Research Article |
Corresponding author: Samuel Gómez ( samuelgomez@ola.icmyl.unam.mx ) Academic editor: Danielle Defaye
© 2018 Samuel Gómez.
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:
Gómez S (2018) New species of the genus Mesocletodes Sars, 1909 from the deep Gulf of California (Copepoda, Harpacticoida). ZooKeys 751: 75-112. https://doi.org/10.3897/zookeys.751.20387
|
Investigations on the effects of the oxygen minimum zone on the distribution, abundance, and diversity of deep-sea benthic and pelagic fauna of the Gulf of California and Eastern Tropical Pacific has received attention recently. However, very little is known about the diversity of deep-sea benthic harpacticoids from this region, and only three species, Ancorabolus hendrickxi Gómez & Conroy-Dalton, 2002, Ceratonotus elongatus Gómez & Díaz, 2017 and Dendropsyllus californiensis Gómez & Díaz, 2017, have been described so far. The genus Mesocletodes Sars, 1909 is one of the most common and abundant genera of deep-sea harpacticoids. This genus has been traditionally subdivided into two species groups, the abyssicola and the inermis groups, based on the presence/absence of a dorsal cuticular process on the cephalothorax and anal somite, but some species have been reported to deviate from this scheme. As a result of their investigations, other researchers proposed the monophyly of the abyssicola group, and suggested the probable monophyly of two other species-groups. In this paper, the descriptions of three new species of the genus Mesocletodes from the deep sea of the Gulf of California are presented with some notes on their relationships with other species. Some comments on the monophyly of the genus are given.
Argestidae , Mesocletodes , deep sea, new species, taxonomy
Great effort has been deployed since the late 1980’s to study the biodiversity of the deep sea of the Gulf of California and Eastern Tropical Pacific (a complete list of contributions is available upon request). Extensive oceanographic campaigns (Talud I-XIII cruises) on board the research vessel “El Puma” of the Universidad Nacional Autónoma de México (
The genus Mesocletodes Sars, 1909 is one of the most common and abundant genera of the family Argestidae in deep-sea samples (
In this paper three new species of Mesocletodes from the deep sea of the Gulf of California are proposed. Additionally, some comments on the monophyly of Mesocletodes are provided.
Sediment samples for meiofaunal analyses were taken in August 2000 in the Southern Gulf of California from Carmen basin to off Nayarit State, and in February 2007 in the Southern Trough of Guaymas Basin, during Talud IV and Talud X cruises, respectively, on board the research vessel “El Puma” of the Universidad Nacional Autónoma de México (
Abbreviations used in the text:
acro acrotheke;
ae aesthetasc;
ENP endopod;
EXP exopod;
EXP (ENP)1 (2,3) first (second, third) exopodal (endopodal) segment;
P1–P6 first to sixth legs;
se pinnate, naked, setiform;
sp spinose, spiniform;
STE subapical tubulate extension.
The type material was deposited in the Copepoda collection of the Instituto de Ciencias del Mar y Limnología, Unidad Académica Mazatlán (ICML-EMUCOP). The map showing the sampling locations where the new species were found were prepared with GeoMapApp (http://www.geomapapp.org/) and the Global Multi-Resolution Topography (GMRT) default basemap (
One female holotype as follows: habitus, left antennule and right antenna left intact and preserved in alcohol (ICML-EMUCOP-270800-02), right antennule, left antenna, mandibles, maxillules, maxillae and maxillipeds dissected and mounted onto four slides (ICML-EMUCOP-270800-01); Talud IV cruise; August 27, 2000; coll. S. Gómez.
Southern Carmen Basin, Gulf of California, México (25°54.7'N, 110°11'W), 2018 m depth (see Fig.
