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. 2024 Dec 25;63:e37. doi: 10.6620/ZS.2024.63-37

Checklist and DNA Barcoding of the Scorpaenidae (Teleostei: Scorpaeniformes) in Taiwan

Tak-Kei Chou 1, Wen-Chien Huang 1, Wei-Cheng Jhuang 2,3, Te-Yu Liao 1,*
PMCID: PMC12399982  PMID: 40900687

Abstract

Species of the family Scorpaenidae are easily misidentified due to their similar appearances, a result of camouflaging to their surroundings. In recent years, many species from this family have been described, and generic placements of some species have been revised. Previously, there were 80 species belonging to 29 genera of the Scorpaenidae recorded in Taiwanese waters. However, their taxonomy has not been revised for decades. It is necessary to update the checklist of the Scorpaenidae occurring in Taiwanese waters based on updated morphological and molecular data. In the present study, we revised the Taiwanese scorpaenids based on 296 specimens and updated the checklist, amounting to a total of 85 species of 29 genera, of which Sebastapistes mauritiana (Cuvier) is a new record, and three species from the genera Phenacoscorpius, Scorpaenopsis, and Sebastapistes are unable to be identified to any species. Using molecular analysis, we conducted the first comprehensive DNA barcoding study of the Scorpaenidae from Taiwanese waters based on a partial cytochrome c oxidase I (COI) gene of 655 bps. A total of 118 COI sequences were generated from voucher specimens of 66 species (28 genera) identified based on morphological characters. The COI sequences of Parascorpaena maculipinnis, Scorpaena pepo, and Scorpaenopsis orientalis are new to online databases. According to the Kimura-2 Parameter (K2P) genetic distance, the mean interspecific variation (15.61%) was distinctly greater than the mean intraspecific variation (0.22%), suggesting a barcoding gap. The maximum likelihood tree showed that all lineages were supported by high bootstrap values.

Keywords: Diversity, Ichthyofauna, Mitochondrion, Stonefish, Taxonomy

BACKGROUND

Members of the family Scorpaenidae are known for the venomous spines on their dorsal, pelvic, and anal fins (Allen and Eschmeyer 1973; Nelson 2006), and their ability to camouflage with their surroundings using variable color patterns, tentacles, flaps, and barbels (Poss 1999; Randall and Eschmeyer 2002; Krzyżak and Korzeniewski 2021). This family is distributed in a variety of habitats, such as intertidal and sublittoral zones, sandy substrates, coral reefs, rocky reefs, and continental shelves with a depth range from 0 to 1,600 m (Masuda et al. 1984; Fedorov et al. 2003; Poss and Eschmeyer 2003; Randall 2005a).

The Scorpaenidae are a species-rich family, composed of 36 genera and more than 350 valid species (Nelson et al. 2016; Fricke et al. 2024), but their definition has been contentious (Matsubara 1943; Washington et al. 1984; Ishida 1994; Poss 1999; Imamura 2004; Shinohara and Imamura 2005; Smith et al. 2018). In the present study, the definition of the Scorpaenidae follows the classification of Smith et al. (2018) which was based on morphological and molecular characteristics of a large number of Scorpaeniformes specimens. Poss (1999) and Smith et al. (2018) proposed that this family is characterized by the presence of a suborbital stay firmly connected to the preopercle, spines on head, gill membranes free from isthmus, compressed body, presence of scales, 24 to 30 vertebrae, venomous glands of the dorsal, anal, and pelvic fin spines, dorsal fin with XII to XIII spines (rarely VIII), anal fin with III spines (rarely II), pelvic fins with one spine and five rays, and a well-developed pectoral fin with lower rays unbranched. Morphological identification of scorpaenids has been considered difficult because of limited diagnostic characters and variable color patterns based on surroundings for most species (Poss 1999; Randall and Eschmeyer 2002; Randall 2005a b; Krzyżak and Korzeniewski 2021). In the last two decades, most taxonomic work on Scorpaenidae was conducted based on specimens from the western Pacific, including descriptions of new species and/or synonymizations of nominal species in the genera Brachypterois, Dendrochirus, Pterois, Scorpaena, Scorpaenopsis, Sebastapistes, Sebastiscus, Parascorpaena, Lythrichthys, Neomerinthe,

Phenacoscorpius, Pteroidichthys, Scorpaena, and Scorpaenodes (Randall and Eschmeyer 2002; Allen and Erdmann 2008; Motomura 2008 2009; Motomura and Senou 2008; Motomura et al. 2010a b 2014 2015 2016; Matsunuma et al. 2013 2017; Motomura and Kanade 2015; Matsunuma and Motomura 2015 2019; Morishita et al. 2018; Wibowo and Motomura 2019a b; Hoshino and Motomura 2021; Wada et al. 2021; Chou and Liao 2022; Chou et al. 2023; Matsumoto et al. 2023; Matsumoto and Motomura 2024). Among these studies, only Matsunuma et al. (2017), Wada et al. (2021), Chou and Liao (2022), and Chouet al. (2023) provided molecular characters of scorpaenids.

