Latitudinal variation in abundance of herbivorous fishes: a comparison of temperate and tropical reefs

The aim of the study was to provide comparable estimates of abundance of herbivorous reef fishes at temperate and tropical localities using a standardized methodology. Faunas of herbivorous fish were sampled on the rocky reefs of temperate northern New Zealand and on the coral reefs of the northern Great Barrier Reef (GBR), Australia, and the San Blas Archipelago in the Caribbean. A pilot study established the most appropriate habitat setting and the scale and magnitude of replication for the sampling program in temperate waters. Herbivorous fishes, including members of families endemic to the southern hemisphere (Odacidae and Aplodactylidae), were most abundant in turbulent, shallow water (0 to 6 m) and had patchy distributions within this habitat. A hierarchical sampling program using 10-min transect counts within the 0 to 6 m depth stratum examined abundance patterns at a range of spatial scales including mainland and island coasts, localities separated by up to 100 km and sites separated by up to 10 km. This program identified a characteristic fauna of seven species of herbivorous fishes with mean total abundances ranging from 23 to 30 individuals per 10-min transect. Species composition of the fauna varied between islands and coasts. A similar methodology was used to sample the major families of herbivorous fish in a number of sites in each of the tropical regions. These sampling programs revealed a fauna dominated by acanthurids and scarids in both the GBR and Caribbean localities. Estimates of abundance from these regions were similar, with a mean of 108 individuals recorded on the GBR and 129 per 10-min transect in the Caribbean. Species richness varied between each region, with 44 taxa recorded from the GBR and 11 from the Caribbean. Abundances of temperate water herbivores in New Zealand were found to be 75 to 80% lower than those recorded from shallow water habitats sampled on coral reefs. This was not related to species richness, since both New Zealand and the Caribbean locality had patterns of low richness. We suggest that the differences in abundance found by our study between temperate and tropical regions are not restricted to herbivorous fishes, but are representative of general latitudinal trends in reef fish faunas. Received: 4 November 1996 / Accepted: 15 December 1996     

Exudation of dissolved organic carbon by brown algae

The release via exudation of dissolved organic carbon by Ascophyllum nodosum (L.) Le Jol. was investigated under laboratory conditions. In contrast to results previously reported by Sieburth (1969), continuous release of organic matter was found with only 1 specimen out of 21 tested either in stagnant sea water or in sea water flowing through the system. The holdfast of the plants was found to release relatively large amounts of organic material; however, this release lasted only for short periods, and was most probably due to damage caused by removing the plants from their substrate. The possible reasons for the failure to detect exudation are discussed. With Fucus vesiculosus L., no reliable results could be obtained, most probably due to particulate matter sloughing off the plants during the washings and during the experiments under conditions of photosynthesis.     

The trophic status of herbivorous fishes on coral reefs

Dietary analyses of 17 species of nominally herbivorous fishes on the northern Great Barrier Reef were carried out to resolve the questions to what extent diet in nominally herbivorous reef fishes is dominated by living plant material and whether short-chain fatty acid profiles in the alimentary tract predict diet. The fishes included members of the families Acanthuridae, Scaridae and Kyphosidae. The analyses revealed consumption of a wide range of food resources including macroscopic algae, turfing algae, pelagic and planktonic animal matter, organic detritus and sediments. In only 7 of the 17 species was living algae the dominant dietary item. Gastrointestinal short-chain fatty acids (SCFA) profiles reflecting the fermentative activities of gut symbionts were used as a framework for the initial analysis of dietary patterns. The feeding patterns implied by the SCFA profiles were validated by direct observation of food material through gut content analysis. We identified four main trophic groupings differentiated by fermentation profiles and diet. These were: (1) species with low levels of SCFA in gut samples (mean mM 9.2ǂ.8) but a high proportion of isovalerate (mean 15.4%dž.7): Acanthurus nigricauda, A. olivaceus, Ctenochaetus striatus, Chlorurus microrhinos, C.sordidus, Scarus schlegeli; (2) species that fed on turfing, filamentous and small thallate algae and displayed moderate levels of SCFA (mean mM 21.9DŽ.9): Acanthurus nigricans, A.lineatus, Naso tuberosus; (3) species that fed on large thallate (macroscopic) algae with high levels of SCFA (mean mM 41.2Dž.5) with very low proportions of isovalerate (mean 0.5%ǂ.1): Kyphosus cinerascens, K.vaigiensis, Naso unicornis, Zebrasoma scopas; and (4) species with diets dominated by planktonic animal material with moderate levels of SCFA (mean mM 21.3ǃ.5): Naso annulatus, N. brevirostris, N. hexacanthus, N. vlamingii. We conclude that a wide range of dietary items are taken by the 17 nominal herbivores, and that dietary groupings do not reflect taxonomic relationships. There was evidence of convergence in feeding modes and diet between phylogenetically distinct taxa and divergence within particular lineages.     

