A New Species of Diaphus Associated with Seamounts of the Emperor Chain, North-Western Pacific Ocean (Teleostei: Myctophiformes: Myctophidae) †
by
and
Artem M. Prokofiev
1,2, 
Olga R. Emelyanova
3,
Alexei M. Orlov
2,4,5,6,7,8,* and
Svetlana Y. Orlova
3,9 1
Laboratory of Ecology of Aquatic Communities and Invasions, A.N. Severtsov Institute of Ecology and Evolution of the Russian Academy of Sciences, Leninsky Prospekt 33, 119071 Moscow, Russia
2
Laboratory of Oceanic Ichthyofauna, P.P. Shirshov Institute of Oceanology of the Russian Academy of Sciences, 117218 Moscow, Russia
3
Laboratory of Molecular Genetics, Russian Federal Research Institute of Fisheries and Oceanography, 107140 Moscow, Russia
4
Department of Marine Fishes of the Russian Far East, Russian Federal Research Institute of Fisheries and Oceanography, 107140 Moscow, Russia
5
Department of Ichthyology and Hydrobiology, Tomsk State University, 634050 Tomsk, Russia
6
Department of Ichthyology, Dagestan State University, 367000 Makhachkala, Russia
7
Laboratory of Marine Biology, Caspian Institute of Biological Resources, Dagestan Federal Research Center of the Russian Academy of Sciences, 367000 Makhachkala, Russia
8
Laboratory of Behavior of Lower Vertebrates, A.N. Severtsov Institute of Ecology and Evolution of the Russian Academy of Sciences, 119071 Moscow, Russia
9
Laboratory of Ecology of Coastal Bottom Communities, P.P. Shirshov Institute of Oceanology of the Russian Academy of Sciences, 117218 Moscow, Russia
*
Author to whom correspondence should be addressed.
†
Publications: urn:lsid:zoobank.org:pub:09E84AEF-4422-4E55-9959-A728620DC465.
J. Mar. Sci. Eng. 2022, 10(1), 65; https://doi.org/10.3390/jmse10010065
Submission received: 10 November 2021
/
Revised: 4 December 2021
/
Accepted: 21 December 2021
/
Published: 5 January 2022
(This article belongs to the Special Issue Deep-Sea Fish and Fisheries)
Abstract
:A new species, Diaphus balanovi, is described based on 35 specimens collected over the Emperor Seamount Chain in the north-western Pacific Ocean. It belongs to the D. fulgens species complex and is most similar to D. kuroshio both morphologically and genetically. Nevertheless, the new species can be distinguished from D. kuroshio by its higher gill-rakers count, large luminous scale at PLO, large Dn, somewhat higher position of SAO3, otolith shape, and larger absolute size. The CO1 mtDNA sequence of D. balanovi differs by 16 substitutions from that of D. kuroshio. Diaphus balanovi may represent a benthopelagic derivate of D. kuroshio endemic to the Emperor Seamounts.
1. Introduction
The genus Diaphus is the largest genus of the lanternfish family Myctophidae, with some 76 recognized recent species distributed circumglobally in temperate and tropical latitudes [1,2,3] that account for about one-third of the species diversity of the family [4]. Although this genus inhabits mainly mesopelagic habitats, certain species are associated with the slope and seamounts. These species usually reach a much larger size than their mesopelagic relatives, and often have a restricted or patchy distribution [1,5], though they may be locally abundant [6,7].
There is insufficient knowledge regarding the lanternfish fauna associated with the seamounts of the North Pacific. Among nearly 90 myctophid species recorded over Emperor and Hawaiian underwater ridges and nearby oceanic waters, 29 species of Diaphus were listed, but identification of some of these species requires confirmation [8,9,10]. Although published data on the species composition and abundance of the Diaphus species in the northwestern Pacific are rather controversial [11,12,13,14,15], D. theta and D. kuroshio were reported as the most common and abundant species within the transition zone, including waters over the Emperor Seamounts.
In April 2019, during a research cruise onboard R/V “Professor Kaganovsky” [16] several specimens of an unidentified Diaphus species were found in catches on the Lira and Ojin Seamounts [17], along with Diaphus kuroshio and D. metopoclampus. Although these specimens can be identified as D. kuroshio using the taxonomic key developed by Becker [1], they can be distinguished from D. kuroshio both morphologically and genetically. The main purpose of this paper is to describe a new species and to provide its phylogenetic position among congeners, based on the analysis of the CO1 mtDNA gene.