In Taiwan, the first study on the diversity of the Scorpaenidae was Chen (1969)’s synopsis of the vertebrates of Taiwan, in which 13 species from nine genera were recorded. Later, Chen (1981) revised the taxonomy of the Scorpaenidae from Taiwan and recorded 42 species of 25 genera, and Shen et al. (1993) updated the list to 48 species of 26 genera. Recent taxonomic studies of the Scorpaenidae have greatly expanded the biodiversity of this fish group in Taiwan (Randall and Eschmeyer 2002; Chen 2003; Motomura et al. 2007 2009a b 2010a 2011; Shao et al. 2008; Motomura and Senou 2009; Chen et al. 2010; Shen and Wu 2011; Morishita et al. 2018; Koeda and Ho 2019; Koeda et al. 2019; Chou 2021; Chou and Tang 2021; Wada et al. 2021; Chou and Liao 2022), amounting to a total of 80 species placed in 29 genera. Despite the increased number of recorded species, there was no taxonomic revision of the Scorpaenidae made from Taiwanese waters after Chen (1981). Furthermore, generic placements of some species have changed (Randall and Poss 2002; Motomura et al. 2010b; Poss et al. 2010; Wada et al. 2021), and some species in the western Pacific were considered junior synonyms or misidentifications (Nakabo 2002; Motomura et al. 2009a b 2010b 2015; Wibowo and Motomura 2019a b; Hoshino and Motomura 2021; Wada et al. 2021). These recent taxonomic works imply that the checklist of scorpaenids in Taiwan needs revision based on museum collections and recent taxonomic literature. In addition, DNA barcoding is needed to provide molecular references for the scorpaenids of Taiwan.

In the present study, we aimed to: (a) provide a reliable DNA barcoding reference of the Scorpaenidae from Taiwanese waters based on morphological examination; and (b) update the checklist of the Scorpaenidae present in Taiwan.

MATERIALS AND METHODS

Sampling and morphological analyses

Fresh specimens were collected in Taiwanese waters from 2017 to 2022 using hand nets and angling, as well as purchasing from local fish markets. Several islands around Taiwan were also surveyed, including Penghu, Liuqiu, Lanyu (Orchid Island), and Green Islands (Fig. 1). Fresh specimens were fixed in 10% neutral buffered formalin and preserved in 70% ethanol thereafter. Some examined materials were loaned from museum collections of the Academia Sinica Institute of Zoology, Taipei (ASIZP), and the National Museum of Marine Biology and Aquarium (NMMB-P), Pingtung. At least five specimens per species were examined whenever possible. The definition of habitat types in Taiwan followed Shao et al. (2008). Terminology and definitions of morphometrics and meristics generally followed Motomura (2004a b), Motomura et al. (2005a–c), and Motomura and Johnson (2006). The terminology of head spines followed Randall and Eschmeyer (2002). The meristics were examined on the left side of fish. The last two rays of the dorsal and anal fins were counted as one. The vertebra count was determined by X-radiographs. Measurements were carried out using a digital caliper with 1 mm precision. At least two tissue samples per species (when possible) were taken from the fin clips or dorsal muscle and preserved in 95% ethanol. Some tissue samples were loaned from the cryobank of the Research Center for Biodiversity of Academia Sinica. Voucher specimens collected in this study were deposited in ASIZP, Department of Oceanography, National Sun Yat-sen University, Kaohsiung (DOS), and NMMB-P.

Fig. 1.

Fig. 1.

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Map of sampling sites in this study.

