BULLETIN OF MARINE SCIENCE, 67(1): 449–463, 2000
NEW TAXA PAPER
EUNICE AND PALOLA (EUNICIDAE: POLYCHAETA) FROM THE
EASTERN BRAZILIAN COAST (13°00'–22°30'S)
Joana Zanol, Paulo Cesar Paiva and Fabiano da Silva Attolini
ABSTRACT
Seventeen species of Eunicidae belonging to the genera Eunice and Palola were found
on the eastern Brazilian Coast. Among them are two new species: Eunice marcusi and
Palola brasiliensis. The coast line is dominated by calcareous bottoms and its fauna is
one of the poorest known in the western Atlantic.
The eastern Brazilian coast polychaete fauna is among the least known in the western
Atlantic. The coast (ca 13°00'–22°30'S) is dominated by biogenic calcareous bottoms of
living and dead algae, coral reefs and other biodetritic fragments (Lana, 1996). This substrate is suitable for reef-boring organisms, such as Eunicidae which is the dominant
polychaete family in the area. Despite their large body size and the large number of characters, eunicid taxonomy is quite problematic. Several original descriptions were rather
brief, leading to a great number of indeterminable species. Fauchald (1992a,b) reviewed
the genera Eunice and Palola and redescribed them in a more realistic framework that
allowed for the identification of many rare species previously reported as synonyms of
‘well known’ species. Hence, in this study, of the total of 17 species recorded, two are
new: Eunice marcusi and Palola brasiliensis, and many are known only from original
description or very few samples.
MATERIALS AND METHODS
Material used for this study was collected by oceanographic surveys using the vessels: HV AN(Diretoria de Hidrografía e Navegação—Brazilian Navy) and SV A STRO -G AROUPA
(PETROBRÁS). Location and depth of sampling stations are presented in Tables 1, 2 and 3. Sampling gear included Van Veen grabs, dredges and beam trawls. Shallow water samples (intertidal to
10 m) were collected by SCUBA. All samples were sieved (0.5 mm mesh-size) on board ship and
fixed in 10% buffered formalin. All material, including holotypes and paratypes, were deposited at
the Departamento de Zoologia, Instituto de Biologia, Universidade Federal do Rio de Janeiro
(IBUFRJ).
TARES
SYSTEMATICS
Eunice cariboea Grube, 1856
Eunice cariboea Grube, 1856: 57; Fauchald, 1992a: 98–102, figs. 29g–q.
Eunice (Nicidion) cariboea Hartman, 1944: 123–124, pl. 7: figs. 157–163, pl. 8: fig. 178; Nonato
and Luna, 1970: 83, fig. 59.
Nicidion incerta Hansen, 1882: 8, pl. 2: figs. 19–21.
Material Examined.—sta. 25BT, 1 spec.; sta. 35VV, 1 spec.; sta. 10C, 3 spec.; sta. 11C,
2 spec.; sta. 15C, 2 spec.; sta. 16C, 1 spec.; sta. 21C, 3 spec.; sta. 29 C, 3 spec.; sta. 32C,
449
BULLETIN OF MARINE SCIENCE, VOL. 67, NO. 1, 2000
450
Table 1. Station list of the Bacia de Campos Program.
Stations
St. 3W
St. 10W
St. 12W
St. 15W
St. 23W
St. 25W
St. 28W
St. 30W
St. 31W
St. 32W
St. 50W
St. 2S
St. 3S
St. 9S
St. 10S
St. 15S
St. 21S
St. 25S
St. 27S
St. 28S
St. 29S
St. 30S
St. 32S
St. 35S
St. 37S
St. 43S
Depth (m)
20
40
70
25
15
30
60
18
150
40
135
27
29
23
44
30
98
38
48
69
106
20
47
71
110
98
Longitude
21° 20'42"S
21° 41'48"S
21° 42'50"S
22° 10'40"S
22° 22'03"S
22° 23'20"S
22° 24'32"S
22° 35'36"S
22° 54'53"S
22° 40'47"S
23° 37'42"S
21° 21'02"S
21° 21'08"S
21° 40'02"S
21° 41'25"S
22° 10'34"S
22° 06'52"S
22° 23'43"S
22° 24'05"S
22° 25'55"S
22° 25'50"S
22° 35'47"S
22° 40'35"S
22° 55'22"S
23° 03'44"S
23° 16'15"S
Latitude
40° 35'42"W
40° 20'42"W
40° 11'06"W
41° 39'15"W
41° 40'48"W
41° 18'50"W
40° 43'47"W
41° 55'06"W
40° 47'58"W
41° 43'57''W
41° 23'58"W
40° 42'45"W
40° 35'01"W
40° 32'46"W
40° 20'46"W
40° 58'55"W
40° 05'01"W
41° 19'08"W
41° 05'08"W
40° 42'44"W
40° 35'27"W
41° 35'01"W
41° 43'21"W
41° 13'15"W
40° 55'47''W
41° 15'15"W
Sediment
biodetritic calcareous
medium sand
biodetritic calcareous
fine sand
coarse sand
fine sand with biodetritic calcareous
biodetritic calcareous
coarse sand with biodetritic calcareous
biodetritic calcareous
very fine sand
biodetritic calcareous
medium sand
biodetritic calcareous
coarse sand
medium sand
coarse sand
biodetritic calcareous
fine sand with biodetritic calcareous
biodetritic calcareous
biodetritic calcareous
biodetritic calcareous
coarse sand with biodetritic calcareous
medium sand with biodetritic calcareous
biodetritic calcareous
biodetritic calcareous
biodetritic calcareous
1 spec.; sta. 15 W, 1 spec.; sta. 30W, 1 spec.; sta. 32W, 3 spec.; sta. 2S, 1 spec.; Abrolhos10 m, 3 spec.; Abrolhos/Siriba-3 m, 10 spec.; Abrolhos/Redonda-intertidal, 6 spec.
Remarks.—The start of the subacicular hooks in part of this material differs from that
reported previuosly. They begin from setiger 25 to up to setiger 48, showing a greater
variation (26 to 32) than observed by Fauchald (1992a). The only abranchiate species
whose subacicular hooks appear this far posteriorly is E. imogena, also described from
the eastern Brazilian coast by Monro (1924), which has hooks beginning after setiger 50.
