ABSTRACT The Embioptera fauna of Mato Grosso do Sul is still poorly documented. This study provides the first contribution to the study of the order in this state, focusing on the family Archembiidae. Three new species are described, Biguembia paxixi sp. n. Malacosembia morena sp. n. and Pararhagadochir hirsuta sp. n. Additionally, the female of Biguembia cocum Szumik, is recorded and described. Pararhagadochir flavicollis (Enderlein) is recorded for the first time in Brazil, while Archembia dilata Ross, Pararhagadochir castaneus Salvatierra, and Pararhagadochir confusa Ross, are recorded for the first time in the state. Finally, the type localities of Archembia oruma Szumik, Biguembia cocum Szumik, and Gibocercus urucumi are briefly discussed. A key to species of Malacosembia is provided; B. paxixi sp. n. and P. hirsuta sp. n. are added to previous keys. http://www.zoobank.org/urn:lsid:zoobank.org:pub:EB3A91FA-5954-40AC-80C6-A32C084674FF

The aim of this retrospective observational study was to evaluate the prevalence and distribution of dental anomalies in orthodontic patients with different malocclusions. Pretreatment diagnostic records of 766 orthodontic patients were retrospectively evaluated in a single-center setting. Dental anomalies were classified according to sex, orthodontic malocclusion type (Class I, n = 122; Class II, n = 568; Class III, n = 76), and jaw localization. Statistical analyses were performed using Fisher's exact test, Yates' correction, and Pearson's chi-square test. No statistically significant association was found between sex and the prevalence of dental anomalies (P = 0.739). At least one dental anomaly was detected in 71.8% of the patients. Root dilaceration was the most frequently observed anomaly (35%). The prevalence rates of numerical, dimensional, shape, eruption, and structural anomalies were 13.97%, 4.04%, 53.91%, 24.15%, and 30.67%, respectively. No significant difference in the prevalence of dental anomalies was observed among different malocclusion groups (P = 0.055). Dental anomalies were frequently observed in this orthodontic population. The prevalence and distribution of anomalies varied according to anomaly type and jaw localization, highlighting the importance of radiographic evaluation in orthodontic patients. These findings may support earlier detection of diagnostic challenges and more individualized treatment planning in routine orthodontic practice. Early identification of dental anomalies may facilitate comprehensive orthodontic treatment planning and help prevent potential functional and structural complications.

Spinal cord injury (SCI) is a devastating neurological disorder marked by profound disturbances in cytokine signaling, redox balance, mitochondrial homeostasis, and glial-neuronal communication. Although many therapeutic strategies have been explored to attenuate secondary injury, effective molecularly targeted interventions remain limited. Increasing evidence identifies signal transducer and activator of transcription 3 (STAT3) as a central signaling node in the neurochemical response to SCI. Recent studies indicate that STAT3 exhibits pronounced spatiotemporal and cell-type-specific activation after SCI. Depending on the upstream trigger and cellular compartment involved, STAT3 can amplify or restrain neuroinflammation, shape astrocyte and microglial reactivity, influence mitochondrial bioenergetics and oxidative stress, modulate ferroptosis and apoptosis, and alter the regenerative state of the injured spinal cord. In this review, we frame STAT3 not simply as a downstream effector of the JAK/STAT cascade, but as an integrative regulator of SCI neurochemistry that links cytokine-driven signaling to metabolic stress, glial remodeling, and axonal repair. We emphasize how injury phase, cell type, and subcellular localization influence STAT3-dependent outcomes, discuss emerging therapeutic strategies that converge on STAT3-centered pathways, and outline the key challenges that must be addressed for precise translational targeting.

