The genetic determination of alternate stages in polyphenic insects.

Molt-based transitions in form are a central feature of insect life that have enabled adaptation to diverse and changing environments. The endocrine regulation of these transitions is well established, but an understanding of their genetic regulation has only recently emerged from insect models. The pupal and adult stages of metamorphosing insects are determined by the stage specifying transcription factors broad-complex (br) and Ecdysone inducible protein 93 (E93), respectively. A probable larval determinant, chronologically inappropriate metamorphosis (chinmo), has just recently been characterized. Expression of these three transcription factors in the metamorphosing insects is regulated by juvenile hormone with ecdysteroid hormones, and by mutual repression between the stage-specific transcription factors. This review explores the hypothesis that variations in the onset, duration, and tissue-specific expression of chinmo, br, and E93 underlie other polyphenisms that have arisen throughout insects, including the castes of social insects, aquatic stages of mayflies, and the neoteny of endoparasites. The mechanisms that constrain how chinmo, br, and E93 expression may vary will also constrain the ways that insect life history may evolve. I find that four types of expression changes are associated with novel insect forms: (1) heterochronic shift in the turnover of expression, (2) expansion or contraction of expression, (3) tissue-specific expression, and (4) redeployment of stage-specific expression. While there is more to be learned about chinmo, br, and E93 function in diverse insect taxa, the studies outlined here show that insect stages are modular units in developmental time and a substrate for evolutionary forces to act upon.

Metabolic mechanisms of species-specific developmental tempo.

Development consists of a highly ordered suite of steps and transitions, like choreography. Although these sequences are often evolutionarily conserved, they can display species variations in duration and speed, thereby modifying final organ size or function. Despite their evolutionary significance, the mechanisms underlying species-specific scaling of developmental tempo have remained unclear. Here, we will review recent findings that implicate global cellular mechanisms, particularly intermediary and protein metabolism, as species-specific modifiers of developmental tempo. In various systems, from somitic cell oscillations to neuronal development, metabolic pathways display species differences. These have been linked to mitochondrial metabolism, which can influence the species-specific speed of developmental transitions. Thus, intermediary metabolic pathways regulate developmental tempo together with other global processes, including proteostasis and chromatin remodeling. By linking metabolism and the evolution of developmental trajectories, these findings provide opportunities to decipher how species-specific cellular timing can influence organism fitness.

Brain developmental and cortical connectivity changes in transgenic monkeys carrying the human-specific duplicated gene SRGAP2C.

Human-specific duplicated genes contributed to phenotypic innovations during the origin of our own species, such as an enlarged brain and highly developed cognitive abilities. While prior studies on transgenic mice carrying the human-specific SRGAP2C gene have shown enhanced brain connectivity, the relevance to humans remains unclear due to the significant evolutionary gap between humans and rodents. In this study, to investigate the phenotypic outcome and underlying genetic mechanism of SRGAP2C, we generated transgenic cynomolgus macaques (Macaca fascicularis) carrying the human-specific SRGAP2C gene. Longitudinal MRI imaging revealed delayed brain development with region-specific volume changes, accompanied by altered myelination levels in the temporal and occipital regions. On a cellular level, the transgenic monkeys exhibited increased deep-layer neurons during fetal neurogenesis and delayed synaptic maturation in adolescence. Moreover, transcriptome analysis detected neotenic expression in molecular pathways related to neuron ensheathment, synaptic connections, extracellular matrix and energy metabolism. Cognitively, the transgenic monkeys demonstrated improved motor planning and execution skills. Together, our findings provide new insights into the mechanisms by which the newly evolved gene shapes the unique development and circuitry of the human brain.

A new species of Rhagophthalmus Motschulsky, 1854 (Coleoptera, Rhagophthalmidae) from Laos represents the smallest known member of the genus.

