Catalogue |
Corresponding author: Gilmar Perbiche-Neves ( gilmarperbiche83@gmail.com ) Academic editor: Danielle Defaye
© 2015 Gilmar Perbiche-Neves, Geoffrey Allan Boxshall, Daniel Previattelli, Marcos Gomes Nogueira, Carlos Eduardo Falavigna Da Rocha.
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:
Perbiche-Neves G, Boxshall GA, Previattelli D, Nogueira MG, da Rocha CEF (2015) Identification guide to some Diaptomid species (Crustacea, Copepoda, Calanoida, Diaptomidae) of “de la Plata” River Basin (South America). ZooKeys 497: 1-111. https://doi.org/10.3897/zookeys.497.8091
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An identification guide is presented for species of calanoid copepod family Diaptomidae from “de la Plata” River Basin (Argentina, Brazil, Bolivia, Paraguay and Uruguay). It was based on material collected during the summer and winter of 2010 from 43 sites across the eastern part and the lower stretches of this basin, the second largest in South America and the fourth in the world. The guide contains identification keys and species diagnoses for males and females, richly supported by scanning electronic micrographs and/or line drawings of 19 species. It also includes some general remarks on the taxonomy and phylogenetic relationships of these species. The key was adjusted to be useful for these species only, with separate keys for each sex, and is the first for females of South America. One species classified herein as incertae sedis was not included in the analysis. At least ten other species have previously been recorded in the basin but were not present in our samples. This is the first attempt to compile comprehensive taxonomic information on this group of copepods in this region, and it is expected to become a useful tool for biologists and young taxonomists interested in the crustacean biota of the Neotropical region.
Copepoda, plankton, taxonomy, rivers, reservoirs, Notodiaptomus, macro scale distributions
Copepods are typically small crustaceans that are widely distributed in virtually all aquatic habitats from hot springs to glacial meltwater pools, and from deep ocean trenches to high altitude lakes (
In continental or inland waters, copepods are predominantly represented by three orders: Calanoida, Cyclopoida, and Harpacticoida. In general terms, members of the Calanoida tend to be planktonic forms whereas the Cyclopoida are represented by planktonic as well as littoral, epibenthic and benthic species. Harpacticoids are essentially benthic and interstitial. We use the term planktonic to include the potamoplankton, referring to organisms found in rivers.
All the calanoids collected during our study belong to the subfamily Diaptominae, the most diverse taxon within the family Diaptomidae (
In South America, copepods of continental waters began to be studied from the 1890s, and several major works appeared in the 1920s and 1930s (e.g.
Although some of the studies were made many years ago, we found that the following papers, used jointly, are quite useful for identifying diaptomids in the region:
Net zooplankton samples were taken across “de la Plata” river basin in lotic stretches and in reservoirs (upstream to dam – intermediate zone –, and close to dam – lentic zone) (Figure
List of sampling sites. For each site a code is provided based on a 3 or 4 letter abbreviation of the name of the reservoir or river, and a 1 letter suffix. The suffixes used for reservoirs are: U = upstream and D = dam, and for rivers: H = high stretch, M = middle stretch, L = low stretch). Country abbreviations are: ARG = Argentina, BOL = Bolivia, BRA = Brazil, PAR = Paraguay, and URU = Uruguay. Geographical coordinates and altitude (m.a.s.l.) are also given.
N° | Codes | River/Reservoir | Coordinates | Alt. (m) |
---|---|---|---|---|
1 | EMB-U | Emborcação Reservoir – BRA | 18°22'40.47"S, 47°44'3.58"W | 634 |
2 | EMB-D | Emborcação Reservoir – BRA | 18°29'33.09"S, 47°58'17.22"W | 648 |
3 | SSIM-U | São Simão Reservoir – BRA | 18°40'22.54"S, 50° 4'17.76"W | 402 |
4 | SSIM-D | São Simão Reservoir – BRA | 18°59'15.59"S, 50°30'18.93"W | 406 |
5 | FUR-U | Furnas Reservoir – BRA | 20°58'35.58"S, 45°31'24.18"W | 771 |
6 | FUR-D | Furnas Reservoir – BRA | 20°39'36.51"S, 46°18'12.16"W | 769 |
7 | AVER-U | Água Vermelha Reservoir – BRA | 19°55'42.17"S, 49°45'5.31"W | 388 |
8 | AVER-D | Água Vermelha Reservoir – BRA | 19°52'03.73"S, 50°19'28.77"W | 388 |
9 | BBON-U | Barra Bonita Reservoir – BRA | 22°40'04.08"S, 48°21'05.01"W | 463 |
10 | BBON-D | Barra Bonita Reservoir – BRA | 22°31'43.07"S, 48°31'26.05"W | 454 |
11 | TIRM-U | Três Irmãos Reservoir – BRA | 20°57'21.57"S, 50°36'34.83"W | 320 |
12 | TIRM-D | Três Irmãos Reservoir – BRA | 20°41'57.09"S, 51°05'58.43"W | 326 |
13 | JUR-U | Jurumirim Reservoir – BRA | 23°19'25.07"S, 48°42'11.07"W | 572 |
14 | JUR-D | Jurumirim Reservoir – BRA | 23°13'41.07"S, 49°13'28.03"W | 566 |
15 | ROS-U | Rosana Reservoir – BRA | 22°36'28.27"S, 52°09'43.75"W | 262 |
16 | ROS-D | Rosana Reservoir – BRA | 22°36'04.71"S, 52°49'48.15"W | 261 |
17 | FAR-U | Foz do Areia Reservoir – BRA | 26°03'41.64"S, 51°24'02.25"W | 754 |
18 | FAR-D | Foz do Areia Reservoir – BRA | 25°59'57.06"S, 51°38'52.27"W | 749 |
19 | SCAX-U | Salto Caxias Reservoir – BRA | 25°30'32.11"S, 53°18'24.26"W | 333 |
20 | SCAX-D | Salto Caxias Reservoir – BRA | 25°31'50.96"S, 53°28'45.76"W | 319 |
21 | ISOL-U | Ilha Solteira Reservoir – BRA | 20°10'29.60"S, 51° 2'7.06"W | 332 |
22 | ISOL-D | Ilha Solteira Reservoir – BRA | 20°22'10.87"S, 51°20'37.65"W | 321 |
23 | ITA-U | Itaipu Reservoir – BRA/PAR | 24°29'10.77"S, 54°19'42.38"W | 217 |
24 | ITA-D | Itaipu Reservoir – BRA/PAR | 25°25'09.67"S, 54°32'14.47"W | 220 |
25 | YACI-U | Yaciretá Reservoir – ARG/PAR | 27°24'24.13"S, 56°15'19.86"W | 71 |
26 | YACI-D | Yaciretá Reservoir – ARG/PAR | 27°30'9.12"S, 56°31'56.69"W | 78 |
27 | RPAR-M1 | Paraná River – middle stretch – ARG | 28°30'10.12"S, 59°03'03.24"W | 43 |
28 | RPAR-M2 | Paraná River – middle stretch – ARG | 30°01'07.73"S, 59°33'50.86"W | 26 |
29 | RPAR-M3 | Paraná River – middle stretch – ARG | 31°38'29.94"S, 60°23'21.53"W | 21 |
30 | RPAR-L1 | Paraná River – low stretch – ARG | 32°44'02.61"S, 60°43'23.95"W | 5 |
31 | RPAR-L2 | Paraná River – low stretch – ARG | 33°41'19.94"S, 59°37'30.79"W | 5 |
32 | RPAR-L3 | Paraná River – low stretch – ARG | 33°56'31.07"S, 58°27'46.80"W | 5 |
33 | RPLA | Rio de la Plata – URU/ARG | 34°26'49.57"S, 57°36'27.99"W | 1 |
34 | RPAG-H | Paraguay River – high stretch – BOL/BRA | 18°58'07.73"S, 57°38'55.75"W | 94 |
35 | RPAG-M | Paraguay River – middle stretch – PAR/BRA | 21°41'09.18"S, 57°52'59.85"W | 71 |
36 | RPAG-L | Paraguay River – low stretch – ARG/PAR | 26°51'15.42"S, 58°19'21.45"W | 54 |
37 | MAC-U | Machadinho Reservoir – BRA | 27°32'26.71"S, 51°37'52.31"W | 476 |
38 | MAC-D | Machadinho Reservoir – BRA | 27°29'27.77"S, 51°46'26.50"W | 484 |
39 | RURU-M1 | Uruguay – middle stretch – ARG/BRA | 27°17'15.23"S, 54°11'31.66"W | 112 |
40 | RURU-M2 | Uruguay – middle stretch – ARG/BRA | 28°32'38.40"S, 56° 1'24.69"W | 54 |
41 | SGRA-U | Salto Grande Reservoir – URU/ARG | 30°46'27.52"S, 57°47'55.53"W | 33 |
42 | SGRA-D | Salto Grande Reservoir – URU/ARG | 31°15'31.41"S, 57°55'33.66"W | 34 |
43 | RURU-L | Uruguay River – low stretch – URU/ARG | 33°48'07.39"S, 58°26'07.48"W | 8 |
The samples were collected by vertical hauls with a conical plankton net with a 68 µm mesh, equipped with an anti-reflux adaptation following
Back in the laboratory, the copepods were sorted and analysed under a stereo- microscope (Zeiss Stemi SV6 and Zeiss Discovery V20), or a binocular microscope (Zeiss Standard 20 and 25). Copepods were dissected with fine needles and semi-permanent slides were made in glycerine or in 70% lactophenol. Only adults were analysed.
