Crustaceana 88 (9) 1003-1029
DESCRIPTION OF ELAPHOIDELLA PARAMUNA N. SP.
(CANTHOCAMPTIDAE), A NEW HARPACTICOID
COPEPOD FROM COLOMBIA
BY
SANTIAGO GAVIRIA1,3 ) and DANIELLE DEFAYE2 )
1 ) Department of Limnology and Oceanography, University of Vienna & Technisches Büro für
Biologie, Fred-Raymondgasse 19/2/4, A-1220 Vienna, Austria
2 ) Muséum national d’Histoire Naturelle, Département Milieux et Peuplements aquatiques, CP 53,
61 rue de Buffon, F-75005 Paris, France
ABSTRACT
A new species of harpacticoid copepod was discovered on submerged mosses of a high Andean
lake in Colombia. The diagnostic characters of the male and female canthocamptid Elaphoidella
paramuna n. sp. are based on the chaetotaxy of legs 1 to 4, morphology of leg 5, armature of anal
urosomite, size and armature of anal operculum and morphology, armature and ornamentation of
caudal rami. It belongs to the group X of Lang (1948); its relationships with Elaphoidella pectinata,
E. armata and E. brevifurcata are discussed. The 35 species and subspecies of Elaphoidella known
in the Neotropical region are assigned to groups I, II, VII, VIII and X of Lang’s system (1948).
Identification keys to Colombian Elaphoidella species are provided.
Key words. — Systematics, Neotropical region, Andean lakes, biodiversity, meiofauna
RESUMEN
Una nueva especie de copépodo harpacticoideo fue descubierta en musgos sumergidos de
una laguna altoandina de Colombia. Los caracteres diagnósticos del macho y la hembra del
cantocámptido Elaphoidella paramuna n. sp. se basan en la quetotaxia de las patas 1 a 4, la
morfología de la pata 5, la armadura del urosomito anal, el tamaño y la armadura del opérculo anal,
y la morfología, armadura y ornamentación de la rama caudal. La nueva especie pertenece al grupo
X de Lang (1948); se discute su relación con Elaphoidella pectinata, E. armata y E. brevifurcata.
Las 35 especies y subespecies del género Elaphoidella conocidas en el Neotrópico fueron asignadas
a los grupos I, II, VII, VIII y X del sistema de Lang (1948). Se suministran claves de identificación
de las especies Colombianas del género Elaphoidella.
3 ) Corresponding author; e-mail: santiago.gaviria@gmx.at
© Koninklijke Brill NV, Leiden, 2015
DOI 10.1163/15685403-00003464
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SANTIAGO GAVIRIA & DANIELLE DEFAYE
INTRODUCTION
Copepods of the family Canthocamptidae (about 980 species) (Defaye &
Dussart, 2011; Gaviria-Melo et al., 2015) constitute the most species-rich family of
harpacticoid copepods inhabiting continental water-bodies. Canthocamptids also
include some marine representatives (i.e., Mesochra Boeck, 1865, Cletocamptus
Schmankevitsch, 1875).
Within the family, the genus Elaphoidella includes about 230 species and subspecies, followed by Attheyella (about 100 species and subspecies) (Defaye & Dussart, 2011; Gaviria-Melo et al., 2015), and constitutes the most speciose genus
of the family. Representatives of the genus inhabit all types of water-bodies,
from surface- to groundwaters, as well as semiterrestrial environments. As most
freshwater harpacticoid copepods, they show a benthic mode of life in streams
and lakes, also inhabiting hyporheic, phreatic and groundwaters. Species living
in terrestrial and semi-terrestrial environments inhabit mosses, leaf-litter, humid
soils and phytotelmata. Brancelj (2009) showed that the number of species of
Elaphoidella in most countries varies between 0 and 5, exceeding 10 in 5 European countries (Bulgaria, France, Italy, Romania and Slovenia). Mori & Brancelj
(2008) highlighted that one-third of the known species of Elaphoidella has been
recorded from Europe. These high numbers in Europe are related to the presence
of groundwater species and the large extension of karstic systems, in connection
with the number of taxonomic experts in the region. Recently, efforts to study
the groundwater species of the genus outside Europe have been made: Brancelj
et al. (2010) described Elaphoidella namnaoensis Brancelj, Watiroyram & Sanoamuang, 2010 from a cave in Thailand, and Watiroyram et al. (2015) described two
species, Elaphoidella jaesornensis Watiroyram, Brancelj & Sanoamuang, 2015
and Elaphoidella thailandensis Watiroyram, Brancelj & Sanoamuang, 2015, from
caves in the same country.
Several attempts have been made to find morphological affinities and to establish phylogenetically related groups of species of this very species-rich genus.
Lang (1948) established 10 groups that are still valid. Petkovski & Brancelj
(1988) added an 11th group in the description of Elaphoidella serbica Petkovski
& Brancelj, 1988, and Gaviria (1993) suggested an additional group to accommodate Elaphoidella colombiana Gaviria, 1993. Apostolov (1985) split Elaphoidella
into four genera based on the number of articles of the swimming legs and on the
structure of the fifth leg. Later he created a new genus, Praelaphoidella Apostolov,
1991. These genera have not been accepted yet because no natural groups were
considered and because important morphological characters such as the structures
used for reproduction were disregarded. Moreover, some species were not taken
into consideration (Reid, 1990; Defaye & Dussart, 2011). Nonetheless, the compilation of the species and elaboration of a species list was very valuable at that time
ELAPHOIDELLA PARAMUNA NOV.
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(Reid, 1990; Wells, 2007). The genera cannot be accepted before a revision of the
genus is done (see also Defaye & Dussart, 2011).
In the Neotropical region, 35 species and subspecies of Elaphoidella are known,
belonging to five different groups of Lang defined mainly by the size of the
basoendopod and the chaetotaxy of the fifth pair of legs of the female, the number
of segments of the endopod of the first leg in both male and female, and the
structure of the armament of the last segment of the endopod of the fourth pair
of legs in the male.
Group I: Elaphoidella cabezasi Petkovski, 1982, Elaphoidella jojoi Petkovski,
1982, Elaphoidella quemadoi Petkovski, 1982 and Elaphoidella sabanillae Petkovski, 1982.
