Apposition Eyes Research Articles

Anatomy and ultrastructural details of the eye of the passalid beetle Ceracupes yui Okano 1988 (Scarabaeoidea; Passalidae)

One of the least studied eyes of any beetle taxon are those of the scarabaeoid family Passalidae. Some members of this family of around 600 species worldwide are known to have superposition eyes (Aceraius grandis; A. hikidai) while others have apposition eyes (Cylindrocaulus patalis; Ceracupes yui). In C. yui of nearly 3 cm body length (this paper) the retinal layer is very thin and occupies approximately half of an ommatidium's total length, the latter amounting to 284 and 266 μm in the respective dorsal and ventral eye regions. The two eye regions are almost completely separated by a prominent cuticular canthus, a feature usually associated with the presence of a tracheal tapetum, a clear-zone between dioptric and light-perceiving structures and a regular array of smooth facets. In C. yui the facets are smooth (but not very regular) and a tracheal tapetum and a clear-zone are absent. The rhabdoms, formed by 8–9 retinula cells, are complicated, multilobed structures with widths and lengths of around 15 and 80 μm, respectively. The combination of some superposition and mostly apposition eye features, e.g., extensive corneal exocones, relatively small number of ommatidia, absence of a clear-zone and tracheal bush, suggest an adaptation of this species' eye to the fossorial lifestyle of C. yui, and, thus, a manifestation of the passalid eye's plasticity.

Investigation of the ultrastructures and retinal arrangements of larval stomatopod eyes

Though the transparent apposition eyes of larval stomatopod crustaceans lack most of the unique retinal specializations known from their adult counterparts, increasing evidence suggests that these tiny pelagic organisms possess their own version of retinal complexity. In this paper, we examined the structural organization of larval eyes in six species of stomatopod crustaceans across three stomatopod superfamilies using transmission electron microscopy. The primary focus was to examine retinular cell arrangement of the larval eyes and characterize the presence of an eighth retinular cell (R8), which is typically responsible for UV vision in crustaceans. For all species investigated, we identified R8 photoreceptor cells positioned distal to the main rhabdom of R1-7 cells. This is the first evidence that R8 photoreceptor cells exist in larval stomatopod retinas, and among the first identified in any larval crustacean. Considering recent studies that identified UV sensitivity in larval stomatopods, we propose that this sensitivity is driven by this putative R8 photoreceptor cell. Additionally, we identified a potentially unique crystalline cone structure in each of the species examined, the function of which is still not understood.

Microfabrication of bioinspired curved artificial compound eyes: a review

Eyes are one of the most important organs for natural creatures to perceive external information. The curved compound eyes of insects endowing with merits of a wide field of view (FOV), high sensitivity and detecting quick moving targets have aroused extensive concern. There are mainly two categories of compound eyes in nature. One is apposition eyes which can avoid light crosstalk and the other is superposition ones with higher sensitivity. With the rapid development of micron and nano machining technologies, the fabrication techniques of compound eyes have experienced changes from waveguide self-writing initially to advanced femtosecond laser method through thermal reflow and other processes. The diameters of compound eyes varies from the original several hundreds micrometers to the smallest 5 µm currently, which is much closer to the natural ones. Therefore, a critical review is presented to systematically summary and compare the micromachining processes of curved artificial compound eyes in recent years from the perspective of microfabrication technology. Finally, we describe a broad applications including navigation and location, digital camera and rapid detection of moving objects as well as existing challenges and future perspective of curved artificial compound eyes. It is hoped that this review can help researchers to gain a better overall understanding of microfabrication in this field and push forward it to a new stage.

Nocturnal Bees Feed on Diurnal Leftovers and Pay the Price of Day – Night Lifestyle Transition