DNA extraction, amplification, and sequencing

DNA was extracted from tissue samples using GeneMark Easy Tissue & Cell Genomic DNA Purification Kit following the manufacturer’s protocol. The polymerase chain reaction (PCR) was used to amplify the cytochrome c oxidase subunit I (COI) gene. PCR products were amplified in a 25 μL volume containing 3 μL of 10X Taq Buffer, 2 μL of dNTP mixture at 10 mM, 1 μL of forward and reverse primers at 5 μM, 0.13 μL of Pro Taq Plus DNA polymerase (Protech Technology Enterprise, Taiwan), 1 μL of template DNA, and 16.87 μL of ultrapure water. The COI fragment was amplified using Ward et al.’s (2005) universal COI primers, and ScorF (5'- CTCAGCCATCCTACCTGTGG-3') and ScorR (5'- ACTTCTGGGTGRCCGAAGAA-3') designated by the present study. The thermal PCR condition of COI was composed of an initial denaturation step at 95°C for 4 min, then 35 cycles of 94°C for 30 s, 48°C for 30 s and 72°C for 1 min, and a final extension at 72°C for 10 min. PCR products were visualized on 2% agarose gels and subsequently purified using SAP-Exo Kit (Jena Bioscience). PCR products were sequenced in both forward and reverse directions by a biotechnology company (Genomics, Taiwan). The forward and reverse sequences were edited and assembled using BioEdit ver. 7.2.5 (Hall 1999). All sequences were deposited in Genbank (Table S1).

Sequence analysis

DNA sequences were aligned using ClustalW (Thompson et al. 1994) in BioEdit ver. 7.2.5 (Hall 1999). The substitution saturation of COI mutation was tested using DAMBE ver. 7.0.10 (Xia 2018). The Kimura-two parameter (K2P) model was implemented for COI gene in phylogenetic reconstruction and distance metrics among species. Phylogenetic analysis of COI sequences was conducted with the Maximum likelihood (ML) method using MEGA ver. 10.1.1 (Kumar et al. 2018). Branch support value was assessed using the bootstrapping criterion with 1,000 replicates. Synanceia verrucosa (accession number: JQ432179) of the Synanceiidae was chosen as outgroup for phylogenetic analysis. Genetic divergences at different taxonomic levels (inter-generic, inter-specific, intraspecific) were calculated using MEGA ver. 10.1.1 (Kumar et al. 2018). All COI sequences were compared with those from public databases using Basic Local Alignment Search Tool (BLAST) on GenBank and the BOLD Identification System (IDS) (Ratnasingham and Hebert 2007; Johnson et al. 2008). Sequences with similarity values greater than 98% were considered to be conspecific (Huang et al. 2023).

Ethics statement

The study was conducted in strict accordance with the Wildlife Conservation Act in Taiwan. No ethical approval was required for this study since all species in this study were not protected species and not listed in CITES. Some specimens were collected from Liuqiu and Kenting with the approvals of the Dapeng Bay National Scenic Area Administration, Tourism Bureau (Project #NAMR110029; Collection Permit #11005508700) and the Kenting National Park Headquarters (Project #NAMR110029; Collection Permit #1091002868), respectively. All individuals were not involved in animal experiments.

RESULTS

A total of 296 scorpaenid specimens belonging to 85 species from 29 genera (Table 1) were examined and literature reviewed in the present study (Table S1, Fig. S1), of which one species, Sebastapistes mauritiana (Cuvier, 1829), was newly recorded, and three species of Phenacoscorpius, Scorpaenopsis, and Sebastapistes were not able to be identified to any known species. In total, 118 COI sequences belonging to 66 species of 28 genera were generated, in which sequences of Parascorpaena maculipinnis, Scorpaena pepo, and Scorpaenopsis orientalis were new to online databases (GenBank and BOLD systems). Eight species in four genera were only examined morphologically without molecular data (Table S1).

Table 1.

Checklist of the Scorpaenidae from Taiwanese waters

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DNA barcoding

All data are available in GenBank with accession numbers and catalog numbers of voucher specimens listed in table S1. After alignment, the consensus length of all COI fragments was 655 bps. The saturation was tested for the entire fragment and each codon position of the COI sequences using DAMBE v. 7.0.10 (Xia 2018), and no signs of saturation were detected. No insertion, deletion, and stop codon were found.