Rhagophthalmus Motschulsky, 1854 is the most speciose genus in Rhagophthalmidae, distributed in the region encompassing South, East, and Southeast Asia. Here, we describe R.nanussp. nov. from the Houaphanh Province of eastern Laos, which represents the smallest known species in Rhagophthalmus and one of the smallest in Rhagophthalmidae. We compare it with the morphologically similar and geographically close congeners and provide a preliminary identification key to adult males of Rhagophthalmus species from mainland Southeast Asia. Additionally, we discuss the morphology and variability of male genitalia within Rhagophthalmus.

Transcriptomic heterochrony and completely cleistogamous flower development in the mycoheterotrophic orchid Gastrodia.

Cleistogamy, in which plants can reproduce via self-fertilization within permanently closed flowers, has evolved in > 30 angiosperm lineages; however, consistent with Darwin's doubts about its existence, complete cleistogamy - the production of only cleistogamous flowers - has rarely been recognized. Thus far, the achlorophyllous orchid genus, Gastrodia, is the only known genus with several plausible completely cleistogamous species. Here, we analyzed the floral developmental transcriptomes of two recently evolved, completely cleistogamous Gastrodia species and their chasmogamous sister species to elucidate the possible changes involved in producing common cleistogamous traits. The ABBA-BABA test did not support introgression and protein sequence convergence as evolutionary mechanisms leading to cleistogamy, leaving convergence in gene expression as a plausible mechanism. Regarding transcriptomic differentiation, the two cleistogamous species had common modifications in the expression of developmental regulators, exhibiting a gene family-wide signature of convergent expression changes in MADS-box genes. Our transcriptomic pseudotime analysis revealed a prolonged juvenile state and eventual maturation, a heterochronic pattern consistent with partial neoteny, in cleistogamous flower development. These findings indicate that transcriptomic partial neoteny, arising from changes in the expression of conserved developmental regulators, might have contributed to the rapid and repeated evolution of cleistogamous flowers in Gastrodia.

Would the Cephalic Development in the Purebred Arabian Horse and Its Crosses Indicate a Paedomorphic Process?

This study examined paedomorphosis in PAH and F1 crossbreds. A sample of 99 horses was selected from 40 different breeders and consisted of three groups: stallions (n = 16), mares (n = 53), and geldings (n = 30), ranging from 10 months to 27 years in age. All horses presented a concave celloid lateral left head profile in the acquired photographic images. The hypothesis proposed in this study suggested the lateral profile of the head in juveniles was representational in the adult form due to the neonate's facial bones (part of the splanchnocranium) developing at a different rate to those of the skull. The methodology utilized geometric morphometrics to identify 23 landmarks so as to identify profile curvature indicative between the three groups (stallions, mares, and geldings). Principal component analysis reduced the number of variables to 14 examinable landmarks. Using a two-NPMANOVA and multivariate regression test, it was demonstrated that an isometric relationship between the concave celloid profile in the juvenile and its adult counterpart existed. This result supported the hypothesis that PAH and F1 crossbreds expressed a paedomorphic trait due to the adult form retaining the concave celloid profile identified in the juvenile.

Human-specific regulation of neural maturation identified by cross-primate transcriptomics.

Unique aspects of human behavior are often attributed to differences in the relative size and organization of the human brain: these structural aspects originate during early development. Recent studies indicate that human neurodevelopment is considerably slower than that in other nonhuman primates, a finding that is termed neoteny. One aspect of neoteny is the slow onset of action potentials. However, which molecular mechanisms play a role in this process remain unclear. To examine the evolutionary constraints on the rate of neuronal maturation, we have generated transcriptional data tracking five time points, from the neural progenitor state to 8-week-old neurons, in primates spanning the catarrhine lineage, including Macaca mulatta, Gorilla gorilla, Pan paniscus, Pan troglodytes, and Homo sapiens. Despite finding an overall similarity of many transcriptional signatures, species-specific and clade-specific distinctions were observed. Among the genes that exhibited human-specific regulation, we identified a key pioneer transcription factor, GATA3, that was uniquely upregulated in humans during the neuronal maturation process. We further examined the regulatory nature of GATA3 in human cells and observed that downregulation quickened the speed of developing spontaneous action potentials, thereby modulating the human neotenic phenotype. These results provide evidence for the divergence of gene regulation as a key molecular mechanism underlying human neoteny.

