Abstract Tanaidacea contains about 1600 species across four superfamilies (Apseudoidea, Neotanaoidea, Paratanaoidea, and Tanaidoidea). Phylogenetic relationships among the four superfamilies had been investigated molecularly with one to several genes, or with transcriptome data, but not with mitochondrial genome (mitogenome) data. Mitogenome data were available only for Pseudotanais sp. (Paratanaoidea) and Arctotanais alascensis (Tanaidoidea). We sequenced and annotated a complete mitogenome sequence for Sinelobus kisui (Tanaidoidea) and nearly complete mitogenome sequences for Apseudes nipponicus , Apseudes ranma , Carpoapseudes spinigena (Apseudoidea), and Neotanais sp. (Neotanaoidea). We detected the typical two rRNA and 13 protein-coding genes (PCGs) in all five species, but failed to detect one to five tRNAs in each species. The gene orders of the four superfamilies differed from the pancrustacean ground pattern and from one another. In maximum-likelihood phylogenetic trees based on amino acid and nucleotide datasets for the 13 PCGs and two rRNA genes, Tanaidacea emerged as a weakly supported sister clade to Isopoda. Within Tanaidacea, a fully supported Apseudoidea was the sister group to a moderately supported clade comprising the other three superfamilies; in the latter clade, a weakly supported Tanaidoidea clade was successively sister to Paratanaoidea and then to Neotanaoidea. The mitogenome-based topology among the four superfamilies differed from previous topologies based on 18S rRNA or transcriptome data (Apseudoidea, (Paratanaoidea, (Neotanaoidea, Tanaidoidea))). This contradiction suggests possible effects of mito–nuclear discordance or long-branch attraction for Pseudotanais sp. and S. kisui .
Abstract Despite the recorded presence in Hawai'i of Cephalonomia hyalinipennis, a presumed parasitoid of the coffee berry borer (Hypothenemus hampei), it has never been recovered from infested coffee berries. Here we show, using morphometric and molecular data, that what has been recorded in Hawai'i as C. hyalinipennis is actually a complex of morphologically similar species, each distinct from specimens attacking the coffee berry borer in Chiapas, and from the C. hyalinipennis types from Florida. The taxon from Chiapas utilizing the coffee berry borer is also found to be distinct from C. hyalinipennis sensu stricto, and is here described as C. soconuscensis sp. nov. The five species found in Hawai'i are described as Cephalonomia lynnea sp. nov., a parasitoid of Hypothenemus eruditus; Cephalonomia luiji sp. nov.; Cephalonomia liula sp. nov., a parasitoid of a Leptophloeus sp. known to predate on the coffee berry borer; Cephalonomia kala sp. nov.; and Cephalonomia zander sp. nov., a parasitoid of an Ozognathus sp. Laboratory testing showed that C. lynnea has the ability to develop on coffee berry borer larvae, but it does not appear to do so in the wild. In addition to morphological data, DNA barcoding was used to provide further support for distinguishing species in the C. hyalinipennis species complex. The large dataset of morphometric measurements showing reliability of different ratios, and degenerate PCR primers developed to permit assembly of COI barcodes even from old museum specimens, should enable more rapid discernment within such cryptic species complexes in the future.
In the highly diverse, endemic Madagascan lineage of Syntomini, the "black-and-white" general body pattern appears to be present in a few closely unrelated clades (and in six existing genera), causing uncertainties in their systematics. Based on materials from five natural history collections, we confirm the monophyletic character of the morphologically variable and largely montane genus Tritonaclia Hampson, 1898, with six existing species, inferred from both molecular and morphological evidence. The taxonomy of the genus is revisited, with the description of two new species: T. vonifotsy Wiorek, sp. nov. and T. ombilahy Wiorek & Przyby & lstrok;owicz, sp. nov.. Naclia melania Oberth & uuml;r, 1923 syn. nov. is treated as a junior subjective synonym of Glaucopis tollini Keferstein, 1870, and lectotypes of both taxa, now placed in Tritonaclia, are designated. We provide determination keys for the genus, based on adults, and their male and female genitalia. All available data on the distribution and ecology of Tritonaclia species are presented, including new geolocalisation of some important insect collecting places in Madagascar. Additionally, we provide descriptions and illustrations of male scent scales (androconial patches) on fore- and hindwings, present in two Tritonaclia species: T. tollini and T. vonifotsy Wiorek, sp. nov. Such structures are recorded in the tribe Syntomini for the first time.