Copepods were identified with the aid of a number of publications dealing with the taxonomy of Diaptomidae (
All original zooplankton samples are deposited in the Continental Water Microcrustacean Collection–CMAC, Department of Zoology, University of the State of São Paulo–UNESP, Botucatu campus. Vouchers of some species were deposited in the Museum of Zoology, University of São Paulo–MZUSP (São Paulo, Brazil), with registration numbers as follows: Argyrodiaptomus azevedoi (MZUSP 32928); Argyrodiaptomus falcifer (MZUSP 32929); Argyrodiaptomus denticulatus (MZUSP28393); Notodiaptomus coniferoides (MZUSP28389); Notodiaptomus carteri (MZUSP28390); Notodiaptomus santafesinus (MZUSP28391); Odontodiaptomus thomseni (MZUSP28392); Notodiaptomus anisitsi (MZUSP 32930); Notodiaptomus cearensis (MZUSP 32931); Notodiaptomus conifer (MZUSP 32932); Notodiaptomus dentatus (MZUSP 32933); Notodiaptomus henseni (MZUSP 32934); Notodiaptomus iheringi (MZUSP 32935); Notodiaptomus incompositus (MZUSP 32936), and Notodiaptomus spiniger (MZUSP 32937).
Vouchers of the species “Diaptomus” curvatus and “Diaptomus” frutosae were deposited in other collections including the National Institute of Amazonian Research–INPA (Manaus, Brazil), the Bernardino Rivadalvia Museum (Argentina), and the Natural History Museum (UK) (
The keys presented in this guide were constructed using only differential characters that emerged as being consistently useful for the identification of genera and species. Similarly, the diagnoses were designed to focus on the basic features traditionally used to characterize diaptomid genera and species, as provided in the specialized taxonomic literature. The morphological structures used in this study are illustrated for both males (Figure
Key to the morphological structures of adult male diaptomids. A antero-lateral view of posterior end of prosome, urosome and caudal ramus from left side showing with left leg 5 (P5L) attached, illustrating characters A1 to A23 (Table
Key to the morphological structures of adult female diaptomids. A Caudal view of left fifth leg (P5L), illustrating characters A1–12 (Table
Key to male characters numbered in Figure
N | Structure name | Abbrev. | N | Structure name | Abbrev. |
---|---|---|---|---|---|
A | Distal prosome, urosome and P5 | C | Right geniculate antennule | A1R | |
1 | Pediger 5 | Ped5 | 1 | First segment of A1R | |
2 | Urosome somite 1 | Ur1 | 2 | Second segment of A1R | |
3 | Urosome somite 2 | Ur2 | 3 | Thirteenth segment of A1R | |
4 | Urosome somite 3 | Ur3 | 4 | Modified seta on segment 13 of A1R | |
5 | Urosome somite 4 | Ur4 | 5 | Fourteenth segment of A1R | |
6 | Caudal rami | CR | 6 | Spinous process on segment 14 of A1R | |
7 | Caudal setae | CS | 7 | Fifteenth segment of A1R | |
8 | Coxa of left P5 | CxP5L | 8 | Spinous process on segment 15 of A1R | |
9 | Sensilla of basipodite of left P5 | 9 | Sixteenth segment of A1R | ||
10 | Basipodite of left P5 | BspP5L | 10 | Twentieth segment of A1R | |
11 | Setae of basipodite of left P5 | 11 | Spinous process on segment 20 of A1R | ||
12 | Endopodite of left P5 | EnpP5L | D | Fifth leg anterior(P5) | |
13 | Apical setation of endopodite of left P5 | 1 | Coxa of right P5 | CxP5R | |
14 | First segment of exopodite of left P5 | Exp1P5L | 2 | Basipodite of right P5 | BspP5R |
15 | Second segment of exopodite of left P5 | Exp2PrL | 3 | Knob processes on internal margin of BspP5R | |
16 | Coxa of right P5 | CxP5R | 4 | Endopodite of right P5 | EnpP5R |
17 | Basipodite of right P5 | BspP5R | 5 | Coxa of left P5 | CxP5L |
18 | Lateral seta of basipodite of right P5 | 6 | Basipodite of left P5 | BspP5L | |
19 | Endopodite of right P5 | EnpP5R | 7 | Knob processes on internal margin of BspP5 | |
20 | First segment of exopodite of right P5 | Exp1P5R | 8 | Second segment of exopodite of left P5 | Exp2PrL |
21 | Second segment of exopodite of right P5 | Exp2P5R | E | Adult male | |
22 | Lateral spine of Exp2P5R | 1 | Left antennule | A1L | |
23 | Terminal claw of Exp2P5R | 2 | Right geniculate antennule | A1R | |
24 | Fifth leg | P5 | 3 | Spinous process on segment 20 of A1R | |
B | Right P5 | P5R | 4 | Pediger 3 | Ped3 |
1 | Coxa of right P5 | CxP5R | 5 | Pediger 4 | Ped4 |
2 | Distal projection of CxP5R | 6 | Genital segment or urosome segment 1 | GS or Ur1 | |
3 | Sensilla on top of distal projection of CxP5R | 7 | Urosome somite 2 | Ur2 | |
4 | Basipodite of right P5 | BspP5R | 8 | Rows of spinules on urosome somite 3 | |
5 | Lateral setae of BspP5R | 9 | Caudal rami | CR | |
6 | First segment of exopodite of right P5 | Exp1P5R | 10 | Caudal setae | CS |
7 | Distal projection of Exp1P5R | 11 | Right P5 | P5R | |
8 | Second segment of exopodite of right P5 | Exp2P5R | F | Border between pediger 3 and 4 | |
9 | Lateral spine of Exp2P5R | 1 | Row of spinules on distal margin of pediger 3 | ||
10 | Terminal claw of Exp2P5R |
Key to female characters numbered in Figure
N | Structure name | Abrev. | N | Structure name | Abrev. |
---|---|---|---|---|---|
A | P5 Left - caudal | D | Adult female | ||
1 | Coxa of left P5 | CxP5L | 1 | Left antennule | A1L |
2 | Basipodite of left P5 | BspP5L | 2 | Pediger 3 | Ped3 |
3 | Lateral setae of BspP5L | 3 | Pediger 4 | Ped4 | |
4 | Endopodite of left P5 | EnpP5L | 4 | Lateral wing of prosome pediger 5, with apical sensilla | |
5 | Terminal spinules on endopodite of left P5 | 5 | Expansions of genital double somite, with sensilla | ||
6 | First segment of exopodite of left P5 | Exp1P5L | 6 | Genital double somite | GS |
7 | Second segment of exopodite of left P5 | Exp2P5L | 7 | Caudal rami | CR |
8 | Lateral seta of Exp2P5L | 8 | Caudal setae | CS | |
9 | External seta of Exp3P5L | E | Distal part of prosome and genital segment | ||
10 | Internal seta of Exp3P5L | 1 | Prosome pediger 3 | Ped3 | |
11 | Terminal claw of left P5 | 2 | Lines of spinules on distal dorsal margin of Ped3 | ||
12 | Rows of spinules on terminal claw of left P5 | 3 | Prosome pediger 4 | Ped4 | |
B | P5 left – frontal | 4 | Rows of spinules on distal margin of Ped4 | ||
1 | Coxa of left P5 | CxP5L | 5 | Left lateral wing of pediger 5, with apical sensila | |
2 | Distal projection of CxP5L | 6 | Sensila of right lateral wing of pediger 5 | ||
3 | Sensilla on apex of distal projection of CxP5L | 7 | Left lateral wing of pediger 5 | ||
4 | Basipodite of right P5 | BspP5R | 8 | Genital double somite | GS |
5 | Lateral seta of BspP5L | 9 | Sensilla at apex of right lateral expansion of GS | ||
6 | Endopodite of left P5 | EnpP5L | 10 | Expansions of genital segment, with sensilla | |
7 | Concavity in endopodite of left P5 | F | Distal part of prosome and genital double somite | ||
8 | Terminal spinules on endopodite of left P5 | 1 | Prosome 5, with rows of spinules on distal margin | ||