1. Bees exemplify flights under bright sunlight. A few species across bee families have evolved nocturnality, displaying remarkable adaptations to overcome limitations of their daylight-suited apposition eyes. Phase inversion to nocturnality in a minority of bees that co-exist with diurnal bees provide a unique opportunity to study ecological benefits that mediate total temporal niche shifts. While floral traits and sensory modalities associated with the evolution of classical nocturnal pollination syndromes, e.g. by bats and moths, are well-studied, nocturnality in bees represent a poorly understood, recently invaded, extreme niche. 2. To test the competitive release hypothesis, we examine how nocturnality shapes foraging by comparing pollen loads, nest pollen and flower visitation of sympatric nocturnal and diurnal carpenter bees. We predicted that nocturnal bees primarily use night-blooming flowers, show little/no resource overlap with diurnal species and competitive release favours night-time pollen collection for provisioning. 3. Contrarily, we found substantial resource overlap between nocturnal and diurnal bees. Flower opening times, floral longevity and plant abundance did not define nocturnal flower use. Smaller pollen loads on nocturnal foragers suggests subsistence on resource leftovers largely from diurnal flowers. Greater pollen types/diversity on nocturnal foragers indicates lower floral constancy compared to diurnal congenerics. Reduced activity during new moon compared to full moon suggests constraints to nocturnal foraging. 4. Invasion and sustenance within the nocturnal niche is characterised by: i)opportunistic foraging on residual resources as indicated by smaller pollen loads, extensive utilisation of day-blooming flowers and substantial overlap with diurnal bees, ii)generalisation at two levels – between and within foraging trips as indicated by lower floral constancy, iii)reduced foraging on darker nights, indicating visual constraints despite sensitive optics. This together with smaller populations and univoltine breeding in nocturnal compared to multivoltine diurnal counterparts, suggest that nocturnality imposes substantial fitness costs. 5. In conclusion, the evolution of nocturnality in bees is accompanied by resource generalisation instead of specialisation. Reduced floral constancy suggests differences in foraging strategies of nocturnal and diurnal bees which merits further investigation. The relative roles of competition, floral rewards and predators should be examined to fully understand the evolution and maintenance of nocturnality in bees.

Insights into a 429-million-year-old compound eye

In all arthropods the plesiomorphic (ancestral character state) kind of visual system commonly is considered to be the compound eye. Here we are able to show the excellently preserved internal structures of the compound eye of a 429 Mya old Silurian trilobite, Aulacopleura koninckii (Barrande, 1846). It shows the characteristic elements of a modern apposition eye, consisting of 8 (visible) receptor cells, a rhabdom, a thick lens, screening pigment (cells), and in contrast to a modern type, putatively just a very thin crystalline cone. Functionally the latter underlines the idea of a primarily calcitic character of the lens because of its high refractive properties. Perhaps the trilobite was translucent. We show that this Palaeozoic trilobite in principle was equipped with a fully modern type of visual system, a compound eye comparable to that of living bees, dragonflies and many diurnal crustaceans. It is an example of excellent preservation, and we hope that this manuscript will be a starting point for more research work on fossil evidence, and to develop a deeper understanding of the evolution of vision.

The giant freshwater prawnMacrobrachium rosenbergiialters background colour preference after metamorphosis from larvae to postlarvae: In association with nature of phototaxis

The giant freshwater prawn Macrobrachium rosenbergii larvae have apposition eyes and are positively phototactic, whereas the postlarvae (PL) have superposition eyes and are negatively phototactic. M. rosenbergii has colour vision as early as larval stage. We discovered that M. rosenbergii alters background colour preference after metamorphosis from larvae to PL in association with nature of phototaxis. The test circular glass aquaria covered with a pair of two-colour papers contained with a group of 100 larvae or 20 Pl, and the number of individuals in each colour background was recorded five times for each colour pair. The background colours tested were light blue, green, yellow, red, white and black. The numbers of larvae or PL in each colour background of different pairs were analysed by the Thurstone's law of comparative judgment. In the larvae, significant bias towards yellow was evident. In the PL, of the four pairings of black with other colours, all biased to black. The mean z-scores were highest for yellow in the larvae, and for black in the PL. To determine the possible background brightness preference of the larvae and PL, six different colour backgrounds were presented in pairs. The larvae significantly preferred light blue over dark blue, white over yellow and white over black. The PL exhibited reversed preference. The relationship between z-scores and light reflectance levels of five colour papers was significantly positive in the larvae and negative in the PL. The observed background colour preference was probably due to relative brightness rather than chromaticity difference.

Evolutionary constraints on flicker fusion frequency in Lepidoptera.

