Research Article |
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Corresponding author: Jimin Lee ( leejm@kiost.ac.kr ) Academic editor: Kai Horst George
© 2025 Kyuhee Cho, Jong Guk Kim, Jimin Lee.
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:
Cho K, Kim JG, Lee J (2025) Discovery of Laophontidae (Copepoda, Harpacticoida) from marine plastic debris: Pseudonychocamptus setadefectus sp. nov. and Heterolaophonte discophora (Willey, 1929). ZooKeys 1251: 323-352. https://doi.org/10.3897/zookeys.1251.160858
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Two species belonging to the family Laophontidae Scott T., 1905 were identified among harpacticoid copepods collected from marine plastic debris (MPD) stranded along the Korean coastline. These species were assigned to the genera Pseudonychocamptus Lang, 1944 and Heterolaophonte Lang, 1948. Pseudonychocamptus setadefectus sp. nov. conforms to the generic diagnosis, displaying sexual dimorphism in the armature of the swimming legs. It is distinguished from its six congeners by possessing only two endites on the maxillary syncoxa, notably lacking the proximal endite bearing the seta that is present in all other known species within the genus. Although most morphologically similar to P. colomboi Ceccherelli, 1988, the new species differs in several key characteristics, including the number of setae on the maxillary endopod, the relative length ratios of setae on the female P5 exopod, the presence of an inner seta on the male P5 exopod, and the asymmetry of the male P6. The second species, Heterolaophonte discophora (Willey, 1929), was previously reported from the Atlantic coast of Canada and the Pacific coasts of the USA and Japan. The Korean specimens closely correspond to these previous records, yet exhibit intraspecific variation, particularly in the setal count on the female P3 enp-2. Additional morphological differences were also observed, including variations in the segmentation of the male P3 endopod and in the number of setae on the maxillary endopod. Based on the sexually dimorphic traits observed in males, we propose the subdivision of the genus Heterolaophonte into four distinct species groups. The discovery of P. setadefectus sp. nov. and H. discophora on anthropogenic MPD underscores the potential role of such substrates as microhabitats for benthic harpacticoid copepods.
Biodiversity, Crustacea, Heterolaophonte, meiofauna, Pseudonychocamptus, taxonomy
The family Laophontidae Scott T., 1905 represents one of the most diverse groups of benthic harpacticoid copepods, encompassing two subfamilies, Esolinae Huys & Lee, 2000 and Laophontinae Scott T., 1905, and 77 genera with 353 species (
Although recent taxonomic revisions [e.g.,
Within the Korean fauna, up to 38 species belonging to 17 genera of the family Laophontidae have been reported (
Conversely, earlier records of known 23 species in Korean fauna (e.g.,
Recent studies have identified marine plastic debris (MPD) as both a rafting vector for non-indigenous species and a novel ecological niche for a variety of marine invertebrates (
Samples were collected from various types of floating MPD at Gohado, Naechi, and Jeju Island, Republic of Korea. Additional specimens of Heterolaophonte were obtained by washing macroalgae collected from Masan-ri, Pohang. Copepods were extracted by rinsing the surfaces of debris with tap water using a compressed air sprayer. The dislodged organisms were filtered through a 60-μm mesh hand net and fixed in 90% ethanol at the sampling site. In the laboratory, laophontid harpacticoids were sorted from the preserved material under a stereomicroscope (M165 C; Leica Microsystems, Wetzlar, Germany). Specimens were dissected and temporarily mounted in lactic acid on reverse slides for microscopic examination. Morphometric measurements and illustrations were performed using a differential interference contrast light microscope (BX51; Olympus, Tokyo, Japan) equipped with a camera lucida. All scale bars in the figures are indicated in micrometers.
The dissected body parts were subsequently transferred to and permanently mounted in a lactophenol:glycerin solution (1:5) on H-S slides (Double slide plate, BSDS-011R; Biosolution, Republic of Korea) (cf.
The morphological terminology used in the text and figures follows
Order Harpacticoida Sars, 1903
Family Laophontidae Scott T., 1905
Subfamily Laophontinae Scott T., 1905
Genus Pseudonychocamptus Lang, 1944
Gohado (34°45'56.04"N, 126°22'22.38"E), Mokpo, Jeollanam-do, Republic of Korea.
