Research Article |
Corresponding author: Frank Fiers ( frank.fiers@naturalsciences.be ) Academic editor: Oana Teodora Moldovan
© 2015 Frank Fiers, Moïssou Lagnika.
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:
Fiers F, Lagnika M (2015) Four new representatives of the genus Allocyclops Kiefer, 1932 from semi-consolidated subsoil aquifers in Benin (Copepoda, Cyclopoida, Cyclopidae). Subterranean Biology 16: 1-36. https://doi.org/10.3897/subtbiol.16.4467
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Several species of cyclopoid copepods were collected from improved and unimproved hang-dug-wells in the Republic of Bénin over the years 2009–2014. Fifty five wells located in seven different districts were sampled: 15 wells in the district of Pobè (Department Plateau) and 1 well in Kétou (Department Plateau), 4 wells in Porto-Novo District (Department Ouémé) and, 15 wells in Lokossa District (Department Mono),18 wells in Parakou District (Department Borgou), 1 well in Abomey-Calavi District (Department Atlantique) and 1 in Zogbodomè District (Department Zou). Among them, 4 new species of the genus Allocyclops Kiefer, 1932 were found and are described herein: Allocyclops spinifer sp. n., A. nudus sp. n., A. pilosus sp. n. and A. sakitii sp. n. They are compared with the three African species previously described. Allocyclops appears to be a sub-Saharan taxon specialized to thrive in the variable environment of subsoil aquifers in laterite deck beds. An identification key to the 7 different African Allocyclops species is provided.
Copepoda , Cyclopidae , subterraneous, Bénin, new species, key, laterites
Within the context of a research program in the Republic of Bénin on the biodiversity of the subterranean realm in relation to water quality, pollution vulnerably and local use 48 wells were regularly sampled over a period of one year. Seven wells located in the close vicinity of the monitored wells were occasionally visited. Six of the seven areas are located in the drainage basin of the Ouémé: the area of Parakou in the north of the basin and the area of Kétou, Zogbodomè, Pobè, Abomey-Calavi and Porto-Novo in the south. The seventh study area, the district of Lokossa, belongs to the lower part of the Mono-Couffo drainage basin (see Fig.
Map of Bénin indicating the study areas (black triangles): Lokossa stands for 15 sampled wells, Porto Novo for 4 sampled wells, Pobé for 15 sampled wells and Parakou for 18 sampled wells. Grey dashed lines contour the limits of the 4 different drainage basins, from http://eaubenin.bj/docs/Bassins_Versants/Atlas_Hydrographique_Rapport_final.pdf (accessed November 2014).
The Republic of Bénin, a country 650 km long and 325 km at its widest, in South-West Africa is a sub-Saharan savannah region enduring a tropical climatologic regime: humid in the south, semi-arid in the north (
African continental waters have been a very attractive study subject in zooplankton research. Large regions, however, of this vast continent remain unexplored (
Among the copepods collected during this intensive study (i.e. mostly representatives of Afrocyclops Sars, 1927, Thermocyclops Kiefer, 1927, Mesocyclops Sars, 1914) no less than four different species which undoubtedly belong to the cyclopid genus Allocyclops were detected. They are described herein and compared with the 3 other African Allocyclops previously known.
In the following Tables
Sample localities and principal well characteristics in the district of Lokossa: presence of Allocyclops species (dark grey background), other copepods (light grey background) and absence of Cyclopoida (white background).
Station | Total depth (m) | Ø (m) | Water level fluct. (m) | Wall character. | Well closure | Principal use | Location |
---|---|---|---|---|---|---|---|
P1 | 31,90 | 2,00 | 1,25 | cemented | slab | potable | 06°42'29.5"N, 01°48'11.8"E |
P2 | 41,30 | 1,45 | 3,00 | cemented | slab | potable | 06°42'29.2"N, 01°48'11.9"E |
P3 | 46,15 | 2,00 | 4,80 | cemented | slab | potable | 06°45'24.9"N, 01°49'11.7"E |
P4 | 27,75 | 2,00 | 5,00 | cemented | slab | potable | 06°42'41.0"N, 01°44'46.7"E |
P5 | 14,10 | 1,20 | 6,30 | natural | open | potable | 06°43'12.5"N, 01°44'02.7"E |
P6 | 13,15 | 1,10 | 3,70 | natural | open | potable | 06°43'24.8"N, 01°44'39.2"E |
P7 | 4,45 | 1,00 | 2,00 | cemented | iron | domestic | 06°38'22.8"N, 01°42'54.7"E |
P8 | 5,00 | 1,45 | 0,50 | cemented | slab | domestic | 06°39'01.3"N, 01°40'32.1"E |
P9 | 5,60 | 0,90 | 1,20 | cemented | open | domestic | 06°38'43.2"N, 01°42'26.8"E |
P10 | 11,65 | 2,10 | 2,90 | cemented | slab | domestic | 06°40'03.5"N, 01°47'43.4"E |
P11 | 10,70 | 0,90 | 2,10 | natural | open | domestic | 06°38'54.6"N, 01°46'47.6"E |
P12 | 4,80 | 1,40 | 3,70 | cemented | open | domestic | 06°38'14.8"N, 01°46'54.3"E |
P13 | 5,70 | 1,20 | 2,60 | cemented | slab | domestic | 06°41'29.0"N, 01°39'02.2"E |
P14 | 8,85 | 2,00 | 6,35 | cemented | slab | domestic | 06°42'50.1"N, 01°38'17.8"E |
P15 | 2,80 | 1,30 | 0,50 | natural | open | domestic | 06°43'04.1"N, 01°40'59.8"E |
Sample localities and principal well characteristics in the district of Parakou: presence of Allocyclops species (dark grey background), other copepods (light grey background) and absence of Cyclopoida (white background).
