Research Article |
Corresponding author: Eduardo Suárez-Morales ( esuarez@ecosur.mx ) Academic editor: Danielle Defaye
© 2022 Juan M. Fuentes-Reinés, Eduardo Suárez-Morales, Marcelo Silva-Briano.
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:
Fuentes-Reinés JM, Suárez-Morales E, Silva-Briano M (2022) A new species of Nitokra Boeck, 1865 (Copepoda, Harpacticoida, Ameiridae) from the Caribbean coast of Colombia. ZooKeys 1128: 111-127. https://doi.org/10.3897/zookeys.1128.86210
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Biological samples obtained from a coastal system of northern Colombia yielded male and female specimens of an undescribed harpacticoid copepod of the diverse ameirid genus Nitokra Boeck, 1865. The new species is a member of the genus group III. We describe the new species based on adult male and female individuals. Nitokra puebloviejensis sp. nov., appears to be most closely related to N. vietnamensis Tran & Chang, 2012, but they can be separated by the following characters: 1) number of setal elements on second segment of mandibular palp, 2) P1ENP/EXP ratio, 3) relative lengths of P2, P3ENP/EXP, 4) number of elements on male P5EXP and ENP, and 5) segmentation of male antennule. In addition, N. puebloviejensis sp. nov. can be confused with two other congeners: N. taylori Gómez, Carrasco & Morales-Serna, 2012 from South Africa and Colombia and N. kastjanensis Kornev & Chertoprud, 2008 from the White Sea, but the new species can be distinguished from them by: 1) number of setae on the maxillule coxa, 2) P1ENP/EXP ratio, 3) P2,P3ENP/EXP ratio, 4) female and male P5 setophore, 5) setation pattern of female P5EXP and ENP, 6) structure of female P6, 7) ornamentation of female anal operculum, 8) number of setae on male P5EXP, and 9) the male antennule segmentation. Most importantly, the presence of a group of five short setae on the medial surface of the maxilliped syncoxa allows the new species to be readily distinguished from its congeners. Only two subspecies and one species of this genus have been hitherto recorded from Colombia. A key to the 23 known American species of Nitokra is provided.
Benthic copepods, brackish waters, harpacticoids, new species, northern Colombia, taxonomy
The family Ameiridae Monard, 1927 is one of the most diverse of the copepod order Harpacticoida, and it has been divided into two subfamilies: Stenocopiinae Lang, 1944, and Ameirinae Boeck, 1865, the latter being the most diverse. Currently, the family comprises 49 genera and about 303 species (
The knowledge on the diversity of Nitokra in Colombia is still scarce. Hitherto, only one species and three subspecies have been recorded in the country: N. lacustris colombianus from Bahía Solano, Choco, N. l. sinoi from Ciénaga Grande de Santa Marta, Magdalena and Laguna Navío Quebrado, la Guajira, and N. affinis colombiensis and N. taylori from Laguna Navío Quebrado, La Guajira (
The Ciénaga Grande de Santa Marta, a large costal system of northern Colombia, was biologically surveyed during 2017 as part of an ongoing effort aiming to increase our knowledge of the Colombian aquatic biodiversity. The samples obtained yielded male and female specimens of an undescribed species of Nitokra. The new species is described and compared it with its closest congeners. A key to the 23 species of Nitokra known to occur in the Americas is also provided.
Biological samples were obtained monthly from littoral habitats of the Ciénaga Grande de Santa Marta, northern Colombia (10°52'11.25"N, 74°19'31.64"W) in July, 2022; samples were collected manually from areas with mangrove vegetation using a 25 L bucket. Water salinity, pH, and temperature were measured in situ with a WTW350i Multimeter.
