Progoniada and Goniadella (Annelida: Polychaeta: Goniadidae) from the outer continental shelf and slope off south-eastern Brazil

Abstract

Species of the genera Progoniada and Goniadella are recorded from off the Brazilian coast, from depths of 93 to 808 m. The description of Progoniadaregularis is complemented with new data on the number of chevrons and micrognaths. Goniadellarevizee sp. nov. is described; it differs from the five known species of the genus mainly in having acicular chaetae above the dorsal cirrus, together with 20–21 uniramous chaetigers. The proboscideal papillae of both species are illustrated by scanning electron micrographs.

Full text

Progoniada and Goniadella (Annelida: Polychaeta:
Goniadidae) from the outer continental shelf and slope o¡
south-eastern Brazil
Alexandra Elaine Rizzo* and Antonia Cec|
¤lia Zacagnini Amaral
Departamento de Zoologia, Instituto de Biologia, Universidade Estadual de Campinas^UNICAMP, CP 6109, 13083-970,
Campinas, SP^Brazil *Corresponding author, e-mail: aerizzo@hotmail.com
Species of the genera Progoniada and Goniadella are recorded from o¡ the Brazilian coast, from depths of
93 to 808 m. The description of Progoniada regularis is complemented with new data on the number of
chevrons and micrognaths. Goniadella revizee sp. nov. is described; it di¡ers from the ¢ve known species
of the genus mainly in having acicular chaetae above the dorsal cirrus, together with 20^21 uniramous
chaetigers. The proboscideal papillae of both species are illustrated by scanning electron micrographs.
INTRODUCTION
The family Goniadidae Kinberg, 1865 presently
comprises ten genera with about 73 species (Hilbig, 1994).
Records from the Brazilian coast include 16 species
belonging to the genera Glycinde,Goniada,Goniadides,
Progoniada and Ophioglycera (Hartman, 1965; Steiner,
2000). The complete list of Brazilian coastal species is
available on the site http://www.ib.unicamp.br/pesquisa/
projetos/biota/bentos___marinho/index.htm. The present
contribution adds new records of species of Progoniada and
Goniadella from o¡ the south-eastern coast of Brazil.
The goniadids are characterized by possession of an
annulated, conical prostomium and uniramous and sub-
biramous parapodia. The single exception is Progoniada,in
which all parapodia are uniramous. The end of the
proboscis includes paragnaths arranged in a dorsal and a
ventral arc; two macrognaths with several cusps in a
latero-ventral position and several micrognaths com-
posed of two basally jointed, Hþw-shaped pieces, and of
X- or Y-shaped single pieces, in the dorsal and/or
ventral positions. The proboscis is densely covered by
proboscideal papillae, which can be very diversi¢ed. The
genera Progoniada,Goniadella and Goniada have a paired
row of V-shaped structures, termed chevrons (Hartman,
1950).
Important taxononomic characters distinguishing
genera and species of Goniadidae include the number of
annulations of the prostomium, the shape and arrange-
ment of the proboscideal papillae, the presence or absence
of chevrons, the number and shape of the micrognaths in
the dorsal and ventral arc, the number of uniramous
chaetigers, the number and shape of the parapodial
lobes and the type of noto- and neuro-chaetae (Wilson,
2000).
Herein, we complement the description of Progoniada
regularis and describe a new species of Gonadiella. Both
species were collected in waters o¡ south-eastern Brazil
during the REVIZEE Programme ‘Avaliac a
‹o do Potencial
Sustenta
¤vel dos Recursos Vivos na Zona Econo
“mica
Exclusiva’.
MATERIALS AND METHODS
Specimens of Progoniada regularis and Goniadella revizee sp.
nov. were collected during the REVIZEE Programme/
South Score/Benthos sampling cruises of the RV ‘W.
Besnard’ in December 1997 to March 1999, on the outer
continental shelf and slope o¡ south-eastern Brazil. This
area extends from Ilha Grande Bay in southern Rio de
Janeiro State (24820.5270S43846.7590W) south to
Tr a m a n d a |¤in Rio Grande do Sul State (29828.750S
48809.350W). Water depths ranged from 60 to 808 m. The
samples were collected with vanVeen and box corer grabs
and with a dredge. The polychaetes were sorted from the
sediment, washed, ¢xed with 4% formalin, stored in 70%
alcohol, and identi¢ed.
For goniadids, the proboscis was dissected with
ophthalmic scissors, to examine the papillae, macro- and
micrognaths and chevrons. Measurements and line
drawings were made using ZEISS optical microscopy and
stereomicroscopy. Observations using scanning electron
microscopy (SEM) were made at the Laborato
¤rio
de Microscopia Eletro
“nica, Instituto de Biologia,
Universidade Estadual de Campinas (UNICAMP), with
JEOLJSM-5800 LV1equipment, after critical-point drying
with Balzers CPD 301equipment (temperature 308C, pres-
sure 80 kg cm2) and coating with 25 nm gold.The material
examined was deposited in the Museu de Histo
¤ria Natural of
UNICAMP, in the Polychaeta Collection (MHN-BPO).
RESULTS AND DISCUSSION
SYSTEMATICS
Fa m i l y GONIADIDAE Kinberg, 1865
Progoniada Hartman, 1965
Type species
Progoniada regularis Hartman, 1965.
Diagnosis
Proboscideal papillae of only a few types. Chevrons
present. First segment apodous and without chaetae, with
J. Mar. Biol. Ass. U.K. (2004), 84,47^58
Printed in the United Kingdom
Journal of the Marine Biological Association of the United Kingdom (2004)

