Research Article |
Corresponding author: Samuel Gómez ( samuelgomez@ola.icmyl.unam.mx ) Academic editor: Kai Horst George
© 2022 Samuel Gómez, Beatriz Yáñez-Rivera.
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, Yáñez-Rivera B (2022) The genus Cletocamptus (Harpacticoida, Canthocamptidae): a reappraisal, with proposal of a new subfamily, a new genus, and a new species. ZooKeys 1080: 165-208. https://doi.org/10.3897/zookeys.1080.71192
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A new species of Cletocamptus closely related to C. helobius was found in sediment samples taken from a polluted estuarine system in north-western Mexico. The genus Cletocamptus was relegated to species incertae sedis in 1986, and this finding prompted us to evaluate the current taxonomic position of the genus within the Canthocamptidae. The latter has been subdivided in several, seemingly unnatural subfamilies in the past to better understand the relationships between its constituent taxa. In this study we propose a new subfamily, the Cletocamptinae subfam. nov. for Amphibiperita, Cletocamptus, and Cletocamptoides gen. nov., defined by the synapomorphic subdistal ventral spinules on the rostrum. The genus Cletocamptoides gen. nov. is proposed for C. helobius, C. merbokensis, and C. biushelo sp. nov., and is supported by the ‘cletodid’ shape of the body and the reduced one-segmented endopod of the fourth swimming leg. Cletocamptus includes all the other species with long slender spinules on the posterior margin of prosomites and with the sexually modified inner spine on the second endopodal segment of the second swimming leg in the males. Amphibiperita retained the primitive female fifth leg with exopod and baseoendopod separated, and the primitive prehensile endopod of the first leg, but is defined by the loss of the antennary exopod. Other (syn)apomorphies are given, and the evolution of the mandibular palp is briefly discussed. Additionally, a diagnosis for the new subfamily, Cletocamptinae subfam. nov., an amended narrower diagnosis for Cletocamptus, the diagnosis for Cletocamptoides gen. nov., and a phylogenetic analysis supporting the proposal of these new taxa, are given.
Copepoda, Crustacea, diversity, meiofauna, phylogeny, systematics
Several canthocamptid species were found in meiofauna samples taken during a short-term study on the effects of organic pollution on the diversity and abundance of harpacticoids. Cletocamptus sinaloensis Gómez, Fleeger, Rocha-Olivares & Foltz, 2004 was by far the most abundant canthocamptid, followed by Mesochra cf. pygmaea (Claus, 1863), but the most intriguing was a form of Cletocamptus Schmankevitsch, 1875 closely related to C. helobius Fleeger, 1980, the latter known from salt marshes in Louisiana (
Sediment samples were taken at several sampling stations along Urías system (north-western Mexico) (see
BENP baseoendopod;
ENP endopod;
EXP exopod;
EXP (ENP)1 (2, 3) first (second, third) exopodal (endopodal) segment;
P1–P6 first to sixth legs.
A phylogenetic analysis was performed with 39 characters and 36 taxa (Tables
Character | Description |
---|---|
1 | Body shape: cletodid, with somitic constrictions between somites (1), non-cletodid, without somitic constrictions between somites (0) |
2 | Posterior ornament of the cephalothorax: unornamented (0), finely serrated/crenulated (1), coarsely serrated (2), short spinules (3), long spinules (4) |
3 | Posterior ornament of free prosomites: unornamented (0), finely serrated/crenulated (1), coarsely crenulated (2), coarsely serrated (3), short spinules (4), long spinules (5) |
4 | Sensilla-bearing socles: absence (0), presence (1) |
5 | Rostrum: not fused to cephalothorax (0), fused to cephalothorax (1) |
6 | Rostrum ventral ornament: unornamented (0), spinules or setules (1) |
7 | Number of segments of the female antennule 1: nine (0), eight (1), seven (2), six (3), five (4), four (5) |
8 | Number of abexopodal setae on the allobasis of the antenna: two (0), one (1) none (2) |
9 | Number of segments of the antennary exopod: three (0), two (1), one (2), one- seta (3), absent (4) |
10 | Number of segments of the mandibular palp: one (0), two or more (1) |
11 | Ramification of the mandibular palp: biramous (exopod and endopod discernible even if rami fused to basis) (0), monoramous (only endopod discernible even if fused to basis) (1) |
12 | Number of setae on the mandibular palp: four (0), three (1), two (2), one (3), unarmed (4) |
13 | Number of setae on the syncoxa of the maxilliped: four (0), one (1), unarmed (2) |
14 | Shape of P1 ENP: prehensile (0), not prehensile (1) |
15 | Number of segments of P1 ENP: three (0), two (1), one (2), represented by spine or seta (3), absent (4) |
16 | Inner setae on P1 ENP1 when two or three-segmented: present (0), absent (1) |
