Population regulation by post-settlement mortality in two temperate reef fishes

 Input of individuals dispersing into open populations can be highly variable, yet the consequences of such variation for subsequent population densities are not well understood. I explored the influence of variable input (”supply”) on subsequent densities of juveniles and adults in open local populations of two temperate reef fishes, the bluebanded goby (Lythrypnus dalli) and the blackeye goby (Coryphopterus nicholsii). Variable recruitment was simulated by stocking a natural range of densities of young fishes on replicate patch reefs. Density and mortality of the stocked cohorts were followed over time, until the fishes reached maturity. Over the first day of the experiments, mortality of both species was significantly density-dependent; however, there was still a very strong relationship between density on day 1 and density on day 0 (i.e., simulated recruitment was still an excellent predictor of population density). At this point in the study, the main effects of density-dependent mortality were to reduce mean densities and variation about the mean. Over the period from the start of the experiments until the time when maturity was reached by each species (about 1 and 3 months for Lythrypnus and Coryphopterus, respectively), mortality was strongly density-dependent. Such strong density-dependent mortality virtually eliminated any linear relationship between adult density and ”recruit” density. However, for both species, the relationship between these two variables was well fit by an asymptotic curve, with the asymptotic density of adults equal to c. 3/m² for Coryphopterus, and c. 10/m² for Lythrypnus. Natural recruitment (via settlement of larvae) to the reefs over the period of the study (9 months) was above the asymptotic densities of adults for the two species, even though the study did not encompass the periods of peak annual recruitment of either species. This suggests that adult populations of these two gobies may often be limited, and regulated, by post-settlement processes, rather than by input of settlers. Other studies have shown that mortality of the two species is density-independent, or only weakly density-dependent, on reefs from which predators have been excluded. Hence, it appears that predators cause density-dependent mortality in these fishes. Received: 26 November 1996 / Accepted: 5 April 1997     

Hybridization of reef fishes at the Indo-Pacific biogeographic barrier: a case study

Hybridization is recognized as an important source of genetic variation. In some reef fishes, including the Acanthuridae, hybridization has been detected due to intermediate colouration. This study used a molecular genetic approach to investigate hybridization in two Acanthurid species: Acanthurus leucosternon and Acanthurus nigricans, which have Indian and Pacific Ocean distributions respectively and are sympatric in the eastern Indian Ocean. In this area a putatitve hybrid, Acanthurus cf. leucosternon has been recognized based on intermediate colouration and restriction to the sympatric region of otherwise allopatric putative parental species. This study aimed to test this hypothesis using genetic tools. The three species were sampled from Cocos (Keeling) and Christmas Islands, the biogeographic boundary where many Indian and Pacific Ocean biota meet. Representatives from allopatric populations of both parental species and outgroups were also sampled. Mitochondrial COI and intron 1 of the nuclear ribosomal protein S7 were sequenced from 13 and 30 specimens respectively. Although sample sizes in this study are relatively small and more genetic data, including an extended phylogeographic sampling, is required to further evaluate these findings, the COI results support hybrid origins of Acanthurus cf. leucosternon, but S7 data are inconclusive due to the possibility of incomplete lineage sorting. The fourfold more abundant Acanthurus nigricans is most often the maternal parent. Inter-fertile hybrids apparently backcross with rare Acanthurus leucosternon males, transferring Acanthurus nigricans mitochondria to this species. These results suggest that Acanthurus leucosternon may eventually be lost from these islands, due to their relative rarity and introgressive hybridization.     

Temporal diversification of Mesoamerican cichlid fishes across a major biogeographic boundary

The Mexican Neovolcanic Plateau sharply divides the vertebrate fauna of Mesoamerica where the climate of both the neotropics and temperate North America gradually blend. Only a few vertebrate groups such as the Heroine cichlids, distributed from South America to the Rio Grande in North America, are found both north and south of the Neovolcanic Plateau. To better understand the geography and temporal diversification of cichlids at this geologic boundary, we used mitochondrial DNA sequences of the cytochrome b (cyt b) gene to reconstruct the relationships of 52 of the approximately 80 species of Heroine cichlids in Mesoamerica. Our analysis suggests several cichlids in South America should be considered as part of the Mesoamerican Heroine clade because they and the cichlids north of the Isthmus of Panama are clearly supported as monophyletic with respect to all other Neotropical cichlids. We also recovered a group containing species in Paratheraps+Paraneetroplus+Vieja as the sister clade to Herichthys. Herichthys is the only cichlid clade north of the Mexican Plateau and it is monophyletic. Non-parametric rate smoothing of cichlid cyt b sequence resulted in an estimated divergence time of approximately 6 million years for Herichthys. This temporal diversification is concordant with divergence times estimated for anurans in the genus Bufo, a group that exhibits a similar geographic distribution. Our results indicate the 5-million-year-old extension of the Mexican Neovolcanic Plateau to the Gulf Coast of Mexico has strongly influenced the current transition between the vertebrate faunas of the Neotropics and Nearctic.     

