Jaw lever analysis of Hawaiian gobioid stream fishes: A simulation study of morphological diversity and functional performance

Differences in feeding behavior and performance among the five native Hawaiian gobioid stream fishes (Sicyopterus stimpsoni, Lentipes concolor, Awaous guamensis, Stenogobius hawaiiensis, and Eleotris sandwicensis) have been proposed based on the skeletal anatomy of their jaws and dietary specialization. However, performance of the feeding apparatus likely depends on the proportions and configurations of the jaw muscles and the arrangement of the jaw skeleton. We used a published mathematical model of muscle function to evaluate potential differences in jaw closing performance and their correlations with morphology among these species. For example, high output force calculated for the adductor mandibulae muscles (A2 and A3) of both A. guamensis and E. sandwicensis matched expectations based on the morphology of these species because these muscles are larger than in the other species. In contrast, Stenogobius hawaiiensis exhibited an alternative morphological strategy for achieving high relative output forces of both A2 and A3, in which the placement and configuration of the muscles conveyed high mechanical advantage despite only moderate cross‐sectional areas. These differing anatomical pathways to similar functional performance suggest a pattern of many‐to‐one mapping of morphology to performance. In addition, a functional differentiation between A2 and A3 was evident for all species, in which A2 was better suited for producing forceful jaw closing and A3 for rapid jaw closing. Thus, the diversity of feeding performance of Hawaiian stream gobies seems to reflect a maintenance of functional breadth through the retention of some primitive traits in combination with novel functional capacities in several species. J. Morphol. 2009. © 2009 Wiley‐Liss, Inc.     

Predicting ecological and phenotypic differentiation in the wild: a case of piscivorous fish in a fishless environment

Environmental variation drives ecological and phenotypic change. How predictable is differentiation in response to environmental change? Answering this question requires the development and testing of multifarious a priori predictions in natural systems. We employ this approach using Gobiomorus dormitor populations that have colonized inland blue holes differing in the availability of fish prey. We evaluated predictions of differences in demographics, habitat use, diet, locomotor and trophic morphology, and feeding kinematics and performance between G. dormitor populations inhabiting blue holes with and without fish prey. Populations of G. dormitor independently diverged between prey regimes, with broad agreement between observed differences and a priori predictions. For example, in populations lacking fish prey, we observed male‐biased sex ratios, a greater use of shallow‐water habitat, and larger population diet breadths as a result of greater individual diet specialization. Furthermore, we found predictable differences in body shape, mouth morphology, suction generation capacity, strike kinematics, and feeding performance on different prey types, consistent with the adaptation of G. dormitor to piscivory when coexisting with fish prey and to feeding on small invertebrates in their absence. The results of the present study suggest great potential in our ability to predict population responses to changing environments, which is an increasingly important capability in a human‐dominated, ever‐changing world. © 2015 The Linnean Society of London, Biological Journal of the Linnean Society, 2015, 114 , 588–607.     

Jaw muscle fiber type distribution in Hawaiian gobioid stream fishes: histochemical correlations with feeding ecology and behavior

Differences in fiber type distribution in the axial muscles of Hawaiian gobioid stream fishes have previously been linked to differences in locomotor performance, behavior, and diet across species. Using ATPase assays, we examined fiber types of the jaw opening sternohyoideus muscle across five species, as well as fiber types of three jaw closing muscles (adductor mandibulae A1, A2, and A3). The jaw muscles of some species of Hawaiian stream gobies contained substantial red fiber components. Some jaw muscles always had greater proportions of white muscle fibers than other jaw muscles, independent of species. In addition, comparing across species, the dietary generalists (Awaous guamensis and Stenogobius hawaiiensis) had a lower proportion of white muscle fibers in all jaw muscles than the dietary specialists (Lentipes concolor, Sicyopterus stimpsoni, and Eleotris sandwicensis). Among Hawaiian stream gobies, generalist diets may favor a wider range of muscle performance, provided by a mix of white and red muscle fibers, than is typical of dietary specialists, which may have a higher proportion of fast-twitch white fibers in jaw muscles to help meet the demands of rapid predatory strikes or feeding in fast-flowing habitats.     

