Treehoppers comprise roughly 3,450 species and 428 genera worldwide, and their pronotal forms have been hypothesised to serve functions ranging from crypsis and mimicry to aposematism, pheromone dispersal, and even electroreception. Beyond their architectural pronota, these insects are notable for maternal care and ant attendance, behaviours that have made them favourites of behavioural ecologists. While the New World hosts the family’s best-known representatives, the Oriental zoogeographic region is regarded as a centre of diversity for the subfamily Centrotinae, which contains approximately 1,500 species. In fact, every membracid distributed across Asia is a centrotine, with a single known exception: Stictocephala bisonia, a Nearctic native adventive in Europe and recently discovered in Asia. Understanding Singapore’s centrotines therefore contributes directly to mapping diversity in one of the group’s great evolutionary heartlands.

Singapore’s treehoppers have a distinguished but fragmented history. The first local species were collected by Alfred Russel Wallace and described by Walker in 1857, accounting for three species. The most thorough account to date came from Funkhouser in 1918, who noted new species and inventoried several others, bringing the tally to 24. Scattered contributions from Buckton, Matsumura, Distant, Goding, and Kato added a handful more, but no further work was published after the early twentieth century, leaving the exact state of the current fauna ambiguous. This gap matters because Singapore’s landscape has been transformed since 1965: only approximately 0.28 percent of the original vegetative cover, once lowland dipterocarp forest, mangroves, and freshwater swamp, remains. Vestigial habitats, secondary forest, parks, and planted streetscapes now serve as refuges for local insect diversity, making an updated inventory both timely and urgent.

The study examined 497 specimens digitally, physically, or both, with the majority, 466 specimens, drawn from the museum’s Zoological Reference Collection. These specimens derive from opportunistic collection events rather than targeted surveys, underscoring how much remains to be found. ZRC material was examined under a binocular stereo-microscope, and 31 representative specimens were photographed to anchor the new visual key. Additional type and non-type material was assessed digitally from the Smithsonian National Museum of Natural History, including 16 holotypes, and from the Natural History Museum in London, including two lectotypes. Several specimens could not be confidently assigned to any named species, either falling outside the observed intraspecific variation of well-delineated species or defying assessment against descriptions that lacked sufficient diagnostic information. These are retained as morphospecies pending future resolution.

The annotated checklist consolidates previous combinations, synonyms, type localities, recorded distributions, host plants, and ant attendants for every locally occurring species, drawing on the great catalogues of Funkhouser, Metcalf and Wade, and McKamey, with names standardised against the World Auchenorrhyncha Database. Among the new records for Singapore are Evanchon maculatum, previously known from Indonesia; Gargara luconica, from Indonesia, Malaysia, and the Philippines; Tricentrus acuticornis, from China, Indonesia, and the Philippines; T. banguensis, from India, Indonesia, and Malaysia; T. fulgidus, from China, Indonesia, Malaysia, and Vietnam; T. intermedius, from Sabah; Nilautama minutispina, from Indonesia and Malaysia; and Telingana recurvata, from Indonesia and Sarawak. Many species carry Singapore as their type locality, including Centruchus laticornis, Centrochares ridleyana, Gargara nervosa, G. projecta, G. sordida, G. triangulata, Tricentrus brunneus, T. spinicornis, T. truncaticornis, Micreune formidanda, and Ceraon orientale, several of which were described from specimens collected in 1917.

Host plant data emerged for 37 species, and the patterns are striking. More than a third of species with known hosts, 14 species spread across four tribes, were found on Melastoma malabathricum, a shrub widespread across Singapore’s forest edges and disturbed habitats. The authors suggest this strong concomitance may reflect ecological availability rather than phylogenetic constraint. Other important host taxa include Fabaceae, collectively hosting 11 species, Moraceae with 8, and Myrtaceae with 7. Most species proved polyphagous, with 22 feeding on more than one plant family. The extreme case is Leptocentrus taurus, an exceptionally polyphagous species with 72 recorded host plants across 23 families, from cashew and mango to hibiscus and tomato. The authors caution that these patterns rest on opportunistic records covering only the species for which host data exist, and that expanded surveys across a fuller representation of each tribe will be needed to contextualise them.

Ant attendance was documented for only 11 species, most notably involving the green weaver ant Oecophylla smaragdina, which tends treehoppers for the honeydew they excrete. The ecological stakes are considerable: ant attendance is known to decrease predation and, in some instances, may even be required for nymphal survival. Notably, almost all Indian treehopper species were recorded as ant attended in earlier monographic work, and if similar patterns apply throughout the Old World, ant attendance in Singapore’s fauna is likely heavily undersampled. First host plant and ant attendant records were contributed for Evanchon maculatum, Gargara luconica, and Nilautama minutispina, baseline ecological data that remain absent for many Southeast Asian species. The authors interpret all such records as preliminary, awaiting confirmation through targeted field observations, since museum labels rarely distinguish true hosts, on which nymphs develop or oviposition occurs, from incidental plant associations.

The illustrated identification key is a technical achievement in its own right. Whereas early taxonomic studies of membracids relied primarily on pronotal characters, later works recognised the diagnostic value of frontoclypeal lobes, forewing venation, and leg chaetotaxy, particularly the presence of ab- and adlateral cucullate setae on the mesothoracic femur. The new key incorporates this updated methodology and was designed with a fan-shaped structure, minimising the number of steps required for identification. It accounts for sexual dimorphism, ensuring delineations apply to both sexes, with separate steps for species such as Tricentrus acuticornis, whose males lack the suprahumerals present in females. Images of two non-Singaporean species were incorporated to visualise diagnostic genus-level characters for locally occurring species whose specimens could not be sourced. Only 40 of the 52 species are included, however, as 12 were excluded because their diagnostic characters could not be confidently assessed from available literature, making the key preliminary pending future redescriptions anchored upon type material.

Those exclusions expose a deeper crisis in Oriental treehopper taxonomy. Species identification proved challenging without access to types, because descriptions from the late 1800s to early 1900s seldom provide useful diagnostic information. The problem is most acute in the hyper-diverse genera Gargara, with 174 species, Tricentrus, with 238, and Leptocentrus, with 90, which are the largest yet most poorly studied taxa in the family. Goding’s early twentieth-century keys to the Old World Membracidae, published over 20 years, are limited by unclear terminology and were likely not based on types. Even the better-established Indian and Chinese monographs conflict: independent treatments of the same species occasionally differ, as seen in Tricentrus albomaculatus and T. gibbosulus, where workers examining type material reported contrasting features. The authors conclude that species redescriptions based on type material are sorely needed to resolve current ambiguities.

Morphology-based species counts may also be inflated, since treehoppers display intraspecific variation driven by sexual dimorphism, host-plant specificity, or geographic isolation. Yet underestimates can arise too: the classic Enchenopa binotata complex from the Neotropics, once defined as a single species by morphology, is now recognised as up to 15 species differentiated by substrate-borne mating signals and host specificity, and Oriental cryptic complexes have been disentangled through integrative approaches as well. The foremost local candidate for such investigation is Leptocentrus taurus, whose unusually wide distribution, exceptional polyphagy, pronotal variation, and contrasting sets of synonyms leave it in taxonomic limbo. Nymphal morphology offers a promising alternative, as nymphs of highly polymorphic species can be remarkably monomorphic and diagnostic, but nymphs are rarely present in collections and comparative Oriental data are non-existent for most taxa.

With only 0.28 percent of Singapore’s original vegetative cover intact, it is possible that some species recorded during the 1800s and 1900s have become locally extinct, and future targeted surveys, particularly within primary forest habitats, will help ascertain the continued presence of previously recorded taxa. The uncovering of new records suggests treehopper diversity is undersampled in adjacent Southeast Asian states as well, where broad re-examinations have not occurred since original descriptions decades ago. For now, Singapore’s 52 helmeted inhabitants, from the forked pronota of Micreune formidanda to the weaver-ant-tended colonies of Centrochares ridleyana, stand as a reminder that even in one of the world’s most urbanised nations, evolutionary artistry persists in the margins, awaiting the observers patient enough to look.

