Chironomus bonus was first described from Paravani Lake in 1974 and has also been recorded in Armenia, France, Switzerland and Bulgaria, but neither GenBank nor BOLD previously contained DNA data for it. The study combined chromosome analysis with mitochondrial DNA sequencing to characterize the Georgian specimens and compare them with other members of the Chironomus plumosus species group. Six larvae were examined cytologically.

Following fixation in an ethanol–glacial acetic acid solution, chromosome preparations were made with the ethanol-orcein technique and examined under a Carl Zeiss Axio Imager A2 microscope. Banding sequences were identified for the chromosome arms using established photomaps and mapping systems, allowing the authors to compare the Georgian material directly with previously published karyotypes. All six larvae had the same basic chromosome complement: a diploid number of 2n = 8 plus an additional B chromosome.

Their chromosomes formed the AB, CD, EF and G arm combinations associated with the thummi cytocomplex. The AB and CD chromosomes were metacentric, EF was submetacentric and G was telocentric. Clearly identifiable centromeric bands, one nucleolus, two Balbiani rings on the G arm and a further Balbiani ring on arm B completed the observed profile.

The principal visible distinction from C. plumosus was the additional B chromosome, which was present in almost all examined C. bonus larvae. Chromosome examination was central to species identification because the C. plumosus group comprises 14 recognized species that can be similar in external form and often occur together in the same body of water. The source review describes karyological analysis as the reliable method for identifying species in the group.

DNA extraction was therefore performed on four larvae that had already been studied by chromosome analysis. Their DNA was extracted with a commercial preparation kit, and the mitochondrial COI barcoding region was amplified with primers 911 and 912 in 25-µL PCR reactions. Products were purified, checked by agarose gel electrophoresis and sequenced in both directions by Sanger sequencing.

The analysis produced three complete COI sequences, deposited in GenBank as MZ014021, MZ014022 and MZ014023. Their lengths were 627, 658 and 650 base pairs, respectively, and each represented a different haplotype within the small Georgian sample. For phylogenetic comparison, the researchers assembled 61 nucleotide sequences from GenBank and BOLD, including members of the C. plumosus group and the outgroup Pagastiella orophila.

The alignment was performed in MUSCLE with the invertebrate mitochondrial genetic code in MEGA 6, leaving 579 positions after gaps and missing data were removed. Pairwise evolutionary distances were calculated with the Kimura two-parameter model. Bayesian phylogenetic analysis was performed in BEAST v1.10.4 using the HKY+G substitution model, a strict clock and a Yule speciation process.

The resulting tree contained several strongly supported but internally complex groupings. The three Georgian C. bonus sequences joined C. plumosus sequences from western and eastern Europe in a Palearctic cluster that also included available C. usenicus sequences from Russia. A separate Far Eastern cluster combined C. borokensis with sequences labelled C. plumosus from China, South Korea and Japan, while a Nearctic cluster mixed C. entis and C. plumosus sequences from North America.

The C. bonus sequences did not form an independent branch in this analysis. Genetic distances were low within the three principal clusters but higher between them. Within the Palearctic cluster, Kimura two-parameter distances ranged from 0 to 2.595%; excluding an Iranian C. plumosus sequence reduced the range to 0–0.914%.

The three C. bonus sequences differed from one another by 0.182–0.364%. The Nearctic C. entis–C. plumosus cluster contained distances of 0–1.285%, while the Far Eastern C. borokensis–C. suwai cluster ranged from 0 to 2.217%, or 0–1.839% when one Chinese sequence was omitted. Average distances between clusters were 3.55% for the Palearctic and Far Eastern groups, 3.75% for the Palearctic and Nearctic groups, and 5.98% for the Far Eastern and Nearctic groups.

The authors compared those between-cluster values with a reference range of up to 3% for intraspecific COI distances in Chironomus, while emphasizing that COI results require support from other forms of identification. The 3.75% separation between Palearctic and Nearctic sequences labelled C. plumosus was interpreted as evidence that Nearctic populations assigned to that name may represent a separate species. The authors also cautioned that several database sequences had been identified through morphology alone.

Their clustering patterns suggested that some records labelled C. plumosus in the Far East or North America could belong to C. borokensis, C. suwai or C. entis, although the study did not directly resolve the identity of every database specimen. Comparison of the Georgian sequences with other members of the Palearctic cluster identified six substitutions that could distinguish C. bonus. Four were nonsynonymous and two were synonymous.

A substitution at position 340 was present in all three Georgian C. bonus sequences and was the only change found in the complete dataset that clearly distinguished them from the other species. The remaining substitutions occurred in individual sequences and were also detected elsewhere in the comparison. The authors discussed two possible explanations for the close mitochondrial similarity between C. bonus and European C. plumosus: a relatively recent separation with few accumulated differences, or an older separation followed by mitochondrial introgression through interspecific hybridisation.

The available results did not distinguish between those scenarios. Model-based divergence estimates depended on assumed mitochondrial substitution rates of 1.5%, 2.3% or 3.54% per million years. The earliest split among the three principal clusters was estimated at 5.75–3.43 million years ago, placing it in the Late Miocene to early Pliocene.

The common ancestor of all three clusters was dated to 2.88–1.72 million years ago. Estimated separation between the Palearctic and Far Eastern clusters was 1.97–1.17 million years, while common ancestors for the Palearctic, Far Eastern and Nearctic clusters were estimated at 1.288–0.759, 0.906–0.539 and 0.638–0.378 million years ago, respectively. The Palearctic cluster excluding the Iranian sequence had an estimated common ancestor at 0.517–0.307 million years ago.

The authors proposed that diversification within the group may have been associated with the expansion of separate stagnant-water habitats during the Pliocene. Lakes and ponds can differ in depth, temperature, mineralisation, size and other environmental conditions, potentially producing differences in breeding timing and reproductive isolation. This remains an evolutionary interpretation rather than a directly observed outcome of the Georgian study.

The timing estimates were also provisional because COI data were available for only seven of the 14 species in the C. plumosus group, and the analysis contained ambiguous clusters. Nevertheless, the study established a distinctive karyological identity for the Georgian C. bonus population and supplied the species’ first COI reference sequences, showing why chromosome and mitochondrial evidence were evaluated together rather than treating either marker as sufficient on its own.

Subject of Research: Karyotypic and mitochondrial COI characterization of Chironomus bonus from the South Caucasus

Article Title: Karyotype characteristics and gene COI sequences of Chironomus bonus Shilova et Dzhvarsheishvili, 1974 ( Diptera , Chironomidae ) from the South Caucasus (Republic of Georgia, Paravani river)

Article References: Karmokov, M. K. (2022). Karyotype characteristics and gene COI sequences of Chironomus bonus Shilova et Dzhvarsheishvili, 1974 (Diptera, Chironomidae) from the South Caucasus (Republic of Georgia, Paravani river). Comparative Cytogenetics, 16(1), 19-38. https://doi.org/10.3897/compcytogen.v16.i1.79182

Image Credits: AI Generated

DOI: 10.3897/CompCytogen.v16.i1.79182

Keywords: Chironomus bonus, Chironomus plumosus group, mitochondrial COI, karyotype, B chromosome, South Caucasus, Phylogeography, Mitochondrial divergence, Non-biting midges, characteristics, gene, sequences

The genus Ametropus belongs to the mayfly family Ametropodidae, a small and peculiar lineage within the order Ephemeroptera. Its representatives are among the most specialized mayflies in the world, adapted for life buried in the sandy bottoms of large rivers. The nymphs are psammophilous, meaning they dwell within sand, and behave as semi-burrowers, hiding in muddy or sandy substrata where their slender, densely hairy bodies blend seamlessly with the sediment. This lifestyle makes them notoriously difficult to collect, and their populations are typically rare and scattered even within their broad distribution range. As a result, records of Ametropus are few and far between, and each new occurrence carries significant biogeographic weight.

The history of Ametropus in the Caucasus is a story of uncertainty. The Reverend Alfred Edwin Eaton reported in 1885 that specimens of the genus were found mixed within samples collected from Russia, the Caucasus, and Armenia, though the exact localities remained unclear. Decades later, in 1940 and again in the 1940s, Sadovsky collected Ametropus sp. from the Kura River in Georgia, and the specimens were considered potentially new taxa due to taxonomic ambiguity. However, no voucher material or precise locality data survived, and the 2018 comprehensive checklist of Georgian mayflies, compiled by Gabelashvili and colleagues, flagged these old records as doubtful. Since then, no other record of Ametropus had been documented from the whole South Caucasus or nearby regions further south, leaving the very presence of the genus in the region an open question.

That question has now been answered. On April 17, 2022, researchers collected benthic samples from three plots spaced 30 meters apart in the shallow, murky waters of the Alazani River at an altitude of 95 meters above sea level. The Alazani, a 240-kilometer-long river originating from the southern slopes of the Greater Caucasus Mountains at roughly 2800 meters, is the largest left tributary of the Kura River, and its lower reach forms the border between Georgia and Azerbaijan. At the sampling sites, the river flows slowly, typical of a lowland watercourse, with a bottom composed mainly of clay and muddy silt, interspersed with sand. The water is poorly transparent, algae are negligible, and the benthic communities are homogeneous but dense, a habitat that perfectly matches the known preferences of Ametropus larvae.