2. Materials and Methods
Voucher specimens were deposited in the Shirshov Institute of Oceanology, Russian Academy of Sciences, Moscow (IORAS), and Atlantic Branch of the Russian Federal Research Institute of Fisheries and Oceanography, Kaliningrad (AtlantNIRO). Catalog numbers and label data are presented under species description. Holotype and paratypes have been registered in ZooBank (urn:lsid:zoobank.org:act:99C12AF4-1E9B-4AB3-B7EE-3CBD95EC8851). Tissue samples used for genetic analysis are listed in Table 1. Methods of counts and measurements are standard [18], with the following additions: eye-maxilla distance was measured from the posteriormost border of the eye to the hindmost tip of the upper jaw; and the dorsal-adipose distance was measured from base of last dorsal-fin ray to beginning of the adipose fin. Measurements were taken point to point. As distal portions of fin rays are usually missing in museum specimens, branched and unbranched rays were not separated in fin-ray counts as is usual in myctophid descriptions [19,20]. Lateral-line organs are horseshoe-like structures, lying below lateral-line scales (which are usually missing in museum specimens). Photophore and luminous tissue definitions follow Nafpaktitis [19]. Otolith terminology follows Schwarzhans [3]. Other abbreviations used are SL, standard length; and R/V, research vessel.
Molecular Data and Phylogenetic Analysis
All tissue samples were fixed in volumes of 96% ethanol at least five times larger than the sample volume. Fixed samples were stored at −20 °C; ethanol was changed approximately one month after collection, and again after one year. DNA was extracted using the Wizard SV 96 Genomic DNA Purification System (Promega Corporation, Madison, WI, USA) according to the manufacturer’s manual. All molecular genetic studies (DNA extraction, polymerase chain reaction (PCR), PCR product purification, and nucleotide sequencing) were performed using standard molecular genetic techniques [21]. Cytochrome oxidase subunit I (CO1) fragment of 530 b.p. was amplified with a primer complex of VF2_t1, FishF2_t1, FishR2_t1, FR1d_t1 [21,22]. Amplification was conducted in a volume of 15 μL with 90 ng total DNA, buffer 1x, 2.5 mM MgCl2, 0.2 mM dNTP, 0.5 mM of each primer, and 0.75 U μL−1 Color Taq polymerase. Cycling consisted of 5 min at 95 °C, followed by thirty-five cycles of 30 s each at 95 °C, 45 s at 52 °C, 60 s at 72 °C, and a final extension for 12 min at 72 °C. All resulting amplicons were purified by ethanol precipitation [23].
Purified fragments were sequenced from both strands by Applied Biosystems BigDye Terminator v3.1. kit (Applied Biosystems, Foster City, CA, USA) with capillary electrophoresis on ABI3500 Genetic Analyzer (Thermo Fisher Scientific, Waltham, MA, USA) in VNIRO Laboratory of Molecular Genetics.
The resulting sequences were assembled in Geneious 6.5.0 (Biomatters, Auckland, New Zealand) [24] and aligned with the “ClustalW” built-in algorithm. They were subsequently translated into the necessary format for constructing a haplotype network in the PopArt program (Allan Wilson Centre Imaging Evolution Initiative, Otago, New Zealand) [25]. The FaBox 1.41 converter was used to convert the fasta file to the format required for calculation [26]. A network of haplotypes was constructed based on the maximum parsimony method using TCS v.1.21 software (Computational Science Laboratory, Provo, UT, USA). DnaSP v. 5.10.01 software (University of Barcelona, Barcelona, Spain) was used for the analysis of the average number of nucleotide substitutions and the number of haplotypes in samples [27].
Data processing was performed, and genetic distances were calculated by using Geneious 6.0.5 software (Biomatters, Auckland, New Zealand) based on the Neighbor–Joining method with Lobianchia dofleini and L. gemellari as outgroups [28] with the use of a Genetic Distance HKY model and 1000 bootstrap replicates [29].
Data on CO1 sequences of outgroups and congeners of a new species (sister groups involved in analysis for comparative purpose) were taken from the open BOLD (https://www.boldsystems.org/) database (date of last accession 9 November 2021).
3. Results
Diaphus balanovi sp. nov.
urn:lsid:zoobank.org:act:99C12AF4-1E9B-4AB3-B7EE-3CBD95EC8851
Holotype: IORAS S.0124, mature male, 96 mm SL (Figure 1A), 37°56′36″–37°57′24″ N, 170°24′42″–170°24′54″ E, 1030 m depth, bottom temperature 3.35 °C, R/V “Professor Kaganovsky”, bottom trawling no. 109, 12 April 2019, DNA sample no. 31.
Figure 1.
Diaphus balanovi sp. nov., holotype, IORAS S.0124, 96 mm SL. (A) Habitus. (B) Scheme of disposition of photophores.
Figure 1.
Diaphus balanovi sp. nov., holotype, IORAS S.0124, 96 mm SL. (A) Habitus. (B) Scheme of disposition of photophores.
