No impact of a presumed manipulative parasite on the responses and susceptibility of fish to simulated predation

Parasites manipulating their host to facilitate trophic transmission is a widespread and diverse phenomenon. Trematode eye‐flukes in the family Diplostomidae infect a variety of fish species as metacercariae, many residing in the eyes. A recently described diplostomid, Tylodelphys darbyi, from the South Island of New Zealand has been found to infect common bully Gobiomorphus cotidianus, an endemic freshwater fish. Within the fish, the metacercariae move about freely in the liquid parts of the eye and are quite large. We hypothesized that increasing intensity of T. darbyi infection will result in increasing visual impairment, thus reducing the ability of G. cotidianus to identify and react to a predatory threat. To test this hypothesis, we performed experiments to (a) examine the fish's reaction to a purely visual predator cue and (b) test their ability to avoid simulated predation under natural levels of infection. Among the 64 fish used in our experiments, T. darbyi had a prevalence of 98.7% with an average of 17.6 worms per fish. However, there was no relationship between T. darbyi intensity and either the fish's reaction to a visual predator stimulus or their ability to escape a simulated predator. Our findings indicate that despite being present in large numbers in the eyes of its fish host, the parasite appears incapable of improving its chances of trophic transmission to its avian definitive host. The results also suggest that the fish G. cotidianus could be using other senses (e.g., olfaction and lateral line) to compensate for visual impairment, and detect and respond to predators.     

Eye morphology and visual acuity in the pipevine swallowtail (Battus philenor) studied with a new method of measuring interommatidial angles

Because of the important role sensory systems play in the behaviour of animals, information on sensory capabilities is of great value to behavioural ecologists in the development of hypotheses to explain behaviour. In compound eyes, interommatidial angles are a key determinant of visual acuity but methods for measuring these angles are often demanding and limited to live animals with a pseudopupil. Here we present a new technique for measuring interommatidial angles that is less demanding in terms of technology than other techniques but still accurate. It allows measurements in eyes without a pseudopupil such as dark eyes or even museum specimens. We call this technique the radius of curvature estimation (RCE) method. We describe RCE and validate the method by comparing results from RCE with those from pseudopupil analysis for the butterfly Asterocampa leilia. As an application of RCE we measure the eyes of the butterfly Battus philenor, a species whose visually guided behaviour is well known but whose eye structure and visual acuity are unknown. We discuss the results of the eye morphology in B. philenor in relation to their behaviour and ecology. We contend that RCE fills a gap in the repertoire of techniques available to study peripheral determinants of spatial resolution in compound eyes, because it can be applied on species with dark eyes. RCE then opens up for sampling a larger number of specimens, which, in combination with being able to use museum specimens, makes it possible to quantitatively test ecologically and evolutionarily driven hypotheses about vision in animals in a new way.     

<Emphasis Type="Italic">echinus</Emphasis>, required for interommatidial cell sorting and cell death in the <Emphasis Type="Italic">Drosophila</Emphasis> pupal retina,encodes a protein with homology to ubiquitin-specific proteases

Background  

Programmed cell death is used to remove excess cells between ommatidia in the Drosophila pupal retina. This death is required to establish the crystalline, hexagonal packing of ommatidia that characterizes the adult fly eye. In previously described echinus mutants, interommatidial cell sorting, which precedes cell death, occurred relatively normally. Interommatidial cell death was partially suppressed, resulting in adult eyes that contained excess pigment cells, and in which ommatidia were mildly disordered. These results have suggested that echinus functions in the pupal retina primarily to promote interommatidial cell death.     

Changes in visual fields associated with web reduction in the spider family uloboridae

When visual fields of the primitive orb-weaver, Waitkera waitkerensis, are reconstructed using measurements taken from intact lenses and cross and longitudinal sections of the prosoma, they show that this species has complete visual surveillance, but that none of the visual fields of its eight eyes overlap. The more advanced orb-weaver, Uloborus glomosus, also has eight eyes, but each eye has a greater visual angle, giving this species a complex pattern of overlapping visual fields. Uloborids that spin reduced webs are characterized by reduction or loss of the four anterior eyes and other carapace modifications necessary for them to effectively monitor and manipulate their reduced webs. The eyes of these uloborids have greater visual angles than those of orb-weavers, resulting primarily from perimetric expansion of their retinal hemispheres. Additionally, the axes of their visual fields are more ventrally directed due to greater dorsal than ventral retinal expansion and to ventral redirection of the entire eye. Consequently, even though the anterior lateral eyes of the triangle-weaver Hyptiotes cavatus lack retinae, the species' six functional eyes permit complete visual surveillance and exhibit visual overlap. The single-line-weaver, Miagrammopes animotus, has lost its four anterior eyes, and with them much of the anterior vision and all of the visual overlap found in the other species. However, changes similar to those of H. cavatus permit this species to retain most if its dorsal and ventral visual surveillance. Thus, ocular changes act in consort to maintain relatively complete visual surveillance in the face of eye loss and other major carapace modifications necessary for the operation of reduced webs.     

