Morphological Study of Left-Right Head Asymmetry in Doubledaya bucculenta (Coleoptera: Erotylidae: Languriinae).

Left-right asymmetry in paired organs is well documented across various species, including the claws of fiddler crabs and snail-eating snakes' dentition. However, the mechanisms underlying these asymmetries remain largely elusive. This study investigates Doubledaya bucculenta (Coleoptera: Erotylidae), a lizard beetle species known for pronounced left-sided asymmetry in adult female mandible and gena. Given that insect mouthparts comprise multiple functionally significant appendages, we aimed to clarify the degree of asymmetry extending beyond the mandibles and genae. Phenotypic morphology was assessed through trait measurement and asymmetry index calculations. Our detailed morphometric analyses revealed left-longer asymmetry not only in mandibles and genae but also in maxillae and labium. Notably, the degree of asymmetry in other mouthparts was generally less pronounced compared to that in outer mandibles, suggesting a potential influence of left mandible development on other mouthparts. Additionally, male mandibles exhibited region-specific asymmetry, potentially indicative of constrained evolutionary adaptations. This study enhances a comprehensive understanding of adult phenotype morphology and offers insights into the developmental basis of asymmetrical mouthparts.

Development of the Pronotal Explanate Margin, a Novel Evolutionary Trait in Tortoise Beetles.

Most tortoise beetles, belonging to the subfamily Cassidinae (Coleoptera: Chrysomelidae), possess distinctive explanate margins, comprising elongations of the pronotum and elytra outer margins. These margins flatten against the ground, serving as a unique defensive mechanism against predators. To understand the developmental and evolutionary origins of explanate margins, we examined the development of the pronotal part of these structures in two tortoise beetle species: Thlaspida biramosa (Boheman) (tribe Cassidini) and Laccoptera nepalensis (Boheman) (tribe Aspidimorphini). Although final (fifth) instar larvae of both species exhibited no external prothoracic structures associated with explanate margins, pupae possessed a plate-shaped structure projecting anterolaterally on their pronotum. This plate-shaped structure was identified as the pupal primordium of the pronotal explanate margin, as the explanate margin emerged from inside the structure during eclosion. In prepupae of T. biramosa, the primordial tissue exhibited three region-specific folding and furrowing patterns, beneath larval cuticles. These epithelial structures expanded within minutes at the onset of pupation, resulting in pupal primordial formation. Thus, pronotal explanate margins originate from pronotal epithelia, with the furrowing and folding patterns of the epithelia shaping pupal and even adult pronotal explanate margins. The presence of morphologically similar pupal pronotal projections in various Cassidinae suggests that the development of pronotal explanate margins is likely common in these beetles.

DNA but not always morphology help to recognise monophyletic genera within 'Cercyon' terrestrial water scavenger beetles: a case study of Asiacyon gen. nov. (Coleoptera: Hydrophilidae).