Subject of Research: The treehoppers (Hemiptera, Membracidae) of Singapore: An annotated checklist, new records, and an illustrated identification key

Article Title: The treehoppers ( Hemiptera , Membracidae ) of Singapore: An annotated checklist, new records, and an illustrated identification key

Article References: Tan, V., & Hwang, W. S. (2026). The treehoppers (Hemiptera, Membracidae) of Singapore: An annotated checklist, new records, and an illustrated identification key. ZooKeys, 1295, 161-225. https://doi.org/10.3897/zookeys.1295.204613

Image Credits: AI Generated

DOI: 10.3897/zookeys.1295.204613

Keywords: treehoppers, Hemiptera, Membracidae, Singapore, annotated, checklist, records, illustrated, identification, scientific research

The authors examined dry male and female specimens under a stereoscopic microscope and measured morphological characters with an ocular micrometer. They macerated genital segments in 10% sodium hydroxide and used a KEYENCE VHX-6000 imaging system to obtain color photographs of adult habitus and genitalia. Illustrations were imported into Adobe Photoshop CS8 for labeling and plate composition. Morphological terminology largely followed the 2011 description of the genus, while terminology for leg setal rows followed Rakitov’s 1997 system.

Scaphomonoides robustus closely resembles S. redstripeus in general appearance. Both have an ochraceous body, a yellowish head with a thin dark-brown transverse band on each side of the ocelli, an orange arcuate band between the eyes, and paired orange arcuate bands on the pronotum. The new species has a yellowish-brown face, while the second orange pronotal band is complete in S. robustus but disconnected in the middle in S. redstripeus.

The clearest differences occur in the male genitalia. In S. robustus, the aedeagus bears a pair of long, stout processes arising from the ventral margin near its base and extending beyond the shaft, while the shaft apex bears two pairs of spiniform processes. In S. redstripeus, the paired processes are moderately long and slender, the shaft tapers to an acute apex without processes, and the elongate dorsal apodeme is bifurcate distally. The new species also has a rectangular subgenital plate with 11 stout lateral setae and a valve nearly 1.9 times as broad as its median length; S. redstripeus has eight macrosetae along the subgenital plate margin and a valve nearly 1.5 times as broad as its median length.

A key to the two Chinese species separates males by the continuity of the transverse orange bands and by aedeagal structure. S. redstripeus has incomplete or interrupted bands and a slender, distally tapered aedeagal shaft with a bifurcate dorsal apodeme. S. robustus has complete bands, a shaft with spiniform apical processes and stout basal processes, and a dorsal apodeme that is not bifurcate.

The review also expands the generic redescription of Scaphomonoides. Diagnostic male genital features include an aedeagus with paired processes from the ventral margin, an elongate basal apodeme, a short subgenital plate, and a Y-shaped connective with a very short stem. Externally, the head is slightly narrower than the pronotum’s greatest width; the ocelli lie on the anterior margin, and the vertex is longer medially than beside the eye, with a distinctly angulate anterior margin. The fore femur has two dorsoapical setae, one stout anteromedial seta, 14 intercalary setae and several short basal anteroventral setae. The hind femur has an apical setal formula of 2+2+1.

The authors redescribed S. redstripeus because the original 2005 figures lacked a ventral view of the aedeagus and described the dorsal apodeme as the aedeagal shaft. New illustrations and descriptions document the male genitalia and female structures. Among the female characters, the seventh sternite has a sharply cleft, medially concave posterior margin, the first ovipositor valvulae have strigate dorsal sculpture, and the second valvulae have prominent denticles in the distal half that become smaller toward the rounded apex.

Examined specimens of S. redstripeus came from Hainan, Jiangxi and Yunnan provinces in China, with collection dates spanning 1997 to 2021. Sites included Jianfengling, Jiulianshan, Limushan and Diaoluoshan in Hainan, Jiuliansha in Jiangxi and Mengla County in Yunnan. The species is also recorded from Vietnam. Adult body length including the tegmen measured 5.07–5.44 mm in males and 5.18–5.42 mm in females. The sole known specimen of S. robustus, the male holotype from Mengxiu in Ruili, measured 5.04 mm.

The study is a specimen-based taxonomic review rather than an ecological or population-level investigation. Its assessment of S. robustus is limited to the external and male genital characters of one specimen, so the authors report the new species as known only from Yunnan and do not establish its broader distribution or natural history. The holotype is the only specimen listed for S. robustus, whereas the redescription and distribution of S. redstripeus are supported by additional male and female material from China and the previously recorded occurrence in Vietnam.

Subject of Research: Morphological taxonomy and redescription of Chinese Scaphomonoides leafhoppers

Article Title: Review of Chinese species of the leafhopper genus Scaphomonoides Li, 2011 ( Hemiptera , Cicadellidae , Deltocephalinae , Scaphoideini ), with description of a new species

Article References: Duan, K.-T., Webb, M. D., & Xing, J.-C. (2025). Review of Chinese species of the leafhopper genus Scaphomonoides Li, 2011 (Hemiptera, Cicadellidae, Deltocephalinae, Scaphoideini), with description of a new species. Deutsche Entomologische Zeitschrift, 72(2), 459-463. https://doi.org/10.3897/dez.72.157167

Image Credits: AI Generated

DOI: 10.3897/dez.72.157167

Keywords: Scaphomonoides robustus, Scaphomonoides redstripeus, leafhopper taxonomy, new species, morphological review, male genitalia, Yunnan Province, China, Hemiptera, Cicadellidae, Review, Chinese

These galls are not tumors in the usual sense. Both the strobilus and the adelgid gall are shortened shoots whose needles have been transformed into covering scales, and the transformation stays within the normal morphological variation of the plant. Crucially, the gall is produced not by the aphid's saliva alone but by the specific response of the growing spruce bud to being penetrated by that saliva, which is why monophagous soft scales feeding on the very same branches never produce galls at all.

The central insight of the gallophilous theory concerns the timing of gall opening. Spruce galls do not split open when the aphids inside need them to; they open according to the physiology of the particular tree and branch. Even trees of the same species in the same locality differ in light exposure, water and nutrient supply, age, and crown architecture, so the opening of adelgid galls stretches over months.

Around St. Petersburg in northwestern Russia, this process runs from mid-June to late August; on Hokkaido in Japan, from mid-June to early September. Galls ripen first on the lower branches and then on the upper ones, and in some cases individual cavities within a single gall open at different times, or not at all, dooming the nymphs inside to death.

Similar extended opening is documented in other gall-forming aphids, including the hickory phylloxera Phylloxera devastatrix, whose galls open from early May to mid-June, and the poplar aphid Pemphigus spyrothecae, whose galls open from late August to November and sometimes never. This prolonged, asynchronous opening, the author argues, made effective panmixia impossible for the ancestral aphids locked inside. On any given day, only a small fraction of males and females emerged from opened galls, and these tiny insects, with their relatively weak powers of purposeful flight, could not find each other in sufficient numbers to sustain random mating.

Sexually mature females still trapped in unopened galls were physically isolated from males that had already emerged. Winged females that did escape scattered across suitable food plants, and most remained unfertilized. In a situation that repeated itself every warm season for millions of years, females capable of parthenogenetic reproduction gained a decisive selective advantage: they no longer needed to synchronize their reproductive development with anyone else, and each could lay viable eggs whether she emerged early, late, or not at all.

But parthenogenesis alone could not solve the deeper problem of restoring genetic recombination. The theory proposes that the solution came through migration. The third generation of these ancestral aphids settled on secondary host plants such as larches, firs, pines, and other conifers, where sap sucking never produces closed galls and there are no physical barriers to mating.

The remaining difficulty was re-synchronizing the appearance of males and females in mass numbers at the same place. The only reliable cue available was the simultaneous end of shoot growth on conifers, which occurs by late June or early July in modern conditions. At that moment, nutrition deteriorates sharply, and winged sexuparae emerge en masse from the desynchronized parthenogenetic lines on the secondary hosts.

These sexuparae switch from thelytoky to deuterotoky, producing eggs of both sexes, and fly back to spruce, where they lay eggs on the undersides of needles. From these eggs a bisexual generation develops relatively synchronously, and mass cross-fertilization with all its genetic benefits is finally achieved, concentrated in one place at one time. This framework also explains several otherwise puzzling features of the aphid bisexual generation.

Because it always develops in the second half of the year, when feeding conditions on woody plants deteriorate, sexual individuals are small, have reduced fertility, often producing only a single egg that fills the entire abdominal cavity, and in many cases do not feed at all. The loss of wings in the sexual generation and in the early parthenogenetic generations on secondary hosts also follows logically: to achieve synchrony, both generations must stay put, the first until fertilization and the second until shoot growth ends. The cytogenetic machinery itself reinforces the cycle.

Aphid males produce only two sperm per meiosis, both carrying X chromosomes, so all offspring of fertilization are XX females. Parthenogenetic females, through a special mini-meiosis that eliminates one X chromosome set, can produce embryos with either one or two X chromosomes, generating males as well as females. The bisexual generation is therefore necessarily followed by a parthenogenetic one, a system the author suggests became so deeply entrenched over hundreds of millions of years that abandoning it became impossible even after aphids shifted to flowering plants.

