by these predators since the biomass dominants are large or deep-burrowing species.