Fish distributions along depth gradients of a sea mountain range conform to the mid‐domain effect

Species richness often peaks in the middle of bounded geographic domains (e.g. latitude, altitude or depth). Hump‐shaped richness distributions may be due to deterministic processes, such as adaptations to environmental variation. Alternatively, such distributions might also be due to stochastic process. The mid‐domain effect (MDE) posits that hump‐shaped richness distributions arise when species ranges are randomly arranged within the limits of the domain. We tested whether the MDE could account for the richness of bottom‐associated (demersal) fishes between 200 and 800 m on the Chatham Rise, New Zealand. We quantified the depth distributions of 59 fish species from 1891 research trawl catches made between 1991 and 2007. Results showed a broad plateau of high species richness near the centre of the domain (between 300 and 700 m), which was consistent with expectations of the MDE. Further, empirical species richness was better explained statistically by predictions of the MDE than models incorporating additional abiotic predictor variables. Our results deviated from previous studies that identified a greater richness of fishes in warmer, shallower depths with higher primary production. However, our study was conducted entirely below the euphotic zone, at depths where gradients are relatively weak, suggesting that support for the mid‐domain effect may increase across oceanic domains characterised by weak environmental gradients.     

Patterns of diversity, altitudinal range and body size among freshwater fishes in the Yangtze River basin, China

Aim To document patterns in diversity, altitudinal range and body size of freshwater fishes along an elevational gradient in the Yangtze River basin. Location The Yangtze River basin, China. Methods We used published data to compile the distribution, altitudinal range and body size of freshwater fishes. Correlation, regression, clustering and graphical analyses were used to explore patterns in diversity, altitudinal range and body size of freshwater fishes in 100‐m elevation zones from 0 to 5200 m. Results Species richness patterns across the elevational gradient for total, non‐endemic and endemic fishes were different. The ratio of endemics to total richness peaked at mid elevation. Land area on a 500‐m interval scale explained a significant amount of the variation in species richness. Species density displayed two peaks at mid‐elevation zones. The cluster analysis revealed five distinct assemblages across the elevation gradient. The relationship between elevational range size and the midpoint of the elevational range revealed a triangular distribution. The frequency distribution of log maximum standard length data displayed an atypical right‐skewed pattern. Intermediate body sizes occurred across the greatest range of elevation while small and large body sizes possessed only small elevational amplitudes. The size‐elevation relationship between the two major families revealed a very strong pattern of body size constraint among the Cobitidae with no corresponding elevational constraint and a lot of body size and elevational diversification among the Cyprinidae. Main conclusion The data failed to support either Rapoport's rule or Bergmann's rule.     

Vertical gradients in species richness and community composition across the twilight zone in the North Pacific Subtropical Gyre

Although metazoan animals in the mesopelagic zone play critical roles in deep pelagic food webs and in the attenuation of carbon in midwaters, the diversity of these assemblages is not fully known. A metabarcoding survey of mesozooplankton diversity across the epipelagic, mesopelagic and upper bathypelagic zones (0–1500 m) in the North Pacific Subtropical Gyre revealed far higher estimates of species richness than expected given prior morphology‐based studies in the region (4,024 OTUs, 10‐fold increase), despite conservative bioinformatic processing. Operational taxonomic unit (OTU) richness of the full assemblage peaked at lower epipelagic–upper mesopelagic depths (100–300 m), with slight shoaling of maximal richness at night due to diel vertical migration, in contrast to expectations of a deep mesopelagic diversity maximum as reported for several plankton groups in early systematic and zoogeographic studies. Four distinct depth‐stratified species assemblages were identified, with faunal transitions occurring at 100 m, 300 m and 500 m. Highest diversity occurred in the smallest zooplankton size fractions (0.2–0.5 mm), which had significantly lower % OTUs classified due to poor representation in reference databases, suggesting a deep reservoir of poorly understood diversity in the smallest metazoan animals. A diverse meroplankton assemblage also was detected (350 OTUs), including larvae of both shallow and deep living benthic species. Our results provide some of the first insights into the hidden diversity present in zooplankton assemblages in midwaters, and a molecular reappraisal of vertical gradients in species richness, depth distributions and community composition for the full zooplankton assemblage across the epipelagic, mesopelagic and upper bathypelagic zones.     

