ISSN 0365-4508
Nunquan aliud natura sapienta dicit
J. 14,321
In silvis academi qnoerere rerum,
Quanquam Socraticis madet sermonibus
Ladisl. Netto, ex Hor
VOL. LX
N.l
RIO DE JANEIRO
Janeiro/Março
2002
ARQUIVOS
DO
MUSEU NACIONAL
VOLUME 60
NÚMERO 1
JANEIRO/MARÇO
2002
RIO DE JANEIRO
Arq. Mus. Nac.
Rio de Janeiro
v.6o
n.i
p.1-50
jan./mar.2002
ISSN 0365-4508
COMISSÃO DE PUBLICAÇÕES
MUSEU NACIONAL/UFRJ
Editor
C.N.Ricci
Conselho Editorial - Museu Nacional
V.F.Barbosa, U.Caramaschi, V.G.L.Esteves, V.M.M.Fonseca, A.W.A.Kellner, V.C.Klein, M.A.R.Maia, G.L.F.Mejdalani, M.A.Monné,
M.D.B.G.Oliveira, C.N.Ricci, A.M.N. Vilaça, P.S.Young
Conselho Científico
M.B.M.Abaurre (UNICAMP), C.Ades (USP), M.G.M.Ávila (UFPE), M.T.P.Azevedo (SMA-SP), M.E.C.B.O.Babinski (USP), G.Bacoccoli
(PETROBRÁS), S.G.Baines (UnB), D.C.A.Barberena (UFRGS), M.A.V.Barros (I.Bot.SP), D.C.Bicudo (I.Bot.SP), L.Bisol (PUCRS),
C.R.Brandão (UNICAMP), I.M.Brito (UFRJ), K.S.Brown Jr. (UNICAMP), M.C.O.Bruno (USP), H.F.A.Camargo (USP), M.S.F.S.Capelato
(I.Bot.SP), C.J.B.Carvalho (UFPR), L.d’A.F.Carvalho (JBRJ), C.M.G.Corrêa (MPEG), C.G.Costa (JBRJ), N.M.C.Cruz (CPRM), H.Dayan
(UFRJ), V.R.D.Eickstedt (I.But.SP), C.Fonseca (UFRGS), L.Forneris (USP), E.F.Guimarães (JBRJ), S.M.P.B.Guimarães (I.Bot.SP), J.Jurberg
(FIOCRUZ), M.A.Kato (UNICAMP), J.E.Kraus (USP), A.Langguth (UFPB), M.Lemle (UFRJ), A.A.Lise (PUCRS), M.C.Loureiro (UFV),
J.Loyola e Silva (UFPR), S.MUucas (I.But.SP), L.M.C.Machado (IAB), G.M.A.S.Maior (UFPE), M.C.D.Mansur (FZB), L.A.Marcuschi
(UFPE), M.D.Marques (USP), G.Martinelli (JBRJ), H.RMatthews (UFCE), R.H.Maués (UFPA), J.C.Melatti (UnB), T.S.Melhem (I.Bot.SP),
R. P.Mello (FIOCRUZ), J.O.Menegheti (UFRGS), U.T.B.Meneses (USP), N.A.Menezes (USP), N.L.Menezes (USP), O.H.H.Mielke (UFPR),
A.E.Migotto (USP), J.L.Morais (USP), E.G.Neves (USP), F.C.Novaes (MPEG), P.E.A.M.Ohveira (UFU), M.G.S.Peirano (UnB), J.F.Pereira
(HB), L.M.Pessôa (UFRJ), M.Pinna (USP), S.R.R.Queiroz (UNICAMP), R.E.Reis (PUCRS), C.F.D.Rocha (UERJ), M.A.C.Rodrigues (UERJ),
S. A.Rodrigues (USP), M.C.A.P.Rosa (UFRJ), A.B.Rylands (UFMG), F.M.Salzano (UFRGS), J.F.P.Sanchis (UFMG), C.L.SanfAna (I.Bot.SP),
H.Sarian (USP), E.Schlenz (USP), P.I.Schmitz (IAP-RS), P.A.C.Senna (U.F.S.Car.) A.LSilva (USP), F.L.Silveira (USP), U.R.M.Souza (USP),
J.W.Thomé (PUCRS), D.P.Uchôa (USP), S.A.Vanin (USP), L.Vidal (USP), H.M.Watanabe (I.Bot.SP), OYano (I.Bot.SP).
Diagramação, composição e arte-final
C.N.Ricci, L.M.S.Ribeiro
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Os Editores
THE VERRUCIDAE (CRUSTACEA, CIRRIPEDIA) FROM THE WESTERN COAST OF
NORTH AMERICA, WITH A REVISION ON THE GENUS ALTIVERRUCA «
(With 30 figures)
PAULO S. YOUNG 1 (2)
Museu Nacional
Universidade Federal do Rio de Janeiro
The barnacle fauna from the Northeastern Pacific
has been summarized by PILSBRY (1916),
CORNWALL (1951; 1955), NEWMAN (1982), ROSS
(1962), NEWMAN & ABBOTT (1980) and YOUNG &
ROSS (2000). However none of these authors
recorded any verrucids, thus suggesting they are
unknown in the Northeastern Pacific, although well
known in other parts of the world. ZEVINA &
GALKIN (1993) described the unique species
Altiverruca beringiana occurring north of 30°N in
the North Pacific. ZEVINA (1987) described A.
galapagosa from the Galapagos Is. Not one species
of Verrucidae has been reported from the eastern
Pacific coast of México, Califórnia and Oregon.
The verrucids from the Northeastern Pacific contained
in the collections of the Scripps Institution of
Oceanography and Califórnia Academy of Sciences
are the focus of this study. Eight species of verrucids
are recognized, of which six are new species. Therefore,
the apparent lack of verrucids in this area appears to
be related to the lack of studies of this taxon.
One of the most common verrucids found in the
deep-sea is Altiverruca with more than 40 species
described. Herein this genus is reevaluated and
divided into three new taxa.
Suborder Verrucomorpha Pilsbry, 1916
Family Verrucidae Darwin, 1854
Genus Altiverruca Pilsbry, 1916 s.l.
Remarks - YOUNG (1998a) revised the Verrucidae,
recognizing eight genera, viz: Altiverruca Pilsbry,
1916, Cameravemica Pilsbry, 1916, Newmaniverruca
Young, 1998, Costatoverruca Young, 1998,
Rostratoverruca Broch, 1922, Brochiverruca Zevina,
1993, and Metaverruca Pilsbiy, 1916. In his revision
the erect shells ( Altiverruca ) were separated from
the box-like and flat shells.
YOUNG (1998a) listed 43 described species and
subspecies in Altiverruca s.l., 10 of which are
considered synonyms. At the same moment,
BUCKERIDGE (1997) described one more species of
Altiverruca, enlarging the list to 44 species. In this
study I revalidate six species previously considered
to be synonyms of Altiverruca gibbosa, and describe
five new species of this group bringing the total to
49 taxa, of which 45 are considered valid species.
The box-like shells of most verrucid genera appear
to have evolved from erect shells such as Altiverruca
s.l. (YOUNG, 1998a; NEWMAN, 2000). The
generalized species of Altiverruca exhibit some
plesiomorphic characters: (1) erect shell, (2) opercular
plates with poorly developed ridges, (3) shell plates
with simple sutures, (4) absence of radii-like
structures, (5) absence of pits or ridges for insertion
of the adductor muscle, and (6) having thin basal
margins that are not inflected.
In reevaluating descriptions of the species of
Altiverruca I recognized three groups, which can be
characterized only by externai characters. Many of
these species are poorly described and figured, and
most of them lack any description of the appendages.
Certainly, with more accurate descriptions new
characters should be used in the diagnoses of these
genera or even in recognizing new groups of species.
My reassessment of Altiverruca s.l. suggests it
should be divided into Altiverruca s.s., and two new
genera described below.
Altiverruca Pilsbry, 1916 s.s.
Verruca Section D: Altiverruca Pilsbry, 1916:40 (part).
Verruca (Altiverruca) - BROCH, 1931:45 (part);
FOSTER, 1978:68 (part).
Altiverruca - ZEVINA, 1987a: 1813 (part);
BUCKERIDGE, 1994:92 (part); YOUNG, 1998a:
77 (part).
Spongoverruca Zevina, 1987a: 1813.
Type species- Verruca hoeki Pilsbiy, 1907, by original
designation (PILSBRY, 1916: 40), Recent, Anegada
Passage, Lesser Antilles, 18°30’N, 63°31W, 907m.
Diagnosis - Shell small and erect, wall plates
delicate. Rostrum nearly rectangular, shorter than
carina, usually with a flat or ribbed surface between
the uppermost articular ridge and the base of
scutum. Rostrum and carina suture straight to
undulated. Wall plates without longitudinal ribs,
usually with outward projecting growth lines.
Opercular plates erect, perpendicular to basis and
almost parallel to fixed-scutum and tergum. Scutum
and tergum with weak ridges, usually with one or
two, and not more than three on each plate. Fixed
1 Received on March 14, 2000. Accepted on September 22, 2001.
2 Museu Nacional/UFRJ, Departamento de Invertebrados, Quinta da Boa Vista, São Cristóvão, 20940-040. Rio de Janeiro, RJ, Brazil. psyoung@acd.ufrj.br.
6
P.S.YOUNG
scutum without adductor ridge or myophore. Bases
of wall plates not inflected.
Species - A. incerta (Hoek, 1883), A. obliqua (Hoek,
1883), A. quadrangularis (Hoek, 1883), ? A. inermis
(Aurivillius, 1898), A. erecta (Gruvel, 1900), A.
longicarinata { Gruvel, 1900), ? A. plana (Gruvel, 1907),
A hoeki (Pilsbry, 1907), A spongicola (Gruvel, 1911),
A. joubini (Gruvel, 1912), A. casula (Hoek, 1913), A.
omata (Nilsson-Cantell, 1929), ? A. aves (Zevina,
1975), A. gira (Zevina, 1987), A. galapagosa Zevina,
1987, A. angustiterga Zevina, 1987, A. sculpturata
Zevina, 1987, A. sublima Zevina, 1987, A. longa
Zevina, 1988, A. tchesunovi Zevina, 1988, A. vitrea
Zevina, 1988, A. galkini Zevina, 1990, ?A. beringiana
Zevina & Galkin, 1992, A. laeviscuta Buckeridge,
1994, A. vertica Foster & Buckeridge, 1995 (=A.
obliqua) , and two new species described below.
Remarks - Altiverruca s.s. encompasses those
delicate species with the erect shell, having the
opercular plates perpendicular to the basis, without
fully developed articular ridges and the sutures of
the rostrum and carina feebly developed. This group
of characters appears to be all simplesiomorphies
from which evolved the more flattened shell and
more elaborate shell and opercular plates of the
other species of Verrucidae.
This genus includes many species with unfortunately
brief descriptions and in many cases they are badly
or not figured at all. Therefore, after examination of
the type specimens some rearrangements may be
necessary. Based on the original descriptions and
figures of all species, I note that some of the species
included in this genus do not conform with all of the
diagnostics characters: A. plana is indifferently
figured, not showing any ridge on the opercular plates
and the suture between the rostrum and carina; the
description of this species is possibly based on
juveniles. A. inermis has the tergum figured with four
conspicuous articular ridges. A. aves has the ridges
on the opercular plates well developed. A. beringiana
has the rostro-carinal suture undulated similar to
those of Crístallinaverruca. For the present, these
species are tentatively included in Altiverruca s.s.
Altiverruca s.s. has a worldwide distribution with
individuais occurring between 900 and 4950
meters. Figure 1 lists the type localities of all
Altiverruca s.s. excluding the present synonyms.
40° -
20 °
1 A. incerta
11 A. casula
21/4. vitrea
2 A. obliqua
12 A. ornata
22 A. galkini
3 A. quadrangularis
13 A. aves
23 A. beringiana
4 A. inermis
14 A galapagosa
24 A. laeviscuta
5 A. erecta
15 A. angustiterga
25 A. vertica
6 A. longicarinata
16 A. sculpturata
26 A. vansyoci sp. nov.
7 A. plana
17 A. sublima
27 A. sala sp. nov.
8 A. hoeki
18 A. gira
9 A. spongicola
19 4. longa
10 A. joubini
20 A. tchesunovi
100 °
20° 0 o 20° 60° 100° 140° 180° 140°
Fig. 1- Type localities of all species included in Altiverruca s.s.
100 °
Arq. Mus. Nac., Rio de Janeiro, v.60, n.l, p.5-40, jan./mar.2002
THE VERRUCIDAE FROM THE WESTERN COAST OF NORTH AMERICA...
7
Altivermca beringiana Zevina & Galkin, 1993
(Figs.2-4)
Altiverruca beringiana Zevina & Galkin, 1992;
140:fig.2; 1993:62, fig.2 (translation of 1992);
YOUNG, 1998a:77.
Material - Gulf of Alaska, Kodiak Seamount,
56°51.110’N, 149°15.830W, 2051m, R/V Atlantis,
Alvin dive 3440, P.Londsdale and L.Levin cols. 02
Aug 1999, 6 specimens, rc: 1.5-2.6mm, SIO-BIC C
10239, MNRJ 14333.
Supplementary description - The present specimens
exhibit different stages of development (Fig.2). All
of them are white and erect with the operculum
very steep, almost parallel to the fixed scutum and
tergum. The primordial valves are evident, even in
the larger specimen. The number of articular ridges
of the opercular plates and on the rostro-carinal
suture changes during growth.
In smaller individuais, the tergum is quadrangular
and has only the axial ridge, which is well marked
at the carinal side (Fig.2a). During growth this ridge
becomes to be conspicuous and well marked on both
sides (Fig.2c). In the middle of the articular margin
of the tergum another large, low ridge appears in
the older specimens. The scutum has a conspicuous
axial ridge and during growth, an articular ridge of
the same width as the axial ridge also develops
(Fig.2a-c). A scutal flat area appears above the
articular ridge in the larger specimens.
The rostro-carinal suture (Fig.2a-c) is straight in
young specimens and with the development of the
rostrum and carina the ridges on the plates become
indented.
Both rostral and carinal apexes are projected.
Zevina and Galkin (1992) observed a constancy of
a more projected carina, but the present specimens
have all the States: rostrum more projected, carina
more projected or both with the same projection.
The fixed tergum and scutum (Fig.2d) are similar
to those noted before.
Fig.2- Altiverruca beringiana Zevina & Galkin: (a-c) rostro-carinal view of three specimens in three distinct development
stages; (d) fixed tergum and fixed scutum view of c; (e) tergum and scutum, internai view; (c) carina; (fs) fixed-scutum; (ft)
fixed-tergum; (r) rostrum; (s) scutum; (t) tergum.
Arq. Mus. Nac., Rio de Janeiro, v.60, n.l, p.5-40, jan./mar.2002
P.S.YOUNG
Labrum (Fig.3a) with more than 80 sharp teeth along
margin. Palp (Fig.3a) small, elongated, with fewsimple
setae. Mandible (Fig.3b) with two large teeth and a
wide denticulated lower margin; second tooth with
denticle on upper margin. Maxilla I (Fig.3c) with
cutting border having lower third well projected; two
large spines on upper margin followed by two median-size
spines, lower margin with 4 large and median spines.
