Coexistence of coral reef fishes — a lottery for living space

Synopsis Data are summarised from studies of two reef fish communities — pomacentrids territorial on rubble patches, and fishes resident in small isolated colonies of coral. In each case there is evidence that availability of living sites limits numbers of fishes, and that similar species of fish use the same kinds of spaces. Priority of arrival as recruits, rather than subtle differences in requirements or competitive abilities of adults, appears to determine which species holds each site. Faced with a limited and patchy supply of living space, most reef fishes are sedentary as adults, and produce frequent clutches of pelagic larvae over extended breeding seasons In this way they maximise their chances of getting offspring into suitable living sites as such sites appear. It is argued that by adopting this strategy, reef fishes are preadapted for forming inter-specific lotteries for living space if several species with similar requirements occur together. Such lotteries among similar species may be a feature common to many reef fish communities, and may explain the typically high within-site diversity found in them.This paper forms part of the proceedings of a mini-symposium convened at Cornell University, Ithaca, N.Y., 18–19 May 1976, entitled Patterns of Community Structure in Fishes (G. S. Helfman, ed.).     

The role of herbivorous fishes and urchins in coral reef communities

Synopsis Herbivorous fishes and invertebrates are conspicious elements of coral reef communities where they predominate both in numbers and biomass. Herbivores and the coral reef algae on which they feed represent a co-evolved system of defense and counter-defense. Algal species have developed toxic, structural, spatial and temporal defense or escape mechanisms, while the herbivores employ strategies that involve anatomical, physiological and behavioral adaptations. Current research demonstrates that many reef fishes are highly selective in the algae they consume. Food selection in these fishes may be correlated with their morphological and digestive capabilities to rupture algal cell walls. Sea urchins select more in accordance with relative abundance, although certain algal species are clearly avoided.The determinants of community structure on coral reefs have yet to be established but evidence indicates a strong influence by herbivores. Reef herbivores may reduce the abundance of certain competitively superior algae, thus allowing corals and cementing coralline algae to survive. We discuss how the foraging activities of tropical marine herbivores affect the distribution and abundance of algae and how these activities contribute to the development of coral reef structure and the fish assemblages which are intimately associated with reef structure.This paper forms a part of the proceedings of a mini-symposium convened at Cornell University, Ithaca, N.Y., 18–19 May 1976, entitled Patterns of Community Structure in Fishes (G. S. Helfman, ed.).     

The basis of fish community structure: marine and freshwater comparisons

Synopsis Freshwater and marine fish communities are described and compared for arctic, boreal and tropical latitudes. Details of habitat characteristics, species numbers, and diel and seasonal differences in distribution are given for each community type. The order of increasing richness of fish species in these environments is (1) arctic lakes, (2) arctic marine, (3) boreal lakes, (4) tropical lakes, (5) boreal marine and (6) tropical marine. The richness of numbers of species can be related to a series of factors, each of which may function at some threshold value. These factors include climatic perturbation, solar radiation, spatial heterogeneity, available nutrient supply, availability of cover, and geological time. Discontinuities in the availability of some factors can be partially compensated for by torpor or aestivation; this effectively removes the fish from the community for a period of time. Increased diversity may also be effected through the diurnal/nocturnal shift in activity in some fish communities.The development of an organic matrix, notably macrophyte beds or coral reefs, may contribute significantly to an increase in diversity within fish communities. This matrix operates by an increase in spatial heterogeneity and in biological interactions. The apparent lack of resilience of high diversity fish communities can be related to the characteristics of the underlying organic matrix. A change in the matrix will cause a change in the level of fish diversity that can be maintained in the system.This paper forms part of the proceedings of a mini-symposium convened at Cornell University, Ithaca, N.Y., 18–19 May 1976, entitled Patterns of Community Structure in Fishes (G.S. Helfman, ed.).     

