Occurrence and reproduction of tropical fishes in ocean warming hotspots of Japanese temperate reefs

Reproduction of tropical species beyond their geographic range associated with ocean warming is regarded as the key indicator of a range shift. However, the lack of historical breeding records poses challenges for detecting distinct range shifts of tropical fishes. To obtain baseline data of the current status of the occurrence and breeding activity of tropical pomacentrid and apogonid fishes in ocean warming hotspots of temperate reefs (Kochi and Wakayama, 33°N) of Japan, we conducted a two-year underwater visual survey and synthesized those data with recently published information. By combining data from the present as well as past studies, the results confirmed the occurrence of 52 pomacentrid and 34 apogonid species, whereas breeding activity was confirmed for 19 and 16 species, respectively. Species richness and abundance of recruitment periphery and breeding active species were high at the warmer site adjacent to the Kuroshio Current. Most observed species were found beyond their known geographic range. Some species showing active breeding were widespread tropical fishes (e.g., Amphiprion clarkii, Pomacentrus coelestis and Apogon notatus) and probably have established breeding populations irrespective of recent global warming. The winter sea water temperature around the study sites will continue to rise, increasing by >2 °C by the end of the century; therefore, our results are highly relevant and represent the first step to elucidate the potential range extension of tropical fishes into temperate reefs with climate change.     

Biogeography of sponge chemical ecology: comparisons of tropical and temperate defenses

Examples from both marine and terrestrial systems have supported the hypothesis that predation is higher in tropical than in temperate habitats and that, as a consequence, tropical species have evolved more effective defenses to deter predators. Although this hypothesis was first proposed for marine sponges over 25 years ago, our study provides the first experimental test of latitudinal differences in the effectiveness of sponge chemical defenses. We collected 20 common sponge species belonging to 14 genera from tropical Guam and temperate Northeast Spanish coasts (Indo-Pacific and Mediterranean biogeographic areas) and conducted field-based feeding experiments with large and small fish predators in both geographic areas. We use the term global deterrence to describe the deterrent activity of a sponge extract against all of the predators used in our experiments and to test the hypothesis that sponges from Guam are chemically better defended than their Mediterranean counterparts. Sympatric and allopatric deterrence refer to the average deterrent activity of a sponge against sympatric or allopatric predators. All of the sponges investigated in this study showed deterrent properties against some predators. However, 35% of the sponge species were deterrent in at least one but not in all the experiments, supporting the idea that predators can respond to chemical defenses in a species-specific manner. Tropical and temperate sponges have comparable global, sympatric, and allopatric deterrence, suggesting not only that chemical defenses from tropical and temperate sponges are equally strong but also that they are equally effective against sympatric and allopatric predators. Rather than supporting geographic trends in the production of chemical defenses, our data suggest a recurrent selection for chemical defenses in sponges as a general life-history strategy.     

Developing unified theories in ecology as exemplified with diversity gradients

A scientific field matures as its theoretical underpinnings consolidate around unified theories: conceptual structures consisting of a few general propositions that encompass a wide domain of phenomena and from which can be derived an array of models. We demonstrate this process with a synthetic theory of ecological gradients and species richness. Our unified theory rests on four propositions. First, variation in some environmental factor effects variation in the number of individuals creating a gradient. Second, in a uniform environment of fixed area, more individuals lead to more species. Third, the variance of an environmental factor increases with its mean for sites of equal area. Fourth, all nonmonotonic relationships (i.e., hump shaped or U shaped) require a trade-off in organismal performance or in population characteristics with respect to the environmental gradient. We identify 17 models that link environmental gradients with diversity, show their relationship to our framework, and describe issues surrounding their empirical testing. We illustrate how a general theory can be used to build new models such as that for the U-shaped productivity-diversity relationship. Finally, we discuss how our theory could be unified further with other theories of diversity and indicate other areas of ecology that are ripe for unification. By providing an example of the process of theory unification, we hope to encourage such efforts throughout ecology.     

Evolutionary ecology of mast-seeding in temperate and tropical oaks (Quercus spp.)