Habitat use patterns of fishes across the mangrove-seagrass-coral reef seascape at Ishigaki Island,southern Japan

To clarify seascape-scale habitat use patterns of fishes in the Ryukyu Islands (southern Japan), visual censuses were conducted in the mangrove estuary, sand area, seagrass bed, coral rubble area, branching coral area on the reef flat, and tabular coral area on the outer reef slope at Ishigaki Island in August and November 2004, and May, August and November 2005. During the study period a total of 319 species were observed. Species richness and abundance were highest in the branching and tabular coral areas, followed in order by the seagrass bed and mangrove estuary, and coral rubble and sand areas, in each month. Cluster analysis resulted in a clear grouping of assemblage structures by habitat type rather than by census month. SIMPER analysis showed that fish assemblages in the tabular coral area were mainly characterized by Acanthurus nigrofuscus, Pomacentrus lepidogenys, P. philippinus and P. vaiuli, the branching coral area by Chromis viridis and Pomacentrus moluccensis, the coral rubble area by Amblyeleotris steinitzi and Ctenogobiops pomastictus, the seagrass bed by Cheilio inermis, Lethrinus atkinsoni and Stethojulis strigiventer, the sand area by Valenciennea longipinnis, and the mangrove estuary by Gerres oyena, Lutjanus fulvus and Yongeichthys criniger. Moreover, fishes exhibited two habitat use strategies, inhabiting either a single or several specific habitats throughout their benthic life history stages, or having a possible ontogenetic habitat shift from the mangrove estuary or seagrass bed to coral-dominated habitats (e.g., Lethrinus atkinsoni, Lethrinus obsoletus, Lutjanus fulviflamma, Lutjanus fulvus, Lutjanus gibbus, Lutjanus monostigma and Parupeneus barberinus), suggesting that the mangrove estuary and seagrass bed have a nursery function.     

Behaviour of temperate and sub-tropical reef fishes towards a stationary SCUBA diver

Using remote underwater stereo-video systems we examined fish behaviour towards a stationary SCUBA diver at temperate (Rottnest Island) and sub-tropical (Houtman Abrolhos Islands) reefs in Western Australia. Changes in species richness, relative abundance, fish length, and the mean distance of individual fish from stereo-video cameras, in the presence and absence of a SCUBA diver, were assessed to infer changes in behaviour. Results show that a stationary SCUBA diver may obtain accurate measures of species richness and of the composition of fish assemblages in an area. However, the usefulness of these measures to reflect changes in fish behaviour appears limited as responses of fish towards the stationary SCUBA diver were highly species specific. Several species differed in their mean relative abundance (Heterodontus portusjacksoni, Coris auricularis, Thalassoma lunare), length (Ophthalmolepsis lineolatus, C. auricularis), and the distance to which they would approach the stereo-video systems (Kyphosus sydneyanus, Scarus schlegeli) when a SCUBA diver was present. Here, species-specific changes in the behaviour of several common and abundant fish species towards a stationary SCUBA diver advises caution to avoid biases when interpreting results obtained by SCUBA divers.     