only lateral cirri. All parapodia uniramous, with only
composite chaetae, falcigerous and/or spinigerous.
Progoniada regularis Hartman, 1965
(Figures 1^3; Table 1)
Progoniada regularis Hartman, 1965: 100^101, plate 16;
Hartman & Fauchald, 1971: 76.
Material examined
A total of 89 specimens: Station 6646, 25843.780S
45816.0 60W, 198 m, 14 December 1997, MHN-BPO/AR-
51, 1 specimen. Station 6660, 24817. 67 8 0S43848.1980W,
314m, 9 January 1998, MHN-BPO/AR-30, 2 specimens.
Station 6669, 24807.3470S44842.1420W, 1 01 m , 1 1 Ja n u a r y
1998, MHN-BPO/AR-36, 6 specimens. Station 6671,
24832.910S44827.460W, 260 m, 11 January 1998, MHN-
BPO/AR-31, 4 specimens. Station 6672, 26827.750S
44830.3510W, 165 m, 11 January 1998, MHN-BPO/AR-32,
1 specimen. Station 6676, 24849.6990S44844.9650W,
153 m, 12 January 1998, MHN-BPO/AR-38, 1 specimen.
Station 6677, 24840.7470S44850.8220W, 13 7 m , 1 2 J a nu a r y
1998, MHN-BPO/AR-39, 3 specimens. Station 6678,
24846.3570S45811.13 5 0W, 99 m, 12 January 1998, MHN-
BPO/AR-33, 2 specimens. Station 6681, 25811. 0 0 5 0S
44856.60W, 168 m, 12 January 1998, MHN-BPO/AR-40, 1
specimen. Station 6686, 25836.9880S45813.571 0W, 1 5 3 m ,
13 January 1998, MHN-BPO/AR-34, 3 specimens. Station
6690, 27800.880S46840.210W, 280 m, 18 January 199 8,
MHN-BPO/AR-37, 1 specimen. Station 6706, 25848.600S
45844.500W, 184 m, 21 January 1998, MHN-BPO/AR-35,
1 specimen. Station 6737, 24813.35 0S43825.900W, 4 7 6 m ,
14 February 1998, MHN-BPO/AR-317, 1 specimen.
Station 6738, 24811.6 0 0S43826.200W, 330 m, 14 February
1998, MHN-BPO/AR-321, 2 specimens. Station 6739,
24802.580S43830.800W, 147 m, 14 February 1998, MHN-
BPO/AR-311, 2 specimens. Station 6741, 23849.900S
43814.400W, 138 m, 15 February 1998, MHN-BPO/AR-
286, 5 specimens. Station 6744, 23851.5 0 0S42849.900W,
254 m,15 February 1998, MHN-BPO/AR-316,7 specimens.
Station 6749, 23844.200S42829.800W, 325 m, 16 February
1998, MHN-BPO/AR-282, 2 specimens. Station 6753,
23836.540S42809.860W, 187 m, 16 February 1998, MHN-
BPO/AR-322, 7 specimens. Station 6762, 23826.200S
41815.820W, 146 m, 28 February 1998, MHN-BPO/AR-
309, 6 specimens. Station 6769, 22802.870S40805.930W,
93 m, 13 March 199 8, MHN-BPO/AR-328, 3 specimens.
Station 6783, 27809.900S46852.830W, 350 m, 14 March
1998, MHN-BPO/AR-329, 10 specimens. Station 6784,
27809.510S47804.850W, 195 m, 14 March 1998, MHN-
BPO/AR-340, 4 specimens. Station 6791, 27848.780S
47810.630W, 358 m, 16 March 1998, MHN-BPO/AR-333,
2 specimens. Station 6793, 27846.490S47840.450W, 14 0 m ,
16 March 1998, MHN-BPO/AR-335, 2 specimens.
Station 6797, 28808.150S47822.610W, 190 m, 17 March
1998, MHN-BPO/AR-337, 1 specimen. Station 6801,
28834.160S47840.380W, 166 m, 17 March 1998, MHN-
BPO/AR-332, 2 specimens. Station 6821, 29849.490S
48812.760W, 232 m, 24 March 1999, MHN-BPO/AR-330,
7 specimens.
Description
Thirteen complete specimens, with lengths from 3.42 to
13.62 mm and widths from 0.16 to 0.32 mm, and 76 incom-
plete specimens. Number of chaetigers from 27 to 83 in
complete specimens. Whitish body; pigment, when
present, brownish, irregular, subdistally on the prechaetal
lobes (Figure 3G); punctiform intersegmental pigment
present from the middle ventral region (Figure 3D). Speci-
mens with slender cylindrical body, of uniform width.
Conical prostomium as long as the ¢rst six chaetigers,
