Four new species of Halicyclops Norman, 1903 (Copepoda, Cyclopoida) from Brazil
Carlos Eduardo Falavigna d a Rocha
Departamenlo de Zoologia, Institulo de Biocic?ncias
-
USP, Cai-ua Posfal 20520, 01000 Sdo Paulo, Brazil
Keywords: taxonomy, estuarine crustaceans, Copepoda, Halicyclops, Neotropical Region, Brazil
Abstract
Four new species of Halicyclops are described and figured from plankton samples of the brackish waters of
Pomonga River, Sergipe State (NE Brazil). The ornamentation of the median apical setae of the furcal rami is
proposed as useful diagnostic character for species distinction.
Introduction
The genus Halicyclops is poorly known in the
Neotropical Region. Four species were reported
from there until now: H. exiguus Kiefer, 1934 and
H. similis Kiefer, 1935 from a brackish lake of
Haiti, H. venezuelaensis Lindberg, 1954 from
Venezuela (Lindberg, 1954) and the Amazonas
River mouth (Cipolli & Carvalho, 1973) and H.
crassicornis Herbst, 1955 from brackish water on
the Brazilian coast (Lagoa Rodrigo de Freitas, Rio
de Janeiro and CananCia mangrove).
The plankton samples analysed here were obtained from the Pomonga River (Sergipe State, NE
Brazil) during the course of a biological survey
sponsored by the Sergipe State Government and
Universidade Federal de Sergipe. River Pomonga
meanders across a flat area. It is more of a canal
running paralelly t o the coast in a direction NE-SE,
connecting the rivers Sergipe and Japaratuba
(Fig. 1). Thus, it is influenced by the tides of the
both river estuaries, which meet near our station 3.
The section under the influence of the Japaratuba
River has depths around 1.0 m at high tide, water
rich in suspension of particulated organic material,
salinities ranging from 1.0 t o 13.0 %o during the
year, high temperatures (25.0-32.0 OC), and absence of mangrove trees on the margins, except near
Hydrobiologia 1 19, 1 0 7 1 I7 ( 1984).
O Dr W . Junk Publishers, Dordrecht. Printed in the Netherlands
the confluence of the waters. The other section is
deeper (2.0-7.0 m) and wider, with more saline waters and extensive, marginal mangrove areas. Fresh
water is provided by rain, some springs along the
canal, principally near the confluence area, and
influx of water from the Japaratuba River. During
the rainy period (March-August) diluted waters
occupy the canal from the confluence of the tides to
the mouth in the Japaratuba. Sometimes the whole
canal is under their influence (I .0 to 5.50/,,, in July,
1978).
Materials and methods
The samples were collected using a conic-cylindric net with nylon gauze Monyl Ny 120 HD, which
was pulled horizontally at the surface during one
minute. The material was fixed with formaline of
4%. Three collecting stations were situated in front
of the boat-piers (1- Barreira d'hgua, 2- Papagaio
and 3- Siriba) and the fourth was near Pedrinhas.
Specimens were measured in lateral position.
The body lengths d o not include the furcal setae.
The following abbreviations have been used in
the descriptive text: MZUSP- Museu de Zoologia,
Universidade de SBo Paulo, Brazil; Tx5- fifth thoracic segment; P I . . . P6- first . . . sixth pair of legs;
Fig. I. Partial map of estuary of Sergipe River showing the
sampling stations in Pomonga River.
B1-coxa; B2- base; En1 . . . En3- first. . . third endopodaljoint; Ex1 . . . Ex3-first. . . thirdexopodal
joint.
The terms 'basal portion' (bp) and 'terminal portion (tp) (Fig. 19) of the median apical setae of
furcal rami refer, the former, to the part between
thejoint on the ramus and the constriction near the
middle of the seta and the latter t o that between this
constriction and the distal tip.
CYCLOPIDAE
Halicyclops Norman, 1903
Halicyclops pilosus sp. n.
(Figs. 2-22)
Type material. Holotype (1 female, MZUSP
5224) and allotype (1 male, MZUSP 5225) from
station 4, Pomonga River, Sergipe State, N E Brazil, 1 June 1978, C.E.F. d a Rocha coll. Paratypes:
2 females and 1 male from station 1 , 2 females and
1 male from station 2, 2 females from station 3,
8 females and 5 males from station 4, same river,
and date, 4 females and 3 males from station 4,
same river, 30 June 1978, C.E.F. d a Rocha coll. Six
paratypes in MZUSP (ref. no. 5226).