9 | First segment of exopodite of left P5 | Exp1P5L | 2 | Dorsal/lateral setules on Ped4 and Ped5 | |
10 | Second segment of exopodite of left P5 | Exp2P5L | 3 | Incomplete suture between Ped4 and Ped5 | |
11 | Lateral seta of Exp2P5L | 4 | Left lateral wing of Ped5 | ||
12 | Third segment of exopodite of left P5 | Exp3P5L | 5 | Sensilla at apex of left lateral wing of Ped5 | |
13 | External seta of Exp3P5L | 6 | Right lateral wing of Ped5 | ||
14 | Internal seta of Exp3P5L | 7 | Sensilla at apex of right lateral wing of Ped5 | ||
15 | Terminal claw of left P5 | 8 | Genital double somite | GS | |
16 | Rows of spinules on terminal claw of left P5 | 9 | Sensilla on lateral margin expansion of GS | ||
C | Second exopodite and terminal claw | 10 | P5 | P5 | |
1 | Second segment of exopodite of right P5 | Exp2P5R | 11 | Right distal margin expansion of GS | |
2 | Lateral seta of Exp2P5R | ||||
3 | External seta of Exp3P5R | ||||
4 | Internal seta of Exp3P5R | ||||
5 | Terminal claw of left P5 | ||||
6 | Rows of spinules on terminal claw of left P5 |
Morphological structures were illustrated using phase contrast microscopy (Zeiss Standard) with the aid of a drawing tube. The pencil drawings were inked in nankeen ink, then scanned and corrected for smudges and other imperfections in Adobe Photoshop 7.0 in order to obtain high-quality illustrations.
We remind users of the keys that many structures illustrated in Figures
The scanning electron microscopy (SEM) was carried out in the “Electron Microscopy Center (CME)” of the University of the State of São Paulo–UNESP – Botucatu, Brazil. Material was prepared by packing each sample of individuals in hollow cylindrical polyethylene compartments, within which the copepods were washed, fixed and dehydrated. Washing was performed using 0.1 M phosphate buffer at pH 7.3 (3 washes, each for 5 min). After washing specimens were immediately fixed by immersion in 0.5% osmium tetroxide (in water) for 20 min. Dehydration was performed progressively via a graded series of ethanol as follows: 7.5%, 15%, 30% and 50% (two changes at each concentration for 5 min), then 70% (3 changes each for 10 min), and 90% and 100% (2 changes at each, for 5 min). Subsequently, the material was critical point dried in a BALZERS UNION CTD-020 equipment, using liquid carbon dioxide as the exchange medium. After drying, specimens were dissected when necessary to reveal diagnostic structures, and attached to stubs using adhesive tape. Sputter coating with 15 nm layer of gold was carried out in a BALZERS UNION MED-10 coater. Observations were made on a Philips SEM-515 microscope and images were edited in Photoshop 7.0 (Adobe).
1 | BspP5 with groups of small spinules on inner margin of both or one pediger | Argyrodiaptomus (5) |
– | BspP5 with smooth inner margin | 2 |
2 | CR with semi-circular protuberance at inner distal corner | Odontodiaptomus thomseni (Brehm, 1933) |
– | CR without semi-circular protuberance at inner distal corner | 3 |
3 | Right Exp2P5 subtriangular, 1.1–1.2 times wider than long (Fig. |
“Diaptomus” frutosae Perbiche-Neves & Boxshall, 2013 |
– | Right Exp2P5 >1.2 longer than wide | 4 |
4 | Right Exp2P5 with lateral spine inserted proximally on margin of segment | “Diaptomus” curvatus Perbiche-Neves, Boxshall & Paggi, 2013 |
– | Right Exp2p5 with lateral spine inserted in middle of segment or sub-terminally | Notodiaptomus (8) |
5 | Lateral spine of Exp2P5R inserted at about 1/3 distance between base of segment and terminal claw (Fig. |
Argyrodiaptomus azevedoi (Wright, 1935) |
– | Lateral spine of Exp2P5R inserted at more than 1/3 distance between base of segment and terminal claw | 6 |
6 | Insertion of lateral spine of Exp2P5R separated from base of terminal claw by gap greater than width of base of lateral spine (Fig. |
7 |
– | Lateral spine of right Exp2P5 inserted distally very close to base of terminal claw (Fig. |
Argyrodiaptomus furcatus (Sars, 1901) |
7 | Posterior margins of Ped 4 and Ped5 lacking ornamentation of spinules; A1R with spinous process on segment 14; internal margin of right BspP5 lacking proximal expansion | Argyrodiaptomus falcifer (Daday, 1905) |
– | Posterior margins of Ped4 and Ped5 ornamented with dorsal spinules; segment 14 of A1R lacking spinous process; internal margin of right BspP5 with proximal expansion reaching and overlapping left BspP5 | Argyrodiaptomus denticulatus (Pesta, 1927) |
8 | Lateral spine of right Exp2P5 less than 1/5 (20%) length of terminal claw (Fig. |
9 |
– | Lateral spine of right Exp2P5 more than 1/5 (20%) length of terminal claw | 13 |
9 | Lateral spine of right Exp2P5 inserted proximally, distant from base of terminal claw (e.g. Fig. |
10 |
– | Lateral spine of right Exp2P5 inserted distally, close to base of terminal claw (Fig. |
Notodiaptomus coniferoides (Wright, 1927) |
10 | First segment of A1R lacking rows of spinules | 11 |
– | First segment of A1R ornamented with rows of spinules | Notodiaptomus iheringi (Wright, 1935) |
11 | Spinous process on anterior margin of segment 15 of A1R short, not reaching proximal border of segment 16 (Fig. |
12 |
– | Spinous process on anterior margin of segment 15 of A1R long (reaching about midlength of segment 16) (Fig. |
Notodiaptomus conifer (Sars, 1901) |
12 | Segment 11 of left A1 lacking setae; small granulations absent from fissure in surface of right BspP5; internal margin of right BspP5 with proximal expansion reaching and overlapping left P5 | Notodiaptomus isabelae (Wright, 1936) |
– | Segment 11 of left A1 armed with two setae (Fig. |
Notodiaptomus cearensis (Wright, 1936) |
13 | Lateral spine of Exp2P5R short, between 1/5 and 1/3 the length of terminal claw (Fig. |
14 |
– | Lateral spine of Exp2P5R relatively long, more than 1/3 the length of terminal claw (Fig. |
16 |
14 | Proximal inner margins of left and right BspP5 lacking sclerotized semi-circular processes | 15 |
– | Proximal inner margin of BspP5 bearing one large sclerotized semi-circular process on right leg and two smaller processes on same segment of left leg (Figs |
Notodiaptomus henseni (Dahl, 1894) |
15 | Right Exp1P5 longer than wide; at least four small sclerotized processes present on posterior surface of right Exp2P5 (Fig. |
Notodiaptomus dentatus Paggi, 2001 |
– | Right Exp1P5 wider than long; no sclerotized processes present on posterior surface of right Exp2P5, but single inner margin process present (Fig. |
Notodiaptomus incompositus (Brian, 1926) |
16 | Lateral spine of right Exp2P5 located about at midlength of external margin (Fig. |
Notodiaptomus anisitsi (Daday, 1905) |
– | Lateral spine of right Exp2P5 not strongly curved, located on distal margin; segments 15 and 16 of A1R lacking well developed spinous processes; posterior margins of Ped4 and Ped5 lacking ornamentation of spinules | 17 |