Phylogenetic measures reveal eco-evolutionary drivers of biodiversity along a depth gradient

Energy and environmental stability are positively correlated with species richness along broad-scale spatial gradients in terrestrial ecosystems, so their relative importance in generating and preserving diversity cannot be readily disentangled. This study seeks to exploit the negative correlation between energy and stability along the oceanic depth gradient to better understand their relative contribution in shaping broadscale biodiversity patterns. We develop a conceptual framework by simulating speciation and extinction along energy and stability gradients to generate expected patterns of biodiversity for a suite of complementary phylogenetic diversity metrics. Using a time-calibrated molecular phylogeny for New Zealand marine ray-finned fishes and a replicated community ecological sampling design, we then modelled these metrics along large-scale depth and latitude gradients. Our results indicate that energy-rich shallow waters may be an engine of diversity for percomorphs, but also suggest that recent speciation occurs in ancient fish lineages in the deep sea, hence questioning the role of energy as a key driver of speciation. Despite potentially facing high extinction early in their evolution, ancient phylogenetic lineages specialized for the deep-sea were likely preserved by environmental stability during the Cenozoic. Furthermore, intermediate depths might be a ‘museum’ (or zone of overlap) for distinct lineages that occur predominantly in either shallow or deep-sea waters. These intermediate depths (500–900 m) may form a ‘phylogenetic diversity bank’, perhaps providing a refuge during ancient (Mesozoic) extreme anoxic events affecting the deep sea and more recent (Pliocene–Pleistocene) climatic events occurring in shallow ecosystems. Finally, the phylogenetic structures observed in fish communities at intermediate depths suggest other processes might restrict the co-occurrence of closely related species. Overall, by combining a conceptual framework with models of empirical phylogenetic diversity patterns, our study paves the way for understanding the determinants of biodiversity across the largest habitat on earth.     

Increasing variation in taxonomic distinctness reveals clusters of specialists in the deep sea

Changes in biodiversity with latitude or along a given environmental gradient have been described in many studies, including for marine ecosystems. Currently there is no scientific consensus, however, regarding macroecological patterns of diversity vs depth. Here, we describe variation in the biodiversity of fishes along a depth gradient from 0 to 2000 m in the region of the Norfolk Ridge and Lord Howe Rise (Western Pacific), using data obtained during the NORFANZ voyage. We modelled α diversity (richness), β diversity (using Jaccard's coefficient), evenness, taxonomic distinctness and taxonomic resemblances among fish communities. Although α diversity did not change appreciably with depth, β diversity decreased significantly in deeper strata. Both taxonomic resemblances and Jaccard similarities diminished with depth, indicating convergence in community structure. In addition, average taxonomic distinctness showed no clear pattern with depth, but taxonomic trees constructed among species within deeper samples had more variable path‐lengths than those in shallower samples. The presence of taxonomically distinct clusters of highly related species at depth indicates specialised niches that have developed in a relatively extreme (dark, pressurized) yet stable environment. We propose that reduced β diversity and increased variation in taxonomic distinctness might serve as indicators of ecological communities living in harsh environments – a hypothesis that should be tested in other systems, such as deserts, high altitudes or latitudes.     