Samples were collected using a kick-net with a 500-micrometer mesh size according to the AQEM sampling method, within a depth range of 15 to 80 centimeters. Among the mayfly specimens sorted from the material, five individuals were initially identified as Ametropus sp. The nymph morphology of this genus is unique among mayflies and cannot be confused with any other, which made the initial recognition straightforward. Based on morphological characteristics described by Sadovsky, Bauernfeind and Soldan, and Eiseler, the specimens were further identified as Ametropus fragilis. The general coloration is yellowish-brown, with slender and hairy bodies. The antennae bear 16 segments, and the legs are slender with long claws, the forelegs much shorter than the middle and hind pairs. Fore coxae carry a base-attached spinous patch, and seven pairs of single, lateral plate-like gills, each fringed with long setae on their margins, line the body. Maxillary and labial palps are three-segmented and covered with dense hair. In all respects, the newly collected individuals are morphologically highly similar to the Ametropus specimens described and illustrated by Sadovsky 80 years ago, confirming a striking continuity across the decades.

To resolve the species identity with certainty, the team turned to DNA barcoding of the cytochrome oxidase subunit I gene, the standard molecular tool for species identification in animals. Genomic DNA was extracted from tissue samples using a commercial kit, and partial COI sequences were amplified by polymerase chain reaction using the primer pairs LCO1490-JJ and HCO2198-JJ. Thermal cycling involved an initial denaturation step followed by two cycle sets with progressively lowered annealing temperatures, a touch-down protocol designed to maximize amplification specificity. Amplicons were visualized on agarose gels, and sequencing of unpurified PCR products in both directions was conducted at the Beijing Genomics Institute in Hong Kong. Three barcode sequences of 626 nucleotide base pairs, free from indels or stop codons, were obtained from three specimens and uploaded to the Barcode of Life Data System, where they were assigned processing IDs EPGEO001-23, EPGEO002-23, and EPGEO003-23.

The molecular results were nearly identical among the Georgian specimens, with uncorrected p-distances ranging from 0.15 to 0.61 percent, confirming they belong to a single population. The BOLD System identified the sequences under a new Barcode Index Number, BOLD:AFB6526. What came next, however, was entirely unexpected. The closest genetic match to the Caucasian specimens was not the European Ametropus fragilis but the Nearctic species Ametropus neavei from Canada, with a mean p-distance of just 3.54 percent. A larger genetic distance of 4.3 percent was found between the Georgian specimens and the single available sequence of A. fragilis from Slovakia. In other words, the mayflies of the Alazani River are genetically closer to their North American counterparts than to their European relatives, a pattern that defies simple geographic logic.

Ametropus fragilis is considered the sole species of the family Ametropodidae in Europe, with a distribution spanning Germany, Italy, Croatia, Poland, Austria, Slovakia, Hungary, Bulgaria, Romania, Estonia, Latvia, and Lithuania, with additional records from Russia, China, Belarus, and France. According to Jacob, North American A. albrighti should be treated as a synonym of A. fragilis, making the species Holarctic in scope. Yet the species is usually rare throughout its range, with very small population densities spread over a very large distribution area. Taking this into consideration, large genetic variation of around 5 percent divergence in the COI barcode region between populations would be expected. The observed pattern, however, makes interpretation difficult. The most intuitive explanation is recent translocation and gene flow, perhaps mediated by human activity or natural dispersal events. On the other hand, given the population characteristics and lifestyle peculiarities of Ametropus, the pattern may also indicate the existence of multiple divergent lineages of Eurasian and North American Ametropus, pointing to the need for a thorough taxonomic revision of the genus across the Holarctic region.

The implications of the new record extend well beyond Georgia. Since the Alazani River lies right on the border between Georgia and Azerbaijan, the species can now be said to occur in Azerbaijan as well. Taking the old literature into consideration, the Ametropus specimens observed in Iran and Armenia most probably also belong to the same species. The researchers have provided an updated distribution map, building on earlier work, showing the current known range of A. fragilis across Europe and neighboring areas, with countries where the species has become extinct marked separately from those where data is unavailable. The map reveals a fragmented European presence and highlights how much remains unknown about this elusive insect.

The rediscovery of Ametropus fragilis in Georgia is a testament to the value of renewed field surveys in understudied regions. The Georgian mayfly fauna, now known to include 77 species following recent additions, continues to yield surprises, and the Caucasus Barcode of Life project, under which the specimens are stored at Ilia State University in Tbilisi, is systematically cataloging the region’s biodiversity through molecular tools. For a genus that has hovered in taxonomic limbo for nearly a century in the South Caucasus, the confirmation of its presence, backed by both morphology and molecular evidence, represents a significant step forward. Yet the genetic closeness of Caucasian and North American populations serves as a reminder that even well-known Holarctic species may conceal hidden diversity, and that the full story of Ametropus is far from complete.

Subject of Research: Rediscovery and DNA barcoding of the mayfly Ametropus fragilis in Georgia after 80 years

Article Title: The record of Ametropus fragilis Albarda, 1878 ( Ephemeroptera , Ametropodidae ) from Georgia

Article References: Memishishi, A., Bikashvili, A., Japoshvili, B., & Mumladze, L. (2023). The record of Ametropus fragilis Albarda, 1878 (Ephemeroptera, Ametropodidae) from Georgia. Caucasiana, 2, 77-82. https://doi.org/10.3897/caucasiana.2.e102622

Image Credits: AI Generated

DOI: 10.3897/caucasiana.2.e102622

Keywords: Ametropus fragilis, Ephemeroptera, mayfly, Georgia, Alazani River, DNA barcoding, COI gene, South Caucasus, Holarctic distribution, Vashlovani National Park, biodiversity, taxonomy

The South Caucasus sits at the biogeographic junction between Europe and Asia and likely acted as a dispersal corridor during the Pliocene, yet well-dated carnivoran assemblages from this region remain rare. The Jradzor locality in Armenia comprises a stratigraphically constrained Pliocene vertebrate record with several fossiliferous horizons dated by Lazarev and colleagues in 2023. The carnivoran material derives from horizons JZ–3s, JZ–13 and JZ–15, dated to 4.16–3.98 Ma, and from horizon JZ–7, dated to 3.3 ± 0.03 Ma. The horizon JZ–7 is interpreted as a lahar deposit, a high-energy volcaniclastic mudflow that transported remains over relatively short distances, whereas the remaining layers represent the medium tail of pyroclastic flow. The associated vertebrate assemblage is dominated by taxa typically linked to open habitats, suggesting predominantly open landscapes in the local catchment, including Cremohipparion longipes, Gazellospira sp., Caprini indet. and a new hyracoid. Further, two additional forms from the JZ–7 horizon, Bufotes sp. and Mimomys cf. davakosi, suggest also the presence of open habitats. Here, new occurrences of carnivorans from Jradzor across multiple fossil horizons are documented, using dental and mandibular functional morphology together with morphometrics to discuss trophic ecology and community structure of the carnivoran guild in a regional biogeographic framework.

This study is based on anatomical and taxonomic analysis of the Late Pliocene carnivoran fauna from Jradzor, Armenia. The referred material, including 15 specimens, is stored in the collections of the Institute of Geological Sciences of the National Academy of Sciences of Republic of Armenia. Extant and fossil specimens used for the comparative study are stored in collections across Switzerland, Italy, France and Spain. Measurements of the cranium and mandible follow established protocols, taken with a digital caliper to the nearest 0.1 mm. A Principal Component Analysis was performed on the mesiodistal length and buccolingual width of the p3, p4 and m1 of the mandible of Chasmaporthetes from Jradzor, in comparison with extant hyaenids and selected fossil species from Eurasia, Moldova, France, the Zanda Basin and North America. The deciduous premolar of the hyaenid from Jradzor was compared with those of Pliocrocuta perrieri and Pachycrocuta brevirostris from the Late Pliocene and Early Pleistocene. Analyses were performed in the R environment using RStudio software, with the PCA performed using the prcomp function and plots obtained using ggplot.

Among the five carnivoran taxa reported from Jradzor, the canid Nyctereutes urartuensis is represented by a fairly well-preserved cranium and a right lower jaw from horizon JZ–7, dated to 3.3 ± 0.03 Ma. The cranium is considerably elongated rostrocaudally and large, characterised by a marked postorbital constriction, a vulpine crease on the dorsal surface of the zygomatic processes of the frontals, marked parasagittal and sagittal crests, rugose parietals, squared-shaped nuchal crests, an evident subangular lobe, short canine crowns and enlarged crushing surfaces of the molars with additional or enlarged cuspulids. The attribution to the genus Nyctereutes is maintained by the presence of general features of Vulpini, such as the vulpine crease, and diagnostic features for the genus, including the short-crowned canines and the presence of a rounded and evident subangular lobe on the caudoventral side of the mandible. The development of features shown by the Jradzor sample is intermediate between primitive raccoon dogs like Nyctereutes tingi and Nyctereutes donnezani and more derived forms like Nyctereutes sinensis and Nyctereutes megamastoides. Analyses of morphological, morphometric, endocranial and ecological parameters suggest the Jradzor specimens belonged to a new species, described in a companion paper by Bartolini-Lucenti and colleagues in 2026.