The visual fields of two ground‐foraging birds,House Finches and House Sparrows,allow for simultaneous foraging and anti‐predator vigilance

In birds, differences in the extent and position of the binocular visual field reflect adaptations to varying foraging strategies, and the extent of the lateral portion of the field may reflect anti‐predator strategies. The goal of this study was to describe and compare the visual fields of two ground‐foraging passerines, House Finch Carpodacus mexicanus and House Sparrow Passer domesticus. We found that both species have a binocular field type that is associated with the accurate control of bill position when pecking. Both species have eye movements of relatively large amplitude, which can produce substantial variations in the configuration of the binocular fields. We propose that in these ground foragers, their relatively wide binocular fields could function to increase foraging efficiency by locating multiple rather than single food items prior to pecking events. The lateral fields of both species are wide enough to facilitate the detection of predators or conspecifics while head‐down foraging. This suggests that foraging and scanning are not mutually exclusive activities in these species, as previously assumed. Furthermore, we found some slight, but significant, differences between species: House Sparrow binocular fields are both wider and vertically taller, and the blind area is wider than in House Finches. These differences may be related to variations in the degree of eye movements and position of the orbits in the skull.     

The eyes of Limulus polyphemus (Xiphosura, Chelicerata) and their afferent and efferent projections

The visual system of the American horseshoe crab Limulus polyphemus (L. polyphemus) is an important preparation for studying the photoresponse, the circadian modulation of the photoresponse and visual information processing. Given its unique position in phylogeny the structure of its visual system also informs studies of the relationships among arthropods and the characteristics of eurarthropods. Much has been learned about the organization of the relatively simple L. polyphemus visual system, but much remains to be discovered. This review summarizes current knowledge of the structure of L. polyphemus eyes and the organization of their afferent and efferent projections and points to important unanswered questions.     

Effects of Turbidity and Visual vs. Chemical Cues on Anti‐Predator Response in the Endangered Fountain Darter (Etheostoma fonticola)

Altered turbidity resulting from anthropogenic stressors is a global problem. Threatened by climate change, pollution, and increased recreational usage, the streams and rivers of central Texas are no exception. The impacts of turbidity include behavioral effects as turbidity degrades visual information, which can impair an animal's ability to accurately detect and respond to a predator. Here, we tested the impact of simulated turbidity on anti‐predator response in the endangered fountain darter, Etheostoma fonticola. We examined the response of E. fonticola to four predator cue treatments (chemical, visual, chemical and visual, and no cues) using a native predator, the green sunfish (Lepomis cyanellus). All cue treatments were tested across two vision levels: clear and impaired, to simulate the visual effects of low turbidity (~30 NTU). Our results indicate that E. fonticola requires a combination of visual and chemical stimuli to respond to a native fish predator. In the absence of one or the other sensory modality, E. fonticola did not show an anti‐predator response. Also, anti‐predator response to a combination of visual and chemical stimuli was only present at the clear vision level. When vision was impaired owing to simulated turbidity, a combination of visual and chemical stimuli did not produce a significant anti‐predator response. These results indicate that blocked or compromised vision hampers anti‐predator response in E. fonticola, which may be of concern regarding the future management of this endangered species.     

Compound eyes of insects and crustaceans: Some examples that show there is still a lot of work left to be done

Similarities and differences between the 2 main kinds of compound eye (apposition and superposition) are briefly explained before several promising topics for research on compound eyes are being introduced. Research on the embryology and molecular control of the development of the insect clear‐zone eye with superposition optics is one of the suggestions, because almost all of the developmental work on insect eyes in the past has focused on eyes with apposition optics. Age‐ and habitat‐related ultrastructural studies of the retinal organization are another suggestion and the deer cad Lipoptena cervi, which has an aerial phase during which it is winged followed by a several months long parasitic phase during which it is wingless, is mentioned as a candidate species. Sexual dimorphism expressing itself in many species as a difference in eye structure and function provides another promising field for compound eye researchers and so is a focus on compound eye miniaturization in very small insects, especially those that are aquatic and belong to species, in which clear‐zone eyes are diagnostic or are tiny insects that are not aquatic, but belong to taxa like the Diptera for instance, in which open rather than closed rhabdoms are the rule. Structures like interommatidial hairs and glands as well as corneal microridges are yet another field that could yield interesting results and in the past has received insufficient consideration. Finally, the dearth of information on distance vision and depth perception is mentioned and a plea is made to examine the photic environment inside the foam shelters of spittle bugs, chrysales of pupae and other structures shielding insects and crustaceans.     