17 | Lateral spine of right Exp2P5 inserted some distance from base of terminal claw (Fig. |
Notodiaptomus carteri (Lowndes, 1934) |
– | Lateral spine of right Exp2P5 inserted almost adjacent to base of terminal claw (Figs |
18 |
18 | Distal margin of segment 20 of A1R with simple distally-tapering process (Fig. |
Notodiaptomus spiniger (Brian, 1925) |
– | Distal margin of segment 20 of A1R with short, apically bifid process (Fig. |
Notodiaptomus santafesinus (Ringuelet & Martinez de Ferrato, 1967) |
1 | Lateral wings with two lobes on each side (Fig. |
Argyrodiaptomus (2) |
– | Lateral wings with one lobe on each side, with two pairs of sensillae, one large pair at apex and smaller pair on internal part of dorsal surface (Fig. |
5 |
2 | Posterior margins of Ped4 and Ped5 ornamented with rows of small dorsal spinules | Argyrodiaptomus denticulatus |
– | Posterior margins of Ped4 and Ped5 without rows of spinules | 3 |
3 | EnpP5 long, reaching distal end of inner margin of Exp1P5 (Fig. |
Argyrodiaptomus azevedoi |
– | EnpP5 short, not reaching distal end of inner margin of Exp1P5 | 4 |
4 | Longest distal spine on EnpP5 about half as long as endopod (Fig. |
Argyrodiaptomus falcifer |
– | Longest distal spine on EnpP5 less than half length of ramus; sensillae on swellings of GS asymmetrical, left sensilla larger and more robust than right, approximately 2.8 times longer than wide and directed more strongly backwards than right sensilla | Argyrodiaptomus furcatus |
5 | GS strongly asymmetrical, with well-developed posterior lobe on right side, extending beyond distal margin of GS; small sclerotized lobes present posteriorly on dorsal surface on left side of GS (Fig. |
Odontodiaptomus thomseni |
– | Posterior lobe on right side of GS weakly developed or absent; no small processes present dorsally on left side of GS or on left lateral margin of Ur3; CxP5 without small sclerotized process on distal lobe | 6 |
6 | Lateral wings larger and projecting further posteriorly on left side than on right (Fig. |
“Diaptomus” curvatus |
– | Lateral wings of Ped5 symmetrical or slightly asymmetrical; swellings on GS without hemispherical lobes on both sides | 7 |
7 | Transverse row of spinules marking position of posterior margin of Ped5; GS asymmetrical, with left swelling larger than right, and hemispherical in form; sensilla on left swelling directed slightly posteriorly, sensilla on right directed anteriorly (Fig. |
“Diaptomus” frutosae |
– | These characters not combined | Notodiaptomus (8) |
8 | Posterior margins of Ped4 and/or Ped5 with rows of spinules on dorsal surface | 9 |
– | Posterior margins of Ped4 and Ped5 lacking spinule rows dorsally | 15 |
9 | Rows of spinules present on dorsal surface of posterior margins of both Ped4 and Ped5 | 10 |
– | Rows of spinules present dorsally on posterior margin of Ped5 only | 11 |
10 | Single row of strong spinules present on posterior margin of Ped3, irregular rows of medium-sized spinules present on Ped4, and row of small spinules present on Ped5; lateral wings of Ped5 asymmetrical and elongate, left wing curved slightly anteriorly and right wing directed posteriorly (Fig. |
Notodiaptomus anisitsi |
– | Multiple rows of small spinules present on posterior margin of Ped4; lateral wings of Ped5 slightly asymmetrical, not elongate; GS with hemispherical swelling only on left side, left sensilla directed posteriorly; EnpP5 long, reaching beyond middle of inner margin of Exp1P5 (Fig. |
Notodiaptomus spiniger |
11 | Irregular and multiple rows of hair-like spinules present on dorsal and lateral surfaces of Ped5 | Notodiaptomus henseni |
– | Simple row or rows of spinules present dorsally along posterior margin of Ped5, spinules short and robust, not hair-like | 12 |
12 | Row of small spinules present, marking plane of incomplete suture between Ped4 and Ped5; lateral wings of Ped5 asymmetrical, projecting laterally (Fig. |
Notodiaptomus incompositus |
– | Lateral wings of Pr5 not projecting laterally; asymmetrical in shape | 13 |
13 | Lateral wings of Ped5 asymmetrical, left sensilla directed posteriorly (Fig. |
Notodiaptomus dentatus |
– | Lateral wings symmetrical or slightly asymmetrical; sensilla on lateral wing on left side of Ped5 directed posterolaterally, not posteriorly; right margin of GS lacking postero-distal lobe; internal seta on tip of Exp3P5 less than 4 times as long as external seta | 14 |
14 | Complete single row and short double row of spinules present dorsally near posterior margin of Ped4 (Figs |
Notodiaptomus iheringi |
– | Lines of spinules present on Ped4 border; lateral wings of Ped5 clearly asymmetrical, with apical (outer) sensilla about 2.2 to 2.3 times longer than wide; inner sensilla on lateral wings well developed (Fig. |
Notodiaptomus isabelae |
15 | Dorsal projection present on Ped4 (Fig. |
16 |
– | Ped4 lacking such dorsal projection | 18 |
16 | Inner right sensilla on lateral wing of Ped5 short, setule-like; GS asymmetrical, right sensilla about 2.3 times longer than wide, inserted on hemispherical lobe; right margin of GS with rounded lobe at posterior end (Fig. |
Notodiaptomus carteri |
– | Inner right sensilla of Ped5 not setule-like; GS right sensilla present but inserted in different position, no hemispherical lobe present on right side of GS at insertion of sensilla | 17 |
17 | Lateral wings of Ped5 projecting posteriorly; inner sensilla on Ped5 short with broad base (Fig. |
Notodiaptomus coniferoides |
– | Lateral wings of Ped5 not projecting posteriorly; position of inner sensilla on Ped5 variable; GS about 1.6 times longer than wide; sensilla on left swelling of GS directed slightly posteriorly, right sensilla directed slightly anteriorly; sensilla on CxP5 about 1.5 times longer than wide; external seta on Exp3P5 about half length of internal seta | Notodiaptomus santafesinus |
18 | Sensillae on both swellings on GS directed perpendicular to body axis; right margin of GS with small rounded lobe at posterior extremity (Fig. |
Notodiaptomus cearensis |
– | Sensilla on right swelling on GS directed posteriorly; right margin of GS without posterior lobe (Fig. |
Notodiaptomus conifer |
Diaptomus azevedoi Wright, 1935
Adult male, body length (excluding caudal setae) 1704 µm. Segment 11 of A1R with modified seta about twice length of modified seta on segment 10; segment 13 with long modified seta extending well beyond mid-point of segment 14 (Figs
Argyrodiaptomus azevedoi male. A, B, C Different views of Right P5 D Segments 12, 13 and 14 of A1R E, F Different views of Left P5, showing details of spinular ornamentation G Right Exp2P5 H Right P5 I Basal segments of right and left P5 J Segments 20, 21 and 22 of A1R K Detail showing spinules on BspP5R L Endopod of A2, showing pore and spinular ornamentation.