Description. Female. Total length 565-610 p m
( N = 9), holotype 570 pm. Prosome:urosome ratio
= 1.5 1-1.79: 1 (N = 9). Cephalothorax (Fig. 2) anteriorly triangular in dorsal view, the greatest width
in the posterior region. First urosomal segment
(Tx5) protruding from prosome. Posterior borders
of this segment and of all prosomal segments
smooth.
Genital segment (Fig. 3) nearly as broad as long
(100:90 pm) in the proximal third, where there are
two lateral, triangular protuberances. P6 inserted
dorso-laterally, constituted at a n inner plumose seta and two reduced spines. Posterior border of this
segment and of the two following urosomal segments denticulate; dorsal portion of the fourth urosoma1 segment slightly expanded. Anal segment
with denticules only ventrally.
Furcal rami (Figs. 3, 4) long as wide (19 pm).
Lateral seta plumose, about 1.4 times longer than
furca. Dorsal seta plumose, arising from subterminal papilla and 2.7 times the length of ramus. Outer
apical seta plumose, a little longer than ramus
(25 pm). Inner apical seta shorter than ramus and
outermost seta (0.8: 1 and 0.6: 1, respectively). Inner
median apical seta (Fig. 5) with basal portion naked
externally and with few setules on inner margin;
terminal portion with setules progressively more
slender towards the tip of the seta. Outer median
apical seta (Fig. 5) about one third the length of
inner median apical seta, setulose externally and
with short setules internally on terminal portion.
Antennule (Fig. 6) six-jointed, reaching middle
of cephalothorax. Fourth joint about 1.3 times as
long as wide. Antenna and feeding appendages as
on Figs. 7-1 1.
P1-P4 (Figs. 12-15) biramous; each ramus 3jointed. P2 and P 3 differing only in size. B1 with
inner feather-like seta. B2 with outer seta and hairs
on distal edge, some as long as or longer than the
E n l . B2 P1 bearing a n inner spine, 1.22 times
longer than endopod and ornamented as on Fig. 12.
Ex1 with outer spine, inner seta and some external
spinules. Ex2 with outer spine and inner seta. Ex3
PlLP4 with spinal formula 3.4.4.3 and setal formula 5.5.5.5. Exopodal spines of P I and P2 long and
Figs.2 17. Haliryclopspilosus sp. n. Female 2. habitus, dorsal; 3. urosome, dorsal; 4. furcal ramus, lateral; 5. median apical furcal setae,
dorsal; 6. antennule; 7. antenna; 8. mandibule; 9. rnaxillule; 10. maxilla; 11. maxilliped; 12. PI: 13. intercoxal plate and seta on BI P I ; 14.
P2 and intercoxal plate; 15. P4 and intercoxal plate; 16. En3 P4; 17. P5.
1 --\
Figs. 18-22. Halicyclopspilosus sp. n. Male 18. habitus, dorsal;
19. posterior end of urosome, dorsal(bp = basal portion and tp =
terminal portion of median apical furcal setae); 20. En2 and En3
P4 (allotype); 21. Endopod of P4 (another exemplar); 22. anterior part of urosome showing P5 and P6, lateral.
slender. Innermost spine of Ex3 P3-P4 plumose
internally and spinulose externally. En1 of all legs
with inner seta; En2 with inner seta on P I and 2
setae on P2-P4. En1 and En2 with short hairs on
distal edges. En3 P1 with 2 spines and 4 setae; En3
P2-P3 with 3 spines and 3 setae, the proximal one
modified (plumose proximally and setulose terminally). En3 P4 (Fig. 16) 1.7 times as long as wide,
with 3 spinulose spines and 2 spiniform setae. Inner
apical spine 1.33 times the length of joint and 1.62
times that of outer apical spine. Inner setae equal in
length, plumose proximally and spinulose terminally, both not reaching the tip of innermost spine. In
all legs the normal setae are densely plumose. Protuberances of distal edge of intercoxal plate of P1
with hairs (Fig. 13).