Station | Total depth (m) | Ø (m) | Water level fluct. (m) | Wall character. | Closure | Principal use | Location |
---|---|---|---|---|---|---|---|
A1 | 7,78 | 1,2 | 1,18 | cemented | iron | potable | 09°16'22.7"N, 02°34'54.7"E |
A2 | 6,95 | 1,05 | 0,85 | cemented | open | domestic | 09°19'40.3"N, 02°37'28.8"E |
A3 | 5,9 | 1,25 | 1,15 | natural | iron | domestic | 09°20'53.9"N, 02°36'44.0"E |
A4 | 13,15 | 1,5 | 4,35 | cemented | iron | potable | 09°18'42.1"N, 02°36'07.6"E |
A5 | 8,4 | 1,2 | 3,1 | natural | open | domestic | 09°20'51.8"N, 02°36'14.8"E |
A6 | 8,85 | 1,2 | 0,2 | cemented | open | domestic | 09°20'18.7"N, 02°37'36.3"E |
B1 | 7,9 | 0,9 | 3,5 | cemented | iron | potable | 09°19'57.2"N, 02°39'40.6"E |
B2 | 6,6 | 1,45 | 0,8 | cemented | iron | potable | 09°19'55.2"N, 02°39'40.5"E |
B3 | 7,5 | 1,4 | 1,6 | natural | slab | domestic | 09°20'03.5"N, 02°37'47.2"E |
B4 | 8,0 | 1,2 | 3,1 | cemented | open | domestic | 09°20'08.1"N, 02°37'45.4"E |
B5 | 10,45 | 1,15 | 5,95 | natural | open | domestic | 09°20'18.0"N, 02°37'51.1"E |
B6 | 16,15 | 1,4 | 10,55 | cemented | iron | gardening | 09°21'33.4"N, 02°38'32.4"E |
C1 | 8,8 | 1,1 | 2,1 | cemented | iron | domestic | 09°23'28.7"N, 02°37'21.4"E |
C2 | 14,2 | 1,8 | 5,5 | cemented | iron | potable | 09°23'05.5"N, 02°37'23.2"E |
C3 | 10,8 | 1,2 | 5,8 | cemented | iron | potable | 09°21'58.4"N, 02°37'30.6"E |
C4 | 5,5 | 1 | 0,9 | natural | open | domestic | 09°21'45.4"N, 02°37'46.6"E |
C5 | 12,85 | 2 | 7,25 | cemented | iron | potable | 09°21'56.9"N, 02°36.41.2"E |
C6 | 11,2 | 1,1 | 5,3 | cemented | slab | potable | 09°21'01.7"N, 02°37'37.6"E |
Sample localities and principal well characteristics in the district of Pobè: presence of Allocyclops species (dark grey background), other copepods (light grey background) and absence of Cyclopoida (white background).
Station | Total depth (m) | Ø (m) | Water level fluct. (m) | Wall character. | Well closure | Principal use | Location |
---|---|---|---|---|---|---|---|
Pb1 | 17,30 | 1,40 | 2,00 | natural | open | potable | 06°58'57.3"N, 02°40'32.4"E |
Pb2 | 36,08 | 1,40 | 3,70 | cemented | slab | potable | 06°58'57.1"N, 02°39'54.9"E |
Pb3 | 40,80 | 1,40 | 3,60 | cemented | slab | potable | 06°58'49.7"N, 02°39'19.6"E |
Pb4 | 50,50 | 1,45 | 3,20 | cemented | slab | potable | 06°57'48.1"N, 02°38'50.2"E |
Pb5 | 16,20 | 1,50 | 2,90 | cemented | slab | potable | 06°59'13.6"N, 02°39'56.6"E |
Pb6 | 19,60 | 1,05 | 17,45 | cemented | wood | domestic | 07°12'03.5"N, 02°36'04.4"E |
Pb7 | 6,10 | 0,60 | 3,75 | natural | iron | domestic | 07°09'28.7"N, 02°39'09.2"E |
Pb8 | 26,27 | 1,50 | 7,30 | cemented | open | domestic | 07°02'10.1"N, 02°42'16.7"E |
Pb9 | 19,25 | 1,40 | 4,65 | cemented | wood | potable | 07°09'23.5"N, 02°44'09.5"E |
Pb10 | 17,60 | 1,60 | 5,75 | cemented | open | potable | 07°09°18.2"N, 02°44'10.6"E |
Pb11 | 13,90 | 1,10 | 7,45 | natural | iron | domestic | 07°08'59.4"N, 02°45'02.1"E |
Pb12 | 14,90 | 1,25 | 1,20 | cemented | slab | potable | 06°58'05.0"N, 02°41'47.9"E |
Pb13 | 34,52 | 1,55 | 4,50 | cemented | slab | potable | 06°57'48.4"N, 02°42'10.1"E |
Pb14 | 36,00 | 1,52 | 2,20 | cemented | slab | potable | 06°57'26.0"N, 02°42'25.9"E |
Pb15 | 31,00 | 1,20 | 2,70 | cemented | open | potable | 06°57'39.6"N, 02°42'14.4"E |
Sample localities and principal well characteristics in the district of Porto-Novo: presence of Allocyclops species (dark grey background).