SMURFs: standard monitoring units for the recruitment of temperate reef fishes

I evaluated a standard monitoring unit for the recruitment of reef fishes (SMURF) as a tool for ascertaining spatial and temporal patterns of reef fish recruitment in central California, USA. SMURFs consisted of a 1.0×0.35 m dia. cylinder of fine mesh plastic grid that contained a folded section of larger mesh plastic grid. SMURFs collected new recruits of 20 species of fish with 92% of the individuals collected from 10 species, mostly rockfish (genus Sebastes). An experiment varying depth of SMURFs in the water column (surface, mid-depth, or bottom) showed that surface SMURFs collected the greatest diversity of species and significantly greater abundance for eight species, with two species having significantly greater abundance on mid-depth SMURFs and three species having significantly greater abundance on bottom SMURFs. A comparison of cumulated recruitment from SMURFs that varied in sampling frequency (removal of new recruits every 1-3, 7, or 28 days) suggested that increasing the time between sampling caused a significant decrease in recruitment estimates for some species but not for others. To determine how well temporal patterns of recruitment to SMURFs reflected patterns to nearby reefs, I compared within season temporal patterns of recruitment to SMURFs with that at nearby reefs, estimated by visual transect surveys conducted on scuba. Temporal patterns of recruitment to SMURFs were significantly and positively related to early recruitment on reefs for one group of benthic-algal associated rockfish species when diver surveys were lagged by 30 days (r=0.87) and for another group of canopy-algal associated rockfish species when lagged by 5 days (r=0.72). SMURFs appeared to be an effective and efficient method for indexing relative rates of delivery of competent juveniles for many temperate nearshore reef fishes.     

Comparative phylogeography in a genus of coral reef fishes: biogeographic and genetic concordance in the Caribbean

Geographic barriers that limit the movement of individuals between populations may create or maintain phylogenetically discrete lineages. Such barriers are often inferred from geographic surveys of a single mitochondrial marker to identify phylogenetic splits. Mitochondrial DNA, however, has an effective population size one-fourth that of nuclear DNA, which can facilitate the rapid evolution of monophyletic mtDNA lineages in the absence of geographic barriers. The identification of geographic barriers will thus be more robust if barriers are proposed a priori, and tested with multiple independent genetic markers in multiple species. Here, we tested two proposed marine biogeographic breaks located at the Mona Passage in the Caribbean Sea and at the southern end of Exuma Sound in the Bahamas. We sequenced mitochondrial cytochrome b (400 bp) and nuclear rag1 (573 bp) for nine species and colour forms (183 individuals total) within the teleost genus Elacatinus (Gobiidae) that span the proposed breaks. Our results showed that Mona Passage separated mtcyb and rag1 lineages, with no genetic exchange between populations separated by just 23 km. However, the Central Bahamas barrier was only weakly supported by our data. Importantly, neither barrier coincided with deep genetic splits. This suggests that these two barriers did not initially isolate regional populations, but instead disrupt ongoing gene flow between regions. Our inferred relationships further suggested a division of the Caribbean region into northwestern and southeastern regions, a pattern reflected by some freshwater and terrestrial vertebrates. Our results, coupled with genetic and demographic data from other reef fishes and corals, provide robust support for the Mona Passage as a long-term biogeographic barrier for Caribbean animals.     

Behavioral thermoregulatory abilities of tropical coral reef fishes: a comparison with temperate freshwater and marine fishes

Synopsis Eight species in six different families of tropical marine reef fishes from the Indo-West Pacific region (Naso lituratus, Zebrasoma.flavescens, Balistes fuscus, B. vidua, Forcipiger longirostris, Echidna zebra, Cromileptes altivelis, Canthigaster jactator) were tested for ability to thermoregulate behaviorally in electronic shuttleboxes. All of these species preferred mean temperatures between 20 and 30°C, but differed considerably in thermoregulatory precision. All species avoided lethal high or low temperatures (i.e., they did not die during the tests), and some species thermoregulated as precisely as temperate species. Some temperate species prefer higher temperatures (above 30°C) than do these tropical reef species.     

Diel distribution patterns of fishes in a temperate large lowland river

During the day, juveniles and typical inshore species in the lower Oder River, Germany, inhabited the littoral (0·05–1·50 m deep) while potamal species inhabited the mid‐channel (40–100 m from the banks). During the night, two behavioural groups were distinguished: 1) potamal species and large individuals of other species, moved from offshore to the littoral, and 2) nocturnal foraging species, became active and increased in inshore and offshore habitats. Inshore, the most significant differences at night was the increasing density of silver bream Blicca bjoerkna increasing both total biomass and total length of fishes caught. Offshore, the increasing density of whitefin gudgeon Gobio albipinnatus at night was most significant. Additional night electrofishing improved the assessment of the abundance, age and size structure of typical potamal fish species. It should be incorporated in large river fish monitoring programmes.     

