Intertidal Fish Species Research Articles

Shallow water fish display low phenotypic plasticity to ocean warming and extreme weather events regardless of previous thermal history

Shallow water environments have long been recognized by scientists as sentinels for climate change. By analysing the impacts of ocean warming and marine heatwaves (MHW) in species from these areas, we can estimate their plasticity and hence vulnerability to thermal challenges. Pomatoschistus microps is a benthic intertidal fish species inhabiting coastal lagoons where temperature fluctuations are common. Here, we tested the effects of “Present” and “Future summer” scenarios (22 °C and 25 °C) and their respective heatwaves (27 °C and 30 °C) versus a “Control” scenario of 19 °C on warm (summer)- and cold (winter)- acclimatized fish. Then, we estimated phenotypic plasticity of critical thermal maximum (CTmax), oxygen consumption and cellular stress responses (CSR). Temperature seasonal variation and body weight (as proxy for body size) effects on fish thermal tolerance were also determined. Fish exposed to higher temperature treatments exhibited higher thermal tolerance, with this pattern being consistent for both warm- and cold-acclimatized fish. However, this difference was subtle (<4.6 %), suggesting a low capacity for acclimation. Nonetheless, warm-acclimatized fish (collected in summer) displayed significantly higher CTmax than cold-acclimatized fish (collected in winter), indicating that CTmax is influenced by seasonal thermal variation. Weight also represents a constraint factor for P. microps thermal tolerance, as heavier animals displayed lower CTmax. No alterations in O2 consumption, neither in CSR biomarkers were detected across temperature treatments, suggesting that fish were otherwise relatively insensitive to thermal fluctuations, independently of thermal history, within the thermal scenarios tested. Overall, the studied population of P. microps seems well adapted to temperature variations in their natural environment, exhibiting a large thermal safety margin (average of 11.02 °C).

Lineage-level species distribution model to assess the impact of climate change on the habitat suitability of Boleophthalmus pectinirostris

Global climate change has profound impacts on the habitats of marine organisms, and predicting the habitat changes of species under climate change conditions is crucial for species sustainability. Boleophthalmus pectinirostris is an intertidal fish species that holds significant ecological and economic value. To better protect and manage its resources, this study aimed to predict its current potential distribution and habitat changes under different climate scenarios in the future. This study firstly quantified the hypervolume niches of the three lineages (AE1, AE2, and AES lineages) and compared the niche differentiation among them. Furthermore, this study constructed species-level and lineage-level species distribution models (SDMs) to assess the impact of climate change on the habitat suitability of B. pectinirostris. The result of the niche differentiation assessment showed that there was marked differentiation in niches among the three lineages. The responses of different lineages to environmental variables were different, suggesting that lineage-level models may provide more accurate prediction results. According to the model predictions, the AES may have greater resilience to climate change and may experience habitat expansion in the future, while the AE1 and the AE2 may face habitat loss in some regions. Climate change-driven shifts in oceanic conditions were anticipated to affect the distribution and community structure of marine organisms. This study assessed the impact of climate change on the suitable habitat range of three lineages of B. pectinirostris using SDMs. Consistent with previous studies, the results of our study indicated that lineage-level SDMs may be more reliable than species-level SDMs for species with population differentiation in terms of the accuracy of predictions. In addition, considering the vulnerability of the AE1 and AE2 lineages to climate change, conserving these two lineages should be given a higher priority. The results of this study will provide important information for the future management and conservation of this species.

Microhabitat partitioning between sympatric intertidal fish species highlights the importance of sediment composition in gravel beach conservation.

Gravel beaches in the Mediterranean ecoregion represent an economically important and unique habitat type. Yet, burgeoning tourism, intensive coastal development and artificial nourishment of beaches may jeopardize their ecological communities. To date, species that reside on gravel beaches and the consequences of beach alterations are poorly understood, which hampers the development of a sustainable coastal tourism industry along the region's shorelines. Using a simple collection method based on dredging buckets through the intertidal section of beaches, we quantified the microhabitat association of two sympatric clingfish species in the genus Gouania at seven natural and an artificial gravel beach based on sediment characteristics. We hypothesized that slender (G. pigra) and stout (G. adriatica) morphotypes would partition interstitial niche space based on sediment size, which may affect the vulnerability of the species to changes in gravel beach composition due to coastal development. We detected substantial differences in gravel composition within and among the sampled beaches which suggests scope for microhabitat partitioning in Gouania. Indeed, we found significant relationships between species identity and the presence/absence and abundance of individuals in hauls based on their positioning on PC1. Our results suggest that modifications of gravel beaches through coastal development, including beach nourishment, intensifying coastal erosion, or artificial beach creation, may have detrimental consequences for the two species if sediment types or sizes are altered. We posit that, given the simplicity and efficacy of our sampling method and the sensitivity of Gouania species to prevailing gravel composition, the genus could serve as an important indicator for gravel beach management in the Mediterranean ecoregion.