Diversity and stability of herbivorous fishes on coral reefs

Biodiversity may provide insurance against ecosystem collapse by stabilizing assemblages that perform particular ecological functions (the "portfolio effect"). However, the extent to which this occurs in nature and the importance of different mechanisms that generate portfolio effects remain controversial. On coral reefs, herbivory helps maintain coral dominated states, so volatility in levels of herbivory has important implications for reef ecosystems. Here, we used an extensive time series of abundances on 35 reefs of the Great Barrier Reef of Australia to quantify the strength of the portfolio effect for herbivorous fishes. Then, we disentangled the contributions of two mechanisms that underlie it (compensatory interactions and differential responses to environmental fluctuations ["response diversity"]) by fitting a community-dynamic model that explicitly includes terms for both mechanisms. We found that portfolio effects operate strongly in herbivorous fishes, as shown by nearly independent fluctuations in abundances over time. Moreover, we found strong evidence for high response diversity, with nearly independent responses to environmental fluctuations. In contrast, we found little evidence that the portfolio effect in this system was enhanced by compensatory ecological interactions. Our results show that portfolio effects are driven principally by response diversity for herbivorous fishes on coral reefs. We conclude that portfolio effects can be very strong in nature and that, for coral reefs in particular, response diversity may help maintain herbivory above the threshold levels that trigger regime shifts.     

The trophic status of herbivorous fishes on coral reefs

Estimates of feeding rates, alimentary tract structure and temporal patterns of food processing obtained from twelve species of nominally herbivorous fishes on the northern Great Barrier Reef were compared. These included members of the families Acanthuridae, Scaridae and Kyphosidae. Based on an analysis of diet and short-chain fatty acid (SCFA) profiles from a previous study we initially partitioned the twelve species into four dietary categories, as follows: (a) Category 1: herbivores with a diet of macroscopic brown algae and high SCFA profiles in the hindgut region (Naso unicornis, Kyphosus vaigiensis); (b) Category 2: herbivores feeding on turfing and filamentous red and green algae with moderate SCFA profiles in the hindgut region (N. tonganus, K. cinerascens, Zebrasoma scopas, Acanthurus lineatus); (c) Category 3: zooplankton feeders with moderate SCFA profiles (N. vlamingii, N. brevirostris); (d) Category 4: species feeding on detrital and sedimentary materials with low levels of SCFA (Chlorurus microrhinos, Scarus schlegeli, Ctenochaetus striatus, A. olivaceus). The purpose of this comparison was to determine whether measures of feeding activity, alimentary tract structure, and food processing were concordant with diet. A dichotomy in feeding rates was observed. Species with a diet of algae and zooplankton (categories 1–3) had slower feeding rates than those feeding on detrital aggregates and sediment (category 4). The pattern of food processing also followed the same dichotomy with species of categories 1–3 retaining food in the alimentary tract overnight and commencing the feeding day with substantial amounts of food in the intestine and hindgut. Category-4 species commenced the feeding day with empty alimentary tracts suggesting a rapid turnover of gut contents. Within the herbivorous and zooplankton-feeding species neither alimentary tract structure nor food processing mode were predicted by diet or SCFA profiles. A hindgut fermentation chamber was present in K. vaigiensis but not in N. unicornis, a species with high levels of SCFA in the hindgut region and a diet of brown macroscopic algae. In contrast N. vlamingii, with a diet dominated by animal matter, retained large amounts of food material in a hindgut chamber over the entire feeding cycle. In tropical perciform fishes, herbivory and fermentation are not associated with the alimentary tract structures that characterise herbivorous terrestrial vertebrates. Estimates of the abundance of the different groupings of nominally herbivorous fishes indicated that the dominant elements in the reef grazing and browsing fauna were consumers of detrital and sedimentary materials. These could not be classified as herbivores. Members of this group were dominant in all habitats investigated. Explicitly herbivorous taxa were a minority component in all habitats investigated.Communicated by G.F. Humphrey, Sydney     