Diversity of the Paedomorphic Snail-Eating Click-Beetle Genus Malacogaster Bassi, 1834 (Elateridae: Agrypninae: Drilini) in the Mediterranean.

The soft-bodied click-beetle genus Malacogaster Bassi, 1834 from the Mediterranean region has never been taxonomically revised to date. Information on its morphology, intra- and interspecific variability, systematics and distribution is fragmented and most species have not been properly studied since their description. Therefore, in this study we summarize all available information on the genus Malacogaster. Altogether, we recognize 10 valid species from the area including the Canary Islands, Iberian Peninsula, Balearic Islands, northern coast of Africa, Sardinia, and Sicily. Malacogaster ruficollis Dodero, 1925, stat. nov., which was originally described as a variety of M. bassii Lucas, 1870 and later synonymized with it, is considered a separate species. Malacogaster parallelocollis Reitter, 1894, syn. nov. and M. olcesei var. reductus Pic, 1951, syn. nov. are synonymized with M. maculiventris Reitter, 1894. Malacogaster notativentris Pic, 1951, syn. nov. and M. olcesei Pic, 1951, syn. nov. are synonymized with M. passerinii Bassi, 1834. Lectotypes are designated for M. maculiventris Reitter, 1894, M. nigripes heydeni Reitter, 1894, M. parallelocollis Reitter, 1894, M. thoracica Redtenbacher, 1858, M. olcesei Pic, 1951, and M. rubripes Peyerimhoff, 1949 to fix their identity.

Evolutionary cardiology and experimental research / Cardiología evolutiva e investigación experimental

Abstract Evolutionary medicine studies the role of evolution in health problems. Diseases are considered as phenotypes generated by the expression of sets of genes and a complex interplay with the environment. The main mechanisms involved in evolutionary medicine are antagonistic pleiotropy, ecological antagonistic pleiotropy, atavisms and heterochrony. Antagonistic pleiotropism refers to genes that are beneficial during certain stages of development but become detrimental in others. Ecological antagonistic pleiotropy refers to the misadaptation to current lifestyle conditions which are different from those in which humans evolved. These mechanisms participate in the development of congestive heart failure, hypertension and atherosclerosis. Atavistic conditions or genes are expressed in our ancestors but have remained silent during evolution being suddenly expressed without an apparent cause during the appearance of a disease is another mechanism in evolutionary cardiology. The change in the heart metabolism from fatty acid to glucose dependent can be considered as an atavistic condition that appears in the heart after a stroke and may underlie impaired cardiomyocyte regeneration. Heterochrony is the expression of genes that cause the appearance of traits at a different timing during development and is therefore related to atavisms. Evolutionary medicine explains the interactions of pathogens and the host in infectious diseases where the cardiac tissue becomes a target. Mechanisms involved in evolutionary medicine participate in the generation of diseases and may be approached experimentally. Therefore, to better understand health problems and therapeutical approaches, an evolutionary medicine approach in experimental medicine may prove useful.

Resumen La medicina evolutiva estudia el papel de la evolución en los problemas de salud. Las enfermedades son fenotipos generados por la expresión de genes y una interacción compleja con el medio ambiente. Los principales mecanismos implicados son la pleiotropía antagonista, la pleiotropía antagonista ecológica, los atavismos y la heterocronía. El pleiotropismo antagonista se refiere a situaciones donde los genes que son beneficiosos durante ciertas etapas del desarrollo resultan perjudiciales en otras. La pleiotropía antagonista ecológica se refiere a la mala adaptación a las condiciones de vida actuales, que difieren de aquellas en las que los humanos evolucionaron. Estos mecanismos participan en el desarrollo de insuficiencia cardiaca congestiva, hipertensión y aterosclerosis. Las condiciones o genes atávicos fueron características que se expresaron en nuestros antepasados pero han permanecido silenciadas durante la evolución, expresándose repentinamente durante una enfermedad; un ejemplo es el cambio metabólico en el corazón de dependiente de ácidos grasos a dependiente de glucosa en condiciones de hipoxia que aparece después de un infarto y puede subyacer a la dificultad de la regeneración de los cardiomiocitos. La heterocronía es la expresión de genes que provocan la aparición de rasgos en un momento diferente durante el desarrollo. La medicina evolutiva también explica las interacciones entre los patógenos y el huésped en enfermedades infecciosas. Los mecanismos implicados en la medicina evolutiva participan en la generación de enfermedades y pueden abordarse experimentalmente. Por tanto, la medicina experimental puede enriquecer la medicina evolutiva y el origen de muchos problemas de salud.