Paratypes: IORAS S.0117, nine specimens, 84–94 mm SL, 36°48′8″–36°47′7″ N, 171°23′06″–171°23′30″ E, 642–653 m depth, bottom temperature 4.7 °C, R/V “Professor Kaganovsky”, bottom trawling no. 108, DNA samples nos. 96–99. IORAS S.0118, 25 specimens, 83–98 mm SL, 36°48′48″–36°47′54″ N, 171°23′00″–171°23′30″ E, 643–649 m depth, bottom temperature 4.9 °C, R/V “Professor Kaganovsky”, bottom trawling no. 107, DNA sample no. 130.
Comparative material examined:Diaphus kuroshio: IORAS S.0112, 44 mm SL, 37°56′36″–37°57′24″ N, 170°24′42–170°24′54″ E, 1030 m depth, bottom temperature 3.35 °C, R/V “Professor Kaganovsky”, bottom trawling no. 109, 12 April 2019, DNA sample no. 28. IORAS, uncatalogued, three specimens, 33–47 mm SL, Kuroshio Zone, samplings of A.S. Sokolovskiy 1965–1970, no more precise data, see Parin and Sokolovskiy [30]. Diaphus rafinesquii: AtlantNIRO, uncatalogued, 36 specimens, 28–82 mm SL, from different stations in Gulf Stream Zone, off Morocco and Mauritania. Diaphus theta: IORAS S.0131, 10 specimens, 48–56 mm SL, 43°10′00″–43°09′12″ N, 152°24′00″–152°29′00″ W, 5100 m depth, trawling depth 37 m, surface temperature 9.1 °C, R/V “Professor Kaganovsky”, mid-water trawling no. 99, 28 March 2019, DNA samples nos. A19–A25. IORAS S.0132, 44 specimens, 34–65 mm SL, 41°16′12″–41°15′06″ N, 161°12′00–161°16′48″ W, 5500 m depth, trawling depth 32 m, surface temperature 8.7 °C, R/V “Professor Kaganovsky”, mid-water trawling no. 100, 31 March 2019, DNA samples nos. A61–A70.
Diagnosis: The new species belongs to the Diaphus fulgens species complex as defined by Nafpaktitis et al. [2] and is readily distinguished from most of its members, except D. rafinesquii (North Atlantic and the Mediterranean Sea) and D. kuroshio (Kuroshio Zone in the North Pacific), by a higher gill-raker count (24–28, almost always 25 or more vs. 14–20). This count exceeds the limits known for D. rafinesquii and D. kuroshio (22–25). The new species shares a close resemblance to D. kuroshio in the disposition of photophores (VO5, SAO1, and SAO2 on the same line, AOa1 below level of SAO2, Prc4 its own diameter below lateral line), but differs from that species by the large luminous scale at PLO, large Dn, somewhat higher position of SAO3, otolith shape, and larger absolute size (at least 98 mm SL vs. about 70 mm). Otolith of Diaphus balanovi can be distinguished from the otoliths of other species united in the Diaphus theta otolith group of Schwarzhans [3] by following characters in combination: compressed shape with very weakly and uniformly convex outer face, rather steep and irregular predorsal rim, pronounced postdorsal angle, moderately long rostrum (18–19% of otolith length), and seven or eight (usually eight) denticles along the ventral rim.
Description: Measurements are shown in Table 2. The snout is very deep and short, almost truncate, 4.0 (2.9–4.0) in the eye diameter. The eye is large, 2.9 (2.5–3.0) in head. The mouth is large, with the upper jaw 1.5 (1.4–1.7) in head length; the distance from the posterior border of the eye to the posterior tip of the maxilla is 2.7 (2.4–3.6) in the upper-jaw length and 1.4 (1.3–2.0) in the eye diameter. Jaw teeth are villiform, in a narrow band; the teeth of the innermost row in the posterior half of the premaxilla are somewhat enlarged, broadened at the bases and hooked at the tips; the teeth of the innermost row on the dentary bone are very slightly enlarged. The vomer bears few minute teeth; palatines with a narrow band of villiform teeth tapering caudally; mesopterygoid with a large elongate-oval patch of uniformly small villiform teeth. The postero-dorsal margin of the operculum concave; opercular lobe tapering with distalmost point rounded (from tapering and pointed to short and rounded). Gill rakers on the first arch are 28 (25–28, 24 in 1 of 35 specimens), flattened, their tips rounded, dorsal and ventral surface bearing villiform denticles, and their frequency distributions are: 8 + 1 + 15 (1), 8 + 1 + 16 (15), 8 + 1 + 17 (14), 7 + 1 + 18 (1), 8 + 1 + 18 (2), 9 + 1 + 18 (2). Gill filaments from the upper half of the first ceratobranchial 1.5× those from lower half in length. Pseudobranchial filaments are long but not numerous, 14 (12–20) in number.
The dorsal and anal fins have 13 (12–14) rays. The origin of the dorsal fin is slightly in advance of the base of the pelvic fin (rarely over to just behind). The origin of the anal fin is behind the base of the dorsal fin. The pectoral fin has 10 (10–12) rays, the pelvic fin has eight (eight, rarely seven or nine (one side only)) rays. The tips of the pectoral and pelvic fins are broken in all specimens available. The base of the adipose fin is over the end of the anal fin. Lateral line organs 35 (33–36).