The Structures and Dynamics of Fish Communities in an Okinawan Coral Reef: Effects of Coral-based Habitat Structures at Sites with Rocky and Sandy Sea Bottoms

Although there have been many studies on ecological factors responsible for the organization of reef fish communities, most of the studies have focused on isolated habitats. However, findings from isolated habitats cannot necessarily be applied to fish communities in other habitats (e.g., a continuous habitat). In this study, therefore, we examined the structures of fish communities (abundance, species richness and species composition), and the dynamics of fish communities (seasonal changes in abundance, species richness and species composition) over a 2-year period in two different habitats (continuous habitat and isolated habitat) in an Okinawan coral reef. We established eight permanent quadrats (8m × 8m) on a rocky reef flat (continuous habitat) and rock reef patches surrounded by a sandy sea bottom (isolated habitat). The abundance and species richness of fishes such as pomacentrids, labrids, chaetodontids and acanthurids were greater in quadrats located in the continuous habitat, whereas those of blenniids, gobiids and mullids were greater in quadrats located in the isolated habitat. This caused marked differences between the fish community structure at the two sites. Seasonal and annual changes in fish community structure were relatively small at the continuous habitat site (>0.7 similarity based on C index) but were large at the isolated habitat site (C < 0.4), indicating that the fish community structure was relatively stable at the continuous habitat site but unstable at the isolated habitat site throughout the study period. Spatial differences between the fish community structures in the four quadrats at the continuous habitat site were small (C > 0.65 in most sites), but these differences were large at the isolated habitat site (C < 0.4). Our findings suggest that habitat structure (spatial arrangement of habitats) affects both spatial and seasonal differences in the reef fish community structure. The results also suggest that the main mechanisms underlying organization of reef fish communities in continuous and isolated habitats are different.     

The use of artificial reefs in enhancing fish communities in Singapore

Intense development of the coastal zone in Singapore has resulted in the degradation of much of the marine ecosystem. In order to restore and enhance fish communities of denuded areas, an artificial reef consisting of a tyre reef and a concrete reef, was established in the vicinity of the southern islands of Singapore. Results from fish visual censuses after the establishment of the artificial reef indicated an increase in numbers of juveniles and adults. A total of 37 and 32 fish species were recorded over a period of 1/2 years at the concrete and tyre reefs respectively. The dominant fish families were Pomacentridae, Labridae, Chaetodontidae, Apogonidae, Gobiidae and Nemipteridae. The artificial reefs also serve as a nursery ground for some species (e.g. Neopomacentrus sp.) which are important primary consumers of algae on natural reefs. Greater numbers of target (food-important) fishes were observed at the concrete reef while the tyre reef harboured more juveniles and smallersized adults. The results indicate that the concrete modules were more effective than the tyre reef in terms of fish abundance per unit volume. Such structures can enhance the biological resources of relatively unproductive areas.     

Money-in-the-bank: a model for coral reef fish coexistence

Synopsis An alternative to the reef fish lottery model is proposed for explaining instances of coexistence of reef fishes without apparent spatial resource partitioning. This model is termed money-in-the-bank because of a financial analogy used to explain it. It stresses the importance of habitats that can support only one of two or more closely related species that coexist elsewhere. Populations living in such monospecific habitats could, according to the model, produce enough larvae to repopulate these habitats plus an excess that may settle in the multispecific habitats. Possible examples among cardinalfishes are given.This paper forms part of the proceedings of a mini-symposium convened at Cornell University, Ithaca, N.Y., 18–19 May 1976, entitled Patterns of Community Structure in Fishes (G. S. Helfman, ed.).     

Coral reef fish communities: a compromise view

Synopsis The fishes associated with coral reefs offer excellent opportunities for the study of the factors that determine the species composition of complex, highly interactive communities. Amenable to sampling and direct observations many patch reefs are small enough to be studied as entire units and yet diverse enough to include a wide range of interactions.Coral reef fishes appear to be highly specialized in morphology, color, behavior and life cycles and yet colonization experiments and repeated censusing have shown a surprisingly high variation in the fishes that are associated in similar habitats or in the same habitat at different times. This has led to two different views: (i) the chaos view that holds that the species composition is due to random factors and chance colonization, and (ii) the order view that resource sharing adaptations determine which species can live together.This paper reviews some of the obvious adaptations of reef fishes. An examination of the maximum sizes reached by infaunal reef dwellers shows that the largest individual of each species differs by a constant proportion from the next larger and next smaller species. This suggests growth limitations by interspecies competition. A hypothetical model showing how this might work is offered and it is shown that as long as there are more species than can be accommodated in the community at any one time the number of combinations is great enough to give the appearance of randomness even though the individual species may have precise environmental requirements.This paper forms part of the proceedings of a mini-symposium convened at Cornell University, Ithaca, N.Y., 18–19 May 1976, entitled Patterns of Community Structure in Fishes (G. S. Hellman, ed.).     

