Reproduction,hatching success,and early naupliar survival in Centropages typicus

The broadcast spawner, Centropages typicus, is a very successful copepod species in many coastal areas of the Mediterranean Sea and North Atlantic Ocean. This review assembles the large amount of information on the reproduction and early life history of C. typicus that has emerged since the 1970s and has made this species one of the best-studied copepods, similar in that regard to species of Acartia and Calanus. Observations on mating behavior and the female gametogenic and oogenic cycles are presented, together with information on seasonal cycles of egg production rates in Mediterranean and Atlantic populations from various regions. These studies indicate a strong latitudinal gradient, with continuous reproduction and the main spawning season occurring earlier (late winter/spring) in warmer waters such as the Mediterranean Sea, compared to northern areas such as the North Sea and in the Kattegat, where C. typicus actively reproduces mainly in late summer and fall with reproduction ceasing altogether in winter in the German Bight. These observations strongly suggest that temperature is the controlling factor for reproductive activity in this species. Egg development times are also temperature dependent but do not vary with latitude, and there is as yet no conclusive evidence that diapause egg production occurs in C. typicus. Laboratory experiments have shown that food quantity and quality both affect fecundity and offspring fitness, but most of these studies have focused on diatom and dinoflagellate diets and non-algal prey have been strongly underrepresented, despite their importance for this omnivorous copepod. Large fluctuations in hatching success and naupliar survival have been reported in field surveys and have subsequently been related to maternal feeding history and food quality or toxicity in laboratory experiments. We identify future lines of research that will help to explain the interannual variability in breeding intensity and recruitment of C. typicus in the field based on better understanding of the species’ reproductive biology.     

Temporal variability of Centropages typicus in the Mediterranean Sea over seasonal-to-decadal scales

Centropages typicus is one of the most common, abundant and best studied calanoid copepods in neritic waters of the Mediterranean Sea, which means it can provide useful information about the long-term dynamics of the Mediterranean epipelagic ecosystem. This paper presents the first comparative overview of the seasonal and long-term variability of C. typicus in different Mediterranean regions. This review is based on quantitative information from the published literature and novel data from five ongoing zooplankton time-series carried out in the Mallorca Island (Balearic Sea), the Bay of Villefranche (Ligurian Sea), the Gulf of Naples (Tyrrhenian Sea), the Gulf of Trieste (North Adriatic Sea), and the Saronikos Gulf (Aegean Sea). In most Mediterranean regions, C. typicus has a perennial occurrence, with peaks of abundance that reflect the succession of different generations. Throughout the Mediterranean, the annual cycle of C. typicus is characterized by minima in winter and major peaks in April–June, which is earlier than those observed in European Atlantic waters, where the peaks are more frequently recorded in summer and fall. In the regions investigated, the annual cycle shows remarkable similarities in terms of timing, but notable differences in the peak height; populations are far more abundant in coastal north-western regions and less abundant in the eastern basin. In the long-term, changes in C. typicus phenology observed in the Bay of Villefranche and in the Gulf of Naples are related to the North Atlantic Oscillation (NAO) index. In these two regions, the species responds to climate forcing similarly in terms of average seasonal patterns (bi-modal patterns in years of positive NAO, unimodal patterns in years of negative NAO) but oppositely in terms of quantity, indicating different influence of the NAO on the two regions. At decadal scales, C. typicus populations show high interannual variability with marked geographical differences. In some areas, the patterns are clearly characterized by alternate phases of higher and lower annual abundances, at higher frequency (mainly 1–2 years) in the Gulf of Naples, and lower frequency in the Saronikos Gulf (mainly 4–5 years) and in the Gulf of Trieste (mainly 5–6 years). Synchronous phases of increasing or decreasing abundance are discernable only for a few sites and short periods, for example from 1998 to 2000 in the Gulf of Naples, Gulf of Trieste and Saronikos Gulf. The regional differences observed in the long-term patterns of C. typicus populations suggest that the temporal dynamics of this species are significantly more affected by local conditions than by any possible common driving force acting at basin scale through teleconnections.     