Luminous organs (Figure 1 and Figure 2) are characterized by: Dn is round, directed forward, deeply recessed in cup-shaped structure, slightly exceeding the nasal rosette in diameter; Vn is sexually dimorphic, and is large, elongate, thickened and reaching ventral border of nasal organ anteriorly, extending backward to So in adult males, and is small and thin at the antero-ventral margin of eye in females (Figure 2). Therefore, it is nearly as large as the PLO, situated mid-way between the verticals of the mid-eye and posterior border of pupil (mid-way between these verticals to almost on vertical of posterior border of pupil). Ant and Suo absent.
Figure 2.
Diaphus balanovi sp. nov., luminous glands of the head (Dn, Vn and So). (A) Male, holotype, IORAS S.0124, 96 mm SL. (B) Female, paratype, IORAS S.0118, 96 mm SL. Scale bar: 5 mm.
Figure 2.
Diaphus balanovi sp. nov., luminous glands of the head (Dn, Vn and So). (A) Male, holotype, IORAS S.0124, 96 mm SL. (B) Female, paratype, IORAS S.0118, 96 mm SL. Scale bar: 5 mm.
The PLO is closer to the base of the pectoral fin than to the lateral line (on the same distance or slightly closer to pectoral-fin base). The luminous scale at the PLO much exceeds the diameter of the photophore. The VLO is equidistant from the lateral line and base of the pelvic fin. The VO3 is elevated, mid-way between the VLO and pelvic-fin base. The SAO series forms an obtuse angle between the SAO2 and SAO3. The VO5, SAO1 and SAO2 are on the same diagonal line, with the SAO2 being closer to the SAO1 than to the SAO3. The SAO3 is immediately below the lateral line (immediately to less than its own diameter), and is one lateral-line scale in advance of the anal-fin origin. The AO is 5 + 5. The first AOa is abruptly elevated, at half of its own diameter below the horizontal through SAO2; the last AOa is elevated, at the level of the SAO2. The line through the AOa1 and AOa2 run far below the SAO3. The Pol is about one of its own diameter below the lateral line. The AOp is behind the base of the anal fin. The Prc interspace is progressively wider, with the space between the Prc1 and Prc2 being twice that of the interspace between the Prc1 and last AOp, and the Prc4 being its own diameter below the lateral line.
Otolith (based on four paratypes S.0118 of 88–90 mm SL) (Figure 3): Length × depth = 5.0–5.5 × 3.5–4.0 mm (ratio: 1.3–1.5). The predorsal rim is weakly to moderately expanded, and somewhat irregular; the postdorsal angle is strongly developed, situated above the posterior half of cauda; and the postdorsal depression is moderately to very shallow. The rostrum and antirostrum are conspicuous; the rostrum length is 5.3–5.6 times in the otolith length; and the excisura is broad and sharp. The posterior rim is convex, with the cauda not closely approached to it. The ventral rim has seven to eight (usually eight) massive blunt denticles (some of which can be poorly expressed). The inner face is almost flat; the sulcus are 1.3 times in the otolith length, ostium:cauda length = 1.75; ostial colliculum length:height = 2.7, caudal colliculum length:height = 2.5; the dorsal rim of the ostium lobate, the ostial colliculum is somewhat reduced dorsally; and the pseudocolliculum is well-developed. Dorsal depression is moderate, and the ventral furrow is distinct. The outer face is weakly and regularly convex, with the umbo indistinct; the ratio of the otolith depth to thickness is 4.4.
Figure 3.
Diaphus balanovi sp. nov., right otolith (sagitta), inner face (top), and dorsal view (bottom). Scale bar: 1 mm.
Figure 3.
Diaphus balanovi sp. nov., right otolith (sagitta), inner face (top), and dorsal view (bottom). Scale bar: 1 mm.
Etymology. The new species is named after Dr. Andrei A. Balanov, an ichthyologist of A.V. Zhirmunsky National Scientific Center of Marine Biology, Far Eastern Branch of the Russian Academy of Sciences, Vladivostok, in honor of his significant contribution to studies of the North Pacific ichthyofauna, including mesopelagic fishes and fishes of the Emperor Seamount Chain in particular.