The Effects of Satiation on Strike Mode and Prey Capture Kinematics in the Largemouth bass, Micropterus salmoides

The feeding mechanism and kinematics of prey capture have been studied in many fishes. However, the effects of satiation on the strike mode and prey capture kinematics have never been directly measured. We analyze 12 kinematic variables to determine the effects of satiation on prey capture in five largemouth bass, Micropterus salmoides, by using high speed videography. We also present the first experimental test for modulatory capabilities in response to satiation, by using the ram-suction index. Significant changes in the kinematic variables of maximum lower jaw depression, maximum gape distance, maximum hyoid depression, time to maximum hyoid depression, and time from maximum hyoid depression to recovery were seen with the effects of satiation. Change in the kinematic variables imply a decrease in jaw opening velocity and the magnitude of suction velocity created during repetitive strikes by M. salmoides with increasing satiation. The bass primarily uses a ram strike mode, with some suction bites occasionally. Ram-suction index analyses suggests that M. salmoides does not modulate strike mode in response to satiation. However, the bass modulate prey capture kinematics without altering strike mode with the effects of satiation. Prey capture success decreases in each bass, as the probability of a successful prey capture event becomes lower, with increasing satiation. These findings demonstrate that satiation can have major effects on prey capture kinematics and future studies of feeding kinematics should account for satiation in their analyses.     

Zooplankton capture by a coral reef fish: an adaptive response to evasive prey

Synopsis High-speed cinematography and video using modified Schlieren optics and laser illumination helped elicit details of prey capture mechanisms used by Chromis viridis while feeding on calanoid copepods and Artemia. Chromis viridis is capable of a ram-jaw, low-suction feeding, as well as a typical suction feeding behavior described for other species of planktivores. By adjusting the degree of jaw protrusion and amount of suction used during a feeding strike, this fish can modulate its feeding strikes according to the prey type being encountered. The ram-jaw feeding mode enables C. viridis to capture highly evasive calanoid copepods within 6 to 10 msec. The use of specialized feeding behavior for evasive prey and the ability to vary feeding behavior are adaptations for feeding on evasive prey.     

Feeding kinematics and performance of Hawaiian stream gobies,Awaous guamensis and Lentipes concolor: Linkage of functional morphology and ecology

Distributions of Hawaiian stream fishes are typically interrupted by waterfalls that divide streams into multiple segments. Larvae hatch upstream, are flushed into the ocean, and must climb these waterfalls to reach adult habitats when returning back to freshwater as part of an amphidromous life cycle. Stream surveys and studies of climbing performance show that Lentipes concolor Gill can reach fast‐flowing upper stream segments but that Awaous guamensis Valenciennes reaches only slower, lower stream segments. Gut content analyses for these two species indicate considerable overlap in diet, suggesting that feeding kinematics and performance of these two species might be comparable. Alternatively, feeding kinematics and performance of these species might be expected to differ in relation to the different flow regimes in their habitat (feeding in faster stream currents for L. concolor versus in slower currents for A. guamensis). To test these alternative hypotheses, we compared food capturing kinematics and performance during suction feeding behaviors of A. guamensis and L. concolor using morphological data and high‐speed video. Lentipes concolor showed both a significantly larger gape angle and faster jaw opening than A. guamensis. Geometric models calculated that despite the inverse relationship of gape size and suction pressure generation, the fast jaw motions of L. concolor allow it to achieve higher pressure differentials than A. guamensis. Such elevated suction pressure would enhance the ability of L. concolor to successfully capture food in the fast stream reaches it typically inhabits. Differences in jaw morphology may contribute to these differences in performance, as the lever ratio for jaw opening is about 10% lower in L. concolor compared with A. guamensis, suiting the jaws of L. concolor better for fast opening. J. Morphol. 2009. © 2008 Wiley‐Liss, Inc.     