Mast-seeding is the synchronous production of large seed crops within a population or community of species every two or more years. This paper addresses three non-mutually exclusive hypotheses explaining the evolution of mast-seeding in temperate tree species, especially the genus Quercus: (1) mast-seeding is a consequence of mast-flowering which evolves to increased pollination efficiency in mast-flowering years; (2) mast-seeding has evolved as an anti-predator adaptation by which large seed crops during mast years satiate the seed predators and allow survival of some of the seeds; (3) selection on seed size by habitat can indirectly affect the evolution of masting if trees with large seeds require more time to accumulate reserves to mature those seeds. I find support for the pollination hypothesis in several wind-pollinated temperate tree species but not oaks. However, oaks show evidence favoring the predation and seed size hypotheses. I then develop a model to illustrate the relationships among the three hypotheses in their effects on the evolution of masting. Finally, using data from herbaria and Floras, the influence of selection via flowering, fruiting, and seed size in the evolution of masting in tropical oaks is discussed. I conclude that the need for a supra-annual cue to synchronize flowering and fruiting as well as the larger seed size found in many tropical oak species should contribute to the evolution of masting to a greater extent than seen among temperate oaks.     

Fruit fall in tropical and temperate forests: implications for frugivore diversity

There have been few attempts to compare fruit productivity throughout the world, although this is indispensable for understanding the global variations in frugivore diversity. The purposes of this study are (1) to reveal the patterns in fruit fall in tropical and temperate forests, (2) to examine the environmental factors (location, climate, and total litterfall) affecting these patterns, and (3) to assess the effect of fruit fall on frugivore diversity by using bird and primate data. Fruit fall was compared among 53 forests, from around the equator to the cool-temperate zone at 62°N, in Asia, Africa, North and South America, and Australia. Average ± SD of fruit fall (kg/ha/year) was 454 ± 258 in tropical, and 362 ± 352 in temperate forests. Fruit fall was exceptionally high in Australia (812 ± 461). When Australia was excluded, fruit fall significantly decreased with increasing absolute latitude and altitude, and fruit fall in tropical forest was 1.7 times larger than that in temperate forests (265 ± 227). Total litterfall affected fruit fall significantly, explaining 32, 28, and 64% of the variations of fruit fall in the entire data, tropical data, and temperate data, respectively. The fruit fall/litterfall ratio did not differ between temperate and tropical forests but was significantly higher in Australia than in other regions. Among climatic parameters (annual temperature, precipitation, actual evapotranspiration), a positive relationship was found between temperature and fruit fall in the entire dataset and within temperate forests. Fruit fall seemed to explain the temperate/tropical difference in frugivorous primate diversity to some extent, but not for frugivorous bird diversity. This study shows that the difference in fruit fall in tropical and temperate forests is smaller than that in frugivore diversity, and that it could explain at least part of the frugivore diversity.     

Elevational gradients, area and tropical island diversity: an example from the palms of New Guinea

The factors causing spatial variation in species richness remain poorly known. In this study, factors affecting species richness of palms (Palmae/Arecaceae) were studied along the elevational gradient of New Guinea. Interpolated elevational ranges were calculated from a database of all known collections for 145 species in 32 genera. The amount of land area at different elevations greatly affects the species richness gradient. If assessed in equal-elevation bands species richness appears to decline monotonically, but when assessed in equal-area bands species richness shows a pronounced mid-elevation peak, due to the large proportion of lowlands in New Guinea. By randomising species ranges within the total elevational gradient for palms and accounting for area, we found the mid-elevation peak to be consistent with a mid-domain effect caused by the upper and lower limits to palm distribution. Our study illustrates the importance of accounting for area in macroecological studies of richness gradients and introduces a novel yet simple method for doing this through the use of equal-area bands. Together, the effect of area and the mid-domain effect explain the majority of variation in species richness of New Guinea palms. We support calls for the multivariate assessment of the mid-domain effect on an equal footing with other potential explanations of species richness.     