Sampling sites and type localities of Mesocletodes simplex sp. n. (full circle), and M. brevisetosus sp. n. and M. unisetosus sp. n. (full triangle). Figure prepared with GeoMapApp (http://www.geomapapp.org/) and the Global Multi-Resolution Topography (GMRT) default basemap (
Body subcylindrical. Cuticula of body surface reticulated. Free thoracic somites and urosomites, except for anal somite, with posterior margin coarsely serrated. Cephalothorax with dorsal process curved posteriorly. P3–P5-bearing somites and second half of genital-double somite with bifid cuticular processes dorsally. Genital somite and third urosomite incompletely fused dorsolaterally. Anal somite with upward bifid cuticular process dorsally. Caudal rami 17 times as long as wide in lateral view, with seven setae. Antennule octa-segmented; second segment with strong protrusion bearing one strong element. Antenna with basis, with uni-segmented exopod bearing two setae subequal in length. Gnathobase of mandible with grinding face, palp uni-segmented, endopodal lobe with four setae. P2–P4ENP1 with inner seta. P1–P4ENP2 with 3, 3, 2, 2, setae respectively. P5 endopodal lobe represented by two setae; inner seta of the P5EXP very small, issuing subapically.
Body: total length 1420 µm measured from anterior margin of rostrum to posterior margin of caudal rami, subcylindrical, tapering slightly posteriorly, without clear demarcation between prosome and urosome, cuticula of body surface reticulated (Fig.
Mesocletodes brevisetosus sp. n., female holotype. A habitus, lateral B cephalothorax, lateral, reticulated pattern omitted C dorsal cuticular process of cephalothorax showing situation of sensilla, reticulated pattern omitted DP2–P5-bearing somites, lateral, reticulated pattern omitted E genital double-somite, fourth and fifth urosomites, and anal somite, lateral, reticulated pattern omitted F left caudal ramus, lateral, reticulated pattern omitted G distal part of left caudal ramus, lateral, reticulated pattern omitted.
Anal somite (Figs
Caudal rami slender, exceedingly elongated, 17 times as long as wide in lateral view, as long as P4-bearing somite and entire urosome combined, gently curved upwards from lateral view, covered with small spinules (Fig.
Antennule (Fig.
Antenna (Fig.
Mandible (Fig.
Maxillule (Fig.
Maxilla (Fig.
Maxilliped (Fig.
P1 (Fig.
P2–P4 (Figs
P5 (Fig.
Armature formula as follows:
Unknown.
The specific epithet is derived from the Latin adjective brevis, meaning short, and the Latin noun seta, meaning hair, and refers to the reduced innermost seta of the female P5EXP. The name is a noun in the nominative singular.
Mesocletodes brevisetosus sp. n. seems to be more closely related to M. dorsiprocessus than to M. bicornis, both from the Angola Basin, by the combination of several characters: 1) serrated posterior margin of the cephalothorax and P2-bearing somite to penultimate urosomite (serrated posterior margin on cephalothorax, P2-bearing somite to first half of genital double-somite in M. bicornis), 2) relative length of the setae of the antennary exopod (subequal in M. dorsiprocessus and in M. brevisetosus sp. n., but one of them noticeably reduced in M. bicornis), 3) presence of an inner seta on P2–P4ENP1 in M. dorsiprocessus and M. brevisetosus sp. n., but absent in M. bicornis, 4) number of setal complements on P1–P4ENP2 (3, 3, 2, 2 in M. dorsiprocessus and M. brevisetosus sp. n., but 2, 4, 4, 4 in M. bicornis), 5) number of setae on the female P5 endopodal lobe (two setae in M. dorsiprocessus and M. brevisetosus sp. n., but three in M. bicornis), 6) position of the inner seta of the female P5EXP (issuing subapically in M. dorsiprocessus and M. brevisetosus sp. n., but situated in distal third of inner margin of ramus in M. bicornis), and 7) degree of development of the endopodal lobe of the female P5 (without any trace of endopodal lobe in M. dorsiprocessus and M. brevisetosus sp. n., but endopodal lobe discernible in M. bicornis). Mesocletodes dorsiprocessus and M. brevisetosus sp. n. can be separated based on the number of setae on the endopodal lobe of the mandibular palp (one seta in M. dorsiprocessus, but four elements in M. brevisetosus sp. n.), the relative length of the innermost seta of the female P5EXP (well-developed in M. dorsiprocessus, but very reduced in M. brevisetosus sp. n.), appearance of the cuticula (plain in M. dorsiprocessus, but reticulated in M. brevisetosus sp. n.), armature complement of the seventh antennular segment (six in M. dorsiprocessus, but four in M. brevisetosus sp. n.), and number of setae on the syncoxa of the maxilliped (one in M. dorsiprocessus, but two in M. brevisetosus sp. n.).
One dissected female holotype mounted onto five slides (ICML-EMUCOP-130207-01); Talud X cruise; February 13, 2007; coll. S. Gómez.