17 | Number of segments of P2 ENP when present: three (0), two (1), one (2) |
18 | Inner seta of P2 ENP1 when ENP two- or three-segmented: present (0), absent (1) |
19 | Inner seta of P2 ENP2 when ENP three-segmented: present (0), absent (1) |
20 | Number of setae of P2 ENP2 when ENP two-segmented: five (0), four (1), three (2), two (3), one (4), unarmed (5) |
21 | Number of setae of P2 ENP3 when ENP three-segmented: five (0), four (1), three (2), two (3) |
22 | Number of setae of P3 EXP3 when EXP three-segmented: eight (0), seven (1), six (2), five (3), four (4) |
23 | Number of segments of the female P3 ENP when present: three (0), two (1), one (2) |
24 | Inner seta of P3 ENP1 when ENP two- or three-segmented: present (0), absent (1) |
25 | Number of setae of P3 ENP1 when ENP one-segmented: three (0), two (1) |
26 | Number of setae of the female P3 ENP2 when ENP two-segmented: six (0), five (1), four (2), three (3), two (4) |
27 | Number of setae of P4 EXP3 when EXP three-segmented: eight (0), seven (1), six (2), five (3), four (4) |
28 | Number of segments of P4 ENP when present: three (0), two (1), one (2), represented by spine or seta (3) |
29 | Inner seta of P4 ENP1 when ENP two-segmented: present (0), absent (1) |
30 | Number of setae of P4 ENP1 when ENP one-segmented: four (0), three (1), two (2), one (3) |
31 | Number of setae of P4 ENP2 when ENP two-segmented: five (0), four (1), three (2), two (3), one (4) |
32 | Subdistal spine of P2 ENP: present (0), absent (1) |
33 | Subdistal spine of P3 ENP: present (0), absent (1) |
34 | Female P5 BENP/EXP fusion: not fused (0), fused (1) |
35 | Male rostrum dimorphism: present (1), absent (0) |
36 | Male P5 BENP/EXP fusion: not fused (0), fused (1) |
37 | Number of setae of male P5 ENP lobe: six (0), three (1), two (2), one (3), unarmed (4) |
38 | Number of segments of male P3: three (0), two (1) |
39 | Shape of inner apophysis of male P3: arrow-head tip (0), simple (1) |
Subfamily | Species | Characters | ||||||||||||||||||||||||||||||||||||||
---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
1–10 | 11–20 | 21–30 | 31–39 | |||||||||||||||||||||||||||||||||||||
1 | 2 | 3 | 4 | 5 | 6 | 7 | 8 | 9 | 0 | 1 | 2 | 3 | 4 | 5 | 6 | 7 | 8 | 9 | 0 | 1 | 2 | 3 | 4 | 5 | 6 | 7 | 8 | 9 | 0 | 1 | 2 | 3 | 4 | 5 | 6 | 7 | 8 | 9 | ||
Cletocamptinae subfam. nov. | Amphibiperita neotropica | 0 | 3 | 4 | 0 | 0 | 1 | 3 | 1 | 4 | 1 | 1 | 4 | 1 | 0 | 1 | 0 | 1 | 0 | - | 2 | - | 2 | 1 | 1 | - | 2 | 2 | 1 | 1 | - | 2 | 1 | 1 | 0 | 1 | 1 | 1 | 1 | 1 |
Cletocamptoides biushelo sp. nov. | 1 | 1 | 1 | 0 | 0 | 1 | 3 | 0 | 3 | 0 | 1 | 3 | 1 | 1 | 1 | 1 | 1 | 0 | - | 2 | - | 3 | 1 | 1 | - | 3 | 4 | 2 | - | 2 | - | 0 | 0 | 1 | ? | ? | ? | ? | ? | |
Cletocamptoides helobius comb. nov. | 1 | ? | ? | 0 | 0 | 1 | 3 | 1 | 3 | 0 | 1 | 3 | 2 | 1 | 1 | 1 | 1 | 0 | - | 2 | - | 3 | 1 | 1 | - | 3 | 4 | 2 | - | 3 | - | 0 | 0 | 1 | ? | 1 | 1 | 1 | 1 | |
Cletocamptoides merbokensis comb. nov. | 1 | 1 | 1 | 1 | 0 | 1 | 3 | 1 | 2 | 0 | 0 | 2 | 2 | 1 | 1 | 1 | 1 | 0 | - | 3 | - | 3 | 1 | 1 | - | 4 | 4 | 2 | - | 2 | - | 1 | 1 | 1 | 0 | 1 | 1 | 0 | 1 | |
Cletocamptus albuquerquensis | 0 | 4 | 5 | 0 | 0 | 1 | 3 | 0 | 2 | 0 | 1 | 4 | 1 | 1 | 1 | 0 | 1 | 1 | - | 2 | - | 3 | 1 | 1 | - | 1 | 4 | 1 | 1 | - | 3 | 0 | 0 | 1 | 1 | 1 | 1 | 1 | 1 | |
Cletocamptus assimilis | 0 | 4 | 5 | 0 | 0 | 1 | 3 | 0 | 2 | 0 | 1 | 4 | 1 | 1 | 1 | 0 | 1 | 1 | - | 2 | - | 3 | 1 | 1 | - | 3 | 4 | 1 | 1 | - | 3 | 0 | 0 | 1 | ? | ? | ? | ? | ? | |
Cletocamptus axi | 0 | 4 | 5 | 0 | 0 | 1 | 3 | 0 | 2 | 0 | 1 | 4 | 1 | 1 | 1 | 0 | 1 | 1 | - | 2 | - | 3 | 1 | 1 | - | 3 | 4 | 1 | 1 | - | 3 | 0 | 0 | 1 | 1 | 1 | 1 | 0 | 1 | |
Cletocamptus cecsurirensis | 0 | 4 | 5 | 0 | 0 | 1 | 3 | 0 | 2 | 0 | 1 | 4 | 1 | 1 | 1 | 0 | 1 | 1 | - | 2 | - | 3 | 1 | 1 | - | 3 | 4 | 1 | 1 | - | 3 | 0 | 0 | 1 | 1 | 1 | 1 | 0 | 1 | |
Cletocamptus confluens | 0 | 4 | 5 | 0 | 0 | 1 | 3 | 0 | 2 | 0 | 1 | 3 | 1 | 1 | 1 | 0 | 1 | 1 | - | 2 | - | 3 | 1 | 1 | - | 1 | 4 | 1 | 1 | - | 3 | 0 | 0 | 1 | ? | 1 | 1 | 1 | 1 | |
Cletocamptus deborahdexterae | 0 | 4 | 5 | 0 | 0 | 1 | 3 | 0 | 2 | 0 | 1 | 4 | 1 | 1 | 1 | 0 | 1 | 1 | - | 2 | - | 3 | 1 | 1 | - | 3 | 4 | 1 | 1 | - | 3 | 0 | 0 | 1 | 1 | 1 | 1 | 0 | 1 | |
Cletocamptus dominicanus | 0 | 1 | 1 | 0 | 0 | 1 | 3 | 0 | 2 | 0 | 1 | 4 | 1 | 1 | 1 | 0 | 1 | 1 | - | 2 | - | 3 | 1 | 1 | - | 1 | 4 | 2 | - | 2 | - | 0 | 0 | 1 | 1 | 1 | 1 | 1 | 1 | |
Cletocamptus fourchensis | 0 | 4 | 5 | 0 | 0 | 1 | 3 | 0 | 2 | 0 | 1 | 4 | 1 | 1 | 1 | 1 | 1 | 1 | - | 2 | - | 3 | 1 | 1 | - | 3 | 4 | 1 | 1 | - | 3 | 0 | 0 | 1 | 1 | 1 | 1 | 0 | 1 | |
Cletocamptus goenchim | 0 | 4 | 5 | 0 | 0 | 1 | 3 | 0 | 2 | 0 | 1 | 4 | 1 | 1 | 1 | 0 | 1 | 1 | - | 2 | - | 2 | 1 | 1 | - | 1 | 3 | 1 | 1 | - | 3 | 0 | 0 | 1 | 1 | 1 | 1 | 0 | 1 | |