Out to sea: ocean currents and patterns of asymmetric gene flow in an intertidal fish species.

Passive dispersal via wind or ocean currents can drive asymmetric gene flow, which influences patterns of genetic variation and the capacity of populations to evolve in response to environmental change. The mangrove rivulus fish (Kryptolebias marmoratus), hereafter "rivulus," is an intertidal fish species restricted to the highly fragmented New World mangrove forests of Central America, the Caribbean, the Bahamas, and Florida. Mangrove patches are biological islands with dramatic differences in both abiotic and biotic conditions compared to adjacent habitat. Over 1,000 individual rivulus across 17 populations throughout its range were genotyped at 32 highly polymorphic microsatellites. Range-wide population genetic structure was evaluated with five complementary approaches that found eight distinct population clusters. However, an analysis of molecular variance indicated significant population genetic structure among regions, populations within regions, sampling locations within populations, and individuals within sampling locations, indicating that rivulus has both broad- and fine-scale genetic differentiation. Integrating range-wide genetic data with biophysical modeling based on 10years of ocean current data showed that ocean currents and the distance between populations over water drive gene flow patterns on broad scales. Directional migration estimates suggested some significant asymmetries in gene flow that also were mediated by ocean currents and distance. Specifically, populations in the center of the range (Florida Keys) were identified as sinks that received migrants (and alleles) from other populations but failed to export individuals. These populations thus harbor genetic variation, perhaps even from extirpated populations across the range, but ocean currents and complex arrangements of landmasses might prevent the distribution of that genetic variation elsewhere. Hence, the inherent asymmetry of ocean currents shown to impact both genetic differentiation and directional migration rates may be responsible for the complex distribution of genetic variation across the range and observed patterns of metapopulation structure.

Site fidelity of fish on a rocky intertidal in the south of Portugal

The short-term site fidelity of intertidal fish species was studied in a rocky intertidal zone in southern Portugal using visible implant elastomer tagging. Eleven fish species were caught, tagged and released, seven of which were recaptured. The larger tide pool with the highest complexity level and located at a lower position in the shore level did not have recaptures, while in others the individual fish species recaptured rate ranged from 11.1 to 38.9%, suggesting that the shore height is a dominant factor influencing site fidelity. Although the breeding season seems not to be the only explanation for the recapture rate, the higher mean recapture rates of Parablennius sanguinolentus, Lipophrys pholis and Gobius cobitis may be related to the parental behaviour of these species during that period. Site fidelity of juveniles of Scorpaena porcus in tidepools is reported for the first time, showing the importance of this habitat as a nursery ground.

Metal and metalloid concentrations in the southern African endemic inter- and infratidal super klipfish, Clinus superciliosus, from the west and south coasts of South Africa

Clinus superciliosus was collected for element analysis from six localities along the South African west and south coasts. Concentrations in muscle and liver were determined, considering size and sex. No significant positive correlation between size and concentrations were detected, except for Mn and Sn in liver from Simons Town marina, while no significant differences in sex were detected. The majority of element concentrations were significantly higher in fish from Tsitsikamma in the Garden Route and the small town Chintsa, while some concentrations were significantly higher in muscle at Simons Town marina. Land-use activities had a limited role in element bioaccumulation in klipfish. Element concentrations were influenced by large scale oceanographic processes (currents; upwelling) and localised seasonal geogenic derived run-off. Limited data on element accumulation patterns of intertidal fish species in South Africa, highlights the need for long-term monitoring and further studies on different resident and transient intertidal fish species.

Freshening effect on the osmotic response of the Antarctic spiny plunderfish Harpagifer antarcticus.