All other characters are very similar in all specimens analyzed, including the inflated
body of the anterior region (setigers 9–10), the color pattern and antennal size and width.
Distribution on the Brazilian Coast.—Rio de Janeiro, Espírito Santo, Bahia, Sergipe
and Alagoas States, from intertidal to 105 m, associated with calcareous algae and sand
bottoms.
Eunice donathi Carrera-Parra and Salazar-Vallejo, 1998
Eunice donathi Carrera-Parra and Salazar-Vallejo, 1998: 150, figs. 1f–k.
Material Examined.—sta. 8C, 1 spec.; sta. 18C, 3 spec.; sta. 24C, 2 spec.; sta. 29C, 4 spec.;
sta. 33C, 1 spec.; sta. 35C, 1 spec.; sta. 3S, 1 spec.
ZANOL ET AL.: EUNICIDAE FROM THE EASTERN BRAZILIAN COAST
Table 2. Station list of the REVIZEE Program and locations.
Stations
St. 3BT
St. 21BT
St. 25BT
St. 2D
St. 3D
St. 6D
St. 7D
St. 23D
St. 32D
St. 39D
St. 21VV
St. 33VV
St. 35VV
St. 2C
St. 5C
St. 7C
St. 8C
St. 10C
St. 11C
St. 12C
St. 13C
St. 14C
St. 15C
St. 16C
St. 17C
St. 18C
St. 20C
St. 21C
St. 22C
St. 24C
St. 25C
St. 27C
St. 28C
St. 29C
St. 30C
St. 32C
St. 33C
St. 34C
St. 35C
St. 36C
St. 37C
St. 44C
St. 45C
St. 46C
St. 47C
Depth (m)
82
56
45
82
82
60
60
37
22
90
27
28
25
50
50
58
50
50
50
50
40
66
60
53
55
65
67
55
110
62
65
60
54
58
50
54
55
55
55
52
60
65
125
108
60
Longitude
22° 52'42"S
20° 38'00"S
19° 59'54"S
22° 53'16"S
22° 52'40"S
22° 18'01"S
22° 19'58"S
20° 21'17"S
18° 52'28"S
19° 28'33"S
28° 38'14''S
18° 53'16''S
18° 52'00"S
13° 38'26''S
15° 34'11"S
16° 19'55"S
17° 34'21"S
17° 05'54"S
17° 03'59"S
17° 02'24"S
16° 47'14"S
17° 47'49"S
18° 01'22"S
18° 01'24"S
18° 34'00"S
18° 35'37"S
19° 16'08"S
20° 42'21"S
20° 36'13"S
20° 21'03"S
19° 31'11''S
19° 45'36"S
19° 48'47"S
19° 48'01"S
28° 08'46"S
20° 40'24"S
20° 35'03"S
26° 46'02"S
20° 52'00"S
21° 31'00"S
22° 22'10"S
20° 51'24"S
20° 57'05"S
20° 40'34"S
20° 36'51"S
Latitude
41° 09'15"W
40° 01'00"W
39° 54'09"W
41° 09'15"W
41° 09'15"W
40° 48'50"W
40° 50'54"W
40° 05'49"W
39° 35'25"W
38° 22'28"W
40° 01'18''W
39° 13'52''W
38° 58'00"W
38° 45'40''W
38° 51'36"W
38° 14'39"W
38° 25'21"W
36° 45'08"W
36° 48'28"W
37° 36'26"W
38° 41'33"W
35° 52'50"W
35° 53'28"W
37° 21'55"W
38° 04'00"W
37° 54'45"W
38° 00'54"W
35° 27'25"W
35° 51'25"W
36° 38'14"W
38° 46'06''W
39° 31'36"W
37° 56'33"W
37° 46'22"W
37° 29'06"W
37° 42'37"W
38° 04'55"W
40° 05'59"W
40° 10'00"W
40° 18'00"W
37° 35'31"W
33° 38'34"W
34° 00'20"W
34° 35'22"W
34° 53'39"W
Sediment
sand with biodetritic calcareous
silt with biodetritic calcareous
biodetritic calcareous
mud
sand with biodetritic calcareous
sand with biodetritic calcareous
sand with biodetritic calcareous
calcareous algae
sand with biodetritic calcareous
calcareous mud
calcareous mud
biodedritic calcareous
biodetritic calcareous
biodedritic calcareous
biodetritic calcareous
sand with biodetritic calcareous
biodetritic calcareous
biodetritic calcareous
calcareous algae
calcareous algae
calcareous algae
calcareous algae
calcareous algae
calcareous algae
biodetritic calcareous
biodetritic calcareous
biodetritic calcareous
calcareous algae
calcareous algae
calcareous algae
calcareous algae
calcareous algae
calcareous algae
calcareous algae
biodetritic calcareous
calcareous algae
calcareous algae
calcareous algae
calcareous algae
calcareous algae
calcareous algae
calcareous algae
calcareous algae
sand with biodetritic calcareous
calcareous algae
451
BULLETIN OF MARINE SCIENCE, VOL. 67, NO. 1, 2000
452
Table 3. Direct samplings locations in Abrolhos Archipelago.
Location
Siriba Island
Redonda Island
Coral reef
Depth (m)
3
0−3
10
Longitude
17° 58'07"S
17° 57'57"S
17° 58'S
Latitude
38° 42' 46"W
38° 42'46"W
38° 40'W
Sediment
coral reef debris
under stones
coral reef
Description.—Only anterior fragments 2.5–4 mm wide including setae, 21 to 73 mm
long and up to 166 setigers were available. Prostomium as long as peristomium. Body
reddish with several white spots. Peristomial ring less colored. Setigers 4 and 5 (in one
specimen also setiger 6) with a white crossbar. Antennae smooth, outer lateral antennae
(AI) to the middle of first peristomial ring, inner lateral antennae (AII) to setigers 1-4 and
unpaired median antenna (AIII) to setigers 4–7. Eyes posterior to AI and lateral to AII.
Peristomial cirri ovate and short not reaching the margin of the first peristomial segment.