with eight distinctly visible rings (Figures 1A & 3A).
Prostomial appendages biarticulate, with robust basal
articulation and a median constriction; distal articulation
short, about one-fourth as long as basal articulation
(Figure 1B). Eyes not visible. Nuchal organs inconspicuous,
ciliated, located in latero-dorsal position in the basal
prostomial ring. Four small ciliated cavities on each
prostomial ring, two ventral and two dorsal, except on
distal and basal ring (Figure 1A). All proboscids invagi-
nated, with macro- and micrognaths located between
chaetigers 13 and 14. Proboscis with two macrognaths in
latero-ventral position; three micrognaths in ventral and
about 10^14 in dorsal arc, or fewer in juvenile specimens.
Macrognaths with three or four cusps. Micrognaths with a
single X-shaped piece, or two basally jointed H þw-
shaped pieces (Figure 3C), usually arranged in the
following manner: (a) in ventral position: three groups of
the large Hþw-shaped pieces; (b) in dorsal position: a
pair of small X-shaped pieces, located near the macro-
gnaths, externally; two large pairs, with H þw-shaped
pieces; a small X-shaped pair, rarely accompanied by a
smaller U-shaped piece; and three groups with Hþw-
shaped median pieces. Paired rows of chevrons of the inva-
ginated proboscis located at the level of chaetiger 6 or 7,
with about 14 to 22 slender V-shaped pieces (Figure 3B);
juvenile specimens with eight pairs. Proboscis surface
covered with low papillae, sub-triangular, with one
subapical ciliated pore (Figure 2A). Proboscideal papillae
more numerous on the dorsal region, composed of about
5^6 series of cordiform papillae with distal ends pointed,
on narrow base (Figure 2A, F^G). Latero-ventral region
with three larger series of smaller papillae with distal ends
pointed, on wide base (Figure 2B); the ¢rst with cordiform
papillae and the distal end laterally pointed (Figure 2C),
the second, tricuspid with the median cusp bi¢d (Figure
2D); the third with papillae rounded at base with distal
end slightly pointed (Figure 2E). A posterior row,
ventral, with lateral papillae, slightly spread on narrow
bases and with distal ends rounded (Figure 2H^I).
Papillae near chevrons smaller than other papillae, with
distal ends rounded. First segment apodous, without
chaetae, and with lateral cirri only (Figures 1A & 3A).
Parapodia with prechaetal lobe digitiform and postchaetal
lobe rounded; postchaetal lobe about one-half to one-third
length of prechaetal lobe (Figures 1C & 3E,G). Dorsal and
ventral cirri digitiform to subtriangular, about as long as
postchaetal lobe, or slightly shorter. Parapodia progres-
sively slightly longer and more slender from anterior to
posterior region. Chaetigers usually with six chaetae: two
falcigers with short distal appendages, located in the outer
region of the bundle (Figures 1D & 3H), two falcigers with
distal appendages long, in the intermediate region
(Figure 3I), and two long spinigers, in the inner part of
the bundle (Figures 1E & 3J). Last chaetigers rudimen-
tary, with fewer chaetae. Pygidial cirri not visible, probably
lost (Figure 3F).
48 A.E. Rizzo and A.C.Z. Amaral Progoniada and Goniadella (Goniadidae) from Brazil
Journal of the Marine Biological Association of the United Kingdom (2004)

