The phylogeny of diving beetles (Coleoptera: Dytiscidae) and the evolution of sexual conflict

A model of evolution based on conflicts of interest between the sexes over mating decisions is examined in relation to diving beetles (Dytiscidae). The model predicts the following evolutionary sequence: (1) cost to females of mating increases, (2) females evolve behavioural resistance to male mating attempts, (3) males evolve devices to overcome female resistance, and (4) females evolve morphological counter-adaptations to the male devices. This model is tested using species of Dytiscidae, in which (1) some species have a very long mating duration while others mate quickly, (2) females of some species resist male mating attempts by swift and erratic swimming when seized by a male, (3) males of some species possess a grasping device in the form of sucker-shaped setae on the legs used to adhere to the pronota or elytra of females prior to mating, and (4) females of some species have a modified dorsal cuticle with irregular sculpturing which appears to interfere with the male adhesive setae. The predicted order of evolution of some of these features was tested in a cladistic analysis of 52 taxa in Dytiscidae and Hygrobiidae using characters from adult and larval morphology and a portion of the gene wingless . The combined analysis resulted in nine most parsimonious cladograms. The consensus cladogram of these indicates that male sucker setae arose a single time in a clade of Dytiscinae. Nested within this clade are five groups with an independently derived, modified dorsal cuticle in females. This pattern of characters in Dytiscinae is consistent with the prediction implied by the model of sexual selection. The utility of wingless as a marker for phylogenetic analysis of diving beetles is discussed, and the resulting phylogeny is compared with previous analyses and current classification.  © 2003 The Linnean Society of London, Biological Journal of the Linnean Society , 2003, 79 , 359–388.     

Sexual dimorphism and sexual conflict in the diving beetle Agabus uliginosus (L.) (Coleoptera: Dytiscidae)

Sexual conflict can drive intersexual arms races, with female resistance and male persistence traits coevolving antagonistically. Such arms races are well documented in some diving beetles, although the extent of sexual conflict in this family remains unclear. The European dytiscid Agabus uliginosus has a strikingly dimorphic female; individuals from most regions are smooth and male‐like, whereas those from some populations have a strongly roughened dorsum, a trait that has attracted the name dispar. We demonstrate that rough and smooth females differ consistently in the development of dorsal surface microreticulation, and that these females are associated with males that differ in the development of their persistence traits. These findings extend the occurrence of pre‐insemination sexual conflict and associated intrasexual dimorphism in Dytiscidae, and suggest that such mating systems are relatively widespread in these beetles.     

Intraspecific variation and intersexual correlation in secondary sexual characters of three diving beetles (Coleoptera: Dytiscidae)

Male diving beetles of the subfamily Dytiscinae possess tarsi with adhesive discs that they strike on the female dorsum during mating interactions. Females of many species are dimorphic, being either smooth or structured dorsally. Darwin suggested the female structures were an aid for the male but in this study we investigate these characteristics in the light of sexual conflicts. The intraspecific variation in the numbers and size distribution of male tarsal discs, and in body measurements were recorded for three dytiscine species, all with dimorphic females. The number of protarsal discs in the two Dytiscus species varied much more than previously reported. In addition, only a small part of the variation could be explained by body size. In Graphoderus we found highly significant differences in male secondary sexual characters among populations. A multivariate analysis significantly correlated male secondary sexual characters with the proportion of granulate females in the populations. These observations are consistent with the theory of arms races and female counter adaptations. Covariation between male and female characters is predicted from a framework of sexual conflict over mating rate. At the same time our study gives a new perspective on the function of dytiscine female dorsal irregularities debated ever since Darwin.     

The inheritance of intrasexual dimorphism in female diving beetles (Coleoptera: Dytiscidae)

Many species of Dytiscus diving beetles exhibit intrasexual dimorphism, e.g., the elytra is smooth in some females and grooved in others. However, the expression of the grooves and whether they are a product of heredity or the environment remain unknown. One Japanese species, Dytiscus sharpi sharpi Wehncke, 1875 , also shows female dimorphism, with grooved and smooth morphs, while D. sharpi validus Régimbart, 1899, only has a single morph (the grooved type). A hybrid of the two species should therefore provide a means of sorting out how the grooves are inherited. We found two independent wetlands of D. sharpi sharpi in Chiba Prefecture, Japan. One was a place where a high proportion of grooved females lived, and the others had high proportions of smooth females. After five to eight generations of beetles from two populations with different proportions of grooved females were reared under aquarium conditions constituting a common garden design, i.e., water temperature, water depth, and presence of a plant for oviposition, the differences remained. We mated smooth virgin females of D. sharpi sharpi with males of D. sharpi validus to obtain hybrid offspring. The elytral traits of the hybrid females produced only grooved forms. These results suggested that the female dimorphism is determined by genetics, and that the grooved morph was dominant over the smooth one, independent of environmental factors. In addition, the hybrid insects did not differ from the two subspecies insects in larval survivorship, pupation success, or sex ratio. They also showed neither morphological abnormality nor reduced survival.     

