Food habits of fishes in the surf zone of a sandy beach at Sanrimatsubara, Fukuoka Prefecture, Japan

To clarify the feeding habits of fishes in surf zones, the gut contents of 19 fish species collected in the surf zone of a sandy beach at Sanrimatsubara, western Japan, were examined. Ontogenetic changes in food preference were recognized in seven species (Mugil cephalus cephalus, Lateolabrax latus, Sillago japonica, Paralichthys olivaceus, Paraplagusia japonica, Takifugu poecilonotus, and Takifugu niphobles). A cluster analysis based on dietary overlaps showed that the surf zone fish assemblage comprised six trophic groups (zooplankton, benthic and epiphytic crustacean, detritus, polychaete, fish, and insect feeders). Of these, the most abundant trophic group was zooplankton feeders, along with benthic and epiphytic crustacean feeders.     

Seasonal dynamics of fish assemblage structures in a surf zone on an exposed sandy beach in Japan

Seasonal dynamics in fish assemblage structure, comprising postlarval to early adult stages, in the surf zone on an exposed sandy beach on the Kashimanada coast were studied over a 2-year period. A total of 32 species and two higher group taxa were found. In terms of abundance, Salangichthys ishikawae, Mugil cephalus cephalus, Plecoglossus altivelis altivelis, and Engraulis japonicus were the four most dominant species. The results of cluster analysis and canonical correspondence analysis using 12 dominant species in terms of abundance revealed that water temperature and wind factor (wind speed × wind direction) had significant independent associations on species occurrence patterns for 11 species; that is, occurrence of 11 species was significantly related to water temperature and occurrence of 7 species was related to wind factor. Two species (S. ishikawae and P. altivelis altivelis) showed clear monthly increases in size, consequently suggesting their use of the surf zone as a nursery area. In contrast, M. cephalus cephalus and E. japonicus showed few increases in standard length, indicating their use of the surf zone only in their postlarval and/or juvenile stage. It is suggested that numerous fish species use the surf zone as shelter and/or a nursery area, even in the harsh conditions of an exposed sandy beach with great wave action.     

Partitioning of food resources among Sillago japonica, Ditremma temmincki, Tridentiger trigonocephalus, Hippocampus japonicus and Petroscirtes breviceps in an eelgrass, Zostera marina, bed

We carried out dietary analysis on five numerically abundant fishes, Sillago japonica, Ditremma temmincki, Tridentiger trigonocephalus, Hippocampus japonicus and Petroscirtes breviceps in an eelgrass bed in Kwangyang Bay, Korea. Comparisons between species demonstrated that the dietary composition of each fish species was significantly different from that of every other species. Although gammarid amphipods and caprellid amphipods were consumed by all species, their individual contributions to each species' diet varied. Furthermore, polychaetes contributed to the diets of S. japonica and T. trigonocephalusand crab larvae were consumed byD. temmincki. Algae and eelgrass were not consumed by four fish species and made only a minimal contribution to the diet of P. breviceps. The diet of each fish species except H. japonicus underwent size-related changes; smaller fishes consumed gammarid amphipods, mysids and copepods, while larger fishes ate polychaetes, gastropods, isopods and other fishes. Differences in the prey organisms consumed of each individual species could be often related to differences in mouth length and width. S. japonica, D. temmincki, T. trigonocephalus, and P. breviceps underwent also a significant diel changes that could be related to differences in foraging behavior and/or prey availability. Thus, use of vision to detect prey would account for the greater daytime consumption of copepods by S. japonica and of crab larvae by D. temmincki, whereas the nocturnal emergence of gammarid amphipods, polychaetes and isopods from the substrate explained their greater consumption by S. japonica, D. temmincki, T. trigonocephalusand P. breviceps at night. Dietary breadth was greater for species with larger mouth dimensions.     