Predator diversity and density affect levels of predation upon strongly interactive species in temperate rocky reefs

Indirect effects of predators in the classic trophic cascade theory involve the effects of basal species (e.g. primary producers) mediated by predation upon strongly interactive consumers (e.g. grazers). The diversity and density of predators, and the way in which they interact, determine whether and how the effects of different predators on prey combine. Intraguild predation, for instance, was observed to dampen the effects of predators on prey in many ecosystems. In marine systems, species at high trophic levels are particularly susceptible to extinction (at least functionally). The loss of such species, which is mainly attributed to human activities (mostly fishing), is presently decreasing the diversity of marine predators in many areas of the world. Experimental studies that manipulate predator diversity and investigate the effects of this on strongly interactive consumers (i.e. those potentially capable of causing community-wide effects) in marine systems are scant, especially in the rocky sublittoral. I established an experiment that utilised cage enclosures to test whether the diversity and density of fish predators (two sea breams and two wrasses) would affect predation upon juvenile and adult sea urchins, the most important grazers in Mediterranean sublittoral rocky reefs. Changes in species identity (with sea breams producing major effects) and density of predators affected predation upon sea urchins more than changes in species richness per se. Predation upon adult sea urchins decreased in the presence of multiple predators, probably due to interference competition between sea breams and wrasses. This study suggests that factors that influence both fish predator diversity and density in Mediterranean rocky reefs (e.g. fishing and climate change) may have the potential to affect the predators' ability to control sea urchin population density, with possible repercussions for the whole benthic community structure.     

Symbiont diversity in scleractinian corals from tropical reefs and subtropical non-reef communities in Taiwan

We examined zooxanthellae diversity in scleractinian corals from southern Taiwan and the Penghu Archipelago, a tropical coral reef and a subtropical non-reefal community, respectively. Zooxanthellae diversity was investigated in 52 species of scleractinian corals from 26 genera and 13 families, using restriction fragment length polymorphism (RFLP), and phylogenetic analyses of the nuclear small-subunit ribosomal DNA (nssrDNA) and large-subunit ribosomal DNA (nlsrDNA). RFLP and phylogenetic analyses of nuclear-encoded ribosomal RNA genes showed that Symbiodinium clade C was the dominant zooxanthellae in scleractinian corals in the seas around Taiwan; Symbiodinium clade D was also found in some species. Both Symbiodinium clade C and D were found in colonies of seven species of scleractinian corals. Symbiodinium clade D was associated with corals that inhabit either shallow water or the reef edge in deep water, supporting the hypothesis that Symbiodinium clade D is a relatively stress-tolerant zooxanthellae found in marginal habitats.Communicated by Biological Editor H.R. Lasker     

Distribution,movements and diet of nocturnal fishes on temperate reefs

We counted nocturnal fishes both day and night, and monitored the position of tagged individuals on temperate reefs in New South Wales, Australia. Pempheris affinis and P. multiradiata were the most abundant nocturnal planktivores on Sydneys rocky reefs and showed great differences in diel migration behaviour. Both species were observed in deep shelter sites during the day (5–10m), and most emerged into the water column at night. P. multiradiata was found to undergo extensive vertical and horizontal migrations. In contrast, P. affinis remained within daytime depth strata, with tagged individuals often moving less than 20m at night. Tagged adult P. affinis returned to tagging sites for up to 7weeks, indicating high site fidelity. Dietary analysis demonstrated that small and large pempherids differed in diet and the timing of foraging, suggesting a size-based transition from diurnal to nocturnal foraging. Stratified sampling of planktonic assemblages at different depths during the day and night showed spatial variation in the availability of prey items at different times of the day. Amphipods, the main prey of large fish, were only available during the night, and concentrated in shallow water, whereas decapod larvae, consumed mainly by small fish, were abundant day and night. Large P. affinis also fed on polychaetes, which were never found in the stomachs of P. multiradiata, suggesting that these species may have different prey requirements, or that these polychaetes are only found in deep water where foraging P. affinis were abundant. We found no general model for the Pempheridae. The movements and behaviour of nocturnal fishes varied greatly by species, and this may be due to differences in body size, and/or physiological (e.g. visual ability) and ecological constraints.     

Fruiting and flushing phenology in Asian tropical and temperate forests: implications for primate ecology

