Research Article |
Corresponding author: Man-Ki Jeong ( jmgdeux@gmail.com ) Corresponding author: Ho Young Soh ( hysoh@chonnam.ac.kr ) Academic editor: Pavel Stoev
© 2022 Seohwi Choo, Man-Ki Jeong, Ho Young Soh.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Choo S, Jeong M-K, Soh HY (2022) Taxonomic reassessment of chaetognaths (Chaetognatha, Sagittoidea, Aphragmophora) from Korean waters. ZooKeys 1106: 165-211. https://doi.org/10.3897/zookeys.1106.80184
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Since the first record of chaetognaths (arrow worms) reported from Korean waters by Molchanov in 1907, three families, 12 genera and 21 species have been additionally described. Eighteen of the 21 recorded species have been reported under scientific names different from the latest taxonomic system. This study aimed to address this issue by conducting a taxonomic re-evaluation of chaetognaths collected from Korean waters. Furthermore, the taxonomic usefulness of morphological differences in corona ciliata and distribution of ciliary sense receptors were re-examined using specimens stained with 1% Chlorazol black E (CBE) solution. This study includes taxonomically-validated voucher specimens of 18 species from Korean waters. Based on the specimens, re-description including image data and CBE staining pattern, distribution, ecological information and improved key were provided for each species. However, Decipisagitta decipiens, Serratosagitta serratodentata and Sagitta pseudoserratodentata from Korean waters is still questioned because of the paucity of the voucher specimen and scientific literature.
Arrow worms, chaetognaths, East China Sea, key, staining solution, taxonomy, voucher specimens
The chaetognaths are marine mesoplanktonic carnivores present in most marine habitats and play an important role in the food web of pelagic ecosystems comprising connecting planktonic organisms of higher trophic levels. They have two sets of retractable chitinous grasping spines flanking a ventral mouth. They mostly feed on copepods, cladocerans, amphipods, krill and fish larvae depending on their size and developmental stage (Vega-Pérez 1995;
The phylum Chaetognatha was first mentioned as “arrow-shaped worms” by Slabber M (1769) and
Despite the long taxonomic history of Chaetognatha and its ecological importance, taxonomic research on Korean species is extremely limited. The first record of chaetognaths in Korean waters was presented by
Therefore, in this study, we aimed to accomplish the following: 1) to secure the first taxonomically verified voucher specimens of chaetognath from Korean waters and disclose them to public institutions; 2) to create the first comprehensive report of taxonomic features, including morphology, ecology and image information on Korean chaetognath species, based on newly-obtained voucher specimens; and 3) to provide an updated key to species for chaetognath taxa in Korea.
To understand the current status of Korean record on chaetognaths, a total of 14 taxonomic and ecological papers published since 1940 to date were investigated (
Korean chaetognath species list reported in previous studies. The species list consists of
Taxa | ES | KS | nECS | YS | ES, KS, YS and nECS |
---|---|---|---|---|---|
Class Sagittidae | |||||
Order Aphragmophora | |||||
Family Krohnittidae | |||||
Genus Krohnitta Ritter-Zahony, 1910 | |||||
Krohnitta pacifica (Aida, 1897) | ● | ● | ● | ● | |
Krohnitta subtilis (Grassi, 1881) | ● | ● | ● | ||
Family Pterosagittidae (not valid) | |||||
Genus Pterosagitta Costa, 1869 | |||||
Pterosagitta draco (Krohn, 1853) | ● | ● | ● | ||
Family Sagittidae | |||||
Genus Aidanosagitta Tokioka,1965 | |||||
Aidanosagitta crassa (Tokioka, 1938) | ● | ● | ● | ● | ● |
Aidanosagitta neglecta (Aida, 1897) | ● | ● | ● | ||
Aidanosagitta regularis (Aida, 1897) | ● | ● | ● | ● | |
Genus Decipisagitta Tokioka, 1965 | |||||
Decipisagitta decipiens (Fowler, 1905) | ● | ● | ● | ||
Genus Ferosagitta Kassatkina, 1971 | |||||
Ferosagitta ferox (Doncaster, 1902) | ● | ● | ● | ||
Ferosagitta robusta (Doncaster, 1902) | ● | ● | ● | ||
Genus Flaccisagitta Tokioka, 1965 | |||||
Flaccisagitta enflata (Grassi, 1881) | ● | ● | ● | ● | ● |
Flaccisagitta hexaptera (D’Orbigny,1902) | ● | ● | ● | ||
Genus Mesosagitta Tokioka, 1965 | |||||
Mesosagitta minima (Grassi, 1881) | ● | ● | ● | ||
Genus Parasagitta Tokioka, 1965 | |||||
Parasagitta elegans (Verrill, 1873) | ● | ● | ● | ● | |
Genus Pseudosagitta Germain & Joubin, 1912 | |||||
Pseudosagitta lyra (Krohn, 1853) | ● | ● | ● | ||
Genus Sagitta Guoy & Gaimard, 1827 | |||||
Sagitta bipunctata Quoy & Gaimard, 1827 | ● | ● | ● | ||
Genus Serratosagitta Tokioka, 1965 | |||||
Serratosagitta pacifica (Tokioka, 1940) | ● | ● | ● | ||
Serratosagitta pseudoserratodentata (Tokioka, 1940) | ● | ● | ● | ||
Serratosagitta serratodentata | ● | ● | ● | ● | |
Genus Zonosagitta Tokioka, 1827 | |||||
Zonosagitta bedoti (Beraneck, 1895) | ● | ● | ● | ● | |
Zonosagitta pulchra (Doncaster, 1902) | ● | ● | |||
Zonosagitta nagae Alvariño, 1967 | ● | ● | ● | ● |
Field surveys were conducted at 20 stations in Korean waters from May 2019 to August 2020. Zooplankton collection was carried out from the bottom layer to the surface layer using a conical net (diameter: 0.6 m, mesh size: 200 μm) and MOCNESS (area of mouth: 1 m2, mesh size: 200 μm). Samples were fixed with 5% formalin and the morphological features were observed using a stereo-optical microscope (DS-Fi3, Nikon, Japan). Furthermore, chaetognath specimens were identified at the species level by referring to the taxonomic terms suggested by
Quantitative and qualitative characters of the chaetognaths used in this study (modified after Gasami et al. 2014).