Seasonal succession of zooplankton in the north basin of Lake Biwa

To examine the seasonal succession of the entire zooplankton community in Lake Biwa, zooplankton biomass (on an areal basis) and its distribution patterns among crustaceans, rotifers and ciliates were studied in the north basin from April 1997 to June 1998. Seasonal changes in phytoplankton and population dynamics of Daphnia galeata were also examined to assess food condition and predation pressure by fish. From March to November, crustaceans dominated zooplankton biomass, but rotifers and ciliates were dominant from December to February. Among crustaceans, Eodiaptomus japonicus was the most abundant species, followed by D. galeata. Zooplankton biomass increased from January to a peak in early April, just before the spring bloom of phytoplankton, then decreased in mid-April when mortality rate of D. galeata increased. From mid-June, zooplankton increased and maintained a high level until the beginning of November. During this period, both birth and mortality rates of D. galeata were relatively high and a number of rotifer and crustacean species were observed. However, their abundances were very limited except for E. japonicus which likely preys on ciliates and rotifers. In Lake Biwa, food sources other than phytoplankton, such as resuspended organic matter from the sediments, seems to play a crucial role in zooplankton succession from winter to early spring, while zooplankton community seems to be regulated mainly by fish predation from summer to fall.     

Winter spatial distribution of fish larvae assemblages relative to the hydrography of the waters surrounding Taiwan

The aim of this study was to investigate the species composition and distribution of fish larvae in relation to hydrographic conditions in the waters surrounding Taiwan Island (TI) in February 2003. In total, 242 kinds of fish larvae belonging to 127 genera and 75 families were recognized. Among these, 109 taxa were identified to the family or genus level, others to the species level. The 12 predominant types, which constituted 71% of the total fish larvae, were Engraulis japonica, Scomber sp., Diaphus spp., Benthosema pterotum, Carangoides ferdau, Embolichthys mitsukurii, Maurolicus sp., unidentified Myctophidae, Gonostoma gracile, Trichiurus lepturus, unidentified Gobiidae, and Myctophum asperum. The distribution of fish larvae showed a clear association with water masses around TI, with higher abundances and lower species richness northwest of TI where the China Coastal Current prevails, and lower abundances and higher species diversity east of TI where the Kuroshio Current dominates. Cluster analysis distinguished three station groups and four species groups, and the distribution patterns of fish larvae also corresponded to hydrographic conditions. The total abundances of fish larvae and eight of the 12 predominant taxa showed significant and positive correlations with zooplankton abundance, which suggests that food source might be a key factor determining the abundance and distribution of fish larvae during the winter.     

The importance of zooplankton in the diets of three native fish species in floodplain waterholes of a dryland river,the Macintyre River,Australia

The zooplankton of freshwater systems has been recognized as an important energy resource for fish of small body size that, in turn, provide energy to piscivorous fish consumers higher up the food web. This study evaluates the importance of zooplankton to the diets of three species of fish living in floodplain waterholes of an Australian dryland river. The species selected for study represent different trophic categories in waterhole food webs: Ambassis agassizii is a microcarnivore, Leiopotherapon unicolor is an omnivore, and Nematalosa erebi is a detritivore. Dietary differences among size classes of each species were also evaluated to understand possible ontogenetic shifts in zooplankton consumption. Ambassis agassizii fed primarily on zooplankton (99.9%, made up mostly of 81.6% Calanoida and 17.4% Moinidae), regardless of the size of individual fish. Leiopotherapon unicolor fed on zooplankton (47%, mostly Daphniidae and Moinidae) and aquatic insects (46.7%). Smaller individuals of Leiopotherapon unicolor (30–49 mm TL—total length) were responsible for 36.1% of the plankton consumed by the species. Nematalosa erebi fed on detritus (84.6%) with zooplankton (Calanoida, Moinidae, and Cyclopoida) contributing only 13.7% of the mean diet. Smaller individuals (40–69 mm TL) were responsible for 98% of the plankton consumed by Nematalosa erebi, and individuals of 40–49 mm (TL) fed exclusively on zooplankton (53.8% Moinidae and 46.2% Calanoida). Although the three fish species had different diets, reflecting differences in species-specific and ontogenetic morphological and behavioral characteristics, zooplankton formed the basis of the diet of all species when young. These results confirm the importance of zooplankton as a major food resource for three fish species and smaller size classes of these species in floodplain waterholes of the Macintyre River, Australia. Guest editors: U. M. Azeiteiro, I. Jenkinson & M. J. Pereira Plankton Studies     

Immunoreactive atrial natriuretic peptide in the fish heart and blood plasma examined by electron microscopy,immunohistochemistry and radioimmunoassay