Station | Total depth (m) | Ø (m) | Water level fluct. (m) | Wall character. | Closure | Principal use | Location |
---|---|---|---|---|---|---|---|
AR | 7,80 | 0,90 | 3,00 | cemented | iron | domestic | 06°29'04.2"N, 02°38'24.3"E |
SA | 7,00 | 0,90 | 2,00 | natural | open | domestic | 06°29'02.4"N, 02°38'26.0"E |
LB | 10,50 | 0,85 | 2,85 | natural | open | domestic | 06°29'05.7"N, 02°38'25.3"E |
AH | 4,00 | 1,00 | 2,50 | natural | iron | domestic | 06°29'18.8"N, 02°38'14.8"E |
The occasionally sampled wells: at Abomey-Calavi located adjacent to the university grounds in the village of Zogbadjè (11.75 m deep, 1 m Ø, 2.10 mean water level, natural walls, left open, domestic purposes; well location: 6°25'32.2"N, 002°20'17.2"E; Department Atlantique); village well at Kéwi in the district of Kétou (open, natural, abandoned well in secondary forest; district centre: 7°21'29"N, 2°36'27"E; Department Plateau) and well at Zogbodomè situated close to the main road (open natural domestic use; district centre 7°05'N, 2°06'E; Department Zou).
Biological samples of the water column were taken with a Cvetkov plankton net (funnel 200 µm mesh size, 150 µm below valve). Baited traps were used to collect bottom dwelling animals. Total depth, diameter (Ø), and mean water depth were measured with a decametre and the exact location with a Garmin® GPS60. Samples were fixed at the spot adding 100 ml of a 7% formaldehyde solution. Sorted out at the laboratory of the Zoology Department of the Abomey-Calavi University, each taxon has been stored separately and preserved in 70% ethanol.
Copepods were first observed in temporary slides (mounted in glycerol). Dissected animal parts are mounted on permanent slides in glycerol, the cover glasses sealed with a polyurethane varnish. Preserved specimens are stored in 75% buffered ethyl alcohol (with 5–10% glycerol). All material is stored in the Crustacea collection of the Royal Belgian Institute of Natural Sciences and inventoried with acc. nr RBINSc COP #. Observations and illustrations were made on a Leitz Diaplan phase contrast microscope equipped with a camera lucida. Abbreviations used in the descriptions: Aesth, aesthetasc; CIV-CV, copepodid stages 4 and 5; EXO, END, exopodite and endopodite, respectively; L/W ratio, length/width ratio; armature distribution of the leg ramal segments: Roman numerals refer to spines, Greek numerals to setae.
Ishèko (Pobè District, Ouémé drainage basin), Well Pb5: West from Pobè center along the principal road to Kétou District (details in Table
Female holotype, dissected and mounted on 3 slides (RBINSc COP 10.304A-C), allotype male, dissected and mounted on 3 slides (RBINSc COP 10.305A-C), 1 female paratype dissected on 2 slides (RBINSc COP 10.306A-B) and 4 females, 5 males and 4 juveniles preserved (RBINSc COP 10.307), all collected 26/07/2012; additional paratypic material: 3 females, 4 males, 5 juveniles (preserved material RBINSc COP 10.353), topotypic, collected 16/04/2013.
derived from the Latin words spinula and ferre, respectively meaning small thorn and to carry, and referring to the spinular ornamentation on pediger 5, on leg 4 and on the caudal rami.
(1) Pobè District, Ouémé drainage basin:
– Well Pb6: 07/02/11: 6 ♀♀, 2 ♂♂ and 2 CV copepodids (♀ dissected RBINSc COP 10.308A-C, preserved material RBINSc COP 10.309); 27/11/2012: 19 ♀♀, 5 ♂♂, 1 CIV & 1 CV ♀ (CV dissected RBINSc COP 10.386A-B, preserved material RBINSc COP 10.327); 04/03/2013: 21 ♀♀, 11 ♂♂, 5 juveniles (preserved material RBINSc COP 10.339); 26/11/2012: 2 ♀♀ (preserved material RBINSc COP 10.377); 20/09/2013: 9 ♀♀, 2 ♂♂ (preserved material RBINSc COP 10.376).
– Well Pb12: 24/07/2012: 3 ♀♀, damaged (preserved material RBINSc COP 10.328).
(2) Zogbodomè District, Ouémé drainage basin:
– Well Zogbodomè: 21/10/2009: 2 ♀♀, 1 ♂ (preserved material RBINSc COP 10.387).
Female. Habitus (Fig.
Allocyclops spinifer sp. n. A urosome in ventral view B urosome in ventral view C copulatory pore and duct D leg 5 and lateral part of pediger, in ventral view E leg 5 and lateral side of pediger in ventral view. (Female RBINSc COP 10.304: A, C; RBINSc COP 10.306: E; male RBINSc COP 10.305: B, D).
Caudal rami (Fig.
Antennule 11-segmented, not reaching beyond cephalothorax, and having the typical armature distribution i.e.: 1(8)-2(4)-3(6)-4(2)-5(2)-6(2)-7(3)-8(2+Aesth.)-9(2)-10(2+Aesth.)-11(7+Aesth). First segment with crescent row of slender spinules in proximal half (Fig.