Cletocamptus gomezi | 0 | 4 | 5 | 0 | 1 | 1 | 3 | 1 | 2 | 0 | 1 | 4 | 1 | 1 | 1 | 0 | 1 | 1 | - | 1 | - | 3 | 1 | 1 | - | 1 | 4 | 1 | 1 | - | 3 | 0 | 0 | 1 | ? | 1 | 1 | 0 | 1 | |
Cletocamptus levis | 0 | 4 | 5 | 0 | 0 | 1 | 3 | 0 | 2 | 0 | 1 | 4 | 1 | 1 | 1 | 0 | 1 | 1 | - | 2 | - | 3 | 1 | 1 | - | 3 | 4 | 1 | 1 | - | 3 | 0 | 0 | 1 | 1 | 1 | 1 | 0 | 1 | |
Cletocamptus nudus | 0 | 4 | 5 | 0 | 0 | 1 | 3 | 0 | 2 | 0 | 1 | 4 | 1 | 1 | 1 | 0 | 1 | 1 | - | 2 | - | 2 | 1 | 1 | - | 3 | 3 | 1 | 1 | - | 3 | 0 | 0 | 1 | 1 | 1 | 1 | 0 | 1 | |
Cletocamptus pilosus | 0 | 4 | 5 | 0 | 0 | 1 | 3 | 0 | 2 | 0 | 1 | 4 | 1 | 1 | 1 | 0 | 1 | 1 | - | 1 | - | 2 | 1 | 1 | - | 1 | 3 | 1 | 1 | - | 3 | 0 | 0 | 1 | 1 | 1 | 1 | 0 | 1 | |
Cletocamptus retrogressus | 0 | 4 | 5 | 0 | 0 | 1 | 3 | 1 | 2 | 1 | 1 | 3 | 1 | 1 | 1 | 1 | 1 | 1 | - | 1 | - | 2 | 1 | 1 | - | 1 | 3 | 1 | 1 | - | 3 | 0 | 0 | 1 | 1 | 1 | 1 | 0 | 1 | |
Cletocamptus samariensis | 0 | ? | ? | 0 | 0 | 1 | 3 | 0 | 2 | 0 | 1 | 4 | 1 | 1 | 1 | 0 | 1 | 1 | - | 2 | - | 2 | 1 | 1 | - | 3 | 3 | 1 | 1 | - | 3 | 0 | 0 | 1 | 1 | 1 | 1 | 0 | 1 | |
Cletocamptus schmidti | 0 | 4 | 5 | 0 | 0 | 1 | 3 | 0 | 2 | 0 | 1 | 4 | 1 | 1 | 1 | 0 | 1 | 1 | - | 2 | - | 2 | 1 | 1 | - | 3 | 3 | 1 | 1 | - | 3 | 0 | 0 | 1 | 1 | 1 | 1 | 0 | 1 | |
Cletocamptus sinaloensis | 0 | 4 | 5 | 0 | 0 | 1 | 3 | 0 | 2 | 0 | 1 | 4 | 1 | 1 | 1 | 0 | 1 | 1 | - | 2 | - | 3 | 1 | 1 | - | 3 | 4 | 1 | 1 | - | 3 | 0 | 0 | 1 | 1 | 1 | 1 | 0 | 1 | |
Cletocamptus spinulosus | 0 | 0 | 5 | 0 | 0 | 1 | 3 | 0 | 2 | 0 | 1 | 4 | 1 | 1 | 1 | 0 | 1 | 1 | - | 2 | - | 3 | 1 | 1 | - | 3 | 4 | 1 | 1 | - | 3 | 0 | 0 | 1 | 1 | 1 | 1 | 0 | 1 | |
Cletocamptus stimpsoni | 0 | 4 | 4 | 0 | 0 | 1 | 3 | 0 | 2 | 0 | 1 | 4 | 1 | 1 | 1 | 0 | 1 | 1 | - | 2 | - | 2 | 1 | 1 | - | 1 | 3 | 1 | 1 | - | 3 | 0 | 0 | 1 | 1 | 1 | 1 | 0 | 1 | |
Cletocamptus tainoi | 0 | 4 | 4 | 0 | 0 | 1 | 3 | 0 | 2 | 0 | 1 | 4 | 1 | 1 | 1 | 0 | 1 | 1 | - | 2 | - | 3 | 1 | 1 | - | 1 | 4 | 1 | 1 | - | 3 | 0 | 0 | 1 | 1 | 1 | 1 | 1 | 1 | |
Cletocamptus tertius | 0 | 4 | 5 | 0 | 0 | 1 | 3 | 0 | 2 | 0 | 1 | 4 | 1 | 1 | 1 | 0 | 1 | 1 | - | 2 | - | 3 | 1 | 1 | - | 3 | 4 | 1 | 1 | - | 3 | 0 | 0 | 1 | 1 | 1 | 1 | 0 | 1 | |
Canthocamptinae | Canthocamptus incurvisetosus | 0 | 0 | 0 | 0 | 1 | 0 | 1 | ? | 1 | ? | ? | 4 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | - | 0 | 1 | 1 | 0 | - | 1 | 1 | 1 | 0 | - | 0 | 0 | 0 | 0 | 0 | 0 | 2 | 0 | 0 |
Canthocamptus odaeensis | 0 | 0 | 0 | 0 | 1 | 0 | 1 | ? | 1 | 1 | 0 | 4 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | - | 0 | 1 | 1 | 0 | - | 1 | 1 | 1 | 0 | - | 0 | 0 | 0 | 0 | 0 | 0 | 2 | 0 | 0 | |
Elaphoidella bulgarica | 0 | 0 | 0 | 0 | ? | ? | 1 | 1 | 2 | ? | ? | ? | ? | 1 | 0 | 0 | 1 | 0 | 0 | - | - | 2 | 1 | 0 | 1 | - | 2 | 2 | - | 1 | - | 0 | 0 | 0 | 0 | 0 | 4 | 0 | 1 | |
Elaphoidella_ formosanus | 0 | 0 | 0 | 0 | 1 | 0 | 1 | ? | 2 | ? | ? | ? | ? | 1 | 0 | 0 | 1 | 1 | 0 | - | - | 2 | 1 | 1 | 1 | - | 3 | 1 | 0 | - | 1 | 0 | 0 | 0 | 0 | 0 | 4 | 0 | 1 | |
Elaphoidella hirsutus | 0 | 2 | 3 | 0 | 1 | 0 | 1 | ? | 2 | ? | ? | ? | ? | 1 | 0 | 0 | 1 | 0 | 0 | - | - | 2 | 1 | 1 | 0 | - | 2 | 1 | 1 | - | 1 | 0 | 0 | 0 | ? | ? | ? | ? | ? | |
Hemimesochrinae | Isthmiocaris laurae | 0 | 0 | 0 | 0 | 0 | 0 | 3 | 1 | 2 | 0 | 0 | 3 | 1 | 1 | 1 | 1 | 2 | - | - | - | - | 4 | 2 | - | 1 | - | 4 | 2 | - | 3 | - | 0 | 0 | 1 | 0 | 0 | 2 | 0 | 0 |
Isthmiocaris longitelson | 0 | 0 | 0 | 0 | 0 | 0 | 3 | 1 | 2 | 1 | 1 | 4 | 1 | 1 | 2 | - | - | - | - | - | - | 5 | - | - | - | - | - | - | - | - | - | - | - | 1 | 0 | 1 | 3 | 0 | 0 | |
Itunella arenaria | 0 | 0 | 2 | 0 | 0 | 0 | 2 | 1 | 2 | 0 | 0 | 4 | 1 | 1 | 1 | 1 | 2 | - | - | - | - | 3 | 2 | - | 0 | - | 3 | 2 | - | 1 | - | 0 | 0 | 0 | 0 | 0 | 2 | 1 | - | |
Hepactophaninae | Stygepactophanes_ occitanus | 0 | 0 | 0 | 0 | 1 | 0 | 2 | 0 | 2 | 0 | 1 | 4 | 2 | 1 | 1 | 1 | 1 | 1 | - | 2 | - | 4 | 1 | 1 | - | 3 | 3 | 2 | - | 2 | - | 0 | 0 | 0 | ? | ? | ? | ? | ? |
Epactophanes philippinus | 0 | 0 | 0 | 0 | 1 | 0 | 2 | 1 | 2 | 0 | 1 | 4 | 2 | 1 | 1 | 1 | 1 | 1 | - | 4 | - | 3 | 1 | 1 | - | 4 | 3 | 2 | - | 3 | - | 1 | 1 | 0 | 0 | 1 | 4 | 0 | 1 | |
Outgroup | Podogennonta | 0 | 0 | 0 | 0 | ? | ? | 0 | ? | 0 | 1 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | 0 | - | 0 | 0 | 0 | 0 | - | - | 0 | 0 | 0 | - | - | ? | ? | 0 | ? | 0 | 1 | 0 | ? |
Weights | 5 | 1 | 3 | 1 | 1 | A | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 5 | 1 | 1 | 1 | 5 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 1 | 3 | 1 | 1 | 1 | 1 | 1 | 5 | 1 | 1 | 1 | 1 | 1 |
The maximum parsimony analysis was performed using Phylip ver. 3.697 (
In addition to the limited number of canthocamptid taxa used for our phylogenetic analysis, we consulted the available descriptions of 102 species distributed in 45 genera of the different canthocamptid subfamilies for our comparative analysis.