The ML tree is shown in figure 2. All morphologybased species were monophyletic groups supported by high bootstrap values. Three genera of the Scorpaenidae were non-monophyletic, including Pteroidichthys, Scorpaenodes, and Sebastapistes. The pairwise genetic distances at different taxonomic levels are summarized in table 2. The intraspecific K2P distance was between 0 and 1.37%, with a mean of 0.22%. The maximum value was found in Helicolenus hilgendorfii (1.37%). The interspecific K2P distance was from 0.46 to 32.32% with a mean of 15.61%. Several species pairs had interspecific distances lower than 2%, including Neochirus bella vs. Neochirus brachyptera (0.69%), Pterois lunulata vs. Pterois russelii (0.62%), Pterois lunulata vs. Pterois volitans (0.85%), Pterois russelii vs. Pterois volitans (0.85%), and Sebastiscus tertius vs. Sebastiscus vibrantus (0.46%). Overall, the mean interspecific distances were over 70-fold higher than the mean intraspecific distances. Aside from the abovementioned five species pairs, the distribution of genetic distances (Fig. 3) also showed a barcoding gap between intraspecific and interspecific divergences. After blasting in GenBank and BOLD databases, sequences of 22 species of nine genera were found to have more than one species with similarities ≥ 98% (Table S2). Three specimens of three genera, Phenacoscorpius, Scorpaenopsis and Sebastapistes, were unsuccessfully identified to any species and their sequences do not match any species in the online database.

Fig. 2.

Fig. 2.

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The maximum likelihood tree based on 118 COI sequences of 66 species of the Scorpaenidae collected from Taiwanese waters. Numerals on nodes represent bootstrap values. Bootstrap values below 70 are not shown. Right hand side labels mark non-monophyletic genera. Color blocks on lineages denote the taxonomically uncertain taxa, including Phenacoscorpius sp. (yellow), Scorpaenopsis sp. (blue), and Sebastapistes sp. (red).

Table 2.

Summary of K2P genetic distances at different taxonomic levels

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Fig. 3.

Fig. 3.

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Distribution frequency of K2P genetic distances (%) for COI of the Scorpaenidae at different taxonomic levels.

The Scorpaenidae fauna in Taiwanese waters

The updated checklist of the Scorpaenidae from Taiwanese waters is shown in table 1. The counts of dorsal, pectoral, and anal fins of all examined specimens are shown in table 3, while the standard length of all examined specimens is shown in table S1. In the present study, 11 species listed in the checklist have a lack of examined specimens and sequences, as their records are only based upon references. Remarks on the new records, species without examined specimens, taxonomically uncertain species, and species pairs with low genetic distance were provided as follows.

Table 3.

Frequency distribution of spine and ray counts on dorsal, pectoral, and anal fins in Taiwanese species of the Scorpaenidae

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Iracundus signifer Jordan & Evermann, 1903

Remarks: No specimen was examined in this study. The record was based on a specimen (BPBMI23411) collected in Nanwan, Pingtung by J.E. Randall in 1978 (Chen 1981), in which some specimens were collected from Taiwanese waters.

Lythrichthys cypho (Fowler, 1938)

Remarks: No specimen was examined in this study. The record was based on Wada et al. (2021).

Lythrichthys longimanus (Alcock, 1894)

Remarks: No specimen was examined in this study. The record was based on Wada et al. (2021), in which some specimens were collected from Taiwanese waters.

Neochirus bella (Jordan & Hubbs, 1925)

Remarks: This species was proposed as a member of the Dendrochirus (= Neochirus) brachypterus complex due to the similarity in overall body appearance (Matsunuma et al. 2017). Neochirus bella could be distinguished from N. brachyptera by the lower count of longitudinal scale series (ca. 34 in N. bella vs. 45–54 in N. brachyptera). Based on our examined materials, the lower count of longitudinal scale series (29–33) matched the description of N. bella.

Neochirus brachyptera (Cuvier, 1829)

Remarks: The comparison between Neochirus bella and N. brachyptera was shown in the remark of Neochirus bella. Based on our examined materials, the higher count of longitudinal scale series (41–44) matched the description of N. brachyptera.

Neomerinthe ignea Matsumoto, Muto & Motomura, 2023

Remarks: No specimen was examined in this study. The record was based on Matsumoto et al. (2023), in which some specimens were collected from Taiwanese waters.

Neomerinthe megalepis (Fowler, 1938)

Remarks: No specimen was examined in this study. The first record was reported by Chen (1981) based on six specimens (CAS27744, SDSC73-37, SIO80-206, SIO80-207, SIO80-208, SIO80-221) from Tungkang, Pingtung. But these specimens showed more scales in longitudinal series (36–41), a discrepancy from the 25–30 scales in the original description (Fowler 1938; Herre 1952). These specimens are actually N. kaufmani based on the number of longitudinal series. We also examined a specimen (ASIZP0064297) originally identified as N. megalepis and found that it is a misidentification of N. kaufmani. However, some specimens of N. megalepis were collected from Taiwanese waters by Matsumoto and Motomura (2024).