Diversity, abundance and depth distribution of demersal deep-water fishes off Lanzarote and Fuerteventura, Canary Islands

A total of 168 bottom longline operations were carried out at depths between 72 and 1102 m during four cruises off Lanzarote and Fuerteventura, Canary Islands. One hundred and twenty-eight operations were successful, resulting in the capture of 1167 fishes belonging to 55 species. Clear variations in fish total weight and number, species richness and diversity, and the relative abundance of single species occurred among five different depth zones between 100 and 1100 m. The particular platform topography of the Canary Islands may affect the density and diversity of slope-dwelling fishes. Whereas fish body length and weight showed an overall increase with depth, no such trends were found at the level of individual species. One species, Lepidopus caudalus , even revealed a bigger-shallower trend. Among three species of trichiurids, depth distribution patterns were detected that may reflect vertical space partitioning. In October 1995, an unusually high number of Mora moro with advanced gonadal maturity were collected off eastern Fuerteventura at 997 m depth. At lesser depths and during other seasons, only a few individuals of this species were caught in the same area. This indicated a seasonal variability in local abundance that is most probably related to reproduction. Mora moro has an excellent flesh and may represent a valuable fisheries resource. Open problems requiring further research in the study area such as selectivity effects of the fishing gear and possible influences of ecological factors on geographical distribution of the deep-water ichthyofauna, as well as important aspects of the future demersal fisheries management in the Canarian archipelago are briefly outlined.     

Altitudinal variation in fish assemblage diversity in streams of the central Andes of Colombia

This study documents differences in fish assemblages for 32 freshwater streams located between 258 and 2242 m a.s.l. on the eastern slopes of the central range of the Colombian Andes. A total of 2049 fishes belonging to 62 species, 34 genera and 16 families were collected. Species richness declined rapidly with altitude; nearly 90% of the species were recorded between 250 and 1250 m a.s.l. Three of the four physico‐chemical variables, of the water, temperature, dissolved oxygen and pH, explained 53·5% of the variation in species richness along the altitudinal gradient, with temperature the most important (37·6%). An analysis of species composition showed that the distinctiveness of the fish fauna increased with elevation, with the greatest turnover observed between 1000 and 1750 m a.s.l. On this altitudinal gradient, turnover was dominated by the loss of species rather than gain, and dominance by just a few species was greater at higher elevations. Turnover was also observed along the altitudinal gradient in the structure of the three functional groups (torrential, pool and pelagic species). The study focused on understanding the pattern of diversity of fish communities inhabiting the Andes in Colombia. Anthropogenic effects on the altitudinal distribution of fish species in the region, however, are largely unknown and would require further investigations.     

Circumtropical patterns in butterflyfish communities

Synopsis Butterflyfish species richness increases along a longitudinal circumtropical gradient from lows of 3–5 species in the tropical Atlantic and Eastern Pacific to highs of 40 or more in the Indo-Pacific region. Biomass of the fishes increases as species richness increases, and single-site (alpha) diversity increases as does between-site (beta) diversity. There is no evidence of density compensation in richer communities, but at the level of islands and regions, habitat breadth diminishes as species richness increases. Morphologically, species are added to communities both at the boundaries and in the middle of morphospace.     

Explaining bathymetric diversity patterns in marine benthic invertebrates and demersal fishes: physiological contributions to adaptation of life at depth