Mandibles black with white borders. Maxillary formula: Mx I: 1+1, Mx II: 3+3(4), Mx
III: 3(4)+0, Mx IV: 3(4)+6(7), Mx V: 1+1. Left Mx III and IV in a distal arc. Ventral cirri
short, swollen basely after setiger 5 or 6. Branchiae begin from setigers 24–31, present to
the end of fragments. Normally with one filament, but one specimen with two filaments
on setiger 26 and three in setiger 34. Limbate, pectinate and bidentate compound falcigers
present. Pectinate slightly heterodont with ca 11 teeth. Single black, unidentate subacicular
hooks starting in setigers 26 to 31.
Remarks.—The material studied presents some differences from the original description of Carrera-Parra and Salazar-Vallejo (1998). These differences were related mainly
to the start of branchiae (24 to 31 in our material vs 22 in the original description), start of
subacicular hooks (26 to 31 vs 23) and number of branchial filaments (1 to 3 vs 2). Miura
(1986), Orensanz (1990) and Fauchald (1991) discussed how the start of the subacicular
hooks, branchiae and number of branchial filaments depend on the body size. Considering that the original description was based on just one specimen, the observed differences
could be due to ontogenetic variation of these characters or on population variability. The
color pattern is very similar to another fuscus-unidentate species, E. sonorae Fauchald,
1970, but this species has articulate ceratostyles, 6 branchial filaments and antennae much
shorter than those observed in E. donathi.
Distribution on the Brazilian Coast.—E. donathi was known only from the type locality (Quintana Roo, Mexico in the northwestern Caribbean Sea). The distribution is expanded to the Brazilian coast (Bahia, Espírito Santo and Rio de Janeiro States) in calcareous algae bottoms between 20 and 62 m.
Eunice cf. edwinlinkae Carrera-Parra and Salazar-Vallejo, 1998
Eunice edwinlinkae Carrera-Parra and Salazar-Vallejo, 1998: 151, figs. 2A–F
Material Examined.—sta. 13C, 1 spec.; sta. 50W, 1 spec.
Remarks.—E. edwinlinkae belongs to the group of species with flavus-tridentate hooks
and articulate dorsal cirri. It differs from the other species of this group in having very
long antennae, bidentate compound falcigers; in having dorsal cirri much longer than
branchiae and in having up to three branchial filaments. The antennal lengths of the stud-
ZANOL ET AL.: EUNICIDAE FROM THE EASTERN BRAZILIAN COAST
453
ied material varies from the original description, AI to setiger 3 versus setiger 4, AII to
setiger 13 vs setiger 21 and AIII to setiger 26 vs setiger 34.
Distribution on the Brazilian Coast.—E. edwinlinkae was known only from the type
locality in Quintana Roo, Mexico. The distribution is expanded to the Brazilian coast,
between the states of Bahia and Rio de Janeiro, in calcareous algae and biodetritic bottoms between 40 and 135 m.
Eunice filamentosa Grube, 1856
Eunice filamentosa Grube, 1856: 56; Luna, 1980: 175–176; Fauchald, 1992a: 138–140, figs. 45a–g.
?Eunice filamentosa Rullier and Amoureux, 1979: 173.
Material Examined.—sta. 6D, 2 spec.; sta. 7D, 2 spec.; sta. 9S, 2 spec.; sta. 15S, 2
spec.; sta. 27S, 3 spec.; sta. 32S, 3 spec.; Abrolhos Archipelago – 10 m, 1 spec.
Distribution on the Brazilian Coast.—Widely distributed on the coast from São Paulo
to Ceará States, intertidal to 72 m in calcareous algae and sand bottoms.
Eunice fucata Ehlers, 1887
Eunice fucata Ehlers, 1887: 91–93, pl. 25: figs. 8–20
Staurocephalus gregaricus Mayer, 1900: 1.
Mayeria gregaricus Verril, 1900: 650
Material Examined.—sta. 10C, 1 spec.; sta. 11C, 3 spec.; sta. 14C, 3 spec.; sta. 22C, 3
spec.; sta. 47C, 1 spec.; sta. 3W, 2 spec.; sta. 3S, 2 spec.; Abrolhos/Siriba-3 m, 1 spec.
Remarks.—Three complete specimens, 6.5–7.5 mm wide, 120–124 mm long, 219–233
setigers. Body reddish with many discolored dorsal patches. Single unidentate subacicular
hooks starting between setigers 32 to 35, normally light honey-colored. Even though
Fauchald (1992a) remarked that E. fucata has unidentate subacicular hooks with light to
medium brown cores and clear sheaths, the species was included in the fuscus-unidentate
group (Fauchald, 1992a; Carrera-Parra and Salazar-Vallejo, 1998). E. schemacephala
Schmarda (1861), considered indeterminable by Fauchald (1992a), resembles E. fucata
but differs in having black subacicular hooks starting at setiger 40 and branchiae with a
maximum of 8 filaments (Hartman, 1944; Day, 1967; Ibarzabal, 1989). Since the start of
subacicular hooks and number of branchial filaments could be regarded as ontogenetic
variation, the color of the subacicular hooks seems to be the best character to distinguish
between the species.
Distribution on the Brazilian Coast.—E. fucata was described from Florida and is distributed in Caribbean waters where it is known as the Atlantic palolo. The distribution is
expanded to the Brazilian coast (Bahia, Espírito Santo and Rio de Janeiro States) under
stones, associated with calcareous algae from intertidal to 110 m.
454
BULLETIN OF MARINE SCIENCE, VOL. 67, NO. 1, 2000
Eunice marcusi new species
(Fig. 1A–F)
?Leodice sp. (floridana Pourtalès?) Treadwell, 1921: 33–34, figs. 77–84.
Material Examined.—sta. 3BT, 11 spec.; sta. 3W, 2 spec.; sta. 3S, 5 spec.; sta. 10S, 2 spec.