Abstract African tropical forests are highly underexplored regarding microinvertebrate diversity, including tardigrades. Here, we describe two new tardigrade species from the genera Parahypsibius Gąsiorek et al., 2024 and Echiniscus C.A.S. Schultze, 1840, collected in the tropical rainforest of the Kibale National Park in southwestern Uganda. Species delimitation was based on an integrative approach combining morphological, morphometric, and molecular data. Morphological analyses employed light and scanning electron microscopy, while molecular characterization used mitochondrial COI and nuclear 18S rRNA, 28S rRNA, ITS1, and ITS2 markers. Additionally, we report Echiniscus lineatus Pilato, Fontoura, Lisi and Beasley, 2008 from Uganda for the first time, providing new sequences and reconstructing haplotype networks using previously published genetic data of this species. Phylogenetic analyses of subfamily Hypsibiinae Pilato, 1969 and genus Echiniscus clarify the relationships of the new taxa and place them within the broader phylogenetic context. Our study highlights the underestimated diversity of African tardigrades and demonstrates the importance of integrative taxonomy for species delineation. The new records from Kibale National Park provide also baseline data for understanding the distribution and phylogeny of tardigrades of the Central Africa.
Abstract Hainanthereua albilineata gen. nov. et sp. nov. is described and illustrated based on specimens from Hainan Province, China. Morphological examination showed that these specimens belong to Thereuoneminae and share certain similarities with Thereuopodina Verhoeff, 1905. Phylogenetic reconstruction based on five genes (nuclear 18S and 28S rRNA, mitochondrial 12S and 16S rRNA, and cytochrome c oxidase subunit I) indicated that the specimens form a distinct and well-supported clade that is sister group to Thereuonema Verhoeff, 1904, so a new genus is accordingly established. Combining the evolutionary history and biogeographic framework of Thereuoneminae, this study revealed the evolutionary significance of Hainanthereua gen. nov. in the context of the Peninsular Indian Plate as a biotic ferry implicated in the origin of East and Southeast Asian lineages. Morphological similarities between Hainanthereua and Thereuopodina are symplesiomorphies of a clade that includes these two genera, Thereuonema and Thereuopoda Verhoeff, 1904.
Abstract Recent field campaigns conducted in Peru along with the examination of museum specimens allowed us to identify small tarantulas that do not fit with any known Theraphosidae genera. Morphology and additional molecular evidence from the mitochondrial gene COI led us to propose Kiskalla gen. nov. from southern Peru, at Puno region. Three new species of Kiskalla gen. nov. (K. ignacioi sp. nov., K. yeisoni sp. nov. and K. zukuapasanka sp. nov.) are herein described, diagnosed and illustrated. Kiskalla gen. nov. differs from the known Theraphosinae genera in the presence of lateral stripes on the abdomen and a small dorsal arrowhead-shaped patch of type III urticating setae, presence of a large number of spines on all legs, short and stout setae on the dorsal metatarsi encircling the filiform trichobothria, an apical crown of long spines on the metatarsi of all legs in both sexes and scopula of tarsi IV slightly developed, restricted to the margins due to the presence of long ventral setae. Males and females show genitalic features that resemble those of Hapalotremus Simon, 1903, but differ by the development of copulatory bulb keels and the aspect of the spermathecae. We also present data on the natural history and distribution of the species.
Abstract Larvae of the megadiverse Scarabaeoidea play important economic and ecological roles, and their mouthparts provide valuable morphological clues for species identification and dietary inference. However, the underlying relationships between morphological characters and feeding habits remain insufficiently elucidated. In this study, the larval mouthparts of 22 species representing ten major lineages of Scarabaeoidea were examined in detail. Seventeen characters of the larval mouthparts were analyzed within a phylogenetic framework. Three characters were identified as candidate synapomorphies for the major lineages represented here: maxillary stridulatory teeth for Scarabaeidae, mandibular stridulatory ridges for Cetoniinae + (Dynastinae + Rutelinae), and a circular arrangement of epipharyngeal phobae for Aphodiinae + Scarabaeinae. By contrast, several homoplastic mouthpart traits were recurrently associated with larval feeding habits in the sampled taxa, including a setose acroparia, heli on the epipharynx, and a blade-like incisor region in phytophagous lineages, and the circular phoba complex together with the absence of nesium in coprophagous lineages. These results suggest that larval mouthparts in Scarabaeoidea may retain both phylogenetic and trophic signals, providing a morphological basis for interpreting the evolution of feeding habits in scarab larvae.