Progoniada and Goniadella (Goniadidae) from Brazil A.E. Rizzo and A.C.Z. Amaral 49
Journal of the Marine Biological Association of the United Kingdom (2004)
Figure 1. Progoniada regularis (SEM micrographs). (A) Anterior region, left latero-frontal view of the prostomium, showing the
ciliated cavity (cc); (B) prostomial appendages, frontal view; (C) parapodia of the median region, lateral view; (D) falcigerous
chaetae; (E) spinigerous and falcigerous chaetae. (D&E, chaetae from middle region.)

50 A.E. Rizzo and A.C.Z. Amaral Progoniada and Goniadella (Goniadidae) from Brazil
Journal of the Marine Biological Association of the United Kingdom (2004)
Figure 2. Progoniada regularis (SEM micrographs). (A) Proboscideal papillae; (B) latero-ventral papillae, arranged in enlarged
series of shorter papillae with distal ends pointed, on a wide base; (C) papilla with distal ends laterally pointed; (D) tricuspid
papilla with bi¢d median cusp; (E) papilla rounded at base and with distal end slightly pointed; (F) cordiform papillae; (G) detail
of the cordiform papilla; (H) £attened papillae with distal ends rounded, on a narrow base; (I) detail of the £attened papilla.

Progoniada and Goniadella (Goniadidae) from Brazil A.E. Rizzo and A.C.Z. Amaral 51
Journal of the Marine Biological Association of the United Kingdom (2004)
Figure 3. Progoniada regularis. (A) Anterior region, dorsal view; proboscis invaginated, chevrons (ch) seen by transparency; (B)
paired rows of chevrons with 17 pieces each; (C) pieces of micrognaths, Hþw-shaped and basally jointed (left) and X-shaped
pieces accompanied by U-shaped piece (right); (D) median region, ventral view; (E) chaetiger 35, posterior view (dc, dorsal cirrus;
post, postchaetal lobe; pre, prechaetal lobe; vc, ventral cirrus); (F) last chaetigers; (G) chaetiger 50, posterior view; (H) falcigerous
chaetae with short distal appendages; (I) falcigerous chaeta with long distal appendage; (J) spinigerous chaetae.

Remarks
Gambi & Giangrande (1988) identi¢ed as Progoniada sp.
a single specimen collected from shallow depths in the
Adriatic Sea, which was found together with Goniada
maculata rsted, 1843 and Glycinde nordmanni (Malmgren,
1865). This specimen had ¢ve pairs of chevrons, and most
of the composite chaetae without the distal appendages;
some complete spinigers were present. We believe that the
specimen examined by Gambi & Giangrande (1988) is a
juvenile of Goniada sp., because although it had only unira-
mous chaetigers, the sub-biramous chaetigers might not
yet have developed. Species of the genus Progoniada have
only lateral cirri in the ¢rst chaetiger, but the specimen
from Mediterranean Sea had the ¢rst chaetiger complete,
without chaetae. In the area where P. regularis was collected
together with specimens of Goniada brunnea Treadwell, 1906
and G. maculata, we found two small specimens, one
complete and the other incomplete with nine prostomial
rings, one pair of eyes in the basal ring, most of the
chaetae spinigerous and some falcigerous, and 5^6 pairs of
chevrons. We identi¢ed them as juveniles of G. brunnea,
because all the chaetigers were uniramous, and the total
number of chaetigers in the complete specimen was less
than 40 uniramous chaetigers, as found in this species.
According to Gilbert (1984), species belonging to Goniada
have neuropodia with spinigerous chaetae, but inferior
falcigers may be present on some anterior chaetigers.
Besides the type species, the genus Progoniada includes
only P. simplex Hartman, 1971, described from the
Mozambique Basin. Both species were found in deep
waters, between 770 and 5001m. At Bermuda, Hartman
(1965) found some individuals with a short squat body
and long natatory chaetae, together with specimens of
P. regularis. She remarked that the former might be a
distinct species, rather than an epitoke.
The specimens examined for the present report had one
pair of chevron rows, each with about 14 to 22 pieces
(mean 18), except for one juvenile with only eight. The
number of chevrons on each side of the proboscis did not
vary. Juveniles also had fewer micrognaths, some still in
the process of development. Like the chevron number, the
ventral macro- and micrognaths showed little variation.
One specimen had one w-shaped piece with four instead
of three cusps. Micrognaths in the dorsal position
numbered from ten to 14, with the exception of one juve-
nile that had only six plus some pieces in the process of
development.
In the original description, Hartman (1965) mentioned
a ¢xed number (20) of pairs of chevrons and only ¢ve
H-shaped micrognaths in the dorsal arc, besides the three
usually found in the ventral arc.
Progoniada regularis di¡ers from P. simplex in that the
latter has only spinigerous chaetae (Table 1). Examination
of the holotype of P. simplex will be necessary to determine
whether this species distinction is valid.
Bathymetric distribution
Progoniada regularis has been reported from depths of 196
to 5023 m. In this study, the species was collected from 93
to 476 m.
Geographic distribution
Western Atlantic Ocean: o¡ Massachusetts, USA;
Bermuda; o¡ Surinam; north of the mouth of the
Amazon River, o¡ Pernambuco, and from Rio de Janeiro
to Rio Grande do Sul, Brazil.
Goniadella Hartman, 1950
Type species
Eone gracilis Verril, 1873.
Diagnosis
Proboscideal papillae of one or several types. Chevrons
present. First segment apodous, without chaetae and with
one pair of lateral cirri. Anterior parapodia uniramous,
posterior sub-biramous. Prechaetal neuropodial lobe
entire, not bifurcated. Notopodial lobe absent or reduced.
Acicular notochaeta projecting directly from the epidermal
tissue, above or below the dorsal cirrus. Falcigerous and
spinigerous neurochaetae.
Goniadella revizee new species
(Figures 4^6, Table 2)
Type material
Twenty four specimens. Holotype: Station 6741,
23849.900S43814.4 00W, 138 m, 15 February 1998, MHN-
BPO/85-0; paratype series: Station 6741, 23849.900S
43814.400W, 138 m, 15 February 1998 MHN-BPO/85-1 to
MHN-BPO/85-23, 23 specimens.
Other material examined
A total of 95 specimens: Station 6666, 24817.1 2 9 0S
44812.179 0W, 163 m, 10 January 1998, MHN-BPO/AR-42,
52 A.E. Rizzo and A.C.Z. Amaral Progoniada and Goniadella (Goniadidae) from Brazil
Journal of the Marine Biological Association of the United Kingdom (2004)
Table 1. Morphological characteristics of the species of Progoniada recorded in the literature.
Species Depth (m)
Lengthwidth
(mm)
Number of
chaetigers
Numbers of
chevrons
Proboscis:
micrognaths
(ventral; dorsal arc) Chaetae
P. regularis Hartman, 1965 770^5001 14^150.25 475 i 20^20 3; 5 Falcigers and
spinigers
P. simplex Hartman, 1971* 4886^5069 22.51.3 98 i 15^15 3; 10* Spinigers
Material examined 99^476 3.42^13.62
0.16^0.32
27^83 c 14^22 3; 10^13 Falcigers and
spinigers
*, Hartman (1971) may have erred in describing ten ventral micrognaths and three in the dorsal arc, because goniadids have 0, 3 or
more micrognaths in the ventral arc and a variable number in the dorsal arc.We assume that P. simplex has three ventral and ten dorsal
micrognaths in all its populations. c, complete; i, incomplete.