Global warming is having a significant impact around the world, modifying environmental conditions in many areas, including in zones that have been thermally stable for thousands of years, such as Antarctica. Stenothermal sedentary intertidal fish species may suffer due to warming, notably if this causes water freshening from increased freshwater inputs. Acute decreases in salinity, from 33 down to 5, were used to assess osmotic responses to environmental salinity fluctuations in Antarctic spiny plunderfish Harpagifer antarcticus, in particular to evaluate if H. antarcticus is able to cope with freshening and to describe osmoregulatory responses at different levels (haematological variables, muscle water content, gene expression, NKA activity). H. antarcticus were acclimated to a range of salinities (33 as control, 20, 15, 10 and 5) for 1 week. At 5, plasma osmolality and calcium concentration were both at their lowest, while plasma cortisol and percentage muscle water content were at their highest. At the same salinity, gill and intestine Na+ -K+ -ATPase (NKA) activities were at their lowest and highest, respectively. In kidney, NKA activity was highest at intermediate salinities (15 and 10). The salinity-dependent NKA mRNA expression patterns differed depending on the tissue. Marked changes were also observed in the expression of genes coding membrane proteins associated with ion and water transport, such as NKCC2, CFTR and AQP8, and in the expression of mRNA for the regulatory hormone prolactin (PRL) and its receptor (PRLr). Our results demonstrate that freshening causes osmotic imbalances in H. antarcticus, apparently due to reduced capacity of both transport and regulatory mechanisms of key organs to maintain homeostasis. This has implications for fish species that have evolved in stable environmental conditions in the Antarctic, now threatened by climate change.

Acute high temperature exposure impairs hypoxia tolerance in an intertidal fish.

Acute heat shock has previously been shown to improve subsequent low O2 (hypoxia) tolerance in an intertidal fish species, a process known as cross-tolerance, but it is not known whether this is a widespread phenomenon. This study examined whether a rock pool specialist, the triplefin fish Bellapiscis medius, exhibits heat shock induced cross-tolerance to hypoxia, i.e., longer time to loss of equilibrium (LOE) and lower critical O2 saturation (Scrit) after recovering from an acute heat challenge. Non-heat shock controls had a median time to loss of equilibrium (LOE50) of 54.4 min under severe hypoxia (7% of air saturation) and a Scrit of 15.8% air saturation. Contrary to expectations, however, treatments that received an 8 or 10°C heat shock showed a significantly shorter LOE50 in hypoxia (+8°C = 41.5 min; +10°C = 28.7 min) and no significant change in Scrit (+8°C = 17.0% air saturation; +10°C = 18.3% of air saturation). Thus, there was no evidence of heat shock induced cross-tolerance to hypoxia in B. medius because exposure to acute heat shock impaired hypoxia tolerance.

Geographic and ontogenetic variations in parasite communities of intertidal fish species from the south-eastern Pacific coast

Parasite composition can be affected by physiological and ecological changes during host ontogeny. Intertidal fish do not travel long distances and live in the same area throughout their lifetimes, meaning that parasite communities can differ across geographic ranges. The objective of this study was to analyse the parasite communities of three fish species (Hypsoblennius sordidus, Helcogrammoides cunninghami and Scartichthys viridis) collected from the Chilean coast. The composition of parasite species was compared among host ontogenetic stages (larvae, juveniles and mature fish) and geographic areas. A total of 184 larval, 252 juveniles and 217 mature individuals were collected in the northern area (c. 24°S), and 186 larval, 192 juveniles and 112 mature individuals from the central area (c. 33°S). Ectoparasites were most prevalent in fish from the central area, whereas endoparasites were most prevalent in the northern area. The parasite species richness varied significantly between geographical areas for H. sordidus and H. cunninghami, but the parasite composition varied significantly between geographical areas for all fish species analysed. Therefore, the geographical area was the most important factor determining the parasite composition of intertidal fish species. The absence of endoparasites in fish larvae and the increased infestation in juvenile and mature fish may be explained by the shift in habitat from the water column to intertidal pools where prey abundance and availability are higher. On the other hand, hydrographic barriers affecting prey distributions may also offer an explanation as to the differences in parasite composition.

Size, sex and social experience: Experimental tests of multiple factors mediating contest behaviour in a rockpool fish