Description.—Holotype (IBUFRJ-0001) complete, 6 mm wide, 66 mm long, 121
setigers. Paratypes (IBUFRJ), only anterior fragments, 2–6 mm wide, 14–74 mm long,
14–110 setigers; one specimen a ripe female. Prostomium much shorter than peristomium,
appearing to be biarticulate with an oblique division (Fig. 1A). Median sulcus deep, visible dorsally and ventrally. Eyes located between AI and AII. Antennae with moniliform
articulations which become drop-shaped distally, AIII with 6 to 16 articulations. AI to the
middle of first peristomial ring-setiger 2, AII to setigers 2–4, AIII to setigers 4–7.
Ceratophores ring-shaped. Peristomial cirri articulate with great size variation reaching,
at maximum length, the front of prostomium. First peristomial ring three to four times
longer than the second and with a distinct lower lip. Separation between rings distinct
dorsally and ventrally. Maxillary formula: Mx I: 1+1, Mx II: 4+4, Mx III: 6(7)+0, Mx IV:
3(5)+8, Mx V: 1+1(2), Mx VI: 1+1. Left Mx III and IV in a distal arc. Mandibles (Fig. 1B)
well calcified, winged shaped with pointed external teeth. Branchiae palmate, shorter
than dorsal cirrus (Fig. 1C), begin at setiger 4 (5 in only one specimen), continuing to
setiger 110 (112). First branchiae with one filament; maximum 3 to 4 filaments between
setigers 6(9) to 17(52) and decreasing to two and one filament toward posterior end. Last
five branchiae smaller (Fig. 1D) and button shaped. Two smaller forms (probably young)
with only one branchial filament. Dorsal cirri moniliform tapering distally. Ventral cirrus
with swollen bases from setiger 7 to 38(42). Limbate setae slender marginally smooth,
pectinate setae heterodont and compound falcigers bidentate (Fig. 1E) with teeth of similar size. Black acicula paired, the last 30 setigers with one of them very tiny which disap-
Figure 1. Eunice marcusi sp. nov.: a, anterior region; b, mandible; c, parapodium 22; d, parapodium
110; e, compound falciger, anterior setiger; f, subacicular hook.
ZANOL ET AL.: EUNICIDAE FROM THE EASTERN BRAZILIAN COAST
455
pears in the last 9 setigers. Subacicular hooks black and bidentate (Fig. 1F) starting in
setigers 19 (26) to the end of the body, one per setiger. One pair of anal cirri with up to 12
articulations.
Remarks.—The species belongs to B-2 group of Fauchald (1970), fuscus-bidentate with
branchiae beginning anterior to setiger 10 and continuing beyond setiger 100. It is close
to E. dubitata, differing by the ending of the most anterior branchiae and the relative
width of the first antennae. Furthermore, in E. marcusi sp. nov. the prostomium has a
distinct oblique division, a character shared with E. aphroditois (Pallas, 1788), E.
scombrinis Quatrefages, 1866, E. kinbergi Ehlers, 1868 and E. sebastiani Nonato, 1965.
Of these species only E. kinbergi has articulated antennae, but with well developed
branchiae (up to 22 filaments) and subacicular hooks starting from setigers 123–135.
Treadwells description of Leodice sp. (floridana Pourtalès) is very similar to our material, including the lower number of branchial filaments and articulated anal cirri, which
are the main differences between his material and E. floridana (Pourtalès, 1867).
Distribution on the Brazilian Coast.—Rio de Janeiro and Espírito Santo States, between 20 and 44 m in calcareous algae and medium sand bottoms.
Etymology.—The species is named for the great zoologist Ernest Marcus, who dedicated almost all his life to the study of Brazilian invertebrate fauna and was responsible
for a whole generation of invertebrate zoologists in Brazil.
Type locality.—Northern coast of Rio de Janeiro, off Cabo de São Tomé (21°41'25"S,
40°20'46"W), 44 m.
Eunice multicylindri Shisko, 1981
Eunice multicylindri Shisko, 1981: 971–973, figs. 2a–d; Fauchald, 1992a: 227–228, figs. 75f–i.
Material Examined.—sta. 2D, 5 spec.; sta. 7D, 3 spec.; sta. 5C, 1specimen; sta. 17C, 1
spec.; sta. 18C, 2 spec.; sta. 3S, 8 spec.; sta. 9S, 1 spec.; sta. 10S, 6 spec.; sta. 35S, 1 spec.
Remarks.—E. multicylindri is closely related to E. unifrons in having antennae with
cylindrical articulations, bidentate compound falcigers and single subacicular hooks. Differences between them are observed in the peristomial cirri (articulated and tapering,
rather than smooth and digitated as in E. unifrons) and in the disposition of the antennae
(AIII isolated by a gap in the former rather than AI isolated by a gap in the latter).
Distribution on the Brazilian Coast.—This is the first record for the Brazilian coast,
from the state of Rio de Janeiro to Bahia, between 20 and 60 m, in biodetritic, medium
sand and coarse sand bottoms. Distribution extended from Southern California to Brazil.
Eunice mutilata Webster, 1884
Eunice mutilata Webster, 1884: 315–316, pl. 9: figs. 36–40; Hartman, 1944: 113–114, pl. 6: figs.
140–141; Fauchald, 1992a: 232–233
Leodice mutilata Treadwell, 1921:30–33, pl. 3: figs. 5–8
Material Examined.—sta. 10C, 1 spec.; sta. 22C, 2 spec.
Remarks.—The 3 specimens studied have paired acicula on the anterior part of the
body and some of the compound falcigers are marginally serrated, characters only found
in the Treadwell (1921) description.
456
BULLETIN OF MARINE SCIENCE, VOL. 67, NO. 1, 2000
Distribution on the Brazilian Coast.—Espírito Santo and Bahia from 50 to 110 m associated to calcareous algae.
Eunice cf. nicidioformis Treadwell, 1906
Eunice nicidioformis Treadwell, 1906: 1169, figs. 49–51; Fauchald, 1992a: 237–238, figs. 79f–m
Material Examined.—sta. 37S, 1 spec.
Remarks.—The single specimen studied is a fragment with only 40 setigers (length, 20
mm). It was not possible to see all characters, such as the end of the branchial region and
the unusual vascularized sleeve which surrounds the base of the notopodial cirri between
setigers 40–50. The large size of the first articulation of the antennae, which extends
through the third setiger on AIII, and subacicular hooks appearing on setiger 31 makes
this specimen different from the Fauchald (1992a) description. Since Fauchald (1992a)
redescribed types from Hawaii (Pacific Ocean), the identification must be considered
highly questionable.