Geographical gradients of marine herbivorous fishes: patterns and processes

We present new data and the first rigorous analysis of latitudinal and thermal gradients of diversity, density and biomass of marine herbivorous fishes and review proposed explanatory mechanisms. Consistently, negative relationships between latitude, and positive relationships between sea surface temperature (SST), and relative richness and relative abundance of herbivorous fishes were found worldwide. Significant differences in the strength of gradients of richness and abundance with latitude and SST between tropical and extratropical zones were found consistently across ocean basins. Standardized sampling along the western Atlantic also showed negative relationships between latitude and total density and biomass. The trends, however, are driven by different components of the fish assemblages (i.e., scarids in the Caribbean and acanthurids in Brazil). Patterns of abundance along thermal gradients, generally associated with extensive latitudinal gradients, also were found at the local scale. Feeding rate of the ocean surgeonfish Acanthurus bahianus decreases with temperature more rapidly than the mean metabolic rate of teleost fishes. This relationship suggests a temperature-related physiological constraint. From the new standardized and comparative data presented and the review of the explanatory hypotheses, we conclude that temperature-related feeding and digestive processes are most likely involved in the distribution patterns of herbivorous fishes. Electronic Supplementary Material Supplementary material is available for this article at     

The distribution of herbivorous fishes on the Great Barrier Reef

The composition and functionality of ecologically important herbivorous fish assemblages were examined throughout much of Australia’s Great Barrier Reef (GBR). Diversity and abundance of surgeonfishes (Acanthuridae), parrotfishes (Labridae) and rabbitfishes (Siganidae) were strongly associated with position on the continental shelf, whilst effects of latitude were weaker and inconsistent. Species distributions varied considerably amongst taxonomic groups; parrotfishes were mostly widespread whilst distributions of surgeonfishes were often restricted. Most inshore environments supported depauperate herbivore assemblages dominated by different taxa and functional groups compared with assemblages in offshore environments. There were also strong cross-shelf transitions in the main taxa performing each functional role. Overall, this study show that the functional contributions of herbivorous fish assemblages to important ecosystem processes and the contributing taxa vary considerably amongst different GBR environments. Additionally, the two most numerically dominant herbivores actively select detritus, not algae, supporting increasing evidence for the importance of detritus in coral reef ecology.     

Association of color and feeding deterrence by tropical reef fishes

Summary. While many marine molluscs have been suggested to use aposematic coloration to avoid predation, few studies have tested the ability of marine predators to learn to associate colors with distasteful prey. In field experiments, we tested the ability of two populations of reef fishes to discriminate among red, yellow, and black artificial nudibranch models when one color was paired with a feeding deterrent. We offered fishes (1) the models without any feeding deterrents, (2) the models with a feeding deterrent coated onto one color, and (3) the models without deterrents again. If reef fishes learn to associate colors with noxious prey, we expected the color paired with the feeding deterrent to be eaten less frequently in the final assay than the initial assay. In both populations, fishes formed clear associations between color and feeding deterrence. However, when the experiment was repeated in one population, changing the color paired with the feeding deterrent, fishes did not form an association between color and feeding deterrence. In this case, prior learning may have affected subsequent trials. Our study indicates that common colors of nudibranchs are recognizable by fishes and can be associated with noxious prey. Received 24 September 1998; accepted 18 December 1998.     