DNA-based studies have revealed that the terrestrial water scavenger beetle genus Cercyon Leach, 1817 (Coleoptera: Hydrophilidae: Sphaeridiinae: Megasternini) is polyphyletic, grouping similarly looking but unrelated species that were not assigned to other genera due to the absence of unusual morphological characters. In this study, we analyse the morphology, DNA data and species diversity of one of the Asian clades of 'Cercyon ' to test whether a natural, phylogeny-based generic classification can be established. We add DNA data (five nuclear and three mitochondrial fragments) for additional species and specimens of the clade to test its monophyly and reveal phylogenetic relationships among species. We perform a detailed morphological study of all species, including SEM micrographs, to reveal synapomorphies of the DNA-based clades. We demonstrate that the lineage, described here as Asiacyon Mai, Jia, Ryndevich & Fikácek, gen. nov., is strongly supported by DNA data, has limited distribution (eastern Asia), and its species share similar biology (inhabiting fresh or decaying plant tissues), though it can be only diagnosed by a combination of plesiomorphic characters. A detailed treatment is provided for the Chinese species, including species diagnoses, illustrations of habitus and male genitalia, (re)descriptions and a key to species. A total of 14 Chinese species are recognised, of which 11 are described as new: Asiacyon pax sp. nov. (Guangdong, Hong Kong, Hunan), A. pseudincretus sp. nov. (Anhui, Jiangxi, Taiwan, Zhejiang), A. vicincretus sp. nov. (Guangdong, Guangxi), A. zhengyucheni sp. nov. (Yunnan), A. huilanae sp. nov. (Yunnan), A. cornipenis sp. nov. (Yunnan), A. belousovi sp. nov. (Yunnan), A. liangchengi sp. nov. (Yunnan), A. paraequalis sp. nov. (Yunnan), A. pengzhongi sp. nov. (Hunan) and A. xiuzhenae sp. nov. (Hainan, Yunnan). Three Chinese species previously classified in Cercyon are transferred to Asiacyon : A. incretus (d'Orchymont, 1941), comb. nov., A. primoricus (Ryndevich & Prokin, 2017), comb. nov. (newly recorded from China) and A. indicus (d'Orchymont, 1926), comb. nov. (newly recorded from China and Laos). Additionally, seven Asian species previously classified in Cercyon are transferred to Asiacyon based on their external morphology, but not studied in detail: A. aequalis (Sharp, 1884), comb. nov., A. conjiciens (Walker, 1858), comb. nov., A. dilutus (Régimbart, 1903), comb. nov., A. placidus (Sharp, 1884), comb. nov., A. pseudodilutus (Satô, 1979), comb. nov., A. retius (Ryndevich & Prokin, 2017), comb. nov., and A. rubicundus (Sharp, 1884), comb. nov. In total, the genus now comprises 21 named species and several undescribed species from India, Myanmar and Indonesia. ZooBank: urn:lsid:zoobank.org:pub:E3C949A0-34E4-46EF-BA6A-2CD048D583B2.

Integrative taxonomy of Teucriogethes pollen beetles (Coleoptera: Nitidulidae: Meligethinae), with implications on the systematics of the genus Teucrium (Lamiaceae).

In highly specialised phytophagous insects, important insights on phylogeny of the involved insect group can be often inferred from the evolutionary history of their host plants, and vice-versa, as in the case for a new, peculiar species of pollen beetles, recently discovered in Central China (Hubei Province). This new species is described herein as Teucriogethes huangae Liu & Audisio, sp. nov., and information on its ecology and systematic position is provided. The new species, associated as larvae with a strongly isolated member of the genus Teucrium L. (Lamiaceae), T. ornatum Hemsl., exhibiting a problematic taxonomic position, represents in turn an isolated and morphologically rather aberrant taxon in its lineage, probably the largest in size (2.2-2.8mm), and the first endemic representative of its genus in the whole Eastern Palaearctic. An updating of the taxonomic assessment of members of Teucriogethes and on possible evolutionary relationships with their larval host plants, based on an integrative taxonomy approach, is finally presented. A key to identification of all known species is also introduced. ZooBank: urn:lsid:zoobank.org:pub:E349044B-29A5-416C-85CA-56143C7A29BE.

An unusual Cretaceous beetle with affinity to Anamorphidae (Coleoptera: Coccinelloidea).

Coccinelloid beetles have a sparse fossil record in the Mesozoic. Here, we describe and illustrate an unusual coccinelloid beetle, Yassibum yoshitomii gen. et sp. nov., from mid-Cretaceous Kachin amber. Yassibum stands out within the Coccinelloidea due to its notched profemora and the presence of antennal grooves on the elytral epipleura. Based on our phylogenetic analyses, we suggest that Yassibum is most likely related to the family Anamorphidae. The alternative placements are critically evaluated based on our comparison of the morphology.

Effect of temperature on sexual size dimorphism during the developmental period in the broad-horned flour beetle.

Adult size in numerous insects is strongly dependent on temperature. In several cases, a temperature-size rule is observed in which developmental temperature and adult size tradeoff. Although several previous studies have demonstrated the temperature-size rule, only a few have explored the relationship between developmental temperature and weapon traits or sexual size dimorphism. This study was conducted to investigate the size of the broad-horned flour beetle Gnatocerus cornutus when it was developed under different temperatures. G. cornutus males possess weapon traits for male-male combat and exhibit sexual size dimorphism in other morphological traits. Results showed that male weapon size and body size complied with the temperature-size rule. Furthermore, the extent of sex dimorphism in genae width, a weapon-supportive trait, were larger at lower temperatures. Our findings suggest that the temperature-size rule also influences the size of sexual traits.