Structure in temperate predaceous diving beetle communities (Coleoptera: Dytiscidae)

Collection records of adult dytiscid beetles (Coleoptera, Dytiscidae) from 312 sites within the province of Alberta, Canada, were analyzed for patterns of similarity in species distribution. Sites compared on the basis of species occurrence (Jaccard coefficient of similarity and group average cluster analysis) separated into 12 principal clusters, interpreted as communities. However, cluster inclusion occurred at very low levels of similarity and 30% of sites were unclustered outliers. Ecological charade us tics of sites that were recognized as varying between dusters were salinity, productivity, stability, water temperature, substrate type, flow and vegetation. An index relating these qualitatively defined variables to species numbers is presented. An analysis of co-occurrence patterns of species was conducted as an alternative approach to definition of multispecies associations. Each possible combination of co-occurrence between pairs of species occurring in more than 5 collections each was tested for significance (p = 0.005) by means of Yates corrected χ² tests. Results of tests were expressed in binary terms, blank – no significance, x - significant. Species were then clustered on the basis of patterns of significant co-occurrences as above. This analysis was used as the basis for ordination of species in a symmetrical matrix with species ordered so as to maximize density of points of significant co-occurrence (x) around the principal diagonal. Areas in the matrix showing high densities of co-occurrence were interpreted as representing communities. The cluster analysis of species produced a better cluster pattern than obtained for sites but while generally similar, the species clustering pattern was less fine. The principal species groups consisted of those of saline water, alpine/subalpine lotic sites, other lotic habitats, and lentic habitats with two subgroups, species in forest areas and grassland species. The ordination, while reflecting this grouping of species, demonstrated that each group graded into others through loss of certain species and acquisition of others and thus showed the continuum nature of these communities. Restricting ordination to congeneric species indicated that these do not usually show identical patterns of co-occurrence but neither are patterns coincident with mutual exclusion. Within genera, considerable overlap in occurrence between species is the norm but species tend to occupy slightly different positions relative to one another along qualitatively defined ecological gradients. Relative position of species in these ordinations shows good correlation with patterns of geographical distribution lending support to the interpretation of ordination patterns reflecting relative ecological segregation. The dytiscid beetle fauna of north temperate regions is especially rich. Also, the large number of significant co-occurrences observed between species shows that species packing in many habitats of this region is dense. It is speculated that this species richness is possible because of the seasonality of habitats at these latitudes which reduces competition and predation from other groups and produces seasonal pulses of high productivity permitting a species rich fauna to develop because of abundant resources. It is likely that factors other than interspecific competition for resources are important in shaping dytiscid communities in habitats with strong nutrient pulses.     

Female dimorphism in Japanese diving beetle Dytiscus marginalis czerskii (Coleoptera: Dytiscidae) evidenced by mitochondrial gene sequence analysis

Three species of the Japanese diving beetle Dytiscus have been identified: D. dauricus Gebler, 1832; D. marginalis czerskii Zeitzev, 1953; and D. sharpi. At present, the latter consists of the subspecies D. sharpi sharpi Wehncke, 1875 and D. sharpi validus Régimbart, 1899 based on the comparative data of mitochondrial DNA cytochrome‐c oxidase I (COI) sequences. Many Dytiscus species have smooth and grooved elytra, which are female dimorphic traits. For many years it has been thought that Japanese D. marginalis czerskii has a single morph, that is, only grooved females, although there were some collecting reports of smooth females occurring at the foot of Mt. Chokaisan in Yamagata and Akita Prefectures. However, the population of smooth females (smooth population) has not yet been identified by DNA markers. To understand the species status of the smooth population, we sequenced 769 bp of COI of a male derived from a smooth mother insect and compared it with the sequence from a known grooved female. The sequences of 769 bp of the COI gene in the smooth population were identical to that in the grooved female, indicating that Japanese D. marginalis czerskii has female dimorphic traits.     

