Integrative Taxonomy Research Articles

A new species of yellow acorn ant discovered in Italy via integrative taxonomy (Temnothorax luteus-complex, Formicidae)

The Mediterranean, a global hotspot for rare ant species, hosts a significant representation of the global diversity of the mainly Holarctic ant genus Temnothorax. However, several groups still require significant taxonomic efforts. The taxonomy of the T. luteus complex species was revised in 2014 when morphometrics allowed distinguishing two valid species and two synonyms out of four taxa that had been originally described from France. The two species recognized since then are T. luteus, distributed from Iberia to the Alps, and the largely sympatric but much more xerothermophilic T. racovitzai. In Italy, only a few records of the complex were ever published, and the identity of the Italian population was never thoroughly assessed. We combined morphometrics with phylogenomic data to assess the identity of the T. luteus populations that spanned from Sicily to the Italian Alps and discovered that all Italian samples belong to a new cryptic species, which we describe as T. apenninicussp. nov. whose glacial refugium was probably in the southern Apennines.

Intergrative Taxonomic Study of the Frullania parvistipula Complex with a Modern Circumscription of the Section Trachycolea (Frullaniaceae, Marchantiphyta).

Frullania (subg. Trachycolea) sect. Trachycolea has been studied using integrative taxonomy methods and utilizing sampling from almost all areas of distribution of the species previously referred to this section. A phylogenetic analysis based on nuclear ribosomal ITS1-2 and chloroplast trnL-F sequence data and a morphological study reveal a wide range of morphological variability within specimens that has largely disguised the overall taxonomic diversity. Frullania parvistipula, previously regarded as a widespread species, has been found to represent a group of separate species within different sections of F. subg. Trachycolea: F. caucasica and F. conistipula in F. sect. Trachycolea, F. parvistipula in F. sect. Australes, and F. fukuzawana in F. sect. Integristipulae II. Illustrations of the type specimens of F. conistipula, F. fukuzawana, and F. parvistipula, as well as illustrations of the sequenced specimens belonging to two of the discussed species (F. conistipula and F. parvistipula), are provided. The morphological differences separating the highly similar F. caucasica, F. conistipula, F. fukuzawana, F. koponenii, and F. parvistipula are discussed. A dichotomous key is presented for accepted species. New combinations are provided for two taxa.

Major specialized natural products from the endangered plant Heptacodium miconioides, potential medicinal uses and insights into its longstanding unresolved systematic classification

A comprehensive phytochemical investigation of the flower buds and leaves/twigs of Heptacodium miconioides, a cultivated ornamental plant native to China and categorized as ‘vulnerable’, has led to the isolation of 45 structurally diverse compounds, which comprise 18 phenylpropanoids (1–4, 7–20), 11 pentacyclic triterpenoids (5, 6, 21–29), eight secoiridoid glycosides (30–37), three quinic acid derivatives (38–40), and a few miscellaneous components (41–45). Among them, (+)-α-intermedianol (1), (+)-holophyllol A (2), and (−)-pseudolarkaemin A (3) represent previously unreported enantiomeric lignans, while (+)-7′(R)-hydroxymatairesinol (4) is an undescribed naturally occurring lignan. Heptacoacids A (5) and B (6) are undescribed 24-nor-urs-28-oic acid derivatives. Their chemical structures were determined by 2D-NMR, supplemented by evidence from specific rotations and circular dichroism spectra. Given the uncertainty surrounding the systematic position of Heptacodium, integrative taxonomy (ITA), a method utilized to define contentious species, is applied. Chemotaxonomy, a vital aspect of ITA, becomes significant. By employing hierarchical clustering analysis (HCA) and syntenic pattern analysis methods, a taxonomic examination based on the major specialized natural products from the flower buds of H. miconioides and two other Caprifoliaceae plants (i.e., Lonicera japonica and Abelia × grandiflora) could offer enhanced understanding of the systematic placement of Heptacodium. Additionally, compounds 39 and 40 displayed remarkable inhibitory activities against ATP-citrate lyase (ACL), with IC50 values of 0.11 and 1.10 μM, respectively. In summary, the discovery of medical properties and refining systematic classification can establish a sturdy groundwork for conservation efforts aimed at mitigating species diversity loss while addressing human diseases.