The mustelid Baranogale helbingi is represented by a fragment of left maxilla with P3–P4 from horizon JZ–13 and a fragment of left mandibular body with m1 from horizon JZ–15, both dated to 4.16–3.98 Ma. The maxilla fragment displays a large and round infraorbital foramen, with the P2 alveolus being two-rooted, indicating a moderately reduced tooth. The P3 is elongated and irregularly oval-shaped, with a prominent protocone pushed slightly mesially and situated centrally. The elongated and narrow P4 has a straight crown, with a moderately high paracone bearing a thin ridge across the mesial border and a long, rectangular protocone. The large and rather narrow m1 has a low and short paraconid, a prominent and short protoconid, a strongly developed metaconid clearly separated from the protoconid, and a long, trenchant talonid narrower than the trigonid. The Jradzor mustelid differs distinctly from Psalidogale altenburgensis, from both species of the genus Vormela, and from the highly evolved Oxyvormela maisi, instead strongly resembling Baranogale helbingi in all features of the P3–P4 and m1. The plotted index of total length and trigonid breadth of the m1 of B. helbingi individuals showed three non-overlapping sets of data, with the Jradzor specimen interpreted as a female based on its smaller m1 length of 6.6 mm.

A lynx-sized felid is represented by one mandibular corpus with canine and p3–p4 from horizon JZ–3s and one isolated m1. The corpus is stout and high below the premolars, with a mesiodistally short diastema between the canine and the premolars. The m1 is sharp, buccolingually compressed and with an absent metaconid and talonid. The size and morphology of the mandibular corpus and the lower carnassial are consistent with a lynx-sized felid; however, the fragmentary nature of the material does not allow a confident species-level attribution, and the specimens are therefore referred to cf. Lynx sp. pending additional material and a dedicated taxonomic treatment.

Two large hyaenids from horizon JZ–7 represent distinct feeding strategies. The hunting hyaena Chasmaporthetes lunensis is represented by two well-preserved hemi-mandibles belonging to the same individual, displaying a single mental foramen below the p2, a slightly lingually curved tooth row, a deep masseteric fossa, asymmetric crowns of the i2 and i3 with enlarged main cuspids, premolars slightly tilted backwards with sharp protoconids, and an m1 with a vestigial metaconid and a mesiodistally reduced talonid bearing a well-developed hypoconulid. Compared to the stoutly built extant hyaenas, representatives of Chasmaporthetes display more slender dentition and more elongated, slender post-cranial bones, leading to their inclusion in the category of hunting hyaenas for their inferred cursorial abilities. The Jradzor specimens display several morphological affinities with Ch. lunensis, including a curved tooth row, unicuspid talonid, large mesial accessory cuspulid on the p4, and presence of a mesial accessory cuspulid on the p2. In the PCA on lower teeth, the first principal component explains 76.69% and the second about 9.17% of the variability, and the two Jradzor specimens fall perfectly within the variability of Ch. lunensis.

The bone-cracking hyaena Pliocrocuta perrieri is represented by a right p4 and m1 and a right dp4 from horizon JZ–7. The occlusal outlines of the p4 and m1 are buccolingually wide, with the p4 displaying a high protoconid and two accessory cuspulids, and the m1 bearing a large paraconid and protoconid of equal height and an individualised but small metaconid. Pliocrocuta perrieri is a large bone-cracking hyaena commonly recorded in Early to Middle Villafranchian European assemblages, with a known chronological range from 4.2 to 2.0 Ma, distributed across Europe, Asia and Africa. From a phylogenetic viewpoint, Pliocrocuta is considered a sister-taxon of Pachycrocuta brevirostris. The morphology of the dp4 from Jradzor has a fairly extended talonid with evident cuspulids, contrasting with the more generalised deciduous dentition of P. brevirostris. Considering the morphological features of the adult dentition consistent with Pliocrocuta, the deciduous specimen is suggested to belong to the same species, a suggestion metrically confirmed when compared to other deciduous premolars of Pliocrocuta and Pachycrocuta.

Both large hyaenid species recorded from Jradzor are known to have had different dietary habits. Although both were hypercarnivores, Chasmaporthetes lunensis was a hunting hyaena, while Pliocrocuta perrieri was a bone-cracking species. Antón and colleagues in 2006 suggested that the limited fossil record of Ch. lunensis could reflect its solitary lifestyle, while P. perrieri is more abundant and its fossil record could reflect a group-denning scavenger. However, based on the shape of the muzzle and the elongated, gracile fore-limbs, Chasmaporthetes would still have required group action to kill medium and large prey. Considering the similarities in body size and the presence of adaptations for bone cracking in both species, competition between the two for carcasses was likely mitigated by ecological complementarity: if P. perrieri, due to its greater bone-cracking abilities, did not hunt actively and was well adapted to scavenging carcasses, Ch. lunensis would have used its crushing premolars on its own prey rather than consuming scavenged carrion, explaining why P. perrieri has been more abundantly found at sites where hyaenid presence correlates with their capacity to scavenge carcasses of other animals.

The carnivoran assemblage from Jradzor represents a significant contribution to understanding Pliocene faunal dynamics in western Asia and, more broadly, within the framework of European palaeobiogeography. The European Pliocene, often referred to as the dark epoch, remains comparatively less understood than the preceding Miocene and the subsequent Pleistocene, with well-documented carnivoran guilds virtually non-existent or excavated in the 19th century without proper stratigraphic control. The presence of a primitive form of Nyctereutes at Jradzor is particularly noteworthy: the m2/m1 ratio of 0.62 is considerably high and consistent with the range of individual variance of omnivorous species, suggesting a more omnivorous diet than that of its extant relatives and an ecological profile more consistent with jackal-like canids such as the golden jackal, potentially capable of forming transient foraging groups. The record of Baranogale helbingi significantly increases knowledge of this species both temporally and spatially: regarded so far as a European endemic, this archaic mustelid has a much broader geographical scope than previously thought, with the Jradzor material being the earliest from the Caucasus and Asia and the easternmost occurrence of the species. Like other European Ictonychini, B. helbingi vanished during the Early Pleistocene, most likely due to competition with small, ecologically plastic Mustelinae. Taken together, the carnivoran fauna of Jradzor offers a compelling snapshot of Pliocene ecosystems in a historically under-represented region, reflecting a transitional phase in carnivore evolution marked by increasing specialisation, geographic dispersal and adaptation to changing environmental conditions, and highlighting the strong links between the Jradzor assemblage and the Eastern Paratethyan record, particularly the southern margin of the Eastern Paratethys in the dispersal of vertebrate assemblages between Asia and Europe.

Subject of Research: Pliocene carnivoran palaeontology and palaeoecology at the Jradzor locality, Armenia

Article Title: Pliocene carnivorans from Jradzor, Armenia

Article References: Bartolini-Lucenti, S., Madurell‑Malapeira, J., Marciszak, A., Meyer, M., Becker, D., & Vasilyan, D. (2026). Pliocene carnivorans from Jradzor, Armenia. Swiss Journal of Palaeontology, 145, 579-596. https://doi.org/10.3897/sjp.145.180311

Image Credits: AI Generated

DOI: 10.3897/sjp.145.180311

Keywords: Pliocene, Armenia, South Caucasus, carnivorans, Nyctereutes urartuensis, Baranogale helbingi, Chasmaporthetes lunensis, Pliocrocuta perrieri, palaeoecology, biogeography, Jradzor, scientific research

Carpenter moths belong to the family Cossidae, a group of mostly large, heavy-bodied moths whose caterpillars bore into wood, stems, and roots of trees and shrubs. Unlike many familiar butterflies and moths, cossids are often inconspicuous, cryptically colored, and poorly studied, particularly in mountainous and arid regions where many species live highly localized lives. The scientific knowledge of Georgia’s Cossidae began with Romanoff in 1885 and has grown steadily through the work of researchers such as Didmanidze, whose most recent survey with Yakovlev in 2007 reported fourteen species in ten genera. The biology of many of these species, however, remains largely unknown.

The first new record concerns Cecryphalus strix, a nocturnal species first described by Grum-Grshimailo in 1895. The genus Cecryphalus contains only two known species, which inhabit deserts of Central Asia, North Africa, and the South Caucasus. Until now, only C. strix was known to occur in the South Caucasus, reported from Azerbaijan and Armenia. Three male specimens were documented in Georgia: two from Vashlovani National Park in Dedoplistkaro Municipality at 114 meters above sea level, collected on 6 June 2023, and one from Chachuna Managed Reserve at 248 meters, collected on 17 July 2023. The species is otherwise known from Southern Kazakhstan, Kyrgyzstan, Southern Mongolia, Northwestern China, Tajikistan, Uzbekistan, Turkmenistan, Azerbaijan, Northern Iran, and Southern Armenia. Its host plant and immature stages remain unknown.

The second new record is Dieida ledereri, first described by Staudinger in 1871. Dieida is a small genus of only six species of diurnal Palaearctic carpenter moths, and D. ledereri was previously known in the South Caucasus only from Azerbaijan. A single male specimen was hand collected in Vashlovani National Park at 460 meters above sea level on 16 April 2022. The species is otherwise distributed in Turkey, Azerbaijan, Afghanistan, Syria, Iraq, Israel, and Iran. Its host plant, eggs, and caterpillar are unknown, making the Georgian record an important addition to the understanding of its range.