Quantitative characters | Qualitative characters | ||
---|---|---|---|
C1 | Transverse muscles (absent/ present) | Q1 | Total length (min/max) |
C2 | Body firmness (flaccid/ rigid) | Q2 | Body/ tail ratio |
C3 | Body transparency (transparent, opaque, translucent) | Q3 | Number of anterior teeth |
C4 | Collarette (absent/ present) | Q4 | Number of posterior teeth |
C5 | Number of lateral fins (one pair/ two pairs) | Q5 | Number of hooks (min/max) |
C6 | Fin positions. Anterior fins can be present on anterior part of ventral ganglion, middle of ventral ganglion, end of ventral ganglion, long distance behind the end of the ventral ganglion. | ||
C7 | Comparison of anterior fins and posterior fins size | ||
C8 | Rayless zone in the lateral fins (absent/ present) | ||
C9 | Intestinal diverticula (absent/ present) | ||
C10 | Type of hooks (gently curved/ gently curved and serrated/ abruptly curved) | ||
C11 | Type of seminal vesicle (elongated, oval, spherical, conical) | ||
C12 | Positions of seminal vesicle | ||
C13 | Type of eye pigment (E, T, star, H, +, B shaped) | ||
C14 | Type of corona ciliata (following Tokioka (1965)) | ||
C15 | Type of teeth row | ||
C16 | Number of teeth row (only anterior row/ only posterior row/ presented both anterior and posterior) |
Schematic drawing showing the distribution of the ciliary sense organs (modified from
The size of the chaetognaths belonging to the order Aphragmophora ranged between 5 and 80 mm. Aphragmophora had no transverse muscles in the body. In general, species diagnosis in Aphragmophora was mostly based on the size and body appearance, the morphology of the intestine, the chitinous cephalic armature (shape and number of grasping spines and teeth) and the shape and position of the corona ciliata, lateral fins and seminal vesicles.
The transparency of the body is related to the development of longitudinal muscles in the trunk and tail and is a distinguishing characteristic at the generic level (
The fins are used for floating and balancing (
Three families inhabiting Korean waters A Ferosagitta robusta (Sagittidae) B Pterosagitta draco; C Krohnitta subtilis (Krohnittidae). Abbreviations: AF = anterior fin; CC = corona ciliata; COL = collarette; CF = caudal fin; E = eye; GS = grasping spine; LF = lateral fin; PF = posterior fin; SV = seminal vesicle; VG = ventral ganglion.
Another diagnostic character at the genus and species level is the presence of a rayless zone in the lateral fins. For example, Zonosagitta has a long rayless zone on the anterior fins, but Aidanosagitta does not have a rayless zone on either anterior or posterior fins (Fig.
All chaetognaths are hermaphroditic and have both female and male organs. In particular, the shape and location of seminal vesicles are distinct in different species. Seminal vesicles can be elongated along the lateral side of the tail (Fig.
Shape and position of seminal vesicles A Aidanosagitta regularis (elongated) B Ferosagitta robusta (pear shape) C Pseudosagitta lyra (spherical shape) D Zonosagitta nagae (conical shape) E Flaccisagitta enflata (spherical shape) F Serratosagitta pacifica (seminal shape with chitinous teeth).
The digestive apparatus of the chaetognath is in a single line from the mouth to the anus located just anterior to the posterior septum; the intestine extends in the trunk, but is not present in the tail. Classification can be done, based on the presence or absence of two intestinal diverticula located in the most anterior part of the intestine. They are clearly observed in the genera Aidanosagitta (Fig.
The grasping spines are laterally attached to the head of chaetognaths and are used for capturing and swallowing prey. The grasping spines of the family Krohnittidae are sharply curved, while those of Sagittidae are gently curved (Fig.