Phylogenetic systematics and integrative taxonomy contribute to weed biological control

Taxonomy and systematics are at the base of any biological research, providing the scientific names and evolutionary context to access the biological information needed. The biological control of weeds (BCW), like many other fields, often suffers from the “taxonomic impediment” generated by incomplete taxonomic knowledge or lack of available taxonomic experts on the particular taxa involved (i.e., weed plants and their potential biocontrol agents). Further, most groups with described species require revision by specialists using modern taxonomic methods. Systematists in the different taxonomic groups have the knowledge and experience to perform the accurate identification of the species involved in the study, to recognize if they are non-native species, or if they are new to science. Their expertise is crucial for identifying the organisms involved in BCW, including hybrids or cryptic species. The practice of phylogenetic systematics allows the discovery of evolutionary relationships and natural taxa that are valuable in making host range predictions. This contribution emphasizes the crucial role played by phylogenetic systematics and integrative taxonomy in weed biological control.

Integrative taxonomy reveals inflated biodiversity in the European Temnothorax unifasciatus complex (Hymenoptera: Formicidae)

AbstractTemnothorax unifasciatus (Latreille, 1798) is a widely distributed pan‐European species from the Iberian Peninsula to the Caucasus. This taxon's relatively high morphological variability prompts the taxonomists of earlier times and today to mention the morphologically different elements at specific or subspecific ranks. This paper aims to understand the population structure and genetic diversity within this lineage via integrative taxonomy, incorporating molecular phylogenetics, species delimitation analyses and multivariate analyses of continuous morphometric data from across the geographic range of the T. unifasciatus complex. Phylogenetic analyses yielded incongruent trees. The genealogical diversity index (gdi) and the confirmatory analyses on morphological data found only weak, ambiguous delimitations within the unifasciatus complex. The most highly supported scenario splits T. brackoi from the remaining unifasciatus complex with ambiguous support (gdi = 0.56). This scenario is supported by multivariate morphometry with 100% accuracy in classification success. Instead, our results suggest complex morphological and genetic population structuring within the broad range of T. unifasciatus. Therefore, we confirm the validity of two species, T. brackoi Salata & Borowiec, 2019 and T. unifasciatus (Latreille, 1798), and propose five new junior synonymies, T. cordieri (Bondroit, 1918) syn. nov., T. tauricus (Ruzsky, 1902) syn. nov., T. berlandi (Bondroit, 1918) syn. nov., T. unifasciatus staegeri (Bondroit, 1918) syn. nov., T. tuberum ciscaucasicus (Arnol'di, 1977) syn. nov. with the latter. To achieve maximal taxonomic stability, we designated a lectotype for Temnothorax unifasciatus (Latreille, 1798).

Size does matter: integrative taxonomy and size evolution of threatened charopid land snails on Lord Howe Island (Gastropoda: Stylommatophora)

Size does matter: integrative taxonomy and size evolution of threatened charopid land snails on Lord Howe Island (Gastropoda: Stylommatophora)

Delimitation of Ophioderma teres (Lyman, 1860) and Ophioderma unicolor H.L. Clark, 1940 stat. nov. (Echinodermata: Ophiuroidea), including the description of two new species

Ophioderma teres (Lyman, 1860), an ophiuroid previously believed to have a wide distribution in the eastern Pacific, has been found to have an unclear taxonomic identity. While considered a well-known species, recent studies have revealed that O. teres lacks a holotype and has vague boundaries with its congeners Ophioderma teres unicolor H.L. Clark, 1940 and Ophioderma sodipallaresi Caso, 1986, as well as with two additional new morphotypes detected in Mexico and Nicaragua, causing continuous misidentifications. This study utilized an integrative taxonomy approach based on morphologic, morphometric, and molecular evidence to clarify the taxonomic status of O. teres, O. sodipallaresi, O. teres unicolor, and the two new morphotypes. Data integration led to the following results: 1) the neotype designation and redescription of O. teres; 2) the proposal of O. sodipallaresi as a junior synonym of O. teres; 3) the status change of O. unicolor stat. nov. from subspecies to species, and 4) the description of the morphotypes as the new species Ophioderma aija sp. nov. and Ophioderma bichi sp. nov. An identification key to the eastern Pacific species of Ophioderma was also developed. This work contributes to the knowledge of Ophioderma in the region, increasing the number of described species and providing resources for their accurate identification.