Argyrodiaptomus azevedoi male, SEM photographs. A Geniculate right antennule (500 µm) B Segments 12–17 of A1R (300 µm) C Segments 13–14 of A1R (100 µm) D Enp of A2, showing pore and spinular ornamentation (50 µm) E Segments 20–22 of A1R (100 µm) F Inset showing pore on segment 20 of A1R (5 µm) G Segments 12–16 of A1R (100 µm) H A2 (200 µm) I Maxillipeds (200 µm) J Distal endite of maxilliped (50 µm) K Maxilliped (200 µm).
Adult female, body length (excluding caudal setae) 1851 µm. Dorsal surface of Ped4 and Ped5 without spinule rows; complete suture present between Ped4 and Ped5; lateral wings slightly asymmetrical, with right wing larger than left; each wing with two sensillae close to postero-distal corner (Fig.
Argyrodiaptomus azevedoi female, SEM photographs. A Ventral view of fifth pediger bearing P5, and GS with eggs attached (200 µm) B Distal endite of maxilliped (50 µm) C Detail of spinules on basis of antenna (50 µm) D P5L (50 µm) E P5R (50 µm) F Distal part of ExpP5R (50 µm) G Apical spine of Enp3P4 (100 µm) H Genital area (fertilized female) on ventral surface of GS (50 µm) I P5 (100 µm).
In identifying species of Diaptomidae in general, caution must be exercised in using the shape of segment 20 of A1R for species identification because it is often variable within a population. This segment has a falciform process at the distal angle in several species (
Ultrafine-scale ornamentation characters might prove to be valuable in future comparative studies, including 1) the presence of a pore on segment 20 of male A1R (Fig.
This species is widely distributed, extending from northeastern Brazil to the Itaipu Reservoir at the end of the upper Paraná River in southern Brazil (Fig.
Diaptomus denticulatus Pesta, 1927
Adult male, body length 1657 µm. Dorsal surface of Ped5 ornamented with irregular rows of spinules (Figs
Argyrodiaptomus denticulatus male. A Segments 13–16 of A1R, showing spinous processes on segments 13 and 15 (arrowed) B Complete P5, showing proximal lobe on right Bsp overlapping margin of left Bsp C Segment 20 of A1R D Part of boundary between Ped4 and Ped5, showing ornamentation of dorsal spinules.
SEM photographs. Argyrodiaptomus denticulatus male. A Dorsolateral view of right side of Ped4 and Ped5 (100 µm) B Segment 20 of A1R (100 µm) C Segments 11–15 of A1R (200 µm) D A1R (500 µm) E P5R (200 µm) F, G Detail showing spinular ornamentation on process at base of lateral spine of Exp2P5R (F = 10 µm; G = 10 µm) H Right P5 (200 µm).
Adult female, body length 1753 µm. Spinules present dorsally along posterior margins of Ped3 and Ped4 (Fig.
Argyrodiaptomus denticulatus female, SEM photographs. A Dorsolateral view of Ped4 and Ped5, showing ornamentation of spinules (200 µm) B Lateral wings on left side of Ped5, showing sensilla at apex of each projection (100 µm) C Left terminal claw (50 µm) D Right terminal claw (50 µm) E Lateral view of P5 (100 µm).
The male illustrated was collected from the Salto Grande reservoir (SGRA-D), Uruguay River. The female illustrated here belongs to the collection of the Museo Argentino de Ciencias Naturales (Buenos Aires) (MACN-In 29733), and was examined because of the scarcity of females in the samples from de la Plata river basin. This species can be readily identified because of the distinctive structure of its P5 and the presence of dorsal rows of spinules on the male pedigers, which are lacking in other species of Argyrodiaptomus.
This species is common in the lower regions of the basins of the Paraná and Uruguay rivers (Figure
Diaptomus falcifer Daday, 1905
Diaptomus argentinus Wright, 1938
Argyrodiaptomus argentinus (Wright, 1938)
Adult male, body length 1495 µm. Ur4 with patches of fine spinules on dorsal surface (Fig.
Argyrodiaptomus falcifer male, SEM photographs. A Segments 12–15 of A1R, showing detail of spinous process on segment 14 (arrowed) (100 µm) B Entire A1R (500 µm) C Segments 11–15 of A1R (100 µm) D Adult male, lateral view (1000 µm) E Spinous processes on segments 13 and 14 A1R (20 µm) F Segment 20 of A1R, showing falciform process (50 µm) G Complete P5, caudal view (200 µm) H Medial surface of BspP5L, showing spinular ornamentation (50 µm).
Adult female, body length 1648 µm. Dorsal surface of pedigers lacking ornamentation of spinules (Fig.
The specimens illustrated here were caught in the middle stretch (RPAR-M2) of the Paraná River, and the sample contained only a few females and two males.
This species, together with others of this genus, can be very abundant in small scale habitats, such as water pools, and is generally scarce in lotic environments. It was found at only three stations in the present study (Fig.
Diaptomus furcatus Sars, 1901
Adult male, body length 1354 µm. Modified seta of segment 13 of A1R with bifid apex, reaching or exceeding proximal border of segment 14 (Fig.
Argyrodiaptomus furcatus male, SEM photographs. A A1R (500 µm) B A2R (200 µm) C Segments 10–15 and 20–22 of A1R (200 µm) D A1R (500 µm) E P5 (300 µm) F Distal segments of Exp2P5L (50 µm) G Right P1 (200 µm), anterior view H Distal segments of right P4 rami (50 µm), posterior view I Basis of right A2 showing spinules (20 µm).
Adult female, body length 1712 µm. Dorsal surface of Ped4 and Ped5 unornamented, lacking lines of spinules (Figs
The figured specimens were collected from the Iguaçu River at the reservoir of Foz do Areia (FARE-D) (Figure
Diaptomus anisitsi Daday, 1905
Diaptomus inflexus Brian, 1925
Adult male, body length 1144 µm. Row of spinules present along posterior margins of Ped3 and Ped4 (Fig.
Notodiaptomus anisitsi male, SEM photographs. A Ped3, Ped4, and Ped5 (100 µm) B Details of ornamentation on Ped3, Ped4 and Ped5 (100 µm) C Segments 10–22 of A1R (300 µm) D P5L (500 µm) E Segments 9–18 of A1R (200 µm) F A1R (500 µm) G Terminal somites of urosome and Exp2P5R (100 µm) H ExpP5R (100 µm).
Adult female, body length 1309 µm. Single row of spinules present along posterior margin of Ped3; irregular rows of spinules present, marking plane of posterior margin of Ped4; discrete transverse row of minute spinules present across middle of Ped5 (Fig.