The number of teeth rows is an important key to distinguish families (Tokioka 1965 a, b). Sagittidae (including Pterosagitta draco) has two rows of teeth arranged in a comb shape (Fig.
The corona ciliata is related to the sensory organs, presumably involved in chemoreception (
Ventral transverse musculature absent, less glandular structures on body surface. Grasping spines gently or abruptly curved (
1 | One pair of lateral fins | 2 |
– | Two pairs of lateral fins | 4 |
2 | One row of teeth, collarette absent | Krohnittidae (present in Korea) |
– | Two rows of teeth | 3 |
3 | One pair of lateral fins on the tail, Collarette remarkably thick | Pterosagitta draco (present in Korea) |
– | One pair of lateral fins on the trunk | Pterokrohniidae |
4 | Two rows of teeth, neck contraction | Bathybelidae |
– | Two rows of teeth | Sagittidae (present in Korea) |
Small head. Grasping spines abruptly curved. One row of teeth. Collarette either short or absent. One pair of lateral fins arranged on the posterior trunk and tail.
Slender and transparent body. Lateral fins on the body and tail with rayless zone or partially rayed. Intestinal diverticula absent. Seminal vesicles oval or elongated touching both lateral fins and caudal fins.
1 | One pair of fins with rayless zone. Seminal vesicles oval shaped and elongated, touching both paired fins and caudal fins | K. subtilis |
– | One pair of rayed fins. Seminal vesicles oval shaped and placed dorsally at the point where lateral fins meet caudal fins | K. pacifica |
Spadella subtilis: Grassi, 1883: 16 p., table 1.
Krohnia subtilis: Fowler, 1906: 25–26 p., figs 86–88;
Krohnitta subtilis: Burfield & Harvey, 1926: 117 p., figs 45–50.;
Korea Strait (33°30.000'N, 125°54.000'E), 0–90 m depth, oblique towing with conical net, Feb 2020,
Total body length ranged between 10.8 and 11.5 mm and tail 27.3–33.7% of body length. Hooks 8–10. Anterior teeth 14. Slender and transparent body (Fig.
This species is found in the epipelagic (0–200 m depth) and mesopelagic zones (200–500 m depth) of the Pacific, Indian and Atlantic Oceans (
This cosmopolitan species can be found in tropical to temperate waters (
This species is clearly distinguished from K. pacifica by the presence of a rayless zone in the lateral fin. Furthermore, the presence of a pair of lateral fins with a wide rayless zone and a fan-shaped dentition in K. subtilis collected from Korean waters are consistent with the records of
Krohnitta pacifica: Tokioka, 1965: 352–353 p.;
Northern East China Sea (32°29.420'N, 127°29.654' E), 20–100 m depth, oblique towing with MOCNESS, Aug 2020 (one specimen).
Slender and transparent body. Small head. Grasping spines abruptly curved. One row of stout teeth arranged in fan shape. One pair of lateral fins partially rayed, forward and equidistant from caudal septum. Lateral fins positioned at anterior end, at level of caudal septum with rayed lateral fins. Caudal fin damaged. Collarette and intestinal diverticula absent. No seminal vesicle visible.
This species is found in the epipelagic zone (0–200 m depth) of the northern Indian Ocean (
An inhabitant of the surface layer of the warm oceanic waters (
Only immature individuals could be collected in this study. The specimens of K. pacifica we observed had one pair of fins and the structure of the fin was rayed, except for the base part close to the body. These were distinct characteristics of K. pacifica that differentiated it from K. subtilis. Since seminal vesicles were not observed in all observed Korean specimens, they were classified as immature stage (
Wide head. Two rows of teeth. Collarette wide and extending through full body. One pair of rayed lateral fins located on the tail. Intestinal diverticula absent. Only one genus has been described within this family: Pterosagitta (Costa, 1869).
A previous phylogenetic study reported that Pterosagittidae is genetically quite close to Sagittdae (
1 | One pair of rayed lateral fins on the tail. Collarette extremely thick and extends from head to seminal vesicles. Seminal vesicles oval-shaped and elongated, touching both lateral fins and caudal fin | P. draco |
Spadella draco: Grassi, 1883: 15 p.;
Pterosagitta draco: Michael, 1919: 264–265 p., table 18;
Northern East China Sea (32°30.000'N, 127°5.100'E), 0–120 m depth, oblique towing with conical net, Feb 2020,
Total body length ranged between 6.5 and 9.1 mm and tail 38.4–40.1% of body length. Hooks 8. Anterior teeth 10 and posterior teeth 12. Rigid and translucent body (Fig.
This species is located in the epipelagic (0–200 m depth) and mesopelagic zones (200–500 m depth) of the Pacific, Indian and Atlantic Oceans (
This species is widely distributed in warm water masses and appears all year round in Korean waters, except in the Yellow Sea (
The largest specimen collected in this study was 9.1 mm in length, which was at stage 4 maturity. It was smaller than the specimen from New Zealand (16 mm) reported by
Two rows of stout teeth arranged in comb shape. Two pairs of lateral fins on the trunk and tail.