Isopods on isopods: integrative taxonomy of Cabiropidae (Isopoda: Epicaridea: Cryptoniscoidea) parasitic on anthuroid isopods, with descriptions of a new genus and three new species from Japan.

Species in the parasitic isopod family Cabiropidae are known to utilise various isopods as hosts but there are currently no records of members parasitising anthuroid hosts. We describe Anthuroniscus gen. nov. for three new cabiropid species, Anthuroniscus shimomurai sp. nov. , Anthuroniscus dentatus sp. nov. and Anthuroniscus latus sp. nov. , all of which are parasitic on anthuroid isopods. Anthuroniscus gen. nov. differs from the other 14 cabiropid genera and 10 genera treated as family incertae sedis in females having an elongate, dorsally compressed, posteriorly tapering body with six pairs of lateral bulges; and cryptoniscus larvae in the following combination of characters: (1) eyes lacking, (2) antennular article 1 with eight teeth on the posterior margin, (3) uropodal exopod and endopod rectangular rather than tapering, and endopod longer than exopod, and (4) pleotelson trapezoidal, 2× as wide as long. Anthuroniscus shimomurai sp. nov. was parasitic on Mesanthura sp. from Kaichu Doro, Uruma, Okinawa, south-western Japan; A. dentatus sp. nov. on Accalathura sp. from Irabu Island, Miyako Islands, Okinawa; and A. latus sp. nov. on Colanthura nigra from Kanagawa, central Japan. In pairwise comparisons, the three new species showed p -distances of 0.6-1.3% for the 18S rRNA gene (1440 positions); and A. shimomurai sp. nov. and A. latus sp. nov. showed a p -distance of 36.2% for the 16S rRNA gene (412 positions). In an 18S -based maximum-likelihood tree, an Anthuroniscus gen. nov. clade was the sister group to Cryptoniscoidea sp., parasitic on an ostracod species. This is the first study reporting Cabiropidae from Japan and anthuroids as hosts for Cryptoniscoidea. ZooBank: urn:lsid:zoobank.org:pub:2EE042E2-AE48-4B87-B495-8436462146B9.

Integrative taxonomy of the genus Coridius Illiger, 1807 (Hemiptera: Heteroptera: Dinidoridae) reveals hidden diversity and three new species from North-East India.

The genus Coridius Illiger, 1807 (Heteroptera: Dinidoridae) comprises a group of phytophagous terrestrial bugs consisting of 36 species distributed in the Afrotropical and Indo-Malayan regions. In several communities in northeastern India, insects are recognised as a delicacy, medicine, and a nutritional supplement, with Coridius being a popular delicacy. However, Coridius has received little taxonomic attention to date due to large intraspecific variations, inadequate taxonomic treatments, and the rarity of many species. To address this gap, an integrative taxonomy of the genus was performed. Two mitochondrial genes, viz., cytochrome oxidase subunit 1 (COI) and 16S rRNA, were sequenced to reconstruct the phylogenetic relationships within Coridius. We performed both maximum likelihood (ML) and Bayesian inference (BI) to develop a species tree, followed by the Bayesian implementation of the Poisson tree process (bPTP) and Assemble Species by Automatic Partitioning (ASAP) as an additional test to assess species boundaries and delimit operational taxonomic units. A linear discriminant analysis (LDA) of four key morphological characters was then performed to identify species groups. Overall, our analysis supported the establishment of three new species: Coridius adii sp. nov., Coridius esculentus sp. nov., and Coridius insperatus sp. nov., and revealed six distinct lineages within Coridius chinensis (Dallas, 1851). Linear discriminant analysis of morphological characters indicated the clustering of eight species. The species status of Coridius nigriventris (Westwood, 1837) stat. rev, formerly synonymized under Coridius nepalensis (Westwood, 1837), is reinstated in this study. Further, we revised the genus Coridius from India and rediscovered Coridius assamensis (Distant, 1902) and Coridius fuscus (Westwood, 1837) after 100 years.