Abstract Phylogenetic reconstruction of Proctotrupinae, the largest subfamily of Proctotrupidae, was conducted for the first time based on total evidence combining molecular and morphological characters. Based on our analysis, we conclude that the genus Nothoserphus, previously belonging to the tribe Cryptoserphini, should be transferred to the tribe Disogmini. New diagnostic characters for each genus were proposed based on the morphological analysis. The results of the analysis showed that the exceptionally hardened ovipositor sheath is a defining character of Proctotrupidae, and it was estimated that its morphological character states were not reflected in the phylogenetic relationship, probably because of its variability related to parasitic strategy and host habitat.
Abstract Mosquitoes rank among the most deadly organisms worldwide, facilitating >700,000 human deaths annually through transmission of vector-borne pathogens. Culex are famous as vectors of multiple pathogens affecting both animals and humans. This study presents the first mitogenome sequencing and comparative analysis of seven species within Culex. Our findings demonstrated conserved structural features and nucleotide composition across the mitogenomes of these species. This study performed phylogenetic analysis of Culex based on mitochondrial genome data under both homogeneous and heterogeneous models separately, and estimated the divergence times. Phylogenetic analyses revealed that Culex is paraphyletic, with Lutzia nested within it. Both Cx. (Neoculex) and Cx. (Culex) were non-monophyletic. The two species of Cx. (Neoculex) were placed in separate lineages, with Cx. fergusoni as the sister group to all other Culex. Meanwhile, Cx. (Culex) was rendered paraphyletic by the inclusion of Cx. (Culiciomyia) and Cx. (Oculeomyia) within its clade. Divergence time estimation placed the basal split of Culicidae in Late Triassic, followed by the Culicinae-Anophelinae divergence in Late Jurassic (~147 Mya), with all speciation events within Culex postdating these splits and clustering in Neogene. This study provides a fundamental basis for understanding the mitogenomic architecture and phylogenetic relationships within the genus Culex, and also establishes a theoretical foundation for transmission mechanisms and control strategies of common mosquito-borne diseases.
Abstract Recent systematic work has shown that Lamproblattidae may be an ancient lineage of Blattodea. Given their putative position as sister to Xylophagodea, investigations of their little-known biology can uniquely offer new insight into the evolution of wood-feeding and thus, the evolutionary precursors to eusociality. Here, we compiled all literature information on Lamproblattidae in a new appraisal, integrating it with novel data collection. First, we revised the systematics of Lamproblattidae. We provide a key to the family and related taxa, give revised definitions of taxa, remove Eurycanthablatta from Lamproblattidae, describe Lamproblatta mimadelfi sp. nov., and provide a morphology-based phylogeny for Lamproblattidae. The phylogeny weakly demonstrates the monophyly of Lamproblattidae, the paraphyly of Lamproblatta relative to Lamproglandifera, the lack of support for Eurycanthablatta within Kittrickea, and weak support for Anaplecta + Lamproblattidae. Next, we reviewed hundreds of iNaturalist observations to gain new occurrence data for Lamproblattidae. Combining this distribution data with that from a literature review and other publicly available databases, we model the habitat suitability for Lamproblattidae. Finally, we used multiple genetic techniques to reveal new information about the physiology of L. mimadelfi. We find no evidence for the presence of hindgut parabasalians, but we do find evidence of indirect wood-feeding – xylomycophagy. Metabarcoding of gut contents shows that L. mimadelfi is a generalist feeder, but wood-associated fungi are the most consistent and abundant food across samples, comprising about 40% of its diet. We then discuss all the results to hypothesize possible evolutionary sequences for the acquisition of direct wood-feeding in Kittrickea, and the shift from generalist to specialist feeding.