Maxilla II (Fig.3d) quadrangular, with few simple
setae. Cirri with subequal rami, covered by simple
setae (Fig.4a, b). Length of median article of cirrus VI
(Fig.4c) about 2.5 times its width, with two pairs of
setae on anterior margin, larger pair pinnate, and one
or two simple setae on posterior angle; distai apical
setae about 6 times width of article. Cirral counts in
table 1. Caudal appendage (Fig.4d) multiarticulated,
almost with same length of protopod.
Remarks - Altiverruca beríngiana was originally found
off Bering Is., 55°33,6’N, 167°19’E, 4000m. This new
record at Kodiak Seamount, 56°51.110’N,
149°15.830W, 2051m enhances its distribution.
The stability in number of articular ridges on the
opercular valves has been discussed by some
authors. FOSTER & BUCKERIDGE (1995a, b)
observed variation in the number of ridges for some
species. Others suppose this character is more
stable for each species, but if there is variability in
the number of ridges in a species it is largely because
the specimens are in different stages of development
(YOUNG, 1998a, b). The specimens of Altiverruca
beríngiana are a good example. The smaller ones
have a smaller number of articular ridges with the
number increasing during growth. But if the larger
specimens are examined one can observe by the
structure of the opercular plates that it also had an
earlier stage with fewer articular ridges - easily
observed by the absence of the median articular
ridge of tergum along the first growth line. Another
example is the change in the number of ridges on
the plates of Gibbosaverruca navicula (Hoek, 1913),
which depends upon its size. Therefore, changes
in the number of ridges may occur during ontogeny.
Fig.3- Altiverruca beríngiana Zevina & Galkin: (a) labrum and palp; (b) mandible; (c) maxilla I; (d) maxilla II.
Arq. Mus. Nac., Rio de Janeiro, v.60, n.l, p.5-40, jan./mar.2002
THE VERRUCIDAE FROM THE WESTERN COAST OF NORTH AMERICA...
Fig.4- Altivermca beringiana Zevina & Galkin: (a) cirrus I; (b) cirrus II; (c) intermediate article of cirrus VI; (d) caudal appendage
and protopodite.(CI-CVI) cirri I to VI; (a.) anterior ramus; (c.a.) caudal appendage; (p.) posterior ramus.
In addition, I suspect there is little interspecific
variation in the number of ridges, viz, specimens of
the same size do not have a distinct number of
ridges. When this is observed, there is probably
more than one species involved.
TABLE 1
Number of articles on cirri I-VI, and caudal appendage of
Altivermca berimgiana Zevina & Galkin
I
II
III
IV
V
VI CA
RC 5/5
5/5
-
11/10
14/14
13/13 5
LC 4/6
5/4
6/3
10/10
11/10
14/9
(CI-VI) cirri I to VI; (CA) caudal appendage; (RC) right cirri;
(LC) left cirri.
Altivermca galapagosa Zevina, 1987
(Figs.5-7)
Altivermca gálapagossa [sic] Zevina, 1987a: 1814, fig. 1.
Altivermca galapagosa - ZEVINA, 1988:39;
BUCKERIDGE, 1994:95, fig.3a-i; YOUNG,
1998a:77.
Material - Isla Espanola, Galapagos Islands,
1°22.20’S, 89°49.20W, 354m, J.McCosker coll., 07
Jul 1998, 8 specimens, rc: 1.0-4. lmm, on Hydrozoa,
CASIZ 119488, MNRJ 14334.
Description: Shell (Fig.5a, b) white, opercular valves
nearly perpendicular to base of wall, with growth lines
prominent on all plates; shell plates without
longitudinal ridges; basal margin not thickened. Cuticle
not persistent on wall and opercular valves. Rostrum
(Fig.5a, b) nearly rectangular, with an undulated
rostro-carinal articulation, mostly formed by a wide
ridge, a conspicuous ridge curved directed toward axial
ridge of scutum above the upper articular ridge, rostrum
and fixed scutum articulation without conspicuous
radius-like projection, apexslightlyprojected, incurved.
Carina (Fig.5a, b) triangular, with large radius-like
projection toward fixed tergum; apex projecting and
recurved. Fixed tergum (Fig.5b) slightly higher than
fixed scutum, both sides with well-developed alar-like
projections; apex curved toward fixed scutum. Fixed
scutum (Fig.5b) with wide alar-like process directed
toward rostrum and a radii-like process directed
toward fixed tergum, apex curved toward fixed tergum;
intemally, without adductor pit or ridge.
Arq. Mus. Nac., Rio de Janeiro, v.60, n.l, p.5-40, jan./mar.2002
10
P.S.YOUNG
Tergum (Fig.5a, c) quadrangular, with three articular
ridges; axial ridge prominent, intermediate ridge as
wide as axial ridge; upper ridge thin and marginal to
occludent margin, with a conspicuous depression
between upper and intermediate ridges; carinal area
smooth. Internally, smooth and flat; scutal margin
undulated with a median protuberance. Scutum
(Fig.5a, d) smaller than tergum; with two articular
ridges; axial ridge conspicuous, thin; second ridge
as wide as axial ridge, and a flat upper triangular
projection at scutal margin; rostral area smooth.
Internally, with a long concavity along articular
margin, its upper portion for insertion of adductor
muscle; tergal margin straight, except for a notch at
medio-basal portion.
Labrum (Fig.6a) with a series of about 25 simple
sharp teeth. Palp (Fig.6b) short, acuminate, with few
simple setae on inner margin and distai region.
Mandible (Fig.õc, d) with three teeth, distance
between first and second two times distance between
second and third, second and third tooth
denticulated on the upper margin; lower angle
denticulated. Maxilla I (Fig.õe) with notch and lower
part strongly projected; 2 large spines at upper
border, 3 median spines on notch and 2 large and 2
median spines on lower projected border. Maxilla II
(Fig.6f) triangular, anterior margin with conspicuous
concavity medially; covered by long simple setae,
except on the concavity.
Cirrus I (Fig.7a) with equal rami, both rami covered
with several, long simple setae. Cirrus II (Fig.7b)
with unequal rami, anterior ramus about 2/3 length
1 mm
Fig.5- Altivemica galapagosa Zevina: (a) rostro-carinal view; (b) fixed tergum and fixed scutum view; (c-d) tergum and
scutum, internai view; (e) right and left side of an early juvenile with primordial plates; (s) scutum, (t) tergum, (c) carina.
Arq. Mus. Nac., Rio de Janeiro, v.60, n.l, p.5-40, jan./mar.2002
THE VERRUCIDAE FROM THE WESTERN COAST OF NORTH AMERICA...
11
Fig.6- Altiverruca galapagosa Zevina: (a) labrum; (b) palp; (c-d) mandibles; (e) maxilla I; (f) maxilla II. a, d, from one
specimen and others from another specimen figured in figs. 7 and 9.
of posterior; both rami covered by numerous long,
simple setae, distai articles of anterior ramus also
with finely pinnate setae. Rami of cirrus III subequal
and of cirri IV-VI equal in length. Intermediate articles
of cirrus VI (Fig.7c) having their width about 1 /2 their
length; three pairs of setae on anterior margin, large
antero-distal pair finely pinnate; one or two fine
simple setae and multifid setae on posterior angle.
Caudal appendage (Fig.7d) with 7 articles, slightly
longer than protopodite; long simple setae on distai
margins of articles. Penis short, distally clothed with
thin setulae. Number of articles of cirri I-VI and
caudal appendage is presented in table 2.
TABLE 2
Number of articles on cirri I-VI, and caudal appendage of
Altiverruca galapagosa Zevina
I
n
m
IV
V
VI
CA
RC
10/8
6/10
14/12+
11+/18
13+/17+
17/20
3+
LC
9/8
6/10
14/15
16/15+
16+/7+
19/20
7
(CI-VI) cirri I to VI; (CA) caudal appendage; (RC) right cirri;
(LC) left cirri; ; (+) broken ramus.
Earliest juvenile with primordial valves conspicuous
(Fig.5e). Carina asymmetrical, slightly displaced toward
right side; slightly higher than wide, extending up short
distance between terga. Tergum large, symmetrical on
each side, scutal margin somewhat concave.
Arq. Mus. Nac., Rio de Janeiro, v.60, n.l, p.5-40, jan./mar.2002
12
P.S.YOUNG
Fig.7- Altivemica galapagosa Zevina: (a) cirrus I; (b) cirrus II; (c) intermediate article of cirrus VI; (d) caudal appendage,
protopodite and penis. (CI-CVI) cirri I to VI; (a.) anterior ramus; (c.a.) caudal appendage; (p.) posterior ramus; (pe.) penis.
Scutum larger than tergum, almost symmetrical on
each side, only distinct at basal margin,
superimposing over the terga as a large extension
of the articular margin.
Remarks - The specimens available agree with the
original description and figures of ZEVINA (1987a:
1814, fig.l). Altiverruca galapagosa was described
from 5°13.6 S, 107°31.7’W, between 3830-3850m.
The present record (1°22.20’S, 89°49.20’W), at
354m differs somewhat from its original
geographical and depth range. BUCKERIDGE
(1994: 95) also recorded this species from New
Caledónia, from 570-3680m, but further study
may indicate we are dealing with two species.
NEWMAN (2000: 80) compared the primordial plates
of the earliest juvenile stages of the Neoverrucidae,
X-juvenile, Verrucidae and others. The Verrucidae
was typified by the development of Verruca stroemia
(Müller, 1776) (RÜNNSTROM, 1927). He noted that
juveniles had the carina about as wide as high,
shorter than the terga, displaced to one side, not
extending up between the terga, and the terga and
scuta were distinctly asymmetrical on each side.
Verruca stroemia represents a highly derived species
whereas species of Altiverruca s.l. encompass the
most primitive genus of the family. In A. galapagosa
primordial plates are significantly distinct from V.
stroemia : the carina is a slightly higher than wide,
it is displaced a little to the right side and it extends
up between the terga, but more impressive is the
almost symmetrical terga and scuta. The form of
the primordial plates places this juvenile in a stage
between the Neoverrucidae and V. stroemia
juveniles, agreeing with the general view of the
evolution of the group.
Altiverruca sala sp.nov.
(Figs.8-10)
Material - Holotype: Green Seamount, AD1639,
20°48’N, 109°16W, 1850m, P. Londsdale&L. Levin
cols. Sept. 1985, 1 specimen, rc: 3.2mm, SIO-BIC
C 10309. Paratype, same locality, 1875m, 1
specimen, rc: 3.6mm, MNRJ 14340.
Diagnosis - Shell with growth lines prominent on
all plates. Tergum with only an axial ridge. Scutum
with two articular ridges, both having the same
width. Cirrus I with equal rami; cirrus II with
anterior ramus about 2/3 length of posterior.
Arq. Mus. Nac., Rio de Janeiro, v.60, n.l, p.5-40, jan./mar.2002
THE VERRUCIDAE FROM THE WESTERN COAST OF NORTH AMERICA...
13
Intermediate articles of cirrus VI with two pairs of
setae on anterior margin. Caudal appendage 3 times
length of protopodite.
Description - Shell (Fig.8a, b) white, opercular valves
obliqúe to base of wall, with growth lines prominent
on all plates; wall plates without longitudinal ridges;
basal margin not thickened. Cuticle persistent on
shell and opercular valves with rows of bristles along
the growth lines. Rostrum (Fig.8a, b) and carina
similar in size, with slightly undulated rostro-carinal
articulation, not forming articular ridges; rostrum
and fíxed scutum articulation without large radius-like
projection, apex slightly projected, straight. Carina
(Fig.8a, b) without large radius-like projection
toward fixed tergum; apex projected and recurved.
Fixed tergum (Fig.8b) slightly higher than fixed
scutum, both sides with well-developed alar-like
projections; apex straight. Fixed scutum (Fig.8b) with
wide alar-like process directed toward rostrum and
a poorly defined radius-like process directed toward
fixed tergum, apex curved toward fixed tergum;
internally, without pit or ridge for adductor muscle.
Tergum (Fig.8a, c) nearly quadrangular, with only
a prominent axial ridge, having only growth lines
besides the axial ridge with a notch on their
projected surface; carinal area smooth. Internally
flat, and with a large concavity on most of its surface;
scutal margin straight except for a small
protuberance on upper portion. Scutum (Fig.8a, c)
straight, slightly smaller than tergum; with two
articular ridges; axial ridge projected, with both
borders conspicuous, thin; second ridge with same
width as axial ridge, without flat upper triangular
projection at scutal margin; rostral area smooth;
apex acu te. Internally, with a large concavity parallel
to occludent margin, which on upper portion
adductor muscle attaches; tergal margin straight,
except for a small notch at upper portion.
Fig.8- Altivemica sala sp.nov. Holotype: (a) rostro-carinal view; (b) fixed tergum and fixed scutum view; (c) tergum and
scutum, internai view; (c) carina; (fs) fixed-scutum; (ft) fixed-tergum; (r) rostrum; (s) scutum; (t) tergum.
Arq. Mus. Nac., Rio de Janeiro, v.60, n.l, p.5-40, jan./mar.2002
14
P.S.YOUNG
Labrum (Fig.9a) with a series of 43 simple sharp
teeth and multifid scales on the inner border but
large scales covering the outer border. Palp (Fig.9a)
short, acuminate, with few simple setae on inner
margin and distai region. Mandible (Fig.9b) with
three teeth, distance between first and second a
slightly more than distance between second and
third, third tooth tri-cuspid; lower angle
denticulated. Maxilla I (Fig.9c) with lower part
projecting strongly; 2 large and 3 short spines at
upper border, 3 median spines on lower projected
border. Maxilla II (Fig.9d) triangular, anterior
margin with shallow concavity medially; covered by
long simple setae, except on the shallow concavity.
Cirrus I (Fig. 10a) with equal rami, both rami covered
with several long simple setae. Cirrus II (Fig. 10b)
with unequal rami, anterior ramus about 2/3 length
of posterior, articles more protuberant; both rami
covered by numerous long, simple setae. Rami of
cirri III-VI equal in length. Intermediate articles of
cirrus VI (Fig. 10c) with width about 1/2 length; two
pairs of setae on anterior margin, longer pair finely
pinnate, smaller pair simple; one or two fine simple
setae and multifid setae on posterior angle. Caudal
appendage (Fig.lOd) with 16/17 articles, 3 times
length of protopodite. Penis smaller than coxopodite,
clothed with thin setulae. Number of articles of cirri
I-VI and caudal appendage is presented in table 3.
Etymology - from the Latim salum (the open sea).
Remarks - Altiverruca s.s. can be separated into two
groups of species: one encompassing those species
with a rostrum having a smooth or ribbed surface
above the upper articular ridge and directed toward
the scutum basis and the other in which the species
have the upper margin directly connected with the
articulation with the carina. The first group contains
12 species of Altiverruca s.s. and the second,
including A. sala, has 11 species. The latter is
represented by: A. obliqua (Hoek), A. quadrangularis
(Hoek), A. inermis (Aurivillius), A. plana (Gruvel), A.
spongicola (Gruvel), A. angustiterga Zevina, A.
sculpturata Zevina, A. sublima Zevina, A. gira (Zevina),
A. tchesunovi Zevina, A. beringiana Zevina & Galkin
and A. vertica Foster & Buckeridge (= A. obliqua).
Fig.9- Altiverruca sala sp.nov.: (a) labrum and palp; (b) mandible; (c) maxilla I; (d) maxilla II.