New bathyal amphipod species (Amphipoda: Eusiridae: Rhachotropis) from southwestern Pacific through integrative taxonomy

New bathyal amphipod species (Amphipoda: Eusiridae: Rhachotropis) from southwestern Pacific through integrative taxonomy

Integrative taxonomy reveals two new species of karst-dwelling Hemiphyllodactylus Bleeker, 1860 (Squamata: Gekkonidae) from the border region of Laos and Vietnam

An integrative taxonomic analysis of the gekkonid lizard genus Hemiphyllodactylus was conducted using molecular, morphological, color pattern, and ecological data on a clade of 19 species, including two new species from karst landscapes in northern Laos and Vietnam. Hemiphyllodactylus houaphanensis sp. nov. (Laos) and H. vanhoensis sp. nov. (Vietnam) are karst-dwelling species whose discovery further emphasizes the unrealized herpetological diversity and site-specific endemism in karst ecosystems across Southeast Asia and the need for increased field work to protect these biodiverse landscapes.

A treasure trove of endemics: two new species of snake-eyed skinks of the genus Panaspis Cope, 1868 (Squamata, Scincidae) from the Serra da Neve Inselberg, southwestern Angola

Four species of the genus Panaspis – P. cabindae, P. wahlbergi, P. maculicollis and P. mocamedensis – are currently known from Angola. The analysis of recently collected specimens from Serra da Neve Inselberg, an isolated mountain located in northern Namibe Province, revealed unexpected taxonomic diversity in the group. Using an integrative taxonomy approach based on morphological and DNA sequence data, with both mitochondrial (16S) and nuclear (RAG-1) genes, we were able to distinguish two distinct populations, described here as two new species, Panaspis ericaesp. nov. and P. mundavambosp. nov. Both species are assumed to be endemic to the inselberg. This reinforces our notion of southwestern Angola as a hotspot of skink diversity, and highlights the urgent need for the conservation of Serra da Neve.

Camellia neriifolia and Camellia ilicifolia (Theaceae) as separate species: evidence from morphology, anatomy, palynology, molecular systematics

BackgroundThe systematic status of sect. Tuberculata and its taxonomy have recently attracted considerable attention. However, the different bases for defining the characteristics of sect. Tuberculata has led to many disagreements among the plants in this group. Camellia neriifolia and Camellia ilicifolia have been the subject of taxonomic controversy and have been treated as different species or varieties of the same species. Therefore, it is important to use multiple methods, i.e., integrative taxonomy, to determine the taxonomic status of C. neriifolia and C. ilicifolia. This is the first study to systematically explore the taxonomic position of these two plants on the basis of Morphology, Anatomy, Palynology and Molecular Systematics.ResultsExtensive specimen reviews and field surveys showed that many differences exist in C. neriifolia and C. ilicifolia, such as the number of trunk (heavily debarked vs. slightly peeling), leaf type (smooth thin leathery, shiny vs. smooth leathery, obscure or slightly shiny), leaf margin (entire vs. serrate), flower type (subsessile vs. sessile), number of styles (3–4 vs. 3), and sepal (ovate vs. round). Moreover, C. neriifolia has a more distinctive faint yellow flower color, and trunk molting was more severe in C. neriifolia than that in C. ilicifolia. In addition, micromorphological analysis of the leaf epidermis showed that the two species differed in the anticlinal wall, stomatal apparatus, and stomatal cluster, and pollen morphology analyses based on pollen size, germination furrow, and polar and equatorial axes showed that they are both distinct from each other. The results of the phylogenetic tree constructed based on the whole chloroplast genome, protein-coding genes, and ITS2 showed that both C. ilicifolia and C. neriifolia were clustered in different branches and gained high support.ConclusionsThe results combine morphology, anatomy, palynology, and molecular systematics to treat both C. neriifolia and C. ilicifolia as separate species in the sect. Tuberculata, and the species names continue to be used as they were previously. In conclusion, clarifying the taxonomic status of C. neriifolia and C. ilicifolia deepens our understanding of the systematic classification of sect. Tuberculata.

DNA barcoding and phylogenetic relationships of ecologically and commercially important seaweed species from the Azores (NE Atlantic)

Marine natural resources are increasingly demanded to support the livelihoods and wellbeing of humans facing multiple and simultaneous crises across the planet. Seaweeds, in particular, are renowned sources of bioactive marine compounds used for various purposes, including human and animal nutrition, soil fertilization, and healthcare markets. DNA-based tools are being adopted in integrative taxonomy and ecological research, as well as to monitor the supply chain of commercially important species. This research provides novel insights into the population genetics, phylogenetic relationships, and haplotype network of six commercially significant seaweed species that are abundant in the Azores archipelago and currently traded for industrial applications. This study provides a detailed assessment of the genetic diversity of the red seaweeds Asparagopsis armata, Asparagopsis taxiformis, and Pterocladiella capillacea, and the brown seaweeds Halopteris scoparia, Zonaria tournefortii, and Cystoseira pustulata, contributing with 17 newly generated barcodes of COI, rbcL, and 28S rDNA gene records from the Azores region to the reference databases. Understanding the species identification and population dynamics is critical for understanding species composition in biodiversity hotspots. This knowledge supports taxonomy, conservation, environmental protection, and the sustainable use of marine resources.