Macroecological study of Centropages typicus in the North Atlantic Ocean

Centropages typicus is a temperate neritic-coastal species of the North Atlantic Oceans, generally found between the latitudes of the Mediterranean and the Norwegian Sea. Therefore, the species experiences a large number of environments and adjusts its life cycle in response to changes in key abiotic parameters such as temperature. Using data from the Continuous Plankton Recorder (CPR) survey, we review the macroecology of C. typicus and factors that influence its spatial distribution, phenology and year-to-year to decadal variability. The ecological preferences are identified and quantified. Mechanisms that allow the species to occur in such different environments are discussed and hypotheses are proposed as to how the species adapts to its environment. We show that temperature and both quantity and quality of phytoplankton are important factors explaining the space and time variability of C. typicus. These results show that C. typicus will not respond only to temperature increase in the region but also to changes in phytoplankton abundance, structure and composition and timing of occurrence. Methods such as a decision tree can help to forecast expected changes in the distribution of this species with hydro-climatic forcing.     

Metabolism of Centropages species in the Mediterranean Sea and the North Atlantic Ocean

Information on the metabolism rates of Centropages typicus and congeneric species (C. hamatus, C. furcatus, C. brachiatus and C. abdominalis) in neritic areas of the Mediterranean Sea, the North Atlantic Ocean and the Pacific Ocean are reported here. Respiration rates and excretion rates are strongly influenced by abiotic (i.e. temperature, salinity) and biotic factors (i.e. food availability and composition). Differences in the response of respiratory rates to temperature of acclimated, acclimatized and adapted individuals are clearly observed among regions of the Mediterranean Sea and the West and East shores of the Atlantic Ocean. Food supply also strongly affects respiration and excretion rates, as well as the size, sex and stage development of the individuals. The co-measurement of these two rates allows confirmation of the omnivory or carnivory oriented feeding habits of these species. The role of this neritic genus in coastal environment is also discussed.     

Development and growth rates of Centropages typicus

This paper presents an updated review of our knowledge about Centropages typicus development and growth based on a synthesis of all data found in the literature on stage durations, development rates, size, weight, and C:N ratios. Developmental and growth characteristics have been recorded during field time series and cruises, as well as in controlled laboratory experiments. A few laboratory studies have contributed to an understanding of the impact of food quantity, food quality and temperature on growth and development.C. typicus has a sigmoidal development pattern when the contribution of the different stages to total development time is considered, CIV and CV having much longer durations than preceding stages. The development times of this species are closely related to temperature with a nearly equiproportional development. In turn, food conditions, both in quantity and quality, have less effect on development than on growth rate, leading to large variations in weight within stage, even larger than those predicted from changes in length and biovolume. Actual measurements of weight and biochemical composition are required to better understand the growth of C. typicus in relation to food composition. As a consequence, our review shows that weight–length relationships obtained in particular environmental conditions may be inadequate for others.The combination of a temperature-dependent development rate and a food-dependent growth rate may explain why shelf regions are a more favourable environment for C. typicus as compared to offshore regions, as they support large females, production of more and better quality eggs, and probably better survival of the offspring.     

Seasonal and spatial dynamics of Centropages typicus and C. hamatus in the western North Atlantic

Centropages typicus and Centropages hamatus are two of the most abundant copepods on the continental shelf in the mid-Atlantic region of the western North Atlantic. Their range extends from the Scotian Shelf (C. typicus) and the Grand Banks (C. hamatus) in the north to Cape Hatteras in the south. South of Cape Hatteras they have only been observed in inshore waters of North and South Carolina and not offshore on the continental shelf or in coastal waters of eastern Florida. However, C. hamatus has been observed in western Florida. Abundances of both species are greatest in inshore regions in the mid-Atlantic Bight with C. hamatus tending to have a more coastal distribution. In this region seasonal variability is low with high abundances from late fall through mid-summer. In the north year-round presence of both species is confined to inshore areas and offshore banks such as Georges Bank, and Browns Bank, Emerald Bank and Western Bank on the Scotian Shelf. In this northern region there is a pronounced seasonal cycle in abundance with high abundances during late summer and fall. Periods of high reproductive rates are closely linked to blooms of large phytoplankton and food availability rather than temperature appears to be controlling population abundances.     