Their young bite better: On- and off-host selection pressure as drivers for evolutionary-developmental modification in Rhipicephalus ticks.

Distinct life stages may experience different selection pressures influencing phenotypic evolution. Morphological evolution is also constrained by early phenotypes, since early development forms the phenotypic basis of later development. This work investigates evolutionary-developmental modification in three life stages and both sexes of 24 Rhipicephalus species using phylogenetic comparative methods for geometric morphometrics of basis capituli (basal mouthpart structure used for host attachment), and scutum or conscutum areas (proxy for overall body size). Findings indicate species using large hosts at early life stages have distinct basis capituli shapes, correlated with host size, enabling attachment to the tough skins of large hosts. Host-truncate species (one- and two-host) generally retain these adaptive features into later life stages, suggesting neoteny is linked to the evolution of host truncation. In contrast, species using small hosts at early life stages have lost these features. Developmental trajectories differ significantly between host-use strategies (niches), and correlate with distinct clades. In two-host and three-host species using large hosts at early life stages, developmental change is heterotopically accelerated (greater cell mass development) before the first off-host period where selection probably favours large individuals able to better resist dehydration when questing (waiting) for less abundant, less active hosts. In other species, development is heterotopically reduced (neotenic), possibly because dehydration risk is bypassed by prolonged host attachment (one-host species - heterotopic neoteny), or is allometrically repatterned possibly by using highly abundant and active hosts (three-host species using small hosts at early life stages - allometric repatterning). These findings highlight complex trade-offs between on- and off-host factors of free-living ectoparasite ecology, which mediate responses to diverse selection pressures varied by life stage and host-use strategy. It is proposed that these trade-offs shape evolutionary-developmental morphology and diversity of Rhipicephalus ticks.

Characterization of naked mole-rat hematopoiesis reveals unique stem and progenitor cell patterns and neotenic traits.

Naked mole rats (NMRs) are the longest-lived rodents yet their stem cell characteristics remain enigmatic. Here, we comprehensively mapped the NMR hematopoietic landscape and identified unique features likely contributing to longevity. Adult NMRs form red blood cells in spleen and marrow, which comprise a myeloid bias toward granulopoiesis together with decreased B-lymphopoiesis. Remarkably, youthful blood and marrow single-cell transcriptomes and cell compositions are largely maintained until at least middle age. Similar to primates, the primitive stem and progenitor cell (HSPC) compartment is marked by CD34 and THY1. Stem cell polarity is seen for Tubulin but not CDC42, and is not lost until 12 years of age. HSPC respiration rates are as low as in purified human stem cells, in concert with a strong expression signature for fatty acid metabolism. The pool of quiescent stem cells is higher than in mice, and the cell cycle of hematopoietic cells is prolonged. By characterizing the NMR hematopoietic landscape, we identified resilience phenotypes such as an increased quiescent HSPC compartment, absence of age-related decline, and neotenic traits likely geared toward longevity.

[Evolutionary cardiology and experimental research]. / Cardiología evolutiva e investigación experimental.