Parascorpaena maculipinnis Smith, 1957

Remarks: Five specimens were examined in this study (Tables 3, S1). Parascorpaena mcadamsi and P. maculipinnis were distinguished from other species of Parascorpaena by having 15 to 16 pectoral-fin rays, supraocular tentacle absent or very short, presence of a spine below eye, and presence of a distinct black blotch on spinous dorsal fin in male. Parascorpaena maculipinnis can be further distinguished from P. mcadamsi by having three suborbital spines (vs. two in P. mcadamsi) (Chou and Liao 2022).

Phenacoscorpius megalops Fowler, 1938

Remarks: No specimen was examined in this study. The record was based on Chen (1981). All specimens (CAS47299) were collected from Taiwan, and the figure and description in Chen (1981) corroborate the original description of P. megalops.

Phenacoscorpius sp.

(Fig. 4a)

Remarks: The counts of dorsal, pectoral, and anal fins are provided in table 3. The number of total vertebrae is 25. The lateral line is incomplete with only two lateral line scales anteriorly, a diagnostic characteristic of Phenacoscorpius. The palatine teeth are present. The specimen could not be identified to any valid species of Phenacoscorpius according to diagnostic characters (Motomura 2008; Motomura and Last 2009; Motomura et al. 2012a b; Wibowo and Motomura 2017). This species can be clearly distinguished from other congeners by six anal fin rays (except for P. adenensis and P. eschmeyeri), two lateral line scales (vs. more than three in other congeners), and presence of black spots on upper pectoral fin (Fig. 4a) (vs. absence in other congeners). In addition, this species could be distinguished from P. megalops by presence of palatine teeth (vs. absence in P. megalops). The taxonomic status of this specimen is unclear and further study using more specimens is needed.

Fig. 4.

Fig. 4.

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Preserved specimens of the three taxonomically uncertain species. (a) Phenacoscorpius sp., NMMB-P036068, 46 mm SL. (b) Scorpaenopsis sp., DOS08531, 137 mm SL. (c) Sebastapistes sp., DOS08051, 17 mm SL.

Pontinus macrocephalus (Sauvage, 1882)

Remarks: No specimen was examined in this study. Pontinus macrocephalus was morphologically similar to P. tentacularis but could be distinguished by the counts of dorsal-fin rays (10 in P. macrocephalus vs. 9 in P. tentacularis) and pectoral-fin rays (17 in P. macrocephalus vs. 16 in P. tentacularis) (Eschmeyer 1969; Eschmeyer and Randall 1975). We have checked all specimens of P. macrocephalus from the collections in Taiwan, but the counts of dorsal-fin rays and pectoralfin rays are identical to P. tentacularis. The occurrence of P. macrocephalus in Taiwan needs to be confirmed.

Pterois lunulata Temminck & Schlegel, 1843

Remarks: Pterois lunulata, P. russelii and P. volitans differ from other congeners of Pterois by having fewer pectoral-fin rays (< 15). Both P. lunulata and P. russelii lack black spots on dorsal rays, anal rays, and the caudal fin. Pterois lunulata can be distinguished from P. russelii by more dorsal-fin rays (10–11 vs. 11– 12 in P. russelii) and pectoral-fin rays (13–14 vs. 12– 13). According to the overlapping meristic characters, we agreed with Wilcox et al.’s (2018) opinion that P. lunulata might be a junior synonym of P. russelii.

Rhinopias frondosa (Günther, 1892)

Remarks: Chen (2003) first recorded R. aphanes from Penghu, but we re-identified it from the photo as R. frondosa based on the shape of the caudal fin (margin of fin membrane between soft rays strongly notched in R. aphanes vs. weakly notched to non-notched in R. frondosa) (Motomura and Johnson 2006).

Scorpaena pepo Motomura, Poss & Shao, 2007

Remarks: Five specimens were examined in this study (Tables 3, S1). Scorpaena pepo is closely related to S. onaria occurring in Taiwan, but it can be distinguished from S. onaria by its 16 pectoral-fin rays (Table 3) (Motomura et al. 2007).

Scorpaenodes hirsutus (Smith, 1957)

Remarks: No specimen was examined in this study. The first record was reported by Chen (1981) based on a single specimen collected from Hengchun, Pingtung. Hoshino and Motomura (2021) recently reported another specimen from Hong Chai, Pingtung.

Scorpaenodes minor (Smith, 1958)

Remarks: Motomura et al. (2009b) reported the new record of Scorpaenodes albaiensis from East Asia and re-identified Taiwanese S. minor as S. albaiensis. They indicated the two species are closely related and similar in overall body appearance, but S. minor can be distinguished from S. albaiensis by larger and fewer scales on longitudinal series (27–32 vs. 37–42 in S. albaiensis). We have examined two specimens (NMMB-P007032) with 31–32 scales in longitudinal series that conform with S. minor.