4. Discussion
Nafpaktitis [20] defined several species complexes and groups in Diaphus including the Diaphus theta–fulgens species complex which correlates with the So-group of Kawaguchi and Shimizu [11]. This species assemblage can be characterized by the presence of So, sexually dimorphic Vn, hooked and broad-based teeth in the innermost row in the posterior half of the premaxilla, and reduced vomerine dentition. Most of the species (excluding the aberrant D. vanhoeffeni) are also characterized by the recessed Dn directed forward, and Vn well separated from Dn. Later, Nafpaktitis et al. [2] further divided this complex into the D. fulgens and D. theta species complexes based on the position of the first AOa photophore (raised in the D. fulgens complex vs. level with subsequent photophores in the D. theta complex). Diaphus balanovi should be attributed to the D. fulgens species complex and can be readily distinguished from most species (see Table 1 in ref. [2]) by a much higher gill-raker count (24–28 vs. 14–20). The only members resembling the new species in this character are D. rafinesquii from the North Atlantic Ocean and the Mediterranean Sea and D. kuroshio from the North Pacific Ocean, although both compared species additionally share a lower count (22–25) than is known for the new species. Although the range is slightly overlapping, the common limits are different: D. balanovi has normally 25 to 28 gill rakers on the first arch (24 in one of 35 specimens studied). Although 25 rakers were reported for both D. rafinesquii and D. kuroshio [31,32], this count rarely occurs in these species (none of our specimens of D. kuroshio show it, and only seven of 36 specimens of D. rafinequii studied by us possess 25 rakers).
Diaphus balanovi shows a close similarity with D. kuroshio but differs from that species by the increased size of the luminous scale at PLO (its vertical diameter 2.5–3.0 times exceeds the diameter of PLO vs. 0.75–1.5 times in D. kuroshio) (Figure 4), the higher position of SAO3 (less than its diameter below lateral-line canal vs. one to two diameters below), a larger Dn (equal to slightly larger than olfactory organ vs. about half-diameter of the latter in D. kuroshio), typically eight rakers on the first epibranchial (vs. commonly seven in D. kuroshio), details of the otolith structure, and a larger absolute size (maximum known SL 98 mm vs. about 70 mm in D. kuroshio). By the otolith structure, D. balanovi can be attributed to the Diaphus holti subgroup as defined by Schwarzhans [3] to which D. kuroshio is also included. However, D. balanovi is distinct from D. kuroshio by a longer rostrum (18–19% vs. 15% of otolith length), a pronounced postdorsal angle (vs. indistinct, with dorsal rim rather regularly convex), and much less convex outer face (Figure 6a,b in ref. [3]). The otolith of D. balanovi are most similar to that of D. holti, except for the weakly and uniformly convex outer face lacking the umbo (vs. outer face bent along the horizontal axis with the pronounced postcentral umbo in D. holti), and less expressed denticles on the ventral rim (Figures 4 and 5a,b in ref. [3]). However, the otolith subgroups of the D. theta group of Schwarzhans [3] do not correspond to the grouping proposed by Nafpaktitis et al. [2], and D. holti is fairly different from D. balanovi in possessing all AOa on the same level.
Figure 4.
The luminous scale at PLO (arrowed). (A) Diaphus balanovi sp. nov., holotype, IORAS S.0124. (B) Diaphus kuroshio, IORAS S.0112. Scale bars: (A) 3 mm; (B) 1.5 mm.
Figure 4.
The luminous scale at PLO (arrowed). (A) Diaphus balanovi sp. nov., holotype, IORAS S.0124. (B) Diaphus kuroshio, IORAS S.0112. Scale bars: (A) 3 mm; (B) 1.5 mm.
Diaphus balanovi is readily distinguished from D. rafinesquii by the relative position of VO5, SAO1, and SAO2 (on the same line vs. SAO2 below the line connecting VO5 and SAO1 in D. rafinesquii), less elevated AOa1 (line through AOa1 and AOa2 running far below SAO3 vs. through SAO3 or above it in D. rafinesquii), and the higher position of Prc4 (one its diameter below lateral line vs. two or three diameters in D. rafinesquii). The otolith of D. balanovi differs from the otolith of D. rafinesquii by its more pronounced postdorsal angle, and by a very slightly and uniformly convex (vs. bent along the horizontal axis) outer face lacking a distinct umbo (vs. pronounced postcentral umbo in D. rafinesquii) (Figures 7–10 in ref. [3]).
Molecular data strongly suggest a close relationship between D. balanovi and D. kuroshio (Figure 5). The CO1 mtDNA sequence of D. balanovi differs by 16 substitutions from that of D. kuroshio, contrary to 36 substitutions from that of D. holti and 46 substitutions from that of D. parri. The number of mutations between other species of the genus Diaphus analyzed by us ranges from 23 between D. brachycephalus and D. richardsoni to 51 between D. metopoclampus and D. effulgens. At the same time, the number of nucleotide substitutions between representatives of the same species in the vast majority of cases does not exceed five, and only in two cases is eleven (D. holti) or twelve (D. fulgens). This may indicate a recent time of divergence of D. balanovi and D. kuroshio.