Holotype. • 1♀ (MABIK CR00258872), preserved in a vial with 95% ethanol, collected from washings of marine plastic debris (a gunny sack) at the type locality on 14 March 2022 by GH Han. Paratypes. • 2♀♀ (MABIK CR00258874–00258875), each dissected and mounted on 2 H-S slides; • 2♀♀ (MinRB-Hr104-L001), preserved in a vial with 95% ethanol; • 1♂ (MABIK CR00258876), dissected and mounted on 2 H-S slides; and • 1♂ (MinRB-Hr104-L002) and 1♂ (MABIK CR00258873), each preserved in a vial with 95% ethanol. All material was collected from the type locality.
• 1♀ (MInRB-Hr104-S004), dissected and mounted on 2 H-S slides, collected from washings of marine plastic debris (expanded polystyrene buoy) at Chagwido Port, Jeju (33°19'25.86"N, 126°09'56.88"E) on 20 September 2022 by GH Han.
Female. Body length 441–535 μm (n = 6; measured from anterior margin of rostrum to posterior margin of caudal rami). Habitus broad, moderately flattened and gradually tapering posteriorly in dorsal view, separation between prosome and urosome indistinct (Fig.
Caudal rami (Figs
Rostrum (Fig.
Antennule (Fig.
Antenna (Fig.
Mandible (Fig.
Maxillule (Fig.
Maxilla (Fig.
Maxilliped (Fig.
P1–P4 (Figs
P1 (Fig.
P2–P4 (Fig.
Armature formula of P1–P4 as follows:
| Exopod | Endopod | |
|---|---|---|
| P1 | 0.0.022 | 0.020 |
| P2 | 0.1.123 | 0.1-220 [0.120 in ♂] |
| P3 | 0.1.123 | 1.221 [0.120 in ♂] |
| P4 | 0.1.123 | 1.121 [020 in ♂] |
P5 (Fig.
Male (based on the paratypes). Body (Fig.
Urosome (Fig.
Antennule (Fig.
Swimming legs. P1 and P2 similar to those of female, except for P2 enp-1 and enp-2 subequal in length (enp-2 relatively shorter than that of female), with one inner and two apical setae (Fig.
P3 and P4 (Fig.
P5 (Fig.
P6 (Fig.
Morphological variation was observed in the armature formula of the thoracopods in P. setadefectus sp. nov. In females, the P2 enp-2 typically bears two inner setae; however, in one of the five specimens examined, three inner setae were present.
The species name setadefectus is derived from the Latin words seta (bristle) and defectus (lacking, missing), referring to the absence of the proximal endite bearing a seta of maxilla—a distinguishing feature of this species among its congeners. It is an adjective in the singular nominative, gender masculine.
Species of the genus Pseudonychocamptus exhibit marked sexual dimorphism, particularly in the following: the inner seta on P3–P4 enp-1, present in females, is absent in males; and the two-segmented P4 endopod in females is reduced to a single segment bearing two setae in males. Notably, the latter trait represents the most prominent apomorphy of the genus (
The most distinctive feature of P. setadefectus sp. nov., which sets it apart from its six congeners, is the presence of only two endites on the maxilla, lacking the proximal endite represented by a single seta that is present in all other species of the genus. This condition represents a unique and highly unusual trait within Pseudonychocamptus.
The new species, P. setadefectus sp. nov., shares the presence of two inner setae on the female P3 enp-2 with four of the six valid species in the genus. These species are P. abbreviatus (Sars G.O., 1920), P. colomboi Ceccherelli, 1988, P. marinovi Apostolov & Petkovski, 1980, and P. proximus (Sars G.O., 1908). Conversely, P. koreni (Boeck, 1873) and P. spinifer Lang, 1965 possess only one inner seta on the female P3 enp-2. Among these, P. setadefectus sp. nov. is most similar to P. colomboi in that the male P2–P3 enp-2 bears only a single seta. It should be noted, however, that the male of P. marinovi has not yet been described. Previous studies (e.g.,
Although the morphological differences between females of P. setadefectus sp. nov. and P. colomboi are relatively subtle, the new Korean species can be readily distinguished by several characters. The fourth outermost seta on the P5 exopod is ~1.5 × longer than the third outermost seta in P. setadefectus sp. nov. (Fig.