Family Canthocamptidae Brady, 1880
Cletocamptus Schmankevitsch, 1875.
Amphibiperita Fiers & Rutledge, 1990, Cletocamptoides gen. nov.
Canthocamptidae. Body fusiform, without clear distinction between prosome and urosome. Without nuchal organs on cephalothorax or body somites. Female rostrum distinct, rarely fused to cephalothorax, large and triangular with broad proximal margin; ornamented with ventral (sub)distal spinules or (occasionally) setules and with rounded or (rarely) bilobed tip in both sexes. Cephalothorax and/or prosomites with posterior margins serrated or ornamented with short or long slender spinules; posterior margin of urosomites except for anal somite serrated or with short spinules; posterior margin of cephalothorax and body somites without or (rarely) with cuticular sensillum-bearing socles. Anal operculum without dorsal ornamentation or with transverse row of strong large or short, small spinules; posterior margin unornamented, serrated, or with small or large spinules. Female genital somite and third urosomite separated dorsolaterally, completely fused ventrally forming genital double-somite. Female antennule six-, rarely seven-segmented. Antenna with allobasis, with one or two abexopodal setae (proximal element basal, distal seta endopodal); exopod one-segmented, longer than wide or minute, or absent. Mandibular palp one-segmented, very small and wider than long, or longer than wide; basis and endopod incorporated to basis or (rarely) with basis and endopod distinct (uniramous); exopod absent or (rarely) represented by single seta; when palp uniramous, then basis unarmed or with one seta, endopod with three setae at most; when palp one-segmented and longer than wide, then with two basal and two endopodal setae at most, and exopod (when present) represented by single seta; when palp one-segmented very small and wider than long, then basis and endopod not discernible, with one or two (most probably endopodal) setae, with or without surface (most probably exopodal) seta on coxa. Maxillule with endopod and exopod incorporated to basis; praecoxal arthrite with ventral seta thick and strongly spinulose, or slender and pinnate or smooth. Maxilla with two syncoxal endites; endopod completely incorporated to allobasis. Maxilliped subchelate; syncoxa with one seta; basis unarmed; claw with accessory seta. P1 not prehensile, rarely prehensile; P1–P4 EXP three-segmented; female P1–P3 ENP two-segmented, P4 ENP two- or one-segmented; inner exopodal and endopodal setae with or without comb tip. Female P5 EXP and BENP fused, occasionally separated; both baseoendopods of P5 separated. Armature formulae as follows:
P1 | P2 | P3 | P4 | P5 | |
EXP | 0;1;0,1-2,2 | 0;1;1,2,2 | 0;1;1-2,2,2 | 0;0-1;0-2,2,2 | ♀3-5 |
♂3-4 | |||||
ENP | 0-1;1,1-2,1 | 0-1;0-2,1-2,0-1 | ♀0;0-2,1-2,0-1 | 0;0,2,0 | ♀5-6 (7*) |
♂dimorphic: | or | ♂3 | |||
0-1;iap;0,2,0 | 0,2,0 | ||||
or | |||||
0-1;iap,2,0 | |||||
or | |||||
0;iap,1,oap |
Female P6 with one or two setae. Caudal rami with six or seven setae; setae IV and V fused basally or separated.
Sexual dimorphism can be expressed in a) rostrum slenderer in the male, b) the male antennule (chirocer or subchirocer), c) basis of P1 (with or without inner distal process), d) outer spine on P2 ENP2 (thicker and/or shorter than in the female), e) P2 EXP and/or P3 and P4 EXP (segments longer than in the female, outer spines stronger than in the female, rami curved inwards,), f) shape of P2-P4 ENP (segments thicker than in female), g) P3 ENP (two- or three-segmented; when three-segmented then inner apophysis on second segment; when two-segmented then inner apophysis medially or subdistally on second segment, occasionally with additional outer apophysis; inner apophysis simple, without arrow-head tip, variable in length; with or without asprothekes on second segment), h) P5 (both legs fused medially or separated; EXP and BENP fused; both legs separated or fused to somite), i) P6 (composed of two lappets articulated to somite or asymmetrical in which case only one leg functional, the other fused to somite; unarmed or (occasionally) with one or two setae), j) caudal rami (longer than in the female).
Cletocamptus retrogressus Schmankevitsch, 1875 (type by original designation).
Cletocamptus affinis Kiefer, 1957, C. albuquerquensis (Herrick, 1894), C. assimilis Gomez & Gee, 2009, C. axi Mielke, 2000, C. cecsurirensis Gómez, Scheihing & Labarca, 2007, C. chappuisi Gómez, Gerber & Fuentes-Reinés, 2017, Cletocamptus confluens (Schmeil, 1894), C. deborahdexterae Gómez, Fleeger, Rocha-Olivares & Foltz, 2004, C. dominicanus Kiefer, 1934, C. feei (Shen, 1956), C. fourchensis Gómez, Fleeger, Rocha-Olivares & Foltz, 2004, C. goenchim Gómez, Ingole, Sawant & Singh, 2013, C. gomezi Suárez-Morales, Barrera-Moreno & Ciros-Pérez, 2013, C. gravihiatus (Shen & Sung, 1963), C. koreanus Chang, 2013, C. levis Gómez, 2005, C. mongolicus Stĕrba, 1968, C. nudus Gómez, 2005, C. pilosus Gomez & Gee, 2009, C. samariensis Fuentes-Reinés, Zoppi de Roa & Torres, 2015, C. schmidti Mielke, 2000, C. sinaloensis Gómez, Fleeger, Rocha-Olivares & Foltz, 2004, C. spinulosus Gomez & Gee, 2009, C. stimpsoni Gómez, Fleeger, Rocha-Olivares & Foltz, 2004, C. tainoi Gómez, Gerber & Fuentes-Reinés, 2017, C. tertius Gomez & Gee, 2009, C. trichotus Kiefer, 1929.
Species incertae sedis.
Marshia brevicaudata Herrick, 1894.
Cletocamptus bermudae Willey, 1930, C. cfr. bicolor sensu
C. deitersi: in
Canthocamptidae: Cletocamptinae. Body fusiform, without clear distinction between prosome and urosome. Without nuchal organs on cephalothorax or body somites. Female rostrum distinct, rarely fused to cephalothorax, large and triangular with broad proximal margin; ornamented with ventral (sub)distal spinules and with rounded tip in both sexes. Posterior margin of cephalothorax and/or prosomites with long and slender, or short spinules, or serrated; posterior margin of urosomites except for anal somite with short spinules or serrated; body somites without cuticular sensillum-bearing socles; anal operculum without dorsal ornamentation or with transverse row of strong, large, or short, small spinules; posterior margin unornamented, or with small or large spinules. Female genital somite and third urosomite separated dorsolaterally, completely fused ventrally forming genital double-somite. Female antennule six-, rarely seven-segmented (the latter reported for C. gravihiatus requires confirmation). Antenna with allobasis, with one or two abexopodal setae (proximal element basal, distal seta endopodal); exopod one-segmented and longer than wide, or minute, rarely represented by single seta (the latter reported for C. chappuisi requires confirmation). Mandibular palp one-segmented, rarely two-segmented (basis and endopod distinct as in C. retrogressus); when one-segmented, then very small and wider than long, or as long as wide (C. dominicanus), or longer than wide (C. confluens, C. gomezi); when palp one-segmented, then basis and endopod not discernible, rarely discernible (represented by single seta as in C. confluens and C. dominicanus); endopod with two setae, rarely with one single element (C. pilosus); exopod absent; with or without surface (most probably exopodal) seta on coxa, the latter present only in some species with a one-segmented palp wider than long, absent in species with palp as long as wide or longer than wide. Maxillule with endopod and exopod incorporated to basis; praecoxal arthrite with ventral seta thick and strongly spinulose, or slender and pinnate or smooth. Maxilla with two syncoxal endites; endopod completely incorporated to allobasis. Maxilliped subchelate; syncoxa with one seta; basis unarmed; claw with accessory seta. P1 ENP not prehensile; P1–P4 EXP three-segmented; female P1–P3 ENP two-segmented, P4 ENP two- or one-segmented; inner exopodal and endopodal setae with or without comb tip (see
P1 | P2 | P3 | P4 | P5 | |
EXP | 0;1;0,1-2,2 | 0;1;1,2,2 | 0;1;1-2,2,2 | 0;0-1;0-1,2,2 | ♀4-5 |
♂3-4 | |||||
ENP | 0-1;1,1,1 | 0;1-2,1-2,1 | ♀0;1-2,1-2,1 | 0;0,2,0 | ♀5-6 (7+) |
♂dimorphic | or | ♂3 | |||
0;iap;0,2,0 | 0,2,0*** | ||||
or | |||||
0;iap,2,0* | |||||
or | |||||
0;iap,1,oap** |
Female P6 with one or two setae. Caudal rami with six or seven setae; setae IV and V fused basally or separated.