Genetic differentiation,speciation, and phylogeography of cactus flies (Diptera: Neriidae: Odontoloxozus) from Mexico and south‐western USA

Nucleotide sequences from the mitochondrial cytochrome c oxidase subunit I (COI) gene, comprising the standard barcode segment, were used to examine genetic differentiation, systematics, and population structure of cactus flies (Diptera: Neriidae: Odontoloxozus) from Mexico and south‐western USA. Phylogenetic analyses revealed that samples of Odontoloxozus partitioned into two distinct clusters: one comprising the widely distributed Odontoloxozus longicornis (Coquillett) and the other comprising Odontoloxozus pachycericola Mangan & Baldwin, a recently described species from the Cape Region of the Baja California peninsula, which we show is distributed northward to southern California, USA. A mean Kimura two‐parameter genetic distance of 2.8% between O. longicornis and O. pachycericola, and eight diagnostic nucleotide substitutions in the COI gene segment, are consistent with a species‐level separation, thus providing the first independent molecular support for recognizing O. pachycericola as a distinct species. We also show that the only external morphological character considered to separate adults of the two species (number of anepisternal bristles) varies with body size and is therefore uninformative for making species assignments. Analysis of molecular variance indicated significant structure among populations of O. longicornis from three main geographical areas, (1) Arizona, USA and Sonora, Mexico; (2) Santa Catalina Island, California, USA; and (3) central Mexico (Querétaro and Guanajuato), although widely‐separated populations from Arizona and Sonora showed no evidence of structure. A TCS haplotype network showed no shared haplotypes of O. longicornis among the three main regions. The potential roles of vicariance and isolation‐by‐distance in restricting gene flow and promoting genetic differentiation and speciation in Odontoloxozus are discussed. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 110 , 245–256.     

A Comparison of Foraging Behavior in Two “Percher” Dragonflies, Pachydiplax longipennis and Erythemis simplicicollis (Odonata: Libellulidae)

Feeding behaviors of two adult libellulid percher dragonflies, Pachydiplax longipennis and Erythemis simplicicollis, were compared. All results pertain to feeding from natural perches located some distance away from reproductive rendezvous sites. Compared to P. longipennis, E. simplicicollis chose broader and less structurally discrete perches, moved more frequently and over a larger area, and took, on average, much larger prey, although diet overlapped broadly in the two species. Erythemis made more frequent feeding flights but with a much lower success rate than Pachydiplax; consequently the prey capture rate was similar in the two species. Gut contents of mature, but not of immature, E. simplicicollis comprised a significantly greater proportion of body mass than in P. longipennis, apparently confirming the importance of larger prey in the diet of the former.     

Effects of prey density on nocturnal zooplankton predation throughout the ontogeny of juvenile <Emphasis Type="Italic">Platax orbicularis</Emphasis> (Teleostei: Ephippidae)

Juvenile Platax orbicularis switches foraging tactics from diurnal herbivory to nocturnal zooplanktivory within a day. To examine how juvenile fish actively feed on zooplankton prey during nighttime, a field-recorded video analysis was conducted in the reefs off Kuchierabu-jima Island, southern Japan. Juveniles consistently showed fast and sudden attacks that were accurately directed at individual zooplankton prey, and changed feeding frequencies with different prey densities. A negative relationship was observed between feeding frequency and prey density, with higher feeding frequencies occurring at lower prey concentrations, implying a disturbance effect of clouded zooplankton prey on the juvenile fish. A clear transition from a ram-based to a suction-based feeding mode was observed with fish size, suggesting that changes in the feeding behaviors occur even in juveniles fishes, without drastic morphological changes.     

Within-colony feeding selectivity by a corallivorous reef fish: foraging to maximize reward?

Foraging theory predicts that individuals should choose a prey that maximizes energy rewards relative to the energy expended to access, capture, and consume the prey. However, the relative roles of differences in the nutritive value of foods and costs associated with differences in prey accessibility are not always clear. Coral‐feeding fishes are known to be highly selective feeders on particular coral genera or species and even different parts of individual coral colonies. The absence of strong correlations between the nutritional value of corals and prey preferences suggests other factors such as polyp accessibility may be important. Here, we investigated within‐colony feeding selectivity by the corallivorous filefish, Oxymonacanthus longirostris, and if prey accessibility determines foraging patterns. After confirming that this fish primarily feeds on coral polyps, we examined whether fish show a preference for different parts of a common branching coral, Acropora nobilis, both in the field and in the laboratory experiments with simulated corals. We then experimentally tested whether nonuniform patterns of feeding on preferred coral species reflect structural differences between polyps. We found that O. longirostris exhibits nonuniform patterns of foraging in the field, selectively feeding midway along branches. On simulated corals, fish replicated this pattern when food accessibility was equal along the branch. However, when food access varied, fish consistently modified their foraging behavior, preferring to feed where food was most accessible. When foraging patterns were compared with coral morphology, fish preferred larger polyps and less skeletal protection. Our results highlight that patterns of interspecific and intraspecific selectivity can reflect coral morphology, with fish preferring corals or parts of coral colonies with structural characteristics that increase prey accessibility.     