Arq. Mus. Nac., Rio de Janeiro, v.60, n.l, p.5-40, jan./mar.2002
THE VERRUCIDAE FROM THE WESTERN COAST OF NORTH AMERICA...
15
Altiverruca sala sp.nov. has a characteristic straight
scutum with two ridges: the axial and another on the
tergal margin. Only A. obliqua and A. quadrangularis
have similar opercular valves with the same number
of articular ridges. But both species do not have
projected growth lines, the tergum has a wide ridge
above the axial ridge in the former and the latter
species has a more elaborated carino-rostral
articulation. Furthermore, both species were described
from the Atlantic Ocean, but A. quadrangularis was
also recorded from the Philippines and Reunion Is.
(ROSELL, 1989: 24; FOSTER & BUCKERIDGE,
1995b: 367). A. sala sp.nov. is based on only two
specimens collected together with Newmaniverruca
digitiformis sp.nov. on Green Seamount, off Puerto
Vallarta, México, from 1850m.
TABLE 3
Number of articles on cirri I-VI,
and caudal appendage of Altiverruca sala sp.nov.
I
n
ffl
IV
V
VI
CA
RC
8/7
5/7
14/14
16/17
19/20
20/20
16
LC
8/7
5/6
14/14
15/17
18/20
20/20
17
(CI-VI) cirri I to VI; (CA) caudal appendage; (RC) right cirri;
(LC) left cirri.
Altiverruca vansyoci sp.nov.
(Figs. 11-14)
Material - Holotype: Off Oregon coast, 44°4.90’N,
125°24.50’W, 2938m, R/V Yaquina coll., 15 Jun
1972, rc: 5.3mm, on octocoral stem, CASIZ 138571.
Paratypes: Same locality, 8 specimen, rc: 2.6-
5.6mm, CASIZ 115100; MNRJ 14335.
Fig. 10- Altivermca sala sp.nov: (a) cirrus I; (b) cirrus II; (c) intermediate article of cirrus VI; (d) caudal appendage and
protopodite; (CI-CVI) cirri I to VI; (a.) anterior ramus; (c.a.) caudal appendage; (p.) posterior ramus.
Arq. Mus. Nac., Rio de Janeiro, v.60, n.l, p.5-40, jan./mar.2002
16
P.S.YOUNG
Diagnosis - Shell with growth lines not prominent
on all plates. Rostrum with undulated rostro-carinal
articulation, forming one upper articular ridge and
with a large surface above articular ridge varying
from smooth to with low ridges turned toward
scutum. Fixed tergum much higher than fixed
scutum with apex curved toward fixed scutum.
Tergum with only poorly developed axial ridge.
Scutum axial ridge slightly projected and a second
ridge slightly discernable beside axial ridge. Cirri I
and II with anterior rami slightly longer than
posterior. Intermediate articles of cirrus VI with two
pairs of setae on anterior margin. Caudal appendage
two times length of protopodite.
Description - Shell (Fig. 11a, b) white, opercular
valves perpendicular to base of wall, with growth
lines not prominent on all plates; shell plates
without longitudinal ridges; basal margin not
thickened. Cuticle not persistent on shell and
opercular valves. Rostrum (Fig.lla) rectangular,
with an undulated rostro-carinal articulation, forming
one upper articular ridge and sometimes a tooth
below (Figs.lla, 12), with a large surface above
articular ridge varying from smooth to with low
ridges turned toward scutum; rostrum and fixed
scutum articulation without large radius-like
projection, apex usually marginal, but sometimes
displaced anteriorly from margin, slightly projected,
straight. Carina (Fig. 11a) without radius-like projection
toward fixed tergum; apex slightly projected and
straight. Fixed tergum (Fig. 11b) much higher than
fixed scutum, both sides with well-developed
alar-like projections; apex curved toward fixed
scutum. Fixed scutum (Fig. 11b) with wide alar-like
process directed toward rostrum and a poorly
defined radii-like process directed toward fixed
tergum, apex straight; internally, without pit
or adductor ridge.
Tergum (Fig. 1 la, c) nearly quadrangular, with only
axial ridge, slightly prominent; carinal area smooth.
Fig. 11- Altivemica vansyoci sp.nov. Holotype: (a) rostro-carinal view; (b) fixed tergum and fixed scutum view; (c) tergum and
scutum, internai view; (c) carina; (fs) fixed-scutum; (ft) fixed-tergum; (r) rostrum; (s) scutum; (t) tergum.
Arq. Mus. Nac., Rio de Janeiro, v.60, n.l, p.5-40, jan./mar.2002
THE VERRUCIDAE FROM THE WESTERN COAST OF NORTH AMERICA...
17
Fig.12- Altiverruca vansyoci sp.nov.: Rostro-carinal suture of several specimens. Note the variation in the development of the tooth
below the large articular ridge of rostrum; the variation on the apex position, from marginal to displaced; (c) carina; (r) rostrum.
Internally, flat; scutal margin nearly straight.
Scutum (Fig. 1 la, c) smaller than tergum; axial ridge
slightly projected sloping gradually at scutal side,
thin; a second ridge slightly discernable beside axial
ridge, and a small, straight flat upper projection at
scutal margin; rostral area smooth. Internally, with
a large concavity for adductor muscle conspicuous
at occludent margin; tergal margin nearly straight.
Labrum (Fig. 13a) with a series of approximately 70
simple sharp teeth becoming two rows laterally and
some scattered teeth on the median area; large
scales covering the outer border. Palp (Fig. 13a)
short, triangular, wit h few simple setae on outer
margin. Mandible (Fig. 13b) with three teeth,
distance between first and second a slightly more
than distance between second and third, second
and third teeth with upper margin denticulated;
lower angle denticulated. Maxilla I (Fig. 13c) with
lower part projected; 2 large and 3 short spines at
upper border, 4 median spines on lower projected
border. Maxilla II (Fig.l3d) rounded, anterior margin
with shallow concavity medially; covered by long
simple setae, except on the shallow concavity.
Cirrus I (Fig. 14a) with anterior rami a few segments
longer than posterior, both rami covered with
several long simple setae. Cirrus II (Fig. 14b) with
posterior ramus a few segments longer than anterior,
articles more protuberant on anterior ramus; both
rami covered by numerous long, simple setae. Rami
of cirri III-VI equal in length. Intermediate articles
of cirrus VI (Fig. 14c) with width 1 /2 length; two pairs
of setae on anterior margin, longer pair finely
pinnate and smaller pair simple; one or two fine
simple setae on posterior angle. Caudal appendage
(Fig. 14d) with 9-10 articles, two times length of
protopodite; long simple setae on distai margins of
articles. Penis (Fig. 14d) smaller than coxopodite,
clothed with thin setulae. Number of articles of cirri
I-VI and caudal appendage is presented in table 4.
TABLE 4
Number of articles on cirri I-VI, and caudal appendage of
Altiverruca vansyoci sp.nov.
I
n
m
IV
V
VI
CA
RC
9/7
7/9
15/16
15+/15+
16+/14+
18+17+
10
LC
9/8
7/9
15/16
19+/18+
18+/15+
19+14+
9
(CI-VI) cirri I to VI; (CA) caudal appendage; (RC) right cirri;
(LC) left cirri; (+) broken ramus.
Arq. Mus. Nac., Rio de Janeiro, v.60, n.l, p.5-40, jan./mar.2002
18
P.S.YOUNG
Fig. 13- Altiverruca vansyoci sp.nov.: (a) labrum and palp; (b) mandible; (c) maxilla I; (d) maxilla II.
Etymology - Named in honor of Dr. Robert J. Van
Syoc, cirripedologist who was largely responsible
for making the Califórnia Academy of Science
specimens available for this study.
Remarks - Altiverruca vansyoci sp.nov. is included
in the Altiverruca group in which the species have a
rostrum with a flat or ribbed surface above the upper
articular ridge and directed toward the base of the
scutum (see remarks under A. sala sp.nov.). This
group contains 12 species: A. incerta (Hoek), A.
erecta (Gruvel), A. longicarinata (Gruvel), A. hoeki
(Pilsbry), A. joubini (Gruvel), A. casula (Hoek), A.
ornata Nilsson-Cantell, A. aves (Zevina), A.
galapagosa Zevina, A. longa Zevina, A. vitrea Zevina,
A. galkini Zevina. Of these, five species have very
distinct opercular plates, with the terga with two
conspicuous ridges (A. aves, A. galapagosa, A.
incerta, A. longa and A. vitrea ).
Of the remaining species, A. erecta, A. galkini, A.
joubini and A. ornata have the rostro-carinal suture
straight or undulated without an upper articular
ridge on the rostrum as seen in A. vansyoci sp.nov.
Therefore three species of Altiverruca s.s. have similar
opercular valves and rostro-carinal sutures: A.
longicarinata, A. hoeki and A. casula. Altiverruca
longicarinata was described from the Sargasso Sea
(GRUVEL, 1900: 242; 1902: 91) and subsequently
recorded from the Guiana Basin and Mid-Atlantic
Ridge (ZEVINA, 1987b: 1305; YOUNG, 1998c:113).
In this species the rostrum has a smooth surface
above the articular ridge; the apexes of rostrum and
carina are more projected; the mandible has the 2 nd
and 3 rd teeth with smooth upper margin (based on
Zevina’ s specimens). Altiverruca hoeki was described
from Anegada Passage, Lesser Antilles (PILSBRY,
1907) and subsequently recorded from the Grenada
Basin (ZEVINA, 1975). It has the terga squared, the
rostro-carinal suture is less undulated with the upper
ridge less pronounced, the apexes of rostrum and
carina are more projected, the apex of the fixed
tergum is straight; and the fixed scutum with narrow
radii-like process directed toward the fixed tergum,
Arq. Mus. Nac., Rio de Janeiro, v.60, n.l, p.5-40, jan./mar.2002
THE VERRUCIDAE FROM THE WESTERN COAST OF NORTH AMERICA...
19
Fig. 14- Altivermca vansyoci sp.nov.: (a) cirrus I; (b) cirrus II; (c) intermediate article of cirrus VI; (d) caudal appendage, penis
and protopodite; (CI-CVI) cirri I to VI; (a.) anterior ramus; (c.a.) caudal appendage; (p.) posterior ramus; (pe.) penis.
restricted to the lower part. Altiverruca casula was
described from the Ceram Sea, Indonésia (HOEK,
1913). It has a scutum with two conspicuous
ridges, the rostrum with a smooth surface above
the articular ridge; fixed tergum with a large
rounded projection directed toward the fixed-rostrum.
Altiverruca vansyoci, besides the differences
described above, was collected off the coast of
Oregon, 44°4.90’N, 125°24.50’W, far from the
localities of the related species.
Gibbosaverruca gen.nov.
(Fig. 15)
Verruca Section D: Altiverruca Pilsbiy, 1916:40 (part).
Verruca (Altiverruca) - BROCH, 1931:45 (part);
FOSTER, 1978:68 (part).
Altiverruca - ZEVINA, 1987a: 1813 (part);
BUCKERIDGE, 1994:92 (part); YOUNG, 1998a:
77 (part).
Type species - Verruca gibbosa Hoek, 1883, Recent,
Challenger Expedition, Argentine Basin, sta. 317,
48°37’S, 55°17W, 1035 fathoms [1892m].
Etymology - from the Latin gibbosus (protuberant)
and verruca (wart); feminine gender.
Diagnosis - Shell large and inclined, wall plates
massive. Rostrum triangular, same height of carina,
usually without a flat or ribbed surface between
the uppermost articular ridge and the base of
scutum. Rostrum and carina United by strong
ridges, the uppermost on rostrum being the largest.
Shell plates varying from smooth to strongly
ornamented. Operculum obliqúe to basis inclined
more than 45° and also obliqúe to fixed-scutum and
tergum. Scutum and tergum with more than two
conspicuous ridges on each valve. Fixed scutum
without adductor ridge or myophore. Bases of wall
plates not inflected.
Species - G. gibbosa (Hoek, 1883), G. nitida (Hoek,
1883), G. sulcata (Hoek, 1883), G. mitra (Hoek, 1907),
? G. darwini (Pilsbry, 1907), G. navicula (Hoek, 1913),
G. rathbuniana (Pilsbry, 1916), G. costata (Aurivillius,
1898), G. somaliensis (Nilsson-Cantell, 1929), G.
montereyi sp. nov., G. robusta sp.nov., G. mateoi sp.nov.
Arq. Mus. Nac., Rio de Janeiro, v.60, n.l, p.5-40, jan./mar.2002
20
P.S.YOUNG
Remarks - Most of the species included in this
genus were considered synonyms of G. gibbosa
(NILSSON-CANTELL, 1928; 1938; 1955; TARASOV
& ZEVINA, 1957; ZEVINA, 1976; FOSTER &
BUCKERIDGE, 1995a) but others suggested it was
a species complex (NEWMAN & ROSS, 1971;
YOUNG, 1998a).
HOEK (1883), based on the Challenger Expedition
samples, described Gibbosaverruca gibbosa from
the Southern Atlantic, and it has been recorded
several times from different oceans. Five described
species and one subspecies were considered
synonyms of G. gibbosa including Verruca sulcata
Hoek, 1883, described from the South Fiji Basin;
V. mitra Hoek, 1907, from off Antarctic Peninsula;
V. bicomuta Pilsbry, 1916 from East of Nantucket,
Massachusetts; V. darwini Pilsbry, 1907, from
Cape May, New Jersey; V. rathbuniana Pilsbry,
1916, from Southeast of Cape Cod,
Massachusetts; and V. gibbosa somaliensis
Nilsson-Cantell, 1929 from the coast of Somalia
(NILSSON-CANTELL, 1928: 25; TARASOV &
ZEVINA, 1957: 154). When the descriptions and
figures of these species are examined in detail,
there are two conclusions: either Gibbosaverruca
gibbosa is a very plastic species having an
exceptional morphological variation, or it actually
represents a complex of species.
Based on the series of characters listed in table 5,1
propose that Gibbosaverruca gibbosa is a group of
species and all of the previously described species
have to be reevaluated. In the G. gibbosa complex,
considered herein as Gibbosaverruca gen.nov., 6
species should be included: G. nitida (Hoek, 1883),
G. navicula (Hoek, 1913), G. costata (Aurivillius,
1898), and the three new species described bellow.
These species are relatively large with the shell
having pronounced growth lines and well-developed
articular ridges on the opercular plates, rostrum
and carina. G. gibbosa somaliensis is elevated to
species status; G. darwini is tentatively included in
this genus, but it does not have the upper articular
ridge of the rostrum much larger than the one on
the carina. Figure 15 presents the type localities of
all species of Gibbosaverruca gen.nov. excluding the
previous synonyms.
20° 0 o 20° 60° 100° 140° 180° 140°
Fig. 15- Type localities of all species included in Gibbosaverruca s.s.
Arq. Mus. Nac., Rio de Janeiro, v.60, n.l, p.5-40, jan./mar.2002
Arq. Mus. Nac., Rio de Janeiro, v.60, n.l, p.5-40, jan./mar.2002
TABELA 5
DIFFERENTIAL CHARACTERS OF THE SPECIES OF GIBBOSAVERRUCA GIBBOSA GROUP INCLUDING THE NEW SPECIES.
COSTATOVERRUCA BICORNUTA (PILSBRY, 1916) IS NOT INCLUDED, DUE ITS TRANSFERENCE TO OTHER GENUS
species
character^^^^
G. gibbosa
G. sulcata
G. mitra
G darwini
G
rathbuniana
G
somaliensis
G monterey
sp.nov.