Group II: Elaphoidella bidens bidens (Schmeil, 1894), Elaphoidella bidens
subterranea Nogueira M. H., 1959, Elaphoidella grandidieri (Guerne & Richard,
1893), Elaphoidella neotropica Petkovski, 1973, Elaphoidella parvifurcata Petkovski, 1980 and Elaphoidella laciniata (Van Douwe, 1911) (female only).
Group VII: Elaphoidella schubarti Chappuis, 1936, Elaphoidella humboldtii
Löffler, 1963, Elaphoidella botosaneanui Petkovski, 1973, Elaphoidella crenobia Petkovski, 1973, Elaphoidella einslei Petkovski, 1973, Elaphoidella subcrenobia Petkovski, 1980, Elaphoidella turgisetosa Petkovski, 1980, and possibly
Elaphoidella negroensis Kiefer, 1967 and Elaphoidella prohumboldti Petkovski,
1980, both latter species with undescribed males.
Group VIII: Elaphoidella surinamensis (Delachaux, 1924), Elaphoidella malayica Chappuis, 1928, Elaphoidella seweli americana (Chappuis, 1933), Elaphoidella bispina Dussart, 1984, Elaphoidella radkei Reid, 1987, Elaphoidella
paraplesia Kiefer, 1967 and Elaphoidella suarezi Reid, 1987. Elaphoidella malayica originally described from Java, was reported from Martinique (Dussart, 1982).
Group X: Elaphoidella armata (Delachaux, 1917), Elaphoidella pectinata
(Delachaux, 1924), Elaphoidella brevifurcata Chappuis, 1936, Elaphoidella jakobii Nogueira M. H., 1959, Elaphoidella neoarmata Petkovski, 1973, Elaphoidella
karllangi Petkovski, 1973, Elaphoidella synjakobi Petkovski, 1980 and Elaphoidella parajakobii Reid & José, 1987 (female only).
According to Reid & José (1987), Elaphoidella pintoae Reid & Jose, 1987,
described only from a female, shows the most similarities with group II. Nevertheless, due to the differences in chaetotaxy of the swimming legs, it should be placed
in a subgroup of group II until the male is discovered and described.
Elaphoidella colombiana belongs to an additional group that can be named
group XII, as already proposed by Gaviria (1993). Groups I, II, VII, VIII, and X
also include species that are distributed outside the Neotropical region (Karanovic,
2001; Dussart & Defaye, 2011). Within the listed species only two, E. bidens
bidens and E. grandidieri, show a wide distribution in the world, while all others
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SANTIAGO GAVIRIA & DANIELLE DEFAYE
are endemic to their region. Group XI, created in 1988, includes Elaphoidella
serbica Petkovski & Brancelj, 1988 currently only known from Serbia.
The first record of the genus Elaphoidella in Colombia was provided by Löffler
(1972). He reported the presence of a taxon in northern Colombia that he named
Elaphoidella schubarti-group (Löffler, 1972, fig. 5). Unfortunately, the species was
not described and, therefore, no exact definition of the species can be considered.
Years later, Reid (1987) described E. radkei and E. suarezi, collected in outdoor
tanks at the Magdalena Valley in central Colombia. Six years later, a third species,
E. colombiana, was described from a high mountain lake from the Eastern Andean
Cordillera (Gaviria, 1993). During a study on the biodiversity of microcrustaceans
in northwest Colombia (1999-2000), the first author found E. bidens bidens at
Palmitos, Antioquia, in Laguna Cerro del Padre Amaya, and E. grandidieri at
the university campus of the Universidad de Antioquia in Medellín (Gaviria &
Aranguren, 2007). Recently, Fuentes-Reines & Zoppi de Roa (2013) reported E.
grandidieri in Ciénaga Grande de Santa Marta, a Caribbean coastal lagoon. Thus,
five species are known from Colombia, all inhabiting surfacewater bodies, either
in cold waters (E. colombiana) or in warm waters (E. radkei, E. suarezi, E. bidens
bidens and E. grandidieri). The present record includes the description of a new
species from a cold, high mountain lake of the Eastern Andean Cordillera.
MATERIAL AND METHODS
The sample containing harpacticoid copepods was collected at Laguna de
Buitrago, Colombia (for details of the lake, see type locality of the species), using
a handnet of 100 µm mesh size near the banks of the lake. The sample was fixed
with 5% formaldehyde (approx. final concentration).
Specimens were later transferred to lactic acid, measured, dissected in glycerine
and mounted on slides with gelatine-glycerine. Mouthparts of the animals were
dissected using tungsten needles sharpened in an electrolytic bath constituted of
a NaCl solution and a 9 V battery (Camacho & Puch, 1990); “minutiae needles”
were used to dissect appendages. The animals were examined under a Leica DMLB
compound microscope. Illustrations were done using a drawing tube mounted
on the microscope; final plates were elaborated with the Adobe Photoshop CS3
program after scanning the drawings.
The descriptive terminology follows Huys & Boxshall (1991). However, we
used the term “intercoxal plate” (Dussart & Defaye, 2001) to describe the structure
connecting the legs.
Specimens were deposited at the Instituto de Ciencias Naturales, Museo de
Historia Natural, Universidad Nacional de Colombia (ICN-MHN), the Muséum
national d’Histoire naturelle, Paris (MNHN) and the Naturhistorisches Museum
Wien, Vienna (NHMW).
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SYSTEMATIC PART
Order H ARPACTICOIDA Sars, 1903
Family C ANTHOCAMPTIDAE Brady, 1880
Genus Elaphoidella Chappuis, 1928
Elaphoidella paramuna n. sp.
(figs. 1-8)
Material examined.— Holotype, female ICN-MHN-CR 2716, dissected on 3 slides, collected on
13.vi.1989 on submerged Sphagnum moss at Laguna de Buitrago (4°45′ 15′′ N 73°49′ 44′′ W), 3560 m
altitude, 0.81 ha surface area, 3 m mean depth, Chingaza Region, Cundinamarca, Colombia, leg. S.
Gaviria.
The following animals are paratypes, with same locality, date and collector as the holotype:
allotype, male ICN-MHN-CR 2717, dissected on 2 slides; 1 female NHMW 25495, dissected on
3 slides; 1 male NHMW 25496, dissected on 2 slides; 1 female NHMW 25498, dissected on one
slide; 1 female NHMW 25527 undissected on one slide; 1 female and 2 males ICN-MHN-CR 2718,
undissected, in ethanol; 1 female and 2 males NHMW 25497, undissected, in ethanol; and 1 female
MNHN-IU-2013-8002 and 1 male MNHN-IU-2013-8003 dissected, each on one slide.