Sexual dimorphism expressed in a) rostrum (slenderer than in female), b) the male antennule (subchirocer), c) basis of P1 (with inner distal process), d) outer spine on P2 ENP2 (thicker and/or shorter than in the female), e) P3 ENP (three-segmented as in most species, or two-segmented as in C. albuquerquensis, C. chappuisi, C. confluens, C. dominicanus, and C. tainoi; when three-segmented then inner apophysis on second segment; when two-segmented then inner apophysis medially as in C. dominicanus, or subdistally on second segment as in C. albuquerquensis, C. chappuisi, C. confluens, and C. tainoi, but occasionally with additional outer apophysis as in C. confluens), f) P5 (both legs fused medially as in most species, or separated as in C. axi, C. cecsurirensis, C. retrogressus, and C. schmidti; EXP and BENP fused, and both legs articulating with somite, g) P6 (composed of two lappets articulated to somite and unarmed as in most species or, occasionally, with one seta attributable to intraspecific variability as in C. axi, C. schmidti, C. sinaloensis). Additionally, sexual dimorphism can be expressed also (depending on the species) in a) P2 EXP and/or P3 and P4 EXP (segments longer than in the female, outer spines stronger than in the female, and/or rami curved inwards as in C. retrogressus, C. confluens, C. albuquerquensis, C. samariensis, C. spinulosus, C. tainoi, C. trichotus), b) shape of P2–P4 ENP (segments thicker than in female as in C. confluens), c) caudal rami (longer than in the female).
Amphibiperita neotropica Fiers & Rutledge, 1990 (type by original designation).
As in
Cletocamptoides merbokensis (Gee, 1999) comb. nov. (= Cletocamptus merbokensis Gee, 1999)
Cletocamptoides helobius (Fleeger, 1980) comb. nov. (= Cletocamptus helobius Fleeger, 1980), Cletocamptoides biushelo sp. nov.
Canthocamptidae. Body fusiform, without clear distinction between prosome and urosome; body with somitic constrictions between somites. Without nuchal organs on cephalothorax or body somites. Female rostrum distinct, large, and triangular with broad proximal margin; ornamented with ventral (sub)distal spinules or setules; with rounded or bilobed tip. Cephalothorax and/or prosomites with posterior margins serrated; posterior margin of urosomites except for anal somite serrated; posterior margin of cephalothorax and body somites without or with cuticular sensillum-bearing socles. Anal operculum without dorsal ornamentation or with transverse row of mall spinules; posterior margin unornamented or serrated. Female genital somite and third urosomite separated dorsolaterally, completely fused ventrally forming genital double-somite. Female antennule six-segmented. Antenna with allobasis, with one or two abexopodal setae (proximal element basal, distal seta endopodal); exopod one-segmented and minute, or absent. Mandibular palp one-segmented, longer than wide; endopod incorporated to basis, with two basal and two endopodal setae at most; exopod absent or represented by single seta. Maxillule with endopod and exopod incorporated to basis; praecoxal arthrite with ventral seta thick and strongly spinulose, or slender and pinnate. Maxilla with two syncoxal endites; endopod completely incorporated to allobasis. Maxilliped subchelate; syncoxa unarmed or with one seta; basis unarmed; claw with accessory seta. P1 ENP not prehensile; P1–P4 EXP three-segmented; female P1–P3 ENP two-segmented, P4 ENP one-segmented; inner exopodal and endopodal setae with or without comb tip. Female P5 EXP and BENP fused, separated by shallow or deep notch; both baseoendopods of P5 separated. Armature formulae as follows:
P1 | P2 | P3 | P4 | P5 | |
EXP | 0;1;0,2,2 | 0;1;1,2,2 | 0;1;1,2,2 | 0;0-1;0,2,2 | ♀3–4 |
♂3–4 | |||||
ENP | 0;1,1,1 | 0;0-1,1-2,0-1 | ♀0;0-1,1-2,0-1 | 0,2,0 | ♀5–6 |
♂dimorphic: | ♂3 | ||||
0;iap;0,2,0 | |||||
or | |||||
0;iap,2,0 |
Female P6 with two setae. Caudal rami slightly convergent; with six setae; setae IV and V separated; seta IV normal, whip-like, slightly tapering posteriad, or with bulbous base.
Sexual dimorphism can be expressed in a) the male antennule (subchirocer), b) setae on P2 ENP2 (spiniform as in C. merbokensis comb. nov.), c) P3 ENP (two- or three-segmented; when three-segmented then inner apophysis on second segment (C. merbokensis comb. nov.); when two-segmented then inner apophysis subdistally on second segment (C. helobius comb. nov.); inner apophysis simple, without arrow-head tip, variable in length), d) P5 (both legs fused medially (C. merbokensis comb. nov.) or separated (C. helobius comb. nov.); EXP and BENP fused; both legs separated (C. helobius comb. nov.) or fused to somite (C. merbokensis comb. nov.)), e) P6.
The Ancient Greek sufix εἶδος, eîdos, meaning likeness and refers to the resemblance of the new genus with Cletocamptus.
Urías estuary, Mazatlán, Sinaloa State, stn. 5 (23.2056°N, 106.3715°W; 0.6 m depth; organic carbon content 0.99%; organic matter content 1.71%; sand 78.61%; clay 6.72%; silt 14.67%) (see also
Urías estuary, Mazatlán, Sinaloa State, stn. 7 (23.2174°N, 106.3917°W; 3.7 m depth; organic carbon content 5.59%; organic matter content 9.62%; sand 10.78%; clay 37.54%; silt 51.68%) (see also
Female holotype dissected (ICML-EMUCOP-180119-179); 18 Jan. 2019; S. Gómez leg. One additional female from stn 7 (see above) was used for molecular analyses.
Female. Total body length of holotype 411 µm measured from anterior tip of rostrum to posterior margin of caudal rami. Habitus (Fig.
Antennule (Fig.
Antenna (Fig.
Mandible (Fig.
Maxillule (Fig.
Maxilla (Fig.
Maxilliped (Fig.