G. robusta
sp.nov.
G. mateoi
sp.nov.
Scutum: articular
ridges (No)
3
3
2
2
2
2
2
3
2
Scutum: marginal
articular ridge
Vestigial,
marginal
Vestigial,
marginal
Absent
Absent
Absent
Absent
Absent
Absent
Absent
Scutum:
intermediate
articular ridge
Protuberant,
curved, near
and with
same width
of axial ridge
Protuberant,
curved, near
and with
same width
of axial ridge
Protuberant,
curved, near
and with
same width
of axial ridge
Protuberant,
curved, near
and with
same width
of axial ridge
Protuberant,
curved, near
and with
same width
of axial ridge
Protuberant,
curved, near
and with two
times wider
than axial
ridge
Protuberant,
curved, near
and wider
than axial
ridge
Protuberant,
curved, near
and wider
than axial
ridge
Protuberant,
curved, as
wide as axial
ridge
Scutum:
accessory
articular ridge
Absent
Absent
Absent
Absent
Absent
Absent
Absent
Protuberant,
curved, near
and thinner
than 2 nd
articular ridge
Absent
Scutum: axial
ridge
Conspicuous,
with both
sides well
marked
Conspicuous,
with both
sides well
marked
Conspicuous,
with only
tergal side
well marked
Conspicuous,
with only
tergal side
well marked
Conspicuous,
with both
sides well
marked
Conspicuous,
with both
sides well
marked
Conspicuous,
with only
tergal side
well marked
Conspicuous,
with only
tergal side
well marked
Conspicuous,
with both
sides well
marked
Tergum: articular
ridges (No)
3
3
3
3
3
3
3
4
3
Tergum: marginal
articular ridge
Marginal,
thin
Marginal,
wide
Marginal,
wide
Marginal,
thin
Marginal,
thin
Marginal,
thin
Marginal,
wide
Marginal, thin
Marginal,
thin
Tergum:
intermediate
articular ridge
Protuberant,
curved, near
and with
same width
of axial ridge
Protuberant,
curved, near
and with
same width
of axial ridge
Slightly
protuberant,
curved,
medial, with
same width
of axial ridge
Slightly
protuberant,
curved,
medial, wider
than axial
ridge
Slightly
protuberant,
curved,
medial, wider
than axial
ridge
Slightly
protuberant,
curved,
medial, wider
than axial
ridge
Slightly
protuberant,
curved,
medial, wider
than axial
ridge
Slightly
protuberant,
straight,
medial, wider
than axial
ridge
Protuberant,
curved,
medial, as
wide as axial
ridge
Tergum:
accessory
articular ridge
Absent
Absent
Absent
Absent
Absent
Absent
Absent
Protuberant,
straight, near
and with
same width of
axial ridge
Absent
Scutum/tergum
basis length ratio
1/2
2/5
2/3
2/3
2/3
4/5
3/4
1/2
2/3
Rostrum: No and
development of
articular ridges
3, decreasing
in width from
apex to basis
4, of same
width
2 , of same
width
3, l st
smaller, 2nd
largest
3, l st
smaller, 2nd
largest
3, decreasing
in width from
apex to basis
5, uppermost
smaller, 2 nd
larger and
decreasing in
width to
basis
3, medial
wider
3, decreasing
in width from
apex to basis
continue...
THE VERRUCIDAE FROM THE WESTERN COAST OF NORTH AMERICA...
Arq. Mus. Nac., Rio de Janeiro, v.60, n.l, p.5-40, jan./mar.2002
...conclusion
K>
to
species
characters
G. gibbosa
G. sulcata
G mitra
G darwini
G
rathbuniana
G
somáliensis
G monterey
sp.nov.
G. robusta
sp.nov.
G mateoi
sp.nov.
Carina: No and
development of
articular ridges
3, decreasing
in width from
apex to basis
3, l st largest, 3, l st largest,
2 nd and 3 rd 2 nd and 3 rd
equal in width equal in width
3, decreasing
in width from
apex to basis
3, decreasing
in width from
apex to basis
4, decreasing
in width from
apex to basis
5, uppermost
smaller, 2 nd
larger and
decreasing in
width to basis
3, medial
wider
3, decreasing
in width from
apex to basis
Length of fixed-
Tergum
Higher than
fixed scutum
Higher than
fixed scutum
Higher than
fixed scutum
As high as
scutum
Higher than
fixed scutum
Higher than
fixed scutum
Higher than
fixed scutum
Higher than
fixed scutum
Higher than
fixed scutum
Rostrum: flatter
surface directed to
scutum
Absent
Absent
Absent
Present, well
developed
Absent
Absent
Absent
Absent
Present, well
developed,
with ridge
Fixed-scutum:
Alar- / radii-like
processes width
ratio
2 times
1/2
3 times
2 times
2 times
2 times
2 times
3 times
2 times
Fixed-tergum:
carinal/fixed-scutum
alar-like processes
width ratio
1/2
Not visible in
figure
1/2
Equal
1/2
Equal
Equal
Equal
1/2
Rostrum: apex
Incurved
Incurved
Straight
Incurved
Straight
Straight
Recurved
Straight
Incurved
Carina: apex
Recurved
Recurved
Recurved
Straight
Recurved
Recurved
Recurved
Recurved
Recurved
Cirrus I: anterior
and posterior rami
length
Equal
-
-
Subequal
Subequal
Une qual
Equal
Subequal
Subequal
Cirrus II: anterior
and posterior rami
length
1/2
-
-
2/3
4/5
1/2 to 1/3
2/3
2/3
1/2
Cirrus II: anterior
and posterior rami
length
Subequal
-
-
Subequal
Subequal
Equal
Equal
Equal
Equal
Cirrus VI: No of
pairs of setae on
anterior margin of
articles
3
-
-
-
-
3
2
3
Caudal appendage:
No of articles
11
-
-
11
-
18
20
17
9
Caudal appendage
/protopodite of
cirrus VI length
11/2 times
-
-
11/3 times
-
Larger than
protopod
2 times
2 times
Little longer
omoÀSd
THE VERRUCIDAE FROM THE WESTERN COAST OF NORTH AMERICA...
23
Gibbosavermca montereyi sp.nov.
(Figs. 16-18)
Material - Holotype, Monterey Submarine Canyon,
Monterey, Califórnia, 36°46’28.7”N, 121°59’27.5”W,
750m, J. Barry coll. 17 Aug 1992, rc: 13.0, SIO-BIC
C 9890; Paratypes: same locality, 2 specimens, rc:
12.9 and 3.9mm, SIO-BIC C 10304; MNRJ 14336.
Diagnosis - Shell with growth lines prominent on
all plates. Rostrum with five flat articular ridges,
uppermost very small, second largest, decreasing
in size toward basis. Tergum quadrangular, with
three articular ridges; axial ridge well prominent;
intermediate ridge wide. Scutum with two articular
ridges; axial ridge thin; second ridge twice width
of axial ridge. Cirrus I with equal rami. Cirrus II
with anterior ramus about 2/3 length of posterior.
Intermediate articles of cirrus VI with three pairs
of simple setae on anterior margin. Caudal
appendage two times length of protopodite.
Description - Shell (Fig.l6a, b) white, opercular
valves obliqúe to base of wall, with growth lines well
prominent on all plates; rostrum and carina
undulated, forming large and low longitudinal
ridges, fixed tergum and scutum without
longitudinal ridges; basal margin not thickened.
Cuticle yellowish, smooth, persistent on shell and
opercular valves. Rostrum (Fig.l6a) and carina
similar in size, former with five flat articular ridges,
with shallow grooves between them on the rostro-
carinal articulation, uppermost very small, second
largest, decreasing in size toward basis; rostrum
and fixed scutum articulation with large radius-
like projection, apex projected. Carina (Fig. 16a) with
five articular ridges directed toward scutum,
uppermost very small, others of similar width or
with second larger, with shallow grooves between
them, with large radius-like projection toward fixed
tergum; apex curved and projected. Fixed tergum
(Fig. 16b) higher than fixed scutum, both sides with
well-developed alar-like projections; apex straight.
Fig. 16- Gibbosavermca montereyi sp.nov.: (a) rostro-carinal view; (b) fixed tergum and fixed scutum view; (c) tergum and
scutum, internai view.
Arq. Mus. Nac., Rio de Janeiro, v.60, n.l, p.5-40, jan./mar.2002
24
P.S.YOUNG
Fixed scutum (Fig. 16b) with wide alar-like process
directed toward rostrum twice width of radii-like
process directed toward fixed tergum; internally,
with a pit for adductor muscle and a narrow
projected adductor ridge.
Tergum (Fig. 16a, c) quadrangular, with three articular
ridges; axial ridge prominent, narrower than
intermediate ridge; intermediate ridge wide, upper
ridge thin and marginal to occludent margin, with a
large conspicuous depression between upper and
intermediate ridges; carinal area smooth. Internally,
smooth and flat; scutal margin straight except for
protuberance on upper portion. Scutum (Fig. 16a, c)
smaller than tergum; with two articular ridges; axial
ridge somewhat triangular in shape with scutal border
conspicuous and sloping gradually to rostral surface,
thin; second ridge twice width of axial ridge, and a
flat upper triangular projection at scutal margin;
rostral area smooth. Internally, with a large concavity
on upper portion for adductor muscle; tergal margin
straight, except for notch at upper portion.
Labrum (Fig. 17a) with a series of simple sharp teeth,
rarely bifid or multifid. Palp (Fig. 17a) short, acuminate,
with simple setae on inner margin and distai
region. Mandible (Fig. 17b) with three teeth,
distance between first and second twice distance
between second and third, upper margin of third
tooth with accessoiy denticle; lower angle denticulated.
Maxilla I (Fig. 17c) with lower part slightly projected
and with a shallow concavity in middle; two large
and one median spines at upper angle, 10 small
spines in the concavity, and lower projected part,
with two large spines medially positioned, five
median-size spines on upper portion and seven
small spines on lower portion. Maxilla II (Fig. 17d)
triangular, anterior margin with shallow concavity
medially; covered by long simple setae, except on
the shallow concavity.
Cirrus I (Fig. 18a) with equal rami, both rami with
wide basal articles tapering toward distai portion,
covered with several long simple setae. Cirrus II
(Fig. 18b) with unequal rami, anterior ramus about
2/3 length of posterior, articles more protuberant;
both rami covered by numerous long, simple setae.
Fig. 17- Gibbosaverruca montereyi sp.nov.: (a) labrum and palp; (b) mandible; (c) maxilla I; (d) maxilla II.
Arq. Mus. Nac., Rio de Janeiro, v.60, n.l, p.5-40, jan./mar.2002
THE VERRUCIDAE FROM THE WESTERN COAST OF NORTH AMERICA...
25
Rami of cirri III-VI equal in length. Intermediate
articles of cirrus VI (Fig.l8c) with width equal to
length; three pairs of simple setae on anterior
margin; none to two fine simple setae on posterior
angle, multifid scales at distai margin. Caudal
appendage (Fig. 18d, e) with 20 articles, two times
length of protopodite; long simple setae on distai
margins of articles. Penis smaller than coxopodite,
clothed with thin setulae. Number of articles of cirri
I-VI and caudal appendage is presented in table 6.
Etymology - From Monterey, Califórnia, the locality
where it was collected.
Remarks- Gibbosaverruca montereyi sp.nov. and G.
robusta sp.nov. were collected at the same locality,
off Monterey, and both are large verrucids having
rostro-carinal diameters reaching more than lOmm.
At first, I thought they were variants of the same
species, but G. montereyi is easily distinguished be
the presence of two articular ridges on the scutum
and three on the tergum instead of respectively, three
and four in G. robusta. The differential characters of
these new species and that species of the
Gibosaverruca gibbosa group are listed in table 5.
TABLE 6
Number of articles on cirri I-VI, and caudal appendage of
Gibbosaverruca montereyi sp.nov.
I
n
m
IV
V
VI
CA
RC
27/24
16/26
31/34
36/37
37+/42
36+/42
15+
LC
27/25
18/26
29/33
35/37
40/42
41/41
20
(CI-VI) cirri I to VI; (CA) caudal appendage; (RC) right cirri;
(LC) left cirri; (+) broken ramus.
Fig. 18- Gibbosaverruca montereyi sp.nov.: (a) cirrus I; (b) cirrus II; (c) intermediate article of cirrus VI; (d) caudal appendage
and protopodite; (e) intermediate article of caudal appendage; (CI-CVI) cirri I to VI; (a.) anterior ramus; (c.a.) caudal appendage;
(p.) posterior ramus.
Arq. Mus. Nac., Rio de Janeiro, v.60, n.l, p.5-40, jan./mar.2002
26
P.S.YOUNG
Gibbosavermca robusta sp.nov.
(Figs. 19-21)
Material - Holotype: Monterey Submarine Canyon,
Monterey, Califórnia, 36°46’28.7”N, 121°59’27.5”W,
750m, J. Barry coll. 17 Aug 1992, rc: 12.4mm,
SIO-BIC C 10239. Paratypes: same locality, 2
specimens, one specimen lacking rostrum, one
specimen only with carina, opercular plates and
body, SIO-BIC C 10304, MNRJ 14337.
Diagnosis - Shell with growth lines prominent on
all plates. Rostrum with three articular ridges,
intermediate widest. Tergum with four articular
ridges; axial ridge prominent with another similar
ridge beside it, intermediate ridge wider than axial
ridge. Scutum with three articular ridges; axial
ridge thin; second ridge twice width of axial ridge.
Cirrus I with anterior ramus slightly smaller than
posterior. Cirrus II with anterior ramus about 2/3
length of posterior. Intermediate articles of cirrus VI
with two pairs of setae. Caudal appendage two
times length of protopodite.
Description - Shell (Fig.l9a, b) white, opercular
valves obliqúe to base of wall, with growth lines
prominent on all plates; rostrum and carina
undulated, forming large and low longitudinal
ridges, fixed tergum and scutum without
longitudinal ridges; basal margin not thickened.
Cuticle smooth, persistent on shell and opercular
valves. Rostrum (Fig.l9a) and carina similar in
size, former with three articular ridges, with
shallow grooves between them on the rostro-carinal
articulation, intermediate largest; rostrum and
fixed scutum articulation without radius-like
projection, apex straight, projected.
Fig. 19- Gibbosavermca robusta sp.nov.: (a) rostro-carinal view; (b) fixed tergum and fixed scutum view; (c) tergum and
scutum, internai view.
Arq. Mus. Nac., Rio de Janeiro, v.60, n.l, p.5-40, jan./mar.2002
THE VERRUCIDAE FROM THE WESTERN COAST OF NORTH AMERICA...
27
Carina (Fig.l9a) with four articular ridges directed
toward scutum, second largest, others small, with
shallow grooves between them, with large radius-like
projection toward fixed tergum; apex curved and
projected. Fixed tergum (Fig.l9b) higher than fixed
scutum, both sides with well-developed alar-like
projections; apex straight. Fixed scutum (Fig.l9b)
with wide alar-like process directed toward rostrum
three times width of radii-like process directed
toward fixed tergum; intemally, with a pit for adductor
muscle and a small projected adductor ridge.