Fundamental biogeographic patterns across the Mexican Transition Zone: an evolutionary approach

Transition zones, located at the boundaries between biogeographic regions, represent events of biotic hybridization, promoted by historical and ecological changes. They deserve special attention, because they represent areas of intense biotic interaction. In its more general sense, the Mexican Transition Zone is a complex and varied area where Neotropical and Nearctic biotas overlap, from southwestern USA to Mexico and part of Central America, extending south to the Nicaraguan lowlands. In recent years, panbiogeographic analyses have led to restriction of the Mexican Transition Zone to the montane areas of Mexico and to recognize five smaller biotic components within it. A cladistic biogeographic analysis challenged the hypothesis that this transition zone is biogeographically divided along a north‐south axis at the Transmexican Volcanic Belt, as the two major clades found divided Mexico in an east‐west axis. This implies that early Tertiary geological events leading to the convergence of Neotropical and Nearctic elements may be younger (Miocene) than those that led to the east‐west pattern (Paleocene). The Mexican Transition Zone consists of five biogeographic provinces: Sierra Madre Occidental, Sierra Madre Oriental, Transmexican Volcanic Belt, Sierra Madre del Sur, and Chiapas. Within this transition zone, at least four cenocrons have been identified: Paleoamerican, Nearctic, Montane Mesoamerican, and Tropical Mesoamerican. Future studies should continue refining the identification of cenocrons and the reconstruction of a geobiotic scenario, as well as integrating ecological biogeographic studies, to allow a more complete understanding of the patterns and processes that have caused the biotic complexity of this transition zone.     

Variation in coral-associated cryptofaunal communities across spatial scales and environmental gradients

Coral Reefs - Most of the diversity on coral reefs is in the cryptofauna, the hidden organisms that inhabit the interstitial spaces of corals and other habitat-forming benthos. However, little is...     

Uncertainty in the detection of disturbance spatial patterns in temperate forests

The use of individual-based models in the study of the spatial patterns of disturbances has opened new horizons in forest ecosystem research. However, no studies so far have addressed (i) the uncertainty in geostatistical modelling of the spatial relationships in dendrochronological data, (ii) the number of increment cores necessary to study disturbance spatial patterns, and (iii) the choice of an appropriate geostatistical model in relation to disturbance regime. In addressing these issues, we hope to contribute to advances in research methodology as well as to improve interpretations and generalizations from case studies.We used data from the beech-dominated Žofínský Prales forest reserve (Czech Republic), where we cored 3020 trees on 74 ha. Block bootstrap and geostatistics were applied to the data, which covered five decades with highly different disturbance histories. This allowed us to assess the general behavior of various mathematical models. Uncertainty in the spatial patterns and stability of the models was measured as the length of the 95% confidence interval (CI) of model parameters.According to Akaike Information Criterion (AIC), the spherical model fitted best at the range of ca. 20 m, while the exponential model was best at the range of ca. 60 m. However, the best fitting models were not always the most stable. The stability of models grew significantly with sample size. At <500 cores the spherical model was the most stable, while the Gaussian model was very unstable at <300 cores. The pure nugget model produced the most precise nugget estimate. The choice of model should thus be based on the expected spatial relations of the forest ecosystem under study. Sill was the most stable parameter, with an error of ±6–20% for ≥1110 core series. By contrast, practical range was the most sensitive, with an error of at least ±59%. The estimation of the spatial pattern of severe disturbances was more precise than that of fine-scale disturbances.The results suggest that with a sample size of 1000–1400 cores and a properly chosen model, one reaches a certain precision in estimation that does not increase significantly with growing sample size. It appears that in temperate old-growth forests controlled by fine-scale disturbances, it is necessary to have at least 500 cores to estimate sill, nugget and relative nugget, while to estimate practical range at least 1000 cores are needed. When choosing the best model, the stability of the model should be considered together with the value of AIC. Our results indicate the general limits of disturbance spatial pattern studies using dendrochronological and geostatistical methods, which can be only partially overcome by sample size or sampling design.     

Temporal patterns in ambient noise of biological origin from a shallow water temperate reef

A systematic study of the ambient noise in the shallow coastal waters of north-eastern New Zealand shows large temporal variability in acoustic power levels between seasons, moon phase and the time of day. Ambient noise levels were highest during the new moon and the lowest during the full moon. Ambient noise levels were also significantly higher during summer and lower during winter. Bandpass filtering (700-2,000 Hz and 2-15 kHz), combined with snap counts and data from other studies show that the majority of the sound intensity increases could be attributed to two organisms: the sea urchin and the snapping shrimp. The increased intensity of biologically produced sound during dusk, new moon and summer could enhance the biological signature of a reef and transmit it further offshore. Ambient noise generated from the coast, especially reefs, has been implicated as playing a role in guiding pelagic post-larval fish and crustaceans to settlement habitats. Determining a causal link between temporal increases in ambient noise and higher rates of settlement of reef fish and crustaceans would provide support for the importance of ambient underwater sound in guiding the settlement of these organisms.     