Taxonomic Revision of the Dichotomius mormon (Ljungh, 1799) Species Group (Coleoptera: Scarabaeidae: Scarabaeinae), with a Description of a New Species from Bolivia.

Dichotomius Hope, 1838 (Coleoptera: Scarabaeidae: Scarabaeinae) is a very abundant and diverse genus of dung beetles of the New World ecosystems, with about 190 species, distributed in four subgenera. Luederwaldt (Separata Rev Mus Paul 14: 3-13 1929) proposed the division of subgenera into sections (now species groups) based mainly on characters of external morphology. Based on Luederwaldt's proposal, progress has been made in recent years in the taxonomic revision of the genus; however, inconsistencies have been found in the subgeneric division and species groups proposed by this author. Based on the external morphology and the male genital organ of the all type material and other material deposited in fifteen entomological collections, in this paper, the redefinition and taxonomic revision of the Dichotomius mormon species group is carried out. The new definition of the group and the species key are presented. Five species are included in the group: Dichotomius conicollis (Blanchard, 1846) (Bolivia), Dichotomius larseni sp. nov. (Bolivia), Dichotomius mormon (Ljungh, 1799) (Argentina, Brazil, and Paraguay), Dichotomius ohausi (Luederwaldt, 1923) (Brazil, Colombia, Ecuador, and Peru), and Dichotomius talaus (Erichson, 1847) stat. rev. (Bolivia, and Peru). The lectotypes for D. conicollis, Copris ephialtes Mannerheim, 1829, Copris nasuta Castelnau, 1840, Copris protensus Perty, 1830, D. ohausi, and D. talaus were designated. For each of the species that are included in the group, the following information is presented: taxonomic history, species' citation in published literature, description or redescription of males and females, list of material examined, photographs of the external morphology, illustrations of male genital organ and its endophallites, and distribution map.

Genomic, morphological and physiological data support fast ecotypic differentiation and incipient speciation in an alpine diving beetle.

An intricate interplay between evolutionary and demographic processes has frequently resulted in complex patterns of genetic and phenotypic diversity in alpine lineages, posing serious challenges to species delimitation and biodiversity conservation planning. Here we integrate genomic data, geometric morphometric analyses and thermal tolerance experiments to explore the role of Pleistocene climatic changes and adaptation to alpine environments on patterns of genomic and phenotypic variation in diving beetles from the taxonomically complex Agabus bipustulatus species group. Genetic structure and phylogenomic analyses revealed the presence of three geographically cohesive lineages, two representing trans-Palearctic and Iberian populations of the elevation-generalist A. bipustulatus and another corresponding to the strictly-alpine A. nevadensis, a narrow-range endemic taxon from the Sierra Nevada mountain range in southeastern Iberia. The best-supported model of lineage divergence, along with the existence of pervasive genetic introgression and admixture in secondary contact zones, is consistent with a scenario of population isolation and connectivity linked to Quaternary climatic oscillations. Our results suggest that A. nevadensis is an alpine ecotype of A. bipustulatus, whose genotypic, morphological and physiological differentiation likely resulted from an interplay between population isolation and local altitudinal adaptation. Remarkably, within the Iberian Peninsula, such ecotypic differentiation is unique to Sierra Nevada populations and has not been replicated in other alpine populations of A. bipustulatus. Collectively, our study supports fast ecotypic differentiation and incipient speciation processes within the study complex and points to Pleistocene glaciations and local adaptation along elevational gradients as key drivers of biodiversity generation in alpine environments.

Functional morphology of antennae and sensilla of the fungivore beetle, Triplax ainonia Lewis (Coleoptera: Erotylidae).