Alexandrium minutum Halim, the generitype species of the toxigenic Alexandrium, was described from a 1958 bloom in the Eastern Harbour, Alexandria (Egypt), but has never been re-examined with modern methods from the type locality. Here, we establish a monoclonal strain (TIO1796) by cyst germination from Alexandria's port sediment and a second strain (TIO1816) from plankton samples from the Gulf of Suez. Cell morphology of both cultured strains and plankton samples was examined using scanning electron and light microscopy. The plate formula of A. minutum was identified as Po, *4', 6'', 6C, 9S, 5''', 2p, 1''''. Our results confirmed the consistent presence of a ventral pore and a variable connection between Po and 1'. Molecular phylogenetic analyses based on LSU rRNA gene and ITS rRNA region sequences revealed four ribotypes (A-D) of A. minutum. The LSU rRNA gene and ITS rRNA region sequences of both strains were identical and belong to the widespread ribotype A ("Global/Mediterranean clade"). The other ribotypes are restricted to the Asia-Pacific region. Strain TIO1796 (type locality) was non-toxic regarding paralytic shellfish toxin (PST < 0.024 pg · cell-1), whereas TIO1816 produced GTX1/4-dominated toxins (0.48 pg PST · cell-1). These results (i) anchor the name A. minutum to ribotype A via neotypification of strain TIO1796, (ii) document a PST-producing A. minutum in the Red Sea, and (iii) provide evidence supporting that A. minutum constitutes a cryptic species complex analogous to the A. tamarense species complex. The historical decline of A. minutum blooms in Alexandria since the 1990s correlates with near-absent cyst banks in recent sediments.

A new species of Pristimantis is described from the Cordillera del Cóndor, Zamora Chinchipe Province, southeastern Ecuador, based on morphological and molecular data, Pristimantisetsasp. nov. The new species is diagnosed from its congeners by the following combination of characters: female SVL 32.5 mm and male SVL 18.3 mm, dorsolateral folds formed by rows of subconical tubercles, strongly areolate ventral skin, two distinct rows of forearm tubercles, one along the ventrolateral margin and a second along the externolateral margin of the forearm, and a prominent yellow groin blotch in the female holotype. The species belongs to the Pristimantiscryptomelas group, part of the Huicundomantis subgenus, and is closely related to P.nangaritza, P.verrucosus, and P.plateado. Currently, the species is known only from its type locality, where it inhabits low montane evergreen forests at elevations of 1,655–1,830 m. Additionally, we discuss the use of the term “ulnar tubercles” in Pristimantis, noting that it may refer to tubercles occupying different positions on the forearm.

ABSTRACTElsholtzia Willd. is the largest genus in tribe Elsholtzieae (Lamiaceae) and has notable medicinal and economic value in East Asia. During a field survey in Sichuan Province, China, we discovered a morphologically distinct population of Elsholtzia resembling E. lamprophylla but differing in leaf size, indumentum, inflorescence type, and bract shape. To determine its taxonomic status, we conducted phylogenomic analyses using 1115 single‐copy nuclear genes and 240 plastid loci from 45 accessions, representing major lineages of Elsholtzia. Both nuclear coalescent and concatenation‐based trees consistently placed the new species in Clade I (Ser. Fruticosae) with strong support, while plastid data revealed a close relationship with several Ser. Fruticosae members, suggesting possible hybridization or incomplete lineage sorting. Morphometric analyses of 12 vegetative and floral traits clearly separated the new species from E. lamprophylla. Based on morphological and phylogenomic evidence, we describe it as Elsholtzia yajiangensis Y. Wang, L. J. Liu & Q. Wang, sp. nov. It is characterized by its secund spikes, broadly lanceolate bracts, strigose leaves with 5–7 lateral veins, and remotely obtuse‐serrate margins. The species is currently known only from the type locality in Yajiang County, Sichuan, and is provisionally assessed as Data Deficient (DD) based on its narrow distribution and ongoing threats. This discovery highlights that the species diversity of Elsholtzia in the Hengduan Mountains remains underestimated.

Within the family Osphronemidae, mouth-brooding fighting fishes are small, air-breathing fish species that hold their eggs and offspring in their buccal cavities as a parental care behavior, usually found in running waters and distributed in the wild throughout Southeast Asia. This study aims to describe the morphological development and generate identification keys for the larval and juvenile stages of seven wild mouth-brooding fighting fish species found in Thailand, i.e., Bettaapollon, B.ferox, B.pallida, B.pi, B.prima, B.pugnax, and B.simplex from the wild. The broodstocks were collected from type localities or based on characteristics that most closely matched each species description, with healthy fishes selected and breeding continued for our size-series collection, including B.simplex, a previous study. The results showed that mouth-brooders released their offspring when developed to the post-flexion stage within 11–12 (mode = 11) days after fertilization (DAF), except B.pi, which took 18–20 DAF, and the post-flexion larva developed to the juvenile stage within 18–30 days after release. The main characteristics of the new-release post-flexion larva were an oblong and depressed body, a large head, an oval to rounded eye, and rays where the caudal fin began to develop, as well as having fully developed ventral fins with two or three dorsal, central, and ventral stripes and a caudal spot. Myomere numbers and fin rays differed among species across a range of 8–10 dorsal, 10–13 pectoral, 6 ventral, 24–31 anal, and 10–13 caudal fin rays. Diagnostic characters were selected to create a dichotomous identification key, with an illustration provided. In terms of taxonomy, the different larval stages also differed in pigmentation patterns among species, with pigmentation patterns on head (pre-orbital, sub-orbital, post-orbital, and sub-opercular bands) and longitudinal stripes on the side of the body able to be used to distinguish among different development stages as well as different species.