All specimens of D. balanovi were collected by bottom trawls with non-closing openings. This does not exclude the possibility of catching specimens during the reeling-in of gear. However, D. balanovi was quite abundant in catches contrary to the single specimens of the mesopelagic D. kuroshio and D. metopoclampus in the same hauls. This may indicate that the new species can be associated with the seamounts and may have a benthopelagic mode of life. Its larger absolute size in comparison with its closest relative and possible ancestor (D. kuroshio) supports this assumption. Diaphus kuroshio is a mesopelagic species that migrates to the upper 100 m layer at night but does not appear on the surface [32]. No data on the vertical migrations of D. balanovi are known.
Author Contributions
Conceptualization, A.M.P. and A.M.O.; methodology, A.M.P., O.R.E. and S.Y.O.; software, O.R.E., A.M.O. and S.Y.O.; validation, A.M.P., O.R.E., A.M.O. and S.Y.O.; formal analysis, A.M.P., O.R.E. and S.Y.O.; investigation, A.M.P., O.R.E., A.M.O. and S.Y.O.; resources, A.M.P., O.R.E., A.M.O. and S.Y.O.; data curation, A.M.P., O.R.E., A.M.O. and S.Y.O.; writing—original draft preparation, A.M.P. and A.M.O.; writing—review and editing, A.M.P., A.M.O. and S.Y.O.; supervision, A.M.O. All authors have read and agreed to the published version of the manuscript.
Funding
Preparation of this paper was funded by the Ministry of Science and Higher Education, Russian Federation (grant No. 13.1902.21.0012, contract No. 075-15-2020-796).
Institutional Review Board Statement
Not applicable.
Informed Consent Statement
Not applicable.
Data Availability Statement
The specimens described in this study are available at Shirshov Institute of Oceanology, Russian Academy of Sciences, Moscow (IORAS) and at the Atlantic Branch of the Russian Federal Research Institute of Fisheries and Oceanography, Kaliningrad (AtlantNIRO). Voucher IDs: Diaphus balanovi sp. nov.: holotype IORAS S.0124, paratypes IORAS S.0117; Diaphus theta: IORAS S.0112, IORAS S.0131, IORAS S.0132, IORAS uncatalogued; Diaphus rafinesquii: AtlantNIRO uncatalogued. The COI sequences that support the findings of this study have been deposited in NCBI GenBank with the accession codes OL894503 (Diaphus_kuroshio), OL894504-OL894508 (Diaphus balanovi), and OL894509-OL894523 (Diaphus theta). The new species registration of Diaphus balanovi in Zoobank with LSID: urn:lsid:zoobank.org:act:99C12AF4-1E9B-4AB3-B7EE-3CBD95EC8851. The publication LSID: urn:lsid:zoobank.org:pub:09E84AEF-4422-4E55-9959-A728620DC465.
Acknowledgments
The authors are grateful to their colleagues from TINRO (Pacific Branch of VNIRO, Vladivostok, Russia) who took part in the processing of catches in the cruise of the R/V “Professor Kaganovsky” in 2019, storing samples in TINRO and transporting them to Moscow. The authors also thank Laura Haniford (Fish Ecology and Conservation Physiology Laboratory, Department of Biology, Carleton University, Ottawa, ON, Canada) and two anonymous reviewers for their valuable comments allowed for improvement of the manuscript. Special thanks to Kirill Kolonchin (VNIRO) and Aleksey Baitaliuk (TINRO), who provided participation of A.M.O. and S.Y.O. in the cruise.
Conflicts of Interest
The authors declare no conflict of interest.
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Figure 5.
Phylogenetic relationships of Diaphus balanovi sp. nov. and allied species by CO1 mtDNA sequences. Numbers indicate haplotypes (see Table 1 for explanation), numbers of substitutions are given in brackets.
Figure 5.
Phylogenetic relationships of Diaphus balanovi sp. nov. and allied species by CO1 mtDNA sequences. Numbers indicate haplotypes (see Table 1 for explanation), numbers of substitutions are given in brackets.
Table 1.
Information about CO1 sequences and respective samples used for molecular analysis (na = not available).
Table 1.
Information about CO1 sequences and respective samples used for molecular analysis (na = not available).