The antennal sensilla play an important role in many behavioral activities of insects. The fungivorous beetle Triplax ainonia Lewis (Erotylidae) is an important pest which prefers to feed on Pleurotus mushrooms. In order to clarify the types, number, and distribution of the antennal sensilla of male and female T. ainonia, scanning electron microscopy was used. The results showed that there were five sensillum types on the antennae of adults male and female, including Böhm's bristles (BB), sensilla chaetica (three subtypes: SC 1, SC 2, and SC 3), sensilla basiconica (three subtypes: SB 1, SB 2, and SB 3), sensilla trichodea (ST), and sensilla styloconica (SS). Among all the sensilla, the number of SB 2 was the most abundant in both sexes. We found that there was no sexually dimorphic in the sensillum types, but there were differences in the number, lengths, and diameters of some sensilla between males and females. Based on the information of the morphology and distribution of the sensilla, the potential functions of the antennal sensilla of T. ainonia adults were discussed. The results of this study provide a basis for further study on the behavioral ecology and electrophysiology of the fungivore beetles of the Erotylidae.

The morphology of the pygidial glands and the chemical composition of their secretions of four sphodrine ground beetle species (Carabidae: Platyninae).

Ground beetles possess a pair of pygidial glands that produce and release secretions that play an important role in defense against predators. The morphology of these glands and the chemical composition of their products were studied in four species of the tribe Sphodrini: Calathus (Calathus) fuscipes (Goeze, 1777), C. (Neocalathus) cinctus Motschulsky, 1850, C. (N.) melanocephalus (Linnaeus, 1758) and Laemostenus (Antisphodrus) elongatus (Dejean, 1828). The morphological analyzes of the glands of the four taxa mentioned were carried out for the first time using bright-field and nonlinear microscopy. All morphological structures were precisely measured and photographed. The pygidial gland secretions of C. (C.) fuscipes and L. (A.) elongatus were analyzed for the first time using gas chromatography-mass spectrometry. A total of 30 compounds were detected from the extracts of pygidial gland secretions of the four Sphodrini species studied. The simplest chemical mixture was found in L. (A.) elongatus, while the most complex secretion was that of C. (C.) fuscipes. 1-Undecanol, which we were able to detect in all taxa examined here, and dodecyl butyrate, which was detected in the three Calathus species, have never before been detected in the secretions of ground beetles.

Passive wing deployment and retraction in beetles and flapping microrobots.

Birds, bats and many insects can tuck their wings against their bodies when at rest and deploy them to power flight. Whereas birds and bats use well-developed pectoral and wing muscles1,2, how insects control their wing deployment and retraction remains unclear because this varies among insect species. Beetles (Coleoptera) display one of the most complex mechanisms. In rhinoceros beetles, Allomyrina dichotoma, wing deployment is initiated by complete release of the elytra and partial release of the hindwings at their bases. Subsequently, the beetle starts flapping, elevates the hindwing bases and unfolds the hindwing tips in an origami-like fashion. Although the origami-like fold has been extensively explored3-8, limited attention has been given to the hindwing base movements, which are believed to be driven by the thoracic muscles5,9-11. Here we demonstrate that rhinoceros beetles can effortlessly deploy their hindwings without necessitating muscular activity. We show that opening the elytra triggers a spring-like partial release of the hindwings from the body, allowing the clearance needed for the subsequent flapping motion that brings the hindwings into the flight position. After flight, the beetle can use the elytra to push the hindwings back into the resting position, further strengthening the hypothesis of passive deployment. We validated the hypothesis using a flapping microrobot that passively deployed its wings for stable, controlled flight and retracted them neatly upon landing, demonstrating a simple, yet effective, approach to the design of insect-like flying micromachines.

A New Species of Megaceropsis Dechambre, 1976 and a New Record of Megaceropsis quadridentata Dechambre, 1976 (Coleoptera, Melolonthidae, Dynastinae, Oryctini).

Megaceropsis Dechambre, 1976 (Coleoptera, Melolonthidae, Dynastinae, Oryctini) is a South American genus with two known species until now: Megaceropsis quadridentata Dechambre, 1976 and Megaceropsis lecourti Dechambre, 1996. We describe a third species herein: Megaceropsis kleytoni sp. nov., from Brazil. Illustrations, a distributional map, and an identification key including all Megaceropsis species are provided. Additionally, a first record of M. quadridentata from Brazil is presented.

Evolution across the adaptive landscape in a hyperdiverse beetle radiation.