Abstract Therevidae is a diverse family of lower Brachycera, yet its early evolutionary history remains poorly understood due to the absence of available Mesozoic fossils. Here, we describe †Paleothereva longicoxa gen. et sp. nov. from mid-Cretaceous Kachin amber, representing the oldest member of known derived therevids. Phylogenetic analysis and morphological comparisons integrating both extant and fossil species, consistently place this fossil within Taenogera genus-group of Agapophytinae. This finding indicates that Agapophytinae had already diverged by the mid-Cretaceous, suggesting an earlier origin of Therevidae than previously inferred from molecular data. The occurrence of Paleothereva on the Gondwanan-derived West Burma Block further supports the hypothesis that the ancestors of Agapophytinae—or possibly the broader Agapophytinae + Therevinae clade—originated in Gondwana. Ancestral character state reconstruction of female terminalia within therevid flies reveals that Paleothereva and extant relatives likely shared similar oviposition behaviors associating with sandy-soil environments, which may have driven the corresponding specialization and diversification of acanthophorite spines in female. These new findings provide rare, direct evidence illuminating the early evolution of derived therevid lineages.
Acanthocephala, with 40 known species, is the most diverse genus of the tribe Acanthocephalini, which contains some of the largest species of Coreidae. In this contribution, Acanthocephala guatemalena, A. granulosa, A. luctuosa, A. panamensis, and A. thoracica were resurrected as valid species, and a morphological phylogenetic analysis, using continuous and discrete characters, was performed to test the monophyly of the genus. The analysis was carried out under implied weighting, including 34 species of Acanthocephala as ingroup, and 20 species from other genera of Acanthocephalini and Placoscelini as outgroups. Acanthocephala was recovered as a monophyletic group, and sister of the clade (Spilopleura parensis + S. ochracea). To formalize and show the relationships of the species in Acanthocephala, the subgenera A. (Acanthocephala) and A. (Metapodiessa) were revalidated, and three new subgenera were proposed: A. (Spinipedia) subgen. nov., A. (Contrastata) subgen. nov., andA. (Pronoptera) subgen. nov. Additionally, a detailed redescription of Acanthocephala, a subgeneric key, descriptions and redescriptions for all subgenera, photographs of dorsal and ventral habitus of the type species of each subgenus, and photographs of the diagnostic characters, including male and female genitalia are provided.
Coccoderus Buquet, 1840 is one of the largest genera in Torneutini (Cerambycinae), composed of 14 species. A previous morphology-based phylogeny recovered the genus as a monophyletic group. Since then, four species were described in the genus. Herein, we expanded this previous morphological dataset with the inclusion of seven characters and eight species. We performed maximum parsimony, maximum likelihood and Bayesian inference analyses. We used the Statistical Dispersal-Vicariance Analysis and the Bayesian Binary Method to reconstruct the potential ancestral ranges of Coccoderus. Our phylogenetic analysis recovered Coccoderus as a monophyletic group and shows shifts in species relationships, with two clades presenting new placements compared to the previous analysis. Biogeographic analyses identified the Chacoan dominion as a key region in the diversification of the genus, representing its ancestral area. Also, we described Coccoderus trimaculatus sp. nov., from Brazil, increasing the known species of Coccoderus to 15.
The North American checkered beetle (Cleridae) Thanasimus nubilus Klug, 1837 is currently listed as a subspecies of Thanasimus undatulus (Say, 1835). We test the species status of T. nubilus by investigation of DNA barcodes and morphological measurements of multiple specimens of each of the two taxa as well as of multiple specimens of two additional North American species of the genus. We use four species delimitation methods based on gene tree and clustering algorithms (barcode gap threshold clustering, Assemble ly supported treating these two subspecies as distinct species which was also supported by a morphometric analysis. We therefore propose Thanasimus nubilus Klug, 1842 stat. rev. to be reinstated as a valid species.