The effect of leaf source and developmental stage on shoot organogenic potential of sweetgum (Liquidambar styraciflua L.) leaf explants

Leaf explants harvested from shoot proliferating cultures and intact plants of Liquidambar styraciflua Variegata were placed on solidified Woody Plant medium supplemented with 0.1 mgl^-1 (0.5 M) naphthaleneacetic acid and 2.5 mgl^-1 (11.1 M) benzyladenine to initiate shoot meristems directly. Leaves from intact plants produced over 4 times more adventitious shoots than leaves from in vitro shoots and had a greater tendency to form shoots on the lamina. The relative developmental age of leaf tissue used dramatically influenced the shoot organogenic response observed for leaf explants from intact plants of L. styraciflua Variegata and Moraine.-Leaves that were either 20% or 50% of full size and still actively expanding were superior to other developmental stages for shoot organogenesis. As developmental leaf age increased throughout the period of leaf expansion, the number of shoots forming on the petiole stub remained constant, whereas shoot formation on the lamina increased 8 fold. Shoots derived from Variegata leaves rooted well and grew normally as plants. Differences in rooting ability and plant size could be detected between groups that had been separated according to explant source (in vitro vs. intact plant) and the location of shoot formation (petiole vs. lamina).     

Aggression in damselfish: habituation by adults of Pomacentous parritus to juvenile intruders

Synopsis Successful recruitment of juvenile coral reef fishes may depend, in part, upon the aggressive behavior of adults already on the reef. In addition to initial levels of aggression, changes in aggressive behavior of adults, e.g., due to habituation, may have an even greater influence on recruiting juveniles. Adult males of the bicolor damselfish, Pomacentrus partitus, were used as subjects to study habituation of aggression toward conspecific and congeneric (P. variabilis) juveniles. Adults, held in a 1000 liter aquarium, habituated after 4 hours of constant exposure to juveniles restrained in a 1 liter model bottle. Stimulus strength of the juveniles depended on their species identity, size and proximity to the resident adult's shelter. There was a recovery of aggression with a change in stimulus location, but at the same location, a stimulus of greater strength was required to bring about recovery. The implications of these findings for coral reef fish community structure are discussed.     

Appetitive conditioning technique reveals behavioural limits to passage performance in fishes

Numerous environmental features have the potential to act as barriers to fishes in the field. Passage of these barriers depends on two main properties: willingness (behavioural) and physiological capacity. The physiological swimming capacity of fishes has been well studied. However, because most barrier research has focused on the population level effects of man – made barriers, little attention has been given to the behavioural aspects of fish passage. We used appetitive conditioning to create a simple laboratory based protocol to estimate behavioural limits to fish passage. We tested our protocol using barrier height (or passage space) as a model. Our protocol successfully identified behavioural limits in two species, juvenile rainbow trout, Oncorhynchus mykiss, and fathead minnows, Pimephales promelas. Our most interesting observation was that even a partial, submerged barrier had the potential to block fish movements. Our results suggest a potential use for conditioning techniques in assessing factors that constitute behaviour barriers to fishes. This information may help us to better predict fish movements in the field and may help us to design barriers that are more efficient at passing fishes.     

Reproductive strategies of coastal marine fishes in the tropics

Synopsis A synthesis of ethnobiological, behavioral and physical oceanographic information leads to the conclusion that temperate zone models of reproductive strategy are inapplicable to many fishes of the coastal tropics. Intense predation appears to exert heavy selection pressure on fishes that spend their adult lives in coral, mangrove or tropical seagrass communities. Many exhibit spawning behaviors and spawn at times and locations that favor the transport of their pelagic eggs and pelagic larvae offshore where predation is reduced. This creates a countervailing selection pressure — the need to return the larvae to shallow water once they are ready to colonize their post-larval habitats. Accordingly, spawning is often concentrated at times of the year when prevailing winds or currents are at their weakest, thereby reducing the transport of larvae long distances from where they originated. Spawning is also concentrated in the vicinity of nearshore gyres which similarly favor the ultimate return of the larvae to their natal area. Among these species, therefore, offshore larval dispersal does not seem to be an adaptation for dispersal of the species, but rather an evolutionary response to intense predation pressure in the adult habitats. Lunar reproductive periodicity is more common among these species than has previously been recognized, and is one of the strategies employed to enhance the offshore flushing of eggs and larvae.This paper forms part of the proceedings of a mini-symposium convened at Cornell University, Ithaca, N. Y. 18–19 May 1976, entitled Patterns of Community Structure in Fishes (G. S. Helfman, ed.).Contribution No. 524, Hawaii Institute of Marine Biology, University of Hawaii, Honolulu.     