P1 (Fig.
P2–P3 (Figs
P4 (Fig.
P1 | P2 | P3 | P4 | P5 | |
Exopod | 0,1,022 | 0,1,122 | 0,1,122 | 0,1,022 | 3 |
Endopod | 0,210 | 0,111 | 0,111 | 020 | 5 |
P1–P5 armature formulae as follows:
P5 (Fig.
Genital field (Fig.
Male. Unknown.
The selected characters (Table
The heuristic search yielded nine equally most parsimonious trees of 220 steps, and consistency index of 0.39. The extended majority rule consensus tree is shown in Figure
The family Canthocamptidae was created and diagnosed by
In a preliminary note on the revision of the freshwater genus Canthocamptus,
In a preliminary note,
By the early 1950’s,
With 864 species distributed in 58 valid genera (WoRMS 2021), the Canthocamptidae is currently one of the most species-rich families within Harpacticoida. The Canthocamptidae is not a natural assemblage and could be divided into several families (
More recently,
The complexity of the Canthocamptidae is evident and it is due to bad and incomplete descriptions (
The high morphological diversity of the Canthocamptidae makes difficult the detection of (aut)apomorphies to substantiate the establishment and proposal of monophyletic taxa. In this sense, the role of weighted characters for the maximum parsimony approach was essential and allowed high support. The genus Cletocamptus as known before the present study, seems to be composed of two different lineages, the merbokensis-helobius-biushelo-lineage (mhb-lineage; for C. merbokensis, C. helobius, and C. biushelo sp. nov.) and the retrogressus-lineage (for the rest of the species of Cletocamptus). The split of Cletocamptus into two genera is proposed in this study: Cletocamptoides gen. nov. for the mhb-lineage, and Cletocamptus for the retrogressus-lineage.
The reconstruction of the hypothetical ancestor of the Cletocamptinae subfam. nov. is necessary to justify the (aut)(syn)apomorphies for the three genera within the new subfamily. The hypothetical ancestor was reconstructed based on plesiomorphic and (aut)apomorphic states of a selected set of characters, as essential to explain our hypotheses and phylogenetic reasoning.
Most canthocamptids possess a body without somitic constrictions between somites, and few species, e.g., C. merbokensis comb. nov., C. helobius comb. nov., C. biushelo sp. nov., possess a body with somitic constrictions between somites conferring the body a cletodid appearance. The latter is regarded here as the derived (synapomorphic) condition for the mhb-lineage. The ancestral Cletocamptinae subfam. nov. is hypothesized to lack somitic constrictions between somites, tapering gradually posteriad. The latter is considered as the primitive condition and is present in the retrogressus-lineage.
The posterior margin of body somites of most genera of the Canthocamptidae is plain, without any ornamentation, including one species of Cletocamptinae subfam. nov., C. chappuisi, but its condition requires confirmation; it is finely or deeply serrated in some genera, e.g., Antrocamptus Chappuis, 1957, Attheyella, Canthocamptus, Elaphoidella, Moraria, Paramoriaropsis Brancelj, 1991, Spelaeocamptus, and C. dominicanus; it is ornamented with short spinules, e.g., Attheyella, and C. albuquerquensis and C. tainoi, or with long slender spinules, e.g., Cletocamptus as defined here. It is assumed that the plain condition of the posterior margin of body somites is the most primitive within the Canthocamptidae. Species of some genera developed serrate posterior margins that eventually developed into short spinules. The ancestral Cletocamptinae subfam. nov. is hypothesized to have retained the finely serrate posterior margins present in Cletocamptoides gen. nov., and displayed by other canthocamptids (see above) The presence of long slender spinules in the posterior margin of body somites of Cletocamptus as defined here, is assumed to be the derived condition.
Posterior cuticular sensillum-bearing socles have been observed only in C. merbokensis comb. nov., for which this condition is regarded as apomorphic. All other canthocamptids lack cuticular sensillum-bearing socles and is regarded here as a clear plesiomorphy and is the hypothesized condition in the ancestral Cletocamptinae subfam. nov.
The rostrum is separated from the cephalothorax in species of Bryocamptus, Gulcamptus Miura, 1969, Itunella, Mesochra, Moraria, Pordfus Özdikmen, 2008, Boreolimella, Carolinicola, Dahlakocamptus, Heteropsyllus, Perucamptus, Isthmiocaris, and Psammocamptus. It is fused and sometimes reduced in the remaining genera. The well-developed rostrum separated from the cephalothorax is regarded here as the primitive condition for the family, and is assumed to have been retained in the ancestral Cletocamptinae subfam. nov. The condition of the rostrum of C. gomezi, which has been described as fused to the cephalothorax (
The rostrum lacks any spinular ornamentation in the Canthocamptidae; as far as we know, ventral subdistal spinular ornamentation is present only in the Cletocamptinae subfam. nov. for which is a clear autapomorphy and justifies the new subfamily.
As far as we are aware (we could not review all the descriptions for all the species of Canthocamptidae though), the male rostrum is non-dimorphic in most canthocamptids (e.g., Canthocamptinae, Hemimesochrinae and Epactophaninae), and is regarded here as the primitive condition, which was retained in the ancestral Cletocamptinae subfam. nov. The male rostrum is sexually dimorphic in Cletocamptus and Amphibiperita for which is regarded here as synapomorphic.(it is also sexually dimorphic in Taurocletodes tumenae Karaytuğ & Huys, 2004 (the male of T. dubius (Noodt, 1958) remains unknown), but the significance of its presence in Cletocamptus and T. tumenae is uncertain). Cletocamptoides gen. nov. retained the primitive non-dimorphic male rostrum.
The canthocamptid female antennule can be eight- to five-segmented. Following
The canthocamptid antenna has undergone fusion of the basis and the first endopodal segment to form an allobasis. The allobasis of canthocamptids is unarmed or equipped with one or two (one basal, one endopodal) abexopodal setae. The latter is regarded here as the most primitive condition of the allobasis. Two abexopodal setae are retained in most species of Cletocamptinae subfam. nov., but some species (C. gomezi, C. feei, C. retrogressus, C. merbokensis comb. nov., C. helobius comb. nov., A. neotropica) underwent loss of one seta, most probably the endopodal element. The antennary exopod of canthocamptids is mostly one-segmented, but the two-segmented condition is present in species of Bryocamptus, Canthocamptus, Gulcamptus, Maraenobiotus, Mesochra, and Heteropsyllus; it is represented by one seta in species of Stenocaris and is absent in Dahmsopottekina Özdikmen, 2009. The antennary exopod of the Cletocamptinae subfam. nov. is elongate and one-segmented except for C. feei for which the exopod was reported as two-segmented (the latter requires confirmation though), C. chappuisi whose antennary exopod was reported as represented by one seta as in C. helobius comb. nov. and C. biushelo sp. nov., and Amphibiperita in which the antennary exopod is lost. The antennary exopod of C. dominicanus is a minute segment with a single seta. The ancestral antenna of Cletocamptinae subfam. nov. is assumed to possess an allobasis with two abexopodal setae, and a one-segmented exopod; the free endopodal lobe possessed two spines and a slender seta laterally, and three spines and two geniculate setae distally.