Evolutionary medicine studies the role of evolution in health problems. Diseases are considered as phenotypes generated by the expression of sets of genes and a complex interplay with the environment. The main mechanisms involved in evolutionary medicine are antagonistic pleiotropy, ecological antagonistic pleiotropy, atavisms and heterochrony. Antagonistic pleiotropism refers to genes that are beneficial during certain stages of development but become detrimental in others. Ecological antagonistic pleiotropy refers to the misadaptation to current lifestyle conditions which are different from those in which humans evolved. These mechanisms participate in the development of congestive heart failure, hypertension and atherosclerosis. Atavistic conditions or genes are expressed in our ancestors but have remained silent during evolution being suddenly expressed without an apparent cause during the appearance of a disease is another mechanism in evolutionary cardiology. The change in the heart metabolism from fatty acid to glucose dependent can be considered as an atavistic condition that appears in the heart after a stroke and may underlie impaired cardiomyocyte regeneration. Heterochrony is the expression of genes that cause the appearance of traits at a different timing during development and is therefore related to atavisms. Evolutionary medicine explains the interactions of pathogens and the host in infectious diseases where the cardiac tissue becomes a target. Mechanisms involved in evolutionary medicine participate in the generation of diseases and may be approached experimentally. Therefore, to better understand health problems and therapeutical approaches, an evolutionary medicine approach in experimental medicine may prove useful.

La medicina evolutiva estudia el papel de la evolución en los problemas de salud. Las enfermedades son fenotipos generados por la expresión de genes y una interacción compleja con el medio ambiente. Los principales mecanismos implicados son la pleiotropía antagonista, la pleiotropía antagonista ecológica, los atavismos y la heterocronía. El pleiotropismo antagonista se refiere a situaciones donde los genes que son beneficiosos durante ciertas etapas del desarrollo resultan perjudiciales en otras. La pleiotropía antagonista ecológica se refiere a la mala adaptación a las condiciones de vida actuales, que difieren de aquellas en las que los humanos evolucionaron. Estos mecanismos participan en el desarrollo de insuficiencia cardiaca congestiva, hipertensión y aterosclerosis. Las condiciones o genes atávicos fueron características que se expresaron en nuestros antepasados pero han permanecido silenciadas durante la evolución, expresándose repentinamente durante una enfermedad; un ejemplo es el cambio metabólico en el corazón de dependiente de ácidos grasos a dependiente de glucosa en condiciones de hipoxia que aparece después de un infarto y puede subyacer a la dificultad de la regeneración de los cardiomiocitos. La heterocronía es la expresión de genes que provocan la aparición de rasgos en un momento diferente durante el desarrollo. La medicina evolutiva también explica las interacciones entre los patógenos y el huésped en enfermedades infecciosas. Los mecanismos implicados en la medicina evolutiva participan en la generación de enfermedades y pueden abordarse experimentalmente. Por tanto, la medicina experimental puede enriquecer la medicina evolutiva y el origen de muchos problemas de salud.

Rhagophthalmidae Olivier, 1907 (Coleoptera, Elateroidea): described genera and species, current problems, and prospects for the bioluminescent and paedomorphic beetle lineage.

Rhagophthalmidae are a small beetle family known from the eastern Palaearctic and Oriental realms. Rhagophthalmidae are closely related to railroad worms (Phengodidae) and fireflies (Lampyridae) with which they share highly modified paedomorphic females and the ability to emit light. Currently, Rhagophthalmidae include 66 species classified in the following 12 genera: Bicladodrilus Pic, 1921 (two spp.), Bicladum Pic, 1921 (two spp.), Dioptoma Pascoe, 1860 (two spp.), Diplocladon Gorham, 1883 (two spp.), Dodecatoma Westwood, 1849 (eight spp.), Falsophrixothrix Pic, 1937 (six spp.), Haplocladon Gorham, 1883 (two spp.), Menghuoius Kawashima, 2000 (three spp.), Mimoochotyra Pic, 1937 (one sp.), Monodrilus Pic, 1921 (two spp. in two subgenera), Pseudothilmanus Pic, 1918 (two spp.), and Rhagophthalmus Motschulsky, 1854 (34 spp.). The replacement name Haplocladongorhami Kundrata, nom. nov. is proposed for Diplocladonhasseltii Gorham, 1883b (described in subgenus Haplocladon) which is preoccupied by Diplocladonhasseltii Gorham, 1883a. The genus Reductodrilus Pic, 1943 is tentatively placed in Lampyridae: Ototretinae. Lectotypes are designated for Pseudothilmanusalatus Pic, 1918 and P.marginalis Pic, 1918. Interestingly, in the eastern part of their distribution, Rhagophthalmidae have remained within the boundaries of the Sunda Shelf and the Philippines demarcated by the Wallace Line, which separates the Oriental and Australasian realms. This study is intended to be a first step towards a comprehensive revision of the group on both genus and species levels. Additionally, critical problems and prospects for rhagophthalmid research are briefly discussed.