Tergum (Fig. 19a, c) quadrangular, with four articular
ridges; axial ridge prominent with another similar
ridge beside it, intermediate ridge wider than axial
ridge; upper ridge thin and marginal to occludent
margin, with a large conspicuous depression between
upper and intermediate ridges; carinal area smooth.
Internally, smooth and flat; scutal margin straight
except for protuberance on upper portion. Scutum
(Fig. 19a, c) smaller than tergum; with three articular
ridges; axial ridge somewhat triangular in shape with
scutal border conspicuous and sloping gradually to
rostral surface, thin; second ridge twice width of axial
ridge, third ridge thin, and a flat upper triangular
projection at scutal margin; rostral area smooth.
Internally, with a large concavity on upper portion
for adductor muscle; tergal margin straight, except
for notch at upper portion.
Labrum (Fig.20a) with a series of simple sharp teeth,
rarely bifid or multifid. Palp (Fig.20a) short,
acuminate, with simple setae on inner margin and
distai region. Mandible (Fig.20b) with three teeth,
distance between first and second three times
distance between second and third; lower angle
denticulated. Maxilla I (Fig.20c) with lower part
slightly projected; 2 large spines at upper angle, 19
small spines in the middle followed by 4 large spines
medially positioned and 10 small spines on basal
projection. Maxilla II (Fig.20d) triangular, anterior
margin with concavity medially; covered by long
simple setae, except on the shallow concavity.
Papillae of maxillary gland small, rounded.
Fig.20- Gibbosaverruca robusta sp.nov.: (a) labrum and palp; (b) mandible; (c) maxilla I; (d) maxilla II.
Arq. Mus. Nac., Rio de Janeiro, v.60, n.l, p.5-40, jan./mar.2002
28
P.S.YOUNG
Cirrus I (Fig.21a) with subequal rami, anterior a
little smaller than posterior, both rami with wide
basal articles tapering to distai portion, covered with
several long simple setae. Cirrus II (Fig.21b) with
unequal rami, anterior ramus about 2/3 length of
posterior; both rami covered by numerous long,
simple setae. Rami of cirri III-VI equal in length.
Intermediate articles of cirrus VI (Fig.2 lc) with width
equal to length; 2 pairs of long simple setae and 1
simple setulae on anterior margin; 2 or 3 simple
setae on posterior angle, multifid scales at distai
margin. Caudal appendage (Fig.2ld, e) with 17
articles, two times length of protopodite; long simple
setae on distai margins of articles. Penis smaller
than coxopodite, clothed with thin setulae. Number
of articles of cirri I-VI and caudal appendage is
presented in table 7.
Etymology - From the Latin robustus, massive,
referring to the large size of this species.
Remarks - The comparison of this species with others
of the G. gibbosa group is presented in table 5. G.
robusta is easily characterized by its large size, with
the growth lines very prominent on all plates ant the
rostrum having three and tergum four articular ridges.
None of other species of these group have this large
number of ridges on the scutum and tergum.
TABLE 7
Number of articles on cirri I-VI, and caudal appendage of
Gibbosavemica robusta sp.nov.
I
II
III
IV
V
VI
CA
RC
25/24
17/29
30/31
35/38
38/44
30+30+
17
LC
24+/24
20/27
25/33
33/43
38/40
42/44+
17
(CI-VI) cirri I to VI; (CA) caudal appendage; (RC) right cirri;
(LC) left cirri; (+) broken ramus.
Fig.21- Gibbosaverruca robusta sp.nov.: (a) cirrus I; (b) cirrus II; (c) intermediate article of cirrus VI; (d) caudal appendage
and protopodite; (e) intermediate article of caudal appendage; (CI-CVI) cirri I to VI; (a.) anterior ramus; (c.a.) caudal
appendage; (p.) posterior ramus.
Arq. Mus. Nac., Rio de Janeiro, v.60, n.l, p.5-40, jan./mar.2002
THE VERRUCIDAE FROM THE WESTERN COAST OF NORTH AMERICA...
29
Gibbosavermca mateoi sp.nov.
(Figs.22-24)
Material - Holotype: Califórnia, San Mateo County,
Off San Mateo Point, 9.6 miles, 768-823m,
G.D.Hanna col., “N.B. Scofield”, 23 Jan 1953, on
Hyalonema spicule, one specimen, rc: 5.3mm,
CASIZ 061442.
Diagnosis - Shell with growth lines prominent on
all plates. Rostrum with three articular ridges and
one small turned toward scutum. Tergum with three
articular ridges. Scutum with two articular ridges.
Cirrus I with anterior ramus a little smaller than
posterior. Cirrus II anterior ramus about 1/2 length
of posterior. Intermediate articles of cirrus VI with three
pairs of setae on anterior margin. Caudal appendage a
little longer than protopodite.
Description - Shell (Fig.22a, b) white, opercular valves
obliqúe to base of wall, with growth lines prominent
on all plates, without longitudinal ridges; basal margin
not thickened. Cuticle not persistent on shell and
opercular valves. Rostrum (Fig.22a, b) and carina
with similar size, former with three articular ridges
and one small turned toward scutum, with shallow
grooves between them on the rostro-carinal articulation,
upper ridge largest; rostrum and fixed scutum
articulation with small radius-like projection,
apex incurved, projected. Carina (Fig.22a, b)
with three articular ridges directed toward
scutum, first and second of similar size, with shallow
grooves between them; with large radius-like projection
toward fixed tergum; apex recurved and projected. Fixed
tergum (Fig.22a) higher than fixed scutum, both
sides with well-developed alar-like projections;
Fig.22- Gibbosaverruca mateoi sp.nov.: (a) rostro-carinal view; (b) fixed tergum and fixed scutum view; (c) tergum, internai
view, (d) scutum, internai view.
Arq. Mus. Nac., Rio de Janeiro, v.60, n.l, p.5-40, jan./mar.2002
30
P.S.YOUNG
apex curved toward fixed scutum. Fixed scutum
(Fig.22a) with wide alar-like process directed toward
rostrum two times width of radii-like process directed
toward fixed tergum, apex curved to fixed tergum;
internally, with an apical pit for adductor muscle.
Tergum (Fig.22a, c) quadrangular, with three
articular ridges; axial ridge well prominent, curved,
intermediate ridge as wide as axial ridge; upper ridge
thin and marginal to occludent margin, with a large
conspicuous depression between ridges; carinal
area smooth. Internally, smooth and flat; seu tal
margin undulated. Scutum (Fig.22a, c) smaller than
tergum; with two articular ridges; axial ridge
prominent, thin; second ridge as wide as axial ridge,
and a flat upper triangular projection at scutal
margin; rostral area smooth. Internally, with a
large concavity on upper portion for adductor
muscle; tergal margin nearly straight, except by a
medio-basal projection of second ridge.
Labrum (Fig.23a) with a series of 41 simple sharp
teeth. Palp (Fig.23a) acuminate, with simple setae on
margins. Mandible (Fig.23b) with three teeth, third
tooth with a small denticle on upper margin, distance
between first and second 1.5 distance between
second and third; lower angle denticulated. Maxilla
I (Fig.23c) with lower part projecting; two large and
one median-size spine at upper angle, two small,
strong spines in the middle followed by nine médium to
large spines on lower portion of basal projection.
Maxilla II (Fig.23d) triangular, anterior margin with
shallow concavity medially; covered by long simple
setae, except in the shallow concavity.
Cirrus I (Fig.24a) with subequal rami, anterior a little
smaller than posterior, both rami with wide basal
articles tapering to distai portion, covered with
several long simple setae. Cirrus II (Fig.24b) with
unequal rami, anterior ramus about 1/2 length of
posterior; both rami covered by numerous long,
Arq. Mus. Nac., Rio de Janeiro, v.60, n.l, p.5-40, jan./mar.2002
THE VERRUCIDAE FROM THE WESTERN COAST OF NORTH AMERICA...
31
Fig.24- Gibbosaverruca mateoi sp.nov.; (a) cirrus I; (b) cirrus II; (c) intermediate article of cirrus VI; (d) caudal appendage,
penis and protopodite; (CI-CVI) cirri I to VI; (a.) anterior ramus; (c.a.) caudal appendage; (p.) posterior ramus; (pe.) penis.
simple setae and distai articles of anterior ramus
also with pinnate setae. Rami of cirri III-VI equal in
length. Intermediate articles of cirrus VI (Fig.24c)
with width 1/2 the length; three pairs of simple setae
on anterior margin; one or two simple setae on
posterior angle. Caudal appendage (Fig.24d) with 9
articles, a little longer than protopodite; long simple setae
on distai margins of articles. Penis (Fig.24d) longer than
protopodite, apically clothed with thin setulae.
Number of articles of cirri I-VI and caudal appendage
is presented in table 8.
Remarks - Gibbosaverruca mateoi sp.nov. is not a veiy
large species like the other two described above: about
5 mm or rostro-carinal length. It has two articular
ridges on scutum and three on tergum. Therefore, its
number of ridges on the opercular plates are equal to
those of G. mitra, G. darwini, G. rathbuniana, G.
somaliensis, and G. montereyi But G. mateoi presents
the rostrum with a surface above the upper
articular ridge with one small ridge turned toward
scutum that distinguishes it from all other
Gibbosaverruca.
The comparison of this species with others of the
G. gibbosa group is presented in table 5.
TABLE 8
Number of articles on cirri I-VI, and caudal appendage of
Gibbosaverruca mateoi sp.nov.
I
n
m
IV
V
VI
CA
RC
11/10
7/12
17/21
21/22
24/26
26/26
9
LC
11/10
7/1+
3+/20
21/12+
24/23
27/26
9
(CI-VI) cirri I to VI; (CA) caudal appendage; (RC) right cirri;
(LC) left cirri; (+) broken ramus.
Arq. Mus. Nac., Rio de Janeiro, v.60, n.l, p.5-40, jan./mar.2002
32
P.S.YOUNG
Cristallinavemica gen.nov.
(Fig.25)
Vemica Section D: Altivermca Pilsbry, 1916:40 (part).
Verruca (Altivermca) - BROCH, 1931:45 (part);
FOSTER, 1978:68 (part).
Altiverruca - ZEVINA, 1987a: 1813 (part);
BUCKERIDGE, 1994:92 (part); YOUNG, 1998a:
77 (part).
Type species - Verruca cristallina Gruvel, 1907,
Recent, R/V Investigator, Andaman Islands, Cap
Bluff, 768m and Andaman Islands, Cinque
Island, 785m.
Etymology - from the Latin cristallu (limpid) and
verruca (wart); feminine gender.
Diagnosis - Shell moderate in size and inclined, wall
plates massive. Rostrum nearly rectangular to
triangular, same height of carina, rarely with a flat
or ribbed surface between the uppermost articular
ridge and the base of scutum. Rostrum and carina
suturai area formed by several imbricating ridges
similar in size. Shell plates usually smooth except
for articular ridges. Operculum obliqúe to basis
inclined more than 45° and also obliqúe to fixed-
scutum and tergum. Scutum and tergum with
conspicuous two to five ridges, usually ornamented.
Fixed scutum without adductor ridge or myophore.
Bases of wall plates not inflected.
Species - C. cristallina (Gruvel, 1907), C. crenata
(Aurivillius, 1898), C. cassis (Hoek, 1913) (= C.
cristallina), C. bicomuta( Pilsbiy, 1916), C. cristallina
laevis (Broch, 1922) (= C. cristallina), C. regularis
(Nilsson-Cantell, 1929), C. cristallina typica (Broch,
1931) (= C. cristallina), C. allisoni (Rao and Newman,
1972), C. mollae (Zevina, 1990), ? C. jonesae
(Buckeridge, 1997).
Remarks- Cristallina verruca gen.nov. encompasses
species of moderate size that have fully developed
articular ridges on the opercular plates, rostrum
and carina. The ridges on the rostral and carinal
suture are similar in size, making a characteristic
imbricating pattern. All of the species agree with
the proposed diagnosis, except for C. jonesae, which
has an ornamented shell. In figure 25 the type
localities of all the species of Cristallinaverruca
gen.nov. are shown excluding the previous synonyms.
None of the species included in the genus were
studied herein.
100° 60° 20° 0 o 20° 60° 100° 140° 180° 140° 100° 60°
Fig.25- Type localities of all species included in Cristallinaverruca s.s.
Arq. Mus. Nac., Rio de Janeiro, v.60, n.l, p.5-40, jan./mar.2002
THE VERRUCIDAE FROM THE WESTERN COAST OF NORTH AMERICA.
33
Genus Newmanivemica Young, 1998
Newmaniverruca digitiformis sp.nov.
(Figs.26-28)
Material - Holotype: Green Seamount at 21°N on
the East Pacific Rise, off México, 20°48’N, 109°16W,
1875m, P.Londsdale and L.Levin cols., Sept. 1985,
rc: 6.2mm, SIO-BIC C 10224. Paratypes, same
locality, 28 specimens and fragments, rc: 3.4-
lO.Omm, SIO-BIC C 10307, MNRJ 14338. Green
Seamount, AD1639, 1821m, 6 specimens, rc: 2.0
to 6.5mm, SIO-BIC C 10306; AD1639-7, 182lm, 2
specimens, rc: 1.9-5.0mm, SIO-BIC C 10305;
AD1639-9, 1871m, 5 specimens, rc: 1.5 to 5.7mm,
SIO-BIC C 10308.
Diagnosis - Shell box-like, articular ridges forming
digitiforms projections. Rostrum with one small
ridge directed to axial ridge of scutum. Scutum and
tergum with three articular ridges.
Description - Shell (Fig.26a, b) white, box-like,
opercular valves parallel to base of wall, with crenulated
growth lines projecting on all plates, more developed
on the rostro-carinal articular ridges, forming digitiform
projections; parietal plates of shell corrugated, forming
rough longitudinal ridges; basal margin not thickened.
Cuticle yellowish, smooth, persistent on shell
and opercular valves. Rostrum (Fig.26a, b) and
carina similar in size, former with about six well
developed articular ridges and deep grooves between
them on the rostro-carinal articulation, better
developed on the distai margin, and one smaller ridge
going to axial ridge of scutum; parietal plate with
large irregular longitudinal ridges; rostrum and
fixed scutum articulation with a small radius-like
projection, apex curved and slightly projected.
Carina (Fig.26a, b) with about five articular ridges
and deep grooves between them, directed toward
scutum, more developed distally, and one wide flat
ridge on distai margin, with small radius-like
projection toward fixed tergum; parietal plate with
irregular longitudinal ridges; apex projected. Fixed
tergum (Fig.26b) with the same size of fixed scutum,
Fig.26- Newmaniverruca digitiformis sp.nov.: (a) rostro-carinal view; (b) fixed tergum and fixed scutum view; (c) tergum and
scutum, internai view.
Arq. Mus. Nac., Rio de Janeiro, v.60, n.l, p.5-40, jan./mar.2002
34
P.S.YOUNG
both sides with well-developed alar-like projections;
apex projecting slightly backwards. Fixed scutum
(Fig.26b) with wide alar-like projection directed
toward rostrum, small radii-like projection to fixed
tergum; intemally, with only a deep scar for adductor
mu sele, without myophore.
Tergum (Fig.26a, c) quadrangular, with three articular
ridges; axial ridge prominent, but narrower than
intermediate ridge; intermediate ridge large, with the
growth lines crenulated, upper ridge thin and marginal
to occludent margin, with a large conspicuous
depression between upper and intermediate ridges;
carinal area smooth. Intemally, smooth; scutal margin
nearly straight except for notch on upper portion.