1 | Serrated grasping spines | Serratosagitta (present in Korea) |
– | Non-serrated grasping spines | 2 |
2 | Flaccid body | 3 |
– | Rigid body | 6 |
3 | Intestinal diverticula present | 4 |
– | Intestinal diverticula absent | 5 |
4 | Anterior fins beginning at the middle of ventral ganglion, the seminal vesicle closer to lateral fins than caudal fin | Decipisagitta |
– | Anterior fins beginning between ventral ganglion and caudal septum, the seminal vesicle well-separated from lateral fins, but touching caudal fins | Mesosagitta (present in Korea) |
5 | Anterior fins beginning far behind the end of the ventral ganglion | Flaccisagitta (present in Korea) |
– | Anterior fins beginning at the end of ventral ganglion | Pseudosagitta (present in Korea) |
6 | Intestinal diverticula present | 7 |
– | Intestinal diverticula absent | 9 |
7 | Collarette present | 8 |
– | Collarette and rayless zone on lateral fins absent | Parasagitta (present in Korea) |
8 | The head width greater than body width, short collarette, the rayless zone on lateral fins present, the corona ciliata beginning before the eye | Ferosagitta (present in Korea) |
– | The head width smaller than body width, well-developed collarette, completely rayed lateral fins, the corona ciliata beginning behind the eye | Aidanosagitta (present in Korea) |
9 | Anterior part of lateral fins with rayless zone | Zonosagitta (present in Korea) |
– | Rayless zone on lateral fins absent | Sagitta (present in Korea) |
Intestinal diverticula either absent or present. Grasping spines serrated. Two rows of teeth. Two pairs of lateral fins either with rayless zone or completely rayed.
1 | Intestinal diverticula absent | 2 |
– | Intestinal diverticula present | 3 |
2 | Seminal vesicles with forward elongated knob and teeth-like appendages forming 5–10 distal protrusions anteriorly, well-separated from caudal fin but touching posterior fins, the posterior fins and anterior fins of almost same length | S. pacifica |
– | Seminal vesicles touching the posterior fins and caudal fin, the posterior fins longer than anterior fins | S. serratodentata |
3 | Seminal vesicles well-separated from caudal fin, but touching posterior fins, posterior fins and anterior fins of almost same length | S. pseudoserratodentata |
Sagitta serratodentata pacifica: Tokioka, 1959: 72–80 p., fig. 10, table 10;
Sagitta pacifica: Alvariño, 1961: 71 p., fig. A, B, table 2;
Serratosagitta pacifica: Tokioka, 1965: 345–346 p.;
Korea Strait (33°24.504'N, 127°54.600'E), 0–50 m depth, oblique towing with MOCNESS, May 2019,
Total body length ranged within 11.8 and 13.7 mm. Tail 23.4–24.9% of body length. Hooks 6–7. Anterior 10–13 and posterior teeth 16–25. Rigid and opaque body (Fig.
This species is found in the epipelagic (0–200 m depth) and mesopelagic zones (200–500 m depth) of the Pacific and Indian Oceans (
This species mainly inhabits Indo-pacific warm-water masses (
The seminal vesicles are used as an important morphological feature to identify the genus Serratosagitta. S. serratodentata has thick collarette tissue in front of the seminal vesicles and two projections at the anterior-lateral corner. The seminal vesicles touch the end of posterior fins (
Flaccid and opaque body. Collarette absent. Intestinal diverticula present. Grasping spine gently curved and not serrated. Intestinal diverticula present. The anterior fins begin between ventral ganglion and caudal fin and are shorter than posterior fins.
Spadella minima: Grassi, 1883: 15 p.
Sagitta minima: Aida, 1897: 15 p., fig. 5;
Northern East China Sea (32°29.420'N, 127°29.654'E), 20–100 m depth, oblique towing with MOCNESS, Aug 2020 (one specimen).
Small head. Two pairs of lateral fins with rayless zone. Intestinal diverticula small. Seminal vesicles divided into a small anterior knob and elongated posterior part and well-separated from posterior fins, but touching caudal fin. Corona ciliata elongated oval-shaped beginning in neck.
This cosmopolitan species is found in the epipelagic zone (0–200 m depth) of the Pacific, Indian and Atlantic Oceans (
Mesosagitta minima is abundant in mixed waters of the western North Atlantic Ocean (
Only immature individuals could be collected in this study. We easily distinguished M. minima amongst the collected specimens by the relatively small head and unique body shape that thickens towards the tail. These Korean specimens were classified as immature because seminal vesicles and ovaries were absent and undeveloped short eggs (mentioned in description of
Transparent or translucent body. Collarette absent. Intestinal diverticula absent. The anterior fins begin at a far distance behind the end of the ventral ganglion. Seminal vesicles spherical shaped.
1 | Large body (> 40 mm), small eggs reaching the neck | F. hexaptera |
– | Small body (< 20 mm), large eggs reaching the anterior part of posterior fins | F. enflata |
Spadella enflata: Grassi, 1883: 13 p., fig. 7.