The specimens illustrated were collected from the lower Paraná River (RPAR-L2). This species appears to be widely distributed in the south of the basin, from the Iguaçu River into more temperate conditions (Fig.
Diaptomus carteri Lowndes, 1934
Adult male, body length 1484 µm. Modified seta on segment 13 of A1R reaching distal end of segment 14 (Fig.
Adult female, body length 1770 µm. Ped4 and Ped5 separated by incomplete suture; process present on mid-dorsal surface of Ped4 (Fig.
The illustrated specimens were collected from the lower Paraná River (RPAR-L2). The body length is slightly larger than the known range for males, 1315 to 1439 μm, given by
Females of N. carteri can be readily distinguished from congeners by the asymmetrical swellings of the genital double-somite.
Diaptomus cearensis Wright, 1936
Adult male, body length 1008 µm. Ped4 lacking row of spinules along posterior margin. First segment of A1R unornamented (lacking typical spinule row) (Fig.
Adult female, body length 1485 µm. Ped4 and Ped5 fused, without trace of suture; lateral wings symmetrical, both with two pairs of sensillae (Fig.
The illustrated specimens were collected from the Parnaíba River at Emborcação Reservoir (Fig.
Diaptomus conifer Sars, 1901
Adult male, body length 1548 µm. Segment 1 of A1R with spinule row (Fig.
Notodiaptomus conifer male, SEM photographs. A Segments 13–16 of A1R (100 µm) B Segments 12–15 of A1R (100 µm) C Caudal view of P5 (100 µm) D Frontal view of P5 (100 µm) E Segments 13–17 of A1R (200 µm) F, G Segment 1 of A1R with spinule row arrowed (F = 50 µm; G = 20 µm) H P5L, and right Cx, Bsp and Exp1P5 (100 µm) I Detail of surface ornamentation of BspP5R (20 µm) J Segment 1 of Enp of A2 (50 µm).
Adult female, body length 1734 µm. Ped4 and Ped5 separated by complete suture; lateral wings slightly asymmetrical, each wing with sensilla at apex (Fig.
The illustrated specimens were collected from the upper Tiete River, at the Barra Bonita Reservoir. In addition to the well-developed spinous process on segment 15 of A1R of the male, N. conifer can be distinguished from other congeners like N. iheringi and N. cearensis, by its larger body size. These species also differ in their relative lengths of the lateral spines on the right Exp3P5 of the male.
Earlier studies (
Notodiaptomus conifer has been reported from Argentina to the northeast of Brazil, thus suggesting a widespread but scattered occurrence across South America.
Diaptomus conferoides Wright, 1927
Diaptomus lobifer Pesta, 1927
Adult male, body length 1051 µm. Ped2, Ped3 and Ped4 ornamented with spinules near posterior margin (Fig.
Notodiaptomus coniferoides male, SEM photographs. A Pedigers 2–5 (200 µm) B Last 3 urosome somites, CR, and P5R (300 µm) C Segments 9–18 of A1R (200 µm) D Detail showing insertion of lateral spine on Exp2P5R (20 µm) E Exp2P5R (100 µm) F P5 (100 µm) G A1R (300 µm) H A1R from segment 7 to tip (200 µm).
Adult female, body length 1411 µm. Suture between Ped4 and Ped5 incomplete; conical dorsal process present on Ped4 (Figs
The specimens illustrated here were collected in the upper Paraguay River. This is the only calanoid species that was found throughout the Paraguay basin (Fig.
This species has a wide distribution in rivers and associated systems like marginal ponds, but it is not typically recorded in reservoirs. This species is easily distinguishable by the position of the lateral spine on a lobe on the coxa of P5 and by the length of spinous processes on segments 13 and 15 of the male A1R. Some studies have reported this species under the name Notodiaptomus coniferoide (sic.) (see
We recommend a thorough comparative analysis of specimens of N. coniferoides found in the south of Brazil and in the lower Parana River. Comparison of our N. coniferoides with Amazonian specimens indicates that the material described in the present study was relatively smaller and we consider it is necessary to confirm the identity and status of N. coniferoides in de la Plata River Basin.
Adult male, body length 1046 µm. Posterior margin of Ped4 ornamented with more or less regular row of spinules (Fig.
Adult female, body length 1335 µm. Incomplete suture present between Ped4 and Ped5, with spinule row marking plane of fusion (Fig.
The illustrated specimens were collected from the middle stretch of the Paraná River (Fig.
Diaptomus henseni Dahl, 1894
Notodiaptomus oliveirai Matsumura-Tundisi, Espindola, Tundisi, Souza-Soares & Degani, 2010 [new synonym]
Adult male, body length 1123 µm. Dorsal suture between Ped4 and Ped5 incomplete (Fig.
Notodiaptomus henseni male, SEM photographs. A Segments 13–16 of A1R (100 µm) B Ped4 and Ped5, and anterior part of urosome (100 µm) C Detail showing spinular ornamentation of Ped3 and Ped4 (100 µm) D Detail of spinules on Ped2-Ped4 (100 µm) E, F Segment 1 of A1R, showing spinular ornamentation (50 µm) G A2 (100 µm) H Rostrum and spinules on segment 1 of A1L (50 µm) I Segments 11–16 of A1R (100 µm) J Detail of Bsp of A2 (20 µm).
Notodiaptomus henseni male, SEM photographs. A P1, Details of spinular ornamentation on Cx and Bsp (50 µm) B Left and right P5 (150 µm) C Sclerotized processes on left and right BspP5 (10 µm) D P5 (100 µm) E, F Detail of sclerotized processes on left and right BspP5 (E = 20 µm; F = 10 µm) G P5 (200 µm).
Adult female, body length 1275 µm. Complete suture present between Ped4 and Ped5 (Fig.
Notodiaptomus henseni female, SEM photographs. A Dorsal view of posterior part of prosome and GS (300 µm) B Ventral view of anterior end of cephalothorax, with rostrum and basal segments of antennule (150 µm) C Lateral view of posterior part of prosome, with arrows showing surface ornamentation of spinules, GS, urosome and CR (100 µm) D P5 (100 µm) E Dorsal view of segments 1 and 2 of left antennule, showing spinules (50 µm) F Dorsal view of segments 3–5 of left antennule, with some spinules (50 µm).
The specimens depicted here were collected in Furnas Reservoir on the Grande River (Fig.
In addition,
Recently,
Diaptomus iheringi Wright, 1935
Adult male, body length 922 µm. Posterior margin of Ped3 ornamented with rows of spinules; Ped4 and Ped5 ornamented with spinule rows along posterior margins and on lateral surfaces (Fig.
Notodiaptomus iheringi male, SEM photographs. A Rostrum (20 µm) B Ped4 and Ped5, lateral view (50 µm) C Ped3, Ped4, Ped5, lateral view (100 µm) D Detail of spinular ornamentation adjacent to sensilla on Ped5 (50 µm) E Ped4 and Ped5 (100 µm) F Ped3 and Ped4 (20 µm) G, H Segment 1 of A1R, showing row of spinules (50 µm) I Segment 1 of Enp of A2 (50 µm).
Adult female, body length 1093 µm. Incomplete suture present between Ped4 and Ped5 with plane of fusion marked by transverse row of strong spinules, with double row in middle section of dorsal surface (Figs
Our specimens were taken in the Grande River at Furnas Reservoir. In the present study this species was found in southeastern and southern Brazil and in the upper part of the Paraná River basin, with its southerly distribution boundary represented by the Iguaçu River (Fig.
Diaptomus incompositus Brian, 1926
Diaptomus paranaensis Pesta, 1927
Adult male, body length 1029 µm. Rows of spinules present dorsally along posterior margins of Ped3 and Ped4 (Fig.