Paedomorphosis in the Ezo salamander (Hynobius retardatus) rediscovered after almost 90 years.

Although paedomorphosis is widespread across salamander families, only two species have ever been documented to exhibit paedomorphosis in Hynobiidae. One of these two exceptional species is Hynobius retardatus in which paedomorphosis was first reported in 1924, in specimens from Lake Kuttara in Hokkaido. This population became extinct after the last observation in 1932; since then, no paedomorphs of this species have been reported anywhere. Here, we report the rediscovery of paedomorphs of this species. Three paedomorph-like male salamanders were collected from a pond in the south Hokkaido in December 2020 and April 2021; in size, these specimens were similar to metamorphosed adults but they still displayed larval features such as external gills and a well-developed caudal fin. An artificial fertilization experiment demonstrated that they were sexually compatible with metamorphosed females, thus, confirming them to be paedomorphs. Future efforts to find additional paedomorphs in this and other populations are required to assess the prevalence of paedomorphosis in H. retardatus and to improve understanding of the ecology and evolution of paedomorphisis in Urodela.

Progressive reorganization of mitochondrial apparatus in aging skeletal muscle of naked mole rats (Heterocephalus glaber) as revealed by electron microscopy: potential role in continual maintenance of muscle activity.

The authors examined the ultrastructure of mitochondrial apparatus of skeletal muscles of naked mole rats (Heterocephalus glaber) from the age of 6 months to 11 years. The obtained results have demonstrated that the mitochondria in skeletal muscles of naked mole rats aged below 5 years is not well-developed and represented by few separate small mitochondria. Mitochondrial reticulum is absent. Starting from the age of 5 years, a powerful mitochondrial structure is developed. By the age of 11 years, it become obvious that the mitochondrial apparatus formed differs from that in the skeletal muscle of adult rats and mice, but resembles that of cardiomyocytes of rats or naked mole rats cardiomyocytes. From the age of 6 months to 11 years, percentage area of mitochondria in the skeletal muscle of naked mole rat is increasing by five times. The growth of mitochondria is mainly driven by increased number of organelles. Such significant growth of mitochondria is not associated with any abnormal changes in mitochondrial ultrastructure. We suppose that specific structure of mitochondrial apparatus developed in the skeletal muscle of naked mole rats by the age of 11 years is necessary for continual skeletal muscle activity of these small mammals burrowing very long holes in stony earth, resembling continual activity of heart muscle. In any case, ontogenesis of naked mole rat skeletal muscles is much slower than of rats and mice (one more example of neoteny).

When size matters: the first comprehensive anatomical study of a species of "Condylocardiidae", an extremely miniaturized bivalve.