Etymology.— The species name paramuna is derived from “páramo”, the wet
climate zone above the Andean forest, where the lake inhabited by the species
is located. This region may be the main ecosystem inhabited by the species. The
name is to be treated as an adjective agreeing in gender with the (feminine) generic
name.
Diagnosis.— Elaphoidella of rather small size, less than 450 µm. Body squat.
Somites with dorsal posterior edges smooth in most of their length and only long
setules on sides. Anal urosomite without posterolateral spinules. Anal operculum
well developed, with 9 to 13 (female) and 8 to 10 (male) long spinules reaching
middle of caudal rami. Male and female caudal rami subquadrate, without dorsal
carina and with characteristic row of long and curved spinules on mid-dorsal and
inner lateral surface forming a half-crown. Furcal setae: setae II and III slender
and longer than ramus, seta II inserted on distal quarter of ramus, seta III inserted
apically. Legs 1, 2, 4 and exopod of leg 3 similar on male and female, dimorphism
present in number of armature elements of distal segment of endopod of leg 4
(3 in female, 2 in male), size of inner seta of apical segment of endopod leg 2
(longer in female) and size of third segment of exopod of legs 2 and 3 (longer than
corresponding second segment in female, same size in male).
Female characterized by leg 5 with exopod and basoendopod (inner edge) with
3 and 4 armaments, respectively, basoendopod with inner edge extended, as long as
exopod, with wide gap between basoendopod extension and exopod. Somites with
posterior edges smooth, row of spinules near posterior edges of genital-double
somite (dorsally on each side and laterally) and on urosomite 2 and 3 (dorsally on
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each side, laterally and ventrally); urosomite 4 with 7-8 spinules near basis of each
caudal ramus ventrally.
Male leg 5 with fused basoendopods and without armaments on inner edge,
exopod with 3 armaments. Leg 3 endopod second segment with articulated process
on outer margin, apophysis 2 times as long as third segment, ending in 1 barb, third
segment with 2 setae.
Description of female (figs. 1-4, except 3f).— Length of holotype 443 µm
(excluding caudal setae). Habitus cylindrical with cephalosome slightly wider than
thorax and abdomen, cephalosome with small rostrum and rounded integumentary
window (fig. 1a) located just in the middle. Cephalosome, thorax and genital
double-somite with scarce long sensilla distributed as shown in fig. 1a and 1b.
Somites with posterior edges smooth. Thorax somites without spinules. Genital
double-somite as long as the following two somites, copulatory tube long, opening
of copulatory pore at third quarter of somite (fig. 2c), this somite bearing at each
side of dorsal surface a row of long spinules inserted laterodorsally (fig. 1a, b);
third and fourth urosomites with row of spinules dorsally and laterally distributed,
as in genital double-somite but prolonged ventrally (figs. 1b and 2c). Anal
urosomite with 6 spinules inserted near inner base of each caudal ramus ventrally;
anal operculum well developed, slightly convex, with 9 long spinules reaching half
of caudal rami (fig. 1d).
Caudal ramus (figs. 1d) subquadrate, almost as wide as long (ventrally), without
dorsal carina, seta I absent, seta II and III slender, with similar size, longer than
caudal ramus; seta II inserted at last quarter of caudal ramus, with 4 spinules near
its base (fig. 2d); setae III (outermost seta), IV, V and VI inserted apically; setae IV
and VI about of same length as ramus, short and very thin, the latter longer than
the former; seta V strong and very long (fig. 1c), as long as cephalothorax and
urosomites I to IV together; seta VII (dorsal seta) inserted in third quarter of caudal
ramus, its proximal section compound, composed of two small segments. A row
of 3-4 long-arched spinules inserted on dorsolateral inner surface forming a semicrown on ramus, dorsal spinules of crown inserted anterior to dorsal seta, dorsal
surface of caudal ramus with 2 additional spinules (shorter and broader than crown
spinules) inserted near beginning of crown.
Antennule (fig. 3a): 8-segmented, with setation formula (setae and setules) 1,
8, 5, 1 + aesthetasc (seta and aesthetasc with conjoined bases), 1, 3, 2, 6 and 1 +
aesthetasc (seta and aesthetasc with conjoined bases); both aesthetascs as long as
four distal segments of antennule together.
Antenna (fig. 3b) with allobasis. Exopod 1-segmented, with 2 lateral smooth
setae and 2 apical setae, outer apical seta smooth, ornamentation of inner apical
not observed seta (broken seta). Endopod 1-segmented, outer margin with 5 spines,
ELAPHOIDELLA PARAMUNA NOV.
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Fig. 1. Elaphoidella paramuna n. sp. Female. a, Habitus, dorsal; b, habitus, lateral; c, caudal ramus,
seta V; d, caudal rami, dorsal (for Roman numbers, see text).
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SANTIAGO GAVIRIA & DANIELLE DEFAYE
Fig. 2. Elaphoidella paramuna n. sp. Female. a, Labrum and paragnath; b, mandible; c, abdomen,
ventral; d, caudal ramus, lateral.
2 apicalmost short, inner margin with 1 short spine, apical margin with 1 spine, 1
naked normal seta and 3 geniculated setae of inequal length.
Labrum (fig. 2a) armed with a central row of thin teeth and 1 lateral row at each
side of 5 stronger teeth.
ELAPHOIDELLA PARAMUNA NOV.
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Fig. 3. Elaphoidella paramuna n. sp. Female (a-e, g-h), male (f; paratype NHMW 25498). a,
Antennule; b, antenna; c, maxillule; d, maxilla; e, maxilliped; f, maxilliped (male); g, leg 4; h, leg 5.
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SANTIAGO GAVIRIA & DANIELLE DEFAYE
Fig. 4. Elaphoidella paramuna n. sp. Female. a, Leg 1, anterior; b, leg 2 (arrow indicates bulge on
outer margin); c, leg 2, endopod (paratype NHHM 25495); d, leg 3.
Paragnath (fig. 2a) rectangular, its anterior margin ornamented with small teeth,
with round half-plate. Ventral surface with 1 diagonal row of long setae at each
side.
ELAPHOIDELLA PARAMUNA NOV.
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Mandible (fig. 2b), gnathobase with 6 strong, chitinized teeth. Mandibular palp
long, 2-segmented. Basal segment (basis) with 1 seta, distal segment (endopod)
with 1 lateral seta and 4 apical setae.