The feeding ecology of the copepod Centropages typicus (Kröyer)

Here we report on the current knowledge on the feeding ecology of the planktonic copepod Centropages typicus. We describe the foraging process of C. typicus from the detection of prey to their digestion, considering also the effects of several physical variables on the feeding activity of the species. C. typicus is an omnivorous copepod that feeds on a wide spectrum of prey, from small algae (3–4 μm equivalent spherical diameter, ESD) to yolk-sac fish larvae (3.2–3.6 mm length). It uses both suspensivorous and ambush feeding strategies, depending on the characteristics of the prey. In general, C. typicus exhibits selection for large motile prey, such as ciliates or dinoflagellates, both in nature and laboratory, and this selective pattern is enhanced under moderate intensities of turbulence. Daily rations in the field are somewhat lower than those found in the laboratory, which indicates food limitation. This fact, together with the relatively modest capacity of the species to adapt to fluctuations in food availability may explain the geographical distribution of C. typicus, being restricted to near-shelf waters. In general, it does not seem that C. typicus feeding severely impacts planktonic populations. However, the occasional importance of the species in certain ecosystems is also apparent.     

Comparative seasonal dynamics of Centropages typicus at seven coastal monitoring stations in the North Sea,English Channel and Bay of Biscay

We review current knowledge and understanding of the biology and ecology of Centropages typicus in the European shelf-seas (e.g. North Sea, English Channel and Bay of Biscay). Our study is based on observations at seven coastal time-series stations as well as on the Continuous Plankton Recorder dataset. This paper focuses on the influence of the environmental parameters (e.g. temperature and Chla) on the life cycle and distribution of C. typicus and provides a comparison with its congeneric species C. hamatus and C. chierchiae in the study area. Data on abundance, seasonality and egg production have been used to define the temperature and chlorophyll optima for occurrence and reproduction of Centropages spp. within this region of the European shelf-seas.     

Food preferences and rhythms of feeding activity of two co‐existing demersal fish,the longspine snipefish,Macroramphosus scolopax (Linnaeus, 1758), and the boarfish Capros aper (Linnaeus, 1758), on the Mediterranean deep shelf

The feeding behaviour of two potentially competing species, the longspine snipefish, Macroramphosus scolopax, and the boarfish Capros aper was examined. While both species are very abundant along the Mediterranean coast and are regularly caught by demersal trawlers, they are of no commercial value. The diets of boarfish and longspine snipefish were investigated from samples collected between January 2001 and May 2002. Variations in the diet with fish size and season, as well as diet overlap and diversity were explored. Mysid shrimps, amphipods and gastropods were the most important food items in the diet of longspine snipefish. During ontogenetic development, M. scolopax occupies different trophic levels: the diet shifts from being predominantly composed of mysids (Anchialina agilis, Lophogaster typicus, Erythrops sp., Leptomysis spp.) in the smaller longspine snipefish [<6.5 cm total length (TL)] towards decapods (Anapagurus laevis) and amphipods (Leucothoe incisa, Eusirus longipes, Hyperidea) in the larger individuals (>6.5 cm TL). Crustacean decapods and copepods were the most important prey in the stomachs of boarfish. Mysids (Lo. typicus), euphausiids and nematodes were present in the larger individuals (>8 cm TL). A more generalist diet, still containing copepods, crustacean decapods, gastropods (Limacina retroversa) and a large variety of amphipods (e.g. Phtysica marina, Stenotoe bosphorana) and mysids (e.g. A. agilis, Leptomysis spp., Erythrops sp.), dominated the diet of C. aper between 2 and 8 cm TL. Diet overlap between longspine snipefish and boarfish was very low and the differences in stomach species diversity were explained by season and fish size.     

Centropages behaviour: Swimming and vertical migration

The evolutionary success of any species living in a variable environment depends on its capacity to enhance the probability of finding food and mates, and escaping predators. In the case of copepods of the genus Centropages, as in all planktonic copepods, their swimming behaviour is closely tied to these vital aspects, and shows a high degree of plasticity and adaptive capacity. Swimming mechanisms of Centropages change radically during development, mainly in the transition between naupliar stages to the 1st copepodite; nauplii do not produce feeding currents, whereas copepodites do. Adults and late developmental stages of C. typicus, C. hamatus and C. velificatus spend most of the time in slow swimming and resting breaks, with occasional and brief fast swimming (escape reactions) and grooming events. Slow swimming is closely related to the creation of feeding currents, and results from the beating of the cephalic appendages in a “fling and clap” manner. The proportion of time allocated to the different swimming activities depends on sensory cues like type and concentration of food, presence of potential mates, light intensity, hydrodynamic flow, etc.The responses of Centropages to changes in flow velocity fluctuations (small-scale turbulence) are similar to the escape responses (fast swimming) triggered by the presence of potential predators. Centropages generally have standard nocturnal vertical migration patterns involving considerable vertical displacements. This behaviour is closely related to the narrow spectral sensitivity and the low intensity threshold of the genus, and has important consequences for the active vertical transport of matter and energy.The variety of responses of Centropages to environmental changes, and in general all the aspects related to its swimming behaviour seem to be controlled by the trade-off between energetic gains (food intake), losses (swimming energy expenditure), and predation risk. Behavioural plasticity and adaptation appear to be the most relevant characteristics for the success of the genus in a wide range of marine environments.     