Microhabitat association and temporal stability in reef fish assemblages on massive <Emphasis Type="Italic">Porites</Emphasis> microatolls

A massive Porites microatoll generally has three types of microhabitat at the top, side, and base of the microatoll. The purpose of the present study was to analyze microhabitat associations of reef fish on microatolls to determine whether habitat characteristics play an important role in the structuring of reef fish assemblages in a patchy habitat. We also investigated temporal stability of reef fish assemblage structures over a period of 17 months to determine whether fish assemblage structures vary in a random manner. The results of correspondence analysis indicated species-specific habitat associations for pomacentrids (five species) and labrids (seven species). The degree of temporal stability of fish assemblage structures, calculated by Piankas index, was relatively high in a large-sized microatoll (0.503–0.831: 3.6m in diameter), in which microhabitat associations of fishes were clearly observed. The present study suggests that a microhabitat association is one of the important factors responsible for organization of reef fish assemblages in a microatoll.     

Genetic selection of lambda phage clones from a gene clonotheque

Summary A genetic procedure for selection of specific clones, by homologous recombination between clones from a gene clonotheque and sequences cloned into a plasmid, was developed. Resulting clones are isolated in transduction experiments by plating infected Escherichia coli cells under conditions selecting for the antibiotic resistance marker carried by the plasmid. The feasibility of the method was demonstrated in a model test system as well as by isolation of -interferon-specific sequences from the human gene clonotheque.     

Reminiscence about‘Chloropseudomonas ethylicum’ and the FMO-protein

In 1961 the green sulfur bacterium-containing mixed culture known asChloropseudomonas ethylicum was brought to Brookhaven National Laboratory (USA) from Moscow State University (USSR). The water-soluble bacteriochlorophylla-protein (FMO-protein) was extracted, purified and characterized by absorption and circular dichroism spectroscopy, by X-ray crystallography and by primary structure determination.     

Have fishes had their chips? The dilemma of threatened fishes

Synopsis The conservation status and factors threatening fishes worldwide are reviewed in order to introduce a series of one-page articles on Threatened fishes of the world, and to encourage the incorporation of information on threatened fishes into international conservation programmes. Information on fish extinction and threat rates are compared with those of other animal groups, and the unique characteristics of fish conservation problems are highlighted. At present 979 species of fishes are listed as threatened in the IUCN Red List and at least 36 species and three subspecies are listed as recently extinct. It is argued that these figures are probably gross underestimates and that they may mislead conservation authorities and resource users about the seriousness of the situation. Freshwater fishes may be the most threatened group of vertebrates after the Amphibia. Urgent action is required to save many narrowly endemic, stenotopic species from extinction, especially in Africa, Asia and South America. The conservation of common species that drive essential ecological processes is also important. Anthropogenic pressures, especially habitat degradation, the introduction of invasive species and pollution, on inland and coastal waters are particularly severe and many major fish communities are threatened with elimination throughout the world. The conservation of marine fishes is complicated by the fact that it is difficult to ascertain their rarity. The importance of the retention of genetic variation is highlighted, and both orthodox and innovative conservation measures are encouraged. Further research on minimum viable populations, genetics, and the factors that cause fishes to become vulnerable to extinction, is urgently required.Invited editorial     

Trophic and spatial interrelationships in the fish species of an Ontario temperate lake

Synopsis Analysis of the fish faunas of Lake Opinicon and other small, cold temperate Ontario water bodies shows that the component species differ in body size, morphology, abundances, habitats, diurnal and seasonal habitat utilization patterns, diets, dietary changes with age, reproductive strategies, and population turnover rates. These differences are detailed.The number of species occurring in a lake is partly due to historic factors, the number of habitats available, and morphological, behavioral and ecological adaptations that, by channelling their owners towards alternative resources, permit species to co-occur. Diet overlap values between most species are low except for the congeneric bluegill and pump-kinseed sunfishes, where values are moderate. These are the two commonest species in the lake and other parameters must increase the ecological differences between these two species.Lake Opinicon is a highly variable ecosystem. Part of this variability stems from the seasonal nature of the environment and the fact that different resources reach their peak abundance at different times of the year. Ecological overlap levels between fish species fluctuate greatly in the course of the season as species switch from, or move on to, different resources. Population levels in different habitats also vary seasonally. Species adaptations and interaction patterns were presumably evolved over a long period; most of these adaptations undoubtedly developed before the component species colonized the lake.This paper forms part of the proceedings of a mini-symposium convened at Cornell University, Ithaca, N.Y., 18–19 May 1976, entitled Patterns of Community Structure in Fishes (G. S. Helfman, ed.).     