The fossil and comparative evidence supports this reconstruction. The first aphid remains are known from the Triassic, or possibly the Permian, long before flowering plants became significant. The most archaic living aphids, the adelgids, are obligately tied to gymnosperms, retain a functional ovipositor, and lay eggs in every generation.

Their sister family, the phylloxeras, has lost the ovipositor and has no gymnosperm associations, which the author interprets as evidence that phylloxeras arose from an ancient adelgid ancestor only after angiosperms spread in the Cretaceous. Phylloxeras, in turn, are most diverse in North America, where they feed primarily on hickories, the only Nearctic angiosperms of the Cretaceous period known to form galls. The larger internal space of hickory galls compared with the cramped cavities of spruce cones allowed several parthenogenetic generations to develop inside before the gall opened, and made egg-laying inside the gall possible without an ovipositor, explaining the loss of that organ.

The transition to viviparity, the hallmark of the advanced superfamily Aphidoidea, is traced through the eriosomatids, which the author identifies as a probable transitional form. In these aphids, parthenogenetic offspring are occasionally born still enclosed in shells that are homologous to the egg chorion, a plesiomorphic feature pointing from ovoviviparity toward placental viviparity. Moreover, the embryos in migrating eriosomatid females develop simultaneously rather than sequentially, so a female arriving on a secondary host deposits all her offspring within a short period, much like oviparous scale insects.

Once placental viviparity combined with telescopic embryonization, in which an embryo already contains embryos of the next generation, appeared among gall-dwelling eriosomatids, aphids could accelerate generation turnover dramatically regardless of when the gall opened, freeing them from the rigid timing constraints of their spruce-dwelling ancestors. The theory also carries implications beyond aphids. Among terrestrial animals, regular cyclical parthenogenesis has been convincingly demonstrated only in gall wasps, which likewise live in closed galls, and their emergence from galls is similarly extended over time, producing comparable diversity of life cycles.

Other gall-forming hemipterans, such as scale insects, psyllids, and most true bugs, form no closed galls and show no cyclical parthenogenesis, although other forms of parthenogenesis are common among them. The gallophilous theory thus offers a unified explanation for one of the most aberrant reproductive systems in the animal kingdom: the closed gall itself, by fragmenting the mating opportunities of a population across weeks and months, created the selective pressure that first favored parthenogenesis and then locked aphids into a cycle they can no longer escape.

Subject of Research: Evolutionary origin of cyclical parthenogenesis in aphids through life in closed spruce galls

Article Title: Gallophilous theory of cyclical parthenogenesis in aphids ( Homoptera , Aphidinea )

Article References: Gavrilov-Zimin, I. A. (2024). Gallophilous theory of cyclical parthenogenesis in aphids (Homoptera, Aphidinea). Comparative Cytogenetics, 18, 247-276. https://doi.org/10.3897/compcytogen.18.136095

Image Credits: AI Generated

DOI: 10.3897/compcytogen.18.136095

Keywords: aphids, cyclical parthenogenesis, adelgids, spruce galls, phylloxeras, viviparity, telescopic embryonization, host plant alternation, cytogenetics, evolutionary biology, Hemiptera, Adelgidae

The study examined dry specimens collected by sweeping in Yunnan and Fujian provinces. Type specimens are deposited in the Institute of Entomology at Guizhou University in Guiyang. Researchers measured body length from the apex of the vertex to the tip of the forewing, measured the vertex from its apical transverse carina to the tip of its basal emargination, and expressed all measurements in millimeters. Male genital segments were removed, macerated in 10% sodium hydroxide, washed and transferred to glycerol. External morphology and drawings were examined with a Leica MZ 12.5 stereomicroscope, while habitus images were obtained with KEYENCE VHX-6000 and VHX-1000 systems.

Andes cuneatus was established from specimens collected in Mengla County and the Gaoligongshan National Nature Reserve in Yunnan. Its name refers to a wedge-shaped process on the apical part of the periandrium. Male body length ranged from 6.2 to 6.8 millimeters for four specimens, and female length ranged from 6.8 to 7.3 millimeters for four specimens. Diagnostic features include a dark-brown general color, an asymmetric anal segment with a larger right lobe, paired spinose processes near the base of the periandrium, a thick dorsal process, a bifurcated fishtail-shaped apical process and a short endosoma without a spinose process. The authors distinguish the species from A. truncatus by differences involving the proportions of the vertex, the shape of the gonostyli and the apical processes of the periandrium.

Andes luchunensis, named for Lüchun County in Yunnan, was collected in the Huanglianshan National Nature Reserve. Its general color is yellowish brown, the basal half of each forewing bears a claw-shaped marking, and the apical two-fifths to four-fifths of the gonostyli are widened. The male periandrium has two bifurcated spinose processes at the apical part of its ventral margin, while the endosoma bends dorsally and bears three apical spinose processes. Male body length was 7.2–7.5 millimeters for nine specimens, and female length was 7.5–8.6 millimeters for seven specimens. The species differs from A. pallidus in coloration, the number of bifurcated spinose processes and the structure of the apical endosoma.

Andes orbiculatus was described from Wanmulin Provincial Nature Reserve in Fujian. The name refers to a circular lamellar process extending from the proximal ventral margin of the periandrium. Other diagnostic traits include wide yellowish-brown markings and three dark-brown spots on the basal half of the forewing, a lanceolate medioventral process and a strongly asymmetrical anal segment whose left lobe extends downward. Male body length measured 5.7–6.3 millimeters for seven specimens, while female length measured 6.2–6.6 millimeters for seven specimens. The authors distinguish it from A. truncatus by the shape of the medioventral process, asymmetry of the anal tube and non-truncate gonostyli.

Andes scalpratus, collected in Xichou County in Yunnan, has an arc-shaped medioventral process and knife-shaped apical parts of the gonostyli in lateral view. Its aedeagus has three processes, and the apical quarter of the endosoma is densely covered with small teeth. Male body length was 6.7–7.3 millimeters for seven specimens, and female length was 7.8–7.9 millimeters for three specimens. Its name refers to the sharp-edged appearance of the gonostyli. The authors compare the species with A. furcutus, reporting differences in the medioventral process, gonostyli and apical endosoma, which lacks the two processes present in A. furcutus.

The paper also updates a key to the males of Andes species occurring in China. The key incorporates external features such as forewing markings, coloration and the form of the vertex and frons, alongside male terminalia including the pygofer, gonostyli, periandrium and endosoma. The four newly described species are placed among previously recognized species through combinations of these characters. The Andes genus is species-rich and contains 121 described species across the Palaearctic, Oriental, Australian and Afrotropical regions, according to the background cited by the authors.

The paper reports 37 genera and 308 species of Cixiidae recorded from China, compared with approximately 256 genera and 2,660 species worldwide. Within the Andini tribe, which the authors state contains 142 described species globally, the four additions increase the previously reported Chinese fauna by 14%. The authors interpret this increase as suggesting that the tribe may be more species-rich in China than previously suspected, although their conclusions are based on the newly described material and existing records rather than a comprehensive survey of all Chinese habitats.

Host-plant associations remain uncertain for most of the new species. Previous work had proposed that Andes species probably feed on dark, damp mosses or ferns. Adults of A. luchunensis were observed resting on a fern leaf in Huanglianshan National Nature Reserve, which the authors present as suggesting that ferns may serve as hosts for at least that species. The observation does not establish a feeding association or host relationship for the other three newly described species, and the study’s specimen-based taxonomic conclusions are geographically limited to the documented collection sites in Yunnan and Fujian.

Subject of Research: Taxonomic description of four new Andes planthopper species from Yunnan and Fujian, China

Article Title: Four new species of the planthopper genus Andes ( Hemiptera , Auchenorrhyncha , Cixiidae ) from China, with an updated key to species

Article References: Lv, S.-S., Zhi, Y., Yang, L., Long, J.-K., Chang, Z.-M., Zhang, Y.-B., & Chen, X.-S. (2026). Four new species of the planthopper genus Andes (Hemiptera, Auchenorrhyncha, Cixiidae) from China, with an updated key to species. Deutsche Entomologische Zeitschrift, 73(1), 15-29. https://doi.org/10.3897/dez.73.173015

Image Credits: AI Generated

DOI: 10.3897/dez.73.173015

Keywords: Andes planthoppers, new species, taxonomy, morphology, Hemiptera, Cixiidae, insect identification, Yunnan, Fujian, China, Four, species

Kunungua belongs to the plant bug family Miridae, one of the largest families of true bugs, and more specifically to the subfamily Bryocorinae, a principally tropical group comprising more than 1,000 species assigned to approximately 200 genera. Within this subfamily, the tribe Eccritotarsini accounts for more than 60 percent of all described bryocorine species. While the majority of eccritotarsine taxa are confined to the New World, existing data suggest an Oriental origin for the clade, with representatives spreading from the southern part of North America and the Neotropics to the Philippines, New Guinea, and northern Australia.