The single D. ledereri specimen came with an unusual story. The moth was preyed upon by a robber fly of the family Asilidae, which was caught and dropped the specimen on the ground. Unfortunately, the specimen was later damaged during transportation, so no photograph of the undamaged individual exists to fully express its natural colors. The study nevertheless records the appearance of the undamaged moth: it had bipectinate black antennae with two-thirds white on the anterior side, half the length of the forewing, with long black rami. The head, tegula, thorax, and abdomen were densely hirsute and black with a golden terminal part, golden-brown both dorsally and ventrally. The forewings were semitranslucent with a white-ivory basal area, and the hindwings were semitranslucent with white-ivory spots on the submedian area and hind margin.

Because the specimen was damaged, the study provides a detailed description of the male genitalia, which is essential for confirming the identification of cossid species. The tegumen is long and triangular, the uncus small, sclerotized, and triangular peak-shaped, and the socii oval with setae. The arms of the gnathos are long, thin, and fused by membrane, with a triangular membranous gnathos knob. The valvae are lanceolate, six times longer than wide, with a roundish apex; the basal half of the costal margin is convex and the apical half concave, while the apical sixth of the valva is membranous and the basal five-sixths sclerotized. The sacculus is clearly separated from the valvae, slender, and about two-thirds the length of the valvae, gradually narrowing towards its outer end. The aedeagus is cylindrical, slightly longer than the valvae, and without a cornutus, while the vesica is membranous with a large roundish basal part and a gladiate distal part.

The third new record is Dyspessa infuscata, described by Staudinger in 1892. Dyspessa is the most speciose genus of nocturnal carpenter moths, with over 80 species widely distributed in the western Palaearctic. Most species have restricted distributions limited by mountain ranges and deserts, and many are local endemics. Only four species had previously been reported from Georgia: D. alpherakyi, D. pallidata, D. salicicola, and D. ulula. A single female specimen was collected on the Nutsubidze Plateau in Tbilisi at 621 meters above sea level on 24 April 2024. The species is distributed in Turkey, Russia’s North Caucasus, Ukraine’s Crimea, the South Caucasus, and Syria. Because earlier authors did not specify an exact location within the South Caucasus or mention the species in Georgia in later works, this appears to be the first documented record of D. infuscata in the country.

The specimens were collected using light traps, except for the male D. ledereri, which was hand collected. Elevations and GPS coordinates were obtained with a Garmin GPS MAP 64s device, and all specimens are deposited in the personal collection of the author. The moths were examined on the basis of both external and internal morphological characters. Genital dissection followed the technique published by Robinson in 1976 with certain modifications, using a 15 percent potassium hydroxide solution to macerate the full abdomen. Weakly sclerotized structures were stained with chlorazol black and mounted into Euparal. Preserved specimens were photographed with a Canon EOS 60D camera and a Canon EF 60 mm macro lens, with digital images prepared using Zerene Stacker image stacking software and Adobe Photoshop CS6. Genitalia were photographed using a Nikon SMZ745T stereo microscope equipped with a Moticam 2500 camera.

The study places the Georgian discoveries in a broader regional context. Recent collecting efforts in Georgia and adjacent territories have resulted in the descriptions of several new Cossidae species: Holcocerus didmanidzae from Georgia, Stygiodies nupponenorum from Turkey, Isoceras shevnini from Armenia, Phragmataecia effendii from Azerbaijan, and Dieida versicolor from Turkey. Even though the Cossidae fauna of Turkey is relatively well studied compared to that of the South Caucasus, a new genus, Stygia, was recorded there in recent years. Since some of the species listed above are known only from type specimens, and others were described based solely on males or females, investigating the biology and life cycles of these rare species would be highly valuable.

Given the limited study of the Cossidae family in Georgia, the author notes that it is quite possible that rare diurnal genera, such as Stygioides, may still await discovery in the country’s remote arid regions. The new records highlight how much remains to be learned about the moth fauna of the South Caucasus, where arid lowlands, mountain ranges, and isolated valleys create conditions for localized and often endemic species. For a family whose caterpillars bore invisibly inside wood and whose adults fly briefly and cryptically, every documented specimen represents a meaningful expansion of scientific knowledge, and the Georgian carpenter moth checklist now stands at seventeen species in twelve genera.

Subject of Research: First records of two genera and three carpenter moth species (Lepidoptera, Cossidae) in Georgia

Article Title: First records of two genera and three carpenter moths ( Lepidoptera , Cossidae ) from Georgia

Article References: Japaridze, L.-G. (2024). First records of two genera and three carpenter moths (Lepidoptera, Cossidae) from Georgia. Caucasiana, 3, 273-280. https://doi.org/10.3897/caucasiana.3.e140978

Image Credits: AI Generated

DOI: 10.3897/caucasiana.3.e140978

Keywords: Cossidae, carpenter moths, Lepidoptera, Georgia, Cecryphalus strix, Dieida ledereri, Dyspessa infuscata, South Caucasus, biodiversity, new records, Vashlovani National Park, entomology

The specimens were collected on 16 July 2022 at Alvank, at 484 meters above sea level, with coordinates recorded using a Garmin GPS device. In total, ten male specimens of Phragmacossia territa, a species first described by Staudinger in 1879, were taken, seven of them destined for the personal collection of Jari Junnilainen and three for that of Lasha-Giorgi Japaridze. A single female of Stygia mosulensis, described by Daniel in 1965, was also captured at the same site and entered Junnilainen’s collection. All were attracted by a light trap, a standard but remarkably effective tool for surveying nocturnal Lepidoptera.

Carpenter moths belong to the family Cossidae, a lineage whose caterpillars are famous for boring into the wood of trees and shrubs, sometimes causing serious damage to orchards and vineyards. The adults are often heavy-bodied, cryptically patterned insects that can be difficult to tell apart by eye. That is why the team behind the new study, which included Risto Haverinen as collector and researchers working with material distributed between two private collections, relied on both external appearance and internal anatomy to confirm the identifications.

The diagnostic work followed established laboratory protocols. Genital dissection, the gold standard for separating closely related cossid species, was performed using the technique published by Robinson in 1976 with modifications suggested by Fibiger and Goater in 1997. Whole abdomens were macerated in a 15 percent potassium hydroxide solution to clear soft tissue before examination. Photographic documentation was equally rigorous: images of Stygia mosulensis were taken with an Olympus OM-D E-M1 Mark II camera fitted with a 60 millimeter macro lens and stacked with Helicon Focus software, while Phragmacossia territa was photographed with a Canon EOS 90D and a Canon EF 60 millimeter f/2.8 macro lens, with images prepared using Zerene Stacker and Adobe Photoshop.

The results are striking because of what they reveal about biogeography. Phragmacossia is a genus of sixteen known nocturnal species with a mainly Palaearctic distribution, and P. territa itself was already known from the Middle East and North Africa, Central and Southwest Asia, including adjacent areas of Türkiye and Iran. Stygia, by contrast, is a far smaller and stranger genus, with only four known species scattered across Southern Europe, the Canary Islands, Morocco, Iran, Türkiye and Iraq. S. mosulensis had previously been recorded in Iraq, Iran, Greece, southern Bulgaria and southern Türkiye, and probably Morocco, so its appearance in Armenia fills a conspicuous gap between its Turkish and Iranian populations.

Perhaps the most intriguing question raised by the new record concerns the daily rhythm of Stygia mosulensis. Members of the subfamily Stygiinae, to which the genus belongs, are generally reported as diurnal, and related species such as Stygia australis and Stygia hades have been described as day-flying insects. Yet both the historical literature and the new Armenian specimen suggest something different for S. mosulensis: it was collected at night, at a light trap, just as earlier records indicate. The authors propose that this species may in fact be nocturnal or crepuscular, a hypothesis that can only be tested through further investigation into its biology and life cycle.

That uncertainty is compounded by a basic gap in knowledge: the host plant and immature stages of S. mosulensis remain entirely unknown. No one has described its caterpillar, its pupa, or the woody plants its larvae may bore into. For a species now documented on three sides of the Armenian border, the absence of any life-history data is a reminder of how little is known even about insects living within easy reach of research institutions. The same applies to several recently described Caucasus cossids, some of which are known only from their type specimens and others from a single sex.

The new records arrive amid a remarkable run of discoveries in the South Caucasus. Recent collecting has yielded Holcocerus didmanidzae from Georgia, Stygiodies nupponenorum from Türkiye, Isoceras shevnini from Armenia, Phragmataecia effendii from Azerbaijan, Dieida versicolor from Türkiye, and, as recently as 2025, Dyspessa gayane from Armenia. New genera have been documented for Georgia, and new species have been described from both Azerbaijan and Armenia. Yet the family’s regional fauna remains poorly studied, with only general overviews published for Georgia and Azerbaijan to date.

The authors argue that the limited state of knowledge makes further discoveries likely. Given how little the Cossidae of the South Caucasus have been surveyed, they consider it probable that additional representatives of the genus Stygia still await discovery in remote regions of Georgia and Azerbaijan. The Alvank site, perched at the southern edge of Armenia near the Iranian border, sits in exactly the kind of transitional zone where Palaearctic and Middle Eastern faunas meet, making the Meghri area a promising target for future surveys.