Temperature-dependent regulation of reproduction in the diving beetle Dytiscus sharpi (Coleoptera: Dytiscidae)

The effects of temperature on the mating behavior, gonad development, germ cell maturation, and egg spawning of the predaceous diving beetle Dytiscus sharpi (Coleoptera; Dytiscidae), were investigated. By field observations, we found that mating behavior started in October and occurred more frequently from November to December. Under our laboratory breeding conditions, we observed almost the same seasonal variation in mating behavior. We found that temperatures lower than 20 degrees C were required to trigger mating behavior. We also found the same temperature threshold triggered gonadogenesis as well as spermatogenesis. Furthermore, for females, exposure to lower temperatures (<8 degrees C) during the winter was required for egg maturation and spawning in spring; that is, there was a second threshold for successful female reproduction. We conclude that the termination of summer reproductive diapause of D. sharpi is regulated in a temperature-dependent manner, thus effecting the adaptation of D. sharpi to southern warm habitats.     

Functional types of diving beetle (Coleoptera: Hygrobiidae and Dytiscidae), as identified by comparative swimming behaviour

The swimming behaviour of ten species of diving beetles was studied with a video image analysing system, with the aim of testing the interpretation of their size and shape as functional characters reflecting adaptations to different swimming strategies. Velocity, sinuosity of the trajectory, and the relation between the two were studied in an unobstructed aquarium and, for the four largest species, in an aquarium with vertical sticks. Species predicted to be poor swimmers had the lowest average and maximum velocities. A globular species considered to have high manoeuvrability swam in highly sinuous trajectories, and could maintain this sinuosity at a wide range of velocities. One of the larger species, considered to be adapted to high speed swimming, also swam in highly sinuous trajectories, but only at slow velocities; its swimming pattern was considered to be the product of behavioural rather than morphological constraints. For two of the largest species, there was a significant decrease in sinuosity at higher velocities, whilst another was found to have a good compromise between velocity and manoeuvrability. In the aquarium with obstacles all the species reduced their maximum velocity, while the effect on sinuosity varied between species. Although the space limitation of the aquarium and the lack of motivation could have prevented some species from reaching their maximal velocities, a good agreement was found between the predicted and the actual swimming characteristics.     

Evolution of salinity tolerance in the diving beetle tribe Hygrotini (Coleoptera,Dytiscidae)

Some species of the diving beetle tribe Hygrotini (subfamily Hydroporinae) are among the few insects able to tolerate saline concentrations more than twice that of seawater. However, the phylogenetic relationships of the species of Hygrotini, and the origin and evolution of tolerance to salinity in this lineage, are unknown. In this work, we aim to reconstruct how many times salinity tolerance did evolve in Hygrotini, whether this evolution was gradual or if tolerance to hypersalinity could evolve directly from strictly freshwater (FW) species, and to estimate the probabilities of transition between habitats. We build a phylogeny with ca. 45% of the 137 species of Hygrotini, including all major lineages and almost all of the known halophile or tolerant species. We used sequence data of four mitochondrial (COI‐5′, COI‐3′, 16S + tRNA and NADH1) and three nuclear (28S, 18S and H3) gene fragments, plus ecological data to reconstruct the history of the salinity tolerance using Bayesian inference. Our results demonstrate multiple origins of the tolerance to salinity, although most saline and hypersaline species were concentrated in two lineages. The evolution of salinity was gradual, with no direct transitions from FW to hypersaline habitats, but with some reversals from tolerant to FW species. The oldest transition to saline tolerance, at the base of the clade with the highest number of saline species, was dated in the late Eocene‐early Oligocene, a period with decreasing temperature and precipitation. This temporal coincidence suggests a link between increased aridity and the development of tolerance to saline waters, in agreement with recent research in other groups of aquatic Coleoptera.     

Eye and optic lobe metamorphosis in the sunburst diving beetle, Thermonectus marmoratus (Coleoptera: Dytiscidae)

Nearly nothing is known about the transition that visual brain regions undergo during metamorphosis, except for Drosophila in which larval eyes and the underlying neural structure are strongly reduced. We have studied the larvae of the sunburst diving beetle, Thermonectus marmoratus (Coleoptera: Dytiscidae), which are sophisticated visually oriented predators characterized by six elaborate stemmata on each side of the head and an associated large optic lobe. We used general neurohistological staining and 3D reconstruction to determine how the eyes and optic lobe of T. marmoratus change morphologically during metamorphosis. We find that in third (last) instar larvae, the adult neuropils are already forming de novo dorsally and slightly anteriorly to the larval neuropils, while the latter rapidly degenerate. Larval eyes are eventually reduced to distinct areas with dark pigmentation. This complete reorganization, which may be an evolutionarily conserved trait in holometabolous insects, occurs despite the considerable costs that must apply to such a visually complex animal. Our findings are consistent with the concept that stemmata are homologous to the most posterior ommatidia of hemimetabolous insects, an idea also recently supported by molecular data.     