AbstractAggressive contests amongst conspecifics are important to understand from an ecological and evolutionary perspective as contest dynamics can directly influence individual fitness. For some species, individual attributes such as relative body size closely predict the outcome, intensity and duration of contests, whereas for others, prior social experiences play a key role. However, disentangling the relative effects of individual attributes and social experiences is challenging from an experimental perspective, and because of this, few studies have investigated relative effects of multiple factors. Rockpool fishes have been well studied in terms of factors governing abundance, distribution and community structure, but much less so in terms of contest behaviour. This is surprising because contest dynamics are likely to directly affect the distribution of fishes along the rocky shore, and hence indirectly govern population and community composition. Here, we explored multiple factors potentially influencing contest behaviour in a numerically dominant, resident intertidal fish species, Bathygobius cocosensis (Gobiidae). Using a series of staged pairwise contest trials, we investigated the effect of size, sex and social experiences (namely prior residency and winner–loser experiences) on contest dynamics. We found no evidence that prior residency influenced contest outcome, suggesting social experiences play a minor role in contest dynamics. Previous winner/loser experience also did not influence contest outcome, although this is likely a product of low sample size. In addition, the likelihood of winning was unrelated to contestant sex, and the combination of sexes in paired contests did not influence contest intensity or duration. Instead, body size was related to contest outcome, intensity and duration in the majority of experimental trials. These results suggest that body size rather than sex or social experiences is the key determinant of contest dynamics in this species. We suggest that the dynamic biotic and abiotic environment inhabited by intertidal fish may reduce the influence of prior social experiences in modulating contest dynamics.

Regional and Seasonal Differences in Species Composition and Trophic Groups for Tidepool Fishes of a Western Pacific Island – Taiwan

Spatial and temporal variations in the species composition of assemblages are common in many marine organisms, including fishes. Variations in the fish species composition of subtidal coral reefs have been well documented, however much less is known about such differences for intertidal fish assemblages. This is surprising, given that intertidal fishes are more vulnerable to terrestrial human disturbances. It is critical to evaluate the ecology and biology of intertidal fishes before they are severely impacted by coastal development, especially in developing countries such as those in the tropical western Pacific region where coastal development is rapidly increasing. In this study, we investigated the species composition, abundance, biomass and species number (richness) for intertidal fish assemblages in subtropical (northern) and tropical (southern) Taiwan across four seasons by collecting fishes from tidepools using clove oil. We also examined the gut contents of collected fishes to identify their trophic functional groups in order to investigate regional and seasonal variations for different trophic groups. We found significant differences in the species composition of tidepool fish assemblages between subtropical and tropical Taiwan. Bathygobius fuscus, Abudefduf vaigiensis and Istiblennius dussumieri were dominant species in subtropical Taiwan, whereas Bathygobius coalitus, Abudefduf septemfasciatus and Istiblennius lineatus were dominant in tropical Taiwan. Other species such as Bathygobius cocosensis, Abudefduf sordidus and Istiblennius edentulus were common in both regions. For trophic groups, omnivores and detritivores had or showed trends towards higher species numbers and abundances in the subtropical region, whereas herbivores, planktivores and general carnivores had or showed trends towards higher species numbers and biomass in the tropical region. Overall, many intertidal fish species and trophic groups showed differences in abundance, biomass and species number between subtropical and tropical Taiwan. Further studies on large scale geographical gradients in trophic groups and species compositions in the Indo-west Pacific region are encouraged to assist with ecosystem monitoring and assessment. Keywords: Intertidal fishes, spatio-temporal pattern, feeding guild, diet

The complete mitochondrial genome of Bathygobius cocosensis (Perciformes, Gobiidae)

In this study, we sequenced the full mitochondrial genome of Bathygobius cocosensis, an abundant intertidal fish species, which may provide insights into the evolutionary genetics of chaotic genetic patchiness and range expansion in marine systems. The mitochondrial genome is 16,692 bp, and contains 13 protein-coding genes along with 22 tRNA and 2 rRNA genes and a D-loop region, arranged similarly to other Gobiidae species. A Bayesian phylogeny of Gobiidae species indicates close relationships to the genus Glossogobius. The B. cocosensis mitochondrial genome is now available through GenBank (Accession = MG704838).

Chronic warm exposure impairs growth performance and reduces thermal safety margins in the common triplefin fish (Forsterygion lapillum)

Intertidal fish species face gradual chronic changes in temperature and greater extremes of acute thermal exposure through climate-induced warming. As sea temperatures rise, it has been proposed that whole-animal performance will be impaired through oxygen and capacity limited thermal tolerance [OCLTT; reduced aerobic metabolic scope (MS)] and, on acute exposure to high temperatures, thermal safety margins may be reduced because of constrained acclimation capacity of upper thermal limits. Using the New Zealand triplefin fish (Forsterygion lapillum), this study addressed how performance in terms of growth and metabolism (MS) and upper thermal tolerance limits would be affected by chronic exposure to elevated temperature. Growth was measured in fish acclimated (12 weeks) to present and predicted future temperatures and metabolic rates were then determined in fish at acclimation temperatures and with acute thermal ramping. In agreement with the OCLTT hypothesis, chronic exposure to elevated temperature significantly reduced growth performance and MS. However, despite the prospect of impaired growth performance under warmer future summertime conditions, an annual growth model revealed that elevated temperatures may only shift the timing of high growth potential and not the overall annual growth rate. While the upper thermal tolerance (i.e. critical thermal maxima) increased with exposure to warmer temperatures and was associated with depressed metabolic rates during acute thermal ramping, upper thermal tolerance did not differ between present and predicted future summertime temperatures. This suggests that warming may progressively decrease thermal safety margins for hardy generalist species and could limit the available habitat range of intertidal populations.