The extraordinary diversification of beetles on Earth is a textbook example of adaptive evolution. Yet, the tempo and drivers of this super-radiation remain largely unclear. Here, we address this problem by investigating macroevolutionary dynamics in darkling beetles (Coleoptera: Tenebrionidae), one of the most ecomorphologically diverse beetle families (with over 30,000 species). Using multiple genomic datasets and analytical approaches, we resolve the long-standing inconsistency over deep relationships in the family. In conjunction with a landmark-based dataset of body shape morphology, we show that the evolutionary history of darkling beetles is marked by ancient rapid radiations, frequent ecological transitions, and rapid bursts of morphological diversification. On a global scale, our analyses uncovered a significant pulse of phenotypic diversification proximal to the Cretaceous-Palaeogene (K/Pg) mass extinction and convergence of body shape associated with recurrent ecological specializations. On a regional scale, two major Australasian radiations, the Adeliini and the Heleine clade, exhibited contrasting patterns of ecomorphological diversification, representing phylogenetic niche conservatism versus adaptive radiation. Our findings align with the Simpsonian model of adaptive evolution across the macroevolutionary landscape and highlight a significant role of ecological opportunity in driving the immense ecomorphological diversity in a hyperdiverse beetle group.

Odd-Paired is Involved in Morphological Divergence of Snail-Feeding Beetles.

Body shape and size diversity and their evolutionary rates correlate with species richness at the macroevolutionary scale. However, the molecular genetic mechanisms underlying the morphological diversification across related species are poorly understood. In beetles, which account for one-fourth of the known species, adaptation to different trophic niches through morphological diversification appears to have contributed to species radiation. Here, we explored the key genes for the morphological divergence of the slender to stout body shape related to divergent feeding methods on large to small snails within the genus Carabus. We show that the zinc-finger transcription factor encoded by odd-paired (opa) controls morphological variation in the snail-feeding ground beetle Carabus blaptoides. Specifically, opa was identified as the gene underlying the slender to stout morphological difference between subspecies through genetic mapping and functional analysis via gene knockdown. Further analyses revealed that changes in opa cis-regulatory sequences likely contributed to the differences in body shape and size between C. blaptoides subspecies. Among opa cis-regulatory sequences, single nucleotide polymorphisms on the transcription factor binding sites may be associated with the morphological differences between C. blaptoides subspecies. opa was highly conserved in a wide range of taxa, especially in beetles. Therefore, opa may play an important role in adaptive morphological divergence in beetles.

Evolutionary radiation strategy revealed in the Scarabaeidae with evidence of continuous spatiotemporal morphology and phylogenesis.

Evolutionary biology faces the important challenge of determining how to interpret the relationship between selection pressures and evolutionary radiation. The lack of morphological evidence on cross-species research adds to difficulty of this challenge. We proposed a new paradigm for evaluating the evolution of branches through changes in characters on continuous spatiotemporal scales, for better interpreting the impact of biotic/abiotic drivers on the evolutionary radiation. It reveals a causal link between morphological changes and selective pressures: consistent deformation signals for all tested characters on timeline, which provided strong support for the evolutionary hypothesis of relationship between scarabs and biotic/abiotic drivers; the evolutionary strategies under niche differentiation, which were manifested in the responsiveness degree of functional morphological characters with different selection pressure. This morphological information-driven integrative approach sheds light on the mechanism of macroevolution under different selection pressures and is applicable to more biodiversity research.

The development of extremely large male genitalia under spatial limitation.

Extensive research in evolutionary biology has focused on the exaggeration of sexual traits; however, the developmental basis of exaggerated sexual traits has only been determined in a few cases. The evolution of exaggerated sexual traits may involve the relaxation of constraints or developmental processes mitigating constraints. Ground beetles in the subgenus Ohomopterus (genus Carabus) have species-specific genitalia that show coevolutionary divergence between the sexes. Here, we examined the morphogenesis of the remarkably enlarged male and female genitalia of Carabus uenoi by X-ray microcomputed tomography. The morphogenetic processes generating the male and female genitalia at the pupal stage were qualitatively similar to those in closely related species with standard genital sizes. Higher growth rates contributed to the exaggeration of both the male and female genital parts of C. uenoi, possibly related to a gene network commonly upregulated in both sexes. Additionally, the length of the copulatory piece (CP), the enlarged male genital part stored in the aedeagus (AD), reached close to that of the AD at the later developmental stages and thereafter decelerated to grow in parallel with the AD, suggesting a structural constraint on the CP by the outer AD. Then, unlike related species, the lengths of the CP and AD increased at eclosion, suggesting a mechanism leading to further elongation of the male genitalia. These observations suggest that a developmental process allows continuous growth of the male genitalia even under the spatial limitation. These results revealed the spatio-temporal dynamics of the development of exaggerated genital structures under structural constraints.