Progoniada and Goniadella (Goniadidae) from Brazil A.E. Rizzo and A.C.Z. Amaral 53
Journal of the Marine Biological Association of the United Kingdom (2004)
Figure 4. Goniadella revizee sp. nov. (SEM micrographs). (A) Anterior region, dorsal view; (B) basal prostomial ring showing
nuchal organ (no) and ciliated cavity (cc), left lateral view; (C) rows of chevrons with 12 pieces (other four pieces were lost during
SEM processing); (D) acicular notochaetae; (E) transition between uniramous and sub-biramous parapodia (par); beginning of
acicular chaetae (ac) in the notopodium, dorsal view; (F) falcigerous chaetae, with long distal appendage (left) and with short
distal appendage (right).

54 A.E. Rizzo and A.C.Z. Amaral Progoniada and Goniadella (Goniadidae) from Brazil
Journal of the Marine Biological Association of the United Kingdom (2004)
Figure 5. Goniadella revizee sp. nov. (SEM micrographs). (A) Proboscideal papillae of two types with a ciliated subapical pore;
(B) papilla with a single cusp; (C) papilla with bicuspid distal end; (D) micrognaths of the ventral arc, view of the outer part;
(E) papillae located near the base of the proboscis; (F) arc of macro- (ma) and micrognaths (mi) in dorsal (d) and ventral (v)
position; (G) micrognath; (H) macrognath; (I) paired rows of chevrons with 19^20 pieces.

Progoniada and Goniadella (Goniadidae) from Brazil A.E. Rizzo and A.C.Z. Amaral 55
Journal of the Marine Biological Association of the United Kingdom (2004)
Figure 6. Goniadella revizee sp. nov. (A) Prostomial appendages; (B) anterior region, dorsal view; (C) posterior region with inter-
segmental punctiform pigment seen by transparency in dorsal view; (D) chaetiger 6 (dc, dorsal cirrus; post, postchaetal lobe; pre,
prechaetal lobe; vc, ventral cirrus); (E) chaetiger 20; (F) chaetiger 31 (ac: acicular chaetae); (G) chaetiger 40 (D^G, all chaetigers
in posterior view); (H) spinigerous chaeta with long distal appendage (left) and falcigerous chaeta (right).