Summary The immunoreactivity of atrial natriuretic peptide and ultrastructure of cardiocytes were examined in 5 species each of freshwater and seawater teleosts, as well as in 2 species each of elasmobranchs and cyclostomes. Immunoreactivity was strong in the atria of Cyprinus carpio, Anguilla japonica and Conger myriaster, rather weak in atria of Channa maculata, Lepomis macrochirus, Salmo gairdneri, Oplegnathus fasciatus and Eptatretus burgeri, very weak in atria of Pagrus major, Trachurus japonicus and Triakis scyllia, and not detectable in atria of Hexagrammos otakii, Narke japonica and Lampetra japonica. The immunoreactivity of the atrial cardiocytes was generally stronger in freshwater than seawater fish. Ventricular immunoreactivity was detected only in 7 species, always being weaker than that observed in the atrium. Ultrastructurally, however, secretory granules were found in atria and ventricles of all species examined, being more frequent in the former than the latter. By radioimmunoassay, immunoreactive ANP was detected in the extracts of blood plasma and both atrial and ventricular tissues of all species examined. There were no statistically significant differences in the values between freshwater and seawater species.     

Feeding behaviour of major 0 + fish species in a shallow, eutrophic lake (Tjeukemeer, The Netherlands)

The forage base and the food selectivity of 0+ representatives of six abundant freshwater fish species were studied in a shallow, eutrophic Dutch lake. Most species relied on the zooplankton; the size-selective predation in early summer was directed to the smaller copepods and in late summer to larger cladocerans and copepods than concurrently present in the lake. Daphnia spp. and cyclopoid copepods were the main zooplankton taxa for smelt, perch and pikeperch. Energetically, the large cladoceran, Leptodora kindtii, was especially important for pikeperch. Bream and roach preyed upon smaller zooplankton than the other fish species. The influence of the zooplankton predation by abundant 0+ fish was clear from a small mean Daphnia size in September; this size is to be used as an indicator in fishery management. Neomysis integer, the most important macrofauna species, was consumed by perch, pikeperch and ruffe; pikeperch was most size-selective in this respect. The 0+ ruffe was à typically benthivorous fish. Only the 0+ pikeperch became piscivorous, especially in years when smelt was abundant.     

Food consumption and growth of larvae and juveniles of three cyprinid species at different food levels

Synopsis The relationships between food availability, consumption and growth were analyzed from the onset of feeding to an age of 90 days in three cyprinid species. Fish were held at 20 ± 0.5° C and given two (three) constant rations of approximately 30, (40) or 100% dry body weight (dbw) ind^-1 day^-1. Food consisted of living zooplankton, the size of which correlated with fish size. At high food densities consumption rates decreased rapidly with fish size in all three species. At reduced rations, fish consumed most of the food offered until they were larger than 10 mg dbw. In all species and at each feeding level daily rations consumed increased allometrically with body size. Respiration rate, expressed as routine metabolic rate differed insignificantly between the three species. At high ration levels, growth rates of small bleak, Alburnus alburnus, were distinctly lower than those of roach, Rutilus rutilus, and blue bream, Abramis ballerus. At low food supply all three species grew at similar rates. Assimilation efficiency at low food conditions was approximately twice that of the well-fed groups. If the caloric equivalents of oxygen consumption as measured in well-fed fish are applied to fish fed at low rations their energy budgets do not balance. This indicates the limitations of fish larvae in the partitioning of energy for growth or activity at such conditions.     

Seasonal occurrence and food habits of larvae and juveniles of two temperate basses in the Shimanto estuary,Japan

Larvae and juveniles ofLateolabrax japonicus andL. latus occurred from January to May 1986 in the shallow waters of the Shimanto estuary.L. japonicus markedly outnumberedL. latus. Distinct ecological differences were recognized in habitats and food habits between the two species:L. japonicus mainly inhabited eelgrass beds composed ofZostera nana, whileL. latus appeared evenly in both eelgrass beds and non-eelgrass habitats; the former fed on copepods and cladocerans, while the latter fed on copepods and fish larvae. From these habitat and food habit analyses, estuaries were considered to be important as a main habitat forL. japonicus, but not forL. latus. The fact that ecological differences have occurred during the early life stages was inferred to be one of the possible keys to speculate on the speciation of the two species.     