A re-evaluation of the diel feeding hypothesis for marine herbivorous fishes

We re-evaluated the "diel feeding hypothesis" by measuring diel variation in starch, protein, and floridoside in three algal "types" collected from a fringing coral reef at Lizard Island, Great Barrier Reef, Australia. Samples of two species of rhodophyte algae, Gracilaria arcuata and Acanthophora spicifera, and the turf assemblage from the territories of the herbivorous pomacentrid Stegastes nigricans were collected at four time periods through the day: 0630-0730, 1000-1100, 1330-1430, and 1630-1730 hours. We also measured the ability of several species of marine fish (the herbivores Acanthurus nigricans, A. lineatus, A. olivaceus, and Parma alboscapularis and the detritivore Ctenochaetus striatus) to hydrolyse floridoside by estimating !-galactosidase activity in tissue from the anterior intestine. We detected no diel pattern in protein content of the algae but found a significant steady increase in starch content throughout the day. Floridoside content increased in the morning and decreased in the afternoon, a pattern that may be driven by midday photoinhibition of the algae. All the fishes tested could utilise floridoside. Our results support the diel feeding hypothesis. Although floridoside content decreased in the afternoon, our results suggest floridoside was used during the day by the algae to synthesise starch. Thus the algae increased in nutritional value until photoinhibition occurred at midday then subsequently maintained their nutritional value during the afternoon. This pattern of algal nutrients increasing to a midday peak and remaining relatively constant throughout the afternoon correlates well with the diel feeding pattern in many species of marine herbivorous fish.     

Nutritional quality of macrophytes eaten and not eaten by two temperatezone herbivorous fishes: A multivariate comparison

The mean annual chemical compositions (ash, lipid, carbohydrate, protein, nitrogen and carbon) of 23 species of macrophytes (22 seaweeds and 1 seagrass) from a rocky intertidal habitat on the central California coast were determined from December 1981 through December 1982. These data were used to test the hypothesis that the 13 red and green seaweeds eaten by the two principal herbivorous fishes (Cebidichthys violaceus andXiphister mucosus) at the site are higher in nutritional quality than the 9 red and brown seaweeds and the seagrass not consumed by these fishes. A MANOVA using the ash, lipid, carbohydrate and nitrogen data showed that the centroids of the dietary and nondietary species groups were significantly different. In a two-group discriminant analysis that followed, only two species were misclassified as members of the opposite group. Multigroup discriminant analysis of the 23 macrophytes resulted in some overlap among dietary and nondietary species. Species were discriminated on the first canonical axis by ash content and on the second and third axes by lipid and carbohydrate contents. Nitrogen contributed little to the overall discrimination of species in the analysis. The nondietary red algaCorallina vancouveriensis, with its high ash content and therefore relatively low nutritional quality, was clearly separated from all other species in the analysis. Brown algae were of higher nutritional quality, but are not eaten by the two fishes, possibly because these seaweeds produce indigestible carbohydrates and secondary compounds. However, the exclusion from the diets of several red algal species that were virtually indistinguishable from the dietary red algae remains unexplained.Contribution No. 64 from the Ocean Studies InstitutePlease address all correspondence and requests for reprints to Dr. Horn at Fullerton     

Endosymbiotic communities of two herbivorous labroid fishes,Odax cyanomelas andO. pullus