The researchers identified a distinctive lineage within Eccritotarsini, which they refer to as the Prodromus group of genera. This group is characterised by a unique modification of the male genitalia, specifically the aedeagus, which takes the form of a simple sclerotised tubule. In these bugs, the outer section of the aedeagus, known as the phallotheca, and the inner section, the endosoma, lack a clear demarcation. While in other bugs the endosoma is membranous and eversible, in the Prodromus complex it remains consistently everted and sclerotised. The ductus seminis in this group remains entirely membranous and terminates with a secondary gonopore that lacks ornamentation. Some members of this group also exhibit small, non-retractable membranous lobes at the apex of the aedeagus.

Beyond these genitalic features, taxa belonging to the Prodromus group typically display substylate eyes, a deeply punctate pronotum, a generally elongated body with long and comparatively thin legs, and a broad head in dorsal view. The group comprises seventeen genera, including Ambunticoris, Duducoris, Ernestinus, Frontimiris, Grossicoris, Kunungua, Myiocapsus, Namyatovia, Prodromus, Sinervus, Sinervaspartus, Spartacus, Stenopterocorisca, Stonedahlia, Stylopomiris, Taricoris, Thaumastomiris, and Zikaniola. Three genera from this group, Duducoris, Kunungua, and Prodromus, are known to inhabit the Ethiopian Region, with Duducoris and Kunungua restricted to this region while Prodromus has a more widespread distribution that includes the Oriental Region.

Kunungua was established by Carvalho in 1951 to comprise two African species with a wide head and stylate eyes, namely K. boxi from Ghana, the type species of the genus, and K. cinnamomea from Congo. Since then, two more species had been described, K. ukerewensis from Tanganyika and K. pallida from Sudan, but no new information regarding the genus had been published in nearly fifty years. The current revision redescribes all known Kunungua species and adds three new ones: K. atramentomaculata, K. gemina, and K. ornata. A notable feature for distinguishing Kunungua species is their specific colour patterns with distinct bands, while the shape of the parameres plays a crucial role in separating species that may appear superficially similar.

The revised diagnosis of Kunungua reveals a suite of characters that set it apart from related genera. The head is hammer-shaped, only 0.2 to 0.4 times as long as wide, with a longitudinally sulcate vertex and distinctly pedunculate eyes. The collar and pronotal disk are deeply punctate, with dense punctures whose spacing is smaller than the puncture diameter. The calli are prominently raised and divided by a deep longitudinal depression, while the scutellum is very small, only about 0.3 times as wide and 0.3 to 0.4 times as long as the pronotum, and strongly pointed at the apex. The membrane has a thick vein with an acute apical angle, and the membranal cell nearly reaches the apex of the cuneus. Total body length ranges from 3.5 to 4.3 millimetres, with the body elongate at 3.4 to 4.1 times as long as the width of the pronotum.

While working on the updated diagnosis, the researchers conducted a thorough morphological analysis to distinguish Kunungua from its two closest relatives, Prodromus and Duducoris. Their results confirmed that Kunungua warranted recognition as a separate genus, but revealed no distinct differences between Duducoris and Prodromus. Odhiambo had originally described Duducoris in 1962 to accommodate three newly described species from Uganda, differentiating it by a more flattened body, a non-sulcate vertex, eyes only weakly projecting above the vertex, a relatively short narrowed basal part of the antennal segment, weakly raised calli, a pronotum deeply emarginated posteriorly, and shorter legs. However, after examining specimens of various species within both genera, the researchers found that characters such as pronotal convexity, clypeal swelling, the length of the thin basal part of antennal segment I, eye shape, and the degree of posterior pronotal emargination all varied continuously across species of both genera. The species P. ibbaicus and P. kawandanus, for instance, exhibit characters traditionally considered typical for Duducoris, including weakly elevated eyes, a moderately convex frons, a distinctly swollen anteclypeus, and a flattened pronotal disk. The researchers concluded that Duducoris represents a specific form of the highly polymorphic genus Prodromus and synonymised the two, resulting in new combinations for the five species previously contained in the subsumed genus.

With the synonymy established, Prodromus is redefined to contain 31 species distributed across the Paleotropical region. Host plant associations remain poorly known for most members, though P. clypeatus and P. oculatus have been reported to feed on banana, while P. melanonotus and P. thaliae have been documented as inhabitants of Marantochloa and Thalia, members of the Marantaceae family. The only known host for a Kunungua species is Aframomum, a member of the Zingiberaceae, on which the type species K. boxi was collected in Ghana.

Within Kunungua itself, the researchers recognised two species groups based on external characters, particularly colouration. The first comprises K. boxi and K. cinnamomea, which share brown hemelytra, a fuscous membrane, a distinctly swollen clypeus and frons, an extremely short labium reaching only the procoxa, a reddish and comparatively short antennal segment I, a short eye peduncle, and a cinnamon-coloured pronotum. The second group includes the remaining species, characterised by yellowish white hemelytra with specific brownish bands, an antennal segment I that gradually darkens from pale yellow to brown, a transparent membrane, strongly pedunculate eyes, a slightly swollen frons, a labium reaching approximately the middle of the mesothorax, and a distinctly curved distal membranal vein forming a right angle. Interestingly, K. cinnamomea appears to be the only species possessing denticles on the apex of the right paramere, though the poor quality of existing illustrations for K. boxi prevents any conclusions about that species.

The three new species are distinguished primarily by their colour patterns and genitalic structures. Kunungua atramentomaculata, known from two females collected in Haut Uelé in the Democratic Republic of the Congo, bears a brown transverse band on the corium that blurs distally like an ink spot, giving the species its name. Kunungua gemina, described from multiple specimens in Upemba National Park, resembles the dark form of K. ukerewensis in general appearance but differs in having a needle-shaped apex of the right paramere, a pale yellow apically darkened antennal segment I, and a dark brown collar. Kunungua ornata, known from two males in the Congo, displays a wide oblique brown band along the cuneal fracture that gradually narrows towards the costal margin, a feature absent in all congeners. The researchers also excluded K. pallida from Kunungua, finding that this pale greenish yellow species from Sudan possesses sessile eyes, weakly raised calli, a long labium reaching the mesocoxa, and a strongly angulate membranal vein, characters atypical for the genus. With no suitable eccritotarsine genus available to receive it, K. pallida is now treated as a species of uncertain generic placement, pending the discovery of male specimens that would allow proper genitalic comparison.

Subject of Research: Taxonomy and Biodiversity

Article Title: Revision of the genus Kunungua ( Hemiptera , Heteroptera , Miridae ) with descriptions of three new species and new generic synonymy

Article References: Bolshakova, D. S., & Konstantinov, F. V. (2024). Revision of the genus Kunungua (Hemiptera, Heteroptera, Miridae) with descriptions of three new species and new generic synonymy. Deutsche Entomologische Zeitschrift, 71(1), 219-239. https://doi.org/10.3897/dez.71.122735

Image Credits: AI Generated

DOI: 10.3897/dez.71.122735

Keywords: Kunungua, Miridae, plant bugs, taxonomy, new species, Hemiptera, Heteroptera, Eccritotarsini, Prodromus, African entomology, Revision, genus

Greenideinae is a distinctive subfamily within the aphid family Aphididae, comprising roughly 180 species in 16 genera worldwide. Taxonomists currently divide the group into three tribes: Cervaphidini, Greenideini, and Schoutedeniini, with the bulk of the diversity concentrated in Greenideini. What sets these aphids apart morphologically is a combination of unusual features: their dorsal body setae are pointed or forked into two branches, and their siphunculi, the paired tubes projecting from the rear of the abdomen, are elongated and swollen in the wingless forms, bristling with remarkably long, hair-like setae. These are not the drab, featureless aphids most people picture on garden roses.