Maxillule (fig. 3c) with broad precoxa bearing 1 inner smooth seta inserted near
base of arthrite, arthrite proximally narrow, apically with 5 large spines, spines
strongly chitinized. Coxa with 2 smooth subequal setae. Basis with 3 lateral smooth
elements (spines ?), apical margin with 1 smooth outer seta and 1 large inner seta,
inner seta distally bipinnate.
Maxilla (fig. 3d) with 2 endites, proximal endite with 2 short terminal setae, first
seta unipinnate, second seta smooth; distal endite with 1 seta; basis with claw and
2 setae.
Maxilliped (fig. 3e) (paratype NHMW 25498) prehensile, with strong, unarmed
and unornamented syncoxa, inner margin with distal seta, long basis more than 2
times as long as broad, with 5 spinules on outer margin, 2 tiny spinules on inner
margin and naked apical seta inserted near insertion of claw. Endopod consisting
of slightly curved claw. Seta of syncoxa of maxilliped of holotype lost during
dissection.
Legs 1 and 2 (without terminal setae) distinctly shorter than legs 3 and 4.
Leg 1 (fig. 4a): intercoxal plate unarmed. Coxa with 2 rows of minute spinules
on anterior surface distributed as shown in fig. 4a, and 2 spinules near outer distal
corner. Basis with 1 naked spine on outer margin, 1 naked seta inserted near base
of endopod, row of 3 spinules near base of seta and row of 5 spinules near distal
margin of segment. Exopod 3-segmented; first segment with 1 unipinnate spine
and spinules on outer margin; second segment with expanded distal outer corner,
outer margin with 1 unipinnate spine and spinules, inner margin with 1 naked seta;
third segment with 1 unipinnate spine and spinules on outer margin, apical margin
with 1 unipinnate spine and 1 geniculate seta, inner margin with 1 long geniculate
seta inserted subapically. Endopod 2-segmented, a little longer than exopod; first
segment with 1 naked seta on inner margin and a row of spinules on outer margin;
second segment with a long and a short (spinule ?) naked seta on inner margin,
apical margin with 1 geniculate long seta and 1 unipinnate spine, outer margin
with a row of spinules.
Leg 2 (fig. 4b): intercoxal plate with 4 rows of tiny spinules on anterior surface,
2 of them inserted proximally and 2 of them inserted distally; coxa ornamented
with 2 spinules near distal outer corner. Basis with outer unipinnate spine and
row of 5 spinules near base of exopod and hairs at inner corner. Exopod 3segmented; first segment with 1 unipinnate spine and spinules on outer margin and
1 spinule near apical margin beside outer spine; second segment with expanded
distal outer corner, outer margin armed like first segment and inner margin with 1
naked seta; third segment 2 times as long as second segment, with 1 seta inserted
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SANTIAGO GAVIRIA & DANIELLE DEFAYE
medially and 1 subapical spinule on inner margin, apical margin with 1 inner
long unipinnate seta and 1 outer bipinnate spine (inner margin of spine with few
secondary spinules), outer margin of segment with 1 proximal bulge, 2 unipinnate
spines and spinules. Endopod 2-segmented (hairs not described); first segment with
1 spinule on outer margin (spinule absent in paratype NHMW 25495); second
segment with 2 spinules on outer margin (3 in paratype NHHM 25495), 2 bipinnate
setae on apical margin (innermost seta exceeds by far tip of inner seta of inner
margin of exopod), inner margin with 1 naked seta and 1 spinule (spinule absent
in paratype NHHM 25495). Hairs on inner and outer margin of first segment of
endopod variable in number.
Leg 3 (fig. 4d): intercoxal plate unarmed. Coxa with 2 spinules near outer
margin and 2 spinules near distal outer corner of segment. Basis with thin naked
seta on outer margin. Exopod 3-segmented; first segment broader than other
segments, with unipinnate spine and spinules on outer margin, second segment
with distal outer corner strongly expanded, 1 unipinnate spine and spinules on
outer and near distal margins, inner margin with 1 naked seta; third segment
(excluding expanded corner) 2 times as long as second segment, with 2 unipinnate
spines and spinules on outer margin, apical margin with 1 outer bipinnate spine, 1
apical unipinnate seta and 1 inner unipinnate seta and inner margin with 1 naked
seta, anterior surface of segment with spinules near apical margin. Endopod 2segmented; first segment unarmed, very short; second segment with 2 spinules
and a bipinnate seta on outer margin, apical margin with 1 unipinnate inner seta
and 1 bipinnate outer seta (latter with only 1 secondary setula on outer margin),
inner margin with 2 naked spinules, both spinules of same size and inserted at
the same distance from the first segment. Right leg 3: second segment of endopod
with second spinule of outer margin difficult to see because it is inserted behind
first spinule.
Leg 4 (fig. 3g): intercoxal plate and coxa unarmed. Basis with 1 thin naked
seta on outer margin and 4 spinules on anterior surface near apical outer corner.
Exopod 3-segmented; first segment with naked spine and spinules on outer margin,
anterior surface with spinules near apical outer corner; second segment armed
as first segment, additionally with 1 naked seta on inner margin; third segment
2 times as long as second segment with 1 naked spine, 1 unipinnate spine and
spinules on outer margin, apical margin with 1 bipinnate spine and 1 bipinnate
seta, inner margin with 1 bipinnate seta inserted subapically and a naked median
seta. Endopod bisegmented; first segment very short, second segment ending with
two apical naked setae and one subapical inner naked seta.
Leg 5 (fig. 3h): right and left basoendopods separated, expansion of basoendopod as long as exopod, broad gap between basoendopod and exopod, basoendopod
medial margin with protuberance, basoendopod with 4 major bipinnate armaments,
ELAPHOIDELLA PARAMUNA NOV.
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setae I (innermost) and IV short, setae II and III long, the latter longer than the
former, anterior surface with 2 spinules near insertion of seta II, outer margin of
basoendopod with lateral naked seta. Exopod 2 times as long as broad, with narrow base, innermost seta bipinnate and inserted subapically, medial and outer seta
inserted apically, medial seta long and bipinnate, outer seta short and naked.
Legs 1-4 with following formula of major spines (Roman numerals) and setae
(Arabic numerals).