Oreocharisfulvovillosa, a new species of Gesneriaceae, is described and illustrated. This new species has only been discovered on the rock walls of an isolated karst hill at its type locality in Mang City, western Yunnan, China.

BackgroundBiodiversity is at increasing risk, and amphibians are the most threatened vertebrate class, with 40.7% of species globally at risk. For amphibians, the primary cause for their decline is habitat loss. The scale of this global issue becomes most evident in so-called deforestation hotspots, such as the Chiquitano Dry Forest in Bolivia, which harbours a highly diverse fauna and flora with many species still awaiting formal description. Only recently, the frog Oreobates chiquitanus has been described from a single location within this forest. Since then, it was discovered at only two additional sites. An ongoing logging initiative at the type locality led to the deforestation of the sampling site, even before the species was formally described. Ongoing logging activities in the vicinity of all known sites raise questions about the species’ fate. Here, we provide a comprehensive genetic definition of the species and try to assess the genetic diversity and conservation value of the type population.ResultsWe sequenced all available type specimens of Oreobates chiquitanus, providing a reference genome from one of the paratopotypes and mitochondrial assemblies for all type specimens. We further show that all known populations of this species are near recent logging initiatives. The reconstruction of the demographic history indicates that the population was recovering from a dip roughly 300kya. The estimates of relatedness and heterozygosity imply a genetically vital population.ConclusionsThis study exemplifies the importance of collaboration of natural history collections and genomic initiatives like TBG (Translational Biodiversity Genomics) for a better understanding of the anthropogenic impacts in times of global change, to identify and describe biodiversity, to raise awareness, and inform potential conservation measures. We not only provide a comprehensive and permanent molecular definition for O. chiquitanus but also assess the genetic diversity of the population. Therefore, this study provides an important baseline for future studies of genetic erosion in this species. Furthermore, it illustrates the risk of biodiversity loss in the Chiquitano Dry Forest and signifies the need for conservation efforts.Supplementary InformationThe online version contains supplementary material available at 10.1186/s12864-026-12984-5.

The three-branched bamboos (Arundinarieae) form a taxonomically difficult group due to recent rapid radiation and frequent hybridization, with generic and species delimitations of many taxa problematic. Indosasa solearis, described from North Vietnam based on flowering materials, was later transferred to Sinobambusa. But its taxonomic position was controversial due to limited vegetative data. After a detailed morphological study of this bamboo at its type locality, we found that the overall morphology of this species is surprisingly similar to I. parvifolia and I. jinpingensis. Thus, we conducted morphological and phylogenomic analyses to ascertain the taxonomic position of I. solearis and its relationship with those two species. Morphological examination confirmed I. solearis exhibits diagnostic characters of Indosasa, like leptomorph rhizomes, diffuse culms, strongly and asymmetrically raised branch supra-nodal ridges, 4-9 foliage leaves per ultimate branch, pseudospikelets, and each floret with six stamens and three stigmas. No essential difference was found among the three Indosasa species both on macro- and micromorphology. Phylogenies based on single-copy orthologous nuclear gene datasets and single nucleotide polymorphism (SNP) data strongly support I. solearis as an Indosasa member, closely related to I. parvifolia and I. jinpingensis. Based on the combined morphological and molecular evidence, we confirmed that I. solearis belongs to Indosasa, and is conspecific with I. parvifolia and I. jinpingensis. We therefore reinstate I. solearis and synonymize I. parvifolia and I. jinpingensis with it. Furthermore, our gene flow analyses by SNP data revealed the hybrid origin of the genus Sinobambusa, with genomic signatures suggesting an ancient cross between Indosasa and Pleioblastus. This finding highlights the reticulate evolutionary history in the three-branched bamboos.