Flying insects occupy both diurnal and nocturnal niches, and their visual systems encounter distinct challenges in both conditions. Visual adaptations, such as superposition eyes of moths, enhance sensitivity to low light levels but trade off with spatial and temporal resolution. Conversely, apposition eyes of butterflies enable high spatial resolution but are poorly sensitive in dim light. Although diel activity patterns of insects influence visual processing, their role in evolution of visual systems is relatively unexplored. Lepidopteran insects present an excellent system to study how diel activity patterns and phylogenetic position influence the visual transduction system. We addressed this question by comparing electroretinography measurements of temporal response profiles of diverse Lepidoptera to light stimuli that were flickering at different frequencies. Our data show that the eyes of diurnal butterflies are sensitive to visual stimuli of higher temporal frequencies than nocturnal moths. Hesperiid skippers, which are typically diurnal or crepuscular, exhibit intermediate phenotypes with peak sensitivity across broader frequency range. Across all groups, species within families exhibited similar phenotypes irrespective of diel activity. Thus, Lepidopteran photoreceptors may have diversified under phylogenetic constraints, and shifts in their sensitivity to higher temporal frequencies occurred concomitantly with the evolution of diurnal lifestyles.

Trilobite compound eyes with crystalline cones and rhabdoms show mandibulate affinities

Most knowledge about the structure, function, and evolution of early compound eyes is based on investigations in trilobites. However, these studies dealt mainly with the cuticular lenses and little was known about internal anatomy. Only recently some data on crystalline cones and retinula cells were reported for a Cambrian trilobite species. Here, we describe internal eye structures of two other trilobite genera. The Ordovician Asaphus sp. reveals preserved crystalline cones situated underneath the cuticular lenses. The same is true for the Devonian species Archegonus (Waribole) warsteinensis, which in addition shows the fine structure of the rhabdom in the retinula cells. These results suggest that an apposition eye with a crystalline cone is ancestral for Trilobita. The overall similarity of trilobite eyes to those of myriapods, crustaceans, and hexapods corroborates views of a phylogenetic position of trilobites in the stem lineage of Mandibulata.

Palaeoecology of Voulteryon parvulus (Eucrustacea, Polychelida) from the Middle Jurassic of La Voulte-sur-Rh\xf4ne Fossil-Lagerst\xe4tte (France)

Exceptional and extremely rare preservation of soft parts, eyes, or syn-vivo associations provide crucial palaeoecological information on fossil-rich deposits. Here we present exceptionally preserved specimens of the polychelidan lobster Voulteryon parvulus, from the Jurassic of La Voulte-sur-Rhône Fossil-Lagerstätte, France, bearing eyes with hexagonal and square facets, ovaries, and a unique association with epibiont thecideoid brachiopods, giving insights onto the palaeoenvironment of this Lagerstätte. The eyes, mostly covered in hexagonal facets are interpreted as either apposition eyes (poorly adapted to low-light environment) or, less likely, as refractive or parabolic superposition eyes (compatible with dysphotic palaeoenvironments). The interpretation that V. parvulus had apposition eyes suggests an allochthonous, shallow water origin. However, the presence of thecideoid brachiopod ectosymbionts on its carapace, usually associated to dim-light paleoenvironments and/or rock crevices, suggests that V. parvulus lived in a dim-light setting. This would support the less parsimonious interpretation that V. parvulus had superposition eyes. If we accept the hypothesis that V. parvulus had apposition eyes, since the La Voulte palaeoenvironment is considered deep water and had a soft substrate, V. parvulus could have moved into the La Voulte Lagerstätte setting. If this is the case, La Voulte biota would record a combination of multiple palaeoenvironments.

The giant butterfly-moth Paysandisia archon has spectrally rich apposition eyes with unique light-dependent photoreceptor dynamics

The palm borer moth Paysandisia archon (Burmeister, 1880) (fam. Castniidae) is a large, diurnally active palm pest. Its compound eyes consist of ~ 20,000 ommatidia and have apposition optics with interommatidial angles below 1°. The ommatidia contain nine photoreceptor cells and appear structurally similar to those in nymphalid butterflies. Two morphological ommatidial types were identified. Using the butterfly numbering scheme, in type I ommatidia, the distal rhabdom consists exclusively of the rhabdomeres of photoreceptors R1–2; the medial rhabdom has contributions from R1–8. The rhabdom in type II ommatidia is distally split into two sub-rhabdoms, with contributions from photoreceptors R2, R3, R5, R6 and R1, R4, R7, R8, respectively; medially, only R3–8 and not R1–2 contribute to the fused rhabdom. In both types, the pigmented bilobed photoreceptors R9 contribute to the rhabdom basally. Their nuclei reside in one of the lobes. Upon light adaptation, in both ommatidial types, the rhabdoms secede from the crystalline cones and pigment granules invade the gap. Intracellular recordings identified four photoreceptor classes with peak sensitivities in the ultraviolet, blue, green and orange wavelength regions (at 360, 465, 550, 580 nm, respectively). We discuss the eye morphology and optics, the photoreceptor spectral sensitivities, and the adaptation to daytime activity from a phylogenetic perspective.