For now, the ten males of Phragmacossia territa and the lone female of Stygia mosulensis stand as the first evidence that these two genera occur in the South Caucasus, extending the known ranges of both across a mountain barrier that has long shaped the distribution of life in the region. The study, published on 17 June 2025 in Caucasiana under the DOI 10.3897/caucasiana.4.e158970, is a small paper with an outsized message: even in a well-traveled corner of Eurasia, a single night at a light trap can still add entire genera to the map.

Subject of Research: First records of the carpenter moth genera Phragmacossia and Stygia in Armenia and the South Caucasus

Article Title: First records of two genera Phragmacossia Schawerda, 1924 and Stygia Latreille, [1802] in the South Caucasus ( Lepidoptera , Cossidae )

Article References: Japaridze, L.-G., Haverinen, R., Junnilainen, J., Kalashian, M., & Pototskin, A. (2025). First records of two genera Phragmacossia Schawerda, 1924 and Stygia Latreille, [1802] in the South Caucasus (Lepidoptera, Cossidae). Caucasiana, 4, 59-64. https://doi.org/10.3897/caucasiana.4.e158970

Image Credits: AI Generated

DOI: 10.3897/caucasiana.4.e158970

Keywords: entomology, Lepidoptera, Cossidae, Armenia, South Caucasus, biodiversity, new records, First, records, genera, Phragmacossia, Schawerda

The finding, published in the journal Caucasiana, is accompanied by a second milestone: the confirmation of Mystacides azureus, a long-horned caddisfly with a striking metallic blue sheen, as a resident of Georgia. Together, the two records raise the number of caddisfly species known from the country to 250, grouped into 55 genera and 19 families, and they underscore how much remains to be learned about the aquatic insects of the Caucasus.

Caddisflies, the order Trichoptera, are among the most diverse groups of freshwater insects globally, yet their fauna remains poorly studied in many developing countries. Georgia is no exception. The biodiversity of the country’s freshwater ecosystems has long been under-researched, and aquatic insects have been particularly neglected. This neglect is reflected in the pace of recent discovery: more than 100 caddisfly species have been added to the Georgian list in the 2020s alone, and researchers believe many more remain to be found.

The history of caddisfly research in Georgia and the Caucasus stretches back to the nineteenth century, when Robert McLachlan produced some of the earliest records. The most substantial early contributions, however, came from Andrey Martynov, who described 35 new species and recorded 46 species for the first time in Georgia in a series of publications between 1913 and 1934. In the following decades, local and foreign researchers added 18 more species and recorded numerous others for the first time in the country. A new era began in 2019, driven by the exceptionally productive work of János Oláh, who has described over a hundred species from the region.

Despite this progress, much of the Georgian caddisfly fauna remains unexplored. Through their investigation of the country’s caddisflies, the authors of the new study present two additional species: Calamoceras illiesi Malicky & Kumanski, 1974, and Mystacides azureus Linnaeus, 1761, both recorded for the first time in Georgia.

The family Calamoceratidae, to which Calamoceras illiesi belongs, is primarily found in tropical and subtropical regions, with its presence in Europe limited to the genus Calamoceras. The larvae of this family are among the largest case-bearing caddisfly larvae, partly due to their distinctive body shape. They can be differentiated from similar families by morphological characteristics such as a transverse row of about 16 long stout setae in the central part of the labrum, sharp ventral anterior margins of the pronotum and dorsal plates of the mesonotum, and specific blurred margins. Larvae are found sporadically in detritus accumulations of stream pools.

Calamoceras larvae are characterised by a combination of features: a transverse row of 22 to 32 stout setae on the labrum; a hook-shaped fore trochantin; a pronotum with elongated and pointed anterolateral corners; roughly equal-length middle and hind legs with short forelegs; two sclerotised plates on the mesonotum; a membranous metanotum with four very small seta-bearing sclerites; and anal claws that possess an accessory hook. The case is made from pieces of wood. To date, the genus includes three species: C. markusha, C. illiesi, and the recently described C. riffensis from Morocco.

In the new study, researchers collected 2 females and 17 larvae of Calamoceras illiesi from a small, unnamed mountain brook in the Kvereti Managed Reserve, at an altitude of 1,774 metres above sea level. The larvae were caught using a hand net, while the adults were caught with a butterfly net. All specimens were preserved in 95% ethanol and stored in a freezer at −22 °C. When detailed examination of adults was necessary, the abdomens were cleaned in a 10% KOH solution and transferred to glycerol for further preservation. Identification of the larva and imago was based on the descriptions by Sipahiler and Malicky, and all examined material is kept in the collection of the Museum of the Institute of Zoology, Ilia State University, Tbilisi.

Calamoceras illiesi has previously been recorded in Bulgaria, Croatia, Greece, Türkiye, and north-western Russia. In the Caucasus, the genus is represented by a single species, C. illiesi, which occurs in the Western Caucasus, including Novorossiysk and Sochi, and in the North Caucasus, in the southwest of Dagestan. The new record extends the known range of the species into the South Caucasus for the first time. Interestingly, an unknown representative of the genus Calamoceras was recorded by Martynov in 1913 in Akhali Atoni, in western Georgia. The authors assume that this record probably belongs to C. illiesi, but this assumption requires further confirmation, given the high diversity of trichopterans in the region.

In terms of its habitat, despite extensive and repeated sampling, Calamoceras illiesi larvae were found in only one stream, which fully matches the species’ known habitat preferences. The specimens were located at the edge of a small pond, one to two metres wide, formed by a mountain spring, where the slowed flow allows the accumulation of woody debris and leaf litter. Larvae were absent from nearby streams, suggesting a small, microhabitat-dependent population and highlighting its vulnerability. Consequently, protecting this known habitat is essential for the conservation of this rare species.

The second species confirmed in the study, Mystacides azureus, belongs to the family Leptoceridae, commonly known as long-horned caddisflies, one of the largest families within the order, comprising over 1,970 described species classified into 46 extant genera globally. Adults of the family are characterised by extremely long, slender antennae, often two to three times the length of the forewings, which distinguishes them from most other caddisfly families. The head is broad and may bear distinctive sensory projections in some genera. Maxillary palps are five-segmented in both sexes, often modified in males, with the terminal segment annulated or whip-like. The forewings are narrow and covered in dense hairs, with simplified venation.

The genus Mystacides comprises 25 documented species and occurs across temperate regions of the Northern Hemisphere as well as tropical and subtropical areas of Asia. Adults are medium-sized, with elongated slender bodies typically ranging from 7 to 12 millimetres, and wings that often carry a distinct metallic, dark brown to bluish iridescent sheen. The forewings are usually without any pattern or faintly spotted. Maxillary palps are composed of five parts, with sexually dimorphic segmentation in particular species, and the head is hypognathous, with large compound eyes.

Mystacides azureus is one of the most widely distributed species of the genus in Europe, having been recorded in numerous countries including the United Kingdom, Germany, France, Italy, and the Balkan Peninsula. The species is also present in Japan, alongside two newly described species within the M. azureus species group. In the Caucasus region, M. azureus was previously reported from Russia. Notably, the first photographic record of this species in Georgia was made by amateur naturalist Giorgi Makharadze, who posted an image on the Facebook group Wildlife in Georgia in 2020.

In the new study, researchers collected specimens of Mystacides azureus from several locations across Georgia: 2 males and 1 female from Gaghma Dvabzu village in Guria, at 433 metres above sea level; 2 males, 2 females, and 1 larva from Kutaisi in Imereti, at 235 metres above sea level; and 1 male and 1 female from Lia in Samegrelo, at 634 metres above sea level. The initial photograph observation on social media was thus supported by the collection of specimens from multiple locations in the country, providing firm evidence of the species’ occurrence in Georgia.

With the two new species reported in the study, the diversity of Georgian caddisfly fauna has increased to 250 species grouped into 55 genera and 19 families. Of these, 109 species, approximately 44%, are endemic to Georgia, while 141 species, approximately 56%, are endemic to the Caucasus. Among the species-rich families, Philopotamidae stands out with a particularly high level of local endemism, with 16 of 26 species found only in Georgia. An additional 37 Caucasian endemics remain unrecorded in Georgia, reinforcing the need for further research.

The new records highlight both the richness and the fragility of Georgia’s freshwater insect fauna. The discovery of Calamoceras illiesi in a single mountain brook, dependent on a microhabitat of slow-flowing water and accumulated woody debris, serves as a reminder that even common-seeming species can be rare and vulnerable at the local scale. As researchers continue to explore the caddisflies of the Caucasus, the protection of these small, specialised habitats will be essential for the survival of the species that depend on them.