56 A.E. Rizzo and A.C.Z. Amaral Progoniada and Goniadella (Goniadidae) from Brazil
Journal of the Marine Biological Association of the United Kingdom (2004)
Table 2. Morphological characteristics of species of Goniadella.
Species LengthNumber of
Number of
uniramous Number
Proboscis: macro-
gnaths (number of
cuspids)þmicrognaths Chaetae
References Geographic distribution Depth (m) width (mm) chaetigers Eyes parapodia of chevrons (ventral; dorsal) A F S
Goniadella gracilis (Verrill, 1873) sensu Hartman, 1950
Hartman, 1950 Massachusetts, o¡ Gay Head
(type locality)
Intertidal to 6 20 NM 1 pair basal þ
1 pair distal
30^31 26 4þ3; 10
(þ2^3 small)
3^43
^4NM
Pettibone, 1963 Massachusetts; Rhode Island 525 5501.0 4100 1 pair basal þ
1 pair subdistal
27^32 26 3^4þ3; 10 2^4NMNM
Walker, 1972 Liverpool Bay, England 15^37 12^23 NM NM 28^29 20^34 NM 1^22
^42
^3
Goniadella galaica (Rioja, 1923)
Rioja, 1923* Tambo Island, Mar|
¤n, Spain
(type locality)
NM 15^251NM 1pair 22
^23 17^24 3 þ3; 9^10 3 NM NM
Goniadella bobrezkii (Annenkova, 1929)
Annenkova, 1929 Black Sea (type locality) 12.5^17 13^140.5 90 1 pair basal þ
1 pair subdistal
24 17^18 5 þ15 2^3NMNM
Walker, 1972 North Sea and Black Sea NM NM NM 1 pair basalþ
1 pair distal
22^24 17^24 NM 2 2^31
^2
Wol¡ & Stegenga,
1975**
Scheldt River,
The Netherlands
84.00.2
(juvenile)
34 1 pair basalþ
1 pair subdistal
22 4^4* NM þ425
Goniadella falklandica Hartmann-Schro«der, 1986
Hartmann-Schro
«der,
1986*
Falkland Islands (type locality) 535 37 95 Not observed 45^47 15^16 NM þ4; 16* 3 4^64
^7
Goniadella unicirra Campoy & Aguirrezabalaga, 1984
Aguirrezabalaga,
1984
Guipu¤ zcoa Coast, Iberian
Peninsula (type locality)
30^70 NM 44 1 pair basal þ
1 pair subdistal
29 8^93
^4þ4;3 233
Goniadella sp. A Gilbert, 1984
Gilbert, 1984 Gulf of Mexico 19^56 24.50.8 5113 1 pair basal
(In Figure 33^
2f)
14^15 17^23 2 þ3^4; 11 NM NM NM
Goniadella revizee sp. nov.
Material examined Brazil (Rio de Janeiro to
Rio Grande do Sul)
99^808 8.08^20.3
0.14^0.3
54^129 Not observed 20^21 16^22 3 þ3; 8^10 1^32 3
*, Rioja (1923) described three micrognaths in dorsal and 9^10 in ventral position, and Hartmann-Schro
«der (198 6) described four micrognaths in dorsal and 16 in ventral position. They may have
erred, because goniadids have 0, 3 or more micrognaths in the ventral arc and avariable number (usually greater) in the dorsal arc. **, Wol¡ & Stegenga (1975) described eightV-shaped chevrons;
they illustrated the anterior region with two rows of chevrons, with four pieces on each. NM, not mentioned; A, acicular; F, falciger; S, spiniger.