Southern Trough of Guaymas Basin, Gulf of California, México (27°01'N, 110°53’04"W), 1642 m depth (see Fig.
Body subcylindrical. Cephalothorax with dorsal cuticular process curved posteriorly. Genital somite and third urosomite incompletely fused dorsolaterally. Anal somite quadrate from dorsal view, with simple dorsal cuticular process curved posteriorly. Caudal rami subcylindrical, 2.5 times as long as wide, with seven setae. Antennule octa-segmented, second segment with protrusion bearing a long seta, but not as pronounced as in other species of the genus. Antenna with basis and uni-segmented exopod. Gnathobase of mandible with grinding face, and tri-segmented palp. Maxillary syncoxa with two endites, proximal endite with one, distal endite with three elements; endopod uni-segmented, with two setae. Syncoxa of maxilliped with one seta. P4ENP2 with four setae. Outer setae of the P5EXP issuing close to each other.
Body: total length 725 µm measured from anterior margin of rostrum to posterior margin of caudal rami, subcylindrical, tapering slightly posteriorly, without clear demarcation between prosome and urosome (Fig.
Anal somite (Figs
Caudal rami (Figs
Antennule (Fig.
Antenna (Fig.
Mandible (Fig.
Maxillule (Fig.
Maxilla (Fig.
Maxilliped (Fig.
P1 (Fig.
P2–P4 (Figs
P5 (Fig.
Armature formula as follows:
Unknown.
The specific epithet is derived from the Latin adjective simplex, meaning simple, and refers to the simple (not bifurcated) dorsal process on the anal somite. The name is an adjective in the nominative singular.
Mesocletodes simplex sp. n. is attributed here to
One male holotype preserved in alcohol (ICML-EMUCOP-270800-04), one male paratype preserved in alcohol (ICML-EMUCOP-270800-03), and one male paratype dissected and mounted onto seven slides (ICML-EMUCOP-270800-05); Talud IV cruise; August 27, 2000; coll. S. Gómez.
Southern Carmen Basin, Gulf of California, México (25°54.7'N, 110°11'W), 2018 m depth (see Fig.
Body subcylindrical. Cephalothorax, free prosomites and urosomites, except for anal somite, with posterior margin serrated. Cephalothorax dorsoventrally flattened, without dorsal cuticular process. Anal somite quadrate, with dorsal cuticular process. Caudal rami 14 times as long as wide, with seven elements. Antennule octa-segmented, haplocer, second segment with strong protrusion bearing one strong seta and two strong spinules. Antenna with basis, without exopod. Mandibles, maxillules, maxillae and maxillipeds strongly atrophied, nontraceable. P2 and P3ENP1 with one inner seta. P5 endopodal lobe with one seta.
Body: total length ranging from 655 µm to 695 µm (mean= 670 µm; n= 3) measured from anterior margin of rostrum to posterior margin of caudal rami, subcylindrical (Fig.
Anal somite (Figs
Caudal rami (Figs
Mesocletodes unisetosus sp. n., male paratype. A urosome, ventral, P5-bearing somite omitted B right caudal ramus, dorsal C distal part of right caudal ramus, dorsal, sowing seta III, IV, V, and VI D left caudal ramus, lateral E distal part of left caudal ramus, lateral, showing seta III, IV and V F medial part of left caudal ramus, lateral, showing seta I, II and VII.
Antennule (Fig.
Antenna (Fig.
Mandibles, maxillules, maxillae and maxillipeds strongly atrophied, non-traceable.
P1 (Fig.
P2–P4 (Figs
P5 (Fig.
Armature formula as follows:
Unknown.
The specific epithet is derived from the Latin prefix ūni, meaning one, and the Latin noun seta, meaning hair, and refers to the presence of one seta only on the endopodal lobe of the male P5. The name is a noun in the nominative singular.