'Miniaturization' is a widespread phenomenon among the Metazoa. In the molluscan class Bivalvia, records of miniaturization are numerous. Among the Archiheterodonta, Warrana besnardi (Klappenbach, 1963) has attracted attention for its tiny size, which does not exceed 1.5 mm in shell length, and because it belongs to a group with limited anatomical information and often-debated status, the "Condylocardiidae" (which recent molecular studies place deeply nested within the family Carditidae). All species of Warrana Laseron, 1953 are small-bodied, and so miniaturization presumably occurred from a large-bodied ancestor within the Carditidae sensu lato. South American W. besnardi is here studied in detail. Its small size and the enlargement of the anterodorsal region during growth, reflects (and likely led) to infaunal habit, living as a burrowing bivalve that passively feeds on deposit particles entering the pallial cavity anteriorly. Mantle glands, previously reported as a common feature of other archiheterodonts, are missing in W. besnardi, but spongiform tissue in the antero-ventral portion of the mantle lobes presumably represents a blood sinus that might compensate for the great reduction of the ctenidia. Lecithotrophy is reported, with yolky oocytes bearing a thick non-cellular capsule layer; brooding was not observed, and it is here hypothesized that the extreme miniaturization, with the great reduction of ctenidia, is responsible for a shift in the reproductive mode of condylocardiids, contrasting with the commonly reported ovoviviparity of the carditids.

Cellular and Molecular Mechanisms Linking Human Cortical Development and Evolution.

The cerebral cortex is at the core of brain functions that are thought to be particularly developed in the human species. Human cortex specificities stem from divergent features of corticogenesis, leading to increased cortical size and complexity. Underlying cellular mechanisms include prolonged patterns of neuronal generation and maturation, as well as the amplification of specific types of stem/progenitor cells. While the gene regulatory networks of corticogenesis appear to be largely conserved among all mammals including humans, they have evolved in primates, particularly in the human species, through the emergence of rapidly divergent transcriptional regulatory elements, as well as recently duplicated novel genes. These human-specific molecular features together control key cellular milestones of human corticogenesis and are often affected in neurodevelopmental disorders, thus linking human neural development, evolution, and diseases.

Egg retention, viviparity and ovoviviparity in Paraneoptera.

This article is a second part of the themed issue "Aberrant cytogenetic and reproductive patterns in the evolution of Paraneoptera insects", prepared by the Russian-Bulgarian research team. Here, analysis of aberrations related to the egg development is provided based on literature data and the author's own investigations. Evolutionary aspects of ovoviviparity/viviparity are also briefly discussed. Material and methods, terminology and nomenclature of taxonomic names are listed in the first paper of the issue (Gavrilov-Zimin et al. 2021).

The genus Selasia Laporte, 1838 (Coleoptera: Elateridae: Agrypninae) in Sri Lanka.

The neotenic click-beetle genus Selasia Laporte, 1838 has a centre of diversity in the tropical Africa but also includes several species known from the Palearctic and Oriental regions. In this study, we review the Selasia fauna of Sri Lanka. We describe S. ivanae sp. nov., redescribe S. apicalis Pic, 1914, and discuss the systematic placement of S. isabellae Bourgeois, 1909, which is probably a firefly and is considered incertae sedis.

Do holometabolous insects molt spontaneously after adulthood? An exceptional case report in fireflies (Coleoptera: Lampyridae), with discussion of its inferred endocrine regulation especially in relation to neoteny.

It has been a traditionally held view that winged insects stop molting after they reach adulthood. We observed a fascinating phenomenon of a post-imago molt occurring in the neotenic females of a firefly species in Taiwan over the last two years. By rearing Lamprigera minor larvae to adults, four out of the five unmated females studied were found undergoing an extra molt 8-18 days after adult eclosion. They were reproductively mature when the post-imago molt occurred, as evidenced by the eggs inside their bodies. The four females died without oviposition whereas the only normal female laid eggs. A comparison of exuviae of different stages confirmed the existence of post-imago ecdysis. The adult skin differed from the pupal one mainly in the mouthparts and leg structures. No mix of pupal and adult traits was seen in the adult skin. The females retained the same morphology after the extra molt. A close examination of the post-imago molting females revealed that their oviduct openings were all blocked by larval or pupal skin and thus unable to lay eggs. The reproductive stress may invoke an endocrine disorder and lead to an extra molt. We propose that L. minor females retain their prothoracic glands even as adults, allowing them to molt as adults under certain environmental or physiological conditions. Thus, neoteny of L. minor is reflected in both the external morphology as well as the internal physiology. The possible developmental changes associated with the evolution of neoteny are discussed.