Research Article |
Corresponding author: Huyen T. M. Pham ( minhhuyen.ks.nb@gmail.com ) Academic editor: Saskia Brix
© 2022 Huyen T. M. Pham, Ivana Karanovic.
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:
Pham HTM, Karanovic I (2022) Four new Parasterope (Ostracoda, Myodocopina) from the Northwest Pacific and their phylogeny based on 16S rRNA. ZooKeys 1095: 13-42. https://doi.org/10.3897/zookeys.1095.77996
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Parasterope Kornicker, 1975 is a marine ostracod genus with 49 species described so far, which makes it the most diverse representative of the subfamily Cylindroleberidinae, as well as the entire family Cylindroleberididae. Despite its global distribution no species are reported from South Korea. Three new species collected from the Korean coast of the Sea of Japan (Parasterope busanensis sp. nov., P. singula sp. nov., and P. sohi sp. nov.), and one from the Japanese coast of the Pacific Ocean (P. sagami sp. nov.) are described. A taxonomic key to all named species from East Asia is provided. A phylogenetic tree is reconstructed based on partial 16S rRNA sequences of the four new species and other Cylindroleberidinae available from GenBank. Monophyly of Parasterope is supported by high posterior probabilities, but the phylogenetic analyses also indicate that some of the GenBank data attributed to this genus are probably misidentifications. A map of distribution and a checklist of all described Parasterope species are also provided.
Crustacea, East Asia, marine benthos, new species, taxonomic key
Cylindroleberididae is one of the largest myodocopid families with remarkable morphological diversity. It accounts for 225 species classified in 33 genera and the following four subfamilies: Cylindroleberidinae, Cyclasteropinae, Asteropteroninae, and Macroasteropteroninae. The primary defining feature of the family is the presence of seven or eight leaf-like pairs of gills at the posterior end of the body, the presence of a “baleen-comb” on the maxilla and the fifth limb, a mandible with a sword-shaped coxal endite, and a hatchet shaped sixth limb (
The cylindroleberid fauna of East Asia is generally poorly known and mostly consists of species records without many details on their morphology. For example, a systematic study of the Korean ostracods as indicators of water pollution lists nine Cylindroleberidae, all left in the open nomenclature (
In the present paper, we describe four new species belonging to Parasterope that have been collected from seas around Korea and Japan. To facilitate their further identification, we provide a taxonomic key to all East Asian species. In addition, partial 16S rRNA sequences of all new species and partial 18S rRNA sequences of only three new species were successfully amplified. Currently, there are two 16S rRNA sequences, five 18S rRNA sequences, and eleven 28S rRNA sequences available on GenBank (
Samples from Japan were collected from Sagami Bay during the 6th JAMBIO (Japanese Association for Marine Biology) project, on 13 February 2015 with a dredge from shallow waters (130–241m) (
The extraction followed the HotSHOT method described in
List of 16S sequences used for phylogenetic analysis. 1
Genus | Species | GenBank number | |
---|---|---|---|
16S | 18S | ||
Bathyleberis | B. oculata | EU587251 1 | EU591814 1 |
C. marranyin | EU587243 1 | ||
Cylindroleberis | Cylindroleberis J57069 | EU587253 1 | EU587244 1 |
Cylindroleberis NW-2004 | AY624729 2 | ||
UnID | Cylindroleberididae J57076 | EU587257 1 | |
Parasterope | P. busanensis sp. nov. 24_6 | OK048681 | OK048719 |
P. busanensis sp. nov. 24_7 | OK048682 | OK048720 | |
P. gamurru J53224 | EU587255 1 | EU591819 1 | |
P. pollex | AF363309 4 | ||
P. sagami sp. nov. 26_9 | OK048683 | OK048721 | |
P. sagami sp. nov. 27_0 | OK048684 | OK048722 | |
P. singula sp. nov. 1_1 | OK048686 | OK048723 | |
P. singula sp. nov. 1_2 | OK048687 | ||
P. sohi sp. nov. | OK048685 | ||
P. styx | EU587236 1 | ||
Parasterope J57072* | EU587247 1 | ||
Parasterope NW-2004 | AY624728 2 | ||
Postasterope | P. barensi J57079 | EU587258 1 | EU587248 1 |
P. corrugata | EU587259 1 | EU591816 1 | |
Synasterope | Synasterope J57066 | EU587252 1 | EU587250 1 |
Synasterope J57067 | EU591815 1 | ||
Toyoshioleberis | T. magnabucca | MW534153 3 | MZ092883 3 |
Xenoleberis | X. parvus | MW534150 3 | |
X. pacifica 38 | MW534151 3 | MZ092881 3 | |
X. pacifica 39 | MW534152 3 | MZ092882 3 | |
X. pacifica 275 | MW534140 3 | MZ092883 3 | |
X. tanakai 266 | MW534141 3 | ||
X. tanakai 284 | MW534142 3 | MZ092884 3 | |
X. yamadai 272 | MW534143 3 | MZ092879 3 | |
X. yamadai 273 | MW534144 3 | MZ092880 3 | |
X. yamadai 14 | MW534145 3 | MZ092878 3 | |
X. yamadai 30 | MW534147 3 |
Forward and Reverse strands were visually compared and checked for signal quality and low-resolution sites using FinchTV (version 1.4.0) (Geospiza, Inc., Seattle, WA, USA; http://www.geospiza.com). The strands were aligned with MAFFT v.7 127b (
A1 first antenna;
A2 second antenna;
F Furca;
L5 fifth limb;
L6 sixth limb;
L7 seventh limb;
Md mandibula;
Mxl maxillula;
SEM scanning electron microscope.