Terminal joint of P5 (Fig. 17) twice longer than
wide (39:20 pm), with 3 finely serrated spines and a
plumose seta. Innermost spine slightly longer than
terminal joint and outer apical spine (I. 15:1).
Male. Total length 425-466 pm (N = 9), allotype
425 pm. Prosome:urosome ratio = 1.48- 1.8 1:1 ( N =
9).
Cephalothorax (Fig. 18) rounded anteriorly. Urosome with one more segment than in female; posterior borders of segments as in female.
Furcal rami (Fig. 19) slightly wider than long
(18:15 pm). Setae similar in length to those in female. Outer apical seta about twice as long as the
innermost one. Ornamentation of median apical
setae with some differences relatively to those of
female, mainly on its basal portions (Fig. 19).
Antennule prehensile and geniculate, 13-jointed.
Antennae, feeding appendages and P 1-P3 like
those in female.
Endopod P4 (Fig. 20) with proximal seta on En2
and inner setae on En3 similarly modified; in one
paratype, the seta on En2 was plumose, although a
little shorter and thicker (Fig. 21). En3 1.5 times as
long as wide (30:20 pm). Inner apical spine 1.5
times longer than En3 and outer spine.
Terminal joint of P5 (Fig. 22) 1.53 times as long
as wide, with 3 serrated spines and 2 finely plumose
setae. Measurements of spines (from inner to
outer): 40, 33 and 19 pm.
P6 (Fig. 22) with an inner serrate spine, as long
as outer seta (39 pm), both reaching beyond the
posterior border of third urosomal segment. Median seta slender, measuring a half the length of the
outer one.
Etymology. The specific name (from the Latin
'pilosus', hairy) refers to the abundance of hairs on
swimming legs.
Habitat. This species occurred all along the Pomonga River. Salinity and temperature ranges were
1.0-7.4 %, and 27.5-29.0 O C , respectively.
Differential diagnosis. H. pilosus resembles H.
pilifer Lindberg, 1949 from India in the morphology of P5, En3 P4 and in the ornamentation of the
median apical setae of the furcal rami. The main
differences are in the length width ratio of the furcal
rami, the shape of the genital segment and in the
length ratio between the inner and the outer apical
furcal setae.
The abundance of hairs on the swimming legs is
only comparable to that observed by Burckhardt
(19 13) in H. sinensis Kiefer, 1928.
The spine on B2 P1 of H. pilosus is the longest
one known for the genus. Three other species (H.
blachei Lindberg, 1952; H. coulli Herbst, 1977 and
H. fosteri Wilson, 1958) have long spines on their
B2 P I , but not reaching beyond length of the endopod.
Halicyclops ovatus sp. n.
(Figs. 23-40)
Type Material. Holotype (1 female, MZUSP
5227) and allotype (1 male, MZUSP 5228) and 9
paratypes from station 3, Pomonga River, Sergipe
State, NE Brazil, 1 June 1978, C.E.F. da Rocha
coll.
Description. Female. Total length 390-490 p m
( N = 5), holotype 435 pm. Prosome:urosome ratio
= 1.88-2.25: 1 (N = 5). Cephalothorax (Fig. 23)
rounded anteriorly. First urosomal segment (Tx5)
slightly broader than genital segment and not protruding much from prosome. Posterior edges of all
body segments smooth.
Genital segment (Fig. 24) cylindric, slightly wider
than long (70-63 pm) and without lateral protrusions. P6 (Fig. 25) with an inner plumose seta
(smaller than the half of inner spine of P5) and two
outer spinules.
Anal segment (Fig. 26) medially almost divided
into two parts, each one with a row of subterminal
spinules on the ventral surface; the outer spinules
smaller than the inner ones.
Furcal rami (Fig. 26) short, longer than wide
(20: 16 pm), with a ventral row of subterminal spinules. Lateral seta plumose, inserted dorso-laterally
on the middle of ramus, and as long as this. Dorsal
seta plumose too, arising from subterminal papilla
and twice the length of ramus. Outer apical seta
plumose, a little longer than ramus (25 pm), subterminally placed. Inner apical seta slender and
smooth, its length about one-half that of ramus and
0.4 times that of outer apical seta. Inner median
apical seta (Fig. 27) about 2.3 times as long as outer
one; its basal portion naked; the terminal one is
proximally spinulose on both sides, and plumose
apically. Outer median apical seta (Fig. 27) with
basal portion with external edge sparsely spinulose
and terminal portion setulose in both sides.