Allocyclops spinifer sp. n. A antennulary segment 1 B antennulary segments 8–11; A. pilosus sp. n. C antennulary segments 8–11 D antennulary segment 1; A. nudus sp. n. E antennulary segments 8–11 F antennulary segment 1; A. sakitii sp. n. G antennulary segments 8–11 H antennulary segment 1; A. spinifer sp. n. I maxillary basis and endpodite; A. nudus sp. n. J maxillary basis and endopodite; A. pilosus sp. n. K maxillary basis and endopodite; A. sakitii sp. n. L maxillary basis and endopodite (RBINSc COP 10.304: A, B, I; COP 10.310: C, D, K; COP 10.367: E, D, J; COP 10.253: G, H, L).
Antennal general aspect and armature as in A. cavicola: i.e.: 4-segmented, short exopodal seta, 2 abexopodal setae, first endopodite segment with 1, second one with 9, and ultimate segment with 7 elements. Frontal surface of coxobasis (Fig.
Antennary coxobasis: Allocyclops spinifer sp. n. A frontal view B caudal view C caudal view; Allocyclops pilosus sp. n. D frontal view E caudal view F caudal view; Allocyclops nudus sp. n. G frontal view H caudal view; Allocyclops sakitii sp. n. I caudal view J frontal view (RBINSc COP 10.304: A; COP 10.306B, COP 10.305: C; COP 10.310: D, E; COP 10.312: F; COP 10.3334, COP 10.367; COP 10.355: I, J).
Mandible, maxillule and maxilliped as in A. cavicola. General aspect of maxilla as in A. cavicola. Medial claw of maxillary basis (Fig.
Legs 1–4 protopodite (Fig.
Allocyclops spinifer sp. n. A leg 1, frontal view B intercoxal sclerite of leg 1, frontal view C leg 2, frontal view D outer region of coxa and basis of leg2, caudal view E distal segment of leg 3 endopodite, caudal view F leg 4, frontal view G leg 4 protopodite and endopodite, caudal view H distal endopodite segment of leg 3, frontal view. (Female RBINSc COP 10.306: A, B, C, G; female RBINSc COP 10.304: D, F, E; male RBINSc COP 10.305: H).
Legs 1–4 with 2-segmented rami (Fig.
EXO END
Leg 1 I,1 - III,2,3 0,1 - 1,I+1,3
Leg 2 I,1 - III,I+1,4 0,1 - 1,I+1,4
Leg 3 I,1 - III,I+1,4 0,1 - 1,I+1,4
Leg 4 I,0 - II,I+1,4 0,1 - 1,II,3
Outer distal edge of second endopodite segment of legs 1–3 distinctly protruded, of leg 4 small and as large as inner one. Frontal and caudal surface of leg segments of legs 1–2 and frontal surface of leg 3 unadorned. Caudal surface of distal endopodite segment of leg 3, and frontal and caudal surface of leg 4 segments with diffuse pattern of tiny spinules.
Caudal surface of proximal endopodite segment of leg 4 inflated. Distal endopodite segment of leg 4 twice as long as wide with inner terminal spine as long as segment. Outer terminal spine slightly longer than half inner one (ratio: 0.55–0.56). Inner terminal spine set with long slender spinules. Outer subdistal and inner setae, equally long, not reaching noticeably beyond longest (outer) terminal spine.
Leg 5 (Fig.
Leg 6 vestiges located laterally with 3 elements: dorsal one short and setiform, middle and ventral one dwarfed, blunt triangular, with hyaline appearance. Valves unadorned. Copulatory pore located midventrally in anterior third of somite, leading to U-shaped duct (Fig.
Male. Habitus (Fig.
Antennule as described for A. cavicola: i.e.16-segmented with the following armature distribution: 1(8+3 Aesth.)-2(4)-3(2)-4(1)-5(2)-6(2)-7(2)-8(2)-9(2+Aesth.)-10(2)-11(2)-12(2)-13(1+Aesth.)-14(4)-15(3)-16(11+Aesth.). Aesthetascs on segment 1, 9 and 13 long, narrow and linguiform. Mouthparts as in the female. Legs 1–4 with general structure and armature as in the female, but terminal segment of endopodites slightly narrower (see ex. for leg 3: Fig.
Leg 5 (Fig.
Leg 6 (Fig.
Animals from Well Pb 6 collected on 07/02/2011 are smaller with the body length of the females ranging from 875 to 1009 μm, that of the males between 890–895 μm. They display shorter caudal rami (L/W ratio 1/2.6–2.7) with slightly longer setae: distolateral one 1.3–1.4 times longer than the ramus, and medial terminal element 1.1–1.3 times longer than ramus. In all other respects they resemble A. spinifer sp. n. as described above.
Syn. Allocyclops sp. [partim] in
Guéma, Parakou District, Ouémé drainage basin: Well C2 (details in Table
Female holotype, dissected and mounted on 3 slides (RNBNSc COP 10.313A-C), allotype male, dissected and mounted on 2 slides (RBINSc COP 10.312A-B), paratypes: 11 females and 2 males preserved in alcohol (RBINSc COP 10.314), all collected 07/03/2013; and additional (topotypic) paratypes collected on 13/02/2011: 4 females, 1 male, 1 juvenile; on 08/03/2013: 37 females, 15 males, 7 juveniles (preserved material RBINSc COP 10.340).