Bathymetric biodiversity patterns of marine benthic invertebrates and demersal fishes have been identified in the extant fauna of the deep continental margins. Depth zonation is widespread and evident through a transition between shelf and slope fauna from the shelf break to 1000 m, and a transition between slope and abyssal fauna from 2000 to 3000 m; these transitions are characterised by high species turnover. A unimodal pattern of diversity with depth peaks between 1000 and 3000 m, despite the relatively low area represented by these depths. Zonation is thought to result from the colonisation of the deep sea by shallow‐water organisms following multiple mass extinction events throughout the Phanerozoic. The effects of low temperature and high pressure act across hierarchical levels of biological organisation and appear sufficient to limit the distributions of such shallow‐water species. Hydrostatic pressures of bathyal depths have consistently been identified experimentally as the maximum tolerated by shallow‐water and upper bathyal benthic invertebrates at in situ temperatures, and adaptation appears required for passage to deeper water in both benthic invertebrates and demersal fishes. Together, this suggests that a hyperbaric and thermal physiological bottleneck at bathyal depths contributes to bathymetric zonation. The peak of the unimodal diversity–depth pattern typically occurs at these depths even though the area represented by these depths is relatively low. Although it is recognised that, over long evolutionary time scales, shallow‐water diversity patterns are driven by speciation, little consideration has been given to the potential implications for species distribution patterns with depth. Molecular and morphological evidence indicates that cool bathyal waters are the primary site of adaptive radiation in the deep sea, and we hypothesise that bathymetric variation in speciation rates could drive the unimodal diversity–depth pattern over time. Thermal effects on metabolic‐rate‐dependent mutation and on generation times have been proposed to drive differences in speciation rates, which result in modern latitudinal biodiversity patterns over time. Clearly, this thermal mechanism alone cannot explain bathymetric patterns since temperature generally decreases with depth. We hypothesise that demonstrated physiological effects of high hydrostatic pressure and low temperature at bathyal depths, acting on shallow‐water taxa invading the deep sea, may invoke a stress–evolution mechanism by increasing mutagenic activity in germ cells, by inactivating canalisation during embryonic or larval development, by releasing hidden variation or mutagenic activity, or by activating or releasing transposable elements in larvae or adults. In this scenario, increased variation at a physiological bottleneck at bathyal depths results in elevated speciation rate. Adaptation that increases tolerance to high hydrostatic pressure and low temperature allows colonisation of abyssal depths and reduces the stress–evolution response, consequently returning speciation of deeper taxa to the background rate. Over time this mechanism could contribute to the unimodal diversity–depth pattern.     

Red muscle proportions and enzyme activities in deep‐sea demersal fishes

Owing to the paucity of data on the red muscle of deep‐sea fishes, the goal of this study was to determine the proportions of red muscle in demersal fishes and its enzymatic activities to characterize how routine swimming abilities change with depths of occurrence. Cross sectional analysis of the trunk musculature was used to evaluate the proportion of red muscle in 38 species of Californian demersal fishes living at depths between 100 and 3000 m. The activity of metabolic enzymes was also assayed in a sub‐set of 18 species. Benthic fishes had lower proportions of red muscle and lower metabolic enzyme activities than benthopelagic species. Mean proportion of red muscle declined significantly with depth with the greatest range of values in shallow waters and species with low proportions found at all depths. This suggested that while sedentary species occur at all depths, the most active species occur in shallow waters. Citrate synthase activity declined significantly with depth across all species, indicating that the mass‐specific metabolic capacity of red muscle is lower in deep‐sea species. These patterns may be explained by coupling of red and white muscle physiologies, a decrease in physical energy of the environment with depth or by the prevalence of anguilliform body forms and swimming modes in deep‐living species.     

牛山湖两种优势小型鱼类空间分布与沉水植被的关系

运用8种网目规格的成套浮性刺网作为鱼类采样工具,于2005年夏季在长江中游浅水草型湖泊牛山湖进行鱼类定量采样,通过比较不同茂密程度黄丝草生境中的小型鱼类组成、数量和大小结构,探讨此类湖泊小型鱼类的空间分布特征及其与沉水植被的关系.采样期间共捕获13种1124尾鱼,依据其等级丰度和出现频次,鳖和红鳍原鲌为该湖优势上层小型鱼类.在调查的沉水植物生物量范围内,鱼类物种丰富度和Shannon多样性指数与沉水植物生物量之间呈现倒抛物线关系;两种优势小型鱼类的种群丰度均与沉水植物生物量有着显著的线性正相关关系,且其平均个体大小在裸地生境较高、沉水植被茂密区较低,幼鱼更倾向群聚于厚密的黄丝草生境中;其他生境因子(水深和离岸距离)对鳖和红鳍原鲐空间分布的影响不显著.黄丝草植被生境是牛山湖两种优势小型鱼类的重要保护生境,应加强对黄丝草等沉水植被的保护及恢复.     