Zacatecas is in the north-central region of Mexico. Its territorial extension, as well as the unique combination of climatic and physiographic characteristics, favor high biological diversity. However, until now, it has been considered one of the states with lowest herpetofaunal diversity, mainly due to low sampling effort. We provide an updated checklist of the herpetofauna of Zacatecas, including the physiographic provinces and ecoregions where they occur, a summary of their conservation status, and a comparison with neighboring states. Zacatecas has 25 species of native amphibians and 119 native reptiles, with five introduced species (1 frog, 2 lizards, 1 snake and 1 turtle). Of 149 species, four have their type locality in Zacatecas, 16 represent new state records, and 232 records from 79 species are new municipal contributions. More than half of the native herpetofauna (53.5%) of Zacatecas is endemic to Mexico. However, it does not have state endemics, but harbors populations of regional endemics that only inhabit north-central Mexico. Of the native amphibian and reptile species in Zacatecas, 2.8% are listed by IUCN in threatened categories (i.e., Vulnerable, Endangered, Critically Endangered), 13.9% are placed in a protected category by SEMARNAT (i.e., Threatened and in Danger of Extinction), and 33.3% are categorized as high vulnerability by the EVS criteria. Among adjacent states with which it shares a border, Zacatecas is the fourth with the largest territorial extension, the fifth in herpetofauna diversity, and the third with the highest number of country endemic species. Large areas of the state remain underexplored, suggesting that the herpetofauna richness of Zacatecas may increase in the future.

Description of Rhipicephalus hibericus phenology and main hosts in its type locality.

We studied three ciliate species from freshwater habitats: Tetmemena indica Bharti et al., 2019 from Kenya and Tetmemena pustulata (Müller, 1786) Eigner, 1999 and Pleurotricha setifera Engelmann, 1862 from Austria. Tetmemena indica had a rigid body measuring 73×33μm in vivo, a shortened dorsal kinety 4 (DK 4), and a cyst with a smooth surface. Tetmemena pustulata was similar to the original and other populations described near the type locality, characterized by a body size of 97×40μm in vivo, having a bipolar DK 4, and a process-bearing cyst. Pleurotricha setifera measured 156×60μm in vivo, with 24-26 frontal-ventral-transverse cirri, one left and one right marginal cirral row, an adoral zone of membranelles occupying 54% of the cell length on average, and undulating membranes arranged in an Oxytricha-like pattern. The gene phylogenetic analyses of the SSU rRNA sequences revealed that T. indica clustered within the T. saprai/T. indica complex and T. pustulata formed its own lineage within the Tetmemena subclade. Both species differed in SSU rRNA gene sequences, DK 4 length, and cyst morphology. Pleurotricha setifera was morphologically and molecularly similar to Gastrostyla setifera (Engelmann, 1862) Kent, 1882 and Pleurotricha oligocirrata Park et al., 2017. Consequently, we clarified the synonymy of P. setifera and proposed a neotype for T. pustulata.

The genus Merismopedia is morphologically characterized by its distinctive colony arrangement yet exhibits considerable variability among species. Recent studies suggested that Merismopedia is polyphyletic, and its most likely true lineage was provisionally placed within the Microcystaceae. However, the available molecular data for most of the extant species were scarce, and notably, its type species, Me. tranquilla, lacked contemporary distributional and molecular data. In the present study, we sampled multiple localities to capture the potential morphological and genetic variability within Merismopedia. Using culture-dependent and -independent methods, we collected an extensive dataset of 16S rRNA gene and 16S-23S ITS rRNA region sequences. This dataset included the type locality of Me. tranquilla, from which we successfully recovered morphologically authentic isolates and established an epitype. Subsequent phylogenetic analyses of Merismopedia- and Microcrocis-like taxa revealed the placement of Merismopedia sensu stricto within the Geminocystaceae, herein reinstated to Merismopediaceae and further confirmed extensive polyphyly. Based on these findings, we propose two new genera (Neomerismopedia and Planktopedia), and a new species, Me. sepulta. Additionally, we transfer Microcrocis gigas to Merismopedia. We proposed the provisional transfer of Me. hyalina to Synechocystis; however, further revision is needed. These taxonomic revisions provide a foundation for a comprehensive reevaluation of this widespread and morphologically diverse group of Cyanobacteria.