From the Latin word “pilus” meaning hair and refers to the rows of delicate hairs along the medial margin of the caudal rami.
(1) Parakou District, Ouémé drainage basin:
– Well A1: 10/07/12: 5 ♀♀, 1 CV juvenile ♀ (preserved material RBINSc COP 10.321);
– Well B6: 26-10-11: 1 ♀ (preserved material RBINSc COP 10.320); RBINSc COP 10.311);
– Well C6: 13/12/10: 11 ♀♀ (dissected ♀ RBINSc COP 10.316A-B, preserved material RBINSc COP 10.317; 29/10/2011: 33 ♀♀, 1 juvenile (preserved material RBINSc COP 10.342).
(2) Pobè, Plateau District, Ouémé drainage basin:
– Well Pb6: 07/02/11: 4 ♀♀ (preserved material RBINSc COP 10.332); 27/07/2012: 4 ♀♀, 2♂♂ (♀ dissected RBINSc COP 10.385A-B, preserved material RBINSc COP 10.336); 26/11/12 : 2 ♀♀ (preserved material RBINSc COP 10.374); 04/03/2013: 3 ♀♀, 3♂♂, 2 juveniles (preserved material RBINSc COP 10.345); 20/09/2013: 5 ♀♀, 1 CV copepodite (preserved material RBINSc COP 10.375);
– Well Pb7: 27/7/12: 15 ♀♀, 4 ♂♂ (dissected ♀ RBINSc COP 10.325A-B, preserved material RBINSc COP 10.326); 16/04/2013: 28 ♀♀, 2 ♂♂, 1 juvenile (preserved material RBINSc COP 10.346);
– Well Pb 8: July 2013: 2 juveniles (preserved material RBINSc COP 10.343);
– Well Pb 9: 17/04/2013: 2 ♀♀ (preserved material RBINSc COP 10.341);
– Well Pb10: 27/07/2012: 2 ♀♀ (preserved material RBINSc COP 10.347); 30/09/2012: 2 ♀♀, 2 ♂♂ (preserved material RBINSc COP 10.384); 25/11/2012: 3 ♀♀ (preserved material RBINSc COP 10.383);
– Well Pb11: 27/07/2012: 68 ♀♀, 3 ♂♂ (preserved material RBINSc COP 10.344); 30/09/2012: 387 F 5 CV females; 23/02/13: 3 ♀♀, 6 CIV juvenile ♀♀, 4 CV juvenile ♀♀ (dissected adult ♀♀ RBINSc COP 10.322A-C and COP 10.323A-C, preserved material RBINSc COP 10.324);
(3) Porto-Novo, Ouémé District, Ouémé drainage basin:
– Well SA: 01/01/10: 8♀♀, 1 CIV ♀ juvenile (dissected ♀ RBINSc COP 10.318A-B, preserved material RBINSc COP 10.319); 24/10/2010: 625 adult females and males (preserved material RBINSc COP 10.382);
– Well AH: 01/11/2009: 12 ♀♀ with body covered with stalked ectoparasites (preserved material RBINSc COP 10.329 and COP 10.330); 30/03/2013: 1 ♀ (preserved material, RBINSc COP 10.380); 31/01/2014: 6 ♀♀ 2 CV juveniles (preserved material, RBINSc COP 10.381);
– Well AR: 09/12/2010: 1 ♀ (material preserved RBINSc COP 10.331); June 2013: 1 ♀, damaged (preserved RBINSc COP 10.379);
– Well LB: 20/01/2014: 83 ♀♀ 5 ♂♂ 6 copepodites (damaged, material preserved RBINSc COP 10.373); 09/03/2014: 5 ♀♀ (material preserved RBINSc COP 10.374).
(4) Kéwi, Kétou District, Ouémé drainage basin:
– village well: 08/02/11: 17 ♀♀, 1 ♂ (preserved material RBINSc COP 10.315).
Female. Body (Fig.
Caudal rami (Fig.
Antennule 11-segmented with typical armature distribution. First segment with crescent row of slender spinules in proximal half (Fig.
Antennal general aspect and armature as in A. cavicola. Frontal surface of coxobasis with defuse pattern of minute spinules in proximal half on and near outer margin (Fig.
Mandible, maxillule and maxilliped as in A. cavicola. General aspect of maxilla as in A. cavicola. Medial claw of basis rather short and sharp, furnished with 4 large teeth increasing in length medially (Fig.
Legs 1–4 protopodite (Fig.
Allocyclops pilosus sp. n. A leg 1, frontal view B leg 1 intercoxal sclerite C leg 2, frontal view (aberrant number of medial setae on endopodite, normal seta complement shown at the left) D distal endopodite segment of leg 3, frontal view E idem F leg 4, frontal view G leg 4 protopodite and endopodite, caudal view H leg 4 intercoxal sclerite, frontal view I–J distal endopodite segment of leg 4 of other specimens (Females RBINSc COP 10.313: F, G; COP 10.310: A–D, G–H, COP 10.318: I; COP 10.316: J; male RBINSc COP 10.312: E).
Legs 1–4 with 2-segmented rami. Exopodite and endopodite of the legs equally long (Fig.
Leg 5 (Fig.
Leg 6 (Fig.
Male. Prosome and urosome equally long with metasome narrower than in the female (Fig.
Antennule as described for A. cavicola in
Leg 5 (Fig.