Structure and Ultrastructure of the Acrotrichis grandicollis (Coleoptera: Ptiliidae) Compound Eyes and the Eye Features Related to Miniaturisation.

Here is the first description of the visual organ of Acrotrichis grandicollis, a member of Ptiliidae family that includes the smallest free-living insects. The apposition eyes of A. grandicollis have the acone-type crystalline cone; a fused rhabdom is formed by eight retinula cells and surrounded by primary pigment cells. Secondary pigment cells are located only in the distal part of the ommatidium under the lens. The eye consists of about 50 facets about 12 μm in diameter. The A. grandicollis ommatidium ultrastructure has been compared with that of large Coleoptera. The results obtained enabled us to emphasize the compound eye specific features associated with the small body size.

Limit of colour vision in dim light in larvae of the giant freshwater prawn Macrobrachium rosenbergii

Colour vision depends on sufficient ambient light and becomes ineffective at a particular low light intensity. It is not known how decapod crustaceans see colour in dim light. In the present study we investigated the colour vision threshold in larvae of the giant freshwater prawn Macrobrachium rosenbergii in a tank under natural illumination. Plastic beads of different colours (blue, red, yellow and white) in various combinations were suspended in the tank. The larvae swam straight toward the beads and gathered around them. The number of larvae was highest on the blue and white beads. The luminance in the tank was then gradually decreased by covering it with different numbers of layers of black cloth, and the response of the larvae to the beads was video-recorded under infrared illumination. The preference for blue and white beads remained manifest as the luminance was reduced from 418 to 0.07 cd/m2, but not at 0.02 cd/m2, indicating a colour vision threshold between 0.07 and 0.02 cd/m2. The larvae have apposition compound eyes with large optical parameters, comparable to those of apposition eyes of nocturnal insects, which presumably capture more light and show enhanced sensitivity, enabling the larvae to see colour in dim light.

Structure and function of a compound eye, more than half a billion years old

Until now, the fossil record has not been capable of revealing any details of the mechanisms of complex vision at the beginning of metazoan evolution. Here, we describe functional units, at a cellular level, of a compound eye from the base of the Cambrian, more than half a billion years old. Remains of early Cambrian arthropods showed the external lattices of enormous compound eyes, but not the internal structures or anything about how those compound eyes may have functioned. In a phosphatized trilobite eye from the lower Cambrian of the Baltic, we found lithified remnants of cellular systems, typical of a modern focal apposition eye, similar to those of a bee or dragonfly. This shows that sophisticated eyes already existed at the beginning of the fossil record of higher organisms, while the differences between the ancient system and the internal structures of a modern apposition compound eye open important insights into the evolution of vision.

Crustacean Larvae-Vision in the Plankton.

We review the visual systems of crustacean larvae, concentrating on the compound eyes of decapod and stomatopod larvae as well as the functional and behavioral aspects of their vision. Larval compound eyes of these macrurans are all built on fundamentally the same optical plan, the transparent apposition eye, which is eminently suitable for modification into the abundantly diverse optical systems of the adults. Many of these eyes contain a layer of reflective structures overlying the retina that produces a counterilluminating eyeshine, so they are unique in being camouflaged both by their transparency and by their reflection of light spectrally similar to background light to conceal the opaque retina. Besides the pair of compound eyes, at least some crustacean larvae have a non-imaging photoreceptor system based on a naupliar eye and possibly other frontal eyes. Larval compound-eye photoreceptors send axons to a large and well-developed optic lobe consisting of a series of neuropils that are similar to those of adult crustaceans and insects, implying sophisticated analysis of visual stimuli. The visual system fosters a number of advanced and flexible behaviors that permit crustacean larvae to survive extended periods in the plankton and allows them to reach acceptable adult habitats, within which to metamorphose.

On the sighted ancestry of blindness - exceptionally preserved eyes of Mesozoic polychelidan lobsters.