A phylogenetic analysis of the evolution of sexual dimorphism and mating systems in water striders (Hemiptera: Gerridae)

Conflicts over mating decisions characterize the sexual behaviour of many insects, in particular when males encounter females that already carry enough sperm to fertilize their eggs, since a mating often will inflict greater costs than benefits upon females. Therefore, coevolutionary models predict adaptation and counter-adaptation by the sexes in a battle to control the outcome of sexual encounters. A phylogenetic analysis was performed on patterns of sexual dimorphism and mating systems within water striders (Hemiptera, Gerridae). Phylogenetic effects or 'constraints' have significantly shaped patterns of sexual dimorphism in female/ male size ratios, legs and genitalia of males, and the structure of the female abdomen. Males of ancestral gerrids were probably slightly smaller than conspecific females, had powerful fore legs adapted to grasp the female's thorax during mating, and had clasping genitalic structures suited to grasp or pinch the female posteriorly. Most gerrids have a female/male size ratio between 1.05 and 1.14, but more pronounced sexual size ratios (above 1.25) have independently evolved several times in the family, usually in association with extended post-copulatory mate guarding. The comparative, phylogenetic analysis suggests coevolution of female anticlasper and male clasping devices for the clade comprising the subfamilies Cylindrostethinae, Ptilomerinae, and Halobatinae while female anticlasper devices have evolved in the absence of male clasping genitalia in the Gerrinae. The ancestral and most common mating system in gerrids is 'scramble competition polygyny' from which has evolved 'resource defence polygyny' at least four times independently of each other. The phylogenetic effects on patterns of mating behaviour are much less obvious, as exemplified by the large amount of interspecific variation in some genera.     

Range expansion and ancestral niche reconstruction in the Mediterranean diving beetle genus Meladema (Coleoptera,Dytiscidae)

Species of the genus Meladema (Dytiscidae, Colymbetinae) are some of the largest macroinvertebrates in the western Palearctic region, being top predators in fishless streams. Two of the three described species, Meladema imbricata (Wollaston, 1871) and Meladema lanio (Fabricius, 1775) are Macaronesian endemics from the Canary Islands and Madeira, respectively, while the third, Meladema coriacea Laporte, 1835, is widely distributed from Morocco and the Iberian Peninsula to Turkey, including the Canary Islands. Previous phylogenetic analysis using only mitochondrial markers revealed the existence of two cryptic lineages within M. coriacea, one restricted to Corsica and the other including the rest of sampled populations. We reconstruct here the evolutionary history of the species of Meladema using a more comprehensive sampling covering its whole geographical range, adding nuclear markers and Bayesian molecular dating. Using environmental niche modelling, we test for possible differences in climatic preferences among lineages and reconstruct their ancestral climatic niche. Our results strongly supported the existence of four monophyletic lineages represented by the three recognized species plus a fourth cryptic lineage with populations of M. coriacea from the Tyrrhenian islands (Corsica, Sardinia and Montecristo). This pattern is not likely to be the result of mitochondrial artefacts due to Wolbachia infection, as all 11 tested individuals were negative for this parasite. Dating analysis placed the origin of Meladema in the Middle Miocene although diversification among extant Meladema lineages started in the early Pleistocene and took place in a relatively short time period. Phylogeographic analysis inferred a continental origin of Meladema, with an independent colonization of the Macaronesian and Mediterranean islands. From the south‐western Mediterranean region, the continental M. coriacea expanded its range up to Bulgaria and Turkey in the northern basin and to Tunisia in the southern. Results of niche modelling showed that seasonality is the critical factor in shaping the current distribution of Meladema. Island lineages (M. imbricata, M. lanio and the Tyrrhenian lineage of M. coriacea) occur in sites with low seasonality, within the range of the reconstructed ancestral climatic niche of the genus. On the contrary, continental M. coriacea expanded its range to localities outside the ancestral climatic range of the genus, with a higher seasonality and aridity.     