Notodiaptomus incompositus male, SEM photographs. A Dorsal view of Ped3, Ped4, Ped5 and GS (100 µm), showing detail of dorsal rows of spinules B, C P5 (100 µm) D Segments 10–15 of A1R (100 µm) E Dorsal view of Ur4 (50 µm) F Dorsal view of Ur3 (20 µm) G P5L, and Cx and Bsp of P5R (100 µm).
Adult female, body length 1310 µm. Complete suture present between Ped4 and Ped5; transverse row of short spinules present along posterior margin of Ped4 (Fig.
This species was collected in the Machadinho Reservoir (MAC-U) on the Uruguay River and it can be easily identified by the unusual form of the caudal setae. It is distributed across the southern part of de la Plata river basin, including southern Brazil, south of the Iguaçu River (Fig.
Diaptomus isabelae Wright, 1936
Adult male, body length 919 µm. Transverse row of strong spinules present along posterior margin of Ped4 (Fig.
Adult female, body length 1056 µm. Complete suture present between Ped4 and Ped5, ornamented with row of strong spinules dorsally (Fig.
The specimens were collected in the middle section of the Paraná River at Yaciretá Reservoir (Fig.
This species can be easily distinguished from its congeners by the proximal processes on the internal margin of the male right BspP5 and by the shape of the wings and GS of females. A comparison with the description by
Diaptomus santafesinus Ringuelet & Martínez de Ferrato, 1967
Adult male, body length 967 µm. Dorsal and lateral surfaces of Ped3, Ped4 and Ped5 ornamented with scattered setules (Fig.
Notodiaptomus santafesinus male, SEM photographs. A Dorsal view of right side of Ped3, Ped4 and Ped5 (100 µm) B Right P5 (300 µm) C Terminal segments of EnpP4 and ExpP4 (100 µm) D Segments 13–17 of A1R (200 µm) E P5L (100 µm) F Detail of Exp2P5R (50 µm) G Detail of ornamentation of spinules of EnpP4 and ExpP4 (50 µm) H Segments 1 and 2 of A1R (100 µm).
Adult female, body length 1271 µm. Incomplete suture present between Ped4 and Ped5; lacking rows of spinules on posterior margin of pedigers (Fig.
The specimens examined were collected in the lower stretch of the Paraguay River (RPAG–B). This species is found in Argentina in the middle and lower stretches of the Paraná River and it can be considered to be a common species in the zooplankton community of this region (Fig.
Diaptomus spiniger Brian, 1925
Argyrodiaptomus spiniger (Brian, 1925)
Diaptomus toldti Pesta, 1927
Notodiaptomus orellanai Dussart, 1979
Adult male, body length 1466 µm. Patches of spinules present dorsally and laterally along suture between Ped3 and Ped4 (Figs
Notodiaptomus spiniger male. A Dorsal P5 B Ped3 and Ped4, showing details of spinular ornamentation C Details of spinules on corner of Ped4 D Spinous process of segment 13 of A1R E–G P5R H First segment of Enp of antenna 2 I Segment 20 of A1R J–L Different views of P5L M, N Different views of P5R O Detail of mammiform process on internal margin of right BspP5 P Segments 13–16 of A1R Q P5R.
Notodiaptomus spiniger male, SEM photographs. A Dorsal prosome and urosome, showing details of spinular ornamentation (200 µm) B, C P5 (150 µm) D A1R (120 µm) E Dorsal urosome somites 3 and 4, showing details spinular ornamentation (100 µm) F Segments 19 and 20 of A1R (100 µm) G Mammiform process on internal margin of BspP5R (20 µm) H Spinous process on segment 13 of A1R, and segments 14–17 (200 µm).
Adult female, body length 1688 µm. Complete suture present between Ped4 and Ped5; with several irregular rows of spinules present dorsally along posterior margin of Ped4 (Figs
Notodiaptomus spiniger female, SEM photographs. A Dorsal view of adult (500 µm) B Posterior pedigers, showing details of spinule rows, and GS (150 µm) C Dorsal view of posterior pedigers showing details of spinule rows, and GS (150 µm) D P5L (50 µm) E P5L (100 µm) F ExpP5 (50 µm) G Left P1, showing details of rows of spinules on Cx (100 µm).
The illustrated specimens were caught in the upper reaches of the Uruguay River, in the Machadinho Reservoir. The northernmost boundary of its distribution may lie in the northern sector of Paraná State (Brazil), its northernmost record is from the floodplain of the upper Paraná River (Fig.
This species has been the subject of much taxonomic confusion.
There are two other synonyms: “Diaptomus” birabeni Brehm, 1957 and Notodiaptomus orellanai Dussart, 1979 as proposed by Paggi in his MS Thesis in 1994. In each case the new species was established on the basis of morphological characters (
Further study of this species is necessary, including study of museum collections, in order to verify these synonymies and assess the evidence supporting the placement of this species in the genus Notodiaptomus, in part because of the presence of a mammiform process on the internal margin of the right BspP5, a character which conflicts with the current diagnosis of the genus.
Diaptomus thomseni Brehm, 1933
Adult male, body length 1088 µm. Small chitinous knob present distally on internal margin of left CR, other small chitinous processes present on right CR (Fig.
Odontodiaptomus thomseni male, SEM photographs. A Dorsal view of segments 11–16 of A1R (500 µm) B Detail of small chitinous knob on internal margin of left CR (20 µm) C P3, P4 and right P5 (caudal view), showing detail of EnpP5, lateral spine of P5 (200 µm) and were the semi-circular knob in inserted in Exp2P5 in frontal view D Right EnpP5 (2 µm) E Detail of chitinous knob on internal margin of CR (2 µm) F Detail of terminal claw (20 µm).
Adult female, body length 1245 µm. Incomplete suture present between Ped4 and Ped5; surface of Ped4 and Ped5 smooth, lacking rows of spinules dorsally along posterior margin (Fig.
The specimens were collected in the low stretch of the Uruguay River at Salto Grande Reservoir (Fig.
The genus Odontodiaptomus comprises three species (equivalent to the thomseni group of
“Diaptomus” curvatus Perbiche-Neves, Boxshall & Paggi, 2013 in Perbiche-Neves, Boxshall, Paggi, Rocha, Previattelli & Nogueira, 2013
Adult male, body length 923 µm. Ur4 with triangular-shaped dorsal process ornamented with small granulations at left distal corner (Figs
“Diaptomus” curvatus male, SEM photographs. A A1R (150 µm) B Detail of endopod of P5R (25 µm) C Segments 11–14 of A1R (50 µm) D P5 (200 µm) E Segment 20 of A1R (50 µm) F End of urosome and CR, showing detail of dorsal process (50 µm) G Detail of dorsal process on Ur4 (20 µm) H Right P5 (100 µm).
Adult female, body length 1120 µm. Incomplete suture present between Ped4 and Ped5; dorsal surface smooth, without spinules (Fig.
Several individuals of this species were found in the Yaciretá Reservoir (Fig.
“Diaptomus” frutosae Perbiche-Neves & Boxshall, 2013 in Perbiche-Neves, Boxshall, Paggi, Rocha, Previattelli & Nogueira, 2013
Adult male, body length 1292 µm. Irregular row of spinules present dorsally and laterally along posterior margin of Ped4 (Fig.
“Diaptomus” frutosae male, SEM photographs. A P5R, caudal view (300 µm) B P5L (50 µm) C Last segment of ExpP5L (200 µm) D P5 (100 µm) E Cx and Bsp of right P5 (100 µm) F Right P5, lateral view (200 µm) G Segments 12–14 of A1R (50 µm) H Segments 13 and 14, showing details of spinous process on segment 13 (50 µm) I Segment 20 of A1R (20 µm) J A1R (200 µm).