Crossing the impassable: genetic connections in 20 reef fishes across the eastern Pacific barrier

The 'impassable' Eastern Pacific Barrier (EPB), ca 5000 km of deep water separating the eastern from the central Pacific, is the World's widest marine biogeographic barrier. Sequencing of mitochondrial DNA in 20 reef fish morphospecies encountered on both sides of the barrier revealed cryptic speciation in two. Among the other 18 species only two showed significant differentiation (as revealed by haplotype networks and FST statistics) between the eastern and the central Pacific. Coalescence analyses indicated that genetic similarity in the 18 truly transpacific species resulted from different combinations of ages of most recent invasion and of levels of recurrent gene flow, with estimated times of initial separation ranging from approximately 30000 to 1 Myr (ago). There is no suggestion of simultaneous interruptions of gene flow among the species. Migration across the EPB was previously thought to be exclusively eastward, but our evidence showed two invasions from east to west and eight cases in which subsequent gene flow possibly proceeded in the same direction. Thus, the EPB is sporadically permeable to propagules originating on either side.     

Ecological speciation in tropical reef fishes

The high biodiversity in tropical seas provides a long-standing challenge to allopatric speciation models. Physical barriers are few in the ocean and larval dispersal is often extensive, a combination that should reduce opportunities for speciation. Yet coral reefs are among the most species-rich habitats in the world, indicating evolutionary processes beyond conventional allopatry. In a survey of mtDNA sequences of five congeneric west Atlantic reef fishes (wrasses, genus Halichoeres) with similar dispersal potential, we observed phylogeographical patterns that contradict expectations of geographical isolation, and instead indicate a role for ecological speciation. In Halichoeres bivittatus and the species pair Halichoeres radiatus/brasiliensis, we observed strong partitions (3.4% and 2.3% divergence, respectively) between adjacent and ecologically distinct habitats, but high genetic connectivity between similar habitats separated by thousands of kilometres. This habitat partitioning is maintained even at a local scale where H. bivittatus lineages are segregated between cold- and warm-water habitats in both Bermuda and Florida. The concordance of evolutionary partitions with habitat types, rather than conventional biogeographical barriers, indicates parapatric ecological speciation, in which adaptation to alternative environmental conditions in adjacent locations overwhelms the homogenizing effect of dispersal. This mechanism can explain the long-standing enigma of high biodiversity in coral reef faunas.     

Conserved spatial patterns across the protein kinase family

Protein kinases are a large family of enzymes heavily involved in signal transduction, regulation of metabolism, and control of cell growth and differentiation. These functions require precise recognition of widely diverse signals and substrates, and very detailed control of protein kinase activity. Large molecules interact primarily through recognition of surface features. Comparison of surfaces is complicated by both sequence diversity and conformational variability, including multiple possible rotameric states of side chains. We used a recently developed method of protein surface comparison to compare different serine/threonine and tyrosine kinases. As we have shown, two hydrophobic cores inside a protein kinase molecule are connected by a unique formation, called the "spine". It exists only in the active conformation of protein kinases and is dynamically disassembled during the inactivation process. Detection of such structures by any other method was not possible as the residues which comprise the spine do not form any sequence or 3D motifs in a traditional sense.     

Habitat structure,conspecific presence and spatial variation in the recruitment of a temperate reef fish

Pronounced spatial variation in recruitment occurs in many marine invertebrate and fish populations and is thought to be critical to the demography of these species. In this study I examined the importance of habitat structure and the presence of conspecific residents to spatial variation in larval settlement and recruitment in a temperate fish Tautogolabrus adspersus. I define settlement as the movement of individuals from the water column to the benthic habitat, while I refer to recruitment as numbers of individuals surviving some arbitrary period of time after settlement. Experiments in which standard habitats were stocked with conspecifics showed that resident conspecifics were not an important factor contributing to small-scale variability in recruitment. Further correlative analyses demonstrated that large-scale variation in recruitment could not be explained by variability in older age classes. By contrast, manipulations of macroalgal structure within a kelp bed demonstrated that recruitment was significantly higher in habitats with a dense understory of foliose and filamentous algae than in habitats with only crustose algae. Understory algae varied in their pattern of disperison among sites, and the dispersion of fish matched that of the plants. In order to determine the effects of differences in patterns of algal dispersion on the demography of associated T. adspersus populations, I used experimental habitat units to manipulate patterns of dispersion. Settlement was significantly greater to randomly placed versus clumped habitats; however, no differences in recruitment between random and clumped habitats were detected. Because recruitment is a function of the numbers of settlers minus the subsequent loss of settlers, rates of mortality or migration must have been higher in the randomly placed habitats. These results are counter to the current paradigm for reef fishes which suggests that larval settlement is the crucial demographic process producing variability in population abundance. In this experiment patterns of settlement were modified by varying the patch structure of the habitat.Contribution number 278 from the Center for Marine Biology, University of New Hampshire     