Two herbivorous species of the temperate labroid fish family Odacidae were examined for the presence of gut endosymbionts.Odax cyanomelas (Richardson) from southern Australia andO. pullus Schneider from New Zealand both feed on macroalgae, yet appear to lack obvious morphological specializations for herbivory. Specimens ofO. cyanomelas andO. pullus were collected in 1989 from Sydney, Australia, and Leigh, New Zealand, respectively. Both species were found to harbour dense concentrations of prokaryotic and eukaryotic microbes in their lower intestines. The various cell types present were examined by light microscopy, transmission electron microscopy and scanning electron microscopy. Epifluorescence microscope counts were used to quantify the distribution and abundance of the microbiota along the gut ofO. cyanomelas. Major differences were observed in the composition of microbiota between the two species.O. cyanomelas contained spirilla, large rod-shaped bacteria, filamentous bacteria and two forms of trichomonad flagellates.O. pullus also harboured dense, large, rod-shaped bacterial populations, but lacked the other two large prokaryote categories found inO. cyanomelas, and contained diplomonad flagellates. The large rod-shaped bacteria found in both species resembled prokaryotes described from other herbivorous fish and termites.     

Comparative study of metamorphosis in tropical reef fishes

This study explores the types of changes in pigmentation and morphology that occur immediately after settlement in 13 families of tropical reef fishes encompassing 34 species. The morphology of individual fishes was recorded daily from when they were first caught at night as they came into the vicinity of a reef to settle. Changes in pigmentation and morphology were species specific and often varied greatly among species within a family or genus. Pigmentation changes were typically rapid (<36 h) and dramatic. Morphological changes involved the elongation and regression of fin spines and changes in head shape and body depth. Eighteen percent of species experienced changes in snout shape and dorsal spine length of greater than 5%. Similarly, 15% experienced changes in pectoral fin length and head length of greater than 5%. Changes typically occurred gradually over 6 or more days, although in about 44% of the species the major change in one of the measured body dimensions occurred rapidly (within 36 h). Moderately strong positive relationships were found between both growth and developmental rates and the extent of metamorphosis in the damselfishes (Pomacentridae) (r=0.48 and 0.63, respectively). This suggests there may be a minimum level of development necessary to be a fully functional demersal juvenile. Although many of the changes that occur are subtle compared to the preceding development, these changes occur at an important ecological transition. Published online: 16 August 2002     

Quantitative distribution of herbivorous reef fishes in the Gulf of Aqaba (Red Sea)

The distribution of the main herbivorous fishes (Acanthuridae, Scaridae, Siganidae) was studied across a coral reef of the Jordanian coast in the Gulf of Aqaba (Red Sea). Visual counts were realized by diving along transects (200 m long and 5 m wide), parallel to the shore, at 10 stations located from the lagoon to 40 m deep on the outer reef slope. Herbivorous reef fishes reach their highest abundance on the reef front, where 234 fishes were counted per 1,000 m². Their density decreases on the reef flat, with an average of 150 fish 1,000 m^-2, and is lowest on the outer reef slope (69 fish 1,000 m^-2). Surgeonfishes form 63% of the herbivorous ichthyofauna, parrotfishes 35%, and rabbitfishes 2%. Families and species display different distributions according to biota. The Acanthuridae dominate on the reef flat, whereas the Scaridae are more numerous on the outer reef slope. The evolution of the social structure of the main species was observed: the adults generally school in the lagoon and on the reef flat, but are mainly solitary on the reef slope. The distribution of juvenile individuals is more restricted: they are concentrated on the reef front and on the upper part of the reef slope.This study is part of a cooperation programme between the University of Nice (France) and the University of Jordan, to study the ecology of the coral reefs and the surrounding waters of the Jordanian coast (Gulf of Aqaba, Red Sea)     