The mandibular palp of Podogennonta is composed of a basis with four setae; a one-segmented endopod with three proximal lateral, three subdistal lateral, and three sets of setae fused basally with three, two, and two setae, respectively; and a four-segmented exopod with armature formula 2, 1, 1, 2 (
The maxillulary basis, exopod and endopod are fused into a single unit, making difficult the homologation and correct counting of their setae. A careful and more detailed study of this appendage is pending, but it seems that the coxal endite bears two setae, and the basis, exopod and endopod are represented by three setae each, and this is considered as the ancestral state for the maxillule. On the other hand, a very strong and spinulose ventral element is present in several species of Cletocamptus, e.g., C. assimilis, C. axi, C. cecsurirensis, C. deborahdexterae, C. fourchensis, C. levis, C nudus, C. samariensis, C. schmidti, C., and C. sinaloensis, but also in C. helobius comb. nov., and C. biushelo sp. nov., and seems to have appeared independently in different lineages; the ventral seta on the praecoxal arthrite in the other species is visibly slenderer and is regarded here as the plesiomorphic condition found in the ancestral Cletocamptinae subfam. nov. The condition of C. gomezi is not conclusive and requires confirmation.
The architecture of the maxilla and maxilliped is rather constant within the new subfamily. The ancestral maxilla is assumed to bear two syncoxal endites; the endopod is assumed to be completely absorbed into the allobasis and represented by three setae. The ancestral maxilliped is subchelate; the syncoxa bears one seta; the basis is unarmed; the endopod is drawn out into a claw with one accompanying seta.
P1 | P2 | P3 | P4 | P5 | |
EXP | 0;1;0,2,2 | 0;1;1,2,2 | 0;1;2,2,2 | 0;1;2,2,2 | ♀5 |
♂4 | |||||
ENP | 0-1;1,2,1 | 0-1;2,2,1 | ♀0;2,2,1 | 0;0,2,0 | ♀6 |
♂dimorphic: | ♂3 | ||||
0-1;iap;0,2,0 |
The hypothetical ancestral segmentation and armature formulae of P1–P5 is:
The inner setae of P1–P4 EXP are assumed to be whip-like in the ancestral Cletocamptinae subfam. nov., i.e., devoid of a comb tip. See
The pair of P5 in the females are separated (baseoendopods not fused medially) in all species of Cletocamptinae subfam. nov. The female P5 EXP and BENP are fused in all species of Cletocamptinae subfam. nov. except for A. neotropica in which the female P5 EXP and BENP are separated. The ancestral pair of the female P5 are assumed to be separated; the exopod and the baseoendopod of each leg are assumed to be separated; the exopod is assumed to bear five setae as in most species of the new subfamily (four setae are present only in A. neotropica, C. samariensis, C. confluens, and C. merbokensis comb. nov.; three setae are present only in the new species described herein and in C. helobius comb. nov.), and the endopodal lobe is equipped with six setae as in most species of the new subfamily (seven setae have been reported for C. feei but this requires confirmation; five setae are present in C. biushelo sp. nov. and C. helobius comb. nov.). The ancestral pair of the male P5 of Cletocamptinae subfam. nov., are assumed to be fused medially as in most species of the new subfamily (the condition of C. axi, C. cecsurirensis, C. retrogressus, and C. schmidti is not conclusive), and the exopod and baseoendopod are assumed to be fused but separated by a deep notch as in most species for which the males are known (e.g., C. deborahdexterae, C. stimpsoni, and C. sinaloensis); the endopodal lobe is hypothesized to be armed with three setae as in the males of all the species of Cletocamptinae subfam. nov. for which the males are known; the ancestral exopod is assumed to bear four elements (three elements have been reported only for A. neotropica, C. confluens, C. koreanus, C. samariensis, and C. helobius comb. nov.).
The female P6 of the ancestral Cletocamptinae subfam. nov. is assumed to bear two setae; the male P6 is assumed to be represented by two articulate unarmed flaps.
The ancestral caudal rami are assumed here to be longer than broad as in some species of Cletocamptus, and equipped with seven setae.
The genus Cletocamptus was erected by
The ancestral Cletocamptinae subfam. nov. is hypothesized to lack somitic constrictions between somites, tapering gradually posteriad. Cletocamptus retained the plesiomorphic lack of such somitic constrictions.
Cletocamptoides gen. nov. displays the ancestral shape of somites with finely serrated posterior margins (see below), and the long slender spinules displayed by most species of Cletocamptus are assumed here to be the apomorphic condition for the posterior margin of body somites. Also, Cletocamptus retained the plesiomorphic lack of posterior cuticular sensillum-bearing socles on body somites.
Amongst the species of Cletocamptus, only C. gomezi has been described with the rostrum fused to the cephalothorax (
Cletocamptus retained the six-segmented female antennule hypothesized for the ancestral Cletocamptinae subfam. nov. The seven-segmented female antennule of Cletocamptus gravihiatus awaits verification.
Cletocamptus retained the ancestral allobasis of the antenna as well as the two abexopodal setae, and only C. gomezi, C. feei, and C. retrogressus lost one of these setae (one seta only is present also in C. merbokensis comb. nov., C. helobius comb. nov., and A. neotropica). Cletocamptus also retained the plesiomorphic one-segmented elongate antennary exopod (the two-segmented condition of the antennary exopod of C. feei requires to be confirmed) and the architecture of the free endopodal segment as described above for the ancestral Cletocamptinae subfam. nov.
The architecture of the mandibular palp of Cletocamptus is variable and underwent considerable reduction as has been detailed above. Briefly, the mandibular palp of the ancestral Cletocamptus might have been two-segmented, i.e., basis and endopod separated, being the basis and endopod equipped with one and two setae respectively, and the exopod represented by one seta (Fig.
The maxillule is probably the most complicated buccal appendage in Harpacticoida due to the fusion of the basis, endopod and exopod in many taxa, which makes difficult the homologation and correct counting of the setal elements. Pending a more detailed and precise study of this appendage in Cletocamptus, the coxal endite possesses two setae, and the basis, the endopod and exopod are fused into one single unit, with three setae representing each of them. On the other hand, a core of species, C. assimilis, C. axi, C. cecsurirensis, C. deborahdexterae, C. fourchensis, C. levis, C. nudus, C. samariensis, C. schmidti, C., and C. sinaloensis, share the presence of a very strong spinulose ventral seta on the praecoxal arthrite, which is regarded here as derived within the genus (the condition in C. gomezi is inconclusive; a similar seta is present in some species of Cletocamptoides gen. nov.). On the contrary, a visibly slenderer seta in the other species of the genus is regarded here as the plesiomorphic condition (a similar seta is present in C. merbokensis comb. nov.).
The maxilla and maxilliped retained the plesiomorphic architecture of the Cletocamptinae subfam. nov. as described above.