An integrative taxonomy of some close related Palaearctic Macromia spp. (Anisoptera: Macromiidae)

An integrative taxonomy combining morphological and molecular data confirmed that both Macromia amphigena Selys, 1871 and M. manchurica Asahina, 1964 have been assigned to separate species hypotheses, although there is hybridization between the members of these taxa. Scientific names Macromia clio Ris, 1916 and Macromia kubokaiya Asahina, 1964 are shown to be junior synonyms of M. amphigena. Macromia amphigena exhibits a large range of intraspecies morphological variation within its wide distribution, which covers most of East Asia. Macromia berlandi Lieftinck, 1941 is treated as a junior synonym of M. cupricincta Fraser, 1924 based on previous research and the present study. Some traditional diagnostic morphological characters are deemed not suitable for species diagnosis. Integrative taxonomy can help researchers to filter out amphibolous morphological features and find more credible characters for future taxonomy.

Integrative taxonomy reveals unanticipated hidden diversity in the monotypic goosefish genus Lophiomus (Teleostei, Lophiidae), with description of three new species and resurrection of Chirolophius laticeps Ogilby, 1910

Thought to be monotypic for decades, the only species in the goosefish genus Lophiomus Gill, Lm. setigerus (Vahl), shows a wide range of morphological variation and is distributed widely in the Indo-West Pacific (IWP). In this study, datasets for two mitochondrial and two nuclear genes sequences obtained from samples of Lophiomus collected in different localities across the IWP were constructed and analyzed to explore the phylogeny and species diversity within the genus. Our integrated approach with multiline evidence unveiled an unanticipated richness of at least six delimited species of Lophiomus. Herein, based on materials already available from museums and new specimens obtained primarily through the Tropical Deep-Sea Benthos program surveying IWP benthic fauna, we formally describe three new species: Lm. immaculioralis sp. nov., Lm. nigriventris sp. nov., and Lm. carusoi sp. nov. Also, we resurrect Lm. laticeps stat. rev. from synonyms of Lm. setigerus. These species can be diagnosed by genetics, body coloration, patterns on the floor of the mouth, peritoneum pigmentation, morphometric measurements, and meristic counts of cranial spines, dorsal-fin spines, and pectoral-fin and pelvic-fin rays from each other and from Lm. setigerus. The species Lm. setigerus, as well as the genus Lophiomus, are re-described accordingly based on the new results. Amended identification keys to the four extant lophiid genera and to species of Lophiomus are also provided.

Tolypella mongolica (Charophyceae, Characeae): A new dioecious charophyte species from the center of Eurasia and implications for charophyte taxonomy, including the validation of T. porteri and the new combination for T. jwalae

A new dioecious species of Tolypella, T. mongolica sp. nov. from the section Tolypella, was described using an integrative taxonomy. It was found in three distant localities – shallows of a brackish small lake and a channel in the midland steppe in Mongolia and the Altai Mountains, and a small brackish water body in the dry steppe in Southeast Europe. It differs from other species in rbcL and ITS sequences and in distinct complex ornamentation of the oospore surface, varying from sparsely coarsely tuberculate to having the appearance of an aerial view of mountain landscape with acute to mostly sharp edges of elements at probably the ripest stage. Wide intrapopulation variability of expression of this trait, only partly explainable by oospore ripening stage, was found and illustrated for two populations, indicating previous underestimation of this phenomenon in taxonomic research. T. mongolica is richly fertile and appears to be ephemeral as most species of this genus. Only female plants were found in all localities. The absence of male plants could be referred to either their earlier development during the growing season or, less likely, parthenogenesis in this species. The description of the new species suggests taxonomic changes for two dioecious species of charophytes. Tolypella porteri stat. et comb. nov. was validated here. T. jwalae was recognized as a species of Nitella, N. jwalae comb. nov.