The treehopper genus Tricentrus is a widespread group of insects, characterized by typical spines and, in some species, sexually dimorphic suprahumeral horns. With 238 described species worldwide, Tricentrus represents the most species-rich genus in the subfamily Centrotinae. Currently, 72 species are documented in China, yet the phylogenetic relationships and evolutionary history of this genus remain unknown. Here, we use an integrative approach to characterize and describe these treehoppers based on materials collected from China. We name three new species: Tricentrus allochrous Li & Chen sp. nov. and Tricentrus pianmaensis Li & Chen sp. nov. from Yunnan, Tricentrus dexingensis Li & Chen sp. nov. from Xizang, with the revision of one synonym. Combined analysis morphological comparisons, species delimitation and phylogeny reliably separate these new species from known species, and sexual dimorphism with suprahumeral horns in three species is reported for the first time. Divergence time estimation indicates that the Tricentrus differentiated during the Early Cretaceous (110.41 Mya). Ancestral state reconstruction reveals that the most recent common ancestor possessed suprahumeral horns in both sexes and exhibited a short tubular pygofer apex. Our analyses demonstrate four evolutionary transformations in pygofer apex morphology and seven distinct transitions in suprahumeral horn development. These morphological diversifications likely resulted from multiple mechanisms, including sexual selection, life history strategies with associated behavioral adaptations, and incomplete lineage sorting.
Abstract The evolutionary dynamics of morphological traits can often blur the boundaries between interspecific divergence and intraspecific variability, complicating species recognition. This study investigates the variation in secondary sexual traits and the existence of potential speciation processes within what is now considered Berberomeloe insignis (Coleoptera: Meloidae), an endangered blister beetle taxon endemic to southeastern Spain. Despite previous evidence of substantial genetic and phenotypic differentiation, key characters as the morphological variation in secondary sexual traits, such as antennomeres, remain unexplored. Using geometric morphometrics, we analyzed the shape variation of male and female antennomeres VII–XI across all previously recognized lineages of B. insignis. Our results reveal significant morphological differentiation, particularly in antennomeres VII, IX, and XI, which correlate broadly with genetic lineages. Based on the study of newly recorded populations, we confirm that cephalic coloration patterns correspond with mitochondrial lineages, further supporting the existence of geographic lineage differentiation within what was previously considered B. insignis. Climatic niche modeling indicates low climatic niche overlap between the isolated western lineage and the remaining lineages, which also show relatively low to moderate overlap, suggesting that ecological factors could have contributed to the divergence among them. These findings underscore the intricate interplay of genetics and ecology, highlighting the importance of integrating multiple data sources for accurate species delimitation. Based on our results, we describe B. nazari sp. nov. and B. insignis trisanguinatus ssp. nov. reflecting the evolutionary history of this group.
The taxonomy of the subgenus Staphylus (Capilla) is reviewed, including redescriptions of known species, identification keys, and detailed distribution maps. The female genitalia of Staphylus (Capilla) azteca (Scudder, 1872), S. (C.) caribbea (Williams & Bell, 1940), S. (C.) corumba (Williams & Bell, 1940), S. (C.) eryx Evans, 1953, S. (C.) tucumanus (Pl & ouml;tz, 1884) and S. (C.) tyro (Mabille, 1878) are described and illustrated for the first time. Four new species are described: Staphylus (Capilla) nicoleae Lemes sp. nov. from Colombia and Venezuela, S. (C.) ricardoi Lemes, sp. nov. from Peru, S. (C.) neideae Lemes sp. nov. and S. (C.) neivae sp. nov. from Brazil. Pholisora imperspicua Hayward, 1940 is a syn. nov. of Staphylus (C.) lizeri lizeri (Hayward, 1938) and Hesperia melangon epicaste Mabille, 1903 is a syn. nov. of Staphylus (C.) melangon melangon (Mabille, 1883). Neotypes are designated for Staphylus epicaste Mabille, 1903, Nisoniades tucumanus Pl & ouml;tz, 1884 and Staphylus fascia Hayward, 1933. Lectotypes are designated for Pholisora azteca Scudder, 1872, Bolla machuca Schaus, 1913, Helias tyro Mabille, 1878, Staphylus anginus Schaus, 1902, Hesperia melangon 1883, Hesperia musculus Burmeister, 1875 and Helias aurocapilla Staudinger, 1876.