Scutum (Fig.26a, c) smaller than tergum; with three
articular ridges; axial ridge conspicuous, thin, both
sides well marked, intermediate ridge twice width of
axial ridge, upper ridge separated from intermediate
ridge by wide groove; rostral area smooth. Intemally,
smooth; tergal margin nearly straight, except for
protuberance at upper part.
Labrum (Fig.27a) with a series of teeth, each having
three to five acicular cusps, multifid spinules on the
inner surface and scales on the outer surface. Palp
(Fig.27a) acuminate, with few simple setae on inner
margin and distai region. Mandible (Fig.27b) with
three teeth, distance between first and second twice
distance between second and third, upper margin of
second and third teeth denticulated; lower angle
denticulated. Maxilla I (Fig.27c) with lower part
projecting; two large spines at upper angle, 4-6
small spines between upper large spines and lower
projected part, and 6-7 unpaired spines on basal
portion. Maxilla II (Fig.27d) triangular, anterior
margin with shallow concavity medially; covered by
long simple setae, except on the shallow concavity.
Cirrus I (Fig.28a) with unequal rami, anterior ramus 2/
3 length of posterior, covered with several long
simple setae. Cirrus II (Fig.28b) with anterior
ramus about 1/2 length of posterior, articles more
protuberant; both rami covered by numerous long,
simple setae, anterior ramus also with finely
Fig.27- Newmaniverruca digitiformis sp.nov.: (a) labrum and palp; (b) mandible; (c) maxilla I; (d) maxilla II.
Arq. Mus. Nac., Rio de Janeiro, v.60, n.l, p.5-40, jan./mar.2002
THE VERRUCIDAE FROM THE WESTERN COAST OF NORTH AMERICA...
35
bipectinate setae at distai article (Fig.28c). Rami of
cirri III-VI equal in length. Intermediate articles of
cirrus VI (Fig.28d) with width 1/2 length; two pairs
of setae on anterior margin, longer setae finely
pinnate; none to two fine simple setae on posterior
angle, multifid scales at distai margin. Caudal
appendage (Fig.28e, f) with 23 articles, three times
length of protopodite; long simple setae on distai
margins of articles. Penis twice length of protopodite,
clothed with thin setulae. Number of articles of cirri
I-VI and caudal appendage is presented in table 9.
Etymology - from the Latin digitus (finger) and forma
(shape).
TABLE 9
Number of articles on cirri I-VI, and caudal appendage of
Newmanivemica digitiformis sp.nov.
I
n
m
IV
V
VI
CA
RC
13/14
8/17
17/22
23/26
27/27
27/29
19
LC
12/16
8/17
17/21
23/26
26/28
27/29
23
(CI-VI) cirri I to VI; (CA) caudal appendage; (RC) right cirri;
(LC) left cirri.
Remarks - Most species of Newmaniverruca have a
smooth shell without radial ridges on the plates.
Only two species, N. multicostata (Gruvel, 1907) and
N. albatrossiana (Pilsbry, 1912), have the shell
ornamented with conspicuous radial ridges, but
Fig.28- Newmaniverruca digitiformis sp.nov.: (a) cirrus I; (b) cirrus II; (c) distai article of anterior ramus of cirrus II; (d) cirrus
VI and caudal appendage; (e) intermediate article of cirrus VI; (f) intermediate articles of caudal appendage; (CI-CVI) cirri I
to VI; (a.) anterior ramus; (c.a.) caudal appendage; (p.) posterior ramus.
Arq. Mus. Nac., Rio de Janeiro, v.60, n.l, p.5-40, jan./mar.2002
36
RS. YOUNG
none with the growth lines so pronounced and
forming digitiform projections. Furthermore, N.
multicostata differs from N. digitiformis sp.nov. by
having five articular ridges on the scutum, and the
tergum and rostrum do not have the upper ridge
turning toward the axial ridge of scutum. On the
other hand, N. albatrossiana has a scutum with
narrow articular ridges and its rostral area has
longitudinal ridges; the tergum has five articular
ridges and the carinal area has longitudinal ridges;
the rostrum has a veiy wide upper triangular portion,
which is conspicuously observed in apical view.
The specimens do not exhibit variation in the
number of ridges on the scutum and tergum despite
the large range of size available. Only one specimen,
from 182 lm, has its opercular valves in a more
angular position (45°) relative to the base of the wall.
In a recent revision of Verrucidae (YOUNG, 1998a),
Costatovermca was separated from Newmanivemica
and Metaverruca by the rostrum of the former having
radial ridges directed toward the scutum whereas the
latter two genera do not have these ridges. This new
species is intermediate between the latter two genera
and Costatovermca : it has a small apical ridge on the
rostrum, which is not articular, and it is directed
toward the axial ridge of the scutum. This appears to
be an intermediate stage before the development of
the series of upper ridges in Costatovermca. I have
maintained it in Newmanivermca because this ridge
is not well developed when compared with the series
of upper ridges directed toward the scutum in the
species of Costatovermca.
KEY TO THE GENERA OF THE FAMILY VERRUCIDAE
(modified from YOUNG, 1998a)
la. Shell erect; opercular plates perpendicular to base,
more than 45° to base; without adductor ridge or
myophore on fixed-scutum.2
lb. Shell not erect; opercular plates at no more than 45°
to base; with or without adductor ridge or myophore
on fixed-scutum.4
2a. Shell small and erect, wall plates delicate; rostrum and
carina suture linear to undulated, usually without but
rarely with weak articular ridges, opercular plates erect,
perpendicular to basis and almost parallel to fixed-scutum
and tergum. Altiverruca Pilsbry, 1916 s.s.
2b. Shell moderate to large and inclined, wall plates
massive; rostrum and carina very undulated, united
by strong articular ridges; operculum obliqúe to
basis inclined more than 45° and also obliqúe to
fixed-scutum and tergum.3
3a. Large size; rostrum and carina united by strong ridges,
the uppermost on rostrum being the largest
. Gibbosavemica gen.nov.
3b. Moderate size; rostrum and carina suturai area formed
by several imbricating ridges similar in size
. Cristallinaverruca gen.nov.
4a. Fixed tergum and rostrum medially expanded, forming
internai partitioned cavities; opercular plates 45° to
base; with adductor ridge on movable scutum
. Camera verruca Pilsbry, 1916
4b. Fixed tergum and rostrum not medially expanded,
without internai cavities; opercular plates less than
45° to base; without adductor ridge on movable
scutum.5
5a. Shell box-like; opercular plates proportionally large,
their width one half or more the width of shell; with
or without myophore.6
5a. Shell flattened; opercular plates proportionally small,
their width less than one half the width of shell; with
a large myophore. Verruca Schumacher, 1817
6a. Umbo of rostrum and carina marginal.7
6b. Umbo of rostrum not marginal, of carina marginal or
displaced from margin.8
7a. Rostrum without secondary ridges directed toward
tergal base; without myophore; basal margin of shell
not thickened. Newmaniverruca Young, 1998
7b. Rostrum with secondary ridges; with or without
myophore; basal margin of shell not thickened
. Costatovermca Young, 1998
7c. Rostrum without secondary ridges directed toward
tergal base; with myophore; basal margin of shell
thickened. Metavermca Pilsbry, 1916
8a. Umbo of rostrum displaced from margin.
. Rostrato verruca Broch, 1922
8b. Umbo of rostrum and carina displaced from margin
. Brochivermca Zevina, 1993
THE VERTICAL DISTRIBUTION
OF VERRUCID GENERA DISCUSSION
The Verrucidae has a wide bathymetric range from
the intertidal zone to 5700m. In figure 29, the depth
range of all the species of Verrucidae is presented.
Due to certain problems in the identification of
verrucid species I accepted all the names published,
but did not consider whether or not any of them
are synonyms. Therefore, the Verrucidae contains 117
recent species/subspecies distributed in 10 genera:
Vermca (6), Rostratovermca (11), Costatovermca (13),
Newmanivermca (10), Cameravemica (2), Brochivermca
(4), Cristallinavermca gen.nov. (10), Metavermca (22),
Gibbosavermca gen.nov. (13) and Altivermca (26).
The greatest number of species occurs between 300
and 1500m. Above 300m the number of species
decreases significantly, with the occurrences
of the balanomorphs at the shallower depth.
Arq. Mus. Nac., Rio de Janeiro, v.60, n.l, p.5-40, jan./mar.2002
THE VERRUCIDAE FROM THE WESTERN COAST OF NORTH AMERICA...
37
Otherwise, the pedunculates are abundant below
lOOOm, where they dominate. There is a substitution
from balanomorphs to verrucomorphs and from
verrucomorphs to pedunculates along a depth
gradient.
Organizing the depth distribution by genera
another pattern of substitution can be observed
(Fig.30). Most of the genera have different pattems of
distribution. Verruca s.s. predominates in shallow
water (0-100m), followed by Costatoverruca
(250-500m), Metaverruca (1000-2000m),
Gibbosaverruca (2000-3000m) and Altiverruca
(4000-5000m). The apparent dominance of
Cristallinaverruca at 5000-6000m is because there
is only one species recorded for this depth. The
genera considered more derived are found in
shallower waters ( Verruca, Rostratoverruca,
Brochiverruca and Costatoverruca) and that the
ones considered more plesiomorphic occur the
greatest depth ( Altiverruca ).
Number of species
Fig.29- Distribution of the species of the Verrucidae by depth range.
Arq. Mus. Nac., Rio de Janeiro, v.60, n.l, p.5-40, jan./mar.2002
38
P.S.YOUNG
100 %
80%
60%
40%
20 %
0 %
o
o
rH
I
O
O
<N
i
O
o
vo
O
o
o
O
o
o
o
o
o
o
CO
oj-
vo
VD
6
ò
ò
Ó
o
o
o
o
•8
<N
CO
■st
vo
H Verruca
H Newmaniverruca
S Cristallinaverruca
Wi Altiverruca
EU Rostratoverruca
El Cameraverruca
□ Metaverruca
SI Costatoverruca
0 Brochiverruca
□ Gibbosaverruca
Fig.30- Percentage of genera of Verrucidae by depth range.
ACKNOWLEDGEMENTS
I wish to acknowledge Wiiliam Newman (Scripps
Institution of Oceanography) and Robert Van Syoc
(Califórnia Academy of Sciences) for allowing me to
study these collecüons and also to Amold Ross (Scripps
Institution of Oceanography, San Diego), Guilherme
Muricy (Museu Nacional - Rio de Janeiro) and one
anonymous reviewer for their comments on the
manuscript, which significantly improved this paper.
This study was supported by the Conselho Nacional
de Desenvolvimento Científico e Tecnológico
(CNPq) and additional financial support from
Fundação Universitária José Bonifácio (FUJB) and
Fundação de Amparo à Pesquisa do Estado do
Rio de Janeiro (FAPERJ).
ABSTRACT
The genus Altiverruca is divided into three genera:
Altiverruca s.s., Gibbosaverruca gen.nov. and
Cristallinaverruca gen.nov. These are distinguished
mostly by the size of the shell, structure of the rostro-
carinal articulation, shape of the rostrum and the
development of articular ridges on the opercular
plates. A key for the genera of Verrucidae is provided.
The verrucid fauna of the Northeastem Pacific is
analyzed. Altiverruca beringiana, A. galapagosa and
six new species: Altiverruca sala sp.nov., A. vansyoci
sp.nov., Gibbosaverruca montereyi sp.nov., G.
robusta sp.nov., G. mateoi sp.nov. and
Newmaniverruca digitiformis sp.nov. occur within
this region and are described herein.
Arq. Mus. Nac., Rio de Janeiro, v.60, n.l, p.5-40, jan./mar.2002
THE VERRUCIDAE FROM THE WESTERN COAST OF NORTH AMERICA...
39
The depth distribution of the Verrucidae and the
predominant genera at selected intervals are
graphically portrayed. The Verrucidae most
commonly occur between 300 and 1500m and there
are pronounced changes in the distribution of
genera along a depth gradient.
Key words: Crustacea, Cirripedia, Pacific Ocean,
Verrucidae, new genera, taxonomy.
RESUMO
VERRUCIDAE DA COSTA OESTE DA AMÉRICA DO
NORTE, COM REVISÃO DO GÊNERO ALTIVERRUCA
O gênero Altiverruca é dividido em três gêneros:
Altiverruca s.s., Gibbosaverruca gen.nov. e
Cristallinavemica gen.nov. Estes são diferenciados
basicamente pelo tamanho da carapaça, estrutura da
sutura rostro-carenal, forma do rostro e desenvolvimento
das cristas articulares da placa opercular. Uma
chave para a identificação dos gêneros de
Verrucidae é apresentada.
A fauna de verrucídeos do nordeste do Pacífico é
analisada. Altiverruca beringiana, A. galapagosa e
seis espécies novas: Altiverruca sala sp.nov., A.
vansyoci sp.nov., Gibbosaverruca montereyi sp.nov.,
G. robusta sp.nov., G. mateoi sp.nov. e
Newmaniverruca digitiformis sp.nov. occorrem nesta
região e são descritas neste estudo.
A distribuição batimétrica de Verrucidae e dos gêneros
predominantes em intervalos selecionados é
apresentada graficamente. Verrucidae ocorre mais
comumente entre 300 e 1500m e foi observada uma
mudança acentuada na distribuição dos gêneros
ao longo de um gradiente de profundidade.
Palavras-chave: Crustacea, Cirripedia, Oceano
Pacífico, Verrucidae, gêneros novos, taxonomia.
LITERATURE CITED
BROCH, H., 1931 - Papers from Dr. Th. Mortensen’s Pacific
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Vidensk. Meddel. Dansk naturh. Foren., Copenhagen,
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BUCKERIDGE, J.S., 1994 - Cirripedia Thoracica:
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CORNWALL, I.E., 1955 - Canadian Pacific Fauna. 10.
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FOSTER, B.A., 1978 - The marine fauna of New Zealand:
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Inst. Mem., Wellington, 69:1-160.
FOSTER, B.A. & BUCKERIDGE, J.S., 1995a - Barnacles
(Cirripedia: Thoracica) of seas off the Straits of Gibraltar.
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FOSTER, B.A. & BUCKERIDGE, J.S., 1995b - Barnacles
(Cirripedia: Thoracica) of seas off Réunion Island and East
Indies. Buli. Mus. natn. Hist. nat., ser. 4, Paris,
16:345-382.
GRUVEL, A., 1900 - Sur les espèces nouvelles du genre
Verruca provenants du “Talisman”. Buli. Mus. natn. Hist.
nat., Paris, 6:242-244.
GRUVEL, A., 1902 - Cirrhipèdes. Bxpéditions Scientifiques
du “Travailleur” et du “Talisman”, pendant les années
1880, 1881, 1882, 1883. Paris: Massom. p.1-178,
pis.1-7.
HOEK, P.P.C., 1883 - Report on the Cirripedia collected by
H.M.S. Challenger during the years 1873-76. Rept Sei.
Res. Voyage H.M.S. Challenger, Zool., part 25,
London, 8:1-169, 13 pl.
NEWMAN, W. A., 1982 - A review of the extant taxa of the
“Group of Balanus concavus” (Cirripedia, Thoracica) and
a proposal for genus-group ranks. Crustaceana, Leiden,
43:25-36.