Ecologically, the subfamily is closely tied to woody plants. Most greenideine species feed on trees and shrubs in the orders Fagales, Sapindales, and Malvales, drawing sap from oaks, chestnuts, and their relatives in the beech family, as well as from sabias, myrtles, lindens, and birches. This host specialization makes them sensitive indicators of forest composition, and it also means that changes in vegetation, whether through climate shifts or land use, can reshape their distributions in ways that are only now being documented with modern survey effort.

Before this study, South Korea’s Greenideinae fauna consisted of seven species in four genera. Cervaphidini was represented by a single species, Cervaphis quercus, an odd little aphid whose pale yellow body carries ten pairs of branched, projecting processes along its margins, giving it an almost crustacean silhouette. The remaining six species belonged to Greenideini: three species of Greenidea, one species of Mollitrichosiphum, and one species of Eutrichosiphum. That inventory has now grown, and the researchers behind the discovery have paired the new record with something the field has long needed: a fully illustrated identification key covering all eight Korean species.

The new country record came from fieldwork conducted in May 2025, when five apterous, that is wingless, viviparous females were hand-collected from Castanopsis sieboldii at Samgye-ri in Jinju, at coordinates 35.1477 degrees north, 128.0448 degrees east. The specimens were preserved in 95 percent alcohol and later mounted as slide preparations in Canada balsam, following standard techniques, and deposited in the insect collection of the Institute of Agriculture and Life Science at Gyeongsang National University. Under the microscope, the aphids revealed a suite of characters that confirmed their identity as E. sinense rather than the superficially similar E. pasaniae, which was already known from Korea.

The morphological evidence is detailed and diagnostic. The apterous viviparous females of E. sinense measure 2.22 to 2.51 millimeters in body length and are pale yellowish green in life, with dark brown markings on the antennae: the tip of the third segment, the distal half of the fourth, the base of the fifth, and the processus terminalis. The five-segmented antennae run 1.21 to 1.40 millimeters long, with the third segment bearing 15 to 19 setae and measuring 1.23 to 1.34 times the width of the head across the eyes. The ultimate rostral segment, the piercing tip of the feeding mouthparts, is 0.17 to 0.19 millimeters long and carries 8 to 12 accessory setae, and it falls short of reaching the hind coxae, a key separation from E. pasaniae, whose rostrum extends past that point.

Perhaps the most striking measurements concern the siphunculi. In E. sinense these dark brown tubes are exceptionally slender and long, spanning 0.45 to 0.53 times the body length and 15.10 to 19.08 times their own basal width, clothed in many long setae and spinulose but lacking reticulation at the base. The cauda, the tail-like structure behind the siphunculi, carries 7 to 10 setae, most often nine. The legs add further confirmation: the hind coxae and femora are spinulose, the femur is reticulated distally, and the hind tibia is smooth except for spinules on the distal third. Together, these characters distinguish the species from every other greenideine aphid known in Korea.

The known distribution of E. sinense now stretches from India through China and Japan to the Indonesian island of Java, with South Korea as the latest addition. Its host plants are similarly broad across its range, including several Castanopsis oak relatives such as C. eyrei, C. ferox, C. javanica, and C. mollissima, the stone oak Lithocarpus ovalis, and a species of Pterospermum in the mallow family. In Korea, it has so far been found only on Castanopsis sieboldii, an evergreen southern species whose presence on the warmer margins of the peninsula likely explains why the aphid has gone unnoticed until now. Similar logic applies to E. pasaniae, which the same team collected on the same host plant on Jeju Island in 2023 and 2024.

The paper does more than add one species. It provides full morphological descriptions, measurements, and illustrations of the apterous viviparous females of all eight Greenideinae species now known from South Korea, along with documented host plant associations and updated distributional data. The new dichotomous key walks a user through the fauna step by step: Cervaphis quercus is immediately separable by its pale yellow body and branched marginal projections; species with five-segmented antennae and non-reticulated siphunculi split between the dark brown E. pasaniae and the yellow-green E. sinense; and among the six-segmented species, characters such as ventral abdominal spinulation, transverse ridges on the hind tibia, siphunculus length relative to body length, and whether dorsal setae are pointed or bifurcated separate Greenidea nipponica, Mollitrichosiphum luchuanum, G. kuwanai, G. nigra, and G. prunicola.

Why does a single new aphid record matter? Aphids are among the most closely monitored insect groups in temperate Asia because of their agricultural importance, yet their taxonomy still depends on careful morphological work, and country-level checklists remain incomplete. Each new record refines our picture of how these sap-feeding insects track their host plants across East Asia’s forests, and each illustrated key lowers the barrier for future surveys. The voucher specimens, slide preparations, and occurrence data published with the study, including precise coordinates and collection dates, give other researchers the raw material to verify the record and to test whether E. sinense is expanding northward, perhaps in step with a warming climate, or has simply been overlooked among the evergreen oaks of Korea’s southern coast.

Subject of Research: First record of the aphid Eutrichosiphum sinense in South Korea with an illustrated taxonomic key to Korean Greenideinae

Article Title: First record of Eutrichosiphum sinense Raychaudhuri ( Hemiptera , Aphididae , Greenideinae ) from South Korea, with an illustrated key to the Korean Greenideinae species

Article References: Kang, Y., & Lee, W. (2026). First record of Eutrichosiphum sinense Raychaudhuri (Hemiptera, Aphididae, Greenideinae) from South Korea, with an illustrated key to the Korean Greenideinae species. Biodiversity Data Journal, 14, Article e190624. https://doi.org/10.3897/bdj.14.e190624

Image Credits: AI Generated

DOI: 10.3897/BDJ.14.e190624

Keywords: aphids, Greenideinae, Eutrichosiphum sinense, Hemiptera, Aphididae, taxonomy, new record, South Korea, Castanopsis sieboldii, identification key, biodiversity, entomology

Reptalus belongs to the family Cixiidae, a lineage of planthoppers within the order Hemiptera, the true bugs. Cixiids are small, active insects that feed on plant sap through needle-like mouthparts, and their nymphs typically live in soil or leaf litter, often in association with plant roots. The genus Reptalus was established by the Russian taxonomist Emeljanov in 1971, with Cixius quinquecostatus Dufour, 1833 originally designated as its type species. Later work, informed by studies of the group’s morphology, transferred the type fixation to Flata artemisiae Becker, 1865, a change discussed in detail by specialists in 2024. Today the genus comprises four subgenera, Reptalus, Trepalus, Palvanus and Pererepa, and, with the new addition, 35 valid species distributed across the Palaearctic, Nearctic and Oriental regions.

What makes Reptalus recognisable is a distinctive combination of anatomical features. The vertex, the plate on top of the head between the eyes, is extremely broad and rectangular. The pygofer, a ring-like structure of the male genital capsule, is asymmetrical. The anal segment carries a caudal margin that is broadly and irregularly concave, and the apex of the gonostyli bears a characteristic large process on the inner side. Perhaps most telling is the aedeagus, the male intromittent organ, which carries a long process arising from a right-basal periandrial position, often accompanied by a smaller process at its base. These genital structures are the taxonomist’s primary toolkit in this group, because external appearance alone often fails to separate closely related species.

The new species, formally named Reptalus (Pererepa) inaequalis Lv, Li & Chen, sp. nov., was collected by sweeping in Shaowu City at coordinates 27°36’N, 117°49’E, with the holotype and paratypes, four additional males and eight females, gathered on 28 May 2012. All type material is deposited at the Institute of Entomology, Guizhou University in Guiyang. The species is a relatively large member of its genus: males measure 8.5 to 9.5 millimetres in total length and females 9.3 to 10.4 millimetres. Its body is black, with yellowish-brown carinae on the vertex, frons and pronotum, brown eyes, and semi-translucent forewings of light greyish brown marked with a dark brown stigma. The species name derives from the Latin word inaequalis, meaning unequal, a reference to the dorsal margin of the periandrium, which bears an unequally bifurcate spinose process at its apex.

Technically, R. inaequalis is diagnosed by a striking array of genital features. The hind tarsomere shows a chaetotaxy of 8/9 to 10, and the second tarsomere of the hind tarsi carries seven platellae, small plate-like sensilla on the foot. The pygofer is asymmetrical, with a fingerlike process at the middle of the left lobe, and both the anal segment and the gonostyli are asymmetrical, the latter bearing three lamellar processes at the apical part that curve inward. The aedeagus carries seven processes in total: the ventral margin of the periandrium bears two spinous processes, the shorter of which is shaped like a hawk’s beak and directed ventrocephalad, while the dorsal margin carries a forked process of unequal length. The endosoma, curved cephalad, bears three spinous processes at its basal ventral margin. In the female, the posterior vagina and associated structures, including the gonapophyses and the clavate gonoplac, are illustrated in detail, providing a morphological benchmark for future comparisons.