Integrative Taxonomy Suggests Resurrection of Species of the Astyanax bimaculatus Group (Characiformes, Characidae).

Using integrative tools can be effective for species identification, especially in complex groups like Astyanax. Astyanax bimaculatus group is composed of six valid species, including A. lacustris. "A. altiparanae", "A. asuncionensis", and "A. jacuhiensis" are considered as junior synonyms of A. lacustris. Seeking to test the operational taxonomic unit (OTU) status of the junior synonyms of A. lacustris ("A. altiparanae", "A. asuncionensis", and "A. jacuhiensis"), we used analyses through mitochondrial DNA (COI and Cytb), cytogenetic markers (classical and molecular), and morphometry ("truss network"). Analysis of mitochondrial DNA sequences separated A. lacustris from the other synonymized species. The cytogenetic and morphometric analyses did not corroborate the synonymization and suggest that besides A. lacustris, the OTUs A. altiparanae, A. asuncionensis, and A. jacuhiensis are valid species. The analysis of different characters proposed by the integrative taxonomy used on the same individuals could provide greater reliability and minimize the underestimation of biodiversity.

An integrative taxonomy of Asian Caryodaphnopsis (Lauraceae) based on morphology and phylogenomics

AbstractCaryodaphnopsis is a small genus of the Lauraceae family with disjunct distribution in tropical Asia and tropical America. Taxa from each region form a well­supported monophyletic clade. However, the species circumscription in Asian Caryodaphnopsis remains ambiguous due to few and generally unstable morphological characters used for classification. To address this, we conducted an integrative taxonomy of the Asian clade. We investigated 13 populations, sampled 57 accessions covering the entire taxonomic diversity, geographic range, and morphological variation in Asia, and reconstructed a new phylogeny using single‐nucleotide polymorphisms. We obtained a well‐resolved phylogeny of the Asian Caryodaphnopsis. The first diverged clade consists of three species endemic to lowland forests. The second (largest) clade (eight species in total) mainly occurs in limestone mountain forests and has a double‐layered lower leaf epidermis morphology of type 2 (less complete roomy with closed “roof”) and type 3 (complete roomy with closed “roof”). Based on the diagnostic morphological characters, distinct distributional ranges, and monophyly of each clade, we recognize 12 species in the Asian Caryodaphnopsis, including three new species: C. cangyuanensis sp. nov., C. hekouensis sp. nov., and C. rubrinervis sp. nov. We also reinstate C. latifolia, and re‐circumscribe the widespread C. tonkinensis and other species based on fruit and inflorescence morphology and molecular evidence.

Phylogenetics, Molecular Species Delimitation and Geometric Morphometrics of All Reddish-Brown Species in the Genus Neotriplax Lewis, 1887 (Coleoptera: Erotylidae: Tritomini).

To date, five species of reddish-brown Neotriplax have been described, but their highly similar body color and other phenotypic traits make accurate taxonomy challenging. To clarify species-level taxonomy and validate potential new species, the cytochrome oxidase subunit I (COI) was used for phylogenetic analysis and the geometric morphometrics of elytron, pronotum, and hind wing were employed to distinguish all reddish-brown Neotriplax species. Phylogenetic results using maximum likelihood and Bayesian analyses of COI sequences aligned well with the current taxonomy of the Neotriplax species group. Significant K2P divergences, with no overlap between intra- and interspecific genetic distances, were obtained in Neotriplax species. The automatic barcode gap discovery (ABGD), assemble species by automatic partitioning (ASAP), and generalized mixed Yule coalescent (GMYC) approaches concurred, dividing the similar species into eight molecular operational taxonomic units (MOTUs). Geometric morphometric analysis using pronotum, elytron, hind wing shape and wing vein patterns also validated the classification of all eight species. By integrating these analytical approaches with morphological evidence, we successfully delineated the reddish-brown species of Neotriplax into eight species with three new species: N. qinghaiensis sp. nov., N. maoershanensis sp. nov., and N. guangxiensis sp. nov. Furthermore, we documented the first record of N. lewisii in China. This study underscores the utility of an integrative taxonomy approach in species delimitation within Neotriplax and serves as a reference for the taxonomic revision of other morphologically challenging beetles through integrative taxonomy.