Imprint of monsoonal patterns on the fish assemblage in coastal waters of south‐east India: a case study

The composition, species richness and diversity of a coastal fish assemblage from the Kalpakkam coast of south‐east India are described along with temporal distribution patterns related to seasonal fluctuations in dissolved oxygen, salinity, pH, chlorophyll‐a, phytoplankton and zooplankton species richness and density. A total of 244 fish species belonging to 21 orders, 87 families and 163 genera were recorded. The fish assemblage was dominated by reef‐associated species, followed by demersal species. The majority of the species (63%) are widely distributed in the western Indo‐Pacific as well as in the central Indo‐Pacific. Jaccard's coefficient analysis showed three distinct seasonal patterns of fish occurrence: pre‐monsoon (PrM), monsoon (M) and post‐monsoon (PoM). The maximum number of species was during the PrM period, followed by the PoM and M periods. Species occurrence analysis showed Sardinella longiceps to be dominant during PrM and M periods, Leiognathus dussumieri during the M period and Secutor insidiator and Secutor ruconius during the M and PoM periods. Canonical correspondence analysis indicated that salinity and rainfall were the two most influential environmental factors strongly correlated with temporal variation in the fish assemblage. The physico‐chemical conditions, in combination with factors such as greater food availability and shelter, might control the seasonal local distribution of the ichthyofauna in these Indian coastal waters.     

Long-term changes in littoral fish community structure and resilience of total catch to re-oligotrophication in a large,peri-alpine European lake

1. The littoral zone of lakes is used as spawning, shelter, or feeding habitat for many fish species and hence is of key importance for overall lake functioning. Despite this, hardly any studies exist examining the long-term dynamics and response of the littoral fish community, composed mostly of juvenile fish, to environmental change. Here, we study the response of total catch per unit effort (CPUE) and individual species CPUE of such a community to 17 years of oligotrophication and examine whether the species responses can be characterised as synchronous or asynchronous.
2. We analyse a data set of beach seine catches carried out during morning and twilight, late spring and late summer at three sites in large and deep Lake Constance from 1997 to 2014. Generalised additive mixed models were used to explore changes in CPUE of the overall community and of the most frequently occurring species, and Kendall's W test was applied to examine whether the dynamics of fish species were synchronous or asynchronous.
3. Species-specific and total CPUE strongly differed between morning and twilight and between spring and summer indicating an important role of behavioural and life cycle adaptations of species for CPUE. In addition, also the CPUE of some species seeking shelter behind larger stones was lower at sites without these.
4. Total CPUE did not decline suggesting the overall abundance of littoral fish was resilient to declining nutrients. In contrast, fish community composition changed strongly during the study period due to increases in some species (dace, loach, perch) and decreases in others (bream, burbot, chub, ruffe), indicating response diversity of fish to oligotrophication. The type of community dynamics was scale-dependent, whereby significantly synchronous dynamics according to Kendall's W were observed when taking seasonal variability into account. In contrast, significantly asynchronous species dynamics were observed when only the low-frequency variability of species dynamics was considered separately for spring and summer time series.
5. Resilience of littoral fish total CPUE to oligotrophication might have important consequences for ecosystem dynamics and ecosystem services beyond the littoral zone. As small fish often impose strong predation pressure on zooplankton, their resilience might sustain a high top-down control on zooplankton resulting in a further reduction of zooplankton biomass. This could contribute to delayed food web responses and reduced growth of fish with oligotrophication.

     

Genetic identification of fish eggs collected in Sendai Bay and off Johban,Japan

Mitochondrial DNA typing of 52 fish eggs collected in Sendai Bay and off Johban revealed ten haplotypes of the 225–251 bp region of 16S ribosomal DNA. Comparing with known sequences of fishes, we identified them as Trachurus japonicus (Carangidae), Pagrus major, Evynnis japonica (Sparidae), Paralichthys olivaceus, Pseudorhombus sp. (Paralichthidae), Pennahia argentata (Sciaenidae) and Platycephalus sp. (Platycephalidae). The occurrence of Trachurus japonicus and sparid eggs in the study area indicates significant extensions to the ranges of their spawning sites northward. Our method is useful for identifying fish eggs that are difficult to diagnose by morphology, and to uncover cryptic fish egg assemblages in the field.     