Microstructure of settlement-marks in the otoliths of tropical reef fishes

The morphology and ultrastructure of the otolith settlement-mark was examined in 44 tropical reef-fish species spanning nine families. A classification scheme based on similar otolith characteristics is presented. Three major categories are identified based on changes in increment width and optical qualities of the settlement-mark. Of the 44 species examined, 39 possessed “abrupt” settlement-marks (Type I) characterised by a rapid decrease in increment width (up to 50% reduction) over settlement. Type I settlement-marks were found in all nine families examined. The 39 species spanned the whole range of possible larval durations (Pomacentrus moluccensis, 15 d ± 0 SE; Naso hexacanthus, 91.2 d ± 2.97 SE). Four of the 44 species possessed “zonal” settlement-marks (Type II), featuring a band of increments that are wider than pre-settlement increments. Species in this category are the labrids Corisaygula, Thalassoma bifasciatum, T. lunare and an unidentified acanthurid (Acanthurus sp. 2). One species of acanthurid (N. brevirostris) possessed a “gradual” settlement-mark (Type III), manifest as a gradual decrease in increment width during the settlement period. A possible fourth type was identified from the literature. Gnatholepis thompsoni and Coryphopterus glaucofraenum possessed a settlement-mark with increment widths that increased post-settlement. Available data suggest a poor relationship between the structure of the settlement-mark and the magnitude of metamorphosis (previously reported as internal and external morphological change). Evidence suggests that the increment profile over early development and the increment transitions associated with the settlement event are taxon-specific and may enable late-larval stage fishes to be identified to species level. Received: 21 May 1997 / Accepted: 3 February 1999     

Fish guts as chemical reactors: a model of the alimentary canals of marine herbivorous fishes

The alimentary canals of marine herbivorous fishes are variously composed of a few basic structures strung together in series. We model the structures where digestive processes occur as chemical reactors following the model of Penry and Jumars: the stomach as a batch or continuous-flow stirred-tank (CSTR), the intestine as a plug-flow reactor (PFR), and the hindgut caecum as a CSTR. Other structures, where food is mechanically processed-gill rakers, pharyngeal mills, and muscular stomachs-are classified as gates. The optimality criterion for the model is the digestion of the most nutrient in the least amount of time. With the model we are able to predict gut configuration as a function of nutrient concentration and hypothesize that the guts of herbivorous fishes always have a PFR component and may or may not have a CSTR component. The Penry-Jumars model appears to provide a consistent theoretical framework for four main types of digestive mechanisms in marine herbivorous fishes and offers specific testable hypotheses on the feeding ecology and digestive physiology of four representative species of fish as well as other, still unstudied, herbivorous fishes.Contribution No. 68 from the Ocean Studies Institute, Long Beach, California 90840, USA     

Clostridia dominate 16S rRNA gene libraries prepared from the hindgut of temperate marine herbivorous fishes

Bacterial diversity in the microbial communities of posterior gut sections of three temperate marine herbivorous fish species from New Zealand was characterised using Amplified Ribosomal DNA Restriction Analysis, and 16S rRNA gene amplification and sequencing methods. The fish were collected in 1999–2000 in the Hauraki Gulf, New Zealand (35°54’–36°24’S, 174°48’–175°25’E). The gastrointestinal bacterial communities of Kyphosus sydneyanus (Günther, 1886) (F. Kyphosidae), Odax pullus (Forster in Bloch and Schneider, 1801) (F. Labridae) and Aplodactylus arctidens Richardson, 1839 (F. Aplodactylidae) were dominated by five clades of bacteria, four of which belong to recognized clostridial clusters. The clone libraries of K. sydneyanus and O. pullus contained sequences from most of these clades, but were dominated by members of clostridial clusters XI and XIVa, respectively. The clone library of A. arctidens was dominated by members of clostridial cluster XIVb and an unassigned cluster containing Eubacterium desmolans and Papillibacter cinnaminovorans. The finding that strains of Firmicutes dominated the gastrointestinal microbial communities of all three fish species is consistent with the results of similar studies on terrestrial vertebrate herbivores. This work thus contributes to the view that gastrointestinal symbionts in some marine herbivorous fishes may play a similar role to those in terrestrial vertebrate herbivores studied to date.     