Samples were filtered with a plankton net (45 μm mesh size) and then fixed and preserved in 70% ethanol. Copepods were sorted from the original samples and then processed for taxonomical identification, including dissection and mounting of taxonomically relevant appendages. Dissected specimens were mounted in glycerin and sealed with Canada balsam. Drawings of the mounted appendages were prepared with a camera lucida; they were also photographed using a Kodak Easy Share C140 digital camera adapted to a compound microscope. Two adult male individuals were prepared for SEM examination with a JEOL LV 5900 microscope at the
University of Aguascalientes (UAA),
Mexico; one female individual was prepared for SEM examination with a JSM-6010LA microscope at El Colegio de la Frontera Sur, Chetumal, Mexico. The whole specimens were measured in lateral position, from the tip of rostrum to the posterior margin of the caudal rami. Morphological nomenclature follows
Order Harpacticoida G.O. Sars, 1903
Family Ameiridae Boeck, 1865
Subfamily Ameirinae Boeck, 1865
Adult female holotype (INV10139), ethanol-preserved, vial, Ciénaga Grande de Santa Marta, Colombia (10°5211.25"N, 74°19'31.64"W), littoral plankton, coll. J.M. Fuentes-Reinés; adult male allotype (INV CRU10140), ethanol-preserved, vial, same sampling data as holotype. Paratypes: four females (INV CRU10141), and three males (INV CRU10142), same sampling data as holotype and allotype.
Six adult females, four adult males in first authors’ collection. One female and two male individuals prepared for SEM analysis.
Puebloviejo, Ciénaga Grande de Santa Marta, northern Colombia (10°52'11.25"N, 74°19'31.64"W).
The new species is named in reference to the type locality of the new species by adding the toponimic suffix in singular. The gender of the species suffix is feminine to match that of the genus.
Nitokra with 1 inner seta and 5 setae on P1EXP2 and EXP3, respectively and 455 and 777 elements on P2–P4ENP3 and P2–P4EXP3, respectively, plus 111 inner setae on P2–P4ENP1, respectively. Female rostrum hourglass-shaped, distal segment of mandibular palp with 6 setal elements. P1ENP1 almost reaching distal margin of P1EXP3. Modified, club-shaped inner basipodal seta on leg1. Maxilliped with distinctive group of 5 setae inserted medially on the syncoxa.
Body subcylindrical, tapering posteriorly (Fig.
Nitokra puebloviejensis sp. nov. from Puebloviejo, Ciénaga Grande de Santa Marta, Colombia, digital photos. A holotype female, habitus, lateral view B rostrum, ventral view C labrum, ventral view D urosomites and caudal rami, ventral view E anal somite with anal operculum and spine ornamentation, dorsal view F anal somite and caudal rami showing caudal setae I–VII, ventral view. Scale bars: 50 μm (A, C); 10 μm (B); 20 μm (E, F).
Rostrum small, slightly protruding, discernible in dorsal view; roughly hourglass-shaped, with flat tip; rostrum furnished apically with two pairs of short slender sensilla (Figs
Antennule
8-segmented, tapering distally (Fig.
Antenna
(Figs
Nitokra puebloviejensis sp. nov. from Puebloviejo, Ciénaga Grande de Santa Marta, Colombia. Adult female holotype. A rostrum B labrum C mandible with palp D gnathal blade with teeth and dorsal seta E antenna (some setae cut short) F maxillule showing armature of lobes G maxilla H maxilliped showing row of setal elements on syncoxa I detail of accessory seta of maxilliped ENP J antennule showing segmentation. Scale bars: 15 μm (A–D, I); 20 μm (D–F); 25 μm (G, H, J).
Nitokra puebloviejensis sp. nov. from Puebloviejo, Ciénaga Grande de Santa Marta, Colombia. SEM-prepared adult female. A antenna showing EXP and ENP B posterior end of anal somite and insertion of caudal rami showing ornamentation, semi-lateral view C fifth leg exopods (at arrow), ventral view D anal somite and caudal rami, ventral view E anal somite, ventral view F urosome, ventral view.
Mandible
(Fig.
Maxillule
(Fig.
Maxilla
(Fig.
Maxilliped
(Fig.
P1 (Fig.
P2 (Fig.
P3 (Fig.
P4 (Fig.
P5 (Fig.
Nitokra puebloviejensis sp. nov. from Puebloviejo, Ciénaga Grande de Santa Marta, Colombia. A adult female anal and preanal urosomites, dorsal view showing anal operculum and caudal setae I–VII B female anal and preanal urosomites, ventral view C female P5 D female genital field and P6 with setae (arrowed) E male geniculated antennule F modified, club-shaped basipodal spine of male P1 G male leg 6 H male P3, anterior view I male P5, ventral view. Scale bars: 25 μm (A–D, I); 10 μm (E–G); 20 μm (H).
P6 (Fig.