Diversity and composition of estuarine and lagoonal fish assemblages of Socotra Island,Yemen

Estuarine and lagoonal surveys of Socotra Island and selected sites on the Hadhramout coast of Yemen were conducted with the objective of documenting and analysing fish diversity and assemblage structure. A total of 74 species in 35 families were recorded, among which 65 species in 32 families were from Socotra and 20 species in 17 families were from mainland Yemen. Twenty‐one species represent new faunal records for Socotra. Including historic records re‐examined in this study, the total fish species richness of estuaries and lagoons of Socotra Island reaches 76, which is relatively high compared to species inventories of well‐researched coastal estuaries in southern Africa. Five species dominate the occurrence and abundance frequencies: Terapon jarbua, Hyporhamphus sindensis, Aphanius dispar, Ambassis gymnocephala and Chelon macrolepis. Rarefaction and extrapolation analyses suggest that the actual number of fish species inhabiting some of those estuaries might be higher than the one observed. Thus, additional sampling at specific sites should be conducted to record other less conspicuous species. Ordination and multivariate analyses identified four main distinct assemblage clusters. Two groups are geographically well structured and represent northern Socotra and mainland Yemen, respectively. The other two assemblage groups tend to be determined to a greater extent by the synchrony between physical (e.g. estuary opening periods) and biological (e.g. spawning and recruitment periods) variables than by geographical location. Finally, the single intertidal lagoon of Socotra represents by itself a specific fish assemblage. The high proportion of economically important fish species (38) recorded underscores the paramount importance of these coastal water bodies as nursery sites, and for sustaining vital provisioning ecosystem services.     

Temporal dynamics and relationship between climate,limnological variables and zooplankton composition in climate-sensitive Lake Bosumtwi,Ghana

The community composition and the factors affecting seasonal and interannual dynamics of zooplankton in Lake Bosumtwi were studied biweekly at a central index station during 2005 and 2006. The lake zooplankton community was species poor. Mesocyclops bosumtwii was numerically superior seasonally and interannually and was endemic to the lake. Minor constituents included Moina micrura, six rotifer species (except for Hexarthra intermedia) and Chaoborus ceratopogones larvae. Low variance of cyanobacteria-dominated phytoplankton biomass underlined stable zooplankton community structure. Emergence of rare species of rotifers occurred seasonally. The climatic signature on the lake’s stratification and mixing regime was strongly influenced by atmospheric temperature, but weakly by wind strength, because of sheltering of the lake by high crater walls. Increasing mixing depth entrained high TP concentrations from below the thermocline seasonally, but reflected poorly in the phytoplankton biomass behaviour. Total zooplankton abundance did not differ seasonally, but varied markedly from year to year in its timing and magnitude. Herbivores were squeezed between food limitation and high predation pressure from Chaoborus all year round. The low fish planktivory (high fishing pressure) on Chaoborus may create a trophic bottleneck restricting energy transfer efficiency from zooplankton to fish.     

Seasonal variations in the diversity,abundance, and composition of species in an estuarine fish community in the Tropical Eastern Pacific,Mexico

The ichthyofauna of the Chacahua Lagoon in the western Oaxaca State of Mexico was sampled every 2 months, using a trawl net over seven sampling cycles. The estuary mouth closed in late January, generating hypersaline conditions in the system. A total of 33 species belonging to 20 families were recorded; most of them marine species, showing tropical and subtropical affinities. The most abundant species were Diapterus peruvianus, Centropomus robalito, Anchovia macrolepidota, Lile stolifera, and Lutjanus novemfasciatus. Total fish abundance and species richness were not significantly affected by the mouth closing, and this was related to the presence of a permanently open entrance channel with an adjacent lagoon, which allowed easy exchange of fish between these two systems. Canonical correspondence analysis (CCA) revealed that salinity and dissolved oxygen were the most important environmental variables in determining the observed variability in fish community composition. Two fish groups were evident: the fish assemblage of the low-salinity period (open-mouth phase), in which Eucinostomus currani, Sciades guatemalensis, Centropomus armatus, Citharichthys stigmaeus, and Caranx caninus constituted the majority of the catch; and the fish assemblage of the high-salinity period (closed-mouth phase), with A. macrolepidota, L. stolifera, and Harengula thrissina as the most abundant species. Multivariate analyses showed differences in the composition of the fish community between both periods. Other species, such as the permanent residents D. peruvianus and C. robalito, which showed the widest range in tolerance of ambient salinity, were caught during both periods. Seasonal succession of fish populations may be related to differences in life cycle and tolerance of the environment among those species adapting to ecological conditions.     