Development of the compound eyes of dragonflies (Odonata). III. Adult compound eyes

The distribution of ommatidial diameters and interommatidial angles, as determined by measuring the angles between the optic axes of adjacent ommatidia, are mapped across the surface of the compound eyes of a variety of species selected for different adult behaviors, developmental histories, and taxonomic positions. The size of the visual fields, prey capture foveas, foveas composed of large dorsal ommatidia, and other specializations in the numbers of ommatidia that view various directions in the visual field are discussed in relation to adult behavior. Advanced species have less resemblance between their larval and adult eyes than primitive species. In contrast to their larvae, adults increase the monocular resolution of each eye at the expense of binocular vision. Most species have foveas which view in approximately the anterior direction, instead of in a region of binocular overlap, and many species have foveal bands which view along the horizon. Some advanced perching species, which approach their prey and other odonates from below, have an additional vertical foveal band that views along a vertical plane from the anterior direction to a more dorsal direction. The most unusual foveal band is seen in active flying species. The large dorsal ommatidia of the migratory Anax junius, which cover approximately one third of the eye surface, view a narrow region of the visual field that extends along a plane from the most lateral direction of one eye to a dorsal direction, and continues without interruption to the most lateral direction of the other eye.     

Visual fields of orb web and single line web spiders of the family Uloboridae (Arachnida,Araneida)

Summary In the family Uloboridae, web reduction is associated with changes in web monitoring posture and prosomal features. A spider must extend its first pair of legs directly forward to monitor the signal line of a reduced web. This posture is facilitated by shifts in prosomal musculature that cause reduced web uloborids to have a narrower anterior prosoma, a reduced or absent anterior eye row, and prominent posterior lateral eye tubercles. The eye tubercles and larger posterior eyes of these uloborids suggest that web reduction may also be accompanied by ocular changes that compensate for reduction of the anterior eyes by expanding the visual fields of the posterior eyes. A comparison of the visual fields of the eight-eyed, orb web species Octonoba octonaria and a four-eyed, reduced web Miagrammopes species was made to determine if this is true. Physical and optical measurements determined the visual angles of each species' eyes and the pattern of each species' visual surveillance. Despite loss of the anterior four eyes, the Miagrammopes species has a visual coverage similar to that of O. octonaria. This is due to (1) an increase in the visual field of each of the four remaining Miagrammopes eyes, accruing from an extension of the retina and an increase in the lens' rear radius of curvature, and (2) a ventral shift of each visual axis, associated with the development of an eye tubercle and an asymmetrical expansion of the retina. Miagrammopes monitor their simple webs from twigs or moss where they are vulnerable to predation. Therefore, maintenance of visual cover may enable them to detect predators in time to assume or maintain their characteristic, cryptic posture. It may also allow them to observe approaching prey and permit them to adjust web tension or prepare to jerk their webs when prey strikes.     

A phosphatase of undefined function is common to the photoreceptive microvilli of several arthropod species

Summary It has previously been demonstrated, using an ultracytochemical technique, that the photoreceptive microvilli of crab retinae contain a magnesium-dependent phosphatase that hydrolyses the artificial substrate 4-nitrophenylphosphate. Whilst many phosphatases hydrolyse 4-nitrophenylphosphate, the properties of the microvillar enzyme indicated that it is not a conventional acid or alkaline phosphatase. Using the same technique, it is now shown that a similar activity resides in the rhabdomeric microvilli of both the lateral compound eye and the ventral photoreceptors of Limulus polyphemus as well as in the compound eyes of the freshwater crayfish Cherax destructor and the fly Lucilia cuprina. Control cytochemical procedures performed on crayfish and fly showed that in these species too the activity is magnesium-dependent and is not due to a Na⁺/K⁺ ATPase.     

