A new theoretical performance landscape for suction feeding reveals adaptive kinematics in a natural population of reef damselfish.

Understanding how organismal traits determine performance and, ultimately, fitness is a fundamental goal of evolutionary eco-morphology. However, multiple traits can interact in non-linear and context-dependent ways to affect performance, hindering efforts to place natural populations with respect to performance peaks or valleys. Here, we used an established mechanistic model of suction-feeding performance (SIFF) derived from hydrodynamic principles to estimate a theoretical performance landscape for zooplankton prey capture. This performance space can be used to predict prey capture performance for any combination of six morphological and kinematic trait values. We then mapped in situ high-speed video observations of suction feeding in a natural population of a coral reef zooplanktivore, Chromis viridis, onto the performance space to estimate the population's location with respect to the topography of the performance landscape. Although the kinematics of the natural population closely matched regions of high performance in the landscape, the population was not located on a performance peak. Individuals were furthest from performance peaks on the peak gape, ram speed and mouth opening speed trait axes. Moreover, we found that the trait combinations in the observed population were associated with higher performance than expected by chance, suggesting that these combinations are under selection. Our results provide a framework for assessing whether natural populations occupy performance optima.

Trophic separation in planktivorous reef fishes: a new role for mucus?

The feeding apparatus directly influences a species' trophic ecology. In fishes, our understanding of feeding modes is largely derived from studies of rigid structures (i.e. bones, teeth, gill rakers). A recently described lip innovation, however, highlighted the role of soft anatomy in enabling specialized feeding modes. In this study, we explore whether similar diversification may also occur in the soft anatomy of the buccal cavity. Using four key anatomical traits to classify 19 species (14 genera) of wrasses, we evaluated the relationship between anatomical specialization of the buccal cavity and diet. Our data revealed a previously undocumented anatomical adaptation in the mouths of fairy wrasses (Cirrhilabrus): the mucosa throughout the buccal cavity (i.e. anterior to the pharynx) is packed with goblet cells, enabling it to secrete large quantities of mucus in this region; a new trait that, until now, had not been documented in wrasses. This disparity reflects diet differences, with mucus secretion found only in planktivorous Cirrhilabrus that feed predominantly on amorphous organic material (potentially gelatinous organisms). This suggests a cryptic mucus-based resource partitioning in planktivorous wrasses.

Kleptopredation: a mechanism to facilitate planktivory in a benthic mollusc.

Predation occurs when an organism completely or partially consumes its prey. Partial consumption is typical of herbivores but is also common in some marine microbenthic carnivores that feed on colonial organisms. Associations between nudibranch molluscs and colonial hydroids have long been assumed to be simple predator-prey relationships. Here we show that while the aeolid nudibranch Cratena peregrina does prey directly on the hydranths of Eudendrium racemosum, it is stimulated to feed when hydranths have captured and are handling prey, thus ingesting recently captured plankton along with the hydroid polyp such that plankton form at least half of the nudibranch diet. The nudibranch is thus largely planktivorous, facilitated by use of the hydroid for prey capture. At the scale of the colony this combines predation with kleptoparasitism, a type of competition that involves the theft of already-procured items to form a feeding mode that does not fit into existing classifications, which we term kleptopredation. This strategy of subsidized predation helps explain how obligate-feeding nudibranchs obtain sufficient energy for reproduction from an ephemeral food source.

Response of plankton to nutrients, planktivory and terrestrial organic matter: a model analysis of whole-lake experiments.

Terrestrial organic matter can be assimilated by aquatic consumers but implications for biomass and production are unresolved. An ecosystem model was fit to estimate effects of phosphorus (P) load, planktivory, and supply rate of terrestrial particulate organic carbon (TPOC) on phytoplankton and zooplankton in five whole-lake experiments. Phytoplankton biomass increased with P load and planktivory and decreased with TPOC supply rate. Zooplankton biomass increased with P load and responded weakly to planktivory and TPOC supply rate. Zooplankton allochthony (proportion of carbon from terrestrial sources) decreased with P load and planktivory and increased with TPOC supply rate. Lakes with low allochthony (< 0.3) had wide ranges of phytoplankton and zooplankton biomass and production, depending on P load and planktivory. Lakes with high allochthony (> 0.3) had low biomass and production of both phytoplankton and zooplankton. In summary, terrestrial OC inhibits primary production and is a relatively low-quality food source for zooplankton.

Can young-of-the-year invasive fish keep up with young-of-the-year native fish? A comparison of feeding rates between invasive sticklebacks and whitefish.