Distribution on the Brazilian Coast.—Rio de Janeiro State at 60 m associated to calcareous algae.
Eunice ornata Andrews, 1891
Eunice ornata Andrews, 1891: 284–285, pl. 13: figs. 6–13; Fauchald, 1992a: 245–246, figs. 81f–o.
Eunice rubra Nonato and Luna, 1970: 81. [not E. rubra Grube, 1856]
Material Examined.—sta. 3D, 2 spec.; sta. 6D, 1 spec.; sta. 25AV, 2 spec.; sta. 39D, 5
spec.; sta. 7C, 2 spec.; sta. 16C, 1 spec.; sta. 27C, 1 spec.; sta. 28C, 3 spec.; sta. 29C, 1
spec.; sta. 2W, 2 spec.; sta. 3W, 5 spec.; sta. 10W, 2 spec.; sta. 3S, 4 spec.; sta. 10S, 18
spec.; sta. 25S, 12 spec.; sta. 28S, 1 spec.; sta. 30S, 2 spec.; sta. 43S, 4 spec.
Remarks.—According to Fauchald (1992a), The unusual structure of the median and
posterior acicula and the change from between isodont pectinate setae in anterior setigers
to heterodont pectinate setae in posterior setigers is characteristic of E. ornata. Differences between the original description and this material were observed. The branchial
filaments were fewer in number, eyes distinct and antennae, in some cases, not distinctly
moniliform.
Distribution on the Brazilian Coast.—Rio de Janeiro, Espírito Santo, Bahia, Sergipe
and Alagoas between 20 and 100 m in calcareous algae and medium sand bottoms.
Eunice roussaei Quatrefages, 1866
Eunice roussaei Quatrefages, 1866: 309–311, pl. 10: fig. 1–4; Fauchald, 1992a: 288, fig. 97.
Eunice gigantea Cuvier 1830: 200
Material Examined.—sta. 23W, 1 spec.
Distribution on the Brazilian Coast.—Rio de Janeiro State at 15 m in coarse sand.
ZANOL ET AL.: EUNICIDAE FROM THE EASTERN BRAZILIAN COAST
457
Eunice stigmatura (Verrill, 1900)
Leodice stigmatura Verrill, 1900: 641–643; Treadwell, 1921: 20–22, figs. 31-40, pl. 1: figs. 10–13.
Eunice vittata Hartman, 1942: 9 [in part, not Nereis vittata Chiaje, 1829]
Eunice stigmatura Fauchald, 1992a: 311–313, figs. 22,23,46,48,106.
Material Examined.—sta. 27C, 4 spec.; sta. 29C, 1 spec.; sta. 34C, 1 spec.; sta. 12W, 2
spec.; sta.21S, 2 spec.; sta. 35S, 2 spec.; sta. 43S, 1 spec.
Remarks.—E. stigmatura has been considered a synonymous of E. vittata. The branchial distribution is however different. E. stigmatura has branchiae to near the posterior
end and E. vittata branchiae do not extend that far.
Distribution on the Brazilian Coast.—Rio de Janeiro and Espírito Santo States, in
biodetritic and sand bottoms between 58 and 100 m.
Eunice thomasiana Augener, 1922
(Fig. 2A–D)
Eunice thomasiana Augener, 1922: 45; Fauchald, 1992a: 317, figs. 108 a–j, tables 24,25
Material Examined.—sta. 25AV, 3 spec.; sta. 23D, 12 spec.; sta. 33VV, 2 spec.; sta.
11C, 2 spec.; sta. 13C, 1 spec.; sta. 14C, 2 spec.; sta. 15C, 3 spec.; sta. 16C, 2 spec.; sta.
17C, 1 spec.; sta. 20C, 2 spec.; sta. 24C, 4 spec.; sta. 29C, 2 spec.; sta. 30C, 1 spec.; sta.
32C, 1 spec.; sta. 33C, 3 spec.; sta. 36C, 2 spec.
Description.—Of the 43 specimens studied only 9 were complete, 2.5–4.5 mm wide
with setae, 41–73 mm long with 79–115 setigers. Prostomium shorter than peristomium.
Setigers 4 with a white crossbar in most specimens, which extends anteriorly through
setiger 1 (Fig. 2A). Antennae evenly spaced in semicircle, length variable, AI to the middle
of first peristomial ring-setiger 5, AII to setiger 1–10, AIII to setiger 2–8. Ceratostyles
slender and moniliform, increasing in length from AI to AIII (in some specimens AII and
AIII are similar). Ceratophores ring shaped. Eyes behind AI. Separation between peristomial segments visible dorsally and ventrally, peristomial cirri articulated extending to the
anterior border of the first peristomial segment. Mandibles white. Ventral cirri swollen
basely from setigers 6 through setiger 21–30. Notopodial cirri digitiform, articulated,
brown crossbars present on articulations. Branchiae palmate from setigers 4 (in two specimens from setigers 3 and 5) to setiger 39–100. Branchiae single filaments; maximum of
4 filaments (in 4 specimens) present from setigers 9 to 34. Limbate setae marginally
finely serrated, pectinate setae heterodont with ca 15 teeth and bidentate compound
falcigers (Fig. 2D) some with marginally serrated shafts. Acicula (Fig. 2B) paired, brown,
darker in last few setigers, tapering to blunt tips and curved slightly dorsally. Subacicular
hooks (Fig. 2C) black and bidentate, single in most setigers, rarely paired; starting setigers
17–27, through end of body.
Remarks.—The studied material showed great variation regarding the size of antennae,
the beginning of subacicular hooks and the termination of branchiae. These variations
may be due to size differences among specimens as demonstrated for a related species by
458
BULLETIN OF MARINE SCIENCE, VOL. 67, NO. 1, 2000
Figure 2. Eunice thomasiana; a, anterior region; b, acicula; c, subacicular hook; d, compound falciger,
anterior setiger.
Fauchald (1991). The original description (Augener,1922) is very brief and lacks detail.