Visual field structure in the Empress Leilia, Asterocampa leilia (Lepidoptera, Nymphalidae): dimensions and regional variation in acuity

Male Empress Leilia butterflies ( Asterocampa leilia) use a sit-and-wait tactic to locate mates. To see how vision might influence male behavior, we studied the morphology, optics, and receptor physiology of their eyes and found the following. (1) Each eye's visual field is approximately hemispherical with at most a 10 degrees overlap in the fields of the eyes. There are no large sexual differences in visual field dimensions. (2) In both sexes, rhabdoms in the frontal and dorsal ommatidia are longer than those in other eye regions. (3) Interommatidial angles are smallest frontally and around the equator of the eye. Minimum interommatidial angles are 0.9-1 degrees in males and 1.3-1.4 degrees in females. (4) Acceptance angles of ommatidia closely match interommatidial angles in the frontal region of the eye. We conclude that vision in these butterflies is mostly monocular and that males have more acute vision than females, especially in the frontal region (large facets, small interommatidial angles, small acceptance angles, long rhabdoms, and a close match between interommatidial angles and acceptance angles). This study also suggests that perched males direct their most acute vision where females are likely to appear but show no eye modifications that appear clearly related to a mate-locating tactic.     

All the better to see you with: eyes and claws reveal the evolution of divergent ecological roles in giant pterygotid eurypterids

Pterygotid eurypterids have traditionally been interpreted as active, high-level, visual predators; however, recent studies of the visual system and cheliceral morphology of the pterygotid Acutiramus contradict this interpretation. Here, we report similar analyses of the pterygotids Erettopterus, Jaekelopterus and Pterygotus, and the pterygotid sister taxon Slimonia. Representative species of all these genera have more acute vision than A. cummingsi. The visual systems of Jaekelopterus rhenaniae and Pterygotus anglicus are comparable to that of modern predatory arthropods. All species of Jaekelopterus and Pterygotus have robust crushing chelicerae, morphologically distinct from the weaker slicing chelicerae of Acutiramus. Vision in Erettopterus osiliensis and Slimonia acuminata is more acute than in Acutiramus cummingsi, but not to the same degree as in modern active predators, and the morphology of the chelicerae in these genera suggests a grasping function. The pterygotids evolved with a shift in ecology from generalized feeder to specialized predator. Pterygotid eurypterids share a characteristic morphology but, although some were top predators, their ecology differs radically between genera.     

THE EURYPTERID ADELOPHTHALMUS SIEVERTSI (CHELICERATA: EURYPTERIDA) FROM THE LOWER DEVONIAN (EMSIAN) KLERF FORMATION OF WILLWERATH, GERMANY

Abstract: The eurypterid Rhenopterus sievertsi Størmer from the Lower Devonian of Germany is redescribed. Based on new morphological data, including the possession of prosomal limbs of Adelophthalmus‐type and spatulae on the genital operculum, the species is transferred to Adelophthalmus Jordan, in Jordan and von Meyer and thus is the oldest representative of this geographically and stratigraphically widespread genus. Eurypterus? trapezoides Størmer is recognized as a junior synonym of A. sievertsi.     

Visual fields, eye movements, and scanning behavior of a sit-and-wait predator, the black phoebe (Sayornis nigricans)

Foraging mode influences the dominant sensory modality used by a forager and likely the strategies of information gathering used in foraging and anti-predator contexts. We assessed three components of visual information gathering in a sit-and-wait avian predator, the black phoebe (Sayornis nigricans): configuration of the visual field, degree of eye movement, and scanning behavior through head-movement rates. We found that black phoebes have larger lateral visual fields than similarly sized ground-foraging passerines, as well as relatively narrower binocular and blind areas. Black phoebes moved their eyes, but eye movement amplitude was relatively smaller than in other passerines. Black phoebes may compensate for eye movement constraints with head movements. The rate of head movements increased before attacking prey in comparison to non-foraging contexts and before movements between perches. These findings suggest that black phoebes use their lateral visual fields, likely subtended by areas of high acuity in the retina, to track prey items in a three-dimensional space through active head movements. These head movements may increase depth perception, motion detection and tracking. Studying information gathering through head movement changes, rather than body posture changes (head-up, head-down) as generally presented in the literature, may allow us to better understand the mechanisms of information gathering from a comparative perspective.     

Eyes and vision in the coeval Furongian trilobites Sphaerophthalmus alatus (Boeck, 1938) and Ctenopyge (Mesoctenopyge) tumida Westergård, 1922, from Bornholm,Denmark

The two olenid species Sphaerophthalmus alatus (Boeck, 1838) and Ctenopyge (Mesoctenopyge) tumida Westergård, 1922, occur together in the Ctenopyge tumida Zone (Zone 19) of the Furongian of Scandinavia. Material from Bornholm, Denmark, forms the basis of this study of the morphology and partial ontogeny of the eyes. The eyes of both species are directed laterally and have virtually panoramic vision, looking out sideways like those of a rabbit. The eye of S. alatus is comparatively smaller, with fewer lenses and a larger eye parameter; calculations show that this trilobite was adapted for dim light intensity, possibly suggesting a vagrant benthic habit. Ctenopyge (Mesoctenopyge) tumida, with a smaller eye parameter, was adapted for a higher light intensity, and this trilobite was most likely a pelagic swimmer. The two species, although preserved together, inhabited different levels in the water column.     