Phylum Arthropoda Latreille, 1829
Subphylum Crustacea Brünnich, 1772
Class Ostracoda Latreille, 1802
Subclass Myodocopa Sars, 1866
Order Myodocopida Sars, 1866
Family Cylindroleberididae Müller, 1906
Genus Parasterope Kornicker, 1994
1 | Mandible, dorsal margin of basale with mid-bristle | 2 |
– | Mandible, dorsal margin of basale without mid-bristle | 3 |
2 | 6th limb, ventral and postero-ventral margin without plumose bristles | P. nana Poulsen, 1965 |
– | 6th limb, ventral margin of end-joint with 11 or 12 plumose bristles | P. zamboangae Kornicker, 1970 |
3 | Adult female Md with 3 long terminal setae next to exopodite | P. sagami sp. nov. |
– | Adult female Md with 2 long terminal setae next to exopodite | 4 |
4 | 7th limb with 4 bristles (2 on each side) on the terminal segment | P. singula sp. nov. |
– | 7th limb with 6 bristles (3 on each side) on the terminal segment | 5 |
5 | Mandibular basale with 6 spinous end bristles | P. sohi sp. nov. |
– | Mandibular basale with 3 or 4 spinous end bristles | 6 |
6 | Mandibular basale spinous and with cluster of spines near middle | P. mckenziei Kornicker, 1970 |
– | Mandibular basale with no hair on broad surface | 7 |
7 | First antenna 8th joint with small spine-like d-bristle | P. hirutai Chavtur, 1983 |
– | First antenna 8th joint without d-bristle | 8 |
8 | An open row of ~ 7–15 long medial bristles between the posterior ridge and the shell margin | P. obesa Poulsen, 1965 |
– | No bristles or only short, scattered bristles between the posterior ridge and the shell margin | 9 |
9 | 2nd joint of first antenna without lateral bristle; second antenna protopodite with a row of spines on the ventral margin | P. jenseni Poulsen, 1965 |
– | 2nd joint of first antenna with lateral bristle; second antenna protopodite without spine on the ventral margin | P. busanensis sp. nov. |
Holotype
female dissected on one slide, shells on SEM stub (
This species is named in reference to the type locality. It is used as an adjective for a place, with the Latin suffix -ensis.
Surface of the shell mostly smooth. Posterior infold of the carapace with a broad shelf. Dorsal margin rounded in both female and male. Posterior end clearly wider than anterior. Lateral eye well developed with black pigmented ommatidia. A1 8-segmented without d-bristle on the 8th joint. Uropodal lamellae with nine claws.
Female. Shell (Fig.
Light microscopy and scanning electron microscopy images of Parasterope busanensis sp. nov. a–f holotype female adult: a External view b Lateral view from left valve c Spines on dorsal margin d Internal view of anterior part e L5 ventral section f Internal view of posterior part g, h paratype male adult: g External view h Lateral view from right valve i Internal view of left valve.
A1 (Fig.
Bellonci organ (Fig.
Eyes . Lateral eye with many ommatidia obscured by black pigment; 13 ommatidia visible. Medial eye rounded and unpigmented.
A2. Protopodite: rounded without medial bristle. Endopodite (Fig.
Md (Fig.
Mxl (Fig.
L5 (Fig.
L6. Unknown
L7 (Fig.
F (Fig.
Male. All features are comparable to adult female; important differences are listed below.
Shell (Fig.
A1 (Fig.
A2 (Fig.
Md (Fig.
Eyes
(Fig.
Reproductive organs
(Fig.