Migratory flexibility in native Hawai'ian amphidromous fishes

We assessed the prevalence of life history variation across four of the five native amphidromous Hawai'ian gobioids to determine whether some or all exhibit evidence of partial migration. Analysis of otolith Sr.: Ca concentrations affirmed that all are amphidromous and revealed evidence of partial migration in three of the four species. We found that 25% of Lentipes concolor (n = 8), 40% of Eleotris sandwicensis (n = 20) and 29% of Stenogobius hawaiiensis (n = 24) did not exhibit a migratory life-history. In contrast, all individuals of Sicyopterus stimpsoni (n = 55) included in the study went to sea as larvae. Lentipes concolor exhibited the shortest mean larval duration (LD) at 87 days, successively followed by E. sandwicensis (mean LD = 102 days), S. hawaiiensis (mean LD = 114 days) and S. stimpsoni (mean LD = 120 days). These findings offer a fresh perspective on migratory life histories that can help improve efforts to conserve and protect all of these and other at-risk amphidromous species that are subject to escalating anthropogenic pressures in both freshwater and marine environments.     

Diet and prey selection in larval and juvenile Japanese anchovy Engraulisjaponicus in Ariake Bay, Japan

We studied the diet of larval and juvenile Japanese anchovy Engraulis japonicus in the upper Ariake Bay, Japan. Diet was analyzed by examining the digestive tracts; feeding intensity, proportion of empty guts, and prey selectivity were calculated. Anchovy density was negatively influenced by temperature and positively by salinity and prey density. Diet was dominated by Acartia omorii, which was positively selected with two other copepods, Calanus sinicus and Pseudodiaptomus marinus. In contrast, Oithona davisae was highly dominant in the environment but was absent in anchovy guts; thus, this copepod was negatively selected, with two others, Tortanus derjugini and Sinocalanus sinensis. Overall, larger prey were positively selected and smaller ones were negatively selected; value of electivity index correlated negatively with prey size. Larvae [<18 mm of standard length (SL)] showed significantly lower feeding intensities and higher rates of empty guts than juveniles (≥18 mm SL). In juveniles, feeding intensity increased steadily as the fish grew in size, with a corresponding reduction in empty guts. Feeding intensity correlated positively and empty gut correlated negatively with fish size. We suggest that larger prey are important diets for postlarval Japanese anchovy in Ariake Bay.     

Revision of the Euthalia phemius complex (Lepidoptera: Nymphalidae) based on morphology and molecular analyses

Adults of the Euthalia phemius complex, which is composed of three South‐East Asian nymphalid species, Euthalia phemius, Euthalia ipona, and Euthalia euphemia, were genetically analysed by examining mitochondrial and nuclear genes. The E. phemius complex was also examined morphologically, with particular emphasis on wing markings and male genitalia. No significant differences amongst the three species in the complex were detected with respect to either genetic distance or genital morphology. We therefore conclude that the three currently recognized Euthalia species belong to a single species. Accordingly, E. ipona is synonymized with E. phemius. Euthalia euphemia is treated as a subspecies of E. phemius. Type specimens of all taxa and a synonymic list for the E. phemius complex are also given. In addition, we briefly discuss the evolution and biogeography of the species complex. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2012, 164 , 304–327.     