Variability: In most cases the medial triangular depression of the caudal rami is furnished with two parallel sets of fragile hairs as shown in Fig.
Syn. Allocyclops n sp. [partim] in
Ladjifarani or “Nouveau quartier”, Parakou District, Ouémé drainage basin: Well B6 (details in Table
Holotype female, dissected and mounted on 2 slides, RBINSc COP 10.334A-B, allotype male, dissected and mounted on 2 slides, RBINSc COP 10.335A-B; paratypes: dissected female RBINSc 10.367A-B, preserved material: 7 females, 3 males, 5 juveniles (RBINSc COP 10.333) all collected 29/10/2011; additional paratypic material (topotypes): 3 females collected 16/01/11 (preserved RBINSc COP 10.352).
From the Latin adjective “nudus” meaning naked and refers to the absence of hairy or spinular ornamentation on the pedigers surface and caudal rami, contrasting the species from A. spinifer sp. n. and A. pilosa sp. n.
(1) Parakou District, Ouémé drainage basin.
– Well A1: 27/08/2011: 1 ♀, 8 ♂♂ (preserved material RBINSc COP 10.337); 11/01/2011: 4 ♀♀, 1 ♂, 40 juveniles, decomposed condition (preserved material RBINSc COP 10.348); 15/02/2011: 2 ♀♀, 3 ♂♂, 20 juveniles (preserved material RBINSc COP 10.351); 10/07/2012: 48 ♀♀, 12 ♂♂ (preserved material RBINSc COP 10.338);
– Well A5: 16/11/2010: 2♂♂, 4 juveniles (preserved material RBINSc COP 10.350);
– Well C3: 30/10/2011: 1 ♂ (dissected male RBINSc COP 10.368); 14/02/2011: 27 ♀♀, 1 ♂ (preserved material RBINSc COP 10.349).
Female: Body (Fig.
Caudal rami cylindrical with triangular depression along anterior half of medial margin, less than 3 times longer than wide (L/W-ratio 1/2.7–2.8). Anterolateral seta, pinnate, short, without spinules near articulation. Posterolateral element shorter than ramus in holotype (ratio: 0.9/1) ranging from 0.8/1–1.0/1 in paratypes), with long and slender spinules along outer side of the stem, pinnate along the inner side. Spinules present at it insertion with ramus. Medial element longer than ramus (ratio: 1.15–1.25/1) and pinnate. Principal terminal setae with breaking plane and pinnate. Dorsal seta 1.3–1.4 times longer than ramus, and articulating on small basal part. Integument of rami unadorned.
Antennule 11-segmented with typical armament, not reaching to posterolateral edge of cephalothorax. Segment 1 with set of slender spinules and set of short ones (Fig.
Antenna with general appearance and armature as in A. chappuisi (see
Mouthparts as in preceding species. Maxillary basis (Fig.
Leg 1–4 protopodite (Fig.
Leg 1–4 rami 2-segmented with general appearance and armament distribution as in preceding species. Frontal and caudal surface of ramal segments unadorned. Proximal segment of leg 4 endopodite inflated caudally (Fig.
Leg 5 (Fig.
Allocyclops nudus sp. n. A leg 1, frontal view B leg 2, frontal view C leg 4, frontal view D leg 4 protopodite and endopodite, caudal view E leg 3 distal endopodite segment, frontal view F leg 4 endopodite, frontal view (Female RBINSc COP 10.367: A, C, E; Female RBINSc COP 10.334: B, D; male RBINSc COP 10.335: E).
Leg 6 (Fig.
Male: Body narrower than female body (Fig.
Caudal rami cylindrical with short triangular medial depression in anterior half. L/W-ratio: 2.7/1 (allotype, ranging from 2.7–3.0/1 in other male paratypes). Anterolateral seta short, pinnate, without spinules at insertion. Distolateral element shorter than ramus (ratio: 0.8–0.9/1) serrate along outer margin, pinnate along inner one. Spinules at insertion present. Terminal medial element longer and dorsal one longer than ramus:1.15–1.25/1 and 1.25–1.35, respectively. Dorsal seta articulating on basal part. Principal terminal setae with breaking plane and pinnate.
Antennule as described for A. cavicola in
Leg 6 (Fig.
Variability. Except for slight variation (less than 15 µm) in the body length and the L/W ratio of the caudal rami and its armament, no structural differences were observed in the type series nor in the other populations.
at Addrodji, Lokossa District, Mono Couffo drainage basin: Well P2 (details in Table
Female holotype, dissected and mounted on 3 slides, RBINSc COP 10.354; allotype male, dissected and mounted on 2 slides, RBINSc COP 10.356; female paratypes dissected: RBINSc COP 10.355A-C, COP 10.357 A, COP 10. 358 A-C; 14 females and 5 males preserved (RBINSc COP 10.359), all collected 1-12-2012. Additional paratypic material (topotypic), collected 19/06/2012: 185 females, 15 males (preserved material RBINSc COP 10.360).
Named after Nestor G. Sakiti who initiated, in collaboration with Claude Boutin, subterranean biology research in Bénin, and supported the junior author initializing his investigations.
(1) Lokossa, Mono District, Mono-Couffo drainage basin:
– Well P6: 17-06-2012: 1 ♀ (preserved material RBINSc COP 10.364); 06/09/2012: 18 ♀♀ 1 ♂ (preserved material RBINSc COP 10.378); Well PM (about 20 m from well P6, cemented, closed and abandoned, and considered here as identical with well P6): 07/01/2012: 65 ♀♀, 4 ♂♂ (dissected ♀ RBINSc COP 10.370A-C; preserved material RBINSc COP 10.365).