Antennule (Fig. 28) 6-jointed, reaching beyond
middle of cephalothorax when bent backwards; its
fourth joint about 2.9 times as long as wide. Antenna and feeding appendages as on Figs. 29-33.
P1LP4 (Figs. 34-36) biramous; each ramus 3jointed. P2 and P3 differing only in size. B1 with
inner seta. B2 with outer seta. B2 P I bearing an
inner spine extending beyond En1 and ornamented
as in Figure 34. Ex3 PILP4 with spinal formula
3.4.4.3 and setal formula 5.5.5.5. En3 P2-P3 (Fig.
35) with 3 spines and 3 setae, the proximal one
modified (plumose on the base and serrate on
terminal half). En3 P4 (Fig. 36) about 1.5 times as
long as broad with 2 inner modified setae and 3
spines. Inner apical spine little longer than respective joint (1.12: 1) and at least twice the length of the
outer one (28: 12 pm). Setae smooth on basal portion and serrate on terminal half; distal seta longer
and stronger than proximal one, reaching beyond
the apex of inner apical spine. Free edge of intercoxal plate of P1 with 2 naked protuberances; in P4
it is convex in its median portion.
Terminal segment of P5 (Fig. 37) almost square,
as long as wide (20 pm), with 3 slender, naked
spines and a seta, similar in thickness t o the spines.
Measurements of spines (from inner to outer): 35,30
and 30 pm.
Male. Body shape (Fig. 38) similar to that of
female. Total length 390-435 p m (N = 6), allotype
395 pm. Prosome:urosome ratio = 1.75-2.29: 1 (N =
6).
Furcal rami as long as wide. Setae similar in
length and ornamentation t o those in female. Outer
apical seta about 3 times as long as inner one.
Antennule prehensile and geniculate, with thirteen joints. The other cephalothoracic appendages
and swimming legs as in female, except the En2 P4
(Fig. 39), whose proximal inner seta is modified as
the setae of En3 P4 of female.
Terminal joint of P5 (Fig. 40) as long as wide,
with 2 plumose setae and 3 naked spines. The inner
apical spine is longer than other spines and setae
Figs. 23-40. Halicyclops ovatus sp. n. Female 23. habitus, dorsal; 24. urosome, dorsal; 25. P6; 26. posterior end of urosome, ventral; 27.
furcal ramus with only the median apical setae; 28. antennule; 29. antenna; 30. mandibule; 3 I. maxillule; 32. maxilla; 33. maxilliped; 34.
PI and intercoxal plate; 35. P3 and intercoxal plate; 36. P4 and intercoxal plate; 37. P5. Male 38. habitus, dorsal; 39. endopod of P4; 40.
anterior part of urosome with P5 and P6, ventro-lateral.
and about twice the length of the joint. Measurements of spines (from inner to outer): 36, 29 and
22 p m .
P6 (Fig. 40) with a naked spine and two plumose
setae, the outermost 1.5 times the lenght of median
one.
E t ~ ~ m o l o gThe
~ . . specific name (from the Latin
'ovatus', oval) refers t o the shape of prosome of
these animals.
Habitat. H. ovatus is known only from the typelocality. It was found in brown-coloured water,
with 2.2 (XI,, salinity and 29.0 "C temperature.
D(fferentia1 diagnosis. H. ovatus resembles H.
incognitus Herbst, 1962 from the coast of Bretagne,
France, in the general shape of body, size of genital
segment and posterior edges of urosomal segments.
It differs from this and other species of the genus by
the structure of En3 P4, mainly in the length of
distal inner seta, the P5, the shape of the free edge of
the intercoxal plate of the P4 and the ornamentation of the median apical setae of the furcal rami.
Ha/iqj,clops verae sp. n.
(Figs. 41-55)
T l p e material. Holotype (1 female, M Z U S P
5229) from station 2, Pomonga River, Sergipe
State, NE Brazil, 1 J u n e 1978 and a paratype (1
female) from station 1, same river and date, C.E.F.
d a Rocha coll.