Modulation in Feeding Kinematics and Motor Pattern of the Nurse Shark <Emphasis Type="Italic">Ginglymostoma cirratum</Emphasis>

Synopsis Studies of feeding in bony fishes have almost universally demonstrated the ability of individuals to modulate their method of capture in response to differing stimuli. Preliminary evidence indicates that morphologically specialized inertial suction feeding sharks are the most likely fishes to lack inherent modulatory ability. We examined the ability of the nurse shark, Ginglymostoma cirratum, to modulate its feeding behavior based on different food types and sizes. G. cirratum is an inertial suction feeding fish that is apparently stereotyped in its food capture behavior. Electromyography showed no statistical difference between feeding motor patterns based on food type (squid or fish) or size (gape width or twice gape width), although there were slight inter-individual differences in the onset of muscle firing for some muscles. Kinematic analysis showed a statistical difference in variables associated with durations for different food types, with the durations for all variables being faster for squid bites than fish bites, but no difference based on the size of the food item. This apparent lack of modulation may be associated with specialization of the morphology and behavior of G. cirratum for obligate suction prey capture. This functional specialization constrains the method in which G. cirratum captures prey but does not appear to result in dietary specialization. An unusual post capture spit-suck manipulation allows this shark to handle and ingest large prey.     

Prey capture and processing behaviors vary with prey size and shape in Australian and subantarctic fur seals

When hunting at sea, pinnipeds should adapt their foraging behaviors to suit the prey they are targeting. We performed captive feeding trials with two species of otariid seal, Australian fur seals (Arctocephalus pusillus doriferus) and subantarctic fur seals (Arctocephalus tropicalis). This allowed us to record detailed observations of how their foraging behaviors vary when presented with prey items that cover the full range of body shapes and sizes encountered in the wild. Small prey were captured using suction alone, while larger prey items were caught in the teeth using raptorial biting. Small fish and long skinny prey items could then be swallowed whole or processed by shaking, while all prey items with body depths greater than 7.5 cm were processed by shaking at the water's surface. This matched opportunistic observations of feeding in wild Australian fur seals. Use of “shake feeding” as the main prey processing tactic also matches predictions that this method would be one of the only tactics available to aquatic tetrapods that are unable to secure prey using their forelimbs.     

Growth and morphological development of laboratory-reared larval and juvenile climbing perch <Emphasis Type="Italic">Anabas testudineus</Emphasis>

Morphological development, including fin and labyrinth organ, body proportions and pigmentation, in laboratory-reared larval and juvenile climbing perch Anabas testudineus was described and behavioral features under rearing condition were observed. Body lengths (BL) of larvae and juveniles were 1.9 ± 0.1 (mean ± SD) mm just after hatching (day-0), 8.7 ± 1.3 mm on day-19, reaching 18.4 ± 2.1 mm on day-35 after hatching. Aggregate fin ray numbers attained full complements in juveniles larger than 8.3 mm BL. Preflexion larvae started feeding on day-2 following formation of the upper and lower jaws, the yolk being completely absorbed by day-7 after hatching. Teeth appeared in flexion larvae larger than 5 mm BL on day-6, with cannibalism starting shortly after and continuing with further growth. Melanophores on the body increased with growth, a large dark spot developing on the lateral midline around caudal margin of the body in the postflexion and juvenile stages. The labyrinth organ differentiated in postflexion larvae larger than 7.2 mm BL on day-16, with air-breathing starting at the same time. Body proportions attained constant in postflexion larvae larger than 7.0 mm BL, and habitat of fish shifted from bottom to mid-layer. With the exception of fin ray numbers, the above morphological developments corresponded to behavioral shifts that occurred in the postflexion stage (ca. 7 mm BL), their subsequent continuity illustrating that the species possessed most juvenile-equivalent functions from ca. 7 mm BL.     

Stable Isotope Analysis of Amphidromous Hawaiian Gobies Suggests Their Larvae Spend a Substantial Period of Time in Freshwater River Plumes

Synopsis We employed stable isotope analysis (δ¹³C, δ¹⁵N) to evaluate the sources of nutrients used by amphidromous gobiid fishes (Lentipes concolor, Sicyopterus stimpsoni, Awaous guamensis) caught migrating into and living in Hakalau Stream, Hawaii. Although considerable variation amongst the stable isotope values of stream items was noted across all 4 years of our study, the relationships between the fishes were relatively constant. Stable isotope values of recruiting gobies were consistently closer to those of both inshore plankton and freshwater adults than those of offshore plankton, suggesting that the larvae of these species derive much of their nutrition from inshore environments influenced by fresh water. Small differences between the stable values of these species further suggested that their larvae come from different inshore locations. After entering fresh water all species appear to swim rapidly upstream without feeding. Finally, once well upstream, adult L. concolor and A. guamensis appear to assume an omnivorous diet while adult S. stimpsoni rely upon autochthonous production within streams. We propose that freshwater food webs play an integral yet complex role in the lives of both larval and adult amphidromous Hawaiian fishes.     