Extreme range in adult body size reveals hidden trade-offs among sexually selected traits.

Sexually selected weapons used to monopolize mating opportunities are predicted to trade-off with traits used in competition for fertilization. Yet, the limited size range typically found among adults of a species often precludes clear comparisons between population-level and individual-level relative trait investment. The jousting weevil, Brentus anchorago (Coleoptera: Brentidae), varies more than 26-fold in body mass, which is among the most extreme adult body size ranges of any solitary terrestrial species. We reveal a trade-off at a population level: hypermetric scaling in male weapons (slope = 1.59) and a closely mirrored reversal in allocation to postcopulatory traits (slope = 0.54). Yet, at the individual level, we find the opposite pattern; males that invest relatively more in weapons for their size class also invest more in postcopulatory traits. Across 36 dung beetle and 41 brentine weevil species, we find the allometric slope explains more trait variation at larger body size ranges; in brentines, population-level scaling patterns become more detectable in species with a larger range in adult body size. Our findings reveal that population-level allometries and individual-level trade-offs can both be important in shaping relative trait allocation; we highlight that the adult body size range is rarely examined but may be integral to gaining a deeper understanding of trade-offs in reproductive allocation.

Eye morphology in four species of tiger beetles (Coleoptera: Cicindelidae).

Compound eyes undoubtedly represent the widespread eye architecture in the animal kingdom. The insects' compound eye shows a wide variety of designs, and insects use their visual capacity to accomplish several tasks, including avoiding enemies, searching for food and shelter, locating a mate, and acquiring information about the environment and its surroundings. Broad literature data support the concept that visual ability lies in the way the eyes are built. Since the resolution and sensitivity of the compound eye are partly determined by the density of the ommatidia and the size of the facets. Morphological parameters of the compound eyes could influence the function of the visual organ and its capacity to process information, also representing a sensitive indicator of different habitat demands. In this study, we compared compound eyes' parameters in four closely related species of tiger beetles to disclose differences arising from different habitats. Furthermore, to investigate whether there are consistent intersexual differences, we also compared the most relevant parameters of the eye in males and females of four selected species. Our results show sex-related and interspecific differences that occur in examined species.

Sexual dimorphism of Dyschiriini (Coleoptera, Carabidae): Comparative morphological SEM study of palpi sensilla and its possible role in intraspecific chemical communication.

Sexual dimorphism in Dyschiriini (Coleoptera, Carabidae) consists of the presence of an autapomorphic sensory area in apical palpomeres of males, here named as Male Palpi Sensory Area (MPSA). In this work, microstructure of palpi, with focus on MPSA, is characterized and formally described using Scanning Electron Microscopy (SEM). Interspecific variability among 13 species and three subgenera of Dyschirius Bonelli, 1810 and one species of Reicheiodes Ganglbauer, 1891 is assessed. Palpi of studied Dyschiriini presented up to 4 sensilla classes (coeloconica, basiconica, digitiformia, trichodea) in both sexes, while males had one more class (sensilla placodea) found grouped in MPSA. Measurements of sensilla and MPSA are provided. Differences among taxa corresponded to development grade of MPSA and its number of sensilla placodea. The MPSA of Dyschirius (Dyschirius) thoracicus Rossi, 1790 were clearly different to the rest of the studied subgenera and species of Dyschirius and Reicheiodes, whose MPSA were similar and had slight intraspecific variability. We suggest that function of MPSA is likely detection of female pheromones, which would evidence chemical communication between sexes. We hypothesize that evolution of MPSA could be related to burrowing habits of Dyschiriini and its possible sexual behavior in soil tunnels. Study of MPSA may help to elucidate phylogenetic relationships among members of the tribe.

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.