Leg 1
Leg 2
Leg 3
Leg 4
Coxa
0–0
0–0
0–0
0–0
Basis
I–1
I–0
1–0
1–0
Exopod
I – 0, I – 1, I – II – I
I – 0, I – 1, II – II – 1
I – 0, I – 1, II – I + 1 – 2
I – 0, I – 1, II – I + 1 – 2
Endopod
0 – 1, 0 – I + 1 – 2
0 – 0, 0 – 2 – 1
0 – 0, II – 2 – 1
0 – 0, 0 – 2 – 1
Leg 6 (fig. 2c) represented by a small plate with 1 spinule (difficult to see)
located anterior to genital field.
Male (figs. 5-8): length of allotype 335 µm (excluding caudal setae), habitus (fig. 5a, b) cylindrical, with broadest section at midlength of cephalosome.
Cephalosome with rounded integumentary window, located more anteriorly than
female. Cephalosome and thorax with scarce median and lateral sensilla. Somites
with posterior edges smooth. Thorax somites without spinules. First urosomite on
each side of dorsal surface with a small row of spinules inserted near posterior
edge; second, third and fourth urosomites dorsally with row of spinules inserted
as in first urosomite, this row of spinules prolonged laterally and ventrally; anal
somite with ventrally group of 6 spinules near base of each caudal rami (fig. 6c).
Anal operculum as in female but with 10 long spinules instead of 9.
Caudal ramus (figs. 5c, 6a-c) as in female, except size of setae II and III,
insertion position of spinules of half-crown and additional dorsal spinules; setae II
and III 1.5 to 2 times as long as caudal ramus (ventrally), dorsal spinules of halfcrown inserted more apically than in female (insertion of first spinule of half-crown
more posterior than insertion of dorsal seta), additional short spinules dorsally near
spinule half-crown absent (2 spinules in female) (fig. 6b).
Antennules (fig. 7a-e): 8-segmented, geniculated between third and fourth
segment and between fifth and sixth segment, fourth segment expanded, antennules
with setation formula (observed on paratype NHMW 25496) (fig. 7b-e) 1, 5, 2, 1 +
aesthetasc (seta and aesthetasc with conjoined bases), 0, 0, 0, 5 and 1 + aesthetasc
(seta and aesthetasc with conjoined bases).
Antenna as in female. Mandible, maxillule and maxilla not studied. Maxilliped
(fig. 3f) similar to that of female (fig. 3e) except ornamentation. Syncoxa, outer
margin with bipinnate seta inserted near basis (seta probably broken during
dissection), ventral surface with row of tiny spinules (absent in female), basis outer
1016
SANTIAGO GAVIRIA & DANIELLE DEFAYE
Fig. 5. Elaphoidella paramuna n. sp. Male. a, Habitus, dorsal; b, habitus, lateral; c, caudal ramus,
lateral.
margin with 2 groups of spinules (1 group in female), inner margin unornamented
(with 2 spinules on female).
Leg 1 and 2 (without terminal setae) distinctly shorter than leg 4.
Leg 1 (fig. 8a, b): intercoxal plate armed with 1 small row of spinules at each
side of anterior surface distally. Coxa with 1 spinule on outer margin. Basis with
2 rows of minute spinules on anterior surface inserted near proximal margin, 1
unipinnate spine on outer margin, 1 naked seta inserted on anterior surface near
base of endopod, 3 rows of spinules inserted near distal margin as shown in fig. 8a
and hairs on inner margin. Exopod 3-segmented, first segment the widest, as long
as endopod; segments armed as in female. Endopod 2-segmented; first segment
armed as in female; second segment as in female but both naked setae on inner
margin small and equally sized, apical section of geniculate seta broken (in allotype
ICN-MHN-CR2717 and paratype NHMW 25496).
Leg 2 (fig. 8c): intercoxal plate unarmed. Coxa with 2 spinules on outer margin.
Basis with naked spine on outer margin. Exopod 3-segmented, third segment
ELAPHOIDELLA PARAMUNA NOV.
1017
Fig. 6. Elaphoidella paramuna n. sp. Male. a, Caudal rami, dorsal; b, caudal ramus, lateral (except
proximal section of dorsal seta, setae not shown); c, caudal rami, ventral; d, leg 4.
longer than second segment. First segment armed as in female but spine naked;
second segment with strong expansion of apical outer corner (stronger than in
female), armature as in female but spine naked; third segment with 1 naked
1018
SANTIAGO GAVIRIA & DANIELLE DEFAYE
Fig. 7. Elaphoidella paramuna n. sp. Male. a, Antennula (setation not shown, except at segment
I, and aesthetascs with conjoined setae of segment VIII); b, antennula of paratype NHMW 25496
(geniculations lost during slide preparation); c, d and e, antennulary segments II, III and IV; f, legs 5
and 6.
ELAPHOIDELLA PARAMUNA NOV.
1019
Fig. 8. Elaphoidella paramuna n. sp. Male. a, Leg 1; b, leg 1, exopod, seta inner margin; c, leg 2; d,
leg 3 (arrow indicates bulge on outer margin); e, leg 3, endopod, third segment.
median seta and 1 unipinnate subapical seta on outer margin, apical margin with
1 outer unipinnate spine and 1 inner bipinnate longer seta (inner margin of setae
1020
SANTIAGO GAVIRIA & DANIELLE DEFAYE
represented only by 1 element), anterior surface of segment with 1 row of spinules
near apical margin, inner margin with 1 median naked seta much shorter than
corresponding seta of female. Endopod 2-segmented; first segment broad and
unarmed; second segment with 1 subapical naked seta and 1 tiny spinule on inner
margin, apical margin with 1 unipinnate outer setae and 1 bipinnate inner seta,
inner seta longer than outer seta, not reaching apical margin of exopod, the latter
seta distinctly shorter than corresponding seta of female.
Leg 3 (fig. 8d, e): intercoxal plate as in female. Coxa with 1 spinule on outer
margin. Basis with 1 naked thin seta and 2 spinules on outer margin. Exopod 3segmented; first segment with naked spine on outer margin and a distal rounded
prominence on inner margin; second segment with distal outer corner strongly
expanded, 1 naked spine and spinules on outer margin, inner margin with 1 naked
seta; third segment 1.5 times as long as second segment (not considering expanded
corner), outer margin with 1 naked median spine, 1 unipinnate subapical spine
and 1 single spinule near insertion of each spine, proximal margin of segment with
small bulge near expanded corner of second segment, apical margin with 1 spinule,
an unipinnate outer spine and 1 unipinnate inner seta, the latter considerably
shorter than in female (being only as long as second and third segments together),
inner margin with 1 naked median seta and 1 subapical bipinnate seta, subapical
seta considerably shorter than in female (being as long as first, second, and
half third segments together). Endopod 3-segmented; first segment very short,
unarmed; second segment with apophysis on inner margin reaching median section
of third segment of exopod, ending in a barb, inner margin with a thumb-like
process articulated with segment; third segment with 1 short naked seta and 1
unipinnate seta on apical margin.