The new species is closely related to P. iota Kornicker, Harrison-Nelson & Coles, 2007, described from Kapua Channel (Waikïkï, Hawaii). The first joint of the A1 of P. busanensis is without hair on the broad surface, present in P. iota. In addition, the 3rd–8th joints of P. iota each have fairly stout proximal ventral spines, absent in P. busanensis. The distal margin of the Mxl of P. busanensis bears four bristles compared to two in P. iota, and three bristles (one minute) near the ventral margin rather than five in P. iota. The new species shares many characters with P. gamurru Syme & Poore, 2006 described from Queensland, Australia (
The sexual dimorphism found in the new species, the morphology of the dorsal mid-bristle on the basale of Md, and the number of bristles proximal to the a-bristle on the endopodite of the Md, have never been described in Parasterope.
GenBank numbers 16S: OK048681, OK048682; 18S: OK048719, OK048720.
Holotype
female dissected on one slide, shells on SEM stub (
The species name was chosen after the Sagami Bay, from where the species was collected. It is an adjective agreeing with female gender of the genus.
Surface of the shell completely smooth. Posterior infold of the carapace with a broad shelf. Dorsal margin rounded in female and almost perpendicular in male. Lateral eye well developed with black pigmented ommatidia, smaller in female than in male. A1 8-segmented without d-bristle on the 8th joint. Uropodal lamellae with eight claws.
Female. Shell (Fig.
Light microscopy and scanning electron microscopy images of Parasterope sagami sp. nov. a–c holotype female adult: a External view b Lateral view from right valve c Internal view from left valve. d–f paratype male adult: d External view e Lateral view from right valve f Internal view of left valve.
A1 (Fig.
Bellonci orange (Fig.
Eyes
(Fig.
A2 (Fig.
Md (Fig.
Mxl (Fig.
L5 (Fig.
L6 (Fig.
L7 (Fig.
F (Fig.
Male. All features comparable to adult female; important differences are:
Shell (Fig.
A1 (Fig.
Eyes. Lateral eye with 18 ommatidia obscured by black pigment.
A2 (Fig.
Md (Fig.
Reproductive organs. Unknown.
Parasterope sagami differs from all other Parasterope representatives by the combination of the following characters:
Female. Dorsal margin of the Md basale has three long terminal setae next to the exopodite, and it has no mid-bristle. In addition, the A1 g-bristle has a peculiar shape, and there is no lateral bristle on the A1 2nd joint;
Male. The presence of the c- and d-claws bristle-like on the Md endopodite 2nd joint in the adult males is unique for the new species.
In addition to the above characters, P. sagami differs from P. busanensis and the morphology of the endite II of Mxl. Namely, the endite carries three rather than four bristles. Also, the short α-seta on the Mxl exopodite is absent in P. busanensis.
Parasterope jenseni was also described from Sagami Sea, Japan (
Parasterope obesa Poulsen, 1965 from Misaki, Japan has a lateral bristle on the 2nd joint of A1 which is absent in the two above species; 4th joint of the A1 is without lateral spines on the ventral margin otherwise present in the new species; and there are more than ten bristles on the ventral edge of the comb.
GenBank numbers 16S: OK048683, OK048684; 18S: OK048721, OK048722.
Holotype
female dissected on one slide, shells on SEM stub (
The name is a Latin noun, singula, because only a female has been collected. The name is in nominative, feminine singular, agreeing in gender with the genus.
Surface of the shell completely smooth. Anterior end with a deep incisure and posterior infold of the carapace with a broad shelf. Posterior end wider than anterior. Dorsal margin rounded. Lateral eye well developed with black pigmented ommatidia. Uropodal lamellae with nine claws.
Female. Shell (Fig.
A1 (Fig.
Bellonci orange (Fig.
Eyes. Lateral eye with 18 ommatidia.
A2 (Fig.
Md (Fig.
Mxl. Endite I with three bristles. Endite II with three bristles. Basale: one dorsal medial distal bristle, two ventral medial proximal bristles. Endopodite: 1st with one short α-seta and one hairy β-seta; 2nd article with two terminal bristles.
L5 (Fig.
L6 (Fig.
L7 (Fig.
F (Fig.
Male. Not collected.
The presence of rows of small spines on the broad surface near the dorsal margin of the Md basale is a unique feature of Parasterope singula. Additionally, the number of bristles on the L7 (ten bristles: four on the terminal segment (two on each side), six on the proximal segments (three on each side) is less than in all other Parasterope representatives. The new species is similar to P. mckenziei, described from Samar Province, Philippines (McKenzie, 1970) but differs in the following characters: the Md endopodite is without a proximal bristle next to the a-bristle, a character not known in any other Parasterope, and the number of claws on UL (nine vs. seven).
The 2nd joint of the Md endopodite in P. singula carries one oblique row of cleaning setae, also recorded in P. antyx Kornicker, 1989 from Bay of Biscay (Kornicker, 1989), P. gamma Kornicker, Harrison-Nelson & Coles, 2007 from Hawaiian Islands (
Holotype
female dissected on one slide, shells on SEM stub (
The species named after Prof. Ho-young Soh (Faculty of Marin Technology, Chonam National University, Jeonnam, Korea), whom we are greatly indebted to for collecting and providing samples for this publication.