In order to understand the ecological adaptations of primates to survive in temperate forests, we need to know the general patterns of plant phenology in temperate and tropical forests. Comparative analyses have been employed to investigate general trends in the seasonality and abundance of fruit and young leaves in tropical and temperate forests. Previous studies have shown that (1) fruit fall biomass in temperate forest is lower than in tropical forest, (2) non-fleshy species, in particular acorns, comprise the majority of the fruit biomass in temperate forest, (3) the duration of the fruiting season is shorter in temperate forest, and (4) the fruiting peak occurs in autumn in most temperate forests. Through our comparative analyses of the fruiting and flushing phenology between Asian temperate and tropical forests, we revealed that (1) fruiting is more annually periodic (the pattern in one year is similar to that seen in the next year) in temperate forest in terms of the number of fruiting species or trees, (2) there is no consistent difference in interannual variations in fruiting between temperate and tropical forests, although some oak-dominated temperate forests exhibit extremely large interannual variations in fruiting, (3) the timing of the flushing peak is predictable (in spring and early summer), and (4) the duration of the flushing season is shorter. The flushing season in temperate forests (17–28 % of that in tropical forests) was quite limited, even compared to the fruiting season (68 %). These results imply that temperate primates need to survive a long period of scarcity of young leaves and fruits, but the timing is predictable. Therefore, a dependence on low-quality foods, such as mature leaves, buds, bark, and lichens, would be indispensable for temperate primates. Due to the high predictability of the timing of fruiting and flushing in temperate forests, fat accumulation during the fruit-abundant period and fat metabolization during the subsequent fruit-scarce period can be an effective strategy to survive the lean period (winter).     

History,ecology and trends for artificial reefs of the Ligurian sea,Italy

From 1970 to the present 10 artificial reef sites have been developed in coastal waters of the Ligurian Sea, Italy. They range from Ventimiglia, in the west, to La Spezia, in the east, with the largest and best known reef complex being located in the Gulf of Genoa at Loano and consisting of 2,745 m³, about 5,200 t of material and covering a surface of 350 ha. Design and construction practices have advanced from an initial, unsuccessful effort that used automobile bodies (now banned) to current use of custom-designed concrete modules deployed systematically. Funding for reef construction has come since 1983. The earliest aim of reefs was as a physical barrier to protect habitats against illegal otter trawl fishing. Newer objectives include habitat restoration, enhancement of biodiversity and fishing catch, and research to test materials and designs for physical and ecological performance. Reefs also functions as environmental observation stations, with the invasive species Caulerpa taxifolia (Vahl) C. Agardh, being recorded on the reef at Alassio. For some Artificial Reefs (Ars), benthic organisms and fishes, settlement, biomass and development of community are recorded. In Loano AR, immersed in 1986, more than 150 algae species are recorded, more than 200 benthic animal species and 78 species (87 taxa) of fishes. Fifty-six species (61 taxa) of fishes are recorded by visual census, the others are caught only by trammel net and long line. Trammel catches at Loano are on average about 2.32 kg/100 m net. Comparisons among ARs reveal that age of the reef, location and presence of seagrass meadows are crucial for success. An indication of functional equivalence between ARs and natural rocky reefs is seen if both fish and sessile macrobenthos are compared. After 34 years of investigation a database comprising at least one hundred scientific articles based on research programs of up to 15 years, and other unpublished reports, provides information to guide future planning of reefs. On the basis of acquired experience, some management advice is suggested and the best design for the basic module in the Ligurian sea is described. The role of ARs, providing protection of coastal environment against the illegal otter trawling, nursery, microhabitat and food supply, while increasing biodiversity, biomass of benthos and fishes, and facilities for ecotourism, is outlined.

Tree species diversity in temperate and tropical forest gaps: The role of lottery recruitment

How canopy gaps promote tree species coexistence in temperate and tropical forests is reviewed. Given that evidence for traditional resource-based niche partitioning of canopy gaps is weak, lottery recruitment, whereby colonization of vacant living space is random with respect to species identity, may be key to maintaining diversity. Gap formation events are not highly predictable and species’ propagules tend to be patchily distributed. For these reasons the predictability of gap-phase recruitment is low and lottery principles apply. Recruitment limitation from discontinuities in species’ propagule supplies in space and time may permit lottery recruitment in temperate and tropical forests. However, the relative importance of recruitment limitation in species-rich versus species-poor communities is unclear. Although lottery models with purely random recruitment in vacant sites can be applied, relatively complex models appear to better capture the essential features of forest community dynamics. For example, models with recruitment probabilities weighted by species abundance can produce the non-random trajectories often observed in communities. Models with local competitive displacement by individuals of dominant species can also produce non-random trajectories. Models with spatial structure, localized competition, and dispersal limitations may provide further insight into the effects of biotic interactions and recruitment limitation on forest dynamics.