Fauchald (1992a) redescribed Eunice thomasiana based on two syntypes, our material
differs in some details such as maximum number of branchial filaments, the start of
subacicular hooks and the separation between peristomial rings. These differences may
be due to the low number of specimens used in the redescription.
Distribution on the Brazilian Coast.—Rio de Janeiro, Espírito Santo and Bahía between 37 and 67 m in calcareous algae bottoms
Eunice violaceomaculata Ehlers, 1887
Eunice violaceo-maculata Ehlers, 1887: 86–87, pl. 24: figs. 11,12, pl. 25: figs. 1–7; Fauchald,
1992a: 334–337
Material Examined.—sta. 37C, 1 spec.; sta. 31W, 1 spec.; sta. 29S, 1 spec.
Distribution on the Brazilian Coast.—Rio de Janeiro State from 60 to 150 m associated
with calcareous algae.
Eunice cf. websteri Fauchald, 1969
Eunice websteri Fauchald, 1969: 12–14: fig. 6.
Eunice longicirrata Webster, 1884: 318–319, pl. 12: figs. 75-–80; Nonato and Luna, 1970: 80–81,
figs. 60–62.
Leodice longicirrata Treadwell, 1921: 39–40, figs. 107–116, pl. 2: figs. 9–12.
?Eunice biannulata Rullier and Amoureux, 1979: 172 (in part).
Material Examined.—sta. 2C, 1 spec.; sta. 14C, 2 spec.; sta. 25W, 1 spec.; sta. 28W, 3
spec.; sta. 33VV, 1 spec.; sta. 35VV, 2 spec.
Description.—Only anterior fragments (2–4.5 mm) are available. Prostomium deeply
incised. Antennae with cylindrical articulations. AIII with up to 20 articulations (10 on
ZANOL ET AL.: EUNICIDAE FROM THE EASTERN BRAZILIAN COAST
459
one small specimen). Antennae, peristomial cirri and dorsal cirri with narrow brown band
at the joints of the articulations. Articulations are indistinct in anteriormost dorsal cirri,
absent in posterior. Dorsal cirri similar in length to branchial filaments. Peristomial cirri
outreach first peristomial ring anteriorly. Mandibles serrated with 5 teeth (the 3 inner
smaller), strongly calcified. Branchiae pectinate, present from setiger 4 to setiger 34–38,
up to 5 filaments, first and last 6 pairs with single filament. Single subacicular hooks
begin setiger 34–35.
Remarks.—The identification is doubtful since the material examined have branchiae
from setiger 4, instead of 3, with a smaller number of filaments; subacicular hooks appearing anterior to those in Fauchald (1969) redescription. The start of the branchiae on
setiger 4 and the small number of filaments resemble the conditions in E. antillensis
Ehlers, 1887 and E. articulata Ehlers, 1887. However, E. articulata has very long dorsal
cirri and short branchiae with stubby filaments through setiger 55. E. antillensis has the
same distribution of branchiae and subacicular hooks as our material, but has acicula
distally expanded in round tabs and dorsal cirri much longer than the branchial filaments.
The specimens described from the northeastern Brazilian coast as E. longicirrata by Nonato
and Luna (1970) and as E. biannulata (in part) by Rullier and Amoureux (1979) are
considered as junior synonyms of E. websteri. These descriptions also presented some
differences from Fauchald’s (1969) redescription.
Distribution on the Brazilian Coast.—Rio de Janeiro, Espírito Santo, Bahía, Sergipe
and Alagoas States from intertidal to 100 m in calcareous algae bottoms.
Palola brasiliensis new species
(Fig. 3A–F)
Material Examined.—sta. 10C, 3 spec.; sta. 11C, 3 spec.; sta. 15C, 4 spec.; sta. 17C, 2
spec.; sta. 21C, 1 spec.; sta. 21VV, 1 spec.; sta. 24C, 1 spec.; sta. 25C, 2 spec.; sta. 28C, 1
spec.; sta. 29C, 4 spec.; sta. 32D, 1 spec.; sta. 35C, 1 spec.; sta. 37C, 1 spec.; sta. 39D, 1
spec.; sta. 44C, 10 spec.; sta. 45C, 3 spec.; sta. 46C, 1 spec.; sta. 47C, 4 spec.
Description.—Holotype (IBUFRJ-0121) complete specimen 3.5 mm wide, 90 mm long,
259 setigers. Paratypes (IBUFRJ): 2.5 to 3.5 wide, 30 to 90 mm long. Prostomium smaller
than peristomium (Fig. 3A), with shallow median sulcus. Eyes between AI and AII. Antennae wrinkled, arranged in a shallow horseshoe, AII and AIII close together, separated
by a gap from AI. AI to peristomium base-setiger 1, AII to setiger 2–7, AIII to setiger 4–
10. Peristomial cirri reach middle to front of first peristomial ring. Peristomial rings similar in size, separation distinct on all sides. Maxillae II with 4 teeth on the left and 3 on the
right side. Branchiae with single filaments (one specimen with three filaments on one
setiger); where best developed larger than notopodial cirri (Fig. 3B) shorter than half of
the body width, starting on setiger 58–100 and finishing well before end of body, leaving
last 21–172 setigers without branchiae. Ventral cirri from setigers 1–5 digitiform, from
setiger 6–8 to setiger 60–87 bases of ventral cirri inflated (Fig. 3B) forming transverse
ridges, tips tapering; posterior ventral cirri short and tapering. First notopodial cirri almost twice as long as second one. Notopodial cirri digitate, shorter posteriorly. Limbate
setae nearly geniculate (Fig. 3C), tapering abruptly and marginally serrated. Compound
falcigers distinctly bidentate (Figs. 3D,E) with marginally serrated shafts. Setae of anterior and median setigers proportionally larger with shafts less serrated than in posterior
460
BULLETIN OF MARINE SCIENCE, VOL. 67, NO. 1, 2000
Figure 3. Palola brasiliensis sp. nov.: a, anterior region; b, median parapodium; c, limbate setae,
anterior setiger; d, compound falciger, setiger 6; e, compound falciger setiger 188; f, acicula,
setiger 88.
setigers. Compound spinigers and pseudocompound falcigers absent. Acicula single, tapering, distally bent in median setigers (Fig. 3F). Pygidium with two unequal pairs of
cirri, the longer twice the shorter.