The evolution of vertebrate eye size across an environmental gradient: phenotype does not predict genotype in a Trinidadian killifish

Vertebrates exhibit substantial variation in eye size. Eye size correlates positively with visual capacity and behaviors that enhance fitness, such as predator avoidance. This foreshadows a connection between predation and eye size evolution. Yet, the conditions that favor evolutionary shifts in eye size, besides the well‐known role for light availability, are unclear. We tested the influence of predation on the evolution of eye size in Trinidadian killifish, Rivulus hartii. Rivulus are located across a series of communities where they coexist with visually oriented piscivores ("high predation" sites), and no predators (“Rivulus‐only” sites). Wild‐caught Rivulus from high predation sites generally exhibited a smaller relative eye size than communities that lack predators. Yet, such differences were inconsistent across rivers. Second‐generation common garden reared fish revealed repeatable decreases in eye size in Rivulus from high predation sites. We performed additional experiments that tested the importance of light and resources on eye size evolution. Sites that differ in light or resource availability did not differ in eye size. Our results argue that differences in predator‐induced mortality underlie genetically‐based shifts in vertebrate eye size. We discuss the drivers of eye size evolution in light of the nonparallel trends between the phenotypic and common garden results.     

A NEW SPECIES OF DICERORHINUS (RHINOCEROTIDAE) FROM THE PLIO‐PLEISTOCENE OF MYANMAR

Abstract: A skull and mandible of the new species Dicerorhinus gwebinensis sp. nov. of Rhinocerotidae (Mammalia, Perissodactyla) is described. The material is collected from the upper part of the Irrawaddy sediments (Plio‐Pleistocene) in central Myanmar. D. gwebinensis sp. nov. is morphologically more similar to the extant species D. sumatrensis (Sumatran rhinoceros) than to other species of the genus but differs from D. sumatrensis in having the comparatively shorter nasal, the more concave dorsal profile of the skull, the more elevated occiput and presence of molar crista in M3/. This is the first discovery of Dicerorhinus in the upper Miocene to lower Pleistocene of the Indian subcontinent and Mainland Southeast Asia, and fills the chronological and geographical gap of this lineage in Asia. The Dicerorhinus clade probably migrated into Southeast Asia from East Asia by the Pliocene or early Pleistocene. This hypothesis is supported by the scarcity or absence of this clade in the Neogene mammalian fauna of the Indian Subcontinent.     

Ocular changes accompanying eye loss in the spider family uloboridae

In uloborid spiders, eye loss is accompanied by increased visual angles, optical material investment, and potential visual acuity of the retained eyes. Relative to carapace volume, the six-eyed Hyptiotes cavatus and two four-eyed Miagrammopes species have greater retinal hemisphere areas and lens volumes than do the eight-eyed uloborids Waitkera waitkerensis, Uloborus glomosus, and Octonoba sinensis. In Waitkera, in which the eyes have little visual overlap, and in Miagrammopes, in which eye loss simplifies the spiders' patterns of visual overlap, increased retinal cell density enhances potential visual acuity. However, this occurs at the expense of potential retinal cell sensitivity.     

Disentangling mite predator‐prey relationships by multiplex PCR

Gut content analysis using molecular techniques can help elucidate predator‐prey relationships in situations in which other methodologies are not feasible, such as in the case of trophic interactions between minute species such as mites. We designed species‐specific primers for a mite community occurring in Spanish citrus orchards comprising two herbivores, the Tetranychidae Tetranychus urticae and Panonychus citri, and six predatory mites belonging to the Phytoseiidae family; these predatory mites are considered to be these herbivores’ main biological control agents. These primers were successfully multiplexed in a single PCR to test the range of predators feeding on each of the two prey species. We estimated prey DNA detectability success over time (DS₅₀), which depended on the predator‐prey combination and ranged from 0.2 to 18 h. These values were further used to weight prey detection in field samples to disentangle the predatory role played by the most abundant predators (i.e. Euseius stipulatus and Phytoseiulus persimilis). The corrected predation value for E. stipulatus was significantly higher than for P. persimilis. However, because this 1.5‐fold difference was less than that observed regarding their sevenfold difference in abundance, we conclude that P. persimilis is the most effective predator in the system; it preyed on tetranychids almost five times more frequently than E. stipulatus did. The present results demonstrate that molecular tools are appropriate to unravel predator‐prey interactions in tiny species such as mites, which include important agricultural pests and their predators.