In addition to the aforementioned differences, several minor features further distinguish P. setadefectus sp. nov. from P. colomboi: the maxillary endopod bears four setae in P. setadefectus sp. nov., but three in P. colomboi; the maxillipedal claw is accompanied by two accessory setae in the new species, versus one in P. colomboi; the inner setae on the female P2 enp-2 differ in relative length—the distal seta is ~2 × as long as the proximal in P. setadefectus sp. nov., whereas their length ratio is ~3:1 in P. colomboi; all setae, except for the longest apical seta, on the female P5 exopod are ornamented in P. setadefectus sp. nov., but all are bare in P. colomboi; the two setae of the female P6 differ in length by more than twofold in P. setadefectus sp. nov., but they are subequal in P. colomboi; the male P2 enp-1 and enp-2 are subequal in length in P. setadefectus sp. nov., whereas the enp-2 is slightly shorter in P. colomboi; and P. setadefectus sp. nov. possesses a single pair of sensilla near the apex of the rostrum, compared to two pairs in P. colomboi.
Laophonte discophora
Willey, 1929: 531, Abb. 2, 3, 6; Willey 1930: 607, pl. XVIII, figs 16~18 (cited from
Heterolaophonte discophora
Lang, 1948: 1375, fig. 557-1;
Heterolaophonte rotundipes Chappuis, 1958: 420, figs 23–34.
(1) • Gohado (34°45'56.04"N, 126°22'22.38"E), Mokpo, Jeollanam-do, marine rope debris; 01 April 2021; leg. OH Yu, SL Kim and SM Kang: 1♀ (MinRB-Hr105-L001), 3♀♀ (MABIK CR00258880), 1♂ (MinRB-Hr105-L002) and 3♂♂ (MABIK CR00258881), preserved separately in a vial with 95% ethanol; 1♀ (MinRB-Hr105-S003), 2♀♀ (MABIK CR00258882–00258883), 1♂ (MInRB-Hr105-S006) and 2♂♂ (MABIK CR00258884–00258885), each dissected and mounted on 1 or 2 H-S slides.
(2) • Naechi Beach (34°53'17.94"N, 126°00'05.68"E), Sinan, Jeollanam-do, expanded polystyrene buoy; 14 September 2022; leg. GH Han: 1♀ (MABIK CR00258877), dissected and mounted on 2 H-S slides. • Same locality, fishing net; 01 February 2024; leg. SL Kim and SJ Gwak: 5♀♀ (MinRB-Hr105-L003), preserved in a vial with 95% ethanol.
(3) • Masan-ri (36°00'58.99"N, 129°29'05.55"E), Pohang, Gyeongsangbuk-do, green algae; 27 February 1997; leg. J Lee; 1♂ (MinRB-Hr105-L004), preserved in a vial with 95% ethanol. • Same area, coralline algae (36°00'59.90"N, 129°28'58.32"E); 23 April 2005; leg. J Lee: 1♀ (MABIK CR00258878, asymmetry in the number of setae on P2 enp-2), dissected and mounted on 2 H-S slides and 1♀ (MABIK CR00258879), preserved in a vial with 95% ethanol. • Same area (36°00'59.90"N, 129°28'58.32"E), coralline algae; 26 April 2005; leg. J Lee: 2♀♀ (MinRB-Hr105-L005, asymmetry in the number of setae on P2 enp-2), preserved in a vial with 95% ethanol.
Female. Body length 827–1,098 μm (n = 17; measured from anterior margin of rostrum to posterior margin of caudal rami). Habitus slender, cylindrical, and gradually tapering posteriorly in dorsal view, separation between prosome and urosome indistinct (Fig.
Caudal rami (Figs
Rostrum (Fig.
Antennule (Fig.
Antenna (Fig.
Mandible (Fig.
Maxillule (Fig.
Maxilla (Fig.
Maxilliped (Fig.
P1 (Fig.
P2 and P3 (Fig.
P4 (Fig.
Armature formula of P1–P4 as follows:
| Exopod | Endopod | |
|---|---|---|
| P1 | 0.0.022 | 0.020 |
| P2 | 0.1.123 | 0.220 |
| P3 | 0.1.123 | 0.2–321[0.220 in ♂] |
| P4 | 0.1.023 | 0.121 |
P5 (Fig.
Male. Body (Fig.
Urosome (Figs
Antennule (Fig.
P2–P3 (Fig.
P4 (Fig.
P5 (Fig.
P6 (Fig.
Previous descriptions of H. discophora (e.g.,
In a female specimen from Gohado, caudal seta IV on the right caudal ramus was underdeveloped and deeply embedded within the ramus (Fig.