Adult female, body length 1346 µm. Incomplete suture present between Ped4 and Ped5; rows of strong spinules marking position of posterior margin of Ped4 (Figs
“Diaptomus” frutosae female, SEM photographs. A Dorsal view of female (500 µm) B, C Posterior pedigers, GS and urosome, in dorsal view, showing details of spinular ornamentation along posterior margin of Ped4 (B = 100 µm, C = 200 µm) D P5 (100 µm) E Exp3P5 and terminal claw (20 µm) F GS, ventral view (100 µm).
The specimens were found in the middle part of the Paraná River (Fig.
Nineteen species of diaptomid copepods were found during the present survey, but this figure probably is markedly lower than the estimated total for de la Plata river basin. The reason for this discrepancy may be related to sample representativeness due to limitations in terms of seasonality and kind of sampled freshwater habitats (only large rivers and reservoirs). Historical records suggest that there are at least another 15 species to be found in the basin.
There was no gradual decrease in the number of species towards the south of the continent, as might be inferred from
With the exception of the most widespread species, N. coniferoides, N. conifer and N. isabelae, the other diaptomids could be roughly divided into three groups according to their geographical distribution patterns, and their relationship to limnological variables and climatic factors (
As emphasized in the diagnoses of species given above, the detailed analysis of segment 20 of the male A1R revealed substantial variation in the shape and state of development of the distal process of this segment: a falciform process is usually present but other shapes may occur, and many species lack a process on this segment. It is necessary to be cautious in interpreting the wide variability in this process exhibited by species of some diaptomid genera. Based on our observations, it should not be used alone or as a strict diagnostic character for the identity of a particular taxon.
Notodiaptomus spiniger has been considered by some researchers to belong to the genus Argyrodiaptomus (see
In the genus Notodiaptomus characters such as the presence or absence of spinular ornamentation on the dorsal surface of the pedigerous somites and its pattern and shape proved to be highly informative diagnostic features, thus facilitating the discrimination between certain pairs of species or closely related groups of species. The combination of this character with others (for example the state – complete or incomplete – of the suture between Ped4 and Ped5) can provide additional, robust criteria to achieve accurate identifications in this genus.
The fifth leg (P5) of adult diaptomids provides the main set of differential characteristics for most species. However, for some species of the nordestinus complex, for instance, the differences may be minimal, even with reference to the finest scale details.
Body length is useful for the identification of some species of Notodiaptomus. It can, for example, help to separate N. conifer from other species that share the possession of a small lateral spine on Exp2P5R, such as N. iheringi and N. cearensis. Some studies, such as
Notodiaptomus incompositus showed a distribution restricted to the middle and southern parts of the basin. This species shares several morphological characteristics with N. deitersi (Poppe, 1891), the type species of Notodiaptomus, which is included in the nordestinus complex. Notodiaptomus spiniger is not treated as part of this complex, even though it is also restricted to the south of the basin, as are N. dentatus and N. carteri, among others. Although N. isabelae occurred only in the middle and lower basin, there are several published records from the upper part of the Paraná River basin and from other smaller basins nearby within the Brazilian shield, for example, in the State of Minas Gerais, Brazil (
We did not find N. deitersi (Poppe, 1891) in this survey. The species was originally described from the region of Cuiabá (Mato Grosso State, Brazil) and it is the type species of Notodiaptomus (
We decided to treat Notodiaptomus cf. spinuliferus as incertae sedis and not include it in this work because the vouchers deposited in MZUSP (6971) (Fig.
As a result of previous faunistic surveys, some researchers have inferred that particular calanoid species have disappeared from the region (e.g.
Notodiaptomus conifer was reported as missing from Jurumirim Reservoir (Paranapanema River) by
Taxonomically, the identification of N. conifer can be problematic. We recommend comparing the spinous process on segment 15 of the male A1R as well as considering body size. Notodiaptomus conifer has a greater body length than N. iheringi and other species, such as N. henseni, which could be confused with it. The presence of a row of spinules on the first segment of the A1R in N. iheringi allows this species to be distinguished from N. conifer. In N. henseni, the presence of sclerotized processes on the basipodites of both right and left P5 also avoids any confusion with N. conifer. In some N. henseni the presence of a well-developed spinous process on segment 15 of the A1R may cause confusion with N. conifer if this feature is considered alone, so it is always advisable to try to use more than one distinguishing feature, while also taking into account the known geographical distributions.
All these data indicate that care must be taken when reporting the disappearance of a particular species from a habitat; this conclusion should be drawn only after failure to find the species after continued sampling over extended periods of time. The probability of a species being found again seems high when routine sampling is used (e.g.
Studies carried out in Europe and Asia over long time periods have demonstrated that particular species may be dominant at rare intervals over a 20 year time series.
Within de la Plata river basin, the results obtained in this study combined with our review of published data suggest the following recommendations for future research: 1. Continue studying sites where historical collections have been made; 2. Study large and productive reservoirs for which little information is available and where new species records are likely; 3. Conduct studies in places that are unexplored and represent gaps in information, including reservoirs, ponds and wetlands adjacent to tributaries of the Upper Paraguay River, Upper Parnaíba River, Upper Uruguay River and west of the state of Rio Grande do Sul, the region of “Esteros de Iberá” (in Argentina); and 4. Study the transition areas between the northern and southern faunas, in the rivers Iguaçu, Paraná (Yaciretá Reservoir) and middle/upper Uruguay, focusing on flood periods and correlations with bird migration routes.
There are several taxonomic problems still to be resolved within the Neotropical diaptomids and it would be useful to test whether Argyrodiaptomus and Odontodiaptomus are monophyletic. Many species currently placed in Notodiaptomus and “Diaptomus” must eventually be relocated. Diaptomus, for example, continues to serve as a temporary repository for species whose affinities are as yet unresolved, such as the unrelated species “Diaptomus” frutosae and “Diaptomus” curvatus. We also consider that Notodiaptomus spiniger, N. anisitsi and N. coniferoides do not belong in Notodiaptomus. One solution might be the creation of additional new genera to accommodate some of these distinctive Neotropical diaptomids, however, new genera should only be erected after thorough phylogenetic analysis. Phylogenetic analyses based on morphological and molecular data are currently under way by the present authors and will be published elsewhere when completed.
The following genera and species should be reviewed as their taxonomic status is currently equivocal: N. spiniger (synonymy needs verification and its placement in Notodiaptomus requires testing); N. spinuliferus (the material deposited in MZUSP should be redescribed and compared with material found elsewhere and identified as N. spinuliferus); N. amazonicus from Amazonia should be compared with N. amazonicus from de la Plata river basin, including molecular analysis; the morphology of N. coniferoides should be checked by comparison with the holotype from the Amazon; this will provide a more robust basis for determining the true distribution of this species across the continent.
The rarity and seasonallity of some species in the basin, such as N. anisitsi and A. falcifer, and the absence of N. deitersi, O. paulistanus, O. michaelseni, and S. corderoi, suggests that carrying out only two sampling programmes over a year may be inadequate in terms of recording sound data on the seasonal variability of these species. It is necessary to be very careful when using terms like “disappearing species”, since their absence might reflect an inadequate sampling regime.
To FAPESP (2008/02015-7; 2009/00014-6; 2011/18358-3) for financial support. We are deeply grateful to Eduardo Suárez Morales (Mexico), Janet Bradford Grieve (Australia) and Silvina Menu Marque (Argentina) for their detailed revision of the text; to Juan C. Paggi (Argentina), Edinaldo Nelson dos Santos Silva (INPA, Manaus, Brazil), Janet W. Reid (USA), and Margarita Frutos (Argentina) for providing information, suggestions and for help in checking some identifications; to Maria L. Negreiros Fransozo (UNESP, Botucatu, Brazil) for lending optical equipment; and to Tiago Tardivo (CME-UNESP, Botucatu, Brazil) for help with the SEM photographs.