Diel feeding migrations in tropical reef fishes

Summary 1. Many tropical reef fishes that feed during the day rest at night, whereas many that feed at night rest during the day. The feeding grounds of many are some distance from their resting grounds. Thus they migrate between these two locations during twilight as part of a general changeover between diurnal and nocturnal situations.2. At least many of these migrations are predictable, both as to time and to the route taken. The distances traveled vary between species, ranging from just a few meters, to more than several kilometers.3. The pattern of migrations is strongly influenced by the relative threat from predators at different periods of the diel cycle.4. During the day, migrations of reef fishes are limited to intra-reef movements: short vertical movements by certain plankton feeders, and lateral excursions from one part of the reef to another by certain herbivores and plankton feeders. Movements into the open regions that lie adjacent to many reefs are not adaptive in daylight due largely to a danger from predators.5. Despite constant threat from predators during the day, smaller reef fishes remain relatively secure during most of this period by staying close to shelter, or by schooling. However, these defenses are less effective during twilight, when the danger from predators intensifies. The diurnal migrators return to the shelter of their resting places prior to that part of evening twilight when danger is greatest, and the nocturnal migrators usually do not expose themselves for their nightly foraging until after the period of maximum danger has passed. During morning twilight the sequence is reversed.6. The major mechanisms whereby smaller reef fishes reduce predation during the day — schooling and staying close to shelter — are less evident at night. Not only do reef fishes range freely at night into the open regions that are avoided in daylight, but their schools are more loosely defined, and many are active as solitary individuals or in small groups. The tendency for looser associations and ranging farther afield increases on darker nights.7. Most predators that threaten reef fishes are visual feeders whose mode of attack loses effectiveness when light falls below a certain level. Although they operate to some extent under moonlight, they threaten small reef fishes less at night than during the day.8. In addition to whatever other ways a shcool may be adaptive, by reducing variable behavior among its members the school is especially important to migrating species. Responses to the various cues that mark the migration routes may be refined to within acceptable limits for the population as a whole only by coordinated group action.9. Submarine topographical features are important reference points for migrating reef fishes.

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Tägliche Futtersuchwanderungen bei tropischen Riffischen

Kurzfassung Dämmerungszeiten wandern viele tropische Fische zwischen ihren Ruheplätzen auf dem Riff und ihren Futterplätzen. Diese Wanderungen, die sowohl von tag- als auch von nachtaktiven Fischen durchgeführt werden, sind bedingt durch den Licht-Dunkel-Wechsel, und für viele Arten können sogar Zeit und Weg der Wanderung vorhergesagt werden. Das Muster der tagesrhythmischen Wanderungsbewegungen wird vor allem durch das Ausmaß von Bedrohungen durch Raubfeinde beeinflußt. Bewegungen in ungeschützte Gebiete, die vom Riff wegführen, sind am Tage nicht vorteilhaft. Bei Einbruch der Dunkelheit sind diese offenen Gebiete hingegen ein Konzentrationspunkt der Wanderungen aus dem Riff, wobei sich auch der Zusammenhalt der Fischschwärme verringert. Da Schwarmverhalten und Schutzsuchen die beiden wichtigsten Verteidigungseinrichtungen gegenüber Räubern darstellen, bedeutet das Aufgeben dieser Verhaltensweisen zu Beginn der Dämmerung, daß während der Nacht kleinere Riffische weniger bedroht sind als während des Tages. Für wandernde Schwarmfische dürfte eine geringe Variabilität der Verhaltensweisen von adaptiver Bedeutung sein. Nur durch ein koordiniertes Schwarmverhalten ist es offensichtlich möglich, innerhalb bestimmter Grenzen auf die verschieden optischen Marken, welche die Wanderwege kennzeichnen, zu reagieren. Vermutlich spielen die topographischen Gegebenheiten unter Wasser eine entscheidende Rolle für die Orientierung der Riffische.