Heterolaophonte discophora (Willey, 1929) was originally described by
Heterolaophonte discophora was first reported from Korea in the doctoral dissertation by
One of the diagnostic traits of H. discophora is the shape of the P4 exp-3, which is slender and elongated, gradually broadening distally in its proximal part (
Comparative analysis of the Korean specimens with previous records of H. discophora reveals four distinct morphological traits.
In addition, the Korean specimens exhibit several minor differences from previous accounts. They have three setae on the endopod of the maxilla, whereas
Sexual dimorphism in male appendages, particularly the swimming legs, has long been recognized as a crucial source of characters for taxonomic identification and phylogenetic inference within the Laophontidae (
The genus Pseudonychocamptus comprises a morphologically consistent group of species that aligns well with the current general diagnosis. As noted by
The genus Heterolaophonte, originally established by
Sexual dimorphism in the swimming legs of Heterolaophonte can be assessed in the 27 species for which both sexes are known. In these species, male-specific modifications are most frequently observed on the P2 and P3 exopods, with occasional elongation of the P4 exopod compared to females. Two key characteristics are also evident in the P3 endopod: it is generally two-segmented, although some reports describe it as three-segmented—likely reflecting misinterpretation of a weak suture between segments (as discussed in the remarks on H. discophora); and the outermost seta on the P3 enp-2 is commonly transformed into an apophysis. Based on these patterns, a character matrix summarizing sexual dimorphism in the swimming legs across the 27 species is presented in Table
Character matrix summarizing sexual dimorphism in the swimming legs of Heterolaophonte species for which both sexes are available (only 27 species). See character definitions below. Abbreviation and symbol: C = character; ? = character state unconfirmed.
| C1 | C2 | C3 | C4 | C5 | C6 | C7 | C8 | C9 | C10 | C11 | |
|---|---|---|---|---|---|---|---|---|---|---|---|
| H. discophora | 1 | 1 | 0 | 1 | 1 | 0 | 0 | 0 | 2 | 1 | 1 |
| H. livingstoni | 1 | 1 | 0 | 1 | 0 | 0 | 0 | 1 | 2 | 1 | 1 |
| H. tenuispina | 0 | 1 | 0 | 1 | 0 | 0 | 0 | 1 | 2 | 1 | 1 |
| H. stroemii | 1 | 1 | 0 | 1 | 0 | 0 | 0 | 0 | 2 | 1 | 1 |
| H. murmanica | 1 | 1 | 0 | ? | ? | 0 | 0 | 0 | 2 | 1 | 1 |
| H. heejinae | 1 | 1 | 0 | 1 | 1 | 0 | 0 | 0 | 2 | 1 | 1 |
| H. pauciseta | 1 | 1 | 0 | 1 | ? | 0 | 0 | 0 | 1 | 1 | 1 |
| H. brevipes | 1 | 1 | 0 | 1 | 1 | 0 | 0 | 0 | 1 | 1 | 1 |
| H. denticulata | 1 | 1 | 0 | 1 | 0 | 0 | 0 | 0 | 1 | 1 | 1 |
| H. serratula | 1 | 1 | 0 | 1 | 1 | 0 | 0 | 0 | 1 | 1 | 1 |
| H. norvegica | 1 | 1 | 0 | ? | ? | 0 | 0 | 0 | 2 | 1 | 1 |
| H. uncinata | 1 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 2 | 1 | 2 |
| H. curvata | 1 | 1 | 0 | 1 | 1 | 0 | 0 | 0 | 2 | 1 | 2 |
| H. manifera | 1 | 1 | 0 | 1 | ? | 0 | 0 | 0 | 2 | 1 | 2 |
| H. hamata | 1 | 1 | 0 | 1 | 1 | 0 | 0 | 1 | 2 | 1 | 2 |
| H. minuta | 1 | 1 | 0 | 1 | 0 | 0 | 0 | 0 | 2 | 0 | 0 |
| H. laurentica* | 1 | 1 | 0 | ? * | ? * | 0 | 0 | 1 | 2 | 0 | 0 |
| H. lalanai | 1 | 1 | 0 | 0 | 1 | 0 | 0 | 2 | 0 | 0 | 0 |
| H. littoralis | 0 | 0 | 1 | 0 | 1 | 1 | 1 | 1 | 2 | 1 | 3 |
| H. longisetigera | 0 | 0 | 1 | 0 | 1 | 1 | 1 | 1 | 2 | 1 | 3 |
| H. mendax | 0 | 0 | 1 | 0 | 1 | ? | 0 | 1 | 2 | 1 | 3 |
| H. natator | 0 | 0 | 1 | 0 | 1 | 0 | 1 | 1 | 2 | 1 | 3 |
| H. hamondi | 0 | 0 | 1 | 0 | 1 | 1 | 1 | 0 | 0** | 2 | 0 |
| H. letovae | 0 | 0 | 1 | 0 | 1 | 1 | 1 | 0 | 0** | 2 | 0 |
| H. islandica | 0 | 0 | 1 | 0 | 1 | 1 | 1 | 0 | 0** | 2 | 0 |
| H. campbelliensis | 0 | 1 | 1 | 0 | 1 | 1 | 0 | 1 | 2 | ? | ? |
| H. variabilis | 0 | 1 | 1 | 0 | 1 | 1 | 0 | 1 | 2 | 1 | 3 |
These four groups are characterized as follows:
This morphological classification provides a framework to critically reassess previously proposed autapomorphic features of the genus Heterolaophonte, particularly those outlined by