Remarks.—Palola brasiliensis sp. nov. is closely related to P. viridis Gray in Stair 1847.
The differences between them include the peristomial width (greater than peristomium in
P. viridis); maxillae II (P. viridis has 3 teeth on left and 2 right); inflated bases of ventral
cirri (from setiger 15 to setiger 160–200 on P. viridis); median acicula (up to 3 acicula per
parapodium on P. viridis); and pygidium rounded lobes on P. viridis.
Distribution on the Brazilian Coast.—Rio de Janeiro, Espírito Santo and Bahia States
between 22 and 125 m in calcareous algae and mud bottoms.
Etymology.—The wide distribution of the species on the eastern Brazilian coast was
the reason of its specific name.
Type locality.—Southern coast of Bahia (17°05'54"S; 36°45'08"W), 50 m.
Palola sp.
?Eunice paloloides Moore, 1904: 246–249, pl. 7, figs. 5–7.
?Palola paloloides Fauchald, 1992b: 1196–1198, figs. 8a–g.
?Palola siciliensis Nonato and Luna, 1970: 86–87.
Material Examined.—sta. 27S, 2 spec.; sta. 32S, 2 spec.; sta. 43S, 1 spec.
Description.—Only anterior fragments, 3 to 3.5 mm wide, 75 to 80 mm long, up to 224
setigers available. Prostomium shorter than peristomium which is distinctly inflated. AI
to the middle of first peristomial ring, AII and AIII to second peristomial ring, AIII slightly
ZANOL ET AL.: EUNICIDAE FROM THE EASTERN BRAZILIAN COAST
461
longer than AII. Left maxillae II with 3 teeth, right with 2 large teeth and a secondary
distal one. Ventral cirri digitiform; becoming basely inflated after setiger 9. First dorsal
cirrus distinctly longer (less than twice) than on next setigers. Branchiae with one filament, shorter than one-half of body width from setigers 115(122) to last setigers present.
Limbate setae curved, smooth, longer than falcigers. Compound falcigers similar throughout fragment, ratio between width/length of blades less than 0.25. Acicula black or dark
brown, usually paired, few setigers with three acicula.
Remarks.—The species could not be completely identified due to the absence of complete specimens. It presents many characters common to P. paloloides including an accessory distal tooth in right Mx II, a character used by Hartman (1944) to distinguish P.
paloloides from P. siciliensis. P. paloloides differs from our material in having longer
branchial filaments and single acicula in anterior setigers. P. siciliensis was the only species in the genus previously reported from the Brazilian coast (Nonato and Luna, 1970;
Rullier and Amoureux, 1979). But, as pointed out by Fauchald (1992b), in a review of the
types of Palola, this species has been reported for all warm-water areas around the world
and it is likely that these reports may represent other species. Furthermore, in the same
review, he observed that the syntypes of P. siciliensis were devoided of eyes, a character
presented in several reports of the species (Fauvel, 1923; Hartman, 1944; Day, 1967;
Nonato and Luna, 1970).
Distribution on the Brazilian Coast.—Rio de Janeiro State in bottoms of sand and
calcareous fragments at 48 m.
ACKNOWLEDGMENTS
Financial support for P.C.P. was provided by CNPq (Process: 420107/97-5), survey support was
provided by REVIZEE/MMA/CIRM and PETROBRÁS.
LITERATURE CITED
Andrews, E. A. 1891. Report upon the Annelida Polychaeta of Beaufort, North Carolina. Proc. U.S.
Nat’l. Mus. 14: 277–302, pls. 12–18.
Augener, H. 1922. Über litorale Polychaeten von Westindien. Ges. naturf. Freunde Berlin, Sitzber.
1922: 38–53.
Carrera-Parra, L. F. and S. I. Sallazar-Vallejo 1998. A new genus and 12 new species of Eunicidae
(Polychaeta) from the Caribbean Sea. J. Mar. Biol. Ass. U.K. 78: 145–182.
Chiaje, S. delle. 1829. Memorie sulla storia e notomia degli animali senza vertebre del Regno di
Napoli. Naples 4: viii + 214 p.
Cuvier, G. 1830. Le Règne animal distribué daprès son organisation pour servir de base a l’histoire
naturelle des animaux et d’introduction a l’anatomie comparée, N. edit., vol. 3. Deterville Libraire
and Crochard Libraire, Paris. 504 p.
Day, J. H. 1967. A monograph on the Polychaeta of Southern Africa Part I. Errantia. Brit. Mus.
(Nat. Hist.) Publ. Lond. 458 p.
Ehlers, E. 1868. Die Borstenwümer, nach systematischen und anatomischen Untersuchungen
dargestellt. Leipizig. Pages 269–278, 12 pls.
________. 1887. Report on the annelids of the dredging expedition of the U.S. coast survey steamer
BLAKE. Mem. Comp. Zool. Harv. 15 :1–335.
Fauchald, K. 1969. A revision of six species of the Flavus-Bidentatus group of Eunice (Eunicidae:
Polychaeta). Smithson. Contrib. Zool. 6: 15 p.
462
BULLETIN OF MARINE SCIENCE, VOL. 67, NO. 1, 2000
__________. 1970. Polychaetous annelids of the families Eunicidae, Lumbrineridae, Iphitimidae,
Arabellidae, Lysaretidae and Dorvilleidae from Western Mexico. Allan Hancock Fdtn. Monogr.
in Mar. Biol., Univ. So. California. 5: 1–335, 27 pls.
__________. 1991. A morphometric study of Eunicid Polychaetes from Belize, western Caribbean
Sea. Ophelia Suppl. 5: 47–53.
__________. 1992a. A review of the genus Eunice (Polychaeta:Eunicidae) Based upon type material. Smithson. Contrib. Zool. 1–523.
__________. 1992b. Review of the types of Palola (Eunicidae: Polychaeta). J. Nat. Hist. 26: 1177–
1225.
Fauvel, P. 1923. Polychètes errantes. Faune de France (Paris) 5: 1–494, 181 pls.