Do Rapoport's rule,the mid‐domain effect or the source–sink hypotheses predict bathymetric patterns of polychaete richness on the Pacific coast of South America?

Aim We evaluated the bathymetric gradient of benthic polychaete species richness from the Chilean coast, as well as its possible underlying causes. We tested three possible hypotheses to explain the richness gradient: (1) Rapoport's effect; (2) the mid‐domain effect (MDE); and (c) the source–sink hypothesis. Location South‐eastern Pacific coast of Chile. Methods The bathymetric gradient in richness was evaluated using the reported ranges of bathymetric distribution of 498 polychaete species, from the intertidal to abyssal zone (c. 4700 m). Rapoport's effect was evaluated by examining the relationship between bathymetric mid‐point and bathymetric range extent, and species richness and depth. The MDE was tested using the Monte Carlo simulation program. The source–sink hypothesis was tested through nestedness analysis. Results Species richness shows significant exponential decay across the bathymetric gradient. The pattern is characterized by a high presence of short‐ranged species on the continental shelf area; while only a few species reach abyssal depths, and they tend to show extremely wide bathymetric ranges. Our simulation analyses showed that, in general, the pattern is robust to sampling artefacts. This pattern cannot be reproduced by the MDE, which predicts a parabolic richness gradient. Rather, results agree with the predictions of Rapoport's effect. Additionally, the data set is significantly nested at species, genus and family levels, supporting the source–sink hypothesis. Main conclusions The sharp exponential decay in benthic polychaete richness across the bathymetric gradient supports the general idea that abyssal environments should harbour fewer species than shallower zones. This pattern may be the result of colonization–extinction dynamics, characterized by abyssal assemblages acting as ‘sinks’ maintained mainly by shallower ‘sources’. The source–sink hypothesis provides a conceptual and methodological framework that may shed light on the search for general patterns of diversity across large spatial scales.     

Intraspecific variability in the life histories of endemic coral-reef fishes between photic and mesophotic depths across the Central Pacific Ocean

Mesophotic coral ecosystems (MCEs) represent the lowest depth distribution inhabited by many coral reef-associated organisms. Research on fishes associated with MCEs is sparse, leading to a critical lack of knowledge of how reef fish found at mesophotic depths may vary from their shallow reef conspecifics. We investigated intraspecific variability in body condition and growth of three Hawaiian endemics collected from shallow, photic reefs (5–33 m deep) and MCEs (40–75 m) throughout the Hawaiian Archipelago and Johnston Atoll: the detritivorous goldring surgeonfish, Ctenochaetus strigosus, and the planktivorous threespot chromis, Chromis verater, and Hawaiian dascyllus, Dascyllus albisella. Estimates of body condition and size-at-age varied between shallow and mesophotic depths; however, these demographic differences were outweighed by the magnitude of variability found across the latitudinal gradient of locations sampled within the Central Pacific. Body condition and maximum body size were lowest in samples collected from shallow and mesophotic Johnston Atoll sites, with no difference occurring between depths. Samples from the Northwestern Hawaiian Islands tended to have the highest body condition and reached the largest body sizes, with differences between shallow and mesophotic sites highly variable among species. The findings of this study support newly emerging research demonstrating intraspecific variability in the life history of coral-reef fish species whose distributions span shallow and mesophotic reefs. This suggests not only that the conservation and fisheries management should take into consideration differences in the life histories of reef-fish populations across spatial scales, but also that information derived from studies of shallow fishes be applied with caution to conspecific populations in mesophotic coral environments.