The P1 ENP of Cletocamptus is two-segmented and non-prehensile. As said above, this is the derived condition within Canthocamptidae as opposed to the plesiomorphic prehensile endopod (see
Sexual dimorphism in Cletocamptus is expressed in a variety of modifications. These have been outlined above. The most common modifications in the appendages of the males include the subchirocer antennule, the basis of P1 with one inner distal process, the inner spine on P2 ENP2 with several degrees of modifications, the three-segmented P3 ENP with an inner apophysis on second segment and two apical setae on the distal segment, and the fused exopod and baseoendopod of P5. Deviations occur in some species though. The shape of the outer spine on P2 ENP2 varies from slightly modified as in C. albuquerquensis, C. cecsurirensis, C. confluens, C. dominicanus, C. retrogressus, C. samariensis, and C. tainoi, to moderately modified as in C. axi, C. deborahdexterae, C. fourchensis, C. goenchim, C. gomezi, C. levis, C. schmidti, C. spinulosus, C. stimpsoni, and C. tertius, and strongly modified into a recurved spine as in C. pilosus, and C. sinaloensis. The inner spine on the male P2 ENP2 is not dimorphic in A. neotropica and Cletocamptoides gen. nov., and the sexually modified inner spine on P2 ENP2 is regarded here as an apomorphy for Cletocamptus. The male P3 ENP in most species retained the three-segmented condition with an inner apophysis on second segment which is regarded here as plesiomorphic, but the second and third segments underwent fusion leading a two-segmented ramus with the inner apophysis well-developed and situated medially on second segment as in C. dominicanus, or very short and recurved, and situated subdistally on second segment as in C. albuquerquensis, C. chappuisi, C. confluens, and C. tainoi. The former division between the second and third segments of the P3 ENP is still visible in C. dominicanus, which also possesses very long asprothekes (see
The genus Amphibiperita has been diagnosed by
The ancestral Cletocamptinae subfam. nov. lack somitic constrictions between somites (see above). The body shape of Cletocamptoides gen. nov. is cletodid-like, i.e., body somites clearly separated by somitic constrictions, and is a probable apomorphy for the new genus. On the other hand, C. merbokensis comb. nov. is the only species with integumental sensillum-bearing socles along the posterior margin of pro- and urosomites (except anal somite). However, the presence of integumental sensillum-bearing socles are deemed to be apomorphic for C. merbokensis comb. nov., and do not seem to indicate a close relationship with the Cletodidae.
The posterior margin of body somites of most canthocamptid genera can be plain, finely or deeply serrate, or with long slender spinules (see above). The posterior margin of the body somites of Cletocamptoides gen. nov. is finely serrate and is regarded here as plesiomorphic within the Cletocamptinae subfam. nov.
The condition of the antennary exopod and the mandibular palp of C. merbokensis comb. nov. are the most primitive within Cletocamptoides gen. nov. The antennary exopod of C. merbokensis comb. nov. is one-segmented and elongate, and bears three setae (one lateral, two distal); the antennary exopod of C. helobius comb. nov. and C. biushelo sp. nov. is reduced to one minute segment bearing only one seta. Similar antennary exopods are also found in Cletodidae.
The condition of the mandibular palp is more primitive in C. merbokensis comb. nov. than in the other two species of the genus. The mandibular palp of C. merbokensis comb. nov. possesses five setae; three setae are present in C. helobius comb. nov., and C. biushelo sp. nov. Besides the supernumerary setal complement in C. merbokensis comb. nov. relative to the other two species of the new genus, the basis and exopod are still discernible in the former species, being the inner – basal – extension armed with two setae; the basis of the mandibular palp of C. helobius comb. nov. and C. biushelo sp. nov. is completely absorbed into the palp but is still discernible by the presence of one inner seta. The exopod of the mandibular palp of C. merbokensis comb. nov. is indicated by a small outer protuberance with one seta; the exopod in C. helobius comb. nov. and C. biushelo sp. nov. underwent complete reduction and is not discernible. The two apical setae on the mandibular palp of these three species are regarded here as endopodal, and judging by their relative lengths, being the inner seta visibly shorter than the outer, both setae are homologous in these three species and in those species with similar armature (see above). The one-segmented condition of the mandibular palp is the most common in the Cletodidae, but the basal, endopodal and/or exopodal setae are still discernible in some species (e.g., Paracrenhydrosoma normani Gee, 1999b); the basis and endopod are separated, and the exopod is absorbed into the basis and is represented by a seta in Pyrocletodes Coull, 1973 (e.g., P. coulli Dinet, 1976); uniramous mandibular palps (basis and endopod separated) without any trace of the exopod are present also in some species of Paracrenhydrosoma (e.g., P. cornuta Kornev & Chertoprud, 2008, P. kiai Song, Dahms, Lee, Ryu & Khim, 2014), and in the monotypic genus Nannopodella Monard, 1928.
The unisetose condition of the maxillipedal syncoxa is present in most Cletocamptinae subfam. nov., and it is unarmed only in C. merbokensis comb. nov. and C. helobius comb. nov. for which is regarded here as apomorphic. Unarmed maxillipedal syncoxae are also present in species of some other canthocamptids e.g., Attheyella, Lessinocamptus, Paramoriaropsis Brancelj, 1991, Pordfus Özdikmen, 2008, and Termomesochra Itô & Burton, 1980 (Canthocamptinae), and Stygepactophanes and Epactophanes (Epactophaninae); the maxillipedal syncoxa is unisetose in the Cletodidae (
The P1 ENP of Cletocamptoides gen. nov. displays the derived two-segmented condition (see also
One-segmented P4 endopods are present in species of some other canthocamptids, e.g., Antrocamptus Chappuis, 1957, Australocamptus Karanovic, 2004, Gulcamptus Miura, 1969, Hypocamptus, Lessinocamptus Stoch, 1997, Morariopsis, and Elaphoidella (Canthocamptinae), Itunella and Isthmiocaris (Hemimesochrinae), and Stygepactophanes Moeschler & Rouch, 1984 and Epactophanes (Epactophaninae), but as far as we know from the literature available, it is an elongate and well-developed ramus. Within the new subfamily, a one-segmented P4 ENP is present also in C. dominicanus, but it is elongated. The P4 ENP of Cletocamptoides gen. nov. is one-segmented but it has undergone extreme reduction and is an apomorphy, and probable autapomorphy, detected for the new genus.
Both rami of P5 are fused and separated from the supporting somite in the females and males of the genera attributed to the Cletocamptinae subfam. nov., except for the female P5 of Amphibiperita whose female P5 EXP is separated from the baseoendopod. Both P5 are fused in the males, except for some species of Cletocamptus (e.g., C. axi, C. cecsurirensis, C. retrogressus and C. schmidti), and C. helobius comb. nov. The male of C. merbokensis comb. nov. displays the more derived condition of P5, being completely absorbed into the somite, and is considered a potential apomorphy for the species. The male and female P5 of C. helobius comb. nov. and C. biushelo sp. nov. resembles those of C. dominicanus, C. albuquerquensis, C. confluens, C. tainoi, and most probably C. chappuisi, in that both rami are hardly discernible and are separated only by a superficial shallow notch. A similar P5 is present also in Carolinicola Huys & Thistle, 1989 (Hemimesochrinae). The structure of the cletodid P5 is variable; the endopodal lobe and the basis can be fused or separated, and the exopod can be fused to or separated from the baseoendopod (
The Mesochra group as defined by
Thanks go to Abraham Guerrero Ruíz (Centro de Investigación en Alimentación y Desarrollo, Unidad Mazatlán) and Sergio Rendón Rodríguez (Instituto de Ciencias del Mar y Limnología, Unidad Académica Mazatlán) for their help during the sampling campaigns, and to Nataly Ortíz Gálvez and Ángel A. Valenzuela Cruz for their help during the sampling campaigns and for processing the sediment samples. We are grateful to Paulo H. C. Corgosinho, Süphan Karaytuğ for their constructive criticism and comments to the first draft of the manuscript. This is a contribution to project IN202019 Biodiversidad de la meiofauna en un ecosistema costero contaminado del sur de Sinaloa: un enfoque integrativo de técnicas taxonómicas clásicas y moleculares financed by the Programa de Apoyo a Proyectos de Investigación e Innovación Tecnológica (DGAPA-PAPIIT) of the Universidad Nacional Autónoma de México.