Histological analysis is crucial for understanding gastrointestinal (GI) tract homeostasis and disease pathophysiology. Various histological stains are commonly used in research settings for assessing development, disease pathogenesis, and therapeutic impacts. Specifically in the ordered architecture of the GI epithelium, current semi-quantitative analysis of histological staining relies heavily on manual scoring rubrics and often lacks robust spatial assessment. To address this gap, we developed an open-source ImageJ plugin, building on a closed-source predecessor, aimed at analyzing the spatial localization pattern of user-defined points-of-interest, such as positively stained cells, along the GI epithelial units. The plugin, developed using ImageJ 1.53.0 and Java programming language in Eclipse, interfaces with ImageJ and leverages Java libraries for data processing. The workflow involves uploading a microscopy image of the GI tissue of interest, determining base and top orientation landmarks of the GI epithelial units through manual identification, and annotating point-of-interest coordinates using ImageJ. The output includes centroid coordinates for each point-of-interest, the absolute distances of the points from the base and top landmarks, and the normalized distances of the points relative to the total height of the GI unit. The plugin generates histograms for displaying average point-of-interest distances along the GI unit axis. This information facilitates quantification of total points-of interest counts, analysis of height localization within the GI unit, and determination of average GI unit heights. The plugin serves as a crucial tool for robustly assessing various biological mechanisms within the GI tract, including EdU localization, migration distances, changes in cell type localization, and identification of new expression patterns along the GI unit axis. Overall, this open-source ImageJ plugin provides a semi-automated, user-friendly solution for leveraging important insights into spatial localization of tissue histology expression patterns within the GI structured architecture, with streamlined post-processing pipelines for robust large-scale analysis.

The resin bee Trachusa massauahensis Pasteels, 1984, previously known only from the female holotype collected in Massawa on the Eritrean Red Sea coast, is newly reported from Kenya and a small Red Sea island. The previously undescribed male of the species is described here for the first time. The morphological characters observed as well as genetic barcoding support the species’ placement within the subgenus Trachusa (Paraanthidium). Additionally, new material of Trachusa eburneomaculata Pasteels, 1984 is presented from Kenya, expanding the known range of this species, which had also been known only from the type locality. Additional diagnostic characters are provided.

Ischemic stroke remains a leading contributor to cardiovascular disease, and unstable atherosclerotic plaques in the carotid arteries are an important contributor to its pathogenesis. Notably, sex differences exist in plaque composition and morphology; whereas males are more likely to develop unstable plaques, females present with worse outcomes poststroke. As such, this study aims to elucidate sex-specific cellular and molecular mechanisms driving carotid plaque instability. We performed single-cell RNA sequencing on carotid endarterectomy specimens from male and female patients with stable or unstable plaques, profiling over 64 000 cells. To spatially contextualize these findings, we integrated single-cell RNA sequencing data with CosMx high-plex spatial transcriptomics using the human discovery 6k RNA panel to map gene expression and cell type localization within key histological regions defining instability. Single-cell RNA sequencing revealed that vascular smooth muscle cells (VSMCs), T lymphocytes, and macrophages comprised ≈88% of plaque-resident cells, with distinct sex-specific differences in relation to instability. In males, unstable plaques showed depletion of contractile VSMCs, and abundance of inflammatory synthetic and macrophage-like VSMCs, alongside increased inflammatory macrophages and activated naive T lymphocytes. In contrast, female unstable plaques retained relatively more contractile VSMCs but exhibited greater expansion of cytotoxic T lymphocytes and foam-cell macrophages. Sex-specific cell-cell interaction analysis demonstrated increased VSMC-to-stromal interactions and elevated macrophage-macrophage signaling in male and female unstable plaques, respectively. Through spatial transcriptomics, inflammatory synthetic VSMCs identified by single-cell RNA sequencing dominated the lipid core in both male and female unstable plaques. However, only in males did these cells also replace the fibrotic cap, forming immune-rich niches among themselves and with macrophage-like VSMCs. Overall, these findings uncover distinct cellular and spatial mechanisms driving plaque destabilization in males and females, highlighting the need for sex-specific approaches to the diagnosis and treatment of carotid atherosclerosis.