Subject of Research: First record of Calamoceras illiesi and confirmation of Mystacides azureus (Trichoptera) in Georgia

Article Title: The first record of Calamoceras illiesi Malicky & Kumanski, 1974 ( Trichoptera , Calamoceratidae ) and confirmation of the occurrence of Mystacides azureus Linnaeus, 1761 ( Trichoptera , Leptoceridae ) from Georgia

Article References: Memishishi, A., Dzuliashvili, K., & Japoshvili, B. (2025). The first record of Calamoceras illiesi Malicky & Kumanski, 1974 (Trichoptera, Calamoceratidae) and confirmation of the occurrence of Mystacides azureus Linnaeus, 1761 (Trichoptera, Leptoceridae) from Georgia. Caucasiana, 4, 165-171. https://doi.org/10.3897/caucasiana.4.e172790

Image Credits: AI Generated

DOI: 10.3897/caucasiana.4.e172790

Keywords: caddisflies, Trichoptera, Calamoceras illiesi, Mystacides azureus, Georgia, South Caucasus, freshwater biodiversity, endemic species, aquatic insects, species records, first, record

The researchers start from a puzzle that has haunted gender and development work for decades. Empowerment programmes have multiplied, representation targets have been met in many institutions, and yet inequality persists in the arrangements governing access to capital, care, legitimacy and opportunity. Earlier scholars such as Srilatha Batliwala and Andrea Cornwall and Alina Rivas have warned that empowerment can lose its critical force when struggles over power are translated into programmes centred on individual capacity or measurable inclusion. The South African evidence is sobering: women who reach senior leadership still encounter gender stereotypes, work-life pressures and barriers to executive advancement, while digital empowerment initiatives in the public sector remain constrained by institutional and resource limits. Entry, in other words, is not the same as influence.

To explain why, the authors build their argument on three theoretical traditions that have usually travelled separately. The first is Raewyn Connell and James Messerschmidt’s account of hegemonic masculinity, which treats gendered authority not as a personality trait of individual men but as historically specific configurations of practice that legitimate unequal relations across households, organisations, states and global governance. The second is Sara Ahmed’s critique of happiness and her concept of the happy object, which shows how belonging, professional success and recognition become attached to conformity, so that women who challenge unequal arrangements are recast as negative, ungrateful or difficult. The third is Patricia Hill Collins’s Black feminist standpoint epistemology, particularly her idea of the outsider within, which explains how people positioned at the margins of institutions can see contradictions invisible from the centre, even when their insight is not credited. Intersectionality runs through all three as a cross-cutting analytic, shaping who is exposed to sanction, whose testimony is believed and who captures the benefits of reform.

The study uses a theory-synthesis design rather than an empirical test. Following Elisabeth Jaakkola’s guidance on conceptual articles, the authors searched Web of Science, Scopus and Google Scholar across six conceptual domains, from empowerment and leadership to affect and dissent, standpoint epistemology, collective agency and post-socialist settings, updating the literature base in August 2026. Crucially, they deliberately included disconfirming scholarship: evidence that women can reproduce unequal arrangements, that dissent is not automatically progressive, that marginalisation does not by itself generate critical insight, and that collective mobilisation can be captured. Selection closed at theoretical sufficiency, the point at which additional sources no longer altered the core constructs, rival explanations or boundary conditions.

Out of this synthesis emerge three generative relational capacities. Structural diagnosis identifies the institutional mechanisms through which inequality is produced and reproduced, redirecting attention from what individual women lack to how rules, networks and political-economic conditions distribute authority and constraint. Affective refusal contests the emotional expectations, gratitude, agreeableness, loyalty, composure, that protect unequal arrangements, and it need not take the form of open confrontation; strategic silence, documentation and coalition-building can count as refusal where dissent carries real risk. Situated knowledge mobilisation converts socially positioned experience into collectively interpreted, institutionally actionable knowledge, a far more demanding process than consultation, because institutions can collect testimony while retaining control over problem definition and interpretation.

The framework’s most original move is the identification of a translation mechanism the three traditions individually lack. The authors call it collective capacity: the relational, material, epistemic and organisational conditions through which actors interpret problems together, distribute risks, coordinate action and convert diagnosis, refusal and knowledge into sustained institutional leverage. Its elements include shared but revisable problem definitions, trust and solidarity, material and temporal resources, protection from retaliation, representative legitimacy and internal accountability. The logic is starkly practical: dissent concentrated in one individual can be neutralised through removal, promotion or exhaustion, whereas distributed dissent is far harder to separate from the institutional arrangement it targets.

Six propositions formalise the pathway, and each comes with observable indicators that future researchers can test. Structural diagnosis is expected to redirect action from individual deficits towards institutional mechanisms. Affective sanctions are expected to condition whether critique can be expressed and sustained, operating through anticipatory sanction, the self-censorship that occurs before anything is said. Situated knowledge gains influence when its producers retain interpretive ownership and when it connects to recognised decision and accountability processes. Collective capacity translates the three capacities into institutional reconfiguration, meaning actual changes in authority, resources, recognition, knowledge standards, accountability and work organisation. Intersectional distribution determines whether reforms produce structural transformation or merely selective inclusion, since benefits tend to accrue first to women already closest to institutional power. Finally, institutional embedding, durability and recursive feedback determine whether change persists beyond particular leaders or funding cycles.

The authors stress that transformative women’s leadership is defined by institutional consequence rather than by women’s presence or any particular leadership style. They distinguish entry from influence, representation from redistribution, and individual development from institutional change. Training, mentoring and networking may expand capabilities while leaving underlying arrangements intact, and a leader can be transformational in style while leaving every consequential arrangement untouched. Five boundary conditions, material resources, political and institutional conditions, intersectional positioning, protection from retaliation, and collective legitimacy with internal accountability, shape whether the capacities can be exercised, translated and sustained at all.

To demonstrate the framework’s reach, the study uses the South Caucasus, Armenia, Azerbaijan and Georgia, as a comparative boundary illustration rather than an empirical test. Green transitions in the region are frequently framed as technical and economic programmes, but the framework asks who determines transition priorities, who controls investment, land and technology, and which qualifications count as expertise. World Bank assessments show men accounted for 89 percent of engineering students in Armenia, and in Azerbaijan women constituted just 10.7 percent of the energy-sector workforce, concentrated in office-based rather than technical or leadership roles. The framework also reverses the conventional deficit assumption: institutions may lack the ecological knowledge produced through women’s unpaid and informal labour, so a gender-just transition requires two-directional learning, not simply training women to fit existing technical categories.

The conclusion is deliberately uncomfortable for empowerment industries everywhere. Insight without collective protection, voice without authority and policy without embedded practice are not anomalies in this account; they are its predicted outcomes when the translation mechanism is absent. The authors propose that funders and evaluators assess initiatives against six questions: whether women exercise influence over agendas rather than advisory presence, whether decision rights have moved, whether budgets and time have been reallocated, whether situated knowledge shapes problem definition, whether dissent is institutionally protected, and whether changes survive leadership turnover. The propositions remain theoretically derived claims awaiting empirical examination, but the framework relocates the central question of women’s empowerment, asking not whether women have entered institutions, but whether the terms on which they participate are theirs to set, and whether the power to set them endures.

Subject of Research: A theory-synthesis framework explaining how women's leadership in the Global South can move from institutional access to durable institutional transformation

Article Title: Beyond hegemonic orders and ‘happy objects’: reimagining women’s empowerment and intersectional leadership in the global south

Article References: Nodipha, O. G., & Mdhluli, N. I. (2026). Beyond hegemonic orders and ‘happy objects’: reimagining women’s empowerment and intersectional leadership in the global south. Frontiers in Sociology, 11, Article 1957241. https://doi.org/10.3389/fsoc.2026.1957241

Image Credits: AI Generated

DOI: 10.3389/fsoc.2026.1957241

Keywords: women's empowerment, intersectional leadership, hegemonic masculinity, feminist standpoint epistemology, affective refusal, collective capacity, institutional transformation, Global South, South Caucasus, green transitions, gender justice, epistemic injustice

Ghost moths, together with their relatives the swift moths, make up the family Hepialidae, one of the most evolutionarily significant lineages in the entire order Lepidoptera. With 725 described species, Hepialidae is the most diverse family among the primitive moths, and it occupies a pivotal position in the butterfly and moth family tree. The family belongs to the infraorder Exoporia, which is the sister group to the Heteroneura, the vast assemblage that contains the overwhelming majority of living butterflies and moths. Together, these two lineages account for more than ninety percent of all extant lepidopteran species, making the ghost moths a living window into the early evolution of the order.

The biology of Hepialidae is as distinctive as their ancestry. Unlike most moths, whose caterpillars feed openly on leaves, the larvae of most ghost moth species live underground, feeding on the roots of woody or herbaceous plants or consuming fallen foliage in the soil. Another major feeding strategy involves boring into the stems and branches of trees and shrubs, where the larvae feed on callus tissue regrowing at wound surfaces or, in some species, on soft woody tissues. A smaller minority of species feed on plant detritus throughout their development. This subterranean, often cryptic lifestyle is one reason the family remains poorly surveyed in many parts of the world, including the mountainous Caucasus.

Geographically, Hepialidae are found on every major landmass except Antarctica, with notable regional absences in Madagascar, the Caribbean islands and tropical West Africa. Northern Eurasia currently hosts eight genera comprising 21 species, most of which are endemic to the region between Europe and the Caspian Sea or represented only there. The southern boundary of the group runs along the Mediterranean Basin and continues eastwards through Jordan, Türkiye, Georgia and northern Iran. This makes the Caucasus Mountains, which span Georgia, southern Russia, northeastern Türkiye, Armenia and Azerbaijan, a potentially crucial biogeographic frontier for understanding where the western Eurasian ghost moth genera begin and end.