These findings highlight the morphological heterogeneity within Heterolaophonte and suggest that none of the three proposed male traits can reliably serve as autapomorphies for the genus. Clarifying the phylogenetic structure of the genus will require integrative approaches combining detailed morphological assessments with molecular data. In particular, efforts should focus on discovering and describing the male morphology of species currently known only from females, as well as testing morphological hypotheses using independent datasets.
Floating anthropogenic debris, particularly plastics, has increasingly been recognized as significant vector for the passive dispersal of marine organisms (
Heterolaophonte discophora was found on various types of marine debris—specifically a rope, an expanded polystyrene buoy, and a fishing net—all covered with attached macroalgae, including green algae. This, with previous records of the species from habitats associated with coralline or green algae (
Pseudonychocamptus setadefectus sp. nov. was collected from a gunny sack and an expanded polystyrene buoy, neither of which were covered with macroalgae. This indicates no apparent ecological association with macroalgal substrates, in contrast to H. discophora. Among harpacticoid fauna from the MPD, the gunny sack supported a diverse assemblage of copepods, including species of Dactylopusia, Parathalestris Brady & Robertson, 1873, Harpacticus, Paralaophonte, and Sarsamphiascus, as well as fouling macroinvertebrates such as the Mediterranean mussel, Mytilus galloprovincialis Lamarck,1819 and the striped barnacle, Amphibalanus improvisus (Darwin, 1854). Conversely, the polystyrene buoy yielded only a few harpacticoid taxa (Ameira and Folioquinpes Fiers & Rutledge, 1990), with no macroinvertebrates observed on its surface. Although various invertebrates are known to serve as hosts or substrates for certain laophontid species, it remains unclear whether P. setadefectus sp. nov. exhibits such an association. These findings suggest that its occurrence on these substrates may instead be facilitated by morphological adaptations common in Laophontidae, such as the strongly developed claw on the P1 enp-2 and the robust maxilliped, which enable individuals to grasp fibrous or irregular surfaces (
In conclusion, the occurrence of laophontid harpacticoid copepods on MPD underscores the potential role of these substrates not only in enhancing the dispersal capacity of benthic species but also in creating novel microhabitats for colonization.
We are grateful to the editor and two reviewers for their constructive comments and valuable suggestions, which helped us to improve the manuscript. We also appreciate the Macrobenthic Invertebrate Research Team at KIOST for collecting samples and sharing the results of their analysis of macroinvertebrates attached to marine debris.
The authors have declared that no competing interests exist.
No ethical statement was reported.
No use of AI was reported.
This research was supported by the management of Marine and Fishery Bio-resources Center (2025) funded by the National Marine Biodiversity Institute of Korea (MABIK) and by the research program of the Korea Institute of Ocean Science & Technology (Contract No. PEA0304).
Kyuhee Cho conceived and designed the experiments, identified the specimens, prepared materials, figures and/or tables, authored or reviewed drafts of the paper, approved the final draft. Jong Guk Kim helped in the identification, authored or reviewed drafts of the paper, approved the final draft. Jimin Lee conceived and designed the experiments, authored or reviewed drafts of the paper, approved the final draft.
Kyuhee Cho https://orcid.org/0000-0002-5006-5713
Jong Guk Kim https://orcid.org/0000-0001-5299-9838
Jimin Lee https://orcid.org/0000-0001-9004-8275
All of the data that support the findings of this study are available in the main text.