The draft genome of the Temminck's tragopan (Tragopan temminckii) with evolutionary implications.

BACKGROUND: High-quality genome data of birds play a significant role in the systematic study of their origin and adaptive evolution. The Temminck's tragopan (Tragopan temminckii) (Galliformes, Phasianidae), a larger pheasant, is one of the most abundant and widely distributed species of the genus Tragopan, and was defined as class II of the list of national key protected wild animals in China. The absence of a sequenced genome has restricted previous evolutionary trait studies of this taxa. RESULTS: The whole genome of the Temminck's tragopan was sequenced using Illumina and PacBio platform, and then de novo assembled and annotated. The genome size was 1.06 Gb, with a contig N50 of 4.17 Mb. A total of 117.22 Mb (11.00%) repeat sequences were identified. 16,414 genes were predicted using three methods, with 16,099 (98.08%) annotated as functional genes based on five databases. In addition, comparative genome analyses were conducted across 12 Galliformes species. The results indicated that T. temminckii was the first species to branch off from the clade containing Lophura nycthemera, Phasianus colchicus, Chrysolophus pictus, Syrmaticus mikado, Perdix hodgsoniae, and Meleagris gallopavo, with a corresponding divergence time of 31.43 million years ago (MYA). Expanded gene families associated with immune response and energy metabolism were identified. Genes and pathways associated with plumage color and feather development, immune response, and energy metabolism were found in the list of positively selected genes (PSGs). CONCLUSIONS: A genome draft of the Temminck's tragopan was reported, genome feature and comparative genome analysis were described, and genes and pathways related to plumage color and feather development, immune response, and energy metabolism were identified. The genomic data of the Temminck's tragopan considerably contribute to the genome evolution and phylogeny of the genus Tragopan and the whole Galliformes species underlying ecological adaptation strategies.

Critical Role of Simultaneous Reduction of Atmospheric Odd Oxygen for Winter Haze Mitigation.

The lockdown due to COVID-19 created a rare opportunity to examine the nonlinear responses of secondary aerosols, which are formed through atmospheric oxidation of gaseous precursors, to intensive precursor emission reductions. Based on unique observational data sets from six supersites in eastern China during 2019-2021, we found that the lockdown caused considerable decreases (32-61%) in different secondary aerosol components in the study region because of similar-degree precursor reductions. However, due to insufficient combustion-related volatile organic compound (VOC) reduction, odd oxygen (Ox = O3 + NO2) concentration, an indicator of the extent of photochemical processing, showed little change and did not promote more decreases in secondary aerosols. We also found that the Chinese provinces and international cities that experienced reduced Ox during the lockdown usually gained a greater simultaneous PM2.5 decrease than other provinces and cities with an increased Ox. Therefore, we argue that strict VOC control in winter, which has been largely ignored so far, is critical in future policies to mitigate winter haze more efficiently by reducing Ox simultaneously.

Mitochondrial Genomes of Two Pygmy Grasshoppers (Orthoptera: Tetrigoidea) and a Comparative Analysis of Caelifera Mitogenomes.

The nearly complete mitogenomes of Bolivaritettix sikkinensis and Bolivaritettix yuanbaoshanensis were determined and compared with the mitogenomes of other 29 caeliferans. These two newly determined mitogenomes displayed the typical gene organization and composition, codon usage and AT skew of most Caelifera mitogenomes. A comparison of 31 orthopteran sequences revealed that the main causes of the differences in total length are an A+T-rich region and the length of IGS, even the rRNAs, and an AT content proportional to the value of AT-skew in the third position of PCGs. In Tetrigoidea, several tRNA were found to possess smaller TΨC or DHC loops and a number of special and conserved regions or motifs were found in tRNA genes and IGS, which may be useful as a molecular synapomorphy for this genus. ML and BI methods used in phylogenetic analysis revealed that Acridoidea species may have a close relationship with Tetrigoidea species.

Taxonomic status and phylogenetic relationship of tits based on mitogenomes and nuclear segments.

The phylogeny of tits has been studied using various molecular markers, but their phylogenetic relationships remain controversial. To further investigate their taxonomic status and phylogenetic relationships, the entire mitochondrial genomes (mitogenomes) and five nuclear segments were sequenced from 10 species of tits and two outgroups (Sylviparus modestus and Remiz consobrinus), followed by the comparison of mitogenomic characteristics and reconstruction of phylogenetic relationship based on the different datasets. The results revealed the following: the mitogenomes of 10 ingroup tits, each 16,758-16,799bp in length, displayed typical mitogenome organization and the gene order found in most previously determined Passeriformes mitogenomes; close relationships existed between Parus major and P. monticolus, between P. montanus and P. palustris, and between P. ater and P. venustulus; and Pseudopodoces humilis was a sister group to P. spilonotus, P. cyanus, or the clade containing P. major and P. monticolus.

Transcriptome of the pygmy grasshopper Formosatettix qinlingensis (Orthoptera: Tetrigidae).

Formosatettix qinlingensis (Zheng, 1982) is a tiny grasshopper endemic to Qinling in China. For further study of its transcriptomic features, we obtained RNA-Seq data by Illumina HiSeq X Ten sequencing platform. Firstly, transcriptomic analysis showed that transcriptome read numbers of two female and one male samples were 25,043,314, 24,429,905, and 25,034,457, respectively. We assembled 65,977 unigenes, their average length was 1,072.09 bp, and the length of N50 was 2,031 bp. The average lengths of F. qinlingensis female and male unigenes were 911.30 bp, and 941.82 bp, and the N50 lengths were 1,745 bp and 1,735 bp, respectively. Eight databases were used to annotate the functions of unigenes, and 23,268 functional unigenes were obtained. Besides, we also studied the body color, immunity and insecticide resistance of F. qinlingensis. Thirty-nine pigment-related genes were annotated. Some immunity genes and signaling pathways were found, such as JAK-STAT and Toll-LIKE receptor signaling pathways. There are also some insecticide resistance genes and signal pathways, like nAChR, GST and DDT. Further, some of these genes were differentially expressed in female and male samples, including pigment, immunity and insecticide resistance. The transcriptomic study of F. qinlingensis will provide data reference for gene prediction and molecular expression study of other Tetrigidae species in the future. Differential genetic screening of males and females provides a basis for studying sex and immune balance in insects.

Review of the genus Formosatettix Tinkham (Orthoptera: Tetrigidae) with description of new taxa, new synonymy and an updated key to species.

The genus Formosatettix Tinkham is reviewed. Seven new species from China, Formosatettix cliva Deng, sp. nov., Formosatettix guangyuanensis Deng, sp. nov., Formosatettix shuimogouensis Deng, sp. nov., Formosatettix strictivertex Deng, sp. nov., Formosatettix tangjiaheensis Deng, sp. nov., Formosatettix yueqingensis Deng, sp. nov., Formosatettix zheminzhengi Deng, sp. nov. are described and illustrated. One new name is proposed: Formosatettix latifemurus Deng, nom. nov. One new combination is established: Formosatettix nyalamensis (Zheng & Lin, 2015), comb. nov. The following new synonyms are established: Bolivaritettix circocephalus Zheng, 1992 = Formosatettix torulosinota Zheng & Mao, 2002, syn. nov., Criotettix bispinosus (Dalman, 1818) = Formosatettix hainanensis Zheng, 2012, syn. nov., Epitettix guangxiensis (Zheng & Jiang, 1994) = Formosatettix guangxiensis Zheng & Jiang, 1998, syn. nov., Formosatettix longwangshanensis Zheng, 1998 = Formosatettix tianmushanensis Zheng & Li, 2001, syn. nov., Formosatettix serrifemora Deng, 2019 = Formosatettix wulongensis Zha & Ding, 2020, syn. nov., Formosatettix huapingensis Zheng & Jiang, 1997 = Formosatettix nanlingensis Zheng & Cao, 2011, syn. nov. = Formosatettix undulatifemura, Zheng, 2012, syn. nov. = Formosatettix guposhanensis Deng, 2019, syn. nov. In addition, Formosatettix leigongshanensis Zha & Ding, 2020 is briefly commented.

Twenty-three new synonyms of the Eastern common groundhopper, Tetrixjaponica (Bolívar, 1887) (Orthoptera, Tetrigidae).

The Eastern common groundhopper, Tetrixjaponica, is a pygmy grasshopper species widely distributed in the Eastern Palearctic region, and shows a high degree of phenotypic variation. The classification of Tetrixjaponica is difficult and frequently involved errors. Among the many species of Tetrigidae that have been described in China within the last decades, many synonyms of Tetrixjaponica were found. The type specimens of many species deposited in the Chinese museums have been re-examined and as a result, Tetrixjaponica is systematically revised. Based on the results of this review, 23 new synonyms of Tetrixjaponica are proposed: Coptotettixcircinihumerus Zheng & Deng, 2004, syn. nov.; Coptotettixemeiensis Zheng, Lin & Zhang, 2012, syn. nov.; Euparatettixrongshuiensis Zheng, 2005, syn. nov.; Euparatettixzayuensis Zheng, Zeng & Ou, 2011, syn. nov.; Macromotettixnigritubercle Zheng & Jiang, 2006, syn. nov.; Macromotettixyaoshanensis Zheng & Jiang, 2000, syn. nov.; Tetrixalbistriatus Yao & Zheng, 2006, syn. nov.; Tetrixalbomaculatus Zheng & Jiang, 2006, syn. nov.; Tetrixalbomarginis Zheng & Nie, 2005, syn. nov.; Tetrixcenwanglaoshana Zheng, Jiang & Liu, 2005, syn. nov.; Tetrixcliva Zheng & Deng, 2004, syn. nov.; Tetrixduolunensis Zheng, 1996, syn. nov.; Tetrixgrossovalva Zheng, 1994, syn. nov.; Tetrixjiuwanshanensis Zheng, 2005, syn. nov.; Tetrixlatipalpa Cao & Zheng, 2011, syn. nov.; Tetrixliuwanshanensis Deng, Zheng & Wei, 2007, syn. nov.; Tetrixqinlingensis Zheng, Huo & Zhang, 2000, syn. nov.; Tetrixrectimargina Zheng & Jiang, 2004, syn. nov.; Tetrixruyuanensis Liang, 1998, syn. nov.; Tetrixxianensis Zheng, 1996, syn. nov.; Tetrixxinchengensis Deng, Zheng & Wei, 2007, syn. nov.; Tetrixyunlongensis Zheng & Mao, 2002, syn. nov.; Tetrixzhoushanensis Gao, Liu & Yin, 2022, syn. nov. It is expected that there will be the discoveries of more synonyms of this and other Tetriginae species from the Eastern Palearctic.

A de novo assembled genome of the Tibetan Partridge (Perdix hodgsoniae) and its high-altitude adaptation.

The Tibetan Partridge (Perdix hodgsoniae) is an endemic species distributed in high-altitude areas of 3600-5600 m on the Qinghai-Tibet Plateau. To explore how the species is adapted to the high elevation environment, we assembled a draft genome based on both the Illumina and PacBio sequencing platforms with its population genetics and genomics analysis. In total, 134.74 Gb short reads and 30.81 Gb long reads raw data were generated. The 1.05-Gb assembled genome had a contig N50 of 4.56 Mb, with 91.94% complete BUSCOs. The 17 457 genes were annotated, and 11.35% of the genome was composed of repeat sequences. The phylogenetic tree showed that P. hodgsoniae was located at the basal position of the clade, including Golden Pheasant (Chrysolophus pictus), Common Pheasant (Phasianus colchicus), and Mikado Pheasant (Syrmaticus mikado). We found that 1014, 2595, and 2732 of the 6641 one-to-one orthologous genes were under positive selection in P. hodgsoniae, detected using PAML, BUSTED, and aBSREL programs, respectively, of which 965 genes were common under positive selection with 3 different programs. Several positively selected genes and immunity pathways relevant to high-altitude adaptation were detected. Gene family evolution showed that 99 gene families experienced significant expansion events, while 6 gene families were under contraction. The total number of olfactory receptor genes was relatively low in P. hodgsoniae. Genomic data provide an important resource for a further study on the evolutionary history of P. hodgsoniae, which provides a new insight into its high-altitude adaptation mechanisms.

Pygmy grasshoppers of the genus Xistra Bolvar, 1887 (Orthoptera: Tetrigidae: Metrodorinae).

The genus Xistra Bolvar is reviewed. Two new species from China, Xistra hainanensis Deng, sp. nov. and Xistra nigriabdominis Deng, sp. nov. are described and illustrated. Three new combinations are established: Xistra emeiensis (Zheng, 1998), comb. nov., Xistra guangxiensis (Zheng, 1998), comb. nov. and Xistra lochengensis (Zheng, 2005), comb. nov. Four new synonyms are established: Bannatettix ruiliensis Zheng, 1993 = Xistra klinnema Zheng Zeng, 2011, syn. nov., Systolederus emeiensis Zheng, 1998 = Xistra yaanensis Zheng, 2009, syn. nov., Systolederus guangxiensis Zheng, 1998 = Xistra longzhouensis Zheng Jiang, 1998, syn. nov., Mazarredia lochengensis Zheng, 2005 = Xistra oculata Li, Deng Zheng, 2014, syn. nov. In addition, an updated key to all species of the genus Xistra is given.

Comparative mitogenomes and phylogenetic analysis reveal taxonomic relationship of genera Teredorus and Systolederus (Orthoptera, Tetrigoidea).

The genera Teredorus and Systolederus belong to Tetriginae and Metrodorinae respectively. However, species within these two genera have strikingly similar features, made it difficult to identify clearly by morphological characteristics. In this study, we sequenced the mitochondrial genomes (mitogenomes) of two Teredorus species, and compared them with Systolederus mitochondrial sequences. The sequenced mitogenomes of T. hainanensis and T. bashanensis are 14,946 bp and 14,775 bp in size, respectively. The A+T content of mitogenomes is 76.2% (T. hainanensis) and 74.0% (T. bashanensis). Comparative analysis showed that mitochondrial sequences and structure were similar within these two genera. The results of K2P distances and phylogenetic analysis revealed that Systolederus and Teredorus might be likely considered as one genus of Teredorus. It will provide important resources for further understanding of the taxonomy and phylogenetic relationship of Systolederus and Teredorus.

Mitogenomic characterization and phylogeny of Scelimena melli Günther (Orthoptera: Tetrigoidea: Scelimeninae).

The mitochondrial genome (mitogenome) of Scelimena melli, which belongs to Orthoptera, Tetrigoidea, Tetrigidae, Scelimeninae was determined. The mitogenome has a length of 14,598 bp and consists of 37 genes including 13 protein-coding genes (PCGs), two rRNA genes, and 22 tRNA genes. Phylogenetic analysis using 37 mitochondrial genes with other 22 Tetrigoidea species revealed that S. melli had a closer relationship with Paragavialidium sichuanense, but the monophyly of Scelimeninae was not recovered. The mitogenome data of S. melli would provide useful resources for further evolutionary studies of Scelimeninae and Tetrigoidea species.

Molecular phylogeny and species delimitation of the genus Tonkinacris (Orthoptera, Acrididae, Melanoplinae) from China.

Tonkinacris is a small group in Acrididae. While a few species were occasionally sampled in some previous molecular studies, there is no revisionary research devoted to the genus. In this study, we explored the phylogeny of and the relationships among Chinese species of the genus Tonkinacris using the mitochondrial COI barcode and the complete sequences of ITS1 and ITS2 of the nuclear ribosomal DNA. The phylogeny was reconstructed in maximum likelihood and Bayesian inference frameworks, respectively. The overlap range between intraspecific variation and interspecific divergence was assessed via K2P distances. Species boundaries were delimitated using phylogenetic species concept, NJ tree, K2P distance, the statistical parsimony network as well as the GMYC model. The results demonstrate that the Chinese Tonkinacris species is a monophyletic group and the phylogenetic relationship among them is (T. sinensis, (T. meridionalis, (T. decoratus, T. damingshanus))). While T. sinensis, T. meridionalis and T. decoratus were confirmed being good independent species strongly supported by both morphological and molecular evidences, the validity of T. damingshanus was not perfectly supported by molecular evidence in this study.

The De Novo Genome Sequencing of Silver Pheasant (Lophura nycthemera).

Silver pheasant (Lophura nycthemera) belongs to Phasianidae, Galliformes, which exhibits high subspecific differentiation. In this study, we assembled a novel genome based on 98.42 Gb of Illumina sequencing data and 30.20 Gb of PacBio sequencing data. The size of the final assembled genome was 1.01 Gb, with a contig N50 of 6.96 Mb. Illumina paired-end reads (94.96%) were remapped to the contigs. The assemble genome shows high completeness, with a complete BUSCO score of 92.35% using the avian data set. A total of 16,747 genes were predicted from the generated assembly, and 16,486 (98.44%) of the genes were annotated. The average length of genes, exons, and introns were 19,827.53, 233.69, and 1841.19 bp, respectively. Noncoding RNAs included 208 miRNAs, 40 rRNAs, and 264 tRNAs, and a total of 189 pseudogenes were identified; 116.31 Mb (11.47%) of the genome consisted of repeat sequences, with the greatest proportion of LINEs. This assembled genome provides a valuable reference genome for further studies on the evolutionary history and conversion genetics of L. nycthemera and the phylogenomics of the Galliformes lineage.

The first mitogenome of Lesser Whitethroat, Sylvia curruca blythi Ticehurst & Whistler, 1933 (Passeriformes: Sylviidae) and its phylogenetic implications for the genus Sylvia.

The complete mitogenome of the Lesser Whitethroat, Sylvia curruca blythi Ticehurst & Whistler, 1933 was determined, which belongs to Sylviidae, Passeriformes. The mitogenome had a length of 17,959 bp and consisted of 37 genes including 13 PCGs, 2 ribosomal RNAs (rRNA) genes, and 22 transfer RNAs (tRNA) genes. In addition, two control regions (CRs) were also existed in the mitogenome, with Sylvioidea typcial gene arrangement of cytb-trnT-CR1-trnP-nad6-trnE-CR2-trnF-rrnS. Phylogenetic analysis using 37 mitochondrial genes of 17 related species revealed that S. c. blythi had a closer relationship with S. crassirostris, and the monophyly of Sylvia was also recovered. The mitogenome data of S. c. blythi would provide useful resources for further studying the evolution of Sylvia and the subspecies taxonomic revision of S. curruca intraspecific.

Correction: Molecular phylogeny and species delimitation of the genus Tonkinacris (Orthoptera, Acrididae, Melanoplinae) from China.

[This corrects the article DOI: 10.1371/journal.pone.0249431.].

Biomass-burning emissions could significantly enhance the atmospheric oxidizing capacity in continental air pollution.

Volatile organic compounds (VOCs) are important precursors of photochemical pollution. However, a substantial fraction of VOCs, namely, oxygenated VOCs (OVOCs), have not been sufficiently characterized to evaluate their sources in air pollution in China. In this study, a total of 119 VOCs, including 60 OVOCs in particular, were monitored to provide a more comprehensive picture based on different online measurement techniques, proton-transfer-reaction time-of-flight mass spectrometry (PTR-ToF-MS) and online gas chromatography/mass spectrometry (GC/MS), at a receptor site in southeastern China during a photochemically active period. Positive matrix factorization (PMF) and photochemical age-based parameterization were combined to identify and quantify different sources of major VOCs during daytime hours, with the advantage of including VOC decay processes. The results revealed the unexpected role of biomass burning (21%) in terms of ozone (O3) formation potential (OFP) when including the contributions of OVOCs and large contributions (30-32%) of biomass burning to aldehydes, as more OVOCs were measured in this study. We argue that biomass burning could significantly enhance the continental atmospheric oxidizing capacity, in addition to the well-recognized contributions of primary pollutants, which should be seriously considered in photochemical models and air pollution control strategies.

Reverse contralateral proximal tibial plating and cannulated screws fixation for Hoffa fracture: A case report.

Hoffa fracture is a rare type of distal femoral fracture occurs in the coronal plane of either femoral epicondyle. To date, screws in combination with lateral plate fixation is widely accepted to achieve stable fixation and good results. However, up to now there has not been a specially designed anatomical plate for lateral fixation of Hoffa fracture. In this report, we demonstrate a case of Hoffa fracture fixed with reverse application of "L" shaped contralateral proximal tibia plate and cannulated screws, resulting in good one-year results.

Phylogeny and species delimitation of the genus Longgenacris and Fruhstorferiola viridifemorata species group (Orthoptera: Acrididae: Melanoplinae) based on molecular evidence.

Phylogenetic positions of the genus Longgenacris and one of its members, i.e. L. rufiantennus are controversial. The species boundaries within both of L. rufiantennus+Fruhstorferiola tonkinensis and F. viridifemorata species groups are unclear. In this study, we explored the phylogenetic positions of the genus Longgenacris and the species L. rufiantennus and the relationships among F. viridifemorata group based on the 658-base fragment of the mitochondrial gene cytochrome c oxidase subunit I (COI) barcode and the complete sequences of the internal transcribed spacer regions (ITS1 and ITS2) of the nuclear ribosomal DNA. The phylogenies were reconstructed in maximum likelihood framework using IQ-TREE. K2P distances were used to assess the overlap range between intraspecific variation and interspecific divergence. Phylogenetic species concept and NJ tree, K2P distance, the statistical parsimony network as well as the generalized mixed Yule coalescent model (GMYC) were employed to delimitate the species boundaries in L. rufiantennus+F. tonkinensis and F. viridifemorata species groups. The results demonstrated that the genus Longgenacris should be placed in the subfamily Melanoplinae but not Catantopinae, and L. rufiantennus should be a member of the genus Fruhstorferiola but not Longgenacris. Species boundary delimitation confirmed the presence of oversplitting in L. rufiantennus+F. tonkinensis and F. viridifemorata species groups and suggested that each group should be treated as a single species.

Exploration of sources of OVOCs in various atmospheres in southern China.

Oxygenated volatile organic compounds (OVOCs) are critical atmospheric ozone and secondary organic aerosol (SOA) precursors and radical sources, while understanding of OVOC sources in the atmosphere, especially with large anthropogenic emissions, still has large uncertainties. A high-sensitivity proton transfer reaction mass spectrometer (PTR-MS) was deployed in vastly different atmospheres in southern China, including an urban site (SZ-U), a regional site (NA-R), and a background site (NL-B). Four critical OVOCs, i.e., methanol, acetone, methyl ethyl ketone (MEK) and acetaldehyde, five groups of aromatic hydrocarbons, isoprene and acetonitrile were measured with a high time resolution. The featured relative abundance and diurnal variations of the OVOCs indicated that methanol, acetone and MEK had prominent contributions from urban industrial activities, while acetaldehyde was closely related to the photochemical formation at all three sites. The photochemical age-based parameterization method was improved locally and then applied to quantify different sources of daytime OVOCs: anthropogenic secondary and biogenic sources (together 60-73%) were always the dominant source for acetaldehyde in various atmospheres; in addition to a significant background for methanol, acetone and MEK, anthropogenic primary emissions (mostly industrial) were their dominant source at SZ-U (38-73%), while biogenic sources played the key role for them at NL-B (30-43%); biomass burning contributed a small fraction of 5-17% for the four OVOCs at the three sites.

Phylogenetic relationships of Scelimeninae genera (Orthoptera: Tetrigoidea) based on COI, 16S rRNA and 18S rRNA gene sequences.

Scelimeninae is an important subfamily of Tetrigoidea; however, the phylogenetic relationships within Scelimeninae are poorly understood, and its generic classification has remained unstable. In this study, the COI, 16S rRNA and 18S rRNA genes from 24 species in 9 genera within Scelimeninae were amplified and sequenced, the base composition and inter-species genetic distance of the combined sequence of COI, 16S rRNA and 18S rRNA genes were analyzed, and the molecular phylogenetic relationships were reconstructed using Maximum Likelihood (ML) and Bayesian inference (BI) methods. The results of sequence analysis showed that the total length of the combined COI, 16S rRNA and 18S rRNA gene sequence was 3507 bp, including 2345 conservative sites, 1144 variable sites and 901 parsimony-informative sites. The average A+T content was 63.5% and 78.1% in the COI, 16S rRNA sequences, respectively, indicating A+T bias. The average genetic distance between all species was 0.134, and the average genetic distance in the inner group (Scelimeninae) was 0.126. A phylogenetic tree based on the combined sequences of the COI, 16S rRNA and 18S rRNA genes showed that the phylogenetic relationships among 9 Scelimeninae genera were as follows: Criotettix + (((Zhengitettix + Hebarditettix) + (Falconius + (Scelimena + Paragavialidium))) + ((Eucriotettix + Thoradonta) + Loxilobus)). The molecular phylogenetic results generally support the morphological taxonomy; at the genus level, Criotettix, Scelimena, Paragavialidium, Thoradonta and Eucriotettix are monophyletic groups, Scelimena and Paragavialidium form sister groups, and Thoradonta and Eucriotettix also form sister groups, but the relationship between Hebarditettix and Zhengitettix needs further study. At the species level, synonyms may exist between Thoradonta spiculoba and Thoradonta transpicula and Thoradonta nodulosa and Thoradonta obtusilobata, but more studies are required to confirm this inference.