Beyond the new species itself, the review makes a significant contribution by describing and illustrating the female genitalia of Reptalus (Reptalus) arcbogdulus (Dlabola, 1965) and Reptalus (Pererepa) quadricinctus (Matsumura, 1914) for the first time. This matters because recent work on related cixiid genera such as Neocarpia, Oecleopsis and Tsauria has demonstrated that features of the posterior vaginal wall serve as key diagnostic characters for species-level identification. Pronounced sexual dimorphism has also been reported in R. iguchii, where males and females differ visibly in forewing coloration, underscoring how unreliable external appearance can be. Given how scarce female genitalic data have been for Reptalus, the new descriptions establish a reference framework that should make future systematic studies of the genus considerably more rigorous.

The review also delivers a complete checklist and an updated identification key to all seven Chinese species, a practical tool for anyone working with these insects in the field or the museum. The key separates the two subgenera present in China by whether the periandrium bears a dorsal tooth, then works through differences in forewing banding and the number, shape and position of aedeagal processes. Among the species covered, Reptalus (Reptalus) quinquecostatus boasts the widest global distribution, recorded from the United Kingdom and Spain across much of Europe and Asia to China’s Chongqing municipality, while Reptalus (Pererepa) quadricinctus is the most broadly distributed within China itself, known from fourteen provinces and municipalities stretching from Jilin in the northeast to Sichuan and Guizhou in the southwest.

The distributional data collected during the study carry their own headline. The known range of R. quadricinctus has now been extended to include Henan and Hebei provinces, confirming what the authors interpret as an ongoing range expansion within China. The seven Chinese species are spread across six zoogeographical subregions, from the Northeast and North China subregions to the Qinghai–Tibet, Southwest, Central and South subregions, suggesting that the genus has radiated into a wide variety of climates and habitats. The authors note that continued, intensified sampling is expected to extend the distribution of quadricinctus further and may well reveal additional, possibly undescribed taxa, a reminder that even relatively well-studied insect groups in China still hold surprises.

One of the most intriguing findings concerns what these insects eat. Among the documented host associations of Cixiidae, and indeed of planthoppers in general, Reptalus is the most extensively recorded genus, with larvae or adults collected from representatives of seventeen plant orders, ranging from grasses and legumes to roses, pines and nightshades. During the field surveys underpinning this review, adults of the new species R. inaequalis were collected from bamboo, the subfamily Bambusoideae within the grass family Poaceae. This indicates that bamboos may constitute an additional, previously unrecorded host resource for the species, adding a major group of plants to the genus’s already impressive host list and hinting at an ecological association worth investigating further.

For a group of insects most people will never notice, the Reptalus of China now tell a remarkably complete story. Seven species, a fully illustrated key, female genitalia described for three species, and a new bamboo-dwelling addition from Fujian together transform a poorly documented corner of the Chinese fauna into one of the better-characterized planthopper lineages in the region. The work also illustrates a broader truth of modern taxonomy: the characters that define a species often hide in fractions of a millimetre, in the forked tip of a spinose process or the proportions of a vaginal sclerite, and only patient morphological study, supported by careful fieldwork with a simple sweeping net, can bring them to light. As sampling intensifies across China’s provinces, the seven-species count for Reptalus may prove to be less a ceiling than a waypoint.

Subject of Research: Taxonomic review of the cixiid planthopper genus Reptalus in China, including the description of a new bamboo-associated species

Article Title: Review of the Chinese species of the genus Reptalus Emeljanov, 1971 ( Hemiptera , Fulgoromorpha , Cixiidae ), with description of a new species

Article References: Lv, S.-S., Li, Z.-Z., Zhi, Y., Zhang, Y.-B., Chen, X.-S., & Yang, L. (2026). Review of the Chinese species of the genus Reptalus Emeljanov, 1971 (Hemiptera, Fulgoromorpha, Cixiidae), with description of a new species. ZooKeys, 1290, 241-256. https://doi.org/10.3897/zookeys.1290.186750

Image Credits: AI Generated

DOI: 10.3897/zookeys.1290.186750

Keywords: Reptalus, Cixiidae, planthoppers, Hemiptera, new species, taxonomy, China, Fujian, female genitalia, bamboo, ZooKeys, entomology

The discovery may sound modest, involving insects that measure just a few millimeters, but it carries real weight for biodiversity science in the Caucasus. Leptopodidae, commonly called spiny-legged bugs, belong to the infraorder Leptopodomorpha, a group of predaceous bugs that typically hug shorelines and littoral zones. The infraorder contains four extant families, the most diverse being Saldidae with roughly 335 species, while Aepophilidae and Omaniidae are tiny families of highly specialized intertidal species in the Old World. Leptopodidae itself holds at least 55 extant species spread across 38 genera, making it one of the smaller but more enigmatic branches of the true bug tree.

What makes these bugs particularly curious is their lifestyle. Despite being grouped under the umbrella name shore-bugs, Leptopodidae are mainly associated with dry environments, sometimes with no traces of water nearby, and they have even been found in caves. They are small to medium-sized predatory bugs ranging from 2.5 to 8 millimeters in length, varying in shape from nearly globose and slightly flattened to elongate and parallel-sided, and often darkly colored. Their heads are short and broad without dorsal trichobothria, their eyes very large, occupying almost the entire side of the head, and their four-segmented antennae thin and distinct. In Leptopodidae, segments two and three of the four-segmented rostrum are spinose laterally, and the front legs of Palaearctic species are armed with a single row of spines, the trait that gives the family its common name.

Before this study, a single species of the family, Patapius spinosus, was believed to occur in Georgia based on distribution records compiled in the 1990s and later checklists. However, the researchers behind the new paper failed to find any exact record in published sources, suggesting the species was either extrapolated from private collections or never actually documented on Georgian territory. The new findings of Leptopus hispanus and Erianotus lanosus thus represent the first confirmed records of Leptopodidae in the country, rewriting a small but meaningful chapter in the Caucasus bug fauna.

The specimens were collected within the framework of the Caucasus Barcode of Life project, a BMBF-funded initiative aimed at building a genetic library of Caucasus biodiversity. Field teams gathered the insects by hand and sweep-net, fixed them in 96 percent ethanol, and stored them in a freezer at minus 22 degrees Celsius for later DNA barcoding at Ilia State University’s Institute of Ecology in Tbilisi. Identification was carried out using established morphological keys, and high-resolution photographs were captured with a Canon EOS 90D camera and macro lens, with image stacking software used to produce crisp composite images of the tiny specimens.

The genetic work followed standard barcoding protocols. Genomic DNA was extracted from tissue samples using a commercial miniprep kit, and partial sequences of the cytochrome oxidase subunit I gene, the workhorse barcode for animal identification, were amplified by polymerase chain reaction using a well-established primer pair. Thermal cycling conditions involved an initial denaturation step followed by two sets of cycles with gradually decreasing annealing temperatures, and the resulting amplicons were visualized on agarose gels. Sequencing was conducted in both directions at the Beijing Genomics Institute in Hong Kong, and sequence analysis was performed using Geneious Prime software. Extracted DNA was deposited in the scientific collections of Ilia State University, with aliquots destined for the LIB Biobank at Museum Koenig in Bonn, Germany, and the sequences submitted to the Barcode of Life Data System, known as BOLD.

The results are striking. For Erianotus lanosus, three specimens were collected at Gori, at an elevation of 589 meters, under rocks in steppe near the Mtkvari River in September 2021. Three barcodes were obtained, all falling within the same Barcode Index Number, BOLD:AEY2001, with a mean p-distance of just 1 percent. Remarkably, no other barcodes of this species were available in BOLD at the time of publishing, making these the first genetic records for the species worldwide. The nearest neighbor in the database was Valleriola sp. from China, with a mean p-distance of 14.6 percent, a substantial genetic gap confirming the Georgian specimens belong to a distinct lineage.

Leptopus hispanus proved even more widely distributed across the sampling sites. Two specimens were collected at Telovani at 903 meters under roadside rocks in July 2021, three more at Gori in steppe near the Mtkvari River, one at Kodistskaro at 736 meters, one at Vardzia at 1315 meters on the Mtkvari River bank, and one swept from vegetation along the Mashavera River. Five barcodes were obtained, all within BOLD:AEX9285, with a maximum p-distance of only 0.3 percent, indicating a tight genetic cluster. Here too, no barcodes of the species existed in BOLD at the time of publishing. The nearest neighbor was Leptopus marmoratus from France, held in a private status, with a mean p-distance of 13.5 percent, again a clear genetic separation.