Armature formula of female P1–P5 as follows:
Exopod | Endopod | |
---|---|---|
P1 | I-0; I-1; III,2,0 | 0-1;0-1;I,I,2,0 |
P2 | I-0; I-1;III,2,2 | 0-1;0-1; I,2,1 |
P3 | I-0; I-1;III,2,2 | 0-1;0-1; I,2,2 |
P4 | I-0;I-1;III,2,2 | 0-1;0-1;I,2,2 |
P5 | 5 | 5 |
Male. Smaller than female, total body length 578–588 μm (average length = 578 μm, n = 3; allotype specimen length = 588 μm).
Anal operculum, rostrum, antennae (Fig.
Nitokra puebloviejensis sp. nov. from Puebloviejo, Ciénaga Grande de Santa Marta, Colombia. SEM-prepared adult male individual A antennules, semi-lateral view B same, showing detail of proximal segments C antenna, posterior view D same showing exopodal ramus and endopodal segments E maxilliped, ventral view showing row of setal elements on syncoxal medial surface (arrowhead) F leg 1 showing modified, club-shaped basipodal spine (at arrow).
Antennule
(Fig.
Antenna
(Fig.
P1 basis and P3ENP3. P1 basis with modified, club-shaped inner spine (Fig.
P5 (Fig.
P6. With 2 unequal setae, inner one about 3× as long as outer seta. Caudal rami as in female.
One male with 3 setal elements (instead of 4) on P5ENP. Another male was observed to possess 6 setae instead of 5 on P5EXP.
The new species is known only from the type locality, Puebloviejo, Ciénaga Grande de Santa Marta (northern Colombia). The site where it was collected is a shallow mangrove area, 0.7 m deep, with water temperature 26–31 °C; local salinity was 15–20 PSU, and pH values was 7.5–8.1.
There are 23 species of the genus reported from the Americas, as follows: N. typica typica Boeck, 1865, N. spinipes spinipes Boeck, 1865, N. lacustris lacustris (Schmankevitch, 1875), N. hibernica hibernica (Brady, 1880), N. lacustris sinoi Marcus & Por, 1961, N. pusilla Sars, 1911, N. bdellurae (Liddell, 1912), N. affinis affinis Gurney, 1927, N. affinis affinis colombiensis Fuentes-Reinés & Suárez-Morales, 2014, N. dubia G. O. Sars, 1927, N. minor minor Willey, 1930, N. chelifer Wilson, 1932, N. hyperidis Jakobi, 1956, N. fragilis paulistana Jakobi, 1956, N. spinipes armata Lang, 1965, N. affinis californica Lang, 1965, N. lacustris colombiana Reid, 1988, N. sphaeromata Bowman, 1988, N. galapagoensis Mielke, 1993, N. bisetosa Mielke, 1993, N. evergladensis
In a partial revision of Nitokra,
Furthermore, N. puebloviejensis can be confused with N. taylori, another congener known from the Colombian Caribbean, but these two species can be distinguished by the following characters: 1) maxillule coxa with 3 setal elements in N. puebloviejensis (Fig.
The diversity of the ameirid harpacticoid fauna could be underestimated and deserves further study in the Caribbean region.