– Well P7: 13-06-2012: 74 ♀♀, 4 ♂♂, 2 juveniles (dissected ♀ RBINSc COP 10.369A-C; preserved material RBINSc 10.363);
– Well P9: 11-06-2012: 116 ♀♀, 11 ♂♂, 9 juveniles (preserved material RBINSc COP 10.361); 30-11-2012: 9 ♀♀, 2 ♂♂ (preserved material RBINSc COP 10.362 ).
(2) Abomey-Calavi District, Ouémé drainage basin:
– Well near Campus: 27-02-2013: 24 ♀♀, 3 ♂♂, 1 juvenile (preserved material RBINSc COP 10.366).
Female: Body (Fig.
Allocyclops sakitii sp. n. A urosome, ventral view B pediger 5 and genital double-somite, in lateral view C leg 1 intercoxal sclerite, frontal view D leg 2 intercoxal sclerite, frontal view E leg 3 intercoxal sclerite, frontal view F leg 4 intercoxal sclerite, frontal view G pediger 5 and urosomite 2 in ventral view (Female RBINSc COP 10.355: A, B; COP 10.358: C–F; male RBINSc COP 10.356: G).
Caudal rami (Fig.
Antennule 11-segmented, not reaching to posterior edge of cephalothorax. Segment 1 with crescent row of spinules in proximal half (Fig.
Antennary general morphology and armature as in A. chappuisi. Frontal surface of coxobasis with a cluster of minute spinules (6–7 spinules) in middle of outer margin and a short transverse set of slender ones in the proximal half bear the abexopodal margin (Fig.
Mandible, maxillule, maxilliped and general aspect of maxilla as in A. cavicola and the preceding species. Claw of maxillary basis blunt, armed with 8–9 large spinules, increasing in width and length medially. Claw shorter than accessory element (Fig.
Leg 1–4 protopodite without ornamentation on praecoxa (Fig.
Allocyclops sakitii sp. n. A leg 1, frontal view B distal endopodite segment of leg 2, frontal view C leg 3, frontal view D Outer region of coxa and basis of leg 3, caudal view E leg 4, frontal view F leg 4 protopodite and endopodite, caudal view G leg 4 endopodite, caudal view (Female RBINSc COP 10.355: A–C, F; COP 10.358: D, E; male RBINSc COP 10.356: G).
Leg 1–4 rami 2-segmented with general appearance and armament distribution as in preceding species (Fig.
Leg 5 (Fig.
Leg 6 (Fig.
Male: General appearance of body as in the female with prosome 1.5 times longer than urosome. The latter with pediger 6 half as long as wide (Fig.
Caudal rami 2.5–2.7 times longer than wide, distolateral element 1–1.15 times longer than caudal rami, and terminal medial element 1.30–1.45 times longer than ramus.
Antennule as described for A. cavicola in
Leg 5 (Fig.
Leg 6 vestige (Fig.
Variability: Apart from slight variation in body length and L/W-ratio of the caudal rami and the setae on the caudal rami, no structural variability was observed.
The cyclopoid genus Allocyclops originally erected by
An unidentified cyclopine attributed to Allocyclops was reported from Morocco (
Among the 4 species described here, A. nudus sp. n. resembles most closely A. chappuisi, type of the genus, and can be easily confounded with the latter. A. nudus sp. n. is however more robust and distinctly larger than A. chappuisi. Females of the former measure 895–962 µm and males 744–817 µm whereas females and males of A. chappuisi measure 720–740 µm and 650 µm, respectively (
As for A. nudus sp. n., A. sakitii sp. n. resembles A. chappuisi in most aspects. A. sakitii sp. n. differs from the latter by the larger body dimensions (♀: 800–875 µm, ♂: 650–695 µm versus ♀: 720–740 µm, ♂: 650 µm), the longer caudal rami (L/W: ♀ 2.50–2.65 versus 2.33) and the longer distolateral element of the caudal rami (> than ramal length versus < than ramus). A. sakitii sp. n. might also be distinguishable from A. chappuisi by the presence of the hairy ornamentation on the lateral margin of pediger 5. However, presence or absence of ornamentation on pediger 5 in A. chappuisi could not be explicitly confirmed because of the condition of the material (see
A. pilosus sp. n. and A. spinifer sp. n. are readily distinguishable from all the other Allocyclops species by the presence of ornamentation on the caudal rami: pilose in A. pilosus sp. n., serrate in A. spinifer sp. n. The pilose structures on the rami of A. pilosus sp. n. remind to some extent the ornamentation on the caudal rami of A. cavicola but is quite different in the former because of the linear arrangement of the setules and their location restricted to the medial margin of the rami versus the dorsal, lateral and medial location, and the more abundant presence of the structures on the rami of A. cavicola.