2 specimens. Station 6671, 24832.910S44827.460W, 260 m,
11 January 1998, MHN-BPO/AR-50, 1 specimen. Station
6673, 24817.9390S44835.9830W, 133 m, 10 January 1998,
MHN-BPO/AR-43, 3 specimens. Station 6674, 24831.0 80S
44854.000W, 122 m, 10 January 1998, MHN-BPO/AR-4 4,
3 specimens. Station 6676, 24849.6990S44844.9650W,
153 m, 12 January 1998, MHN-BPO-AR45, 1 specimen.
Station 6678, 24846.3570S45811.1 35 0W, 9 9 m, 12 January
1998, MHN-BPO/AR-46, 3 specimens. Station 6679,
25818.874 0S44852.5160W, 808 m, 12 January 1998, MHN-
BPO/AR-47, 2 specimens. Station 6686, 25836.9880S
45813.571 0W, 153 m, 13 January 1998, MHN-BPO/AR-49,
1 specimen. Station 6739, 24802.580S43830.800W, 14 7 m ,
14 February 1998, MHN-BPO/AR-310, 6 specimens.
Station 6744, 23851.50 0S42849.900W, 254 m, 15 February
1998, MHN-BPO/AR-315, 1 specimen. Station 6750,
23840.080S42831.800W, 162 m, 16 February 199 8, MHN-
BPO/AR-312, 1 specimen. Station 6753, 23836.540S
42809.860W, 187 m, 16 February 1998, MHN-BPO/AR-
323, 4 specimens. Station 6754, 23826.700S42814.0’W,
131m, 17 February 1998, MHN-BPO/AR-320, 2 speci-
mens. Station 6759, 23820.00S41822.00W, 11 0 m , 2 8
February 1998, MHN-BPO/AR-319, 32 specimens.
Station 6762, 23826.200S41815.820W, 146 m, 28 February
1998, MHN-BPO/AR-308, 30 specimens. Station 6784,
27809.510S47804.850W, 195 m, 14 March 1998, MHN-
BPO/AR-341, 1 specimen. Station 6797, 28808.150S
47822.610W, 190 m, 17 March 1998, MHN-BPO/AR-336, 1
specimen. Station 6817, 29828.750S48809.350W, 21 0 m , 2 3
March 1999, MHN-BPO/AR-342, 1 specimen.
Description
Holotype complete, with 75 chaetigers, length 10.6 mm,
width 0.28 mm. Paratype series with two complete, 21
incomplete specimens; complete specimens with 66^67
chaetigers, length 8.08^8.83 mm and width 0.24^
0.26 mm. Other material includes 13 complete specimens
with 54 to 129 chaetigers, length 8.25^20.3 mm and
width 0.14^0.3 mm, and 79 incomplete. Pigment, when
present, irregular, brownish, subdistally on the pre-
chaetal lobes and cirri; punctiform intersegmental
pigment present from biramous ventral region. Usually
with brown colouring on entire body, on dorsal and
ventral side.
Body long, anteriorly cylindrical, posteriorly slender.
Conical prostomium with eight rings (Figures 4A & 6B).
Prostomial appendages triarticulate, reaching the second
distal prostomial ring; basal article one-third size of
distal article (Figure 6A). Nuchal organs ciliated, drop-
shaped, located latero-dorsally on basal prostomial ring
four small ciliated cavities on each prostomial ring, two
ventral and two dorsal, except on distal and basal ring
(Figure 4B). Eyes not visible. All proboscids invaginated;
proboscis with two ventral macrognaths, three ventral
micrognaths, and 8^13 dorsal micrognaths (Figure 5F).
Macrognaths with three cusps (Figure 5H). Micrognaths
composed of two basally jointed pieces, one H-shaped and
the other smaller and w-shaped (Figure 5D, G); micro-
gnaths of ventral arc up to twice as large as micrognaths
of dorsal arc. Each row of chevrons with 16 to 22 in
V-shaped pieces (Figures 4C & 5I). Proboscideal papillae
mainly of two types, subtriangular, most with a single cusp
and some bicuspid; both types basally cleft, with a ciliated
subapical pore (Figure 5A^C). Papillae located near the
chevrons, with distal end rounded, and more distal
papillae with pointed ends; all papillae also basally cleft
with a ciliated subapical pore. Basal papillae smaller and
more rounded than others (Figure 5E). Parapodia of
posterior region longer and more slender than those of
anterior region. First segment apodous and achaetigerous,
with one pair of small digitiform lateral cirri (Figures 4B
& 6B). Parapodia composed of one digitiform prechaetal
lobe and one shorter, rounded postchaetal lobe, up to half
the length of the prechaetal lobe; and also of one dorsal
and one ventral cirrus, both digitiform to subtriangular
(Figure 6D,F). Biramous parapodia beginning on chae-
tiger 20^21 (20 in holotype), with the appearance of the
notochaetae (Figure 4E). Notopodium rudimentary, not
prominent, with 1^3 (usually two) simple acicular
chaetae, arising above dorsal cirrus (Figure 6E,G); one of
the chaetae with its tip slightly curved (Figure 4D).
Posterior neuropodia similar to anterior. Neuropodial
bundle usually composed of one pair of outer falcigerous
chaetae, one pair of spinigers with short distal appen-
dages in an intermediate position, and one spiniger
with one long distal appendage in the inner position of
the bundle (Figures 4F & 6H). Noto- and neuro-
podium each supported by one acicula. Anal opening
rounded, distal. Pygidium with one pair of slender
conical anal cirri, reaching in length the last three chae-
tigers (easily lost in ¢xed material).
Etymology
This species is dedicated to the REVIZEE Programme,
which made possible the collection of the material.
Remarks
Goniadella revizee sp. nov., Goniadella sp. A Gilbert, 1984
and Goniadella bobrezkii (Annenkova, 1929) di¡er from
other species of the genus mainly in possessing acicular
notochaetae above the dorsal cirrus on sub-biramous
parapodia, and in regard to the number of uniramous
chaetigers and chevrons. One of the acicular notochaetae
of G. re v i zee sp. nov. has its end slightly curved, similar to
those described for Goniadella unicirra Campoy &
Aguirrezabalaga, 1984 (in Aguirrezabalaga, 1984) and
G. bobrezkii.Goniadella bobrezkii is known from European
waters (Annenkova, 1929; Walker, 1972; Wol¡ & Stegenga,
1975). Goniadella bobrezkii di¡ers from Goniadella revizee sp.
nov. mainly in the number of uniramous chaetigers (24)
and the number of chevrons (17^18), but also in possessing
two pairs of eyes, and macrognaths with ¢ve cusps, besides
15 micrognaths. In the original description of this species,
the presence of ventral micrognaths was not mentioned.
Table 2 lists some morphological characteristics of the
¢ve described species of the genus Goniadella, taken from
the main references, and of Goniadella revizee sp. nov.
Goniadella gracilis (Verrill, 1873) and Goniadella falklandica
Hartmann-Schro
«der, 1986, described from the western
Atlantic, and Goniadella galaica (Rioja, 1923) from o¡
Galicia, Spain, di¡er from Goniadella revizee sp.nov.in
having acicular notochaetae immediately below or on the
dorsal cirrus (Hartman, 1950: plate 5, ¢gure 7). In
Goniadella revizee sp. nov. the dorsal cirrus is next to the
neuropodial lobe in both the uniramous and the biramous
Progoniada and Goniadella (Goniadidae) from Brazil A.E. Rizzo and A.C.Z. Amaral 57
Journal of the Marine Biological Association of the United Kingdom (2004)