| No. Sample | No. Haplotype | Genbank Accession Number | Sampling Date | Locality | Country | Source |
|---|---|---|---|---|---|---|
| Lobianchia dofleini (outgroup 1) | ||||||
| 1 | 38 | SCAFB1054-09 | 8 September 2007 | Nova Scotia | Canada | BOLD Systems |
| 2 | 38 | MAECO457-09 | 28 June 2004 | Mid-Atlantic Ridge | na | BOLD Systems |
| Lobianchia gemellari (outgroup 2) | ||||||
| 3 | 29 | FCFMT219-09 | 15 July 2007 | Mediterranean Sea | Malta | BOLD Systems |
| 4 | 40 | SCAFB1032-09 | 7 September 2007 | Nova Scotia | Canada | BOLD Systems |
| 5 | 39 | MAECO460-09 | 28 June 2004 | Mid-Atlantic Ridge | na | BOLD Systems |
| Diaphus metopoclampus (sister group 1) | ||||||
| 6 | 28 | FCFMT123-09 | 16 June 2007 | Mediterranean Sea | Malta | BOLD Systems |
| 7 | 30 | FOAK120-10 | 15 June 2008 | Tasman Sea | Australia | BOLD Systems |
| 8 | 36 | MAECO111-06 | 30 June2004 | Mid-Atlantic Ridge | na | BOLD Systems |
| Diaphus effulgens (sister group 2) | ||||||
| 9 | 4 | GBGCA11052-15 | 24 May 2000 | New England | USA | BOLD Systems |
| 10 | 4 | UKFBJ499-08 | 24 May 2000 | New England | USA | BOLD Systems |
| 11 | 35 | MAECO105-06 | 28 June 2004 | Mid-Atlantic Ridge | na | BOLD Systems |
| Diaphus holti (sister group 3) | ||||||
| 12 | 33 | GBMIN125128-17 | July 2010 | Balearic Islands | Spain | BOLD Systems |
| 13 | 3 | GBGCA5261-13 | July 2010 | Balearic Islands | Spain | BOLD Systems |
| Diaphus brachycephalus (sister group 4) | ||||||
| 14 | 2 | MFLE267-13 | 26 January 2007 | Caribbean Sea | Belize | BOLD Systems |
| 15 | 2 | FOAO932-15 | 16 September 2014 | Western Indian Ocean | na | BOLD Systems |
| Diaphus fulgens (sister group 5) | ||||||
| 16 | 31 | GBGCA11055-15 | 2 December 2006 | Eastern tropical Pacific | na | BOLD Systems |
| 17 | 34 | LIDMA1136-12 | 10 March 1985 | Eastern tropical Pacific | na | BOLD Systems |
| Diaphus parri (sister group 6) | ||||||
| 18 | 5 | GBGCA11063-15 | na | na | na | BOLD Systems |
| 19 | 32 | GBGCA11065-15 | na | na | na | BOLD Systems |
| 20 | 5 | GBGCA11064-15 | na | na | na | BOLD Systems |
| Diaphus rafinesquii (sister group 7) | ||||||
| 21 | 37 | MAECO115-06 | 30 June 2004 | Mid-Atlantic Ridge | na | BOLD Systems |
| 22 | 8 | MAECO116-06 | 28 June 2004 | Mid-Atlantic Ridge | na | BOLD Systems |
| 23 | 9 | MAECO117-06 | 28 June 2004 | Mid-Atlantic Ridge | na | BOLD Systems |
| Diaphus richardsoni (sister group 8) | ||||||
| 24 | 6 | GBGCA11068-15 | na | na | na | BOLD Systems |
| 25 | 7 | LFSCS086-18 | 25 April 2014 | South China Sea | China | BOLD Systems |
| 26 | 1 | ANGBF22462-19 | 18 December 2010 | Southwestern Pacific | Australia | BOLD Systems |
| Diaphus kuroshio (sister group 9) | ||||||
| 27 | 10 | OL894503 | 12 April 2019 | Emperor Seamounts | na | NCBI |
| Diaphus theta (sister group 10) | ||||||
| 28 | 24 | DSFIB184-12 | 15 September 2011 | Strait of Georgia | Canada | BOLD Systems |
| 29 | 24 | FMV566-11 | 23 May 2006 | Sailish Sea | USA | BOLD Systems |
| 30 | 19 | TZFPB409-05 | 22 October 2005 | SW Vancouver Isl. | Canada | BOLD Systems |
| 31 | 16 | OL894509 | 28 March 2019 | NE Pacific | na | NCBI |
| 32 | 17 | OL894510 | 28 March 2019 | NE Pacific | na | NCBI |
| 33 | 18 | OL894511 | 28 March 2019 | NE Pacific | na | NCBI |
| 34 | 19 | OL894512 | 28 March 2019 | NE Pacific | na | NCBI |
| 35 | 19 | OL894513 | 28 March 2019 | NE Pacific | na | NCBI |
| 36 | 20 | OL894514 | 28 March 2019 | NE Pacific | na | NCBI |
| 37 | 21 | OL894515 | 31 March 2019 | NE Pacific | na | NCBI |
| 38 | 22 | OL894516 | 31 March 2019 | NE Pacific | na | NCBI |
| 39 | 23 | OL894517 | 31 March 2019 | NE Pacific | na | NCBI |
| 40 | 20 | OL894518 | 31 March 2019 | NE Pacific | na | NCBI |
| 41 | 24 | OL894519 | 31 March 2019 | NE Pacific | na | NCBI |
| 42 | 19 | OL894520 | 31 March 2019 | NE Pacific | na | NCBI |
| 43 | 25 | OL894521 | 31 March 2019 | NE Pacific | na | NCBI |
| 44 | 23 | OL894522 | 31 March 2019 | NE Pacific | na | NCBI |
| 45 | 26 | OL894523 | 31 March 2019 | NE Pacific | na | NCBI |
| Diaphus balanovi sp. nov. (main group) | ||||||
| 46 | 11 | OL894504 | 12 April 2019 | Emperor Seamounts | na | NCBI |
| 47 | 12 | OL894505 | 11 April 2019 | Emperor Seamounts | na | NCBI |
| 48 | 13 | OL894506 | 11 April 2019 | Emperor Seamounts | na | NCBI |
| 49 | 14 | OL894507 | 11 April 2019 | Emperor Seamounts | na | NCBI |
| 50 | 15 | OL894508 | 11 April 2019 | Emperor Seamounts | na | NCBI |
Table 2.