Scorpaenodes quadrispinosus Greenfield & Matsuura, 2002

Remarks: No specimen was examined in this study. The Taiwanese record was reported by Motomura et al. (2010a) based on two specimens collected from southern Taiwan.

Scorpaenopsis obtusa Randall & Eschmeyer, 2002

Remarks: No specimen was examined in this study. Motomura et al. (2011) reported a single specimen (NMMB-P0007637) of S. obtusa from Dongsha in the South China Sea, but no specimen was collected from waters around the Taiwan Island. This species can be clearly distinguished from congeners by its distinct short snout. It has been recorded but misidentified as the juvenile of Synanceia verrucosa and Scorpaenopsis diabolus in the plates of Shao et al. (1993, p. 55) and Chen et al. (2010, p. 99), respectively, both from Kenting, Pingtung.

Scorpaenopsis orientalis Randall & Eschmeyer, 2002

Remarks: This species belongs to the Scorpaenopsis oxycephala species group due to its long snout (Randall and Eschmeyer 2002). This species group contains five species, including S. cacopsis, S. cirrosa, S. papuensis, S. orientalis and S. oxycephala. Scorpaenopsis orientalis can be distinguished from others by its 18 pectoral fin rays, and V-shaped and deep interorbital space.

Scorpaenopsis sp.

(Fig. 4b)

Remarks: The counts of dorsal, pectoral, and anal fins are provided in table 3. The specimen has three suborbital spines. This species belongs to the Scorpaenopsis oxycephala species group due to its long snout (Randall and Eschmeyer 2002), and it is morphologically most similar to S. oxycephala and S. papuensis. Scorpaenopsis sp. can be distinguished from S. oxycephala by having fewer scales in longitudinal series (53 vs. 59–67 in S. oxycephala) and differs from S. papuensis by absence of an occipital pit (vs. presence of a shallow occipital pit). The taxonomic status of this specimen is unclear, and further studies are needed.

Sebastapistes mauritiana (Cuvier, 1829)

(Fig. 5)

Remarks: The species is newly recorded from Taiwanese waters based on a specimen in the current study. The counts of dorsal, pectoral, and anal fins are provided in table 3. This species differs from congeners of Sebastapistes by having a strong coronal ridge with a spine (Fig. 4b).

Motomura et al. (2014) have synonymized Scorpaena hatizyoensis with S. mauritiana. There is one specimen of S. hatizyoensis (NTMP0678) collected from Taiwan in 1945 which is supposed to be S. mauritiana. However, we re-identified this specimen as a misidentification of Scorpaena neglecta according to the image of NTMP0678 in the National Taiwan Museum digital archive information system, with a large body size of 370 mm in total length.

Fig. 5.

Fig. 5.

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Fresh specimen of a new record species, Sebastapistes mauritiana, from Taiwanese waters. DOS08337, 71 mm SL. (a) Lateral view, (b) dorsal view. White arrows indicate the coronal ridge.

Sebastapistes sp.

(Fig. 4c)

Remarks: The counts of dorsal, pectoral, and anal fins are shown in table 3. The specimen was identified as a member of Sebastapistes by 12 dorsal-fin spines, presence of teeth on palatines, posterior lacrimal spine directed posteroventrally, complete lateral line, and lack of a deep occipital pit (Motomura et al. 2014). This species and Sebastapistes strongia can be distinguished from other congeners based on pectoral-fin rays usually 15 (vs. usually 16 in other congeners), and presence of several white bands on the mandible (vs. absence), but Sebastapistes sp. can be further distinguished from S. strongia by three suborbital spines (vs. one in S. strongia). The taxonomic status of this specimen is unclear, and further studies are needed.

Sebastes joyneri Günther, 1878

Remarks: No specimen was examined in this study. The record of the species was reported by Chen (1969), but no specimen is known from collections. The presence of S. joyneri in Taiwanese waters is uncertain.

Sebastiscus tertius (Barsukov & Chen, 1978)

Remarks: This species was similar to Sebastiscus vibrantus, and both species can be distinguished from the other two congeners by having 19 pectoral-fin rays (vs. 17 in S. albofasciatus and 18 in S. marmoratus). Sebastiscus tertius differs from S. vibrantus by having a scaled area on the suborbital bone extending over the anterior margins of the orbit (vs. scaled area does not extend over the anterior margins of the orbit), parietal spine equal to nuchal spine (vs. parietal spine longer than nuchal spine), and a shorter pectoral fin base (9.2– 11.7% vs. 11.1–13.2% standard length in S. vibrantus) (Morishita et al. 2018). According to our examined materials, the characters matched the description of S. tertius, including the state of the scaled area on the suborbital bone, equal lengths of parietal and nuchal spines, and the shorter pectoral fin base of 9.9–10.4% standard length.