Sagitta inflata: Ritter-Záhony, 1908: 13–15 p., fig. 4A–D;
Sagitta enflata: Aida, 1897: 15–16 p., fig. 6;
Sagitta enflata f. gardineri: Tokioka, 1959: 91–92 p., table 19
Flaccisagitta enflata: McLelland, 1989: 159 p., figs 7A and 12B
Korea Strait (32°59.175'N, 124°29.595'E), 20–25 m depth, oblique towing with MOCNESS, Nov 2019,
Total body length ranged between 12.7 and 15.4 mm. Tail 14.1–17.6% of body length. Hooks 8–10. Anterior teeth 6–10 and posterior teeth 10–17, respectively. Transparent body, inflated towards middle (Fig.
This cosmopolitan species is found in the epipelagic (0–200 m depth) and mesopelagic zones (200–500 m depth) of the Pacific, Indian and Atlantic Oceans (
This is used as an indicator species of warm currents in water surrounding Korea (
The transparent and flaccid body, star-shaped eye pigments and seminal vesicle morphology were consistent with those recorded in previous studies by
Sagitta hexaptera: Conant, 1896: 213 p.;
Flaccisagitta hexaptera: McLelland, 1989: 159–160 p., figs 7B, C and 12A.
Korea Strait (33°24.504'N, 127°54.600'E), 0–50 m depth, oblique towing with MOCNESS, Nov 2019,
Total body length ranged between 15 and 49 mm. Tail 19–24% of body length. Hooks 4–11. Anterior teeth 2–4 and posterior teeth 2–9, respectively. Large and translucent body (Fig.
This cosmopolitan species is found in the epipelagic (0–200 m depth) and mesopelagic zones (200–500 m depth) of the Pacific, Indian and Atlantic Oceans (
This species is considered as an indicator species of the Kuroshio warm current and fully-grown adults inhabit depths of < 200 m (
Though mainly immature individuals were reported in previous studies from Korea, in this study, adults longer than 40 mm were collected for the first time. The Korean species was consistent with those found in previous studies by
Flaccid and transparent body (but more opaque than Flaccisagitta). Collarette absent. Intestinal diverticula absent. Two pairs of lateral fins partially rayed and connected with a tegumentary bridge. Anterior fins beginning at the rear end of ventral ganglion and longer than posterior fins.
Sagitta lyra: Aida, 1897: 15 p., fig. 4;
Flaccisagitta lyra: McLelland, 1989: 162 p., figs 7D, 7E and 12C.
Korea Strait (33°48.924'N, 126°48.666'E), 40–70 m depth, oblique towing with MOCNESS, May 2019,
Total body length ranged between 16.0 and 60.0 mm. Tail 14–20% of body length. Hooks 8–9. Anterior teeth 4–9 and posterior 8–10, respectively. Large, flaccid and opaque body (Fig.
This species is found in the mesopelagic (500–1,000 m depth) and bathypelagic zones (1,000–2,000 m depth) of the Pacific, Indian and Atlantic Oceans (
This species has a high prevalence in warm waters (
In Korean specimens, the position and length of the fins, distance between the anterior and posterior fins and shape of the seminal vesicles were morphologically consistent with the previous records of Pseudosagitta lyra (
Slender and either opaque or translucent body. Collarette absent or small (almost absent). Intestinal diverticula present or absent. Grasping spines not serrated. Two rows of teeth. Two pairs of lateral fins completely rayed.
Sagitta elegans: Verrill, 1873: 332–333 p.;
East Sea (37°33.198'N, 131°14.598'E), 0–100 m depth, oblique towing with conical net, Feb 2020,
Total body length ranged between 32.5 and 37.0 mm. Tail 14.7–20.6% of body. Hooks 11–12. Anterior teeth 9–10 and posterior teeth 22–29, respectively. Rigid and opaque body (Fig.
This species is found in the Epipelagic (0–200 m depth), mesopelagic (200–500 m depth) and bathypelagic zones (1000–2000 m depth) of the Pacific, Indian and Atlantic Oceans (
A cold-water species, P. elegans is mainly found in the northern part of the Pacific Ocean (
The absence of a rayless zone in the anterior and posterior fins and a collarette in anterior body and the presence of small seminal vesicles extending along the body in the Korean specimens of Parasagitta elegans were consistent with previous records (
Rigid and opaque body. Collarette present. Intestinal diverticula present. Grasping spines not serrated. Two pairs of lateral fins completely or partially rayed.
1 | Seminal vesicles oval shaped, posterior fins with small rayless zone | F. ferox |
– | Seminal vesicles pear shaped, anterior and posterior fins fully rayed | F. robusta |
Sagitta ferox: Doncaster, 1902: 212 p.;
Korea Strait (33°24.504'N, 127°54.600'E), 0–50 m depth, oblique towing with MOCNESS, May 2019 (one specimen).