Pholcid spiders have long been classified into five subfamilies, and this framework ultimately dates back to Eugène Simon’s non-phylogenetic system of 1893. While subfamily relationships and compositions have been updated extensively over the last decades, no new subfamily had to be erected for any of the hundreds of new species newly described since Simon. Here we report two new species from semi-arid Brazilian Caatinga: Caipira mineira Huber sp. nov., and Caipira baiana Huber sp. nov. Genomic data strongly support their sister-group relationship; we thus join them conservatively in a single genus, Caipira Huber gen. nov., even though they show some remarkable morphological differences. This genus is sister to a large clade including all pholcid subfamilies except Pholcinae and Smeringopinae, which necessitates the erection of a new subfamily: Caipirinae subfam. nov. In addition, we formalize the separation of the genus Artema from ‘other Arteminae’. This had previously been suggested by multi-locus genetic data, and is strongly supported by new genomic data. Arteminae is newly circumscribed to include only Artema and Priscula, and the name Physocyclinae subfam. nov. is proposed for ‘other Arteminae’. Pholcidae is thus divided into seven subfamilies, with the following relationships suggested by genomic data: (Pholcinae, Smeringopinae), (Caipirinae, ((Arteminae, Ninetinae), (Physocyclinae, Modisiminae))). Finally, we tested the hypothesis that the Chilean genus Aucana is the closest relative of the new Brazilian species. This is strongly rejected; Aucana is resolved as the only known South American representative of Physocyclinae.
Apolygus (Insecta: Heteroptera: Miridae: Mirinae) is a large and taxonomically challenging trans-Palearctic genus comprising 74 species, and it has never been revised. The greatest diversity of this genus occurs in Asia, and 15 species have been recorded from Russia. Among them, Apolygus limbatus, Apolygus lucorum, and Apolygus spinolae are trans-Palearctic, and A. lucorum and A. spinolae are economically important as pests. Previous studies have shown that A. lucorum and A. spinolae are very similar in morphology and that the barcoding region might not be suitable for their identification. In this study a revision of the Apolygus representatives inhabiting Russia is provided based on morphological and molecular data. Mitochondrial (COI and 16S rRNA) and nuclear markers (ITS1 and 28S3 rRNA) were used for phylogenetic analyses and automatic species delimitation methods were applied. Apolygus lucorum, A. malaisei, A. nigronasutus, A. nigrovirens were treated as junior synonyms of A. spinolae because morphological and molecular data did not show differences between those species. Apolygus shikotan sp. nov. was described as new to science. Morphological data and the COI phylogeny show that A. limbatus might represent two separate species. However, additional specimens and nuclear markers for both groups are needed to test this hypothesis. The synonymy of A. syringae with A. hilaris was supported based on the morphological data. Apolygus gotorum and A. subhilaris are reported from Russia for the first time. The nuclear marker ITS1 is the most reliable marker for species delimitation and identification of Apolygus species.
Tabanus bifarius Loew, 1858 is a horsefly species distributed throughout the Mediterranean region and in the areas of central and southeastern Europe and partly in Asia. In this study, we conducted the first comprehensive phylogeographic and population genetic analysis of T. bifarius collected from 13 localities across Türkiye and Iran, through utilizing mitochondrial (COI) and nuclear DNA (ITS1–5.8S–ITS2) markers. A total of 81 haplotypes and 59 nuclear alleles were identified among 187 sequenced individuals. Both mitochondrial and nuclear DNA patterns and diversity observed in T. bifarius are indicative of complex historical and demographic processes. Among all the populations studied, Eskişehir and Hatay exhibited the highest genetic diversity, which may be due to the region’s topography and transitional zones. Phylogenetic analyses suggested that T. bifarius split from the outgroup species approximately 5.53 million years ago (MYA) during the Late Miocene–Early Pliocene, likely driven by tectonic and climatic events. Subsequent diversification events that occurred during times of climatic and environmental fluctuations in the Late Pliocene and Early Pleistocene also seemed to have significantly affected the species and gave rise to the formation of some important genetic lineages. Our analyses results indicate that T. bifarius exhibits a structured genetic landscape shaped by multiple refugial routes, geographic barriers, and Quaternary climatic oscillations. Further, our findings suggest that the species likely entered Anatolia through three distinct routes: (1) from the Levant region into southern Anatolia via Hatay; (2) from the Caucasus into northeastern Anatolia through Ardahan; and (3) from the Iranian Plateau into eastern Anatolia via Van.