Invasive plants on disturbed Korean sand dunes

The sand dunes in coastal regions of South Korea are important ecosystems because of their small size, the rare species found in this habitat, and the beautiful landscapes they create. This study investigated the current vegetative status of sand dunes on three representative coasts of the Korean peninsula, and on the coasts of Cheju Island, and assessed the conditions caused by invasive plants. The relationships between the degree of invasion and 14 environmental variables were studied. Plots of sand dunes along line transects perpendicular to the coastal lines were established to estimate vegetative species coverage. TWINSPAN (Two-Way Indicator Species Analysis), CCA (Canonical Correspondence Analysis), and DCCA (Detrended Canonical Correspondence Analysis) were performed to classify communities on sand dunes and assess species composition variation. Carex kobomugi, Elymus mollis, and Vitex rotundifolia were found to be the dominant species plotted on the east, the west, and the peripheral coasts of Cheju Island, respectively. Vegetation on the south coast was totally extinct. The 19 communities, including representative C. kobomugi, C. kobomugi–Ixeris repens, C. kobomugi–Oenothera biennis, E. mollis, Lolium multiflorum–Calystegia soldanella, and V. rotundifolia–C. kobomugi, were all classified according to TWINSPAN. Oenothera biennis and L. multiflorum were exotics observed within these native communities. CCA showed that invasive native and exotic species distribution was segregated significantly, according to disturbance level, exotic species number, gravel, sand and silt contents, as well as vegetation size. It further revealed that human disturbance can strongly favor the settlement of invasive and exotic species. Restoration options to reduce exotic plants in the South Korean sand dune areas were found to be the introduction of native plant species from one sand dune into other sand dune areas, prohibition of building and the introduction of exotic soils, and conservation of surrounding sand dune areas.     

Cetacean research in the southern African subregion: a review of previous studies and current knowledge

Cetacean research, in terms of the number of papers, and areas for which data are available, has expanded considerably in the southern African subregion in the past decade, especially in the South-West Indian Ocean. We review cetacean research within this subregion from the 1800s to the present to provide an overview of findings, investigate trends and identify knowledge gaps. Data are presented separately for large whales (those subject to commercial whaling) and smaller cetaceans, and are separated by era and ocean basin. Over 550 peer-reviewed papers and books were identified relating to research on cetaceans within the subregion. More than half (284) have been produced since 1990 and 193 relate specifically to South African waters. The most-studied species are those that are most accessible due to their coastal distributions (southern right whale Eubalaena australis: 45 papers, humpback whale Megaptera novaeangliae: 31 papers, killer whales Orcinus orca: 27 papers, Indo-Pacific bottlenose dolphin Tursiops aduncus: 30 papers, Indo-Pacific humpback dolphin Sousa chinensis (plumbea form): 25 papers) and/or were hunted commercially (sperm whale Physeter macrocephalus: 25 papers). Identified conservation concerns vary throughout the subregion, but include bycatch and directed hunts, oil and gas development, ecotourism activities, shifts in prey resources, and noise and chemical pollution. The inshore stocks of Bryde's whales Balaenoptera edeni, the Indo-Pacific humpback dolphin and the Atlantic humpback dolphin S. teuszii were identified as the populations of highest conservation concern, although there are considerable knowledge gaps relating to deep-water species and almost no data (even on species occurrence) are available for several areas and countries.     