Ilesc~ription.Female. Total length 643 pm, paratype 672 p m . Cephalothorax (Fig. 41) flattened
frontally. First segment of urosome (Tx5) as wide
as the genital one, and protruding from the prosome. Posterior edges of prosomal segments
smooth.
Genital segment (Fig. 42) with two anterior bilobed bulges; measured here, the segment is a s long
as wide. Posterior border of urosomal segments
ventrally denticulate; fourth urosomal segment
with denticules also o n dorsal side. P6 as o n Fig. 43.
Furcal rami (Fig. 42) a s long a s wide (20 pm).
Lateral seta plumose, about 1.5 times longer than
ramus. Dorsal seta 4.3 times the length of ramus.
plumose and inserted o n a subterminal papilla protruding beyond the apex of ramus. Outer apical seta
longer than innermost (50: 10 pm); this is naked and
spiniform. Inner median apical seta (Fig. 44) with
basal portion naked proximally and sparsely spinulose distally; terminal portion with numerous spin-
ules proximally and plumose apically. Outer media n apical seta (Fig. 44) with few external spinules on
basal portion and numerous external spinules on
terminal portion; the apex of the seta is plumose on
its internal margin.
Antennule (Fig. 45) 6-jointed, reaching beyond
half of cephalothorax; length of fourth joint 2.4
times the width. Antenna and feeding appendages
as on Figures 46 t o 50.
P 1 --P4(Figs. 5 1-53) biramous; each ramus with 3
joints. P2 and P3 differing only in size. B1 with
inner seta. B2 with outer seta; only o n PI with a n
inner spine ornamented near the base and reaching
the second-half of En2. Ex3 P l L P 4 with spinal
formula 3.4.4.3. En3 P2-P3 with proximal seta spiniform and spinulose. En3 P4 (Fig. 54) 1.67 times as
long as wide (42:28 pm), with the inner setae similar
t o the spines. Distal seta extending beyond innermost spine apex. Inner apical spine (59 p m ) about
1.4 times the length of segment and 1.3 times that of
outer apical spine.
P5 (Fig. 55) with terminal segment nearly as long
a s wide (30:27 pm), bearing three finely serrate
subequal spines and one long seta (68 pm), about
twice the length of innermost spine.
Ovigerous female (paratype) with 5 eggs in each
sac.
Malt.. Unknown.
E t ~ ~ r n o l oThe
g ~ ~specific
.
name is in memory of Vera
Lucia d a Souza Cruz, a friend and collaborator a t
U niversidade Federal de Sergipe.
Habitat. This species was found in the part influenced by Sergipe River tidal current, a t 7.4 and
3.8 (X,,,salinitiesandatatemperatureof28.0 "C. Both
margins with adjacent extensive mangrove areas.
Diferential diagnosis. H. verae differs from all
the species in the genus by the ornamentation of the
median apical setae of the furcal rami and the armature of En3 P4, with its two inner setae similar t o the
spines. Until now only species with one modified
seta were known. In H. ,japonicu.s Ito, 1956, H.
sinensis Kiefer, 1928 ( a p u d Burckhardt, 1913) and
H. oraeeburnensis Lindberg, 1957 this is the distal
seta, while in H. higoensis Ito, 1957 this is the
proximal one. T h e new species resembles H. pondoensis Wooldridge, 1977 in the length ratio of the
spines and seta of P5, however it differs in the form
of the terminal joint of P5 and in the shape of the
frontal regions of the cephalothorax.
Figs. 41-55. Ha1ic:vclops verae sp. n. Female 41. habitus, dorsal; 42. urosome, dorsal; 43. P6; 44. median apical furcal setae, dorsal; 45.
antennule; 46. antenna; 47. mandibule; 48. maxillule; 49. maxilla; 50. maxilliped; 51. PI; 52. P2; 53. P4; 54. En3 P4; 55. P5.
Haliqclops paradenticulatus sp. n.
(Figs. 56-60)
T,,pe material. Holotype (1 female, M Z U S P
5230) from station 3, Pomonga River, Sergipe
State, N E Brazil, 1 J u n e 1978, C.E.F. d a Rocha
coll.
Dr.sc~ription.Female. Total length 415 pm. Prosome (Fig. 56) rounded anteriorly, with major
width in the posterior region of the cephalothorax.