chaetigers. Moreover, these species have more uniramous
chaetigers than does Goniadella revizee sp.nov.(Table2).
Specimens from the Gulf of Mexico were ¢rst identi¢ed
as Goniadella sp. and G. gracilis. These specimens were later
transferred to Goniadella sp. A by Gilbert (1984), because
they did not share certain taxonomic characters with
G. gracilis, a species known from northern temperate
waters. Goniadella sp. A di¡ers from Goniadella revizee sp.
nov. in having: (a) 14^15 uniramous chaetigers; (b)
macrognaths with two cuspids; (c) straight acicular noto-
chaeta; and (d) a pair of eyes on the basal prostomial ring
(from ¢gure 33^2f, Gilbert, 1984).
Morphological di¡erences were found between the
specimens from the Brazilian south-east coast and those
from the Caribbean region. The specimens from o¡
Brazil have: (a) 20^21 uniramous chaetigers; (b) macro-
gnaths with three cusps; (c) acicular notochaetae with the
ends slightly curved; and (d) eyes not evident. Gilbert
(1984) described the proboscideal papillae of Goniadella
sp. A as being of a single type, sub-triangular with only a
cusp. The SEM observations for the Brazilian population
revealed the existence of two types, with one or two cusps.
Although the specimens from both localities share certain
taxonomic characters, such as the number and position of
the acicular notochaetae and the number of pairs of chev-
rons, we decided to consider them two distinct species.
Our reasoning was based mainly on the di¡erent number
of uniramous chaetigers, which is usually the ¢rst charac-
teristic used to separate species in other genera of gonia-
dids. Additional di¡erences between these two species are
the size (lengthwidth), the number of cusps in the
macrognaths, the presence or absence of eyes, and the
range of depths where both species were found. In our
opinion, the di¡erences between Goniadella gilbertae sp.
nov. and other species are su⁄cient to justify designating
a new species.
Bathymetric distribution
Goniadella revizee sp. nov. occurred from 99 to 808 m.
Geographic distribution
Western Atlantic Ocean: o¡ Brazil, from Rio deJaneiro
to Rio Grande do Sul.
Our thanks to the CNPq (Conselho Nacional de
Desenvolvimento Cient|
¤¢co e Tecnolo
¤gico, Process no. 141504/
98-6), and to the MMA (Ministe
¤rio do Meio Ambiente, dos
Recursos H|¤dricos e da Amazo
“nia Legal) for ¢nancial support.
We appreciate the facilities and assistance of the Departamento
de Zoologia and the sta¡ of the Laborato
¤rio de Microscopia
Eletro
“nica, Instituto de Biologia, UNICAMP. We thank the
anonymous referees whose comments greatly improved the
manuscript. Our thanks also to Maren Boveri, for translating
the work of Annenkova (1929) from German to Portuguese.
Janet W. Reid revised the English text.
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Submitted 12 December 2002. Accepted 10 October 2 003.
58 A.E. Rizzo and A.C.Z. Amaral Progoniada and Goniadella (Goniadidae) from Brazil
Journal of the Marine Biological Association of the United Kingdom (2004)