Grube, A. E. 1856. Annulata Örstediana, Enumeratio Annulatorum, quae in itinere per Indiam
occidentalem et American centralem annis 1845–1848 suscepto legit cl. A.S. Öersted, adjectis
speciebus nonnullis a cl. H. Kröyer in itinere ad Americam meridionalem collectis. Vidensk.
Meddr. Dansk Naturh. Foren. I: 44–62.
Gray, J. E. 1847. [untitled] Pages 17–18 in J. B. Stair, ed. An account of palolo, a sea worm eaten in
the Navigator Islands. Proc. Zool. Soc. Lond. 15.
Hansen, G. A. 1882. Recherches sur les Annélides recueillies par M. le professeur Édouard van
Beneden pendant son voyage au Brésil et à la Plata. Mém. Acad. Roy. des Sci. Belg. Bruxelles.
44: 3–29, 7 pls.
Hartman, O. 1942. A Review of the types of Polychaetous Annelids at the Peabody Museum of
Natural History, Yale Univ. Bull. Bingham Oceanogr. Coll. 8: 1–98, 161 figs.
_________. 1944. Poychaetous Annelids. Pt. 5. Eunicea. Allan Hancock Pac. Exped. 10: 1–231,
18 pls.
Ibarzabal, D. R. 1989. Poliquetos de Punta del Este, Isla de la Juventud, Cuba. IV Orden Eunicida.
Poeyana 384: 1–28.
Lana, P. C. 1996. O Bentos da Costa Brasileira: avaliação crítica e levantamento bibliográfico
(1858–1996). FEMAR, Rio de Janeiro. 432 p.
Luna, J. A. C. 1980. Anelideos Poliquetas do Nordeste do Brasil. IV. Poliquetas Bentônicos (Eunicea)
da Operação Canopus. (Nota preliminar). Trab. Oceanogr. Univ. Fed. PE., Recife. 15: 165–184.
Mayer, A. G. 1900. An Atlantic Palolo, Staurocephalus gregaricus. Bull. Mus. Comp. Zool. Harvard
36: 1–14, 3 pls.
Miura, T. 1986. Japanese Polychaetas of the genera Eunice and Euniphysa: taxonomy and branchial
distributions patterns. Publ. Seto. Mar. Biol. Lab. 31(3–6): 269–325.
Monro, C. C. A. 1924. On the Polychaeta collected by H.M.S. ALERT, 1881–1882: Families
Polynoidae, Sigalionidae, and Eunicidae. Zool. J. Linn. Soc. Lond. 36: 37–64, 24 figs.
Moore, J. P. 1904. New Polychaeta from California. Proc. Acad. Nat. Sci. Phila. 56: 484–503, pls.
37–38.
Nonato, E.F. 1965. Eunice sebastiani sp. nov. (Annelida, Polychaeta). Bolm. Inst. Oceanogr. São
Paulo 14: 133–139.
__________ and J. A. C. Luna. 1970. Anelìdeos Poliquetas do nordeste do Brasil. I. Poliquetas
bentônicos da costa de Alagoas e Sergipe. Bolm. Inst. Oceanogr. São Paulo 19: 57–130.
Orensanz, J. M. 1990. The eunicemorph polychaete annelids from Antarctic and Subantarctic seas.
Antarct. Res. Ser. 52: 1–183.
Pallas, P.S. 1788. Marina varia nova et rariora. Nova Acta Acad. Sci. Imp. St. Petersburg. 2: 229–
249.
Pourtalès, L.F. 1867. Contributions to the fauna of the Gulf Stream at great depths. Bull. Mus.
Comp. Zool. Harvard 1: 103–120.
Quatrefages, A. 1866. Histoire naturelle de Annelès marins et d’eau douce: Annélides et Géphyriens.
Paris: Libr. Encycl. de Rôret 3 vols. and atlas, vol. 1, vii + 588 p., vol. 2: 1–336, and vol. 2: 337–
794.
ZANOL ET AL.: EUNICIDAE FROM THE EASTERN BRAZILIAN COAST
463
Rullier, F. and L. Amoureux. 1979. Campagne de la Calypso au large des côtes Atlantiques de
l’Amérique du Sud (1961-1962). I. 33. Annélides Polychètes. Ann. Inst. Oceanogr. 55 (suppl.):
145–206.
Schmarda, L. K. 1861. Neue wirbelose Thiere beobachtet und Gesammelt auf einer Reise un die
Erde 1853 bis1857. 1. Tuerbellarian, Rotatorien und Anneliden 1: viii+1-164, 21 pls.
Shisko, J. F. 1981. Five new polychaetes of the families Eunicidae and Onuphidae , Collected in
1975 and 1976 during the Southern California Baseline Project. Proc. Biol. Soc. Wash. 94:
968–983, 5 figs.
Treadwell, A. L. 1906. Polychaetous Annelids of the Hawaiian Islands Collected by the Steamer
ALBATROSs in 1902. Bull. U.S. Fish. Comm. 23: 1147–1181, 81 figs.
_____________. 1921. Leodicidae of West Indian Region. Carnegie Inst. Wash. Pub. 15: iv+131,
9 pls.
Verrill, A.E. 1900. Addition to the Turbellaria, Nemertina and Annelida of the Bermudas, with
revisions of some New England genera and species. Trans. Conn. Acad. Arts Sci. 10: 595–671,
figs. 9–11, pl. 70.
Webster, H. E. 1884. Annelida from Bermuda. U.S. Nat’l. Mus. Bull. 25: 307–327, 5 pls.
ADDRESSES: (J.Z., P.C.P) Departamento de Zoologia, Instituto de Biologia, Universidade Federal do
Rio de Janeiro, CCS, Bloco A, Ilha do Fundão, 21941-590, Rio de Janeiro, RJ, Brazil; (F.S.A.)
Instituto Oceanográfico,Universidade de São Paulo, Praça do Oceanográfico 191, 05508-900,
São Paulo, SP, Brazil. CORRESPONDING AUTHOR: (J.Z.) E-mail: <mjrzanol@openlink.com.br>.
View publication stats
Paulo Cesar Paiva