Invasion of non-native species might alter food web structure and the strength of top-down control within lake ecosystems. As top-down control exerted by fish populations is often dominated by young of the year fish, the impact of new fish species might depend on the feeding rates of the juvenile fish. Here we provide comparative analyses of feeding rates of juvenile whitefish (Coregonus wartmanni) - a native and specialised planktivore and an invasive generalist (sticklebacks, Gasterosteus aculeatus). We studied feedings rates of whitefish and sticklebacks in aquaria experiments using 2 cm to 8 cm fish feeding on seven zooplankton species common to Lake Constance. As whitefish hatch several months earlier than sticklebacks, 0+ whitefish are larger than 0+ sticklebacks throughout the year and hence are predicted to have higher feeding rates on especially large zooplankton species. We show that sticklebacks as small as 2 cm were able to feed on the largest zooplankton species of Lake Constance. Further, stickleback feeding rates were similar to both the same size 0+ whitefish and the larger 0+ whitefish co-occurring with smaller 0+ sticklebacks. Hence, 0+ sticklebacks will compete with 0+ whitefish for the same zooplankton species, therefore the invasion of sticklebacks is unlikely to change the relative feeding pressure by individual 0+ fish on zooplankton species.

Phylogenetic perspectives on reef fish functional traits.

Functional traits have been fundamental to the evolution and diversification of entire fish lineages on coral reefs. Yet their relationship with the processes promoting speciation, extinction and the filtering of local species pools remains unclear. We review the current literature exploring the evolution of diet, body size, water column use and geographic range size in reef-associated fishes. Using published and new data, we mapped functional traits on to published phylogenetic trees to uncover evolutionary patterns that have led to the current functional diversity of fishes on coral reefs. When examining reconstructed patterns for diet and feeding mode, we found examples of independent transitions to planktivory across different reef fish families. Such transitions and associated morphological alterations may represent cases in which ecological opportunity for the exploitation of different resources drives speciation and adaptation. In terms of body size, reconstructions showed that both large and small sizes appear multiple times within clades of mid-sized fishes and that extreme body sizes have arisen mostly in the last 10 million years (Myr). The reconstruction of range size revealed many cases of disparate range sizes among sister species. Such range size disparity highlights potential vicariant processes through isolation in peripheral locations. When accounting for peripheral speciation processes in sister pairs, we found a significant relationship between labrid range size and lineage age. The diversity and evolution of traits within lineages is influenced by trait-environment interactions as well as by species and trait-trait interactions, where the presence of a given trait may trigger the development of related traits or behaviours. Our effort to assess the evolution of functional diversity across reef fish clades adds to the burgeoning research focusing on the evolutionary and ecological roles of functional traits. We argue that the combination of a phylogenetic and a functional approach will improve the understanding of the mechanisms of species assembly in extraordinarily rich coral reef communities.

Visual sensitivity and spatial resolution of the planktivorous fish, Atherinomorus forskalii (Atherinidae; Rüppell, 1838), to a polarized grating.

Polarized light detection has been documented in only a small number of fish species. The benefit of polarization vision for fish is not fully understood, nor is the transduction mechanism that underlies it. Past studies proposed that one possible advantage of polarization vision is that it enhances the contrast of zooplankton targets by breaking their transparency. Here, we used an optomotor apparatus to test the responses of the planktivorous Hardyhead silverside fish Atherinomorus forskalii (Atherinidae) to vertical unpolarized (intensity) and polarized gratings. We also tested and compared the spatial and temporal resolutions of A. forskalii in the intensity and polarization domains. A. forskalii responded to the polarization pattern, but only under illumination that included ultraviolet-blue (λ>380nm) wavelengths. The spatial resolution of A. forskalii was measured as a minimum separable angle of 0.57° (a 1-mm prey viewed from 100-mm distance). The temporal resolution to unpolarized vs. polarized gratings was constant, at 33 and 10Hz respectively at most of the stripe widths tested. At the smallest stripe width tested (1mm=the minimal separable angle), which correlates with the size of prey typically consumed by these fish, the temporal resolution to the polarized grating increased to 42Hz. We conclude that A. forskalii is polarization sensitive, may use polarization vision to improve detection of its planktonic prey, and that polarization may be perceived by the fish via a separate visual pathway than intensity.

Fatty Acid Composition and Contents of Seven Commercial Fish Species of Genus Coregonus from Russian Subarctic Water Bodies.