Before the new study, the Hepialidae fauna of Georgia was thought to consist of just four species. Three of these are regional endemics or relics: Zenophassus schamyl, described in 1888, and two species of Triodia, T. laetus and T. mlokossevitschi, described in 1877 and 1884 respectively. The fourth, Korscheltellus lupulinus, is a more broadly distributed European species whose southern distributional limit falls in Georgia. The natural history of the endemic Triodia species remains poorly understood, and the taxonomic status of T. mlokossevitschi is itself uncertain, because its type specimen has never been dissected and described according to modern standards. Only Z. schamyl has been studied in any detail, largely because of its economic importance as a pest of grapevines, a problem documented by Soviet-era researchers in the early and mid-twentieth century.

The newly recognized specimens were all collected by E. Didmanidze and deposited in the Simon Janashia Museum of Georgia. The first, a female, was taken at Bakhmaro in the western region of Guria on 18 July 1961. A male followed from Martvili in Samegrelo-Zemo Svaneti on 4 July 1962, and a second male was collected at 1800 meters above sea level in Khevsureti, in the Mtskheta-Mtianeti region, on 12 July 1970, though without an exact locality. The three specimens span three different regions and three different years, suggesting that Hepialus humuli has been present in Georgia for at least 64 years while remaining almost entirely overlooked by collectors and researchers.

Why would such a conspicuous moth escape notice for so long? The answer likely lies in a combination of behavior and geography. Hepialus humuli is famous among European naturalists for its crepuscular habits: males fly at dusk in short, ghostly patrols, and the species is most active in the narrow window between sunset and full darkness. Combined with its occurrence in remote mountainous regions, this twilight lifestyle makes the species easy to miss with conventional daytime collecting. The species is nonetheless widespread across Europe, extending eastwards to central Siberia, and its known southern distributional limit in western Eurasia runs along the Pyrenees and north of the Mediterranean, along the European Alps, the Balkans and north of the Black Sea. A report of the species from North Ossetia, on the Russian side of the Greater Caucasus, had already suggested that its occurrence in northern Georgia was to be expected.

The biogeographic implications of the Georgian records extend well beyond a single species. The localized endemism of Hepialidae in the Caucasus is comparable to patterns observed in other montane regions along the northern margin of the Mediterranean Basin. Researchers have suggested that the diversification of the northern Eurasian ghost moth genera was shaped by tectonic history, particularly phases of geological uplift associated with the closure of the western Tethys Sea, the ancient ocean that once separated Eurasia from Africa and Arabia. As mountain ranges rose and climates shifted, ancestral populations were fragmented and isolated, giving rise to the endemic lineages that characterize the Caucasus today. Against this backdrop, confirming the presence of a widespread Eurasian genus like Hepialus in the South Caucasus helps to map the boundary zones where broadly distributed faunas meet highly localized endemics.

The distribution of ghost moth genera around the Caucasus is strikingly asymmetric. The western Eurasian genera Gazoryctra, Hepialus, Pharmacis and Phymatopus have previously published ranges lying north of the Caucasus, while Korscheltellus reaches its southern limit in Georgia. Only Triodia extends well south of the mountain chain, reaching as far as southern Israel and Jordan. Even there, however, taxonomy remains unsettled: the status of T. froitzhami from Israel and Jordan is uncertain because its type specimen has not been dissected and no clear external diagnostic differences have been established. The Georgian Hepialus record therefore raises a further question highlighted by the study’s authors: whether the species may also occur further south in Central Asia, particularly in Kazakhstan and adjacent countries, a possibility that had previously been difficult to evaluate without a confirmed South Caucasus occurrence.

The study also carries a practical message for biodiversity science. Museum collections assembled decades ago, sometimes by a single dedicated collector, can hold discoveries of continental significance, but only when they are re-examined with modern taxonomic knowledge. The three Georgian specimens were photographed with a Canon EOS 60D camera fitted with a macro lens, and image stacks processed with focus-stacking software now allow researchers worldwide to compare the Georgian material with European reference specimens. The authors express particular gratitude to the administration of the Simon Janashia Museum of Georgia for granting access to the collection. As systematic surveys of the Caucasus continue, the ghost moths of Georgia may yet yield further surprises, and the humble museum drawer has once again proven to be one of biology’s most productive frontiers.

Subject of Research: First record of the ghost moth genus Hepialus and the species Hepialus humuli in Georgia and the South Caucasus, based on museum specimens

Article Title: New Light on Old Ghosts: First Record of the genus Hepialus Fabricius, 1775 ( Lepidoptera , Hepialidae ) from Georgia

Article References: Japaridze, L.-G., Makharadze, G., Petrov, V., & Grehan, J. R. (2026). New Light on Old Ghosts: First Record of the genus Hepialus Fabricius, 1775 (Lepidoptera, Hepialidae) from Georgia. Caucasiana, 5, 25-29. https://doi.org/10.3897/caucasiana.5.e176387

Image Credits: AI Generated

DOI: 10.3897/caucasiana.5.e176387

Keywords: Hepialidae, Hepialus humuli, ghost moths, Georgia, South Caucasus, Lepidoptera, museum specimens, biogeography, Caucasus Mountains, new record, taxonomy, Simon Janashia Museum

The study was conducted during two contrasting periods in 2024. Between late May and the end of July, the team surveyed lines in the Armavir, Ararat and Kotayk provinces, focusing on areas near the Khosrov Forest State Reserve and the Metsamor and Sardarapat Important Bird Areas. Then, from mid-September to early November, attention shifted to Shirak province, a major raptor migration corridor dominated by high-altitude steppe and agricultural mosaics with sparse tree cover. In total, the surveyors walked transects along 2,071 poles, examining a 40-metre-wide belt centred on each line.

The technical logic behind electrocution is straightforward. On low- and medium-voltage distribution lines, birds die when they bridge the gap between energised conductors, or between a conductor and grounded hardware such as a metal crossarm, while perching or taking off. Concrete poles with closely spaced, uninsulated components are particularly dangerous, which is why the team prioritised them over wooden poles once initial inspections suggested they posed the greater risk. Carcasses found within a five-metre radius of a pole were classified as probable electrocution victims, and each was logged with species, pole type, coordinates and distance from the structure.

The seasonal and regional contrast was stark. During the breeding season across Armavir, Ararat and Kotayk, the team recorded just 24 electrocution cases. In Shirak during autumn migration, they recorded 228. The authors caution that this difference may reflect both seasonal patterns and landscape differences rather than a single cause, but the pattern is consistent with a well-known mechanism: open steppe and farmland with few natural perching structures forces birds onto power infrastructure. Shirak’s lines connect villages across open fields where the only elevated perches are poles.

Species composition tells its own story. Eurasian magpies led the count with 49 records, followed by common kestrels and rooks with 34 each, western jackdaws with 33 and common buzzards with 31. Together these five species accounted for nearly 72 percent of all casualties. Raptors made up 97 individuals, or 38.5 percent of the total, a figure with real conservation weight because many raptors are large-bodied, slow-reproducing species whose populations are sensitive to even modest increases in adult mortality. Among the victims were species listed in the Red Book of Animals of Armenia: a juvenile golden eagle, a Eurasian eagle-owl and lesser kestrels, all classified as Vulnerable, plus four probable Armenian gulls from the colony at Lake Arpi.

Age data, though fragmentary, added a troubling dimension. Of the 53 individuals whose age could be determined, 83 percent were juvenile or immature; within the raptor subset, the figure rose to 91 percent of 34 age-determined birds. Young raptors, still perfecting their hunting and perching skills, appear especially vulnerable to poorly designed poles. Sex could rarely be determined from carcass condition, precluding comparisons between males and females.

Pole type mattered. Of the 252 records, 239, or 94.8 percent, occurred on concrete poles, and 216, or 85.7 percent, were on intermediate poles, the single-pole structures that dominate the network. Anchor poles and switch poles were implicated far less often. The authors stress that this pattern partly reflects their own survey design, which deliberately prioritised concrete lines and excluded wooden-only lines in the second phase, so relative risk between materials cannot be directly compared. Still, the concentration of deaths on a specific, well-defined infrastructure type is precisely what makes targeted retrofitting feasible.

Spatial mapping revealed unmistakable hotspots. The most critical cluster lay around Lake Arpi National Park, a Ramsar-listed wetland that hosts Armenia’s only known breeding site for the Dalmatian pelican and a large colony of Armenian gulls. There, parallel uninsulated 10 kV lines run about 20 metres apart along the lakeshore, cutting through grassland heavily used by raptors and waterbirds. Other hotspots included the stretch between Zorakert and Paghakn villages, the route from Paghakn to Ardenis, and the Bavra–Tavshut highway. Raptor-specific mapping flagged the Hartashen–Tashir line, where a golden eagle, multiple buzzards and kestrels died, and the Karaberd–Maralik section in open steppe. One pole in Shirak accounted for ten carcasses.

The study also produced an unusual bonus dataset: informal carcass persistence observations. Because scavenger removal and observer detection rates could not be formally measured, the reported numbers represent minimum mortality, a limitation the authors acknowledge openly. But by marking fresh carcasses and revisiting them, the team found that buzzard and kestrel remains often persisted for more than a month in open grassland, and one golden eagle carcass stayed on its pole for at least 301 days, from late September 2024 to late July 2025, eventually mummified. High persistence of large raptor remains suggests detection probability in this landscape may be relatively good, though small carcasses likely vanish faster. Notably, only nine carcasses bore visible burn marks, ranging from singed toes to a magpie whose body and surrounding grass were scorched, meaning the absence of burns cannot rule out electrocution as a cause of death.