The ecology of ruffe,Gymnocephalus cernuus (Pisces: Percidae) introduced to Mildevatn,western Norway

Synopsis Diet, habitat use, diel and seasonal activity and a number of population parameters were studied on ruffe,Gymnocephalus cernuus, introduced to Mildevatn, western Norway. This lake is sited outside the natural range of the ruffe and has a lower fish diversity and a different fish species composition than within its native range. From June through September the ruffe was planktivorous and mainly caught at 4 to 6 m depth in the benthic zone. At other times of year ruffe was feeding on zoobenthos and caught deeper in the benthic zone. Ruffe was mainly day active. Zooplankton feeding during summer is the clearest difference compared to ruffe populations living within its natural range. Presence of large zooplankton organisms available for ruffe is suggested as the main reason for the difference found in food choice. The availability of large zooplankton is probably due to community structure caused by a predator and lack of interspecific competition for zooplankton in the deeper parts of the lake. Piscivorous brown trout.Salmo trutta, restrict the habitat of threespined stickleback,Gasterosteus aculeatus, to the zone of littoral vegetation, allowing high densities of larger zooplankton species likeBythotrephes longimanus to be present in the lake. Brown trout is present only in the upper light and well oxygenated parts of the lake, leaving a refuge for the ruffe, where they can feed on the rich zooplankton community.     

STOMACH CONTENTS OF FALSE KILLER WHALES (PSEUDORCA CRASSIDENS) STRANDED ON THE COASTS OF THE STRAIT OF MAGELLAN,TIERRA DEL FUEGO

We examined the stomachs of 25 false killer whales collected from a mass stranding of 181 animals along both coasts of the Strait of Magellan, Chile, in March 1989. The 21 stomachs (nine males and 12 females), with food remains contained 11 prey species (nine cephalopods and two fishes) with a total of 442 individuals. Except for one case, food remains were meager, indicating that the animals had not eaten for some time or through stress had vomited on the beach. Eleven of the 21 animals had mud (often with squid beaks) in the esophagus and first stomach. The prey were identified employing squid beaks, fish otoliths and bones, and their wet weights were estimated using regressions between hard parts and known weights of species. The most important prey were the oceanic and neritic-oceanic squids, Martialia byadesi and Illex argentinus, followed by the neritic fish, Macruronus magellanicus. Of less importance were the oceanic squid, Todaroes fillipovae, the oceanic and epipelagic octopus, Ocytboe sp., and the oceanic squid, Moroteuthis ingens. The rest of the prey were poorly represented and included four oceanic squids and one neritic fish. The prey species of these animals were subantarctic, with two antarctic species, abundant over the Patagonian shelf and adjacent oceanic waters around Tierra del Fuego.     

Inter-annual variation in summer zooplankton community structure in Prydz Bay, Antarctica, from 1999 to 2006

Inter-annual variations in zooplankton community structure in Prydz Bay were investigated using multivariate analysis based on samples collected with a 330-μm mesh, 0.5-m² Norpac net during the austral summer from 1999 to 2006. Two distinct communities, an oceanic and a neritic community, were consistently identified in all surveys. Oceanic communities had higher diversity and were indicated by species such as Haloptilus ocellatus, Heterorhabdus austrinus, Thysanoessa macrura, Rhincalanus gigas, Scolecithricella minor and Oikopleura sp.. Neritic communities were indicated by Euphausia crystallorophias and Stephos longipes and were characterized by fewer but more abundant species. In 1999 and 2006, a transitional community was also distinguished near the continental shelf edge, where ice coverage was more extensive than either the oceanic or neritic regions. Significant inter-annual variations in community structure (mainly involving species abundance rather than species composition) were found in both oceanic and neritic communities, being more obvious in the latter. The timing and amplitude of sea ice retreat (polynya appearance), and its effect on food availability, had strong influences on zooplankton community structure. In oceanic communities during years with earlier ice retreat, the extra time available for phytoplankton blooms to accumulate resulted in a higher proportion of large copepods (Calanoides acutus, Calanus propinquus, Metridia gerlachei) (especially the younger copepodites) in the zooplankton assemblage. In neritic communities, zooplankton such as the ice krill E. crystallorophias, and large copepods (C. acutus, C. propinquus, M. gerlachei), also showed higher abundance and earlier developmental stages in years with larger polynya. On the other hand, in years with later ice retreat, smaller polynya, and less time for phytoplankton blooms to form, the abundance of large copepods was lower and older age classes were more common.