Broad head. Rigid and opaque body. Collarette present. Intestinal diverticula present. Grasping spines gently curved. Two rows of stout teeth arranged in comb shape. Two pairs of lateral fins partially rayed, anterior fins beginning at middle of ventral ganglion. Seminal vesicles oval-shaped with an anterior protruding part touching both lateral and caudal fins.
This species is found in the epipelagic zone (0–200 m depth) of the Pacific and Indian Oceans (
This species inhabits the surface layer of the warm water and is mainly dominant in the Kuroshio Current of the Japanese waters (
Only immature individual was collected in this study. We easily distinguished F. ferox amongst the collected specimens by the presence of distinct head as wide as the trunk and the presence of elongated ovoid seminal vesicles on the body. The Korean specimen was classified as immature because the boundary of the seminal vesicles was not obvious and the inside was mostly empty. This characteristic is consistent with
Sagitta robusta: Fowler, 1906: 19–20 p., figs 59–64;
Yellow Sea (34°5.502'N, 124°36.000'E), 0–75 m depth, oblique towing with conical net, Aug 2019,
Total body length ranged between 10.0 and 11.5 mm. Tail 27.8–35.0% of body length. Hooks 6–8. Anterior teeth 9–10 and posterior teeth 12–20, respectively. Rigid and opaque body (Fig.
This species is found in the epipelagic zone (0–200 m depth) of Pacific and Indian Oceans (
Characteristics of the species in Korean waters, such as the conspicuous intestinal diverticula, head and body width and the seminal vesicles shape, are consistent with previous records (
Rigid and opaque body. Intestinal diverticula present. Collarette present or absent. Grasping spine gently curved and not serrated. Two rows of stout teeth arranged in a comb shape. Two pairs of lateral fins completely rayed.
1 | Seminal vesicles large and oval shaped, well-separated from caudal fins but touching posterior fins | A. neglecta |
– | Seminal vesicles small and elongated | 2 |
2 | Seminal vesicles touching both posterior fins and caudal fin and opening at the anterolateral edge. Collarette covered on ventral ganglion (N type) from head to body (C type) or on the partial body (I type) | A. crassa |
– | Seminal vesicles well-separated from caudal fins, but touching the posterior fins. Thick collarette covered head to tail. Small sized body (< 10 mm) | A. regularis |
Sagitta neglecta: Aida, 1897: 16–17 p., fig. 7;
Korea Strait (34°41.577'N, 127°50.460'E), 0–20 m depth, oblique towing with conical net, Feb 2021 (one specimen).
Rigid and opaque body. Narrow collarette and extending to half distance from neck to ventral ganglion. Intestinal diverticula present. Grasping spine gently curved. Two rows of stout teeth arranged in comb shape. Two pairs of lateral fins completely rayed, anterior fins beginning at end of ventral ganglion. Seminal vesicles oval-shaped with opening at anterolateral edge, position of seminal vesicles well separated from caudal fin, but touching posterior fins.
This species is found in the epipelagic zone (0–200 m depth) of the Pacific and Indian Oceans (
A mature specimen has been reported to be 8 mm is size (
Only one immature individual was collected in this study. Aidanosagitta neglecta is similar to A. regularis in morphological characteristics including collarette. However, the former had much larger seminal vesicles, thus they can be easily distinguished from each other. Aidanosagitta neglecta collected from Korean waters was smaller than the previously reported adult specimens, but seminal vesicles were obviously swollen in an oval shape. Despite the presence of swollen seminal vesicles, the Korean specimen was considered as immature because its size was smaller than the known record of the adult (
Aidanosagitta crassa f. naikaiensis: Tokioka, 1959: 376–377 p., fig. 16, table 15.
Sagitta crassa: Tokioka, 1939: 349–352 p., figs 1–8
Type C (collarette type): East Sea (37°33.198'N, 131°14.598'E), 0–100 m depth, oblique towing with conical net, Feb 2020,
Type C: total body length ranged within 9.9 and 11.2 mm. Tail 26.6–29.8% of body length. Rigid and opaque body (Fig.
A Aidanosagitta crassa Type C (dorsal view) B Aidanosagitta crassa Type N (dorsal view) C head and neck D tail E anterior fin F posterior fin. Abbreviations: AF = anterior fin; CC = corona ciliata; COL = collarette; E = eye; IN = intestine; ID = intestine diverticula; PF = posterior fin; SV = seminal vesicle.
Type N: total body length ranged between 8.1 and8.2 mm. Tail 27.0–30.0% of body length (Fig.
This species is found in the neritic water of the Pacific Ocean (
Aidanosagitta crassa appears in high abundance throughout the year in the relatively low saline waters of the Yellow Sea and coast of Jeju (
Previous researchers classified Aidanosagitta crassa into three types according to the distribution of the collarette: C type, covers from the neck to the body; N type, covers only the ventral ganglion; and I type, covers the ventral ganglion and partially covers the body (
Sagitta regularis: Aida, 1897: 17–18 p., fig. 8;
Korea Strait (33°33.600'N, 127°34.002'E), 0–96 m depth, oblique towing with conical net, Feb 2020,
Total body length ranged within 6.1 and 6.7 mm. Tail 30.5–31.7% of body length. Hooks 6. Anterior teeth 3–4 and posterior teeth 5–6, respectively. Rigid and opaque body (Fig.