Sebastiscus vibrantus Morishita, Kawai & Motomura, 2018

Remarks: The comparison between Sebastiscus tertius and S. vibrantus was provided in the remark of Sebastiscus tertius. According to our examined materials, the characters matched the description of S. vibrantus, including the state of scaled area on suborbital bone, parietal and nuchal spine, and longer pectoral fin base of 12.4–12.7% SL (Morishita et al. 2018).

DISCUSSION

DNA barcoding of the Scorpaenidae

DNA barcoding has been well known as an effective and rapid tool for identification at the species level and discovery of the biodiversity of marine fishes (e.g., Ward et al. 2005; Steinke et al. 2009; Lakra et al. 2011; Zhang and Hanner 2011 2012; Weigt et al. 2012; Wang et al. 2018; Xing et al. 2018; Thu et al. 2019; Fadli et al. 2020; Huang et al. 2023). However, DNA barcoding for species identification also has its limitations. It works only when COI sequences exhibit sufficient interspecific genetic variation. In some cases, the genetic variations of the COI gene are unremarkable, such as tunas (Thunnus spp.) and most hamlets (Hypoplectrus spp.) (García-Machado et al. 2004;

Ward et al. 2005; Viñas and Tudela 2009; Victor 2012; Victor and Marks 2018). Additionally, the application of molecular identification can only work when reliable reference sequences with voucher specimens are available (Ruedas et al. 2000; Harris 2003; Savolainen et al. 2005; Ward et al. 2005 2009; Ratnasingham and Hebert 2007; Mitchell 2008; Pleijel et al. 2008; Wang et al. 2012). Moreover, misidentifications of voucher specimens in the Scorpaenidae could also be a serious concern when no taxonomist is involved in the molecular studies (Poss 1999; Randall and Eschmeyer 2002; Randall 2005a b). In the present study, the sequences of 22 species belonging to nine genera were found to have more than one BLAST result with similarities ≥ 98% (Table S2), probably due to misidentifications from online databases. However, voucher specimen photos of some sequences are not available, and this creates a significant drawback to further verification and re-identification. In this study, a reliable DNA barcoding library of the Scorpaenidae from Taiwanese waters has been established based on voucher specimens.

The application of DNA barcoding for demarcating species relies on a gap between minimum interspecific and maximum intraspecific genetic divergences (i.e., barcode gap) (Hebert et al. 2003a 2004; Barrett and Hebert 2005; Wiemers and Fiedler 2007). For fish, the conspecific chance is very high when the genetic divergence of COI sequences is less than 2% (Hebert et al. 2003a b; Ward et al. 2009). In the current study, the mean interspecific genetic distance (15.61%) was much higher than the mean intraspecific genetic distance (0.22%) (Fig. 3, Table 2). Although a few species pairs of the Scorpaenidae from Taiwanese waters exhibit low genetic variation in COI sequences (less than 2%), reciprocal monophylies were consistently observed in the ML tree (Fig. 2), and these species pairs can be distinguished by their morphological characteristics (Matsunuma et al. 2017; Morishita et al. 2018; Wilcox et al. 2018). The only case of Pterois lunulata vs. P. russelii may represent an exception, in which the former is potentially a junior synonym of the latter (Wilcox et al. 2018). Some species pairs of the genus Sebastes also showed low genetic variation in COI gene, but could be clearly distinguished by their morphology, implying recent diversifications or contemporary/ historic hybridizations between closely related species (Hyde and Vetter 2007; Steinkeet al. 2009; Zhang et al. 2013; Muto and Kai 2023). Most species pairs with low genetic variation observed in this study (except for P. lunulata vs. P. russelii) are probably a consequence of recent diversifications. Additionally, the majority of high interspecific genetic variations were contributed by species pairs within the genus Sebastapistes, such as S. mauritiana vs. S. strongia (32.3%), S. mauritiana vs. S. fowleri (30.4%), S. mauritiana vs. Sebastapistes sp. (30.0%), S. mauritiana vs. S. tinkhami (27.8%), S. mauritiana vs. S. cyanostigma (27.0%), and S. tinkhami vs. Sebastapistes sp. (25.8%). Taxonomic studies of Sebastapistes are scarce, and its taxonomic status needs to be re-examined based on comprehensive sampling (Motomura et al. 2014).