The only species for which the male is known are M. angolaensis, M. elmari Menzel, 2011, M. fladensis, M. nudus and M. unisetosus sp. n. Of these, the female is known only for M. elmari. These species are attributed to
Additionally,
Some years later,
More recently,
No unequivocal apomorphies have been detected so far to justify the monophyly of the family Argestidae, and its monophyletic status has not yet been demonstrated (
Given all the above, there seems to be another approach towards the monophyly of the genus Mesocletodes. It seems plausible that this genus could eventually be attributed to a new subfamily defined by the four synapomorphies currently known for the genus, viz. the presence of a strong protrusion with a strong, bipinnate seta pointing backwards on the second antennular segment, the proximal outer spine of P1EXP3 reduced, the presence of STE’s on the spines of P1EXP3, and the mandibular gnathobase with a strong, grinding tooth, plus the presence of a dorsal cuticular process on the cephalothorax and anal somite.
Under this scheme, the presence of a dorsal process on the cephalothorax and anal somite, and the extreme elongation of the caudal rami between seta III and VII could be regarded as plesiomorphic within the subfamily, and the lack of such processes and the reduction of the caudal rami, as secondary apomorphic reductions. However, this requires more robust, in-depth analyses, accompanied by the diagnosis of this hypothesized subfamily, the re-diagnosis of the genus Mesocletodes, and the proposal of a new genus to include all the remaining species. These two genera could be composed as follows:
Hypothetical genus. M. dorsiprocessus, M. bicornis, and M. brevisetosus sp. n.; defined by the synapomorphic bifid dorsal processes on P3–P5-bearing somites and posterior half of genital double-somite. The bifid dorsal process on the cephalothorax could be regarded as autapomorphic for M. bicornis. The bifid dorsal process on the anal somite would be regarded as plesiomorphic. The monophyly of this taxon was suggested earlier by
Mesocletodes.– M. monensis, M. abyssicola, M. bathybia, M. brevifurca, M. dolichurus, M. katharinae, M. meteorensis, M. robustus, M. soyeri, M. opoteros, M. simplex sp. n., M. quadrispinosa, M. fladensis, M. angolaensis, M. unisetosus sp. n., M. irrasus, M. inermis, M. langi Smirnov, 1946, M. makarovi, M. glaber Por, 1964b, M. guillei, M. farauni Por, 1967, M. commixtus Coull, 1973, M. bodini Soyer, 1975, M. carpinei Soyer, 1975, M. ameliae Soyer, 1975, M. parirrasus Becker Noodt & Schriever, 1979, M. sarsi Becker Noodt & Schriever, 1979, M. parabodini Schriever, 1983, M. trisetosa, M. variabilis Schriever, 1983, M. kunzi Schriever, 1985, M. duosetosus Schriever, 1985, M. elmari, M. tetrasetosus, and M. nudus; defined by the secondary synapomorphic loss of the dorsal process of the cephalothorax and/or anal somite and reduction of the caudal rami. The elongation of the caudal rami between seta III and VII in some of these species, and the bifid dorsal process on the anal somite of M. opoteros are, therefore, plesiomorphic. The presence of the latter in M. opoteros and in the previous genus could support a closer relationship between these two taxa. Four species groups without taxonomic value, for which no apomorphies have been detected, can be envisaged based on the presence/absence of a dorsal cuticular process on the cephalothorax and/or anal somite:
I M. abyssicola, M. brevifurca, M. bathybia, M. dolichurus, M. katharinae, M. meteorensis, M. monensis, M. opoteros, M. robustus, M. simplex sp. n., and M. soyeri
II M. quadrispinosa
III M. angolaensis, M. fladensis, and M. unisetosus sp. n.
IV M. ameliae, M. bodini, M. carpinei, M. commixtus, M. duosetosus, M. elmari, M. farauni, M. glaber, M. guillei, M. inermis, M. irrasus, M. kunzi, M. langi, M. makarovi, M. nudus, M. parabodini, M. parirrasus, M. sarsi, M. tetrasetosus, M. trisetosa, and M. variabilis.
Group I includes
Finally, it is worth mentioning the presence of some other undescribed forms related to M. angolaensis, M. bicornis, M. brevisetosus sp. n., M. dorsiprocessus, M. meteorensis, M. simplex sp. n., and M. unisetosus sp. n. in the Clarion-Clipperton Fracture Zone in the Pacific Ocean (Samantha Tong Jia Wen, Tropical Marine Science Institute, National University of Singapore, pers. comm.).
This study was financed by the Programa de Apoyo a Proyectos de Investigación e Innovación Tecnológica (PAPIIT) of the Dirección General de Asuntos del Personal Académico of the Universidad Nacional Autónoma de México (