Leg 4 (fig. 6d): intercoxal plate quadrate, unarmed. Coxa with spinules on outer
margin. Basis armed as in female. Exopod 3-segmented; first and second segment
armed as in female, second segment with outer distal corner expanded in contrast
to female; third segment 1.5 times as long as second segment (not considering
expanded corner of second segment) with 1 naked outer median spine, 1 unipinnate
subapical spine and spinules on outer margin, apical margin with 1 unipinnate
spine and 1 unipinnate long seta, inner margin with 1 subapical long unipinnate
seta and a naked median seta. Endopod two-segmented as in female but second
segment ending only in an outer short unipinnate seta and a long inner naked seta.
Leg 5 (fig. 7f): right and left basoendopods fused, inner margin unarmed, outer
margin with 1 naked long seta. Exopod as long as broad, inner margin with
bipinnate seta, apical margin with central long bipinnate seta and outer short naked
seta.
Legs 1-4 with following formula of major spines (Roman numerals) and setae
(Arabic numerals). Leg 3, P means process, A means apophysis.
ELAPHOIDELLA PARAMUNA NOV.
Leg 1
Leg 2
Leg 3
Leg 4
Coxa
0–0
0–0
0–0
0–0
Base
I–1
I–0
1–0
1–0
Exopod
I – 0, I – 1, I – I + 1 – 1
I – 0, I – 1, II – I + 1 – 1
I – 0, I – 1, II – I + 1 – 2
I – 0, I – 1, II – I + 1 – 2
1021
Endopod
0 – 1, 0 – I + 1, 2
0 – 0, 0 – 2 – 1
0 – 0, P – A, 0 – 2 – 0
0 – 0, 0 – 2 – 0
Leg 6 (fig. 7f) represented by small plate.
Morphological and meristic variability of males and females.— Females, length
(without caudal setae) 336-443 µm (n = 7). Males, length (without caudal setae)
294-357 µm (n = 6).
Female: Meristic variability was observed in the number of ventral spinules of
the anal segment, varying from 6 (holotype) to 8 (paratype NHMW 25495), in
the number of spinules of the anal operculum varying from 8 (paratype NHMW
25527), 9 (holotype) to 13 (paratype NHMW 25495), and in the number of spinules
of first and second segment of endopod of leg 2: holotype, first segment, outer
margin with 1 spinule; second segment, outer margin with 2 spinules, inner margin
with 1 spinule; paratype NHMW 25495, first segment unarmed, second segment
with 3 spinules on outer margin and without spinules on inner margin (fig. 4c).
Male: Variability was observed in the number of ventral spinules of the anal
somite near base of each caudal ramus, varying from 6 (allotype) to 7 at each side
(paratype NHMW 25496), the number of spinules of the anal operculum, varying
from 8 (paratype NHMW 25496) to 10 (allotype) and the extension of row of
spinules on urosomite 1: paratype NHMW 25496 bears spinules dorsally (at each
side of segment) and laterally, allotype only dorsally.
Remarks.— Elaphoidella paramuna n. sp. fits into the genus Elaphoidella
Chappuis, 1929 particularly by the 8-segmented female antennule, antenna with 1segmented exopodite, mandible with endopodite, natatory legs with 2-segmented
endopods in female and structure of leg 5.
Elaphoidella paramuna n. sp. belongs to the Elaphoidella group X of Lang
(1948). Together with the new one, 12 species of Elaphoidella have been recognized (Petkovski, 1980; Petkovski & Brancelj, 1988; Defaye & Dussart, 2011) as
belonging to this group. Within them, 8 species show a Neotropical distribution.
Species of group X share the following morphological traits. Female: leg 1
with 2-segmented endopod; leg 2 endopod, first segment without (major) armature
elements, second segment with 3 or 4 setae (3 in E. paramuna); leg 3, endopod,
first segment unarmed, second segment with 3, 4 or 5 (major) armaments (3 in E.
paramuna); leg 4, endopod, first segment unarmed, second segment with 1, 2 or 3
setae (3 in E. paramuna); leg 5 with basoendopod expanded at least until half of
exopod segment and armed with 4 (major) armaments, exopod with 3 or 4 (major)
1022
SANTIAGO GAVIRIA & DANIELLE DEFAYE
setae (3 in E. paramuna). Male: leg 2 endopod, second segment with 2 or 3 setae
(3 in E. paramuna); leg 2 and 4 endopods, first segment without setae on inner
margin, leg 4 endopod, second segment with 1, 2 or 3 setae (2 in E. paramuna).
The only difference of the new species to the group diagnosis is the absence
of a dorsal carina on the caudal rami. However, 2 other species of the group, E.
pectinata and E. brevifurcata, also lack such a dorsal carina. Apparently, Lang
(1948) overlooked this character in both taxa during the diagnosis of the group.
Species of Elaphoidella-group X living in the Neotropical region constitute
a phylogenetic group, already recognized by Chappuis (1931), clearly different
from the species of the “Old World”. The former group is characterized by female
leg 5 with basoendopod strongly extended, exceeding the exopod segment, anal
operculum with teeth and caudal rami strongly spinose. At the time of Chappuis,
E. armata and E. pectinata were the only representatives of the group in South
America. Further new descriptions and/or supplementary species records extended
the distribution of the group to Central America and the Caribbean Islands.
Within the Neotropical species of group X, the new species shows the strongest
affinities with E. pectinata (from Costa Rica, French Guyana and Surinam) and
with the Brazilian E. brevifurcata, then with the Peruvian E. armata.
Males and females of E. pectinata, E. armata and E. paramuna share the
following characters: (1) similar segmentation of legs 1, of exopods of legs 2, 3
and 4, as well as of endopods of leg 2; (2) same number of major armaments on
last segment of exopod of leg 2 and (3) same number of setae on exopod of antenna.
Chaetotaxy of exopods of leg 3 and 4 is unknown in E. armata and E. pectinata.