The potential records of the Scorpaenidae in Taiwanese waters

To date, 85 species of 29 genera of the Scorpaenidae have been recorded in Taiwanese waters. Several new records originally known from adjacent waters have been reported in recent studies. Sebastes thompsoni was previously considered to inhabit cold waters in the northwestern Pacific but has been recently observed in northern Taiwan (Chou and Tang 2021); Scorpaenopsis orientalis was considered a Japanese endemic species but has subsequently been discovered in southern Taiwan (Randall and Eschmeyer 2002; Koeda et al. 2019). Scorpaenopsis cotticeps was only known from Japan and the Philippines, and had never been reported from Taiwan until Chou (2021). Lioscorpius longiceps, Lythrichthys dentatus, and Scorpaenodes corallinus, which are distributed in the western Pacific, may also occur in Taiwan (Randall and Lim 2000; Nakabo and Kai 2013; Wada et al.2021; Hoshino et al. 2023). More scorpaenids are waiting to be discovered in Taiwanese waters.

CONCLUSIONS

In this study, we updated the checklist of the Scorpaenidae from Taiwan. In total, 296 specimens of 85 species placed in 29 genera were examined and literature reviewed using morphological and molecular approaches. Among the 85 species, Sebastapistes mauritiana (Cuvier, 1829) is a new record, and the taxonomic status of three species in the genera Phenacoscorpius, Scorpaenopsis, and Sebastapistes remain uncertain. A total of 118 COI sequences belonging to 66 species of 28 genera are generated based on properly identified voucher specimens in the present study. The COI sequences of Parascorpaena maculipinnis, Scorpaena pepo, and Scorpaenopsis orientalis are new to online databases (GenBank and BOLD systems). For K2P distance of the COI gene, the mean interspecific genetic distance (15.61%) was higher than the mean intraspecific genetic distance (0.22%), representing a clear barcode gap which makes DNA barcoding feasible within the Scorpaenidae. Identifying Scorpaenidae species through morphology can be challenging for non-specialists, whereas DNA barcoding offers a rapid and powerful tool that does not require taxonomic expertise.

Supplementary materials

Fig. S1.

Photographs of some sequenced/examined voucher specimens. Following the scientific name are the catalog number, standard length (SL), and GenBank accession number of the specimen. NA for the accession number indicates that the specimen was used solely for morphological examination and was not sequenced.

(download) (4.3MB, pdf)
Table S1.

List of species, body size, catalog number of specimens, and their accession numbers. Species without examined specimen are denoted by a hyphen (-) representing data as “not available” .

(download) (55KB, docx)
Table S2.

List of morphological and molecular identifications. The listed sequences included more than one species after BLAST.

(download) (26.6KB, docx)

Acknowledgments

We are grateful to two anonymous reviewers for their constructive comments; S.-P. Huang (ASIZP), C.-H. Chan, T.-W. Wu, H.-C. Ho (NMMB-P), for their curatorial assistances; C.-C. Ku, C.-N. Tang, H.-C. Chen, J.-S. Lin, N.-W. Lai, S.-L. Ng, Y.-H. Yu for collecting specimens. This study is partly support by a grant from National Science and Technology Council to TYL (111-2611-M-110-026).

Footnotes

Authors’ contributions:TKC and TYL designed the study. TKC, WCH, and WCJ collected, identified, and examined the specimens. TKC, WCH and WCJ prepared the manuscript. All authors revised the manuscript and approved the final version.

Competing interests: The authors declare that they have no competing interests.

Availability of data and materials:All data are available in the paper.

Consent for publication:Not applicable.

Ethics approval consent to participate:Not applicable.

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Associated Data

This section collects any data citations, data availability statements, or supplementary materials included in this article.

Supplementary Materials

Fig. S1.

Photographs of some sequenced/examined voucher specimens. Following the scientific name are the catalog number, standard length (SL), and GenBank accession number of the specimen. NA for the accession number indicates that the specimen was used solely for morphological examination and was not sequenced.

(download) (4.3MB, pdf)
Table S1.

List of species, body size, catalog number of specimens, and their accession numbers. Species without examined specimen are denoted by a hyphen (-) representing data as “not available” .

(download) (55KB, docx)
Table S2.

List of morphological and molecular identifications. The listed sequences included more than one species after BLAST.

(download) (26.6KB, docx)

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