NEWMAN, W. A., 2000 - A new genus and species of bamacle
(Cirripedia, Verrucomorpha) associated with vents of the
Lau Back-Arc Basin: its gross morphology, inferred first
juvenile stage and affinities. Zoosystema, Paris,
22(l):71-84.
NEWMAN, W.A. & ABBOTT., D.P., 1980 - Cirripedia:
the barnacles. MORRIS, R.H.; ABBOTT, D.P. & HADERLIE,
E.C. (Eds.). Intertidal Invertebrate of Califórnia.
Stanford: Stanford University Press. p.502-535, pis.
147-154.
NEWMAN, W.A. & ROSS., A., 1971 - Antarctic Cirripedia.
Antarctic Res. Ser., Baltimore, 14:1-257.
NILSSON-CANTELL, C.A., 1928 - Studies on cirripeds in the
British Museum (Nat. Hist.). Ann. Mag. nat. Hist., Ser.
10, London, 2(7): 1-39.
NILSSON-CANTELL, C.A., 1938 - Cirripedes from the Indian
Ocean in the collection of the Indian Museum, Calcutta.
Mem. Ind. Mus., Calcuta, 13(1): 1-81, 3 pl.
NILSSON-CANTELL, C.A., 1955 - Cirripedia. Repts Swed.
Deep-Sea Bxped. 2, Zool., Stockholm (17):215-220.
PILSBRY, H.A., 1907 - The barnacles (Cirripedia) contained
in the collections of the U.S.National Museum. Buli. U. S.
natn. Mus., Washington, 60:1-122, 11 pl.
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RS. YOUNG
PILSBRY, H.A., 1916 - The sessile barnacles (Cirripedia)
contained in the collections of the U.S. National Museum;
including a Monograph of the American species. Buli.
U. S. natn. Mus., Washington, 93:1-366.
ROSELL, N.C., 1989 - Thoracica Cirripeds from the
Musorstom 2 expedition. Résultats des Campagnes
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144:9-35.
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Berg., Bergen, 65:3-19.
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69:1-268 [in Russian].
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20(l):31-92.
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BOUSQUETS, J.L.; SORIANO, E.G. 86PAPAVERO, N. (Eds.)
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Mediterranean. Trudy Inst. okeanol., Moscow, 100: 233-
258 [in Russian].
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Moscow, 55(8): 1149-1156 [in Russian].
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Thoracica) of the Pacific. 1. The North Pacific. Zool. Zh.,
Moscow, 66(12): 1812-1821 [in Russian].
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(Thoracica) of the Atlantic and Indian Ocean. Zool. Zh.,
Moscow, 66(9): 1304-1313 [in Russian].
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ZEVINA, G.B. 8c GALKIN, S.V., 1992 - Altivemica beringiana
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71:140-144 [in Russian].
ZEVINA, G.B. 86 GALKIN, S.V., 1993 -Altiverruca beringiana
sp.n. (Crustacea, Cirripedia): A first find of Verrucomorphan
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Arq. Mus. Nac., Rio de Janeiro, v.60, n.l, p.5-40, jan./mar.2002
THE GENUS AMPITHOE (CRUSTACEA, AMPHIPODA, AMPITHOIDAE)
FROM THE BRAZILIAN COAST «
(With 8 figures)
CRISTIANA S. SEREJO 1 (2)
MARCUS V. S. LICÍNIO
Museu Nacional
Universidade Federal do Rio de Janeiro
The Ampithoidae are phycophyllous amphipods
commonly found associated with several species of
macroalgae. The species inhabit open ended tubes
constructed among algae with a mucoid cement
secreted from dactyls of pereopods 3 and 4
(CONLAN, 1982). Ampithoids are found in the
shallow water of tropical, subtropical and temperate
zones of the world, and they are quite abundant
along the Brazilian coast. However, only the
cosmopolitan Ampithoe ramondi Audouin, 1826 was
previously recorded from our coast, with its
distribution restricted to Rio de Janeiro and São
Paulo States (SEREJO, 1998; WAKABARA &
SEREJO, 1998). Ampithoe encompasses a large
number of species (61) and is the largest genus
within the Ampithoidae (POORE & LOWRY, 1997).
DANA (1853) first described A. brasiliensis and A.
filicornis from Rio de Janeiro Harbor, both later
considered synonymous with Cymadusa filosa
Savigny, 1816 (BARNARD & KARAMAN, 1991).
Based on material from the Brazilian coast two
species of Ampithoe were herein redescribed: A.
diviswra Shoemaker, 1933 and A. ramondi and a
new species was described, A. seticoxae. Abbreviations
used in the text are the following: (MNRJ) Museu
Nacional/Universidade Federal do Rio de Janeiro.
Brazilian States: (BA) Bahia; (PE) Pernambuco; (RJ)
Rio de Janeiro; (SC) Santa Catarina. Figures: (Ant)
antenna; (Ep) epistome; (Gn) gnathopod; (Hd) head;
(LL) lower lip; (Md) mandible; (Mxl) maxilla; (P)
pereopod; (U) uropod; (UL) upper lip.
MATERIAL AND METHODS
The samples were obtained by SCUBA diving. In the
shallow infralittoral, different types of algae, sponges
and bryozoans were collected and kept in plastic
bags. On the surface, the substrate was dropped in
a large bucket, together with some amount of ethanol
to kill the amphipods. The substrate was then taken
out of the bucket and the debris with the animais
was sieved through a 0.5pm mesh. The sieved
material was then fixed in 70% ethanol and labeled.
Some specimens of each species were dissected, and
the bucal pieces mounted on permanent slides.
Illustrations were made using an Axiolab Zeiss
microscope with a camara lúcida.
SYSTEM ATICS
Family Ampithoidae Stebbing, 1899
Genus Ampithoe Leach, 1814
Diagnosis - Accessory flagellum absent. Outer lobe
of lower lip notched. Mandibular molar well
developed, palp 3-articulate. Gnathopod 2 larger
than gnathopod 1 in males, similar in size and shape
in females. Pereopods 5-7 simple or weekly
prehensile. Pereopod 7 similar to pereopod 6.
Uropod 1, peduncle with small rounded disto-ventral
spur or lacking the spur. Uropod 3, rami broad,
outer ramus with 2 large recurved spines. Telson
with small cusps, expanded into large hooks, or
absent (from POORE & LOWRY, 1997).
Ampithoe divisura Shoemaker, 1933
(Figs.1-3)
Ampithoe divisura Shoemaker, 1933:255, fig.8.
Material examined - Boa Viagem Beach, Guanabara
Bay, RJ, on algae and bryozoans, l-2m depth, 5cf
and 1$ , MNRJ 13158; Gravatás Beach,
Florianópolis, SC, on algae, 2-5m depth, 7d and
109 , MNRJ 13562; Pântano do Sul Beach,
Florianópolis, SC, on algae and sponges, 4-5m, 13d
and 159 , MNRJ 13563.
Diagnosis - Antenna 2 with flagellum shorter than
peduncular article 5. Male gnathopod 2 palm with
blunt tooth, palmar cleft V-shaped. Pereopods 3-4
robust, article 5 about 2/3 length of article 4.
Pereopods 5-7, propodus with 2 distai robust
striated spines. Uropod 1, peduncle with broadly
rounded inter-ramal process.
Description - d (5.3-7.0mm). Antenna 1 slightly
longer than antenna 2, flagellum with 17-20 articles
and aestethascs on each article. Antenna 2
setose, flagellum shorter than peduncular article 5;
1 Received on March 14, 2000. Accepted on September 21, 2002.
2 Museu Nacional/UFRJ, Departamento de Invertebrados. Quinta da Boa Vista, São Cristóvão, 20940-040, Rio de Janeiro, RJ, Brazil. csserejo@acd.ufrj.br.
42
C.S.SEREJO & M.V.S.LICINIO
flagellum with 9-11 articles. Both antennae about 2/5
of body length. Right mandible with 3 spines on setal
row; mandibular palp article 2 subequal to article 3,
article 3 with about 8 distai and sub-distal bipectinate
setae. Lobules of lower lip slender and separated.
Maxilla 1, inner lobe with two long bipectinate setae,
outer lobe with 10 dentate spines, palp with 4 to 6
apical spines and 3 facial setae. Maxilla 2, inner lobe
with sequence of 11-12 submarginal plumose setae;
outer lobe with several long distai setae and marginal
and distai setules. Maxilliped with 11 teeth on inner
margin of outer lobe; outer angle of outer lobe and first
article of palp with one well developed seta each.
Coxa 1 projected forward. Gnathopod 1, basislobate
antero-distally; carpus unproduced at the
posterodistal angle; propodus rectangular, with one
large spine on defining angle of the palm; dactylus
finely serrated on inner side. Gnathopod 2, basis
lobate antero-distally; propodus anterior margin
setose and produced anteriorly in larger specimens
(7.0mm); palm with sparse setae and defined by a
blunt tooth, palmar cleft V-shaped; dactylus finely
serrated on inner side. Pereopods 3-4 robust, article
5 about 2/3 of article 4. Propodus of pereopods 5-7
with four spines, two proximal smooth and two
distai robust striated spines.
Fig. 1- Ampithoe divisura Shoemaker, 1933, cf , 7.2mm, Guanabara Bay, RJ, MNRJ 13158. (Ant) antenna; (Ep) epistome;
(LL) lower lip; (Md) mandible; (Mxl) maxilla; (UL) upper lip.
Arq. Mus. Nac., Rio de Janeiro, v.60, n.l, p.41-50, jan./mar.2002
THE GENUS AMPITHOE (CRUSTACEA, AMPHIPODA, AMPITHOIDAE) FROM THE BRAZILIAN COAST
43
Uropod 1, peduncle with broadly rounded disto-ventral
inter-ramal process; outer ramus with 5-7 marginal
spines; inner ramus with 2 marginal spines. Uropod
2, peduncle with vestigial triangular process; outer
ramus with 3-4 marginal spines; inner ramus with
respectively 3 and 2 marginal spines on the outer and
inner side. Uropod 3, peduncle distai margin with 5
spines. Telson triangular and setose, with rounded tip.
9 (ovigerous, 5.9mm). Antennae slightly shorter
than in males, about 1/3 of body length. Antenna
2, flagellum subequal to peduncular article 5 and
not shorter as in males. Gnathopod 2, palm sinuous,
lacking digitiform process. Uropod 1, peduncular
disto-ventral process inconspicuous.
Remarks - The Brazilian specimens agree in most
aspects to the material from Florida described by
SHOEMAKER (1933). Differences were noted in the
palp of maxilla 1, that presented 4 spines on the
Brazilian specimens, instead of 5 spines; and articles
2-3 of the mandibular palp are subequal in length
and not the second article the longest, as described
in the Florida material (SHOEMAKER, 1933).
Ampithoe divisura is part of the A. ramondi complex,
which is characterized by a large digitiform process on
the male gnathopod 2 palm and was even synonymized
with A. ramondi by BARNARD & KARAMAN (1991).
However, A. divisura can be distinguished from A.
ramondi by some distinct characters such as: the
flagellum of the antenna 2 is shorter than the
peduncular article 5; the male gnathopod 1 carpus is
unproduced at the posterodistal angle; the palm of male
gnathopod 2 has a blunt digitifom process, with palmar
cleft V-shaped; the propodus of pereopods 5-7 has 2
robust distai striate spines; and the uropod 1 has a
broadly rounded inter-ramal process.
Fig.2- Ampithoe divisura Shoemaker, 1933, <3, 7.2mm, Guanabara Bay, RJ, MNRJ 13158. (Gn) gnathopod; (P) pereopod.
Arq. Mus. Nac., Rio de Janeiro, v.60, n.l, p.41-50, jan./mar.2002
44
C.S.SEREJO & M.V.S.LICINIO
Ampithoe divisura is very close to A. kava Myers
(1985), described for the Pacific, in all the
characters listed above, but some differences were
noted. The flagellum of antennae 2 has 9-11
articles, but 6 articles in A. kava and the uropod 1
outer ramus have 5-7 marginal spines, but 4
spines in A. kava (MYERS, 1985). On the other
hand, the specimens from Hawaii described by
BARNARD (1970) as A. ramondi and synonymized
with A. kava by POORE & LOWRY (1997) have the
flagellum of antennae 2 figured with 9 articles in a
6.8mm male. Furthermore, the anterodistal margin
of the propodus male gnathopod 2 is anteriorly
projected as in Brazilian specimens. POORE &
LOWRY (1997), on a detailed redescription of A.
kava, cited the maxilla 1 inner lobe lacking setae
and the palp of outer plate with 8 apico-medial
spines. The Brazilian specimens of A. divisura have
the maxilla 1 with 2 long seta on the inner lobe
and 4 apico-medial spines on the palp of outer lobe.
This was the first record of A. divisura for the
Southwestern Atlantic.
Distribution - Bird Key Reef, Tortugas, Florida, USA
(SHOEMAKER, 1933) (type locality). Brazil: RJ and SC.
Fig.3- Ampithoe divisura Shoemaker, 1933, cf, 7.2mm and? , 5.0mm, Guanabara Bay, RJ, MNRJ 13158. (Gn) gnathopod;
(T) telson; (U) uropod.
Arq. Mus. Nac., Rio de Janeiro, v.60, n.l, p.41-50, jan./mar.2002
THE GENUS AMPITHOE (CRUSTACEA, AMPHIPODA, AMPITHOIDAE) FROM THE BRAZILIAN COAST
45
Ampithoe ramondi Audouin, 1826
(Figs.4-5)
Ampithoe ramondi Audouin, 1826:93; KRAPP-
SCHICKEL, 1978:1, figs. 1-2; 1982:98, figs.66-67;
MYERS, 1985:27, fig.17; RABINDRANATH, 1972:
162, figs. 1-2; SEREJO, 1998:364, fig.lA-I.
Material examined - Santo Aleixo Island, Serinhaém,
PE, 2cf and 59 , MNRJ 10024; Porto da Pedra Beach,
BA, on sponges, 2Cf and 29 , MNRJ 13162; Arraial
do Cabo, RJ, on sponges, 12d" and 309 , MNRJ 9998;
Pântano do Sul Beach, Florianópolis, SC, 4-5m, on
algae and sponges, l9 , MNRJ 13565.
Diagnosis - Antennae 1-2 subequal. Male gnathopod
2 palm with acute tooth, palmar cleft U-shaped.
Pereopods 3-4 robust, article 4-5 subequal in length.
Distai spines on propod of pereopods 5-7 acute and
smooth. Uropod 1 lacking inter-ramal process.
Description - Ó (4.8mm). Antenna 1 larger than
antenna 2, flagellum with about 29 articles, antenna
2, peduncle with articles 4-5 subequal in length,
flagellum with about 17 articles. Lower lip with lobules
slender and separated. Maxilla 1, inner lobe with 4
long bipectinate setae, outer lobe with 10 dentate
spines, palp with 6 apical spines and 4 subdistal setae.
Coxa 1 projected forward. Gnathopod 1, basis
lobated anteriorly; carpus with triangular process
at the posterodistal angle; propod rectangular, with
a large spine defining the palm; dactylus serrated
internally. Gnathopod 2, basis lobated anteriorly;
anterior margin of propod setose and slightly
projected distally; palm defined by an acute
digitiform process, palmar cleft U-shaped, dactylus
serrated internally. Pereopods 3-4 robust, article
4-5 subequal in length. Propod of pereopods 5-7
with acute and smooth spines.