Female of E. brevifurcata is unknown.
Differences between females are shown in table I. The main differences between
E. pectinata and E. armata is the absence of posterolateral spinules on the anal
segment of E. paramuna, the length of the copulatory tube (longer in E. pectinata,
shorter in E. armata), the number of major armaments on exopod of leg 5 (3 in E.
paramuna, 2 in E. pectinata, 4 in E. armata), and form and armature of the caudal
rami (table II).
Differences between males are shown in table III. The main differences of
the new species to E. pectinata, E. brevifurcata and E. armata are the number
of major armaments of leg 5 exopod (2 in E. pectinata and E. brevifurcata, 3
in E. paramuna and E. armata), length of the apophysis of leg 3 (long in E.
brevifurcata and E. pectinata, short in E. paramuna and E. armata) and the
existence or lack of an articulation of the inner process with the second segment of
exopod leg 3 (articulated in E. paramuna and E. pectinata, fused in E. brevifurcata,
unknown in E. armata). The number of armaments of endopod leg 3 is the same
(2) in E. paramuna, E. pectinata and E. brevifurcata and only 1 in E. armata.
Some differences exist in the ornamentation of the somites: E. paramuna and E.
1023
ELAPHOIDELLA PARAMUNA NOV.
TABLE I
Differences between selected females of neotropical species of Elaphoidella-group X
paramuna n. sp. armata
Genital-double segment
Dorsal spinules, each side
Lateral spinules
Position of copulatory pore
Anal segment
Posterolateral spinules
Anal operculum
Size
Edge ornamentation
Number of spines
Legs 2, 3 and 4, endopod, last segment
Number of setae
Leg 4, endopod
Number of segments
Leg 5, exopod
Number of major armaments
Gap between extension of basoendopod and exopod
Basoendopod extension : exopod length ratio
pectinata
Yes
Yes
Third quarter
No
No
Yes
Yes
In the middle Last quarter
No
Yes
Yes
Long
Long spines
10
Short
Cilia
–
Long
Long spines
14
3:4:3
3:?:?
3:4:3
2
2
1a)
3
Broad
1:1
4
Broad
1.5 : 1
2
Narrow
2.5 : 1
Female of Elaphoidella brevifurcata not known.
a) First segment very small, probably overlooked in the description.
brevifurcata share the absence of posterolateral spinules on the anal urosomite
(present in E. pectinata and E. armata) and the presence of dorsal spinules on
each side of the first urosomite (as prolongation of the lateral spinule rows); these
TABLE II
Differences between caudal rami of males and females of selected neotropical species of
Elaphoidella-group X
Sex
Form
Length : width ratio (ventral)
Seta II and III, form
Seta II and III : ramus length ratio
Seta II, insertion
Seta III, insertion
Seta IV, position on apical edge
Dorsal carina
Dorsal subapical teeth
Spinules crown, spinules size
paramuna n. sp. armata
pectinata
brevifurcata
Male and female
Subquadrate
1:1
Slender
>
Last quarter
Apical
Adaxial
No
No
Long
Male and female
Bulb-like
1.4 : 1
Slender
>
Middle
Middle
Central
No
Yes
Long
Male1)
Conical
0.8 : 1
Slender
>
Middle
Apical
Adaxial
No
No
Long
1) Female of Elaphoidella brevifurcata not known.
Male and female
Barrel-like
1.75 : 1
Strong
<
Third quarter
Third quarter
Central
Yes
No
Short
1024
SANTIAGO GAVIRIA & DANIELLE DEFAYE
TABLE III
Differences between males of selected neotropical species of Elaphoidella-group X
Countries found
Thorax, last segment
Dorsal spinules, each side
Lateral spinules
Abdomen, first urosomite
Dorsal spinules, each side
Lateral spinules
Anal segment
Posterolateral spinules
Anal operculum
Form
Edge ornamentation
Number of teeth
Leg 2, endopod
Seta inner margin size
Leg 3, endopod
Second segment,
inner process
Apophysis/third segment
length ratio
Third segment, number
of armaments
Leg 4, endopod
Number of segments
Leg 5, exopod
Number of major armaments
paramuna n. sp. armata
pectinata
brevifurcata
Colombia
Peru, Brazil,
Argentina,
Paraguay
Costa Rica,
Surinam,
French Guyana
Brazil
No
No
No
No
No
No
Yes
Yes
Yes
Yes, dorsal
No
Yes
No
Yes
Yes
Yes, ventral
No
Yes
Yes
No
Long
Long spines
9
Short
Short cilia
–
Long
Long spines
14
Long
Long spines
10
Long
Long
Long
Short
Not fused
–
Not fused
Fused
2:1
2:1
4:1
3.25 : 1
2
1
2
2
2
2
1a)
1a)
3
3
2
2
a) First segment very small, probably overlooked in the description.
dorsal spinules are lacking in E. armata and E. pectinata. The last thoracic somite
lacks spinules in the new species, E. armata and E. pectinata, but they are present
laterally and at each side of the dorsal surface in E. brevifurcata.
The morphology of the anal operculum (except the number of spinules) is not
dimorphic. E. paramuna and E. pectinata show a well-developed anal operculum
with long spines, although E. paramuna has fewer spinules (male 8-10, female 813) than E. pectinata (male and female 14 each). Male of E. brevifurcata shows a
well-developed anal operculum with 10 spines, female of this species is unknown.
In contrast, E. armata shows a small operculum with margin ornamented with
cilia.
Form, armature and ornamentation of the caudal rami are important traits that
differentiate the new species from E. pectinata, E. brevifurcata and E. armata
ELAPHOIDELLA PARAMUNA NOV.
1025
(table II), both male and female. The caudal ramus is subquadrate in E. paramuna
(male and female), barrel-like in E. armata (male and female), bulb-like in E.
pectinata (male and female) and conical in E. brevifurcata (male, female is
unknown). In E. paramuna it is shorter than in E. armata and E. pectinata and
longer than in E. brevifurcata. Particular characters of the caudal rami are the
dorsal carina in E. armata and the dorsal subapical surface tooth in E. pectinata,
both structures absent in the new species.