This species is found in the epipelagic (0–200 m depth) and mesopelagic zones (200–500 m depth) of Pacific and Indian Oceans (
This species is considered a warm-water Indo-Pacific indicator species because many individuals appear in high-temperature and high-salinity water and the distribution range is limited to the areas affected by warm currents (
Amongst the Korean specimens, the collarette was differently inflated, thick and covered the body surface. However, the position and shape of the corona ciliata and fins were consistent with the original description (
Rigid and opaque body. Collarette almost absent. Intestinal diverticula absent or present. Grasping spines not serrated. Two pairs of lateral fins completely rayed.
Sagitta bipunctata: Quoy & Gaimard, 1827: 232–233 p., figs 1, 2, 6, 7;
Yellow sea (33°0.111'N, 125°29.581'E), 0 – 86 m depth, oblique towing, July 2020.
Small, rigid and opaque body. Small head. Collarette almost absent (thin on neck). Eyes square-shaped with no eye pigments observed in this study. Intestinal diverticula absent. Forward end of anterior fins not visible. Posterior fins beginning in front of caudal septum and closing to seminal vesicles, completely rayed. Seminal vesicles elongated with small indentations and touching both lateral and caudal fins.
This species is found in the epipelagic (0–200 m depth) and mesopelagic zones (200–500 m depth) of Pacific, Indian and Atlantic Oceans (
Sagitta bipunctata is known as a cosmopolitan species, appearing in temperate and tropical seas; it is an indicator species of high salinity and the presence of oceanic water (Pierce 1953). The temperature range in sampling stations of this study was 15.08–22.17 °C and the salinity range was 31.77–34.01 psu.
Only one immature individual was collected in this study. Sagitta bipunctata can be distinguished from other Korean species by the following characteristics: absence of intestinal diverticular, presence of completely rayed lateral fins and the restricted position of collarette on the posterior part of the body. The seminal vesicles of Korean specimen were not sufficiently mature compared to the description of
Rigid or flaccid and transparent or opaque body. Collarette small (almost absent). Intestinal diverticula absent. Grasping spines not serrated. Two pairs of lateral fins partially rayed. Anterior part of anterior fins elongated and rayless. Anterior fins longer than posterior fins.
1 | The anterior fins beginning at the middle of ventral ganglion, the seminal vesicles oval shaped touching neither the posterior fins nor caudal fin, small eyes with star-shaped eye pigments | Z. bedoti |
– | The anterior fins beginning at the end of ventral ganglion, the seminal vesicles conical-shaped and touching both lateral fins and caudal fin, small eyes with “E” shaped eye pigments | Z. nagae |
– | The anterior fins beginning at the middle of ventral ganglion, the seminal vesicles elongated with a roundish anterior protruding part, small eyes with star shaped eye pigments | Z. pulchra |
Sagitta bedoti f. minor: Tokioka, 1959: 89–90 p., table 18.
Sagitta bedoti: Béraneck, 1895: 147–152 p.;
Korea Strait (33°0.000'N, 125°18.000'E), 0–75 m depth, oblique towing with conical net, Feb 2020,
Total body length ranged from 16–17 mm. Tail 3.71% of body length. Hooks 7. Anterior teeth 27–28 and posterior teeth 30–35, respectively. Opaque and rigid body (Fig.
This species is found in the epipelagic zone (0–200 m depth) of the Pacific and Indian Oceans (
Zonosagitta bedoti is used as an indicator species in the front area where warm and cold water meet (
Zonosagitta bedoti and Z. nagae are similar in morphology at the immature stage. However, because adult Z. nagae are relatively larger, immature Z. nagae may be misidentified as Z. bedoti. Adults of both species can be distinguished from each other by the shape of seminal vesicles and eye pigments. Zonosagitta nagae has an “E”-shaped eye pigment, while Z. bedoti has a star-shaped eye pigment. The spot pattern on the body surface found through CBE staining is as follows: irregular spot pattern which continued from the head to the ventral ganglion and six spots along the outside of the body were symmetrical around the tail.
Sagitta nagae: Alvariño, 1967: 55–58 p., fig. 34A–D.
Korea Strait (33°29.662'N, 125°30.881'E), oblique towing with MOCNESS, 32–58 m depth, July 2020,
Total body length ranged from 11–15 mm. Hooks 6–8. Anterior teeth 11–15 and posterior teeth 15–35, respectively. Rigid and opaque body (Fig.
This species is found in the epipelagic zone (0–200 m depth) of the Pacific and Indian Oceans (
This species appears year-round in most of the seas around Korea, predominantly in the southern and western seas and shows high abundance in areas where warm and cold currents meet (
The anterior fin of Zonosagitta nagae had a longer rayless zone than the posterior fin, clearly distinguishing it from Z. bedoti. CBE staining showed the following spot pattern on the body surface: an irregular spot pattern continued from the head to the middle of the ventral ganglion; symmetrical dots appeared near the rayless zone of each fin (13 spots on anterior fin, 6 spots on posterior fin).