In several Russian northern lakes and rivers, Arctic cisco Coregonus autumnalis, least cisco C. sardinella, peled C. peled, tugun C. tugun, broad whitefish C. nasus, whitefish C. lavaretus and vendace C. albula were sampled in periods of officially permitted commercial fishery. Special attention was paid to contents (mg g-1 of wet weight) of eicosapentaenoic acid (EPA) and docosahexaenoic acid (DHA) in muscle tissues (filets), which are essential for human nutrition. The highest values of EPA + DHA content in semi-anadromous fish and freshwater fish were recorded for C. autumnalis from the Yenisei River, 17.60 mg g-1 wet weight, and for C. lavaretus from the Sobachye Lake, 16.61 mg g-1 wet weight, respectively. Intra-genus variations of EPA + DHA contents of Coregonus species were from 1.87 to 17.60 mg g-1 wet weight. Since the congeneric species were genetically close to each other, the variations in EPA and DHA contents were thought to be caused primarily by ecological factors: migrational capability, type of feeding and trophic status of aquatic ecosystems. In general, the majority of studied species appeared to be of a high nutritive value for humans, although unfavorable environmental conditions could considerably diminish this value.

Morphological and behavioral antipredatory adaptations of decapod zoeae.

Zoeae of some species of estuarine decapods are retained in the estuary throughout development while others are exported into nearshore coastal waters. The horizontal migrations of decapod zoeae to coastal waters may have evolved to reduce the probability of encountering planktivorous fishes which are most abundant in the estuary. If so, then the morphological vulnerability of zoeae to fish predation should be inversely related to the number of predators occurring where they develop. Six species of estuarine decapod zoeae were offered to Menidia menidia and Fundulus heteroclitus. The behavioral interactions were observed to determine the prey's vulnerability to predation, and the mode of operation and relative effectiveness of their defenses. Feeding trials and behavioral observations both demonstrated that M. menidia 6-16 mm long preferred Uca minax and Callinectes sapidus zoeae, which are exported from the estuary, to Rhithropanopeus harrisii, Sesarma reticulatum and Palaemonetes pugio, which are retained within estuaries. Pinnotheres ostreum zoeae develop in the lower estuary and fish demonstrated an intermediate preference for the zoeae. Menidia menidia 20-40 mm long showed similar preferences for R. harrisii S. reticulatum, P. ostreum and U. minax as did small silversides. Large-mouthed demersal fish, Fundulus heteroclitus 6-10 mm long, also preferred U. minax to R. harrisii, but more readily preyed on zoeae than did M. menidia. The exported species of zoeae have shorter spines and smaller bodies than do retained zoeae, except P. ostreum which is small, spineless and passively sinks when attacked by fish. Other retained species of zoeae also have postcontact behavioral defenses which enhance the effectiveness of their morphological defenses. Zoeae do not evade attacks by fishes, but fishes quickly learned to avoid zoeae, which increases the effectiveness of the zoeae's antipredatory adaptations.

The effects on the plankton community of filter-feeding Sacramento blackfish, Orthodon microlepidotus.

A series of mesocosm experiments was performed to assess the effects on the plankton community of filter-feeding Sacramento blackfish (Cyprinidae; Orthodon microlepidotus). Phytoplankton size-frequency distribution, zooplankton abundance, primary production, potential secondary production, and nutrient concentrations were measured. Blackfish reduce numbers of both evasive and nonevasive zooplankton and large phytoplankton while enhancing nanoplankton densities. Blackfish also significantly increase primary production and potential secondary community production. Levels of dissolved inorganic phosphorus and ammonia-nitrogen also increase. The effects of blackfish are generally similar to those reported for other filter-feeding fish.

Size-dependent foraging efficiency, cannibalism and zooplankton community structure.

To examine size-dependent food web interactions in systems with cannibalism, we compared the abundances of zooplankton and phytoplankton over 2 years in four lakes with cannibalistic perch (Perca fluviatilis) of which two also supported the top predator pike (Esox lucius). The abundance of perch 2 years and older was lower in lakes with pike than in lakes with only perch. In contrast, the abundance of small perch (young-of-the-year and 1-year old) was lower in lakes with only perch suggesting that intense cannibalism reduced these size classes to low levels in lakes lacking pike. Functional response experiments with differently sized perch and zooplankton showed that the attack rate of small perch susceptible to cannibalism was much higher than that of large cannibalising perch. The optimal body size of perch with respect to attack rate was also lower for small zooplankton prey than for large zooplankton. The zooplankton communities in lakes with only perch were dominated by the relatively small species Ceriodaphnia quadrangula and Bosmina spp. and total zooplankton biomass was higher in these lakes than in lakes with both pike and perch. In contrast, the mean size of cladoceran zooplankton was largest in lakes with both pike and perch owing to a dominance of the large zooplankton species Holopedium gibberum in these lakes. We relate these patterns to (1) the low foraging efficiency of large perch on small zooplankton and (2) the low abundance of small zooplanktivorous perch (due to cannibalism) in lakes with only perch. The differences in zooplankton community structure also resulted in different seasonal dynamics of phytoplankton between lakes. Cannibalism introduces a vertical heterogeneity to food webs that causes consumer-resource dynamics that are not predictable from linear food chain models.