Predaceous diving beetles (Coleoptera: Dytiscidae) of Jordan

The first account of the predaceous diving beetles or Dytiscidae of Jordan is presented. Altogether 25 species are listed, although the occurrence of two of them remains doubtful. Nine species are recorded from the territory of Jordan for the first time, and the first precise locality data are provided for an additional seven species. Bidessus anatolicus Wewalka, 1972 and Nebrioporus stearinus (Kolenati, 1845) recorded recently from Jordan based on misidentification are removed from the list. If available, habitat information is provided for recently collected species. The zoogeographical patterns of the region are briefly summarised.     

Phylogeny and diversification of diving beetles (Coleoptera: Dytiscidae)

Dytiscidae is the most diverse family of beetles in which both adults and larvae are aquatic, with examples of extreme morphological and ecological adaptations. Despite continuous attention from systematists and ecologists, existing phylogenetic hypotheses remain unsatisfactory because of limited taxon sampling or low node support. Here we provide a phylogenetic tree inferred from four gene fragments (cox1, rrnL, H3 and SSU, ≈ 4000 aligned base pairs), including 222 species in 116 of 174 known genera and 25 of 26 tribes. We aligned ribosomal genes prior to tree building with parsimony and Bayesian methods using three approaches: progressive pair‐wise alignment with refinement, progressive alignment modeling the evolution of indels, and deletion of hypervariable sites. Results were generally congruent across alignment and tree inference methods. Basal relationships were not well defined, although we identified 28 well supported lineages corresponding to recognized tribes or groups of genera, among which the most prominent novel results were the polyphyly of Dytiscinae; the grouping of Pachydrini with Bidessini, Peschetius with Methlini and Coptotomus within Copelatinae; the monophyly of all Australian Hydroporini (Necterosoma group), and their relationship with the Graptodytes and Deronectes groups plus Hygrotini. We found support for a clade formed by Hydroporinae plus Laccophilini, and their sister relationship with Cybistrini and Copelatinae. The tree provided a framework for the analysis of species diversification in Dytiscidae. We found a positive correlation between the number of species in a lineage and the age of the crown group as estimated through a molecular clock approach, but the correlation with the stem age was non‐significant. Imbalances between sister clades were significant for several nodes, but the residuals of the regression of species numbers with the crown age of the group identified only Bidessini and the Coptotomus + Agaporomorphus clade as lineages with, respectively, above and below expected levels of species diversity. © The Willi Hennig Society 2008.     

Predaceous diving beetle, Dytiscus sharpi sharpi (Coleoptera: Dytiscidae) larvae avoid cannibalism by recognizing prey

Larvae of diving beetles such as the various Dytiscus species (Coleoptera: Dytiscidae) are carnivorous and usually prey on other aquatic animals. Cannibalism among larvae of Dytiscus sharpi sharpi (Wehncke) was observed to begin when they were starved for more than two days under artificial breeding conditions. However, the 2-day starved larvae did not show cannibalism in the presence of intact, motionless, frozen tadpoles, or frozen shrimps. The beetle larvae attacked and captured intact tadpoles faster (15 sec) than other motionless and frozen tadpoles (120 sec), indicating that prey movement was an important factor in stimulating feeding behavior in larvae. Prey density does not have an effect on larval cannibalism. In cases in which preys are present at lower densities than that of larvae, a group of beetle larvae frequently fed on single prey. This feeding behavior, therefore, provides direct evidence of self-other recognition at the species level. Using two traps in one aquarium that allows the larvae to detect only prey smell, one containing tadpoles and another empty, the beetle larvae were attracted to the trap with tadpoles at high frequency, but not to the empty trap. In another experiment, the beetle larvae were not attracted to the trap containing a beetle larva. These results suggest that the larvae of D. sharpi sharpi are capable of recognizing prey scent, which enables the promotion of foraging behavior and the prevention of cannibalism.     

Preference of oviposition plant and hatchability of the diving beetle,Dytiscus sharpi (Coleoptera: Dytiscidae) in the laboratory

For the conservation of the diving beetle, Dytiscus sharpi (Wehncke) (Coleoptera: Dytiscidae), which is included in the Red List of Japan, it is critical to understand its ecological background. In the present study, oviposition preference and hatchability were investigated under laboratory conditions. Possible candidates of oviposition plants were observed in the natural breeding habitats of D. sharpi. Eight aquatic plants were identified, and Oenanthe javanica (Blume) was the major species present in March, when first instar larvae appeared. The number of eggs laid was investigated under the following conditions: (i) eight field plants were authentically recreated at a similar covering rate to that in the natural habitat; (ii) each of the eight plants was set at the same covering rate; (iii) a single plant taken from the field was placed separately. Significant oviposition preference for O. javanica was found when these eight aquatic plants were present in the same aquaria. In addition, hatchability in O. javanica was significantly higher than that in any of the other plants. These results suggest that O. javanica is an important aquatic plant for oviposition by D. sharpi.     