     

Spatial distribution and features of biology of Pacific sleeper shark Somniosus pacificus in the North Pacific

The results of the investigations of spatial and vertical distribution of Pacific sleeper shark Somniosus pacificus in the North Pacific Ocean conducted for many years are presented. In addition, the size distribution and features of biology of the species are studied. The largest abundance of the species is registered in the Bering Sea, western Gulf of Alaska, eastern Aleutian Islands, and Pacific waters of northern Kuril Islands and southeastern Kamchatka. The species is the most abundant near the bottom at the depth from 200 to 700 m and in the pelagic waters at a depth of 100–200 m. The average depths of the catches of Pacific sleeper shark substantially change over the year reaching minimum values in June and maximum values in December. Vertical daily migrations (to the water column at night and to the bottom during the day) are registered. The catches are represented by fish 26–352 cm in length, and sharks 100–200 cm in length prevail. The males are noticeably smaller than the females. In general, condition of the fishes decreases and feeding intensity increases with growth. Food composition substantially changes with the increase of body length: consumption of squids decreases and consumption of crustaceans, fishes, and fishery wastes increases. The food composition is slightly different in the females and males.     

Mesophotic depths as refuge areas for fishery-targeted species on coral reefs

Coral reefs are subjected to unprecedented levels of disturbance with population growth and climate change combining to reduce standing coral cover and stocks of reef fishes. Most of the damage is concentrated in shallow waters (<30 m deep) where humans can comfortably operate and where physical disturbances are most disruptive to marine organisms. Yet coral reefs can extend to depths exceeding 100 m, potentially offering refuge from the threats facing shallower reefs. We deployed baited remote underwater stereo-video systems (stereo-BRUVs) at depths of 10–90 m around the southern Mariana Islands to investigate whether fish species targeted by fishing in the shallows may be accruing benefits from being at depth. We show that biomass, abundance and species richness of fishery-targeted species increased from shallow reef areas to a depth of 60 m, whereas at greater depths, a lack of live coral habitat corresponded to lower numbers of fish. The majority of targeted species were found to have distributions that ranged from shallow depths (10 m) to depths of at least 70 m, emphasising that habitat, not depth, is the limiting factor in their vertical distribution. While the gradient of abundance and biomass versus depth was steepest for predatory species, the first species usually targeted by fishing, we also found that fishery-targeted herbivores prevailed in similar biomass and species richness to 60 m. Compared to shallow marine protected areas, there was clearly greater biomass of fishery-targeted species accrued in mesophotic depths. Particularly some species typically harvested by depth-limited fishing methods (e.g., spearfishing), such as the endangered humphead wrasse Cheilinus undulatus, were found in greater abundance on deeper reefs. We conclude that mesophotic depths provide essential fish habitat and refuge for fishery-targeted species, representing crucial zones for fishery management and research into the resilience of disturbed coral reef ecosystems.     

Biodiversity hotspots, centres of endemicity, and the conservation of coral reefs

On land, biodiversity hotspots typically arise from concentrations of small‐range endemics. For Indo‐Pacific corals and reef fishes, however, centres of high species richness and centres of high endemicity are not concordant. Moreover ranges are not, on average, smaller inside the Central Indo‐Pacific (CI‐P) biodiversity hotspot. The disparity between richness and endemicity arises because corals and reef fishes have strongly skewed range distributions, with many species being very widespread. Consequently, the largest ranges overlap to generate peaks in species richness near the equator and the CI‐P biodiversity hotspot, with only minor contributions from endemics. Furthermore, we find no relationship between the number of coral vs. fish endemics at locations throughout the Indo‐Pacific, even though total richness of the two groups is strongly correlated. The spatial separation of centres of endemicity and biodiversity hotspots in these taxa calls for a two‐pronged management strategy to address conservation needs.     