1 | P4EXP3 with 6 elements | 2 |
1a | P4EXP3 with 7 elements | 3 |
1b | P4EXP3 with 8 elements | 21 |
2 | P2 and P3EXP3 with 7 and 7 elements, respectively | N. bisetosa Mielke, 1993 |
– | P2 and P3EXP3 with 6 and 5 elements, respectively | N. lacustris richardi (Karanovic, 2015) |
3 | P5 female endopodite and exopodite with 6 and 5 elements. respectively | 4 |
– | P5 female endopodite and exopodite with 5 and 5 elements, respectively | 13 |
4 | P4ENP1 with inner seta | 5 |
– | P4ENP1 without inner seta | 8 |
5 | P2EXP3 with seven elements | 6 |
– | P2EXP3 with six elements | 7 |
6 | P1EXP3 almost reaching P1ENP3, anal operculum with 3 or 4 spines | N. taylori Gómez, Carrasco & Morales-Serna, 2012 |
– | P1EXP3 reaching the end margin of P1ENP2, anal operculum without spines | N. bdellurae (Liddell, 1912) |
7 | P1ENP1 reaching the half of P1EXP3, anal operculum about 15 spines, P5EXP male with 5 elements | N. chelifer Wilson, 1932 |
– | P1ENP1 overpassing P1EXP3, anal operculum with 9 spines, P5EXP male with 4 elements | N. typica typica Boeck, 1865 |
8 | P1ENP1 overpassing P1EXP3 | 9 |
– | P1ENP1 not reaching beyond P1EXP3 | 10 |
9 | P2 and P3ENP3 with 4 and 5 elements, respectively; P3EXP3 with 6 elements; anal operculum lacking spines | N. pusilla Sars, 1911 |
– | P2 and P3ENP3 with 2 and 3 elements, respectively; P3EXP3 with 5 elements; anal operculum with about 10 spines | N. hibernica hibernica (Brady, 1880) |
10 | P1ENP1 as long as EXP1 and EXP2 combined | 11 |
10a | P1ENP1 reaching 1/3 or ½ of P1EXP3 | N. lacustris sinoi Marcus & Por, 1961 |
10b | P1ENP1 reaching ½ of P1EXP2 | 12 |
11 | Anal operculum with 7 spines, baseoendopodite of P5 female almost reaching distal end of EXP | N. evergladensis Bruno & Reid, 2002 |
– | Anal operculum with about 10 spines, P5 baseoendopodite reaching middle of EXP | N. dubia Sars, 1927 |
12 | With a spinulate medial surface of the caudal rami and caudal seta II about 2.2 as long as seta I | N. lacustris colombiana Reid, 1988 |
– | Spinulate medial surface absent in the caudal rami and caudal seta II less 2 times as long as seta I | N. lacustris lacustris (Schmankevitch, 1875) |
13 | P4ENP3 with 4 elements | N. hyperidis Jakobi, 1956 |
– | P4ENP3 with 5 elements | 14 |
14 | P2ENP3 with 4 elements, P4ENP2 with 1 element | 15 |
– | P2ENP3 with 3 elements, P4ENP2 unarmed | N. galapagoensis Mielke, 1997 |
15 | P2ENP1 and P4ENP1 without inner set | N. minor minor Willey, 1930 |
– | P2ENP1 and P4ENP1 with inner seta | 16 |
16 | P1ENP1 reaching the margin end of P1EXP2 | N. fragilis paulistana Jakobi, 1956 |
– | P1ENP1 longer than P1EXP2 | 17 |
17 | P1EXP3 reaching the insertion point of inner seta of ENP1 | N. sphaeromata Bowman, 1988 |
– | P1EXP3 going beyond the insertion point of inner seta ENP1 | 18 |
18 | Anal operculum with 6 spines; female and male P6 with 1 and 2 setal elements, respectively; male P5EXP with 5 elements; maxilliped with 4 or 5 setal elements on the syncoxa medial surface | N. puebloviejensis sp. nov. |
– | Characters not as above | 19 |
19 | Anal operculum with 10–15 spines, female and male P6 with 2 and 3 setal elements, respectively; male P5EXP with 6 elements; maxilliped lacking medial setae on the syncoxa | 20 |
20 | Anal operculum with 10–12 spines, posterior margin of genital double somite with row of long lateral spinules | N. spinipes armata Lang, 1965 |
– | Anal operculum with 12–15 spines, posterior part of double genital somite with short row of lateral spinules | N. spinipes spinipes Boeck, 1865 |
21 | Rostrum with long projection | N. affinis colombiensis Fuentes-Reinés & Suárez-Morales, 2014 |
– | Rostrum without long projection | 23 |
23 | P1ENP1 longer than exopodite, posterior edge of antepenultimate somite spinulose dorsal to ventrolateral | N. affinis affinis Gurney, 1927 |
– | P1ENP1 as long as exopodite, posterior edge of antepenultimate somite encircled by spinules | N. affinis californica Lang, 1965 |
Dr Samuel Gómez (UNAM, Unidad Academica Mazatlan) provided useful taxonomic literature during our work on Colombian Nitokra. Araceli Adabache (University of Aguascalientes, Mexico) kindly helped us in processing specimens of the new species for SEM examination and guided our observations. MS-B was supported by University of Aguascalientes project PIB19-2 UAA. We appreciate the editorial processing of our work by the Associate Editor Danielle Defaye.