BackgroundModern representatives of Polychelida (Polychelidae) are considered to be entirely blind and have largely reduced eyes, possibly as an adaptation to deep-sea environments. Fossil species of Polychelida, however, appear to have well-developed compound eyes preserved as anterior bulges with distinct sculpturation.MethodsWe documented the shapes and sizes of eyes and ommatidia based upon exceptionally preserved fossil polychelidans from Binton (Hettangian, United-Kingdom), Osteno (Sinemurian, Italy), Posidonia Shale (Toarcian, Germany), La Voulte-sur-Rhône (Callovian, France), and Solnhofen-type plattenkalks (Kimmeridgian-Tithonian, Germany). For purposes of comparison, sizes of the eyes of several other polychelidans without preserved ommatidia were documented. Sizes of ommatidia and eyes were statistically compared against carapace length, taxonomic group, and outcrop.ResultsNine species possess eyes with square facets; Rosenfeldia oppeli (Woodward, 1866), however, displays hexagonal facets. The sizes of eyes and ommatidia are a function of carapace length. No significant differences were discerned between polychelidans from different outcrops; Eryonidae, however, have significantly smaller eyes than other groups.DiscussionFossil eyes bearing square facets are similar to the reflective superposition eyes found in many extant decapods. As such, they are the earliest example of superposition eyes. As reflective superposition is considered plesiomorphic for Reptantia, this optic type was probably retained in Polychelida. The two smallest specimens, a Palaeopentacheles roettenbacheri (Münster, 1839) and a Hellerocaris falloti (Van Straelen, 1923), are interpreted as juveniles. Both possess square-shaped facets, a typical post-larval feature. The eye morphology of these small specimens, which are far smaller than many extant eryoneicus larvae, suggests that Jurassic polychelidans did not develop via giant eryoneicus larvae. In contrast, another species we examined, Rosenfeldia oppeli (Woodward, 1866), did not possess square-shaped facets, but rather hexagonal ones, which suggests that this species did not possess reflective superposition eyes. The hexagonal facets may indicate either another type of superposition eye (refractive or parabolic superposition), or an apposition eye. As decapod larvae possess apposition eyes with hexagonal facets, it is most parsimonious to consider eyes of R. oppeli as apposition eyes evolved through paedomorphic heterochrony.ConclusionPolychelidan probably originally had reflective superposition. R. oppeli, however, probably gained apposition eyes through paedomorphosis.

Current advances in invertebrate vision: insights from patch-clamp studies of photoreceptors in apposition eyes.

Traditional electrophysiological research on invertebrate photoreceptors has been conducted in vivo, using intracellular recordings from intact compound eyes. The only exception used to be Drosophila melanogaster, which was exhaustively studied by both intracellular recording and patch-clamp methods. Recently, several patch-clamp studies have provided new information on the biophysical properties of photoreceptors of diverse insect species, having both apposition and neural superposition eyes, in the contexts of visual ecology, behavior, and ontogenesis. Here, I discuss these and other relevant results, emphasizing differences between fruit flies and other species, between photoreceptors of diurnal and nocturnal insects, properties of distinct functional types of photoreceptors, postembryonic developmental changes, and relationships between voltage-gated potassium channels and visual ecology.

Arthropod eyes: The early Cambrian fossil record and divergent evolution of visual systems