The co-location of offshore windfarms and decapod fisheries in the UK: Constraints and opportunities

The offshore wind sector in the UK is expanding rapidly and is set to occupy significant areas of the coastal zone, making it necessary to explore the potential for co-location with other economic activities. The presence of turbine foundations introduces hard substrates into areas previously dominated by soft sediments, implying that artificial reef effects may occur, with potential benefits for fisheries. This review focuses on the possibilities for locating fisheries for two commercially important decapods, the brown crab Cancer pagurus and the European lobster Homarus gammarus, within offshore wind farms.Existing understanding of habitat use by C. pagurus and H. gammarus suggests that turbine foundations have the potential to act as artificial reefs, although the responses of these species to noise and electromagnetic fields are poorly understood. Offshore wind farm monitoring programmes provide very limited information, but do suggest that adult C. pagurus associate with turbine foundations, which may also serve as nursery areas. There was insufficient deployment and monitoring of rock armouring to draw conclusions about the association of H. gammarus with offshore wind farm foundations. The limited information currently available demonstrates the need for further research into the ecological and socio-economic issues surrounding fishery co-location potential.     

Inter-annual variability of the epibiotic community on Pagurus bernhardus from Scotland

Epibiont communities of Pagurus bernhardus and its inhabiting shell collected in the same month of two consecutive years near the Isle of Cumbrae (Scotland) were analyzed. The epibionts found were: (1) the protozoans Acineta, Conchacineta, Corynophrya, Zoothamnium, Cothurnia mobiusi, Cothurnia longipes, Chilodochona, Cryptacineta, Ephelota gemmipara and Ephelota plana; (2) the hydrozoans Leuckartiara, Clytia and Phialella; (3) the entoproct Barentsia; (4) the cirripeds Balanus balanus and Balanus crenatus; (5) the polychaetes Pomatoceros triqueter, Circeis amoricana paguri and Hydroides norvegica. Among these epibionts, a protozoan (E. plana), two hydrozoans (Leuckartiara, and Phialella), and the entoproct (Barentsia) have not been described previously as epibionts on crustaceans. The protozoan, entoproct and hydrozoan epibiont species found on P. bernhardus represent the first mention of their presence on this hermit crab. The protozoan epibionts were found only on the crab, while the hydrozoan epibionts were observed on the shell and on the crab. The cirriped species were observed only on the external surface of the shell, and the polychaete species were observed on the external and internal surfaces of the shell, although Circeis was found only internally. An analysis of the distribution and density of each epibiont species on the anatomical units of the crab, as well as on the different shell areas were made. The comparison between the data of the two years showed differences with respect to the size of crabs, diversity of epibiont groups and density of epibionts. There was a significant difference between the two years with respect to the distribution of epibionts on the anatomical units of the crab, both in terms of density and biomass. The influence of some environmental parameters on these differences is considered.     

Different effects of increased water temperature on egg production of Calanus finmarchicus and C. glacialis

Two copepod species, Calanus finmarchicus (a widespread North Atlantic species) and C. glacialis (an Arctic species), are dominant in the zooplankton of Arctic seas. We hypothesized that the anticipated warming in the Arctic might have different effects on the arctic and boreal species. The effect of temperature on egg production rate (EPR) in these species at temperatures of 0, 2.5, 5, 7.5, and 10°C under contrasting feeding conditions was assessed in 5-day-long experiments. The EPR of the fed C. finmarchicus increased with temperature over the entire tested range. On the contrary, the EPR of C. glacialis increased only in the range of 0–5°C and dropped with further temperature growth. The difference in the influence of temperature on reproduction of these two species is statistically significant. Feeding conditions have a considerable effect on the C. finmarchicus EPR. The EPRs of the female C. glacialis that fed or starved for 5 days displayed no significant difference. These results suggest that the C. finmarchicus EPR increases with temperature under favorable feeding conditions, whereas the C. glacialis EPR decreases at a temperature over 5°C independently of the feeding conditions. This allows for prediction of the shift in abundances of these two species in pelagic communities of Arctic seas in the case of a warming scenario.     