Both species are widespread across the Palaearctic region, with distributions classified as Centralasiatic-Mediterranean, stretching from the Mediterranean to Central Asia. From neighboring countries, Erianotus lanosus had been previously reported from Armenia, Azerbaijan, and Turkey, and Leptopus hispanus from the same trio of neighbors. Given this broad distribution and their presence just across the borders, the bugs’ absence from Georgian records long puzzled entomologists. The researchers argue the oversight was not only due to the insects’ small size and secretive lifestyle, but mainly because of a lack of sufficient and targeted research on local Heteroptera, the true bugs of Georgia.

The study also casts doubt on the earlier anecdotal record of Patapius spinosus in Georgia, suggesting it was most likely an extrapolation of data on the species’ occurrence in neighboring Azerbaijan and Armenia, which together with Georgia form the South Caucasus ecoregion. Still, given the wide distribution range of Patapius spinosus and its presence in neighboring Turkey, along with another spiny-legged bug, Leptopus marmoratus, the occurrence of these species on Georgian territory remains highly likely. Further studies are needed to establish the actual distribution and population status of the newly confirmed species, which will allow a more accurate assessment of their protection status, as in some countries they are regarded as rare. For now, the DNA-assisted records stand as a vivid reminder that even well-trodden landscapes can hide species science has never formally met, and that genetic barcodes can turn a handful of specimens under riverbank rocks into a confirmed addition to a nation’s fauna.

Subject of Research: Leptopus hispanus and Erianotus lanosus: the first DNA-assisted records of Leptopodidae (Hemiptera, Heteroptera) for the fauna of Georgia

Article Title: Leptopus hispanus and Erianotus lanosus : the first DNA-assisted records of Leptopodidae ( Hemiptera , Heteroptera ) for the fauna of Georgia

Article References: Chitadze, B., Arsenashvili, E., & Bulbulashvili, N. (2023). Leptopus hispanus and Erianotus lanosus: the first DNA-assisted records of Leptopodidae (Hemiptera, Heteroptera) for the fauna of Georgia. Caucasiana, 2, 25-28. https://doi.org/10.3897/caucasiana.2.e101888

Image Credits: AI Generated

DOI: 10.3897/caucasiana.2.e101888

Keywords: Leptopus, hispanus, Erianotus, lanosus, first, DNA-assisted, records, Leptopodidae, Hemiptera, Heteroptera, fauna, Georgia

The new species belong to two families of planthoppers, the Cixiidae and the Meenoplidae, sap-sucking insects in the order Hemiptera. Three are cixiids: Cixius palmirandus from La Palma, Cixius theseus from El Hierro and Tachycixius gomerobscurus from La Gomera. The fourth, Meenoplus skotinophilus, is a meenoplid from El Hierro. The discovery of Tachycixius gomerobscurus is particularly significant, as it marks the first record of a subterranean-adapted planthopper on La Gomera, an island that lacks lava tube caves altogether because volcanism there ceased around 2.5 million years ago.

The Canary Islands are of purely volcanic origin, with the different islands emerging from the sea during the past 20.2 million years. The seven main islands differ considerably in age, size and altitude, ranging from El Hierro, barely 1.12 million years old, to Fuerteventura’s ancient terrain. This geological youthfulness matters enormously for subterranean life. Lava tubes form in low-viscosity basaltic flows known as pahoehoe, or ropy lava, and range from a few centimeters in diameter to tunnels several meters high and tens of kilometers long. Roots from surface vegetation entering these hollow underground spaces provide a food resource for animals that inhabit the caves throughout their entire life cycle.

More than 230 troglobionts — obligate subterranean arthropods — have been documented in the archipelago, yet the underground environment remains far less well investigated than surface biotopes. Planthoppers are among the insects most frequently encountered in the islands’ lava tubes. While some individuals show no external differences from surface-dwelling relatives and are likely occasional cave visitors, others display characteristic troglomorphies: reduction or loss of compound eyes, reduced and non-functional wings, and light bodily pigmentation. These traits mark them as troglobionts, spending their entire life cycle in subterranean habitats.

Cixius palmirandus, the new species from La Palma, is a case study in extreme cave adaptation. The single known specimen, a male just 3.9 millimeters long, was collected in Cueva Honda de Miranda, a labyrinthic lava tube with over 20 galleries and one kilometer of total development on the island’s eastern slope. The species is completely blind — compound eyes and ocelli are absent — with strongly reduced tegmina and vestigial wings, and a stramineous yellowish coloration. It is certainly unable to fly and is regarded as an obligate cavernicole, restricted to the subterranean environment throughout its life. The gallery where it was found lies relatively close to the cave entrance, where high relative humidity and abundant roots prevail.

Cixius theseus, from El Hierro, is similarly blind and flightless, with a male body length of 2.7 millimeters and a female of 3.3 millimeters. It was discovered not in a cave but in the mesovoid shallow substratum, or MSS — a network of voids and cracks at the interface between soil and rock layer. On El Hierro, this habitat takes a distinctly volcanic form: fields of recent lava and volcanic deposits where a layer of lava clinker is covered by fine ash, maintaining high humidity and rather constant temperature throughout the year. The species was found at 1230 meters altitude in laurel forest dominated by Morella faya and Erica canariensis, within the huge landslide of El Golfo on the island’s northwest. Its name refers to Theseus, the Greek hero and friend of Ariadne — a nod to the blind insect’s thread-like navigation of the subterranean labyrinth.

Tachycixius gomerobscurus presents a milder degree of troglomorphy. Unlike its blind relatives, it retains small but pigmented compound eyes and distinct lateral ocelli, though its hind wings are strongly reduced and vestigial. The species was collected over many years using pitfall traps installed in the colluvial MSS of Reventón Oscuro, a steep slope inside dense, humid laurel forest at 1035 meters altitude in Garajonay National Park. Nearly 40 individuals, both adults and nymphs, have been captured there, making this the most productive site for troglobitic planthoppers in the entire archipelago. The traps, PVC pipes with multiple holes and silicone covers, were baited with liver or blue cheese, though the planthoppers probably fell in by chance rather than being attracted, as they feed by sucking fluids from roots.

Meenoplus skotinophilus, the new meenoplid from El Hierro, occupies an intermediate position between surface-dwelling meenoplids and strongly troglomorphic species. Its compound eyes are small but present and red, its tegmina and wings are well developed, and its name — combining the Greek words for darkness and friend — reflects its cave-dwelling habits. The species was discovered in Cueva de Guinea, a barely 25-meter-long lava tube with a small roof entrance in the lowland area of Frontera. The cave’s high humidity and many roots attached to walls and hanging from the ceiling create a good environment for invertebrate communities. The researchers also obtained mitochondrial COI barcode sequences for three individuals, which proved identical to each other and showed no matches above 85 percent similarity in genetic reference databases, confirming the genetic novelty of the lineage.

With seventeen documented species of strictly hypogean planthoppers, the Canary Islands now hold the highest number of subterranean planthoppers of any region worldwide. Strikingly, twelve of the seventeen species occur on the two youngest islands, La Palma and El Hierro, where volcanic activity prevails and extended areas of suitable cave habitat exist. On La Palma, the most recent eruption in 2021 along the Cumbre Vieja ridge created the new volcano Tajogaite and certainly many new lava tubes. The researchers hypothesize that in the Cixiidae at least five to six, and maximally twelve, successful underground colonization events have taken place, while in the Meenoplidae at least two lineages must have colonized the islands despite no epigean meenoplid species having ever been reported from the archipelago.

The zoogeographic patterns observed carry a sobering evolutionary message. Most troglobitic planthoppers on the Canary Islands have no extant close epigean relatives and are therefore relicts — persistent remnants of formerly widespread faunas. The researchers conclude that the cavernicolous Meenoplidae of Gran Canaria, La Palma and El Hierro, together with several cavernicolous Cixiidae, are indicative of an ancient surface fauna that is now extinct. This pattern recurs across many other arthropod groups in the islands, which are represented only by subterranean-adapted species. All cave planthopper species of the Canary Islands are single-island endemics, known from few locations and few specimens each, and all must be regarded as vulnerable, if not endangered, under IUCN criteria.