A new species of Hemipera Nicoll, 1913 (Digenea: Derogenidae) from fishes of the intertidal rocky zone of Chile.

A new species, Hemipera cribbi sp. nov., is described. This trematode was found in three intertidal fish species: Scartichthys viridis (Valenciennes) (Blenniidae), Gobiesox marmoratus Jenyns (Gobiesocidae) and Myxodes viridis Valenciennes (Clinidae) from the central and southern coast of Chile. Of 233 individuals of S. viridis from the central coast examined, 19 were infected. From the southern coast, nine individuals of S. viridis (one infected), five individuals of G. marmoratus (four infected), and 16 individuals of M. viridis (one fish infected) were examined. Hemipera cribbi sp. nov. is distinguished from the five other congeneric species mainly in the body size, being the smallest and narrowest species in the genus, reaching five times longer than wide. This is the first species of the genus described for the South Pacific Ocean off South America. ITS2 rDNA sequences of Hemipera cribbi sp. nov. from each host and locality were identified.

Diet as a mechanism of coexistence between intertidal fish species of the U.K.

While the syntopic nature of many intertidal fish communities suggest that resources such as food are shared, little has been done to assess the importance of diet on the coexistence of intertidal fish of the U.K. In this study, six intertidal fish species (shanny, Lipophrys pholis, Blenniidae; long-spined scorpion fish, Taurulus bubalis, Cottidae; two-spotted goby, Gobiusculus flavescens, Gobiidae; rock goby, Gobius paganellus, Gobiidae; plaice, Pleuronectes platessa, Pleuronectidae; butterfish, Pholis gunnellus, Pholidae) were collected from two sites along the east coast of England (Filey and Thornwick Bay) and two sites around the coast of the Isle of Anglesey, North Wales (Rhosneigr and Penrhos), during summer and winter. A comparison of the diets of those in the highest abundances (L. pholis, T. bubalis and G. flavescens) found that, in general, prey preferences were dissimilar between species, albeit with some slight overlap, and therefore it could be said that diet acts as an important mechanism of interspecific coexistence.

Effect of UV radiation on habitat selection by Girella laevifrons and Graus nigra (Kyphosidae).

The effect of UV radiation on habitat use of two species of intertidal fishes that inhabit the same pools but exhibit different activity levels and diets was measured: the highly active omnivorous Girella laevifrons and the cryptic carnivorous Graus nigra. Individuals of each species were acclimated to a tank divided in three sections with different illumination; no light (NL), ultraviolet light (UV) and white light (WL), and the time spent and number of visits to each section were recorded. Although both species preferred the NL section, G. laevifrons spent more time in UV and less time in WL compared with G. nigra; G. laevifrons also displayed higher number of visits to UV, suggesting a different tendency in space use in response to UV exposure in intertidal fishes.

Influence of rock‐pool characteristics on the distribution and abundance of inter‐tidal fishes

AbstractRock pools can be found in inter‐tidal marine environments worldwide; however, there have been few studies exploring what drives their, fish species composition, especially in Australia. The rock‐pool environment is highly dynamic and offers a unique natural laboratory to study the habitat choices, physiological limitations and adaptations of inter‐tidal fish species. In this study rock pools of the Sydney region were sampled to determine how the physical (volume, depth, rock cover and vertical position) and biological (algal cover and predator presence) parameters of pools influence fish distribution and abundance. A total of 27 fish species representing 14 families was observed in tide pools at the four study locations. The five most abundant species were Bathygobius cocosensis, Centropogon australis, Enneapterygius atrogulare, Lepidoblennius haplodactylus and Microcanthus strigatus, which together represented 71% of the total number of fish recorded. Larger rock pools containing more algal and rock ledge cover hosted a larger and more diverse population of fish. Furthermore, certain species were only found in pools with specific characteristics, such as the presence of loose shells, a variety algae or rock cover, suggesting a high degree of habitat specificity. By contrast, some species were ubiquitous and thus can likely tolerate a wide variety of physical conditions.