Productivity and nutrient cycling in salt marshes: Contribution to ecosystem health

This study aimed to assess the contribution of different salt marsh halophytes (Spartina maritima, Scirpus maritimus, Halimione portulacoides, Sarcocornia fruticosa, and Sarcocornia perennis) to nutrient cycling and sequestration in warm-temperate salt marshes. Carbon, nitrogen and phosphorus concentration in plant organs and rhizosediment, as well as plant biomass were monitored every two months during one year. Results show that the C retained in the rhizosediment does not seem to be species or site specific. However, some halophytes seem to have a higher contribution to retain C from external sources, namely S. perennis and S. maritima. Regarding N, halophytes colonizing the upper and middle marsh areas had the highest NBPP (net belowground primary production) as well as the retention of N in the rhizosediment. Yet, excluding S. maritimus, all halophytes seem to contribute to the retention of N from external sources. The P retained in the rhizosediment does not seem to be species or site specific. Still, only S. maritima colonizing the lower marsh areas, which also had comparatively lower NBPP, seem to have a higher contribution to retain P from external sources. Additionally, it seems that there is no relation between plants sequestration capacity for nutrients and plant photosynthetic pathway. This work shows that nutrient cycling and accumulation processes by salt marsh halophytes contribute to reduce eutrophication (N and P retention) and also to reduce atmospheric CO₂ (C retention), highlighting salt marsh ecosystems services and the crucial role of halophytes in maintaining ecosystem functions and health.     

Variability in the nutritional value of the major copepods in Cape Cod Bay (Massachusetts, USA) with implications for right whales

The North Atlantic right whale, a seriously endangered species, is found in Cape Cod Bay (Massachusetts, USA) during the winter and early spring. During their residency in these waters, these whales are frequently observed feeding. This study evaluated spatial and temporal changes in the chemical composition (carbon weight and C/N ratio) of the food resource targeted by the right whales in Cape Cod Bay. The three taxa measured (Centropages typicus, Pseudocalanus spp., and Calanus finmarchicus) had highly variable chemical compositions resulting from the different life strategies and from fluctuations in their surrounding environment. The impact of seasonal variability in the energy densities of the food resource of right whales was calculated and compared to the energetic requirements of these whales. Calculations indicated that differences in the nutritional content of the zooplankton prey in Cape Cod Bay could have a considerable effect on the nutrition available to the right whales. Therefore, it is likely that using more precise estimates of the energetic densities of the prey of right whales would lead to a re‐evaluation of the adequacy of the food resource available to these whales in the North Atlantic.     

Habitat partitioning by five congeneric and abundant Choerodon species (Labridae) in a large subtropical marine embayment

The habitats occupied by the juveniles and adults of five morphologically similar, diurnally active and abundant Choerodon species in the large subtropical environment of Shark Bay, a “World Heritage Property” on the west coast of Australia, have been determined. The densities of the two life cycle stages of each Choerodon species in those habitats were used in various analyses to test the hypotheses that: (1) habitats are partitioned among these species and between their juveniles and adults; (2) such habitat partitioning is greatest in the case of the two Western Australian endemic species, i.e. Choerodon rubescens and Choerodon cauteroma; and (3) the extent of habitat partitioning between both of these two species and the only species that is widely distributed in the Indo-West Pacific, i.e. Choerodon schoenleinii, will be less pronounced. Initially, catches of each of the five congeneric species, obtained during other studies in Shark Bay by angling, spearfishing and otter trawling, were collated to elucidate the broad distribution of these species in that embayment. Underwater visual census was then used to determine the densities of the juveniles and adults of each Choerodon species at sites representing the four habitat types in which one or more of these species had been caught, i.e. reefs in marine waters at the western boundary of the bay and seagrass, reefs and rocky shorelines in the two inner gulfs. The compositions of the Choerodon species over marine (entrance channel) reefs and in seagrass were significantly different and each differed significantly from those in both inner gulf reefs and rocky shorelines, which were, however, not significantly different. Choerodon rubescens was restricted to exposed marine reefs, and thus occupied a different habitat and location of the bay than C. cauteroma, the other endemic species, which was almost exclusively confined to habitats found in the inner gulfs. Choerodon cauteroma differed from other Choerodon species in being relatively abundant in each inner gulf habitat and undergoing an ontogenetic change in habitat. Thus, while its juveniles lived predominantly in seagrass, its adults occupied mostly reefs and rocky shorelines. Choerodon cephalotes occurred almost exclusively in seagrass, while C. cyanodus was relatively most abundant along rocky shorelines. Choerodon schoenleinii, the only one of the five Choerodon species to be widely distributed outside Australia, occupied reefs and, to a lesser extent, rocky shorelines in the inner gulfs. The above differences in habitat among Choerodon species would reduce the potential for competition for spatial resources among these abundant and similar species and between the juveniles and adults of one of the species.