Four types of eyes serve the visual neuropils of extant arthropods: compound retinas composed of adjacent facets; a visual surface populated by spaced eyelets; a smooth transparent cuticle providing inwardly directed lens cylinders; and single-lens eyes. The first type is a characteristic of pancrustaceans, the eyes of which comprise lenses arranged as hexagonal or rectilinear arrays, each lens crowning 8–9 photoreceptor neurons. Except for Scutigeromorpha, the second type typifies Myriapoda whose relatively large eyelets surmount numerous photoreceptive rhabdoms stacked together as tiers. Scutigeromorph eyes are facetted, each lens crowning some dozen photoreceptor neurons of a modified apposition-type eye. Extant chelicerate eyes are single-lensed except in xiphosurans, whose lateral eyes comprise a cuticle with a smooth outer surface and an inner one providing regular arrays of lens cylinders. This account discusses whether these disparate eye types speak for or against divergence from one ancestral eye type. Previous considerations of eye evolution, focusing on the eyes of trilobites and on facet proliferation in xiphosurans and myriapods, have proposed that the mode of development of eyes in those taxa is distinct from that of pancrustaceans and is the plesiomorphic condition from which facetted eyes have evolved. But the recent discovery of enormous regularly facetted compound eyes belonging to early Cambrian radiodontans suggests that high-resolution facetted eyes with superior optics may be the ground pattern organization for arthropods, predating the evolution of arthrodization and jointed post-protocerebral appendages. Here we provide evidence that compound eye organization in stem-group euarthropods of the Cambrian can be understood in terms of eye morphologies diverging from this ancestral radiodontan-type ground pattern. We show that in certain Cambrian groups apposition eyes relate to fixed or mobile eyestalks, whereas other groups reveal concomitant evolution of sessile eyes equipped with optics typical of extant xiphosurans. Observations of fossil material, including that of trilobites and eurypterids, support the proposition that the ancestral compound eye was the apposition type. Cambrian arthropods include possible precursors of mandibulate eyes. The latter are the modified compound eyes, now sessile, and their underlying optic lobes exemplified by scutigeromorph chilopods, and the mobile stalked compound eyes and more elaborate optic lobes typifying Pancrustacea. Radical divergence from an ancestral apposition type is demonstrated by the evolution of chelicerate eyes, from doublet sessile-eyed stem-group taxa to special apposition eyes of xiphosurans, the compound eyes of eurypterids, and single-lens eyes of arachnids. Different eye types are discussed with respect to possible modes of life of the extinct species that possessed them, comparing these to extant counterparts and the types of visual centers the eyes might have served.

Absorption-assisted mode transformation in butterfly compound eyes.

The ommatidium of the butterfly's afocal apposition eye exhibits angular performance that can only be achieved by transforming the diffraction pattern of its corneal lens into the fundamental mode of its rhabdom waveguide. A graded index model of the ommatidium has been proposed and verified but the efforts to extract the transformation's underlying physics from it have been hindered by its extreme complexity. Here we numerically investigate the ommatidium model and reveal that the current model, involving only the graded index distribution, is insufficient for the transformation. We further find that adding spatially varying absorption to the existing model dramatically improves its transformation performance, producing near-perfect mode matching with overlap integral exceeding 0.96. Such a combined action of spatially varying index and absorption for microscale mode transformation is new to researchers in optics and biology and will benefit both disciplines.

How do eye size and facet lens size vary by age and sex inAcheta domesticus?

Abstract. Vision is essential to interactions with the environment and survival of many animals. House crickets, Acheta domesticus, are nocturnal insects with compound apposition eyes, and thus may have a specialized visual system selected for low-light vision. Here, an analysis of regional variation in facet diameter around the eye of a cricket was conducted. Eye surface area was estimated and facet diameter was spot-sampled across the eyes of 13 adults and 13 juveniles. Juveniles had smaller facets than adults, and there was significant variation among eye regions in facet size. Specifically, larger facets were found in the posterior eye region. Overall, facets were large given the size of this animal's eye, which is consistent with what one might expect from a nocturnal insect under selection for high sensitivity. Furthermore, juveniles had proportionally larger facets than adults, which may indicate that the insects maximize the sensitivity of their ommatidia early in development, prior to increasing ...

An apposition-like compound eye with a layered rhabdom in the small diving beetle Agabus japonicus (Coleoptera, Dytiscidae).

The fine structure of the compound eyes of the adult diving beetle Agabus japonicus is described with light, scanning, and transmission electron microscopy. The eye of A. japonicus is mango-shaped and consists of about 985 ommatidia. Each ommatidium is composed of a corneal facet lens, an eucone type of crystalline cone, a fused layered rhabdom with a basal rhabdomere, seven retinula cells (including six distal cells and one basal cell), two primary pigment cells and an undetermined number of secondary pigment cells that are restricted to the distalmost region of the eye. A clear-zone, separating dioptric apparatus from photoreceptive structures, is not developed and the eye thus resembles an apposition eye. The cross-sectional areas of the rhabdoms are relatively large indicative of enhanced light-sensitivity. The distal and central region of the rhabdom is layered with interdigitating microvilli suggesting polarization sensitivity. According to the features mentioned above, we suggest that 1) the eye, seemingly of the apposition type, occurs in a taxon for which the clear-zone (superposition) eye is characteristic; 2) the eye possesses adaptations to function in a dim-light environment; 3) the eye may be sensitive to underwater polarized light or linearly water-reflected polarized light.