Fig.4- Ampithoe ramondi Audouin, 1826, d", 4.8mm, Santo Aleixo Island, Serrinhaém, PE, MNRJ 10024. cf, 4.5mm, Porto
da Barra Beach, BA, MNRJ 13162. <3 ,4,5mm, Arraial do Cabo, RJ, MNRJ 9998. (Hd) head; (Gn) gnathopod.
Arq. Mus. Nac., Rio de Janeiro, v.60, n.l, p.41-50, jan./mar.2002
46
C.S. SEREJO & M.V.S.LICINIO
Uropod 1, peduncle lacking inter-ramal process,
outer ramus with 2 and 6 spines, on inner and outer
margins respectively; inner ramus with 3 spines.
Uropod 2, outer ramus with 1 and 3 spines, on inner
and outer margins respectively. Uropod 3 with 5
apical spines. Telson trapezoid.
9 (6.4mm). Gnathopod 2, palm obliqúe, with a
defining spine but lacking digitiform process.
Remarks - Ampithoe ramondi is a quite common
species found on macroalgae and sponges from the
Brazilian coast. This species was observed also as one
of the largest populations living on the sponge Dysidea
robusta Vilanova & Muricy, 2001 at Prainha, Arraial
do Cabo, Rio de Janeiro (SEREJO, 1995; 1998 as D.
Jragilis Johnston). The identity of A. ramondi is still
confusing, mainly because the original description was
based on a female only. The description of the male
gnathopod 2, which is quite variable in this genera, is
important for the identification of Ampithoe species
(MYERS, 1985). The great variability described for
A. ramondi suggests that we are dealing with a
complex of species, which should be revised as
pointed before (SEREJO, 1998).
Distribution - Egypt (type locality). Cosmopolitan
in tropical and warm temperate waters. Brazil: PE,
BA, RJ and SC.
Fig.5- Ampithoe ramondi Audouin, 1826, cf, 4.8mm and 9,4.6mm, Santo Aleixo Island, PE, MNRJ 10024. (Gn)
gnathopod; (P) pereopod; (T) telson; (U) uropod.
Arq. Mus. Nac., Rio de Janeiro, v.60, n.l, p.41-50, jan./mar.2002
THE GENUS AMPITHOE (CRUSTACEA, AMPHIPODA, AMPITHOIDAE) FROM THE BRAZILIAN COAST
47
Ampithoe seticoxae sp.nov.
(Figs.6-8)
Material examined - Holotype, d , 9.0mm, Boa Viagem
Beach, Guanabara Bay, Niterói, Rio de Janeiro, on
algae and biyozoans, l-3m, P.S.Young and C.S.Serejo
cols., IV/1997, MNRJ 13457. Paratypes: same locality,
on algae, 3d , 3 juvenile d , 29 , MNRJ 15127; same
locality, on algae and sponges, ld and 29 , MNRJ
13410; Morcego Beach, Guanabara Bay, RJ, on algae,
l-3m, 1 juvenile d and 29 MNRJ 13411.
Diagnosis - Antennae 1 about 1/3 of body length.
Articles 4-5 and flagellum of antennae 2 densely
clothed with plumose setae. Coxae 1-4 with group
of 2-4 setae on posteroventral margin. Gnathopod
2, propodus rectangular, palm straight, slightly
obliqúe. Propodus of pereopods 5-7 with a row of 5
spines and a disjunct smaller distai spine.
Etymology - The name of the species refers to the
presence of setae on the postero-ventral margin of
coxae 1-4. From the Latin seta and coxa.
Description - d (8.0-9.3mm). Body pitted with
brown spots. Antenna 1 slightly shorter than
antennae 2, reaching about 1/3 of body length,
flagellum with 20 articles. Articles 4-5 and flagellum
of antennae 2 densely clothed with plumose setae.
Mandibular palp articles in rate 2:5:5, article 3 with
2 rows of plumose setae with distinct length. Lobules
of lower lip slender and separated. Maxilla 1, inner
lobe with one tiny seta; outer lobe with 10 spines,
two of them more slender and apically bifurcated;
palp with 4 apical spines and 3 facial setae. Maxilla
2, outer lobe larger than inner lobe; inner lobe with
8 plumose submarginal setae; outer lobe with long
distai setae and hair like setae on externai and distai
margins. Maxilliped with 13 teeth on inner margin
of outer lobe; outer angle of outer lobe lacking setae;
first article of palp with 3-4 long setae.
Mxl 1
Fig.6- Ampithoe seticoxae sp.nov., holotype, Ò , 9.0mm, Guanabara Bay, RJ, MNRJ 13457. (Ant) antenna; (LL) lower
lip; (Md) mandible; (Mxl) maxilla.
Arq. Mus. Nac., Rio de Janeiro, v.60, n.l, p.41-50, jan./mar.2002
48
C.S.SEREJO & M.V.S.LICINIO
Coxa 1 produced anteriorly. Coxae 1-4 with group
of 2-4 setae on posteroventral margin. Gnathopod
1, basis posterior margin with groups of long setae,
anterodistal margin lobate; ischium lobate;
propodus with large spine on posterior margin;
dactylus overlapping palm and finely serrate on
inner side. Gnathopod 2, basis and ischium lobate
antero-distally; propodus rectangular, palm
obliqúe and straight in adult forms, dactylus fitting
palm and serrated on inner side. In some juvenile
males (8.5mm), propodus more oval, with palm
slightly concave. Pereopods 3-4 similar and setose,
carpus slightly shorter than propodus. Pereopod
5, basis broad, not elongated as in pereopods 6-7.
Basis of pereopod 6-7 with a small concavity on
posterodistal angle. Pereopods 5-7, propodus with
row of 5 spines increasing in length distally and
one small distai spine displaced from the row.
Epimera 1-3 with lateral ridge. Uropod 1, peduncle
with row of facial setae. Uropod 2, peduncle with group
of facial setae, outer and inner rami with 2-3 and 2
spines respectively. Uropod 3, peduncle with group of
facial setae, distai margin with a row of 4-5 spines.
Telson trapezoidal, with 2 distai cusps, 3 marginal
setae on each side and 2 long subdistal setae.
9 (7. lmm, ovigerous) . Gnathopod 2, propodus oval,
with a large spine on posterior margin; palm obliqúe;
dactylus serrate on inner margin.
Remarks - Ampithoe seticoxae sp.nov. has some
aspects in common with the group A. lacertosa Bate,
1858, A. valida Smith, 1873 and A. plumulosa
Shoemaker, 1938, discussed by CONLAN &
BOUSFIELD (1982), as follows: coxae 1-2 in males
shallower than coxae 3-5; gnathopod 2, articles 2-
3 with prominent antero-distal lobe, propodus
Fig.7- Ampithoe seticoxae sp.nov., holotype, cf , 9.0mm, Guanabara Bay, RJ, MNRJ 13457. (Gn) gnathopod; (P) pereopod.
Arq. Mus. Nac., Rio de Janeiro, v.60, n.l, p.41-50, jan./mar.2002
THE GENUS AMPITHOE (CRUSTACEA, AMPHIPODA, AMPITHOIDAE) FROM THE BRAZILIAN COAST
49
rectangular with palm nearly transverse and epimera
1-3 with lateral ridge. Within this group, Ampithoe
seticoxae sp.nov. is closer to A. plumulosa in bearing
the antenna 2 clothed ventrally with dense plumose
setae. Despite these similarities, Ampithoe seticoxae
sp.nov. can be distinguished from A. plumulosa by
the antenna 1 shorter than antenna 2, reaching 1/3
of the body length, in A. plumulosa the antenna 1 is
slightly longer than antenna 2; the plumose setae of
antenna 2 appear on article 4 of the peduncle instead
of on article 5; the inner lobe of maxilla 1 has one
seta and not 4 setae; the posterior lobe on article 5
of gnathopod 1 is not distally produced under the
segment 6; coxae 1-4 have a group of setae on
postero-distal margin, which was not described for
A. plumulosa; and by the absence of a ventral oval
keel on the seventh thoracic segment, observed on
A. plumulosa by SHOEMAKER (1938).
Distribution - Guanabara Bay, RJ.
KEY FOR THE AMPITHOE SPECIES
OCCURRING ON THE BRAZILIAN COAST
la. Antenna 2 densely clothed with plumose setae; palm
of male gnathopod 2 lacking digitiform process; coxae
1-4 with group of 2-4 setae on postero-ventral margin
. A. seticoxae sp.nov.
lb. Antenna 2 clothed with simple setae; palm of male
gnathopod 2 with digitiform process; coxae 1-4 lacking
group of 2-4 setae on postero-ventral margin.2
2a. Flagellum of antenna 2 shorter than peduncular
article 5; male gnathopod 2 palm with blunt
digitiform process, palmar cleft V-shaped; uropod 1
with broadly rounded inter-ramal process
. A. divisura Shoemaker
2b. Flagellum of antenna 2 much longer than peduncular
article 5; male gnathopod 2 palm with acute digitiform
process, palmar cleft U-shaped; uropod 1 without
inter-ramal process. A. ramondi Audouin
Fig.8- Ampithoe seticoxae sp.nov., holotype, 3 , 9.0mm, Guanabara Bay, RJ, MNRJ 13457. Paratype, 9,7,lmm,
Guanabara Bay, RJ, MNRJ 15127. (Gn) gnathopod; (T) telson; (U) uropod.
Arq. Mus. Nac., Rio de Janeiro, v.60, n.l, p.41-50, jan./mar.2002
50
C.S. SEREJO & M.V.S.LICINIO
ACKNOWLEDGMENTS
We would like to thank Dr. Paulo S. Young (MNRJ)
for valuable comments on the manuscript and for
editing the plates, and to FUJB (Fundação
Universitária José Bonifácio) for financial support.
ABSTRACT
This study comprises the redescription of two species
of Ampithoe : the cosmopolitan A. ramondi
Audouin, 1826, previously recorded from Brazil and
A. diuisura Shoemaker, 1933, a new record from the
Southwestern Atlantic. Ampithoe seticoxae sp.nov.
is also described based on material from Guanabara
Bay, Rio de Janeiro. A key for the species of Ampithoe
from the Brazilian coast is provided.
Key words: Ampithoe, new species, Brazil.
RESUMO
GÊNERO AMPITHOE (CRUSTACEA, AMPHIPODA,
AMPITHOIDAE) NA COSTA BRASILEIRA
Este estudo compreende a redescrição de duas
espécies de Ampithoe : A. ramondi Audouin, 1826,
espécie cosmopolita previamente registrada para o
Brasil e A. diiÁsura Shoemaker, 1933, novo registro
para o Atlântico Sul Ocidental. Ampithoe seticoxae
sp.nov. é descrita com material proveniente da Baía
da Guanabara, Rio de Janeiro. Uma chave das
espécies de Ampithoe que ocorrem na costa brasileira é
fornecida.
Palavras-chave: Ampithoe, espécie nova, Brasil.
LITERATURE CITED
AUDOUIN, V., 1826 - Animaux articulés. Crustacés. In:
SAVIGNY, J.C. (Ed.) Description de PEgypte, Publiée par
les Ordres de as Majesté TÉmpereur Napoléon-le-Grand,
Histoire Naturelle. v.l, n.4, 93p., pis.II.
BARNARD, J.L., 1970 - Sublittoral Gammaridea (Amphipoda)
of the Hawaiian Islands. Smithson. Contr. ZooL.
Washington, 34:1-286.
BARNARD, J.L. & G.S.KARAMAN, 1991 - The families and
genera of marine gammaridean Amphipoda (except
marine gammaroids), Parts 1 and 2, Rec. Aust. Mus.,
Sydney, Suppl. 13:1-866.
CONLAN, K.E., 1982 - Revision of the gammaridean amphipod
family Ampthoidae using numerical analytical methods.
Can. J. Zool., Ottawa, 60:2015-2027.
CONLAN, K.E. & BOUSFIELD, E., 1982 - The amphipod
superfamily Corophioidea in the northeastem Pacific region.
Family Ampithoidae: systematics and distributional ecology.
Publ. Oceanogr. Biol. Ottawa, 10:41-75.
DANA, J.D., 1853 - Crustacea. Part II. U.S. Explor. Exped..
Philadelphia, 14:689-1618, atlas of 96 pis.
KRAPP-SCHICKEL, G., 1978 - Die gattung Ampithoe
(Crustacea, Amphipoda) im Mittlemeer. Bijdr. Dierk.,
Leiden, 48(1): 1-15.
KRAPP-SCHICKEL, G., 1982 - The Amphipoda of the
Mediterranean, Gammaridea (Acanthonotozomatidae to
Gammaridae). Family Ampithoidae. Mem. Inst.
océanogr., Monaco, 13(1):94-110.
LYONS, J. & MYERS, A.A., 1990 - Amphipoda Gammaridea
from coral rubble in the gulf of Aqaba, Red Sea: Families
Acanthonotozomatidae, Ampeliscidae, Ampithoidae,
Anamixidae, Aoridae and Colomastigidae. J. Nat. Hist.,
London, 24:1197-1225.
MYERS, A.A., 1985 - Shallow-water, coral reef and mangrove
Amphipoda (Gammaridea) of Fiji. Rec. Aust. Mus.,
Sydney, Suppl. 5: 1-144.
POORE, A.G.B. & LOWRY, J.K., 1997 - New ampithoid
amphipods from Port Jackson, New South Wales,
Australia (Crustacea: Amphipoda: Ampithoidae). Invert.
Taxon., Sydney, 11:897-941.
RABINDRANATH, P., 1972 - Marine Gammaridea (Crustacea:
Amphipoda) from the Indian region. Family Ampithoidae.
Mar. Biol., Berlin, 14:161-178.
SEREJO, C.S., 1995 - Fauna de Amphipoda (Crustacea)
associada à esponja Dysidea sp. em Arraial do Cabo,
RJ - Taxonomia e composição da comunidade. Rio de
Janeiro, xvi, 96p. Master Thesis, Museu Nacional/UFRJ.
SEREJO, C.S., 1998 - Gammaridean and Caprellidean fauna
(Crustacea) associated to the sponge Dysidea fragilis
Johnston at Arraial do Cabo, RJ, Brazil. Buli. Mar. Sei.,
Miami, 63(2):363-385.
SHOEMAKER, C.R., 1933 - Two new genera and six new
species of Amphipoda from Tortugas. Pap. Tortugas Lab.,
Florida, 28:247-256.
SHOEMAKER, C.R., 1938 - Three new species of the genus
Ampithoe from the West coast of America. J. Wash. Acad.
Sei., Washington, 28(1): 15-25.
WAKABARA, Y. & SEREJO, C.S., 1998 - Malacostraca -
Peracarida. Amphipoda. Gammaridea and Caprellidea.
In: YOUNG, P.S. (Ed.) Catalogue of Crustacean of Brazil.
Rio de Janeiro: Museu Nacional. (Série Livros 6). p.561-514.
Arq. Mus. Nac., Rio de Janeiro, v.60, n.l, p.41-50, jan./mar.2002
MUSEU NACIONAL
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Programa PROAP/2002
IMPRESSÃO
Divisão Gráfica/PU
UFRJ
Arquivos do Museu Nacional, v.6o, n.i
SUMÁRIO
YOUNC, P.S. The Verrucidae (Crustacea, Cirripedia) from the western coast of North America, with a revision on
the genus Altiverruca . 5
SEREJO