Preparative and analytical extraction of pigments from brown algae with dimethyl sulfoxide

Dimethyl sulfoxide (DMSO) rapidly extracts much of the chlorophyll c and fucoxanthin from the intact thalli of brown algae. Subsequent extraction by acetone rapidly removes most of the chlorophyll a and -carotene. An analytical procedure for these pigments has been developed, based upon this extraction sequence, followed by partition of the acetone extract between hexane and aqueous acetone. Fucoxanthin and chlorophyll c can be recovered from the DMSO extract and the aqueous acetone phase by extraction into ethyl acetate and subsequent evaporation. The DMSO-acetone extraction sequence is also effective with green algae and a few red algae, although the full scope of the method has not been investigated.     

Distribution of algae on tropical rocky shores: spatial and temporal patterns of non-coralline encrusting algae in Hong Kong

Encrusting algae have been described as dominant space occupying species on rocky shores around the world. Despite their abundance, however, most studies classify species under generic names (e.g. Ralfsia sp.) or as a functional group (e.g. encrusting algae), thereby underestimating the number of species present and their ecological importance. Studies on six rocky shores of varying exposure, in Hong Kong, recorded eight common species of encrusting algae. The greatest abundance of encrusting algae was recorded on shores of intermediate exposure, where four distinct zonation bands could be identified; a cyanobacterial Kyrtuthrix-Zone in the upper midlittoral, a Bare-Zone below this, a Mixed-Zone in the lower midlittoral and a Coralline-Zone in the infralittoral fringe. Abundance declined on shores of greater and lower exposure to wave action, where bivalves and barnacles were competitively dominant. Certain species were found in greater abundance on exposed shores (e.g. Dermocarpa sp. and Hildenbrandia occidentalis), while others preferred more sheltered shores (e.g. Hildenbrandia prototypus and Kyrtuthrix maculans). With the exception of some cyanobacterial crusts, the abundance of encrusting algae was always greatest towards the low shore, an area of decreased physical stress and increased herbivore density. Zonation patterns showed seasonal variation associated with the monsoonal climate of Hong Kong. Most species increased in abundance during the cool season, while during the summer months the cover and vertical extent of encrusting algae decreased in relation to summer temperatures, although K. maculans increased in abundance during the summer. On Hong Kong shores, encrusting algae have a high species richness and exhibit within-functional group spatial and temporal variation which is mediated by herbivory and seasonal, physical stresses.     

Temperature tolerance and latitudinal range of brown algae from temperate Pacific South America

Lower and upper survival temperatures of microthalli of 25 species of South American Phaeophyceae isolated from central Peru (14°S) to the Canal Beagle (55°S) were determined using 2-wk exposure for the upper and 4-wk exposure for the lower limit. All species survive 4 wks at -2°C. With respect to the upper limit, species reported only from southernmost South America tolerate 19.9 to 24.5°C (n=8), and species occurring from Cape Horn to central Chile 24.6 to 27.4°C (n=7). Three species that occurred as far north as northern Chile and Peru before the 1982–1983 El Niño event, and whose northern limit was dramatically shifted southwards in 1983, tolerate 20.8 to 25.3°C, whereas five species that have survived in Peru tolerate 25.6 to 28.5°C. Tinocladia falklandica which tolerates 27.8 to 28.1°C but lives only in southernmost South America and Striaria attenuata, which tolerates 31.6 to 31.9°C but occurs at ca. 42°S, are exceptional. Their high temperature tolerance may have no adaptive value in South America. They are restricted to the cold-temperate region due to low temperature requirements for reproduction or for reasons yet unknown. In general, the northern distributional limits of the Phaeophyceae studied along the temperate Pacific coast of South America are reproduction boundaries, except in El Niño years when they are redefined according to the species' upper suvival limits. Temperature tolerance of isolates from northern Chile and Peru agrees well with maximum temperatures reached during the 1983 El Niño.