Ecological and morphological shifts in Lake Michigan fishes: glimpses of the ghost of competition past

Synopsis Both historical patterns and recent evidence of resource partitioning and complementarity within the Lake Michigan fish community provide circumstantial evidence for interspecific competition. But competition is difficult to document in the field without controlled experimentation. In Lake Michigan, controlled experiments on competition within the fish community are nearly impossible, but we still need to understand the interactions among the dominant fishes. For this purpose, I have relied upon hypothesis-based field observation, natural experiments in the field and designed laboratory experiments to evaluate competitive interactions. Resource use patterns and trophic morphology of the bloater, Coregonus hoyi, a native cisco, from samples taken before alewife, Alosa pseudoharengus, became abundant (1960) were compared to more recent data (1979–80). After the alewife density increase, bloaters had significantly fewer and shorter gill rakers. This suggests a morphological shift toward greater benthic foraging efficiency in response to high abundances of an efficient pelagic planktivore, alewife. Resource use comparisons suggested that bloaters now shift from pelagic zooplanktivory to benthic habitats and diets at least two years earlier in their life history than they did before alewife became abundant. This evidence, albeit not experimental, provides strong support for the importance of competition in the structure of the current Lake Michigan fish community.In Lake Michigan, seasonal thermal habitat compression can pack fish into a narrow thermal zone across the lake bottom, leading to increased habitat overlap, reduced prey availability and fish diets containing fewer and smaller prey. Thermal habitat compression, which can occur intermittently through the season, may create competitive bottlenecks which help maintain the observed resource partitioning among these fishes.     

Scientists Prefer them Blind: The History of Hypogean Fish Research

The history of hypogean fish research has been strongly influenced by neo-Lamarckism (including orthogenesis) and typological thinking. Only in the last few decades neo-Darwinism has made any inroads in the research approach to this subject. The majority of the most distinguished and productive hypogean fish researchers have used their research subjects to confirm their own views on evolution rather than to use those subjects as a spring of knowledge to enrich mainstream biological thought. Of these views, I found that the most perversive of all is the notion of evolutionary progress that has led many researchers to envision hypogean fishes as prime examples of regressive evolution. I propose that the utilization of hypogean fishes for the study of convergent evolution should catapult these subjects of research into prime objects of evolutionary ideas.     

The contribution of small individuals to density-body size relationships: examination of energetic equivalence in reef fishes

A key relationship in ecology is that between density and body size, with the emphasis placed on energetic rules constraining the abundance of larger organisms below that of smaller organisms. Most studies have focused upon the density-body size relationship at the species level. However, energy is gathered at an individual level. We therefore examined this relationship in a coral reef fish assemblage, focusing on individuals. Using a comprehensive data set, with over 14,000 observations we found that the relationship between local density and adult body size differs from the linear relationship predicted by the energetic equivalence rule. However, excluding the smallest size classes, the relationship between body size and individual abundance for intermediate to larger fish did not depart from the predicted –0.75. Unlike plants and intermediate to large reef fishes, the smallest fishes appear to have constraints that may reflect different patterns of resource acquisition.     

Spatial patterns and ecology of benthic communities on a high-latitude South Florida (Broward County,USA) reef system

High-latitude reef communities consisting of typical Caribbean fauna of variable composition and density exist on four parallel ridges at varying depths along the Broward County (Florida, USA) coast. Two of these ridges, at 7–13 and 15–30 m in depth, are drowned early Holocene coral reefs of 5 and 7 ky uncorrected radiocarbon age. In this study, community data were collected on each reef using 50-m, line-intercept transects. Using multivariate non-metric statistics, the data show distinct differences in benthic community structure across several spatial gradients. Diversity indices revealed that while all values were low, species diversity (H) was consistent throughout the county, and species richness (d) and evenness (J) increased along a north-south gradient. Scleractinian coral (27 species) cover was low (<6%) in all areas, and Montastrea cavernosa dominated as the major hermatypic scleractinian. Notably absent was the major Caribbean reef-builder Acropora palmata. A rich alcyonacean fauna was present on all studied reefs, and these were typically the most important faunal group determining community structure. Although during the 2-year period 2000–2002 mean water temperatures never fell below 20 °C, scleractinia were small in size.