Sex-specific niche partitioning and sexual size dimorphism in Australian pythons (Morelia spilota imbricata)

Sexual dimorphism is usually interpreted in terms of reproductive adaptations, but the degree of sex divergence also may be affected by sex-based niche partitioning. In gape-limited animals like snakes, the degree of sexual dimorphism in body size (SSD) or relative head size can determine the size spectrum of ingestible prey for each sex. Our studies of one mainland and four insular Western Australian populations of carpet pythons ( Morelia spilota ) reveal remarkable geographical variation in SSD, associated with differences in prey resources available to the snakes. In all five populations, females grew larger than males and had larger heads relative to body length. However, the populations differed in mean body sizes and relative head sizes, as well as in the degree of sexual dimorphism in these traits. Adult males and females also diverged strongly in dietary composition: males consumed small prey (lizards, mice and small birds), while females took larger mammals such as possums and wallabies. Geographic differences in the availability of large mammalian prey were linked to differences in mean adult body sizes of females (the larger sex) and thus contributed to sex-based resource partitioning. For example, in one population adult male snakes ate mice and adult females ate wallabies; in another, birds and lizards were important prey types for both sexes. Thus, the high degree of geographical variation among python populations in sexually dimorphic aspects of body size and shape plausibly results from geographical variation in prey availability.  © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 77 , 113–125.     

Evolution of the Juan Fernández diving beetle,Rhantus selkirki (Coleoptera,Dytiscidae)

Here we provide evidence that confinement in Robinson Crusoe Island (located about 660 km west of continental Chile) over evolutionary time leads to strong morphological modifications in diving beetle (Dytiscidae) larvae. We analysed a large set of morphological larval characters for all currently recognised genera of Colymbetinae as a framework, to infer phylogenetic relationships within the large genus Rhantus Dejean, 1833 and, in particular, of the charismatic Juan Fernández diving beetle, Rhantus selkirki Jäch, Balke & Michat, 2015, comparing our results with a recent phylogeny of the Colymbetinae based on DNA sequence data. We suggest that adaptation to the island's particular habitats resulted in the reversal of certain characters of R. selkirki back to the plesiomorphic states. This may cause the species to be erroneously interpreted as more ‘primitive’ if only morphological characters are analysed. Confinement in the particular, shallow and barely vegetated aquatic habitats of Robinson Crusoe Island for a long time seems to have led to this divergent morphology, particularly in characters related to swimming ability such as several leg and urogomphal setae. In this way, R. selkirki larvae secondarily resemble those of some earlier diverging dytiscid lineages such as Agabinae and Copelatinae, which typically creep on the bottom of water bodies and do not swim well.     

Retinal ultrastructure may mediate polarization sensitivity in larvae of the Sunburst diving beetle, Thermonectus marmoratus (Coleoptera: Dytiscidae)

A number of invertebrates are known to be sensitive to the polarization of light and use this trait in orientation, communication, or prey detection. In these animals polarization sensitivity tends to originate in rhabdomeric photoreceptors that are more or less uniformly straight and parallel. Typically, polarization sensitivity is based on paired sets of photoreceptors with orthogonal orientation of their rhabdomeres. Sunburst diving beetle larvae are active swimmers and highly visual hunters which could potentially profit from polarization sensitivity. These larvae, like those of most Dytiscids, have a cluster of six lens eyes or stemmata (designated E1 through E6) on each side of the head capsule. We examined the ultrastructure of the photoreceptor cells of the principal eyes (E1 and E2) of first instar larvae to determine whether their rhabdomeric organization could support polarization sensitivity. A detailed electron microscopical study shows that the proximal retinas of E1 and E2 are in fact composed of photoreceptors with predominantly parallel microvilli and that neighboring rhabdomeres are oriented approximately perpendicularly to one another. A similar organization is observed in the medial retina of E1, but not in the distal retinas of E1&2. Our findings suggest that T. marmoratus larvae might be able to analyze polarized light. If so, this could be used by freshly hatched larvae to find water or within the water to break the camouflage of common prey items such as mosquito larvae. Physiological and behavioral tests are planned to determine whether larvae of T. marmoratus can actually detect and exploit polarization signals.