The lateral setae (seta II and III) are short and strong in E. armata, but long
and slender in E. pectinata, E. brevifurcata and E. paramuna. Their insertion point
on the ramus is different in each species: seta II is inserted more distally in E.
paramuna than in the other 3 species; seta III is inserted on the apical margin of
the ramus (as in E. brevifurca), in contrast to E. pectinata (median insertion on the
ramus) and to E. armata (on the third quarter of the ramus). The long terminal seta
(seta V) is inserted on the central section of the apical margin in E. armata and E.
pectinata, in contrast to an adaxial insertion (proximal to sagittal body axe) in the
new species and in E. brevifurcata.
The half-crown of spinules on the dorsal and inner lateral surface of the caudal
ramus is composed by long curved spinules in the new species, E. pectinata and E.
brevifurcata. In E. armata the spinules are short and straight.
The morphology of the anal operculum, the form and length of the lateral setae,
and the spinule size on the spinules half-crown of the caudal rami define two
subgroups within the neotropical members of group X: a first subgroup: armata
sub-group characterized by strong lateral setae much shorter than caudal rami,
anal operculum with margin ornamented by short teeth termed cilia by Delachaux
(1917), and half-crown with short spinules; this subgroup includes the Peruvian E.
armata, the Brazilian E. jakobii and the Cuban E. neoarmata and E. synjakobii.
Although the Cuban species E. karllangi shows a female with some setae (II, III
and V) of the caudal rami strongly reduced, other characters such as the presence
of a dorsal carina on the rami of male and female and the ornamentation of the male
caudal rami argue for arranging that species within the armata-subgroup. A second
subgroup with slender setae II and III, longer than caudal rami, and spinule crown
with long curved spinules would include E. pectinata, E. brevifurcata and E.
paramuna. An additional character of this subgroup is the absence of a dorsal
carina on the caudal ramus; this carina is present in the armata-subgroup and part
of the diagnosis of group X; therefore, this character should be amended in the
general diagnosis of the group. The armata-subgroup was created by Petkovski
(1980) during the description of the Cuban species. We propose here to designate
the second subgroup defined as above as the pectinata-subgroup.
1026
SANTIAGO GAVIRIA & DANIELLE DEFAYE
CONCLUSIONS
With the actual discovery, the presently known harpacticoid fauna of Colombian
inland waters (Gaviria & Defaye, 2012) consists of 16 species (11 Canthocamptidae, 3 Parastenocarididae, 1 Phyllognathopodidae and 1 Ameiridae).
Diversity of Elaphoidella in Colombia (5 species) is lower than in Cuba (10)
and Brazil (9), but higher than in Surinam (2) and Argentina (2). Only one
species is known from each of the following Neotropical countries: Mexico, Costa
Rica, Venezuela, French Guyana, Ecuador, Peru and Paraguay. The French islands
Bonaire and Martinique are also inhabited by one species each.
Groundwater, benthic habitats of high Andean lakes in Colombia and aquatic
environments of the Magdalena Valley, the eastern Llanos belonging to the
Orinoco Basin, and the Amazonas are potential habitats for harpacticoid copepods
and particularly for Elaphoidella. Other biotopes still poorly investigated are
phytotelmata and semiterrestrial habitats. Thus, their study would no doubt yield
new species of copepods in the country.
I DENTIFICATION K EY
FOR
C OLOMBIAN S PECIES
OF THE
G ENUS Elaphoidella
Males
(male of Elaphoidella suarezi unknown, male of Elaphoidella bidens not known (Wells, 2007))
1. Anal operculum triangular, posterior margin with 4 strong teeth. Leg 4 exopod, segment 3
outer margin, with proximal short, tooth-like spine . . . . . . . . . . . . . . . . Elaphoidella grandidieri
– Anal operculum convex, posterior edge with at least 8 teeth or spines. Leg 4 exopod, segment
3 outer margin, with proximal spine large, modified or not . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2. Anal operculum short, not reaching posterior edge of anal somite, with 18 to 20 spinules. Leg 4
exopod segment 3, outer margin, both spines modified with strong accessory teeth (fused to
spine), giving the spine a “deer antler-like” appearance . . . . . . . . . . . . Elaphoidella colombiana
– Anal operculum large, exceeding posterior margin of anal somite, with less than 11 spines.
Leg 4 exopod, segment 3 outer margin, with both spines (except size of distal spine being
larger than proximal one) not modified . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
3. Anal operculum not reaching midlength of caudal rami, posterior margin with 8 teeth,
outermost teeth at each side smaller than inner teeth. Leg 4 exopod segment 3 short, apical
margin with outer spine short, modified into stout spine with several claw-like teeth on outer
margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Elaphoidella radkei
– Anal operculum exceeding 3/4 length of caudal rami, posterior edge with 8-10 teeth, all teeth
with same length. Leg 4 exopod 3 with normal size, apical margin with outer spine long, not
modified . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Elaphoidella paramuna n. sp.
Females
(female of Elaphoidella radkei unknown)
1. Anal operculum with edge smooth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Elaphoidella suarezi
– Anal operculum with edge ornamented . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2. Anal operculum not reaching posterior edge of anal somite . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3
ELAPHOIDELLA PARAMUNA NOV.
1027
– Anal operculum extending behind 1/3 of length of caudal rami . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
3. Copulatory tube with conical neck. Anal somite without ventral spines above each caudal
ramus. Leg 5, basoendopod slightly protruded, not reaching midlength of exopod . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Elaphoidella bidens
– Copulatory tube with straight slender neck. Anal somite with 1 ventral spine above each caudal
ramus. Leg 5, basoendopod strongly protruded, exceeding midlength of exopod . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Elaphoidella colombiana
4. Anal operculum strongly convex, margin with dense row of short spinules (>30). Caudal ramus
with seta III inserted on lateral margin . . . . . . . . . . . . . . . . . . . . . . . . . . . Elaphoidella grandidieri
– Anal operculum slightly convex, margin with large spinules (<10). Caudal ramus with seta III
inserted on apical margin . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Elaphoidella paramuna n. sp.
ACKNOWLEDGEMENTS
We thank the “Empresa de Acueducto de Bogotá” (Waterworks of the city of
Bogotá) for their support during sampling and to two anonymous reviewers who
read the manuscript sent to Crustaceana. We also thank T. C. Walter (Washington, DC, U.S.A.) for sending literature. M. Stachowitsch (University of Vienna)
reviewed the English text. A research grant to study the material at the “Muséum
national d’Histoire naturelle” MNHN in Paris was assigned to S. Gaviria in 2012.
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First received 30 June 2015.
Final version accepted 11 August 2015.