Morphometric characters of Diaphus balanovi sp. nov. (n = number of specimens; *—some measurements were taken from fewer than 35 specimens due to their damage in hauls).
Table 2.
Morphometric characters of Diaphus balanovi sp. nov. (n = number of specimens; *—some measurements were taken from fewer than 35 specimens due to their damage in hauls).
| Character | Holotype | Paratypes (n = 34) | Holotype + Paratypes | ||
|---|---|---|---|---|---|
| n * | Mean ± Error of Mean | Standard Deviation | |||
| SL, mm | 96 | 83–98 | - | - | - |
| In % of SL | |||||
| Head length | 30.2 | 27.6–31.8 | 33 | 29.6 ± 0.19 | 1.07 |
| Snout length | 2.6 | 2.6–3.6 | 32 | 3.2 ± 0.05 | 0.28 |
| Horizontal diameter of orbit | 10.4 | 9.6–11.9 | 33 | 10.7 ± 0.10 | 0.58 |
| Interorbital width | 10.4 | 8.5–12.1 | 29 | 10.3 ± 0.15 | 0.81 |
| Length of upper jaw | 19.8 | 17.7–20.5 | 31 | 19.3 ± 0.14 | 0.80 |
| Eye—maxilla distance | 7.3 | 5.5–8.5 | 31 | 6.6 ± 0.13 | 0.72 |
| Greatest body depth | 21.9 | 19.7–24.4 | 28 | 21.8 ± 0.23 | 1.24 |
| Least depth of caudal peduncle | 9.4 | 8.9–10.8 | 30 | 9.7 ± 0.11 | 0.60 |
| Caudal peduncle length | 20.8 | 17.1–21.8 (25.7) | 31 | 20.0 ± 0.29 | 1.60 |
| Distance between origins of pectoral and pelvic fins | 16.7 | 12.8–19.8 | 27 | 16.3 ± 0.34 | 1.76 |
| Distance between origins of pelvic and anal fins | 24.0 | 19.8–26.5 | 27 | 22.2 ± 0.30 | 1.58 |
| Predorsal distance | 43.8 | 42.5–46.9 | 27 | 44.7 ± 0.22 | 1.16 |
| Prepelvic distance | 44.8 | 41.5–48.2 | 29 | 45.7 ± 0.29 | 1.58 |
| Preadipose distance | 80.2 | 77.9–83.5 | 31 | 81.0 ± 0.28 | 1.59 |
| Dorsal-adipose distance | 20.8 | 16.9–20.8 (25.0) | 27 | 19.6 ± 0.32 | 1.64 |
| Length of dorsal-fin base | 14.9 | 13.8–19.6 | 28 | 16.3 ± 0.25 | 1.33 |
| Length of anal-fin base | 14.1 | 12.2–15.4 | 28 | 13.5 ± 0.16 | 0.82 |
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Prokofiev, A.M.; Emelyanova, O.R.; Orlov, A.M.; Orlova, S.Y. A New Species of Diaphus Associated with Seamounts of the Emperor Chain, North-Western Pacific Ocean (Teleostei: Myctophiformes: Myctophidae) . J. Mar. Sci. Eng. 2022, 10, 65. https://doi.org/10.3390/jmse10010065
AMA Style
Prokofiev AM, Emelyanova OR, Orlov AM, Orlova SY. A New Species of Diaphus Associated with Seamounts of the Emperor Chain, North-Western Pacific Ocean (Teleostei: Myctophiformes: Myctophidae) . Journal of Marine Science and Engineering. 2022; 10(1):65. https://doi.org/10.3390/jmse10010065
Chicago/Turabian StyleProkofiev, Artem M., Olga R. Emelyanova, Alexei M. Orlov, and Svetlana Y. Orlova. 2022. "A New Species of Diaphus Associated with Seamounts of the Emperor Chain, North-Western Pacific Ocean (Teleostei: Myctophiformes: Myctophidae) " Journal of Marine Science and Engineering 10, no. 1: 65. https://doi.org/10.3390/jmse10010065
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