Third metasomal segment with a stout spinule at
the middle of each lateral margin. Posterior border
of all body segments with denticules. Dorsal margin
of the third abdominal segment (Fig. 57) with coalescent denticules partially hiding the anal operculum.
Genital segment (Fig. 56) wider than long
(83:70 pm) with two slightly blunt dorso-lateral
protuberances in the proximal portion.
Furcal rami (Fig. 56) as long as wide (18 pm).
Lateral seta plumose and nearly equal in length to
the ramus. Outermost apical seta plumose, about
twice the length of the ramus and 2.8 times longer
than innermost seta. Median apical and dorsal setae were broken in the specimen.
Antennule (Fig. 58) with six joints, the fourth one
Figs. 56 60. Halic~~~clopsparadenricularu~
sp. n. Female 56. habitus, dorsal; 57. anal area, dorsal; 58. antennule: 59. P4 (exopodal setae
removed); 60. anterior part of urosome with P5. lateral.
the longest, being 2.7 times longer than wide. The
distal tip of the antennule reaches beyond the middle of the cephalothorax.
PlLP4 biramous; each ramus 3-jointed. Ex3 with
spinal formula 3.4.4.3. En3 P4 (Fig. 59) nearly as
long as wide (23:2 1 pm), with 3 spinulose spines in
the median portion, and two inner spiniform setae.
Inner apical spine little longer than the outer one
38:30 p m = 1.27: 1) and 1.65 times the length of
segment. Setae plumose on basal third and setulose
distally, with a sharp, smooth point. Distal seta
extending little beyond the apex of innermost spine.
Inner proximal setae of En2 P4 (Fig. 59) and En3
P2-P3 like setae of En3 P4.
P5 (Fig. 60) with terminal segment slightly shorter than wide (25:28 pm). bearing some spinules on
the lateral margins. Innermost spine the longest
(40 pm) and about 1.5 times the length of segment.
Other spines equal in length (28 pm). Seta plumose
and shorter than innermost spine.
Male. Unknown.
Etymology. The specific name 'paradenticulatus'
refers t o the similarity of this species with H. denticulatus Kiefer, 1960.
Habitat. H. paradenticulatus occurred in the
same area as H. ovatus sp. n.
Differential diagnosis. H. paradenticulatus is
very closely related t o H. denticulatus by its dorsal
median hyaline denticules of the posterior border of
the fourth urosomal segment, the shape of the terminal joint of P 5 and the ornamentation of its
spines. T h e main differences are in the length ratios
between the spines of En3 P4 and P5, the length
width ratio of the furcal rami(2: 1 t o H. denticulatus
and I: 1 t o H. paradenticulatus) and the morphology of the genital segment.
The presence of denticules on the posterior edges
of the prosomal segments has been reported in H.
denticulatus and H. stocki Herbst, 1962. However,
only H. paradenticulatus has such structures on the
border of all segments. In H. denticulatus they are
lacking on the cephalothorax border and in H.
stocki on the last prosomal and first urosomal segments.
Taxonomic remarks
Wilson (1958) emphasized the necessity of more
complete descriptions for the better identification
of the ~ & y l o ~species,
s
suggesting that features
not mentioned in the lists of diagnostic characteristics organized by Kiefer (1936) and Lindberg (1949)
should be included in the descriptions.
Here I call attention to the importance of the
apical median setae of the furcal rami. Their ornamentation is peculiar in three of the species studied
here, and can be similar in males and females of the
same species (H. ovatus) or slightly different (H.
pilosus). In the known species, these setae have
been satisfactorily described only in H. pondoensis,
H. sinensis, H. pilifer and H.fosteri. Thus, I suggest
that they be examined with more detail in future
descriptions.
Acknowledgements
I thank Dr. T . K. S. Bjornberg, Dr. A. F. Campaner, Dr. M. A. J. de Carvalho and Dr. F. D. P o r
for their suggestions concerning this research. I a m
indebted to the AdministracTio Estadual d o Meio
Ambiente, Aracaju, Sergipe for financing the trips.
Additional financial support was provided by the
Conselho Nacional de Desenvolvimento Cientifico
e Tecnologico (Proc. no. 100215-80).
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Received 20 September 1982; in revised form 26 January 19x3:
accepted 7 Februarq 1984.