The path forward is well established elsewhere. Insulating energised components, reconfiguring crossarms, installing perch deterrents and routing lines away from key habitats have all been shown to cut electrocution deaths substantially without compromising power delivery. Encouragingly, the survey noted roughly seven kilometres of newer, fully insulated cable lines near villages in Ararat province where no casualties were recorded. The researchers call on Electric Networks of Armenia and other line owners to prioritise retrofitting at the identified hotspots, especially around Lake Arpi and the Shirak migration corridors, and urge the government to embed bird-safe standards into energy planning, technical regulations and environmental assessment. As the authors frame it, power infrastructure represents a form of human–wildlife conflict in which essential development imposes severe but entirely preventable biodiversity losses, and Armenia now has the first evidence base to act on.

Subject of Research: Bird electrocution mortality on medium-voltage distribution power lines in Armenia

Article Title: Avian mortality associated with medium-voltage distribution lines in selected regions of Armenia: results of a pilot survey

Article References: Harutyunyan, L., Kovalev, V., Markosyan, A., Tarasyan, N., Aghajanyan, L., & Gudkov, M. (2026). Avian mortality associated with medium-voltage distribution lines in selected regions of Armenia: results of a pilot survey. Caucasiana, 5, 41-57. https://doi.org/10.3897/caucasiana.5.e188959

Image Credits: AI Generated

DOI: 10.3897/caucasiana.5.e188959

Keywords: bird electrocution, power lines, Armenia, raptors, Shirak province, Lake Arpi National Park, concrete poles, migration, Red Book species, mitigation, South Caucasus, conservation

The team, led by Mariya Antonosyan of the Max Planck Institute for Geoanthropology together with Anahit Hovhannisyan and colleagues, analyzed dental calculus from 77 individuals excavated at 22 archaeological sites across Armenia, spanning the Late Neolithic to the Middle Ages. Dental calculus, the mineralized bacterial biofilm that accumulates on teeth, acts as a remarkable time capsule. As it forms during life, it traps fragments of proteins from everything a person ate, preserving them for millennia in a protective mineral matrix. Using liquid chromatography-tandem mass spectrometry, the researchers extracted and identified these ancient proteins, screening each sample against a curated database of oral and contaminant proteins to ensure that only well-preserved, authentic samples were analyzed.

Of the 77 samples, 60 showed well-preserved oral signatures, indicating their suitability for dietary analysis. Within this group, dairy peptides were identified in 16 individuals from nine sites. The overwhelming majority of these milk proteins were beta-lactoglobulin, a major whey protein found in ruminant milk, detected in 14 samples, while beta-casein appeared in two others. Crucially, some peptides allowed the researchers to identify the species of origin: seven peptide matches pointed to goat milk, four to cattle, and four to sheep, with a larger group of 55 matches that could not distinguish between sheep and cattle. This taxonomic resolution is what transforms a simple detection of milk into a nuanced picture of ancient herd management and dietary choice.

The chronological pattern that emerged is striking. The earliest individual in the study, an adult male aged 45 to 60 from the Late Neolithic site of Masis Blur, showed no direct evidence of milk consumption, nor did any of the Early Bronze Age individuals associated with the Kura-Araxes cultural complex. The earliest direct evidence of milk drinking in the Armenian dataset comes from the Middle Bronze Age, around the 2nd millennium BCE, when all four individuals analyzed, buried at the high-altitude sites of Sotk-2 and Sotk-10 near Lake Sevan, carried milk proteins in their calculus. From that point onward, milk consumption persisted continuously through the Late Bronze, Iron, Classical, and Medieval periods, with both the intensity of consumption and the diversity of exploited animals increasing over time.

Geography mattered as much as chronology. Milk proteins were recovered predominantly from high-altitude sites situated above 1,500 meters above sea level. Among prehistoric individuals with confirmed protein preservation, all seven who yielded dairy peptides came from sites above 1,600 meters, while six individuals from lowland sites showed no milk signal at all. The researchers suggest that milk may have functioned as an adaptive resource in harsh highland environments, where crop cultivation was difficult and livestock offered a reliable source of hydration and nutrition without the need to slaughter valuable animals. A parallel pattern has been observed on the Tibetan Plateau, where ancient milk proteins were found only at high-elevation sites beyond the limits of agriculture. Yet the relationship is not straightforward: seven high-altitude individuals from the Iron Age and Medieval periods passed preservation screening without yielding dairy proteins, indicating that elevation alone cannot explain the mosaic of dietary preferences across landscapes and centuries.

The genetic side of the study tells an equally compelling story. Adult digestion of lactose, the sugar in milk, depends on lactase persistence, a trait controlled by regulatory variants near the LCT gene. The team screened five known lactase persistence-associated variants in 158 modern Armenian individuals and found only the European variant, 13910C>T, at a modest frequency of roughly 6 to 8 percent. The other four variants, including the 13915T>G allele that predominates in the Near East just to Armenia’s south, were entirely absent. This finding resolves previously conflicting estimates for Armenians, which had ranged from 2.7 to 18 percent, and highlights the population’s distinctive genetic position at the crossroads of Europe and the Middle East.

Ancient DNA screening added a temporal dimension to this genetic picture. After imputing lactase genotypes in published ancient genomes from Armenia, the researchers directly examined sequencing reads covering the rs4988235 position using stringent filtering criteria designed to guard against false positives, a critical safeguard because the C>T transition at this site mimics characteristic ancient DNA damage patterns. The earliest confirmed carrier of the derived allele in Armenia dates to the Late Urartian period, around 680 to 550 BCE, approximately 3,000 years ago. Results for eight other samples, including Early Bronze Age specimens, could not be confirmed and may represent imputation artifacts. The confirmed appearance of the allele post-dates a documented Late Bronze and Iron Age genetic influx into the region, suggesting the variant may have been introduced through population interactions rather than arising locally.

Placed side by side, the proteomic and genetic data reveal a profound mismatch. Dairying in the South Caucasus intensified from at least the Middle Bronze Age onward, sustaining mobile pastoral communities, highland herders, and eventually the elite of medieval burial complexes, yet lactase persistence never became widespread in the population. The authors argue that cultural and dietary strategies, rather than genetic adaptation, were the key enablers of milk consumption. Fermentation into products such as matsun, Armenia’s traditional yoghurt, along with cheese-making and the mixing of milk with grains and other foods, reduces lactose content and allows lactose-intolerant people to benefit from dairy without discomfort. Armenia’s rich fermented dairy cuisine, which persists today, may itself be a living legacy of this ancient workaround.

The study also uncovered intriguing social dimensions. In the Late Classical period, individuals from the Odzaberd site displayed the earliest evidence of goat milk consumption in the assemblage, with some showing diversified profiles spanning sheep, goat, and cattle milk, possibly reflecting mixed herding practices or access to trade networks. In the Medieval Aghtsg burial complex, dairy proteins were detected exclusively in two individuals interred in sarcophagi, markers of elevated social status, hinting at preferential elite access to milk, although better preservation conditions inside sarcophagi cannot be ruled out. Meanwhile, the absence of horse milk proteins in the South Caucasus, despite their identification in the neighboring North Caucasus steppe, suggests that horse milking may have been a specifically steppe-based adaptation of mobile horse-reliant groups.

The researchers caution that absence of evidence is not evidence of absence: missing milk proteins may reflect sampling bias, small sample sizes, or individual dietary choices rather than a true lack of consumption. Only one Late Neolithic individual and no Chalcolithic individuals from Armenia were available for analysis, leaving the earliest chapters of the region’s dairy history open. Future work combining larger sample sets, ceramic residue analysis, and stable isotope studies of human bones and teeth promises to refine the picture. Even so, this first integrated genetic and palaeoproteomic reconstruction for a single region across such a wide temporal span delivers a powerful message: for six millennia, the people of the Armenian highlands drank their milk on their own cultural terms, and their genes never needed to catch up.

Subject of Research: Millennial-scale dairy consumption and lactase persistence in ancient Armenia

Article Title: Palaeoproteomic and genetic insights into millennial-scale dairy consumption in Armenia

Article References: Antonosyan, M., Hovhannisyan, A., Roberts, P., Cassidy, L. M., Mkrtchyan, S., Simonyan, H., Bobokhyan, A., Aghikyan, L., Piliposyan, A., Badalyan, R., Yeranyan, N., Gnuni, A., Avetisyan, H., Badalyan, M., Gabrielyan, A., Sarkisian, N., Grigoryan, A., Simonyan, H., Safaryan, M., … Wilkin, S. (2026). Palaeoproteomic and genetic insights into millennial-scale dairy consumption in Armenia. iScience, 29(11), Article 117829. https://doi.org/10.1016/j.isci.2026.117829

Image Credits: AI Generated

DOI: 10.1016/j.isci.2026.117829

Keywords: palaeoproteomics, dental calculus, dairying, lactase persistence, ancient DNA, Armenia, South Caucasus, Bronze Age, fermented dairy, matsun, beta-lactoglobulin, highland pastoralism