Carapace elongated. Surface of the shell completely smooth. Posterior end with a short, rounded incisure, while the anterior has relatively deep incisure. Dorsal and ventral margins rounded. Posterior end wider than anterior. Lateral eye well developed with black pigmented ommatidia.
Female. Shell (Fig.
A1 (Fig.
Bellonci orange (Fig.
Eyes. Lateral eye with 15 ommatidia.
A2 (Fig.
Md (Fig.
Mxl. Endite unknown. Basale: one dorsal medial distal bristle. Endopodite: 1st with one short α-seta and one hairy β-seta; 2nd article with two terminal bristles.
L5. Unknown.
L6. Unknown.
L7. Unknown.
F. Unknown.
Male. Not collected.
The new species Parasterope sohi sp. nov. is poorly known because of the missing L5, L6, L7, and furca but we propose a new name because of the presence of the combination of the following characters: A1 sensory bristle carry 0+6 filaments; exopodite of Md is unusually long; and the Md endopodite has the e-bristle. In majority of Parasterope described so far, the mandible exopodite is at most longer than ½ length of the dorsal margin of the 1st endopodite joint; in P. sohi the exopodite length is greater than the length of the dorsal margin of the 1st endopodite joint. Other species with such a long exopodite include P. lagunicola from French Polynesia (
GenBank number 16S: OK048685. 18S could not be obtained.
A checklist of species the 49 named Parasterope, including four new species described in this study, is presented in Suppl. material
The final alignment of the 16S data set consisted of 33 sequences 445 base pairs long, of which 210 were constant sites (= 47.191% of all sites), 210 were invariant (= 47.191% of all sites), 207 were parsimony informative, and 275 were distinct site patterns. The substitution model TPM2+F+I+G4 was found to be the best fit evolutionary model. Effective sample size for all the continuous parameters (posterior, prior, tree likelihood, tree height, Yule model, and birthrate) estimated by Tracer analysis was far above a recommended number (200). Phylogram is presented on the Fig.
The resulting phylogenetic tree based on the newly obtained sequences and those belonging to the subfamily Cylindroleberidinae reconstructed based on the 16S alignment (Fig.
The unusual clustering of Parasterope and several unidentified Cylindroleberididae and Cylindroleberis on the phylogenetic tree, assigned to other genera, should be considered as misidentification and/or contaminations, and not an indication of polyphyletic nature of Parasterope. This can be supported by the results from previous, more complex, phylogenetic analyses of the entire family Cylindroleberididae (
Due to a very limited number of available sequences attributed to the subfamily Cylindroleberidinae (27 in total) our phylogenetic tree included only 18 species belonging to seven genera. This can be considered a very poor sample, since the subfamily contains ~ 200 species belonging to 23 genera (see
The results of pairwise distance analysis show a fast evolutionary rate of 16S rRNA, in contrary to 18S rRNA. Calculated pairwise distances between 18S sequences were very similar to those of other for Cylindroleberidae, Cypridinidae, and Philomedidae published by
In the
In terms of biodiversity, our results are important as they provide new data for the marine fauna of Korea and Japan. They also point out that samples taken from areas where human impact on ecosystems is high due to dense population and industrial development can yield new species, and enrich our knowledge on the biodiversity of the planet.
We would like thank Dr. Hayato Tanaka (Tokyo Sea Life Park) and the crews of TR/V Toyoshio-maru and R/V Rinkai-maru for providing the Japanese specimens, Prof. Ho-young Soh (Faculty of Marin Technology, Chonam National University, Jeonnam, Korea), Dr. Hyunsu Yoo (Marine Environmental Research and Information Laboratory (MERIL), and the Marine Biodiversity Institute of Korea (MABIK) for providing additional Korean specimens. This study was funded by the Basic Science Research Program through NRF funded by the Ministry of Education (grant number: 2020R1A6A1A06046827), and the National Institute of Biological Resources (
A checklist species of Parasterope Kornicker, 1975
Data type: Checklist
Explanation note: A checklist of species of Parasterope Kornicker, 1975.
Table S1
Data type: gene distances
Explanation note: Pairwise p-distances among 16S sequences of species presented in Fig.
Table S2
Data type: gene distances
Explanation note: Between group means distance of 16S sequences from Cylindroleberidinae genera. IG: intrageneric.
Table S3
Data type: gene distances
Explanation note: Between group means distance of 18S sequences from Cylindroleberidinae genera. IG: intrageneric.
Table S4
Data type: gene distances
Explanation note: Pairwise p-distances among 18S sequences of species presented in Table