Anatomy,histology, and systematic implications of the head ornamentation in the males of four species of Limnonectes (Anura: Dicroglossidae)

The males of four species of the Asian frog genus Limnonectes [Limnonectes dabanus (Smith, 1922a), Limnonectes gyldenstolpei (Andersson, 1916), Limnonectes macrognathus (Boulenger, 1917), and Limnonectes plicatellus (Stoliczka, 1873)] exhibit remarkable ornamentation in the form of a swollen, or cap‐like, structure (caruncle) on the top of their heads. These caruncles vary in their appearance among species, and neither their function nor their actual systematic value is known. We compared their anatomy via dissections, morphometrics, radiography, and histology, and analysed the available mitochondrial DNA sequences as well as new data to place these species within the context of a larger phylogenetic hypothesis for Limnonectes. Despite the externally different morphology, the underlying histological structure is virtually identical. Beneath skin that is densely packed with mucous glands lies a pad of connective tissue overlaying the parietal bone. The actual function of the caruncle, however, remains enigmatic. In addition to the presence of the caruncle, independent evidence from osteological characters and molecular data support the monophyly of a clade comprising of L. dabanus, L. gyldenstolpei, L. macrognathus, and L. plicatellus. The caruncles are therefore interpreted as a robust autapomorphy for this clade, and suggest that the subgenus Elachyglossa should be restricted to the four species in question. © 2014 The Linnean Society of London     

Patterns of morphological evolution of the cephalic region in damselfishes (Perciformes: Pomacentridae) of the Eastern Pacific

Pomacentridae are one of the most abundant fish families inhabiting reefs of tropical and temperate regions. This family, comprising 29 genera, shows a remarkable diversity of habitat preferences, feeding, and behaviours. Twenty‐four species belonging to seven genera have been reported in the Eastern Pacific region. The present study focuses on the relationship between the diet and the cephalic profile in the 24 endemic damselfishes of this region. Feeding habits were determined by means of underwater observations and the gathering of bibliographic data. Variations in cephalic profile were analyzed by means of geometric morphometrics and phylogenetic methods. The present study shows that the 24 species can be grouped into three main trophic guilds: zooplanktivores, algivores, and an intermediate group feeding on small pelagic and benthic preys. Shape variations were low within each genus except for Abudefduf. Phylogenetically adjusted regression reveals that head shape can be explained by differences in feeding habits. The morphometric phylogeny recovered the subfamily Stegastinae and the relationship between Abudefduf troschelii and Chromis species. The cephalic profile of damselfishes contains a clear and strong phylogenetic signal. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 102 , 593–613.     

Planktivory in Pagothenia borchgrevinki (Pisces: Nototheniidae) in McMurdo sound,Antarctica

Summary Specimens of the Antarctic fish P. borchgrevinki were collected from just beneath the sea-ice in McMurdo Sound over the period 8 November to 6 December, 1985. The stomach contents of 200 fish were analysed and compared with the co-occurrence of the prey species in the plankton. Numerically, the dominant prey was a thecosomatid pteropod Limacina helicina (83%), followed by a hyperiid amphipod Hyperiella dilatata (5%). Other prey species in decreasing order of occurrence were Euphausia crystallorophias, Euchaeta antarctica, other copepods, a decapod crustacean larva, chaetognaths, the amphipods Orchomene plebs and Epimiriella macronyx, and unidentified juvenile fish. The order of frequency of occurrence of these prey in the guts of all the fish was H. dilatata (in 73% of the fish), L. helicina (71%), calanoid copepods (55%), chaetognaths (51%), E. crystallorophias (42%), decapod crustacean larva (32%), O. plebs (27%), juvenile fish (20%), and E. macronyx (14%). In volumetric terms, the dominant diet contributors were O. plebs (38%), L. helicina (17%) and chaetognaths (15%). All prey except O. plebs and E. macronyx were taken in concurrent plankton samples. Although planktivory of Pagothenia borchgrevinki is confirmed, it is not certain whether feeding is confined to the immediate sub-ice environment.