Wild-type adeno-associated virus type 2 (wtAAV2) is a small, non-pathogenic DNA virus within the family Parvoviridae that is modified to engineer recombinant AAV vectors used in gene therapy. wtAAV2 genomes localize to nuclear domains enriched in DNA damage response (DDR) factors through mechanisms that remain unknown. We have discovered that the host transcription factor KLF4 (Krüppel-like factor 4) and the DDR protein PARP1 (poly-ADP ribose polymerase 1) are key effectors regulating wtAAV2 nuclear trafficking. In silico analysis revealed that wtAAV2-associated genomic sites are enriched in KLF4 binding motifs. Confocal imaging demonstrated a close spatial association between KLF4 and wtAAV2 nuclear reservoirs, consistent with KLF4's known ability to form nuclear condensates. Mutation of the KLF4 binding site within the wtAAV2 genome and RNAi-mediated knockdown of KLF4 globally reduced the expression of viral Rep68/78 genes and attenuated the localization of the viral genome to cellular DDR sites. Chemical inhibition of the KLF4-interacting protein PARP1 with olaparib decreased the ability of wtAAV2 genomes to localize to cellular DDR sites and transcribe viral genes. Ectopic expression of wild-type PARP1, but not its KLF4-binding-deficient mutant, rescued wtAAV2 gene expression in PARP1-deficient cells. These findings define a novel mechanism by which wtAAV2 exploits the interaction between host transcription factors and DNA repair machinery to establish a persistent nuclear niche. Insertion of KLF4-binding elements into recombinant AAV2 gene therapy vectors is sufficient to enhance transduction of target cells, providing a framework for engineering vectors with improved nuclear targeting and transcriptional activity.IMPORTANCEWild-type adeno-associated virus type 2 (wtAAV2) has emerged as the preferred platform for engineering gene therapy vectors due to its non-pathogenic nature and ability to persist in host cells long term. However, limited understanding of how wtAAV2 genomes navigate the nuclear environment to establish viral reservoirs has hindered the development of efficient recombinant AAV (rAAV) vectors. We demonstrate that the protein KLF4 bound to the wtAAV2 genome recruits the virus to cellular KLF4 sites bound by PARP1. Disruption of either KLF4 binding, PARP1 activity, or KLF4-PARP1 interaction significantly impairs wtAAV2 localization and transcription, highlighting the importance of their function in the non-replicative wtAAV2 life cycle. KLF4 binding sites are sufficient to improve the expression of transgenes from rAAV vectors and increase their association with cellular DNA damage response proteins. This study advances our understanding of wtAAV2-host interactions and opens new avenues for improving rAAV gene therapy platforms.

ABSTRACT Recent research clarified the genetic identity of the ten nominal species of Amazonian Bolitoglossa (subgenus Eladinea) and proposed a remarkable number of undescribed candidate species in this group of salamanders. Here, we provide a genetic analysis of Amazonian Bolitoglossa species using available GenBank data and new DNA sequences of the mitochondrial genes 16S rRNA and cytochrome b, and the nuclear-encoded genes RAG-1 and POMC. Justified by substantial genetic differentiation (uncorrected genetic distance to all other nominal species > 4% for 16S and ≥ 12% for COB), lack of haplotype sharing in nuclear genes, and morphological differences, we describe a candidate species occurring at various sites in Amazonian Peru as Bolitoglossa chrysothyma sp. nov. After about 50 years of zoological research at Panguana, the type locality of the new species, it was observed by researchers for the first time in 2007, suggesting a recent colonization of the area from the slopes of the adjacent Serranía de Sira. When threatened, some individuals displayed a coiled antipredator posture which we speculate could represent a bird dropping mimesis. Aspects of species delimitation in Amazonian Bolitoglossa, their biogeography, possible temporal faunal changes at Panguana, and current threats are discussed. https://zoobank.org/urn:lsid:zoobank.org:pub:FE753BAF-437A-4F00-8FF2-E4BA5E6677DE

The Neogene fossil record of predatory mammals indicates periodic dispersals across the Beringian land bridge. Among the documented immigrant species in the weasel family (Mustelidae), Cernictis hesperus is among the most poorly known genotype species because of the highly fragmentary holotype. Here I report a new specimen representing the species from its type locality in the Pinole Tuff Formation, California. Associated left and right dentaries show that C. hesperus possesses a first premolar, unlike all other known species of the genus. Furthermore, incisor alveoli indicate the presence of a full incisor toothrow that is similarly crowded compared to the cheek teeth. These new morphological characteristics improve the diagnosis of the genus and species for the first time in nearly a century since the initial publication on the taxon.