南四湖沉水植物物种多样性和功能多样性对水深梯度的响应

从物种多样性和功能多样性探讨沉水植物群落对水深的响应可深刻揭示水深对群落构建的影响机制。以南四湖不同水深沉水植物群落为研究对象,对比分析了群落的9个加权功能性状（株高、茎分支数、茎节数、茎直径、根长、根围直径、生物量分配比、比叶面积、植物体磷含量）、5个物种多样性指数（Berger-Parker生态优势度指数、Margalef丰富度指数、Simpson多样性指数、Shannon-Wiener多样性指数和Pielou均匀度指数）和5个功能多样性指数（功能丰富度FRic指数、功能均匀度FEve指数、功能离散度FDiv指数、功能分散度FDis指数和二次熵Rao指数）以及物种多样性和功能多样性关系对水深的响应规律。研究结果表明：（1）水深可显著改变群落株高、根长、根围直径、比叶面积、生物量分配比和植物体磷含量6个加权功能性状;群落茎分支数、茎节数和茎直径3个加权功能性状对水深变化无显著响应;（2）水深可显著影响群落物种多样性和功能多样性,中等水深处沉水植物群落具有较高的Margalef丰富度指数、Shannon-Wiener多样性指数、Berger-Parker生态优势度指数、Pielou均匀度指数以及功能丰富度FRic指数、功能均匀度FEve指数、功能离散度FDis指数、二次熵Rao指数;（3）水深可改变5个物种多样性指数与功能丰富度FRic指数、功能均匀度FEve指数、功能离散度FDiv指数3个功能多样性指数间的相关关系,但对5个物种多样性指数与功能离散度FDis指数、二次熵Rao指数2个功能多样性指数间的相关关系无显著影响。研究结论为：群落不同测度的物种多样性和功能多样性指数及其相关关系对水深变化的响应迥异,在探讨水深对沉水植物群落构建的影响机制时应从多个方面综合考量。     

Community structure and biogeography of shore fishes in the Gulf of Aqaba,Red Sea

Shore fish community structure off the Jordanian Red Sea coast was determined on fringing coral reefs and in a seagrass-dominated bay at 6 m and 12 m depths. A total of 198 fish species belonging to 121 genera and 43 families was recorded. Labridae and Pomacentridae dominated the ichthyofauna in terms of species richness and Pomacentridae were most abundant. Neither diversity nor species richness was correlated to depth. The abundance of fishes was higher at the deep reef slope, due to schooling planktivorous fishes. At 12 m depth abundance of fishes at the seagrass-dominated site was higher than on the coral reefs. Multivariate analysis demonstrated a strong influence on the fish assemblages by depth and benthic habitat. Fish species richness was positively correlated with hard substrate cover and habitat diversity. Abundance of corallivores was positively linked with live hard coral cover. The assemblages of fishes were different on the shallow reef slope, deep reef slope and seagrass meadows. An analysis of the fish fauna showed that the Gulf of Aqaba harbours a higher species richness than previously reported. The comparison with fish communities on other reefs around the Arabian Peninsula and Indian Ocean supported the recognition of an Arabian subprovince within the Indian Ocean. The affinity of the Arabian Gulf ichthyofauna to the Red Sea is not clear. Received in revised form: 2 November 2001 Electronic Publication     

Comparative character of the deep-water and inshore cottoid fishes endemic to Lake Baikal

Lake Baikal's 29 endemic species of cottoid fishes form three groups: shallow-water species in depths to 350 m; eurybathic species from 50 to 1300 m; and abyssal species from 400 to 1600 m. These groups differ in their abilities to withstand high hydrostatic pressure. As in marine deep-water fishes, abyssal cottoids in Baikal have few or no cones in the retina, and some have tubular eyes. Their seismosensory systems predominate, based chiefly on free neuromasts. The proportion of species with canal systems decreases with depth. Diversity of the predominantly gammarid foods also decreases from 45 species in shallow water to five species in deep water, and the lateral line system plays the dominant role in food detection at all depths. Two abyssal cottoids have become secondary pelagic, achieving close to neutral buoyancy through high lipid levels and reduced skeletal mineralization. These forms take advantage of the abundant pelagic planktonic amphipod populations. The adaptations of abyssal forms parallel those seen in deep-water marine fishes.