Conservation concerns are immediate and tangible. Cueva Honda de Miranda, home of Cixius palmirandus, lies in an area heavily transformed into a rural environment with scattered homes and agricultural fields. The sewage sanitation system is non-existent, and wastewater is discharged directly into underground wells, gradually contaminating the subsurface. The detection of American cockroach nymphs in the cave signals local contamination and the potential displacement of native subterranean fauna by invasive species. At Cueva de Guinea, home of Meenoplus skotinophilus, significant tourist activity and facilities equipped with bathrooms lacking sewage connections pose similar threats. Meanwhile, at Cueva de la Curva on El Hierro, breakage of the end of the lava tube by loader machines working on the surface has caused the stable, saturated atmosphere essential for obligate troglobionts to collapse, rendering three species formerly exclusive to that cave very scarce.

Climate change compounds these pressures. Increasing periods of extreme drought have occurred over the past several years, and the stable, saturated atmosphere essential for obligate terrestrial troglobionts is no longer maintained in larger cave passages. Cave planthoppers may cope with short-term adverse conditions by retreating into smaller crevices of the MSS, which are less prone to desiccation, but long-term climate change will have lasting indirect and direct influence on the subterranean environment. Increasing temperatures and lower precipitation will alter surface plant communities, and plant species providing adequate roots for food supply may vanish. Sporadic observations indicate that cave planthoppers may be temperature sensitive, suggesting that rising average annual temperatures could result in their extirpation.

Despite these threats, the outlook for discovery remains bright. The researchers note indications — observations and images without voucher specimens — that more new species await discovery, and it is conceivable that the number of obligate subterranean planthopper species will eventually double. Particularly promising are caves in the north of La Palma, the El Golfo region on El Hierro, and the extended MSS on Tenerife, Gran Canaria and La Gomera. Conversely, the eastern islands of Fuerteventura and Lanzarote are much drier, their caves lack roots inside, and planthoppers have never been found there. Troglobitic planthoppers offer unique opportunities to study adaptation to specialized habitats, including physiology, behaviour and the evolutionary dynamics of subterranean speciation — and they could serve as models for understanding climate change impact on subterranean environments worldwide.

Subject of Research: Discovery of four new cavernicolous planthopper species in the Canary Islands

Article Title: Endless forms most wonderful: Four new cavernicolous planthopper species ( Hemiptera , Fulgoromorpha , Cixiidae and Meenoplidae ) from the Canary Islands

Article References: Hoch, H., López, H., Naranjo, M., Aguín-Pombo, D., & Oromí, P. (2025). Endless forms most wonderful: Four new cavernicolous planthopper species (Hemiptera, Fulgoromorpha, Cixiidae and Meenoplidae) from the Canary Islands. Subterranean Biology, 51, 61-101. https://doi.org/10.3897/subtbiol.51.144111

Image Credits: AI Generated

DOI: 10.3897/subtbiol.51.144111

Keywords: Canary Islands, planthoppers, cave fauna, troglobionts, Cixiidae, Meenoplidae, lava tubes, endemic species, subterranean biology, conservation, Hemiptera, biodiversity

The research team, led by biogeographers working with a time-calibrated phylogeny built from more than 2,500 ultraconserved elements and calibrated with Dominican amber fossils, estimated that the most recent common ancestor of the main kissing bug tribes arose around 59.2 million years ago. That date is older than many previous estimates, which had placed the origin of the subfamily between 45 and 30 million years ago, and it suggests that the initial diversification of Triatominae began significantly earlier than previously recognized, potentially preceding the major Andean uplift that reshaped the South American landscape.

To reconstruct where the group came from, the researchers fitted a dispersal-extinction-cladogenesis model to a presence-absence matrix of 96 species mapped across 11 bioregions derived from a network-based regionalization of the Americas. The model indicated that the ancestral range of the entire clade encompassed multiple areas, with northern South America and parts of Mesoamerica emerging as the most likely, albeit with small probability. This pattern points to a widespread or complex ancestral distribution rather than a single narrowly defined point of origin, though the classical hypothesis that the group arose in Central or northern South America remains the most plausible.

The team then simulated dispersal histories using 100 biogeographic stochastic mappings, generating 6,591 dispersal events whose timing could be compared across replicates. Because the number of branches in a phylogeny increases toward the present, raw event counts naturally rise as more lineages become available to disperse. To correct for this inherent bias, the researchers normalized dispersal frequency by the number of lineages through time, revealing a pattern that raw counts had obscured: a pronounced decline from roughly 60 to 35 million years ago, followed by a gradual recuperation toward the present.

A threshold regression model identified two significant breakpoints at 50 and 30 million years ago, delineating three phases with distinct linear trends. Phase A, spanning 60 to 50 million years ago, showed high normalized dispersal, with lineages moving from the Amazonian region toward northern South America and from northern South America into Mesoamerica, as well as from the Caatinga toward the Cerrado. Phase B, from 50 to 30 million years ago, exhibited a substantial decline and stabilization in dispersal activity, with movements concentrated among Mesoamerica and northern South America and routes reaching the Páramos, Caatinga, and Chaco. Phase C, from 30 million years ago to the present, marked a resurgence in dispersal with the most complex and frequent patterns of the entire reconstruction.

The association between these phases and climate is striking. Phase A coincided with the Paleocene-Eocene transition, a period of significant global warming that included the Paleocene-Eocene Thermal Maximum, and the higher dispersal rate during this interval may have been facilitated by warmer conditions that expanded habitat connectivity and suitability, particularly from tropical source regions like Amazonia. Phase B aligned with global cooling and the transition into the Oligocene, when temperatures dropped sharply. The researchers note that previous analyses of global plant range shifts revealed an approximately 30 percent decrease in movements into the world’s tropics during the Early Eocene Climatic Optimum around 52 million years ago, suggesting large-scale vegetational changes that could have influenced dispersal dynamics. Cooler temperatures might have reduced habitat suitability, increased environmental filtering, or restricted connectivity between regions, thereby limiting dispersal success for the kissing bug lineages.

Phase C encompasses the Miocene, Pliocene, and Pleistocene, a period of fluctuating but generally recovering temperatures compared to the Oligocene minimum, leading into the Quaternary glacial cycles. During this phase, a high frequency of bidirectional dispersals was observed between Mesoamerica and the Nearctic region, likely facilitated by the formation and stabilization of the Isthmus of Panama. Routes from Amazonia extended toward the Páramos and the Caatinga, indicating the continued influence of Amazonia as a source or transit area. Within the Atlantic biomes, a dynamic and reciprocal exchange between the Caatinga and the Atlantic Forest was evident, and dispersal routes following the South American dry diagonal, from Caatinga through Cerrado, Chaco, and Pampas, highlight the importance of drier biomes as corridors for movement. Routes from the Pampas toward both the Atlantic Forest and the Amazon Forest suggest more recent expansions into these areas.

A non-parametric statistical comparison of surface paleotemperatures grouped by dispersal phase revealed significant differences among all three phases, supporting the exploratory association between climate fluctuations and Triatominae dispersal dynamics. The researchers caution that this comparison is exploratory, but the alignment of the biphasic dispersal pattern with major paleotemperature shifts implies a potential influence of climatic change on the evolutionary dispersal processes of this medically important insect group. Individual routes also showed varying temporal peaks, with the bidirectional exchange between Mesoamerica and the Nearctic displaying distinct modes of activity, one peaking around 13 to 15 million years ago and another more recent.

The findings carry epidemiological weight because nearly all Triatominae species serve as vectors for Trypanosoma cruzi, the protozoan that causes Chagas disease, and understanding the historical processes that shaped their distribution may inform how these vectors respond to future environmental change. The study drew on a robust dataset of 18,438 unique occurrence records from the DataTri database, and the regionalization revealed that 54 percent of species were restricted to a single bioregion, underscoring how strongly dispersal history continues to define contemporary patterns of kissing bug diversity across the Americas. By quantifying the timing and directionality of lineage dispersals, the researchers have provided a macroevolutionary framework that connects deep-time biogeography with the present-day geography of one of the most consequential insect vectors in the Neotropics.

Subject of Research: Historical biogeography and dispersal timing of Triatominae (kissing bugs) across the Neotropics

Article Title: Timing and directionality of Triatominae ( Hemiptera , Reduviidae ) lineage dispersals in the Neotropics

Article References: Cardozo, M. D. L. B., Rabinovich, J. E., Lepeco, A., & Löwenberg-Neto, P. (2025). Timing and directionality of Triatominae (Hemiptera, Reduviidae) lineage dispersals in the Neotropics. Frontiers of Biogeography, 18, Article e132274. https://doi.org/10.21425/fob.18.132274

Image Credits: AI Generated

DOI: 10.21425/fob.18.132274

Keywords: Timing, directionality, Triatominae, Hemiptera, Reduviidae, lineage, dispersals, Neotropics, scientific research