The morphology of four new species described herein concerts completely with the amended diagnosis of Allocyclops as proposed by
1 | Leg 4 distal exopodite segment with 5 medial setae; Caudal rami with medial seta twice as long as ramus, at most | 2 |
– | Leg 4 distal exopodite segment with 4 medial setae; Caudal rami with medial seta 2.4–2.5 times longer than ramus | A. beadlei |
2 | Caudal rami 3 times longer than wide at the most (between 2.3–3.0/1); Medial seta on caudal rami 1 to 1.5 times as long as ramus, at the most | 3 |
– | Caudal rami more than 3 times longer than wide (generally 3.15/1); Medial seta on caudal rami 2 times as long as ramus | A. cavicola |
3 | Pediger 5 without ornamentation on lateral and ventrolateral margins | 4 |
– | Pediger 5 furnished with either spinules or slender setules on lateral and ventrolateral margins | 6 |
4 | Medial margin of caudal rami with 2 or 3 transverse rows of thin hairs (visible only when rami are observed dorsally); Leg 4 terminal endopodite segment 2.2–2.3 times longer than wide; Distolateral seta longer than caudal rami (1/1.2–1.3) | A. pilosus sp. n. |
– | Medial margin of caudal rami without ornamentation; Leg 4 terminal endopodite segment 2.1 times longer than wide at the most; Distolateral seta as long as caudal rami at the most (1/0.8–1.0) | 5 |
5 | Caudal rami 2.7–2.8 times longer than wide; Dorsal seta on caudal rami 1.5 times longer than ramus; Medial spine of leg 5 1.5 times longer than middle setiform element | A. nudus sp. n. |
– | Caudal rami 2.5 times longer than wide at the most; Dorsal seta on caudal rami less than 1.5 times longer than ramus; Medial spine of leg 5 shorter than middle setiform element | A. chappuisi |
6 | Lateral and ventrolateral margin of pediger 5 with cluster of spinules; Surface of caudal rami and leg 4 ramal segments with diffuse pattern of minute spinules | A. spinifer sp. n. |
– | Lateral and ventrolateral margin of pediger 5 with cluster of fine hair-like setules; surface of caudal rami and leg 4 ramal segments unadorned | A. sakitii sp. n. |
Of the 52 regularly visited wells, 40 (77%) were populated with a copepod fauna: 39 exclusively with Cyclopidae, 1 exclusively with Harpacticoida. In 13 wells, i.e. 3 in the districts of Lokossa, Pobè, and Parakou and the 4 wells at Porto-Novo, representatives of the genus Allocyclops were the only cyclopines found (Tables
Co-occurrence of two species of Allocyclops has been registered in 3 wells: A. nudus sp. n. and A. pilosus sp. n. in A1 and B6 (Parakou district) and A. spinifer sp. n. and A. pilosus sp. n. in well Pb6 (Pobè district).
A. pilosus sp. n. is the most widespread species. Reported from 16 wells, its presence in the northern district of the study area (Parakou) and the southern districts (Pobè, Porto-Novo, Kétou) spans the entire Ouémé drainage basin. A. spinifer sp. n. appears to be restricted to the district of Pobè but was also detected in the occasionally visited well at Zogbodomè. It might be that the species has a preference to the southern and less elevated parts of central Bénin. Pobè and Zogbodomé are located in the Ouémé drainage basin. The occurrence of A. nudus sp. n. is restricted to the district of Parakou, in the northern and most elevated part of the Ouémé drainage basin. Allocyclops sakitii sp. n. occurs only near Lokossa located in the drainage basin flooded by the rivers Mono and Couffo. The presence of the latter species in the occasionally visited well at Abomey-Calavi (located at the western limits of the Ouémé basin) is probably a result of the mixing of the waters from the Ouémé and Mono-Couffo basins in the coastal zone.
Bénin is well endowed with a dense river system flowing from north to south in central and southern Bénin (Ouémé and Mono-Couffo basins). Drainages in northern Bénin (roughly above 10° N) form tributaries of the large West-African rivers and run north-west (Volta basin) and north-east (Niger basin). Rivers are particularly subject of a flashy response to rainfall. They retain most of the year a low water level (large rivers) or disappear completely at the surface (many second rank tributaries). Villages have always been dependent on presence of a sufficient supply of groundwater during the long periods of drought for which human made hang-dug-wells and natural springs in the laterite soils have been the solution.
Laterite soils are subsoils of equatorial forests, the savannahs of humid tropical regions and of the Sahelian steppes (
The hydrodynamic properties of a laterite aquifer are characterized by extreme variations with rapid lateral throughflow in the upper horizon during the wet season, and a narrow range of natural groundwater levels for much of the year. The majority of wells studied herein show this rather shallow to low water level (Tables
Members of the genus Allocyclops display a curious combination of conservative and advanced features. The combination of the large body size, the fully developed mouthparts and the complete armature compliment on the legs is in contrast with the general image of a subterranean cyclopine. The 11-segmented antennules and the 2-segmented rami of the legs however, are in accordance with a subterranean life style. It is assumed (
Thus far, only A. chappuisi has been explicitly reported from a laterite spring (
The study on the biodiversity in Bénin received support from the Global Taxonomic Initiative (GTI: grant Nr. 2014/SO1/ER1.1/16) a DGD-RBINS program. The authors honour C. Boutin (Paris/Toulouse) for his enthusiasm and continuous encouragement of the African scientists to study the biodiversity in subsoil aquifers in their countries. We are also most indebted to both referees, M. Holyńska and S. Karatuğ, for their constructive comments. The junior author likes to express his gratitude to the High Ministry of Education and Scientific Research of Bénin for financing his Ph.D. program. Many thanks go to all members of the team of the Zoology Department of the Abomey-Calavi University.