The life cycle of Prosorhynchoides carvajali (Trematoda: Bucephalidae) involving species of bivalve and fish hosts in the intertidal zone of central Chile.

We describe the life cycle of the bucephalid Prosorhynchoides carvajali from the intertidal rocky zone of central Chile. To elucidate the life cycle of this digenean, two mytilid bivalves, Semimytilus algosus and Perumytilus purpuratus, and ten intertidal fish species belonging to the families Blenniidae, Tripterygiidae, Labrisomidae, Kyphosidae and Gobiesocidae were analysed for natural infections. In addition, experimental infections of fish were undertaken and molecular analyses were performed of several developmental stages of the digeneans in various host species. Experimental infections of fish were made from infected mytilids to determine which fish species were suitable for the metacercarial stage of Prosorhynchoides. We also determined the abundance and prevalence of metacercariae in natural infections in fish and found that they were lower than in the experimental infections. A molecular analysis showed that sporocysts from S. algosus were identical to metacercariae from five fish species and P. carvajali adults. Sporocysts isolated from P. purpuratus were similar to metacercaria found in one fish species only (G. laevifrons) but were different from P. carvajali, with 1.9-2.0% genetic divergence. Therefore, the complete life cycle of P. carvajali consists of the mytilid species S. algosus as the first intermediate host, at least five intertidal fish species as second intermediate hosts (Scartichthys viridis, Auchenionchus microcirrhis, Hypsoblennius sordidus, Helcogrammoides chilensis and Gobiesox marmoratus), two carnivorous fish as definitive hosts (Auchenionchus microcirrhis and A. variolosus) and one occasional definitive host (Syciases sanguineus). This is the second description of a life cycle of a marine digenean from Chile.

Estructura trófica de la asociación de peces intermareales de la costa rocosa del norte de Chile

The trophic relationships of 13 intertidal fish species collected in northern Chile (20 degrees 18' S and 20 degrees 54' S) from July 1997 to July 1998, the stomach content analysis of 820 specimens, were investigated. The importance of the prey items was assessed by means of gravimetric and frequency of ocurrence methods, together with the Shannon-Wiener (trophic diversity) and Pianka (trophic overlap) indexes. The results allowed to recognize a fish assemblage composed of: 46% of carnivorous species, preying mainly on porcelain crabs. polichaetes and minor crustaceans: 23% of herviborous species, preying mainly on chlorophitic algae: 31% of omnivorous species. preying mainly on chlorophitic algae. Copepoda and Gastropoda. The carnivorous fishes Cheilodactylus variegatus. Helcogrammoides chilensis. Labrisomus philippii and the omnivorous Oplegnatus insignis showed euriphagic tendencies (>2.0 bits), however, the major trophic overlap (>0.90) was observed among herbivorous and omnivorous fishes. Finally, the increase of fish species towards lower latitudes and the increase of herbivory and omnivory levels in the rocky intertidal shore were analyzed.

Theme and variations: amphibious air‐breathing intertidal fishes

Over 70 species of intertidal fishes from 12 families breathe air while emerging from water. Amphibious intertidal fishes generally have no specialized air-breathing organ but rely on vascularized mucosae and cutaneous surfaces in air to exchange both oxygen and carbon dioxide. They differ from air-breathing freshwater fishes in morphology, physiology, ecology and behaviour. Air breathing and terrestrial activity are present to varying degrees in intertidal fish species, correlated with the tidal height of their habitat. The gradient of amphibious lifestyle includes passive remainers that stay in the intertidal zone as tides ebb, active emergers that deliberately leave water in response to poor aquatic conditions and highly mobile amphibious skipper fishes that may spend more time out of water than in it. Normal terrestrial activity is usually aerobic and metabolic rates in air and water are similar. Anaerobic metabolism may be employed during forced exercise or when exposed to aquatic hypoxia. Adaptations for amphibious life include reductions in gill surface area, increased reliance on the skin for respiration and ion exchange, high affinity of haemoglobin for oxygen and adjustments to ventilation and metabolism while in air. Intertidal fishes remain close to water and do not travel far terrestrially, and are unlikely to migrate or colonize new habitats at present, although in the past this may have happened. Many fish species spawn in the intertidal zone, including some that do not breathe air, as eggs and embryos that develop in the intertidal zone benefit from tidal air emergence. With air breathing, amphibious intertidal fishes survive in a variable habitat with minimal adjustments to existing structures. Closely related species in different microhabitats provide unique opportunities for comparative studies.