Sagitta pulchra: Doncaster, 1902: 213 p.;
Korea Strait (34°17.868'N, 128°15.854'E), 0–60 m depth, oblique towing with MOCNESS, Feb 2020 (one specimen).
Rigid and transparent body. Collarette present. Intestinal diverticula absent. Grasping spine gently curved. Two rows of stout teeth arranged in comb shape. Two pairs of lateral fins partially rayed, anterior fins begin at the end of ventral ganglion. Seminal vesicles elongated with an anterior protruding part usually roughly round and touching both lateral fins and caudal fin.
The species is found in the epipelagic zone (0–200 m depth) of the Pacific and Indian Oceans (
Z. pulchra is considered to be a neritic species (
Only one immature individual was collected in this study. The examined Korean specimen belongs to the genus Zonosagitta by the absence of intestinal diverticular and the presence of rayless zone in the anterior part of anterior fins. This Korean specimen was identified as Z. pulchra because its anterior fins were more angled than those of Z. bedoti or Z. nagae. The seminal vesicles of Korean specimen were not sufficiently mature compared to description of
Based on a taxonomic review of newly obtained specimens from Korea, we confirmed the appearance of chaetognath taxa in Korean waters corresponding to one order, three families, 11 genera and 18 species. Taxonomically identified voucher specimens (Krohnitta subtilis, K. pacifica, Pterosagitta draco, Aidanosagitta crassa, A. neglecta, A. regularis, Ferosagitta ferox, F. robusta, Flaccisagitta enflata, F. hexaptera, Mesosagitta minima, Parasagitta elegans, Pseudosagitta lyra, Sagitta bipunctata, Serratosagitta pacifica, Zonosagitta bedoti, Z. nagae and Z. pulchra) were obtained for the first time in Korea and their taxonomic and ecological features were reported. Although the overall morphological characteristics of the six species (A. neglecta, F. ferox, K. pacifica, M. minima, S. bipunctata and Z. pulchra) were mostly consistent with the previously-reported species, their essential characteristics have been briefly described in this study because only immature individuals of these six species were collected. On the contrary, the three species (Decipisagitta decipiens, Serratosagitta serratodentata, S. pseudoserratodentata) mentioned in literature have very poor taxonomic basis for their academic report (drawings, descriptions and voucher specimens), which can result in possible misidentification and their presence in Korean waters is questionable. Most of the samples in this study were collected in summer and winter. Extension of the investigation period to spring and autumn in future studies can facilitate identification of adult specimens of the above-mentioned six species or clarification of the presence or absence of the three suspicious species.
The detailed characteristics of the corona ciliata and fins of chaetognaths are important as taxonomic keys to distinguish genera, but they are difficult to observe under a stereomicroscope. In order to address this problem, we stained the surface of the specimens with CBE, which has rarely been used in the past for observing chaetognaths. The CBE staining of the Korean specimens clearly revealed the features of the corona ciliata, fins and body surface. The corona ciliata of Aidanosagitta is located from behind the eyes to the anterior part of the trunk, whereas that of Sagitta, Serratosagitta, Ferosagitta, Parasagitta and Zonosagitta extends from the front of the eyes to the neck; hence, the two groups could be clearly distinguished. Flaccisagitta is also distinctly differentiable from other genera as the corona ciliata extends from the front of the eyes to the anterior part of the neck. We propose the location and shape of these corona ciliata as additional taxonomic keys to distinguish Korean taxa at the genus level. On the contrary, Krohnitta subtilis had a circular corona ciliata located in front of the eyes, unlike other congeneric species, in which a circular corona ciliata appeared behind the eyes.
The distribution of dots on the body surface was easily confirmed through microscopic observation after CBE staining. The dots pattern is expressed by a regular arrangement of tactile cilia distributed on the body surface of the chaetognaths (
On the contrary, Flaccisagitta did not have similar spot patterns on the body surface between the two Korean species. Irregular spots on the body surface of the Flaccisagitta enflata were observed; however, F. hexaptera did not exhibit any spot patterns, similar to Pseudosagitta lyra. Since the fully-grown body of this species is usually large with a size of 50 mm and more, it is presumed that the relatively thick epidermis prevented effective staining. The regularity of these patterns has also been observed in species such as Sagitta hispida, S. enflata, S. elegans, Spadella schizoptera and Spadella cephaloptera (Feigenbaum, 1977); however, evidence for establishing a connection with genealogy is lacking. Moreover, basic data obtained from a complete individual pattern will be valuable to explain the commonalities at genus levels. Further application of the CBE staining method to other taxa of Aphragmophora will clarify whether new features, such as the location and shape of the corona ciliata and the spot patterns on the body surface, are effective as genus or species grouping features.
We would like to thank Dr. Kasatkina, Dr. Perez and Dr. P Stoev who made constructive and invaluable suggestions and comments. This work was supported by a grant from the National Institute of Biological Resources (