Description of a New Species of Hainania Koller (Teleostei, Cypriniformes, Xenocyprididae) from Guangdong Province, Southern China †
1
State Key Laboratory of Agricultural and Forestry Biosecurity, College of Plant Protection, China Agricultural University, Beijing 100093, China
2
Department of Entomology, College of Plant Protection, China Agricultural University, Beijing 100193, China
3
College of Resources and Environmental Sciences, China Agricultural University, Beijing 100193, China
*
Author to whom correspondence should be addressed.
†
urn:lsid:zoobank.org:act:70B17170-4C76-48CC-8D88-5EAA0BD18CC6.
Diversity 2025, 17(8), 549; https://doi.org/10.3390/d17080549 (registering DOI)
Submission received: 25 June 2025
/
Revised: 23 July 2025
/
Accepted: 24 July 2025
/
Published: 1 August 2025
(This article belongs to the Special Issue Evolution, Systematic and Conservation of Freshwater Fishes)
Abstract
Hainania Koller (Teleostei, Cypriniformes, Xenocyprididae) is known as a monotypic genus of sharpbelly fish that is endemic to Hainan Island, China. We describe Ha. minzhengi sp. nov., the second species of Hainania collected from Guangdong, based on morphology and molecular evidence. Phylogenetic relationships were inferred based on the mitochondrial cytochrome b (Cyt b) gene and cytochrome oxidase subunit 1 (COI) gene, by maximum likelihood and Bayesian methods and different partitioning schemes. Our result supports the sister group relationship between Ha. serrata and Ha. minzhengi sp. nov., but the monophyly of Pseudohemiculter or Hemiculterella was not recovered. A diagnostic key to Chinese species of Hainania and Pseudohemiculter is provided.
1. Introduction
The Hainanese sharpbelly genus Hainania Koller is currently represented by a single species, Ha. serrata, a small pelagic fish inhabiting the upper and middle stretches of rivers on Hainan Island, China. Hainania is characterized by a lateral line abruptly bent behind pectoral fins, running low in concurrence with the ventral profile of the body, rising again over the terminal of the anal-fin base and extending along the middle of the caudal peduncle, an incomplete ventral keel, present only from the anus to the pelvic fin, and last a simple dorsal fin ray, large and hard, with mild serrations along its posterior margin. Hainania was described by Koller in July, 1927 [1] based on specimens from “Kang-kongflusse” (possibly Changjiang River), a river in Hainan Island. Later in the same year (September), Nichols et al. [2] described another Hainanese xenocypridid species with a weakly serrated dorsal spine, Hemiculter serracanthus, from Nodoa, and established a new subgenus Pseudohemiculter to accommodate Hemiculter spp. with an incomplete ventral keel, namely He. hainanensis, He. dispar, and He. serracanthus, with He. hainanensis as the type species. Myers (1931) recognized He. serracanthus as a junior synonym of Ha. serrata and treated Hainania as a subgenus of Hemiculter, thus altering the scientific name of this species to He. (Hainania) serratus [3]. This arrangement was followed by subsequent authors [4], until Wu suggested that the well-developed and weakly serrated dorsal fin spine of He. (Hainania) serratus is unique among congeneric species and valid as a generic diagnostic character, thus restoring the validity of Hainania as a distinct genus in 1964 [5]. The validity of Hainania was widely accepted by the following research [6,7,8,9,10,11,12,13]. The majority of the research treated Hainania as an endemic genus of Hainan Island, China [5,6,7,8,12], but Hainania was also reported from Thailand [10,11] and Vietnam [14].
Although morphological similarity between Hainania and Pseudohemiculter has been addressed by multiple studies [2,5,6,8,13], the phylogenetic position of Hainania within Xenocyprididae remained unresolved. Tang et al. [10] reconstructed a phylogeny of Xenocyprididae (referred as subfamily Oxygastrinae) with four molecular markers, namely mitochondrial Cyt b and COI as well as exon 3 of nuclear RAG1 and opsin (rhodopsin), and placed Ha. serrata within Toxabramis. Cheng et al. [11] utilized part of the Hainania sequences generated in [10] and came up with a congruent conclusion. Deng et al. [12] conducted a phylogenetic analysis in a smaller group (the East Asian hemicultrine fishes) with five target loci, Cyt b, COI, as well as nuclear EGR1, EGR2B, and RH. In this phylogeny, Hainania nested within two Pseudohemiculter species, P. dispar and P. hainanensis [12].
Through an ichthyological survey conducted on the Jianjiang River, an independent coastal river in Guangdong Province of southern China, xenocypridid specimens with a weakly serrated dorsal fin spine, a typical diagnostic feature of Hainania, were obtained. Through intensive morphological and molecular analysis, this population was confirmed as a new species of Hainania, hereby described as Hainania minzhengi sp. nov. A diagnostic key to Chinese species of Hainania and Pseudohemiculter is provided.
2. Materials and Methods
2.1. Sample Collection and Taxon Sampling
Twenty-three specimens of Hainania minzhengi sp. nov. were collected in July 2024 from the Xijianghe River in Shuikou Town, Xinyi City, Maoming City, Guangdong Province, China. Fish were collected through drift nets and angling or bought from local market, photographed with a Canon 5D mark III digital camera (Canon Co., Ltd., Beijing, China) in an aquarium, and then euthanized with eugenol. Right fin tissues of fresh specimens were clipped (n = 13) and preserved in 95% ethanol and stored in a −20 °C freezer before DNA extraction. Specimens were fixed in 5% formaldehyde for 5–7 days and then stored in 75% ethanol for permanent preservation. Type specimens of Hainania minzhengi sp. nov. are deposited either in the Museum of Aquatic Organisms at the Institute of Hydrobiology, Chinese Academy of Sciences (IHB) (n = 20) and the Institute of Zoology, Chinese Academy of Sciences, Beijing, China (ASIZB) (n = 3). All care and use of experimental animals complied with the relevant laws of the Chinese Laboratory of Animal Welfare and Ethics (GB/T 35892–2018). Specimens of Ha. serrata (n = 27), Pseudohemiculter dispar (n = 4), P. hainanensis (n = 4), P. kweichowensis (n = 1), and Hemiculterella wui (n = 1) were collected for morphological and molecular analysis. These specimens are deposited at IHB. Sampling sites of all specimens are illustrated in Figure 1.
Three species of Pseudohemiculter, two species of Hemiculterella, two species of Hemiculter, and two species of Toxabramis were selected as closely related outgroups based on previous phylogeny [13], and Ctenopharyngodon idella was included as a far-related outgroup. Outgroup sequences are either newly generated or obtained from GenBank.
2.2. Morphometric Data Acquiring
Morphometrics were taken point-to-point with Gemred DCB150 digital calipers (Gemred Co., Ltd., Guangxi, China) directly connected to a computer and data was recorded to the nearest 0.1 mm. Specimens with a heavily damaged head or caudal fin were excluded from the measurements. In the end, 19 individuals are used. Measurements and counts were made on the left side of each specimen, following Wu [5] and Chen [6]. All measurements are given as proportions of standard length (SL) or lateral head length (HL). Counts of fin rays and lateral line scales were taken under a ZEISS Stemi 508 stereoscopic microscope (ZEISS Group, Oberkochen, Germany). Measurements and meristic counts were made on the left side whenever possible. One specimen was dissected to examine the pharyngeal teeth. Given the prominent morphological difference in head shape observed, all measurements of the head (highlighted in bold in table in below) of two Hainania species were analyzed by using principal component analysis (PCA). Measurements were initially converted to the percentage of head length (HL) or standard length (SL) before all statistical procedures. Dimensionality reduction was conducted through the function PercentScaler. Based on the covariance matrix of log-transformed statistics, a principal component analysis was conducted by using Past 2.17 (Hammer et al.) [15]. Scatterplots and linear graphs of the PCA loadings on both principle components of each individual were made to visualize between-group differences. All statistical tests were performed using R 4.3.3 (https://www.R-project.org/) and R-packages ggplot2 (https://ggplot2.tidyverse.org) and ggpmisc (https://github.com/aphalo/ggpmisc.).
2.3. DNA Extraction, Amplification, Sequencing, and Phylogenetic Analysis
Total genomic DNA was extracted from the pelvic fin using the TIANamp Genomic DNA Kit (TIANGEN Co., Ltd., Beijing, China), following the standard protocols. The mitochondrial cytochrome b (Cyt b) gene and cytochrome oxidase subunit 1 (COI) gene of selected individuals were amplified by adopting the procedures and primers of Cheng et al. [11]. DNA sequencing was performed on both strands by Beijing Tsingke Biotech Co., Ltd. (Beijing, China). The sequencing results were assembled using SeqMan 7.1.0 (DNAstar lnc., Madison, WI, USA), and other sequences were downloaded from the NCBI database. Newly generated sequences were deposited in GenBank under the accession numbers given in Table 1. Sequences were aligned in MAFFT v.7.245 (Katoh et al.) [16] and manually examined in AliView (Larsson) [17] to ensure the correct reading frame. Alignments were concatenated using Phyx v.1.1 (Brown et al.) [18] to generate the final supermatix for phylogeny.
Phylogenetic trees were constructed under maximum likelihood (ML) methods and Bayesian inference (BI). Two partitioning schemes were used: (1) partition by gene (PGM), and (2) partition by gene and codon position (PCM). ML analyses were carried out using IQ-TREE v.2.1.3 (Nguyen et al.) [19]. Datasets were partitioned and model-tested in ModelFinder (Kalyaanamoorthy et al.) [20] as implemented in IQ-TREE. We found the best partition scheme after merging possible partitions (‘-MFP+MERGE’ command) and determining the best scheme under the Bayesian information criterion (BIC). The best-fitting models were used for phylogenetic reconstructions (‘-p’command). An initial 1000 parsimony trees were generated in IQ-TREE with the command ‘-ninit 1000’, and the 100 trees with the fewest steps were used to initialize the candidate set (-ntop 100), considering all possible nearest neighbor interchanges (-allnni). These 100 trees were maintained in the candidate set during the ML tree search (-nbest 100), and unsuccessful runs were terminated after 1000 iterations (-nstop 1000). Perturbation strength was set to 0.2 (-pers 0.2), as recommended for datasets with many short sequences. We applied nearest-neighbor interchange (NNI) branch swapping to improve the tree search and limit overestimating branch supports due to severe model violations (‘-bnni’ command). Node supports were computed with 1000 UFBoot (‘-B’ command) replicates (Minh et al. & Hoang et al.) [21,22] and SH-aLRT (‘-alrt’ command) (Guindon et al.) [23]. Trees were searched for 10 independent times (‘--runs’ command), and the best topology was selected based on the log-likelihood. BI analysis was performed using MrBayes 3.2.7 (Ronquist et al.) [24]. PartitionFinder v2.1.1 was used to assess the optimal partitioning strategy and substitution model using the “greedy” algorithm with branch lengths estimated as “unlinked” and BIC criterion (Lanfear et al.) [25]. Two independent runs were executed for 200 million generations, with sampling occurring every 1000 generations. Additionally, four independent Markov Chain Monte Carlo (MCMC) chains were employed, consisting of three heated chains and a cold chain, and the initial 30% of samples were discarded as burn-in. When the average standard deviation of split frequencies fell below 0.01, we considered that stationarity had been reached. The phylogenetic trees generated in this study were visualized using Figtreev1.4.4 (Rambaut) [26].
Branches were considered robustly supported if SH-aLRT ≥ 80 and UFBoot ≥ 95 for ML, pp ≥ 0.95 for BI; moderately supported if SH-aLRT ≥ 80 or UFBoot ≥ 95 for ML, 0.95 ≥ pp ≥ 0.8 for BI; and weakly supported if SH-aLRT < 80 and UFBoot < 95 for ML, pp < 0.8 for BI.
Pairwise genetic distances of combined Cyt b and COI genes amongst species of Hainania and closely related Pseudohemiculter were calculated in MEGA v.11 (Tamura et al.) [27] based on the Kimura 2-parameter (K2P) model (Kimura et al.) [28] with transitions + transversions substitutions, uniform rates, pairwise deletion, and selecting all codon positions.
3. Results
3.1. Phylogenetic Analysis
A total of 53 taxa were included in the phylogeny, representing 12 species (Figure 2). The supermatrix comprised 1755 aligned sites (COI: 624 bp and Cyt b: 1131 bp). Both phylogenetic methods (ML and BI) recovered highly consistent topology regardless of partitioning schemes.
The selected taxa of the Hemicultrine group formed a well-supported monophylum, consisting of two clades. Clade I consists of Hemiculterella, Hemiculter, Toxabramis, and Pseudohemiculter kweichowensis. Clade II accommodates two Hainania species and two Pseudohemiculter species, P. hainanensis and P. dispar. The monophyly of both Hainania species were strongly supported, and they were recovered sister to each other. Pseudohemiculter hainanensis is sister to Hainania, and the monophyly of Pseudohemiculter was not recovered, with Hainania nested into Pseudohemiculter (Figure 2). The minimum interspecific pairwise genetic distance based on combined Cyt b and COI genes between the new species and Ha. serrata (4.68%) is significantly greater than the maximum intraspecific pairwise genetic distance of Ha. serrata (1.85%) or Ha. minzhengi sp. nov. (0.46%) (Table 2). Pairwise genetic distance based on separated Cyt b and COI are provided in Supplementary Table S2.
3.2. Systematics
Family Xenocyprididae Günther, 1868
Hainania Koller, 1927
Diagnosis. A small-to-medium sized xenocypridid with lateral line abruptly bent behind pectoral fins, running low in concurrence with the ventral profile of the body, rising again over the terminal of the anal-fin base, and extending along the middle of the caudal peduncle (vs. lateral line without abrupt bend in all other xenocypridid except Anabarilius, Hemiculter, Hemiculterella, Pseudohemiculter, Pseudolaubuca, Siniichthys, and Toxabramis), an incomplete ventral keel, present only from the anus to the pelvic fin (vs. complete ventral keel in Hemiculter, Pseudolaubuca, Siniichthys, and Toxabramis), and a hard and finely serrated dorsal fin spine (vs. smooth dorsal fin spine in Anabarilius, Hemiculterella, and Pseudohemiculter).
Distribution. China (Hainan, Guangdong).
Species included. Hainania minzhengi sp. nov., Hainania serrata Koller, 1927
3.3. Key to Chinese Species of Hainania and Pseudohemiculter
| Hainania |
| Pseudohemiculter |
| Ha. mingzhengi sp. nov. |
| Ha. serrata |
| P. kweichowensis |
| 4 |
| P. dispar |
| P. hainanensis |
3.4. Taxonomic Description
Hainania minzhengi H.-T. Lei, Z.-Y. Gong and X.-K. Li, new species.
ZooBank. urn:lsid:zoobank.org:act:70B17170-4C76-48CC-8D88-5EAA0BD18CC6
Holotype. IHBMM202407001tp, 84.64 mm SL; Xijianghe River [西江河], a tributary of the Jianjiang River in Shuikou Town, Xinyi City, Maoming City, Guangdong Province, China (22.299332° N, 110.850702° E; 38 m elevation); collected by Minzheng Li, 4.VII.2024.
Paratypes. IHBMM202407002-IHBMM202407023, 19, 70.81−104.70 mm SL; ASIZB 248490-ASIZB 248492, 3, 88.39−110.91 mm SL; other data same as holotype.
Diagnosis. Hainania minzhengi can be distinguished from Ha. serrata in larger eyes, eye diameter larger than 30% HL vs. smaller than 30% HL in Ha. serrata; snout length shorter than eye diameter vs. equals or larger in Ha. serrata; pectoral fin extending slightly beyond halfway to pelvic fin origin vs. nearly reaching pelvic fin origin in Ha. serrata; fewer branched anal-fin rays, 13−14 vs. 15 in Ha. serrata; and fewer lateral line scales, 54−52 vs. 56−55 in Ha. serrata (Figure 3, Figure 4 and Figure 5).
Description. Morphometric measurements of 19 individuals of Ha. minzhengi are provided in Supplementary Table S1. Comparison of Ha. minzhengi with Ha. serrata and two closely related Pseudohemiculter species, P. dispar and P. hainanensis is shown in Table 3. See Figure 6 for a general profile of body. Body elongated, laterally compressed, covered in shiny, fragile scales. Body depth greatest before dorsal-fin origin, slightly declining towards caudal fin base. The greatest body depth typically fits more than four times in SL. The lateral line descends steeply downward, running low in concurrence with ventral profile of body, rising again over terminal of anal-fin base and extending along middle of caudal peduncle. Lateral line scales 52−54. Ventral keel incomplete, present only from the pelvic fins to anus. Head small, wedge shaped, length of which similar to body depth. Snout pointed, short, length of which smaller than eye diameter (Figure 6B). Mouth terminal, upper jaw with a tiny notch, providing space for a small symphyseal knob of the lower jaw. Terminal of oral fissure barely reaching anterior margin of eye. Eyes large, approximately one third of head length. No barbels present. Pharyngeal teeth 3-rowed, 5, 4, 2, with hooked tips. Dorsal fin with 3 simple and 7 branched fin rays, origin situated slightly behind mid-point between snout tip and caudal-fin base and slightly posterior to the pelvic fin origin, distal margin truncate. The last simple fin ray enlarged and hardened, with small, densely situated serration on its posterior margin, barely visible with naked eye as they are buried under a thin layer of skin. Pelvic fins with 1 simple and 6 (n = 1) or 7 (n = 18) branched fin rays, origin closer to anal-fin origin than pectoral fin origin. Pectoral fins with 1 simple and 13 (n = 14) or 14 (n = 4) branched fin rays, extending slightly beyond halfway to pelvic-fin origin. Anal fin with 3 simple and 13 (n = 8) or 14 (n = 11) branched fin rays. Caudal fin deeply forked, with 2 simple and 16 branched fin rays, the lower lobe somehow longer than the upper.
Coloration. In life: body silvery, dark greyish dorsally and white ventrally. Caudal fin yellowish with obscure dark margin. Dorsal fin spine grey, other fin rays, and interradial membrane hyaline. Other fins hyaline (Figure 7). In preservative: colors and gloss fade away, body and fins grey to creamy yellow (Figure 3 and Figure 6).
Distribution. This species is presently only known from the Xijianghe River, a tributary of the Jianjiang River in Shuikou Town, Xinyi City, Maoming City, Guangdong Province, China (Figure 8).
Etymology. The specific name “minzhengi” is in honor of Mr. Minzheng Li for his efforts in the specimen collection of this species. In addition, given that Ha. minzhengi is only distributed in Xinyi City, Maoming City, Guangdong Province, China, we propose “信宜海南䱗” (Pinyin: Xin Yi Hai Nan Can) as the Chinese common name of this species.
Field notes. Hainania minzhengi inhabits upper and middle stretches of river, often murky, with moderate velocity and substrate consisting of boulders, gravel, and sand (Figure 8). It is gregarious and feeds on various aquatic invertebrates and algae. Aggression and harmless combat behaviors between individuals were observed from individuals kept in an aquarium. Sympatric fish species found at the same locality include Garra orientailis (Cyprinidae), Squalidus atromaculatus (Gobionidae), Sinibrama melrosei, Opsariichthys hainanensis (Xenocyprididae), Tachysurus intermedius (Bagridae), and Mastacembelus armatus (Mastacembelidae). This species is also commonly captured and sold as food at the local market alongside the afore-mentioned species, accompanied by Traccatichthys tuberculum (Nemacheilidae) and Rhinogobius similis (Oxudercidae).
Morphometry. The results of the principal components analysis (PCA) revealed a separation between the head shape morphometrics of Ha. minzhengi, sp. nov. and Ha. serrata. Both principle components represent morphology. The contribution of the first principle (PC1) component accounted for 61.97% of the total variation, whereas the second principle (PC2) accounted for 19.86%. Overall, Ha. minzhengi, sp. nov. possesses a blunt head and large eyes, as well as a deep and short snout, whereas Ha. serrata has an elongated head shape, small eyes, and long snout. The PCA loadings are presented in Table 4, showing that the snout length, interorbital distance, and eye diameter contribute most to PC 1; head length, head depth, snout length, and interorbital distance contribute most to PC 2. In the factorial map, despite a slight overlap of 80%-equal frequency ellipses of the two species, the p-value of the analysis is 0.001 < 0.05, suggesting a well-supported division of the two species morphologically (Figure 9).
4. Discussion
Since its establishment in 1927, Hainania has been considered Hainan-endemic, except for a few exceptions [10,11,14]. Mai [14] provided the sole Vietnamese record but was reported as “Pseudohemiculter serrata”. We are unable to confirm the identity of Pseudohemiculter serrata due to the unavailability of Vietnamese literature or specimens. Tang et al. [10] and Cheng et al. [11] reconstructed phylogeny using sequences from a “Hainania” specimen collected from Thailand, but it was nested within Toxabramis. In the more recent phylogenetic analysis, Deng et al. [12] recovered a close relationship between Hainania and Pseudohemiculter, thus suggested the Thailand “Hainania” is most likely a misidentified Toxabramis species. Therefore, records of Hainania from Thailand and the Vietnamese remain to be confirmed.
The discovery of Ha. minzhengi sp. nov. confirms that Hainania is not a Hainanese-endemic genus, thus suggesting a possible diversity of Hainania in the coastal river systems of southern China and northern Vietnam. Intensive ichthyological surveys conducted in these regions are required to verify the distribution and diversity of Hainania.
Two Hainania species formed a well-supported monophylum based on the present phylogenetic analyses (Figure 2). The difference in habitat preference between the two Hainania species was observed, with Ha. serrata preferring swift-flowing mainstream stretches of river with a substrate consisting mainly of boulder and rock, in contrast to a preference for tributaries with slower water flow, murky water, and gravel substrate in Ha. minzhengi sp. nov. (Figure 8). The more acuminated head of Ha. serrata might be adaptations for a more torrential habitat. In addition, the development level of dorsal spine serration of Ha. serrata varied amongst different localities, while that of all examined individuals of Ha. minzhengi sp. nov. lay on the weaker end (Figure 4). The reason for such morphological diversity is yet unknown.
Hainania is nested within two Pseudohemiculter species, P. dispar and P. hainanensis, which is congruent with the previous study [12]. Both Hainania and Pseudohemiculter were described in 1927, and Hainania was published two months before Pseudohemiculter. Based on the phylogeny, Pseudohemiculter might be a junior synonym of Hainania, and/or multiple genera need to be established to reflect the natural groups. We decided not to make the change at this stage as the Vietnamese Pseudohemiculter species, P. pacboensis was not sampled in this phylogeny. Further research conducted upon a comprehensive collection of species and populations is necessary to elucidate the phylogenetic relationships and evolutionary history of Hainania, Pseudohemiculter, and their closely related taxa. Phylogenetic analysis is conducted based solely on two mitochondrial gene fragments, which could be unreliable in resolving taxonomic relationships beyond the species level. Utilization of nuclear loci in a larger phylogeny could provide robust support for the current study.
Monophyly of Hemiculter, Hemiculterella, Toxabramis, and Pseudohemiculter were not supported by the present phylogeny as well as in previous research [12,29]. The sister group relationship between P. kweichowensis and Hemiculterella sauvagei is reported for the first time. Both Pseudohemiculter and Hemiculterella are characterized by incomplete ventral keel and possession of hard dorsal fin spine is the key diagnostic character that differentiates Pseudohemiculter from Hemiculterella. Neither Pseudohemiculter nor Hemiculterella are monophyletic based on our result, highlighting the need to explore alternative morphological diagnostic characters to facilitate identification.
Historical biogeography among Hainan Island and northern Vietnam has attracted the attention of ichthyologists [30,31,32,33]. Studies based on Channa gachua, Tanichthys, and Opsariichthys hainanensis all supported the hypothesis that Hainan Island might have originated from part of the northeastern Indo-China Peninsula and adjacent Guangxi, China, instead of from southeastern China mainland, particularly Guangdong [34]. However, most taxon included in previous biogeographic studies typically lack the ability to migrate, and population genetic patterns in a highly migratory clade remain understudied. Hainania minzhengi sp. nov. alongside its Hainanese endemic congener are of great migration ability and potential in bringing new insights into the biogeographic history of Hainanese freshwaters. Current specimens and records of their distributions are insufficient to infer their biogeographic histories with confidence. Future acquisition of a more exhaustive collection of specimens and locality information is required.
5. Comparative Material
Hainania serrata: IHBHK202308001-IHBHK2023080015, 15, 83.9−112.1 mm SL, Longzhou River, Dingan County, Hainan Province, China; IHBHK2023080016- IHBHK2023080022, 7, 93.76−113.48 mm SL, Longzhou River, Chengmai County, Hainan Province, China; IHBSY202505001- IHBSY202505005, 5, 63.81−73.52 mm SL, Caopeng Reservoir, Jiyang District, Sanya City, Hainan Province, China.
Pseudohemiculter dispar: IHBLZ202410001-IHBLZ202410004, 4, 123.4−149.1 mm SL, Liujiang River, Jiangkou Town, Luzhai County, Liuzhou City, Guangxi Zhuang Autonomous Region, China.
Pseudohemiculter hainanensis: IHBHZ202407001, 1, 73.02 mm SL, Dongjiang River, Boluo County, Huizhou City, Guangdong Province, China; IHBBS202301001, 1, 99.51 mm SL, Chengbi River, Youjiang District, Baise City, Guangxi Zhuang Autonomous Region, China; IHBQZ202308001, 1, 66.9 mm SL, Shihou River, Qiongzhong Li and Miao Autonomous County, Hainan Province, China; IHBFCG202405001, 1, 77.64 mm SL, Beilun River, Naliang Town, Fangcheng District, Fangchenggang City, Guangxi Zhuang Autonomous Region, China.
Pseudohemiculter kweichowensis: IHBQN202307001, 1, 118.9 mm SL, Duliujiang River, Sandu Shui Autonomous County, Qiannan Buyi and Miao Autonomous Prefecture, Guizhou Province, China.
Hemiculterella wui: IHBGZ202408001, 1, 77.6 mm SL, Liuxihe River, Conghua District, Guangzhou City, Guangdong Province, China.
Supplementary Materials
The following supporting information can be downloaded at https://www.mdpi.com/article/10.3390/d17080549/s1: Table S1: Morphometric measurements of 19 individuals of Ha. minzhengi sp.nov. Table S2: The uncorrected pairwise genetic distances between Chinese species of Pseudohemiculter and Hainania based on separated Cyt b and COI genes. Maximum intraspecific genetic distances are highlighted in bold. (A) Cyt b; (B) COI.
Author Contributions
Conceptualization, H.L. and X.L.; methodology, H.L.; software, X.L.; validation, H.L.; formal analysis, H.L.; investigation, H.L.; resources, Z.G.; data curation, Z.G.; writing—original draft preparation, H.L.; writing—review and editing, X.L.; visualization, H.L.; supervision, X.L.; project administration, X.L.; funding acquisition, X.L. All authors have read and agreed to the published version of the manuscript.
Funding
This work was supported by the National Key Research and Development Program of China (2023YFC2606600). X. Li was supported by the 2115 Talent Development Program of China Agricultural University. The authors acknowledge the China Agricultural University for providing high-performance computing platforms and support that contributed to the research results reported in this study.
Institutional Review Board Statement
Not applicable.
Data Availability Statement
All newly produced sequences were deposited in GenBank.
Acknowledgments
We would like to express our sincere gratitude to Minzheng Li (Guangzhou, China) and Shubin Wu (Sanya, China) for providing part of the specimens and assistance in sample collection and field works, to Dekui He (IHB) as well as Zhixian Sun, Yutian Fang, and Yahui Zhao (ASIZB) for their constant support and help in specimen preservation, and to Haoyang Xie, Yiyang Xu, and Meiling Qing (IHB) for their assistance in preparation of the illustrations in this study.
Conflicts of Interest
The authors of this work declare that they have no conflicts of interest.
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Figure 1.
Sampling sites of Hainania minzhengi sp. nov. (black triangle), Ha. serrata (black square), Hemiculterella wui (yellow circle), Pseudohemiculter dispar (red asterisk), P. hainanensis (red cross), and P. kweichowensis (red pentagon).
Figure 1.
Sampling sites of Hainania minzhengi sp. nov. (black triangle), Ha. serrata (black square), Hemiculterella wui (yellow circle), Pseudohemiculter dispar (red asterisk), P. hainanensis (red cross), and P. kweichowensis (red pentagon).
Figure 2.
Phylogenetic tree of Hainania, Pseudohemiculter and other closely related genera reconstructed based on combined two mitochondrial (Cyt b and COI) gene fragments. UFB and SH-aLRT supports from maximum likelihood (ML) analyses as well as Bayesian posterior probabilities (BPPs) from Bayesian inference (BI) analyses in two partitioning schemes (PGM and PCM) are noted beside nodes as colored squares. Square 1 for ML and PGM, Square 2 for ML and PCM, Square 3 for BI and PGM, and Square 4 for BI and PCM. Branches were considered robustly supported if SH-aLRT ≥ 80 and UFBoot ≥ 95 for ML, pp ≥ 0.95 for BI (red for ML and blue for BI); moderately supported if SH-aLRT ≥ 80 or UFBoot ≥ 95 for ML, 0.95 ≥ pp ≥ 0.8 for BI (pink for ML and light blue for BI); and weakly supported if SH-aLRT < 80 and UFBoot < 95 for ML, pp < 0.8 for BI (white for ML and light blue for BI). The scale bar represents 0.03 nucleotide substitutions per site.
Figure 2.
Phylogenetic tree of Hainania, Pseudohemiculter and other closely related genera reconstructed based on combined two mitochondrial (Cyt b and COI) gene fragments. UFB and SH-aLRT supports from maximum likelihood (ML) analyses as well as Bayesian posterior probabilities (BPPs) from Bayesian inference (BI) analyses in two partitioning schemes (PGM and PCM) are noted beside nodes as colored squares. Square 1 for ML and PGM, Square 2 for ML and PCM, Square 3 for BI and PGM, and Square 4 for BI and PCM. Branches were considered robustly supported if SH-aLRT ≥ 80 and UFBoot ≥ 95 for ML, pp ≥ 0.95 for BI (red for ML and blue for BI); moderately supported if SH-aLRT ≥ 80 or UFBoot ≥ 95 for ML, 0.95 ≥ pp ≥ 0.8 for BI (pink for ML and light blue for BI); and weakly supported if SH-aLRT < 80 and UFBoot < 95 for ML, pp < 0.8 for BI (white for ML and light blue for BI). The scale bar represents 0.03 nucleotide substitutions per site.
Figure 3.
Morphological comparison of two Hainania species and two closely related Pseudohemiculter species (lateral view of body) in preservative (95% ethanol). (A) Hainania minzhengi sp. nov., IHBMM202407002; (B) Hainania serrata, IHBHK202308011; (C) Pseudohemiculter dispar, IHBLZ202410004; and (D) Pseudohemiculter hainanensis, IHBBS202301001.
Figure 3.
Morphological comparison of two Hainania species and two closely related Pseudohemiculter species (lateral view of body) in preservative (95% ethanol). (A) Hainania minzhengi sp. nov., IHBMM202407002; (B) Hainania serrata, IHBHK202308011; (C) Pseudohemiculter dispar, IHBLZ202410004; and (D) Pseudohemiculter hainanensis, IHBBS202301001.
Figure 4.
Morphological comparison of two Hainania species and two closely related Pseudohemiculter species (detailed morphology of dorsal fin spine) in preservative (95% ethanol). Dorsal fin spines are magnified to illustrate dentations. (A) Hainania minzhengi sp. nov., IHBMM202407002; (B) Hainania serrata, IHBHK202308011; (C) Pseudohemiculter dispar, IHBLZ202410004; and (D) Pseudohemiculter hainanensis, IHBBS202301001.
Figure 4.
Morphological comparison of two Hainania species and two closely related Pseudohemiculter species (detailed morphology of dorsal fin spine) in preservative (95% ethanol). Dorsal fin spines are magnified to illustrate dentations. (A) Hainania minzhengi sp. nov., IHBMM202407002; (B) Hainania serrata, IHBHK202308011; (C) Pseudohemiculter dispar, IHBLZ202410004; and (D) Pseudohemiculter hainanensis, IHBBS202301001.
Figure 5.
Morphological comparison of two Hainania species and two closely related Pseudohemiculter species (detailed lateral profile of head) in preservative (95% ethanol). (A) Hainania minzhengi sp. nov., IHBMM202407002; (B) Hainania serrata, IHBHK202308011; (C) Pseudohemiculter dispar, IHBLZ202410004; and (D) Pseudohemiculter hainanensis, IHBBS202301001.
Figure 5.
Morphological comparison of two Hainania species and two closely related Pseudohemiculter species (detailed lateral profile of head) in preservative (95% ethanol). (A) Hainania minzhengi sp. nov., IHBMM202407002; (B) Hainania serrata, IHBHK202308011; (C) Pseudohemiculter dispar, IHBLZ202410004; and (D) Pseudohemiculter hainanensis, IHBBS202301001.
Figure 6.
Morphological characters of holotype, IHBMM202407001tp of Hainania minzhengi sp. nov. in preservative (10% formalin). (A) Lateral view; (B) detailed lateral profile of head; and (C) detailed profile of ventral keel.
Figure 6.
Morphological characters of holotype, IHBMM202407001tp of Hainania minzhengi sp. nov. in preservative (10% formalin). (A) Lateral view; (B) detailed lateral profile of head; and (C) detailed profile of ventral keel.
Figure 7.
Hainania minzhengi sp. nov. in life, holotype IHBMM202407001tp. Photo from Haoyang Xie.
Figure 8.
Habitats of two Hainania species. Photo from Minzheng Li. (A) Hainania minzhengi sp. nov. (type locality); (B) Ha. serrata.
Figure 8.
Habitats of two Hainania species. Photo from Minzheng Li. (A) Hainania minzhengi sp. nov. (type locality); (B) Ha. serrata.
Figure 9.
Principal component analysis of head shape difference between the two Hainania species (Red: Ha. serrata; Blue: Ha. minzhengi). Circles are 80%-equal frequency ellipses of each species. Line graphs on the edges illustrate the distribution of measurements after dimensionality reduction.
Figure 9.
Principal component analysis of head shape difference between the two Hainania species (Red: Ha. serrata; Blue: Ha. minzhengi). Circles are 80%-equal frequency ellipses of each species. Line graphs on the edges illustrate the distribution of measurements after dimensionality reduction.
Table 1.
Voucher code, sampling localities, and GenBank accession numbers of all sequences used for molecular analyses. Outgroups are underlined.
Table 1.
Voucher code, sampling localities, and GenBank accession numbers of all sequences used for molecular analyses. Outgroups are underlined.
| Number | Species | Locality | Voucher ID | Cyt b | COI |
|---|---|---|---|---|---|
| 1 | Pseudohemiculter hainanensis | Boluo County, Huizhou City, Guangdong Province, China | IHBHZ202407001 | PV164763 | PV165721 |
| 2 | Pseudohemiculter hainanensis | Youjiang District, Baise City, Guangxi Zhuang Autonomous Region, China | IHBBS202301001 | PV164764 | PV165722 |
| 3 | Pseudohemiculter hainanensis | Qiongzhong Li and Miao Autonomous County, Hainan Province, China | IHBQZ202308001 | PV164765 | PV165723 |
| 4 | Pseudohemiculter hainanensis | Naliang Town, Fangcheng District, Fangchenggang City, Guangxi Zhuang Autonomous Region, China | IHBFCG202405001 | PV164766 | PV165724 |
| 5 | Pseudohemiculter dispar | Jiangkou Town, Luzhai County, Liuzhou City, Guangxi Zhuang Autonomous Region, China. | IHBLZ202410001 | PV164767 | PV165725 |
| 6 | Pseudohemiculter dispar | Jiangkou Town, Luzhai County, Liuzhou City, Guangxi Zhuang Autonomous Region, China. | IHBLZ202410002 | PV164768 | PV165726 |
| 7 | Pseudohemiculter dispar | Jiangkou Town, Luzhai County, Liuzhou City, Guangxi Zhuang Autonomous Region, China. | IHBLZ202410003 | PV164769 | PV165727 |
| 8 | Pseudohemiculter dispar | Jiangkou Town, Luzhai County, Liuzhou City, Guangxi Zhuang Autonomous Region, China. | IHBLZ202410004 | PV164770 | PV165728 |
| 9 | Pseudohemiculter kweichowensis | Sandu Shui Autonomous County, Qiannan Buyi and Miao Autonomous Prefecture, Guizhou Province, China | IHBQN202307001 | PV164771 | PV165729 |
| 10 | Hainania serrata | Dingan County, Hainan Province, China | IHBHK202308002 | PV164772 | PV165730 |
| 11 | Hainania serrata | Dingan County, Hainan Province, China | IHBHK202308004 | PV164773 | PV165731 |
| 12 | Hainania serrata | Dingan County, Hainan Province, China | IHBHK202308005 | PV164774 | PV165732 |
| 13 | Hainania serrata | Dingan County, Hainan Province, China | IHBHK202308006 | PV164775 | PV165733 |
| 14 | Hainania serrata | Dingan County, Hainan Province, China | IHBHK202308008 | PV164776 | PV165734 |
| 15 | Hainania serrata | Dingan County, Hainan Province, China | IHBHK202308010 | PV164777 | PV165735 |
| 16 | Hainania serrata | Dingan County, Hainan Province, China | IHBHK202308011 | PV164778 | PV165736 |
| 17 | Hainania serrata | Dingan County, Hainan Province, China | IHBHK202308013 | PV164779 | PV165737 |
| 18 | Hainania serrata | Dingan County, Hainan Province, China | IHBHK202308014 | PX024142 | PV991209 |
| 19 | Hainania serrata | Chengmai County, Hainan Province, China | IHBHK202308015 | PX024143 | PV991208 |
| 20 | Hainania serrata | Chengmai County, Hainan Province, China | IHBHK202308017 | PX024144 | PV991210 |
| 21 | Hainania serrata | Chengmai County, Hainan Province, China | IHBHK202308018 | PX024145 | PV991202 |
| 22 | Hainania serrata | Chengmai County, Hainan Province, China | IHBHK202308019 | PX024146 | PV991211 |
| 23 | Hainania serrata | Chengmai County, Hainan Province, China | IHBHK202308020 | PX024147 | PV991200 |
| 24 | Hainania serrata | Chengmai County, Hainan Province, China | IHBHK202308021 | PX024148 | PV991207 |
| 25 | Hainania serrata | Jiyang District, Sanya City, Hainan Province, China | IHBSY202505001 | PX024149 | PV991201 |
| 26 | Hainania serrata | Jiyang District, Sanya City, Hainan Province, China | IHBSY202505002 | PX024150 | PV991205 |
| 27 | Hainania serrata | Jiyang District, Sanya City, Hainan Province, China | IHBSY202505003 | PX024151 | PV991204 |
| 28 | Hainania serrata | Jiyang District, Sanya City, Hainan Province, China | IHBSY202505004 | PX024152 | PV991206 |
| 29 | Hainania serrata | Jiyang District, Sanya City, Hainan Province, China | IHBSY202505005 | PX024153 | PV991203 |
| 30 | Hainania minzhengi sp. nov. | Shuikou Town, Xinyi City, Maoming City, Guangdong Province, China | IHBMM202407002 | PV164780 | PV165738 |
| 31 | Hainania minzhengi sp. nov. | Shuikou Town, Xinyi City, Maoming City, Guangdong Province, China | IHBMM202407004 | PV164781 | PV165739 |
| 32 | Hainania minzhengi sp. nov. | Shuikou Town, Xinyi City, Maoming City, Guangdong Province, China | IHBMM202407005 | PX024154 | PV991196 |
| 33 | Hainania minzhengi sp. nov. | Shuikou Town, Xinyi City, Maoming City, Guangdong Province, China | IHBMM202407006 | PX024155 | PV991197 |
| 34 | Hainania minzhengi sp. nov. | Shuikou Town, Xinyi City, Maoming City, Guangdong Province, China | IHBMM202407007 | PX024156 | PV991198 |
| 35 | Hainania minzhengi sp. nov. | Shuikou Town, Xinyi City, Maoming City, Guangdong Province, China | IHBMM202407008 | PV164782 | PV165740 |
| 36 | Hainania minzhengi sp. nov. | Shuikou Town, Xinyi City, Maoming City, Guangdong Province, China | IHBMM202407009 | PX024157 | PV991199 |
| 37 | Hainania minzhengi sp. nov. | Shuikou Town, Xinyi City, Maoming City, Guangdong Province, China | IHBMM202407010 | PV164783 | PV165741 |
| 38 | Hainania minzhengi sp. nov. | Shuikou Town, Xinyi City, Maoming City, Guangdong Province, China | IHBMM202407011 | PX024158 | PV991192 |
| 39 | Hainania minzhengi sp. nov. | Shuikou Town, Xinyi City, Maoming City, Guangdong Province, China | IHBMM202407013 | PV164784 | PV165742 |
| 40 | Hainania minzhengi sp. nov. | Shuikou Town, Xinyi City, Maoming City, Guangdong Province, China | ASIZB 248490 | PX024159 | PV991193 |
| 41 | Hainania minzhengi sp. nov. | Shuikou Town, Xinyi City, Maoming City, Guangdong Province, China | ASIZB 248491 | PX024160 | PV991194 |
| 42 | Hainania minzhengi sp. nov. | Shuikou Town, Xinyi City, Maoming City, Guangdong Province, China | ASIZB 248492 | PX024161 | PV991195 |
| 43 | Hainania serrata | Qiongzhong Li and Miao Autonomous County, Hainan Province, China | / | OP251698.1 | OP251568.1 |
| 44 | Hainania serrata | Qiongzhong Li and Miao Autonomous County, Hainan Province, China | / | OP251699.1 | OP251569.1 |
| 45 | Pseudohemiculter hainanensis | Market of Laibin City, Guangxi Zhuang Autonomous Region, China | / | NC_065693.1 | NC_065693.1 |
| 46 | Hemiculterella wui | Conghua District, Guangzhou City, Guangdong Province, China | IHBGZ202408001 | PV164785 | PV165743 |
| 47 | Hemiculterella wui | / | / | NC_084163.1 | NC_084163.1 |
| 48 | Hemiculterella sauvagei | Xin Village, Cuiping Distrct, Yibin City, Sichuan Province, China | / | NC_026693.1 | NC_026693.1 |
| 49 | Toxabramis houdemeri | Fusui County, Chongzuo City, Guangxi Zhuang Autonomous Region, China | / | NC_029348.1 | NC_029348.1 |
| 50 | Toxabramis swinhonis | Dianshan Lake, Qingpu District, Shanghai City, China | / | NC_029249.1 | NC_029249.1 |
| 51 | Hemiculter leucisulus | / | / | NC_022929.1 | NC_022929.1 |
| 52 | Hemiculter yungaoi | Shapo Reservoir, Longhua District, Haikou City, Hainan Province, China | / | OP251707.1 | OP251579.1 |
| 53 | Ctenopharyngodon idella | / | / | NC_010288.1 | NC_010288.1 |
Table 2.
The uncorrected pairwise genetic distances between Chinese species of Pseudohemiculter and Hainania based on combined Cyt b and COI genes. Maximum intraspecific genetic distances are highlighted in bold.
Table 2.
The uncorrected pairwise genetic distances between Chinese species of Pseudohemiculter and Hainania based on combined Cyt b and COI genes. Maximum intraspecific genetic distances are highlighted in bold.
| P. hainanensis | P. dispar | P. kweichowensis | Ha. serrata | Ha. minzhengi sp. nov. | |
|---|---|---|---|---|---|
| P. hainanensis | 0.0544 | ||||
| P. dispar | 0.1023 | 0.0040 | |||
| P. kweichowensis | 0.1217 | 0.1249 | / | ||
| Ha. serrata | 0.0677 | 0.1042 | 0.1195 | 0.0185 | |
| Ha. minzhengi sp. nov. | 0.0673 | 0.0969 | 0.1206 | 0.0483 | 0.0046 |
Table 3.
Morphometric measurements for Hainania minzhengi sp. nov., Ha. serrata, Pseudohemiculter dispar, and P. hainanensis. Characteristics in bold were used in the principal component analysis (PCA).
Table 3.
Morphometric measurements for Hainania minzhengi sp. nov., Ha. serrata, Pseudohemiculter dispar, and P. hainanensis. Characteristics in bold were used in the principal component analysis (PCA).
| Characteristics | Hainania minzhengi (n = 19) | Hainania serrata (n = 22) | Pseudohemiculter dispar (n = 3) | Pseudohemiculter hainanensis (n = 3) | |||||||||
|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
| Holotype | Range | Mean | S.D. | Range | Mean | S.D. | Range | Mean | S.D. | Range | Mean | S.D. | |
| Lateral line scales | 54 | 52–54 | 53 | / | 55–56 | 55 | / | 50–52 | 52 | / | 48–52 | 49 | / |
| Dorsal fin rays | III, 7 | III, 7 | III, 7 | / | III, 7 | III, 7 | / | III, 7 | III, 7 | / | III, 7 | III, 7 | / |
| Anal-fin rays | III, 14 | III, 13–14 | III, 13 | / | III, 15 | III, 15 | / | III, 16 | III, 16 | / | III, 13 | III, 13 | / |
| Pectoral fin rays | I, 13 | I, 13–14 | I, 13 | / | I, 14 | I, 14 | / | I, 14 | I, 14 | / | I, 13 | I, 13 | / |
| Pelvic fin rays | I, 7 | I, 6–7 | I, 7 | / | I, 6 | I, 6 | / | I, 8 | I, 8 | / | I, 7 | I, 7 | / |
| Caudal fin rays | 2 + 16 | 2 + 16 | 2 + 16 | / | 2 + 16 | 2 + 16 | / | 2 + 16 | 2 + 16 | / | 2 + 16 | 2 + 16 | / |
| Standard length (SL, mm) | 84.64 | 69.8–110.9 | 90.7 | 11.5 | 64.5–117.4 | 94.9 | 16.9 | 123.4–149.1 | 139.7 | 14.1 | 66.9–99.5 | 79.8 | 17.3 |
| In percentage of SL (%) | |||||||||||||
| Head length (HL) | 24.3 | 20.6−24.5 | 23.4 | 0.9 | 21.9−26.4 | 24.1 | 1.2 | 21.7–24.2 | 22.8 | 1.2 | 16.8–24.5 | 21.8 | 4.3 |
| Body depth | 23.4 | 14.6−26.5 | 23.2 | 2.4 | 19.4−25.5 | 23.0 | 1.6 | 21.8–25.4 | 24.2 | 2.0 | 25.4–26.3 | 25.7 | 0.5 |
| Predorsal length | 51.6 | 41.7−51.9 | 49.5 | 2.2 | 46.2−52.3 | 50.3 | 1.5 | 49.0–52.3 | 50.9 | 1.7 | 51.9–53.1 | 52.5 | 0.6 |
| Prepelvic length | 48.5 | 35.3−48.4 | 45.9 | 2.8 | 43.8−48.0 | 46.2 | 1.3 | 45.7–48.8 | 47.3 | 1.6 | 48–50 | 49.1 | 1.0 |
| Preanal length | 69.6 | 60.2−71.0 | 67 | 2.3 | 65.2−70.9 | 67.6 | 1.5 | 70.5–73.7 | 72.0 | 1.7 | 67.9–69.5 | 68.6 | 0.8 |
| Dorsal-fin height | 19.5 | 10.6−24.1 | 20.9 | 2.8 | 16.1−25.5 | 21.7 | 2.4 | 17.6–17.7 | 17.7 | 0.1 | 18–18.6 | 18.5 | 0.3 |
| Dorsal-fin base length | 10.5 | 6.5−10.5 | 8.1 | 1.1 | 7.4−10.7 | 9.0 | 0.9 | 8.7–10.5 | 9.6 | 0.9 | 9.4–11.5 | 10.4 | 1.0 |
| Anal-fin height | 12.2 | 7.7−15.5 | 10.3 | 1.5 | 8.1−11.4 | 9.8 | 1.1 | 9.2–10.2 | 9.6 | 0.6 | 11–11.8 | 11.4 | 0.4 |
| Anal-fin base length | 17.7 | 13.6−19.0 | 16.7 | 1.2 | 15.1−18.5 | 16.6 | 1.1 | 15.0–17.6 | 16.3 | 1.3 | 16.9–17.2 | 17.1 | 0.1 |
| Pelvic-fin length | 16.2 | 12.3−16.9 | 14.5 | 1.1 | 19.2−23.8 | 21.5 | 1.3 | 14.1–15.7 | 15.0 | 0.8 | 14–14.9 | 14.4 | 0.5 |
| Pectoral-fin length | 22.0 | 18.7−22.9 | 20.8 | 1.1 | 13.1−18.3 | 15.8 | 1.2 | 18.2–21.1 | 20.1 | 1.6 | 19.8–20.9 | 20.2 | 0.6 |
| Caudal-fin length | 22.5 | 18.9−28.5 | 23.6 | 2.6 | 8.4−10.3 | 9.1 | 0.5 | 21.4–25.5 | 23.5 | 2.0 | 18–23.2 | 20.9 | 2.6 |
| Caudal-peduncle length (CPL) | 15.4 | 13.5−19.1 | 16.4 | 1.3 | 19.2−25.1 | 22.6 | 1.5 | 14.6–16.4 | 15.3 | 1.0 | 14.7–16.9 | 15.7 | 1.2 |
| Caudal-peduncle depth (CPD) | 9.6 | 8.0−10.5 | 9.5 | 0.5 | 18.0−23.4 | 21.6 | 1.5 | 8.8–9 | 8.9 | 0.1 | 8.3–9.1 | 8.8 | 0.4 |
| P–V distance | 24.2 | 21.1−27.4 | 24.1 | 1.4 | 19.2−23.8 | 21.5 | 1.3 | 22.9–25.4 | 24.5 | 1.4 | 23.1–27.2 | 24.7 | 2.2 |
| V–A distance | 21.3 | 19.7−24.2 | 21.5 | 1.2 | 13.1−18.3 | 15.8 | 1.2 | 22.7–24.8 | 23.6 | 1.1 | 18.9–20.6 | 19.9 | 0.8 |
| In percentage of HL (%) | |||||||||||||
| Head depth | 59.4 | 56.5−75.4 | 61.8 | 4.1 | 55.4−63.7 | 59.2 | 2.2 | 61.2–70.6 | 64.5 | 5.4 | 72.5–102.6 | 83.0 | 17.1 |
| Head width | 41.0 | 35.5−44.6 | 40.3 | 2.1 | 37.8−42.7 | 40.0 | 1.5 | 42.6–44.8 | 43.7 | 1.1 | 52–78.7 | 62.0 | 14.6 |
| Snout length | 27.6 | 25.0−29.0 | 26.8 | 1.0 | 26.8−32.9 | 30.2 | 1.5 | 30.7–33.3 | 32.4 | 33.3 | 26.6–40.7 | 32.2 | 7.5 |
| Eye diameter | 33.1 | 30.4−32.9 | 31.1 | 0.6 | 24.6−29.7 | 27.6 | 1.3 | 26.3–29.7 | 28.2 | 1.7 | 29.9–52.2 | 38.8 | 11.8 |
| Interorbital width | 32.6 | 26.2−35.4 | 29.9 | 2.4 | 24.6−33.9 | 27.5 | 2.2 | 27.7–31.2 | 29.5 | 1.8 | 32.7–53.9 | 40.4 | 11.8 |
| Snout length/eye diameter (%) | 83.4 | 80.0−93.0 | 86.2 | 3.2 | 101.0−121.5 | 109.3 | 5.5 | 111.2–117.0 | 115.0 | 3.3 | 77.9–89 | 84.0 | 5.6 |
| CPD/CPL (%) | 62.3 | 48.9–63.1 | 58.4 | 3.8 | 47.9−69.4 | 58.5 | 5.6 | 54.8–60.8 | 58.5 | 3.2 | 53.2–61.6 | 56.6 | 4.4 |
Table 4.
Loadings on the first two principal components extracted from morphometric data of the two Hainania species. Measurements with significant contributions are highlighted in bold.
Table 4.
Loadings on the first two principal components extracted from morphometric data of the two Hainania species. Measurements with significant contributions are highlighted in bold.
| Morphometric Measurements | PC1 | PC2 |
|---|---|---|
| HL/SL | 0.1057 | −0.2265 |
| In percentage of HL | ||
| Head depth | −0.1359 | 0.3471 |
| Head width | −0.0563 | 0.1947 |
| Snout length | 0.3620 | 0.4390 |
| Eye diameter | −0.3858 | 0.1265 |
| Interorbital distance | −0.3553 | 0.7028 |
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MDPI and ACS Style
Lei, H.; Gong, Z.; Li, X. Description of a New Species of Hainania Koller (Teleostei, Cypriniformes, Xenocyprididae) from Guangdong Province, Southern China. Diversity 2025, 17, 549. https://doi.org/10.3390/d17080549
AMA Style
Lei H, Gong Z, Li X. Description of a New Species of Hainania Koller (Teleostei, Cypriniformes, Xenocyprididae) from Guangdong Province, Southern China. Diversity. 2025; 17(8):549. https://doi.org/10.3390/d17080549
Chicago/Turabian StyleLei, Haotian, Ziyu Gong, and Xuankun Li. 2025. "Description of a New Species of Hainania Koller (Teleostei, Cypriniformes, Xenocyprididae) from Guangdong Province, Southern China" Diversity 17, no. 8: 549. https://doi.org/10.3390/d17080549
APA StyleLei, H., Gong, Z., & Li, X. (2025). Description of a New Species of Hainania Koller (Teleostei, Cypriniformes, Xenocyprididae) from Guangdong Province, Southern China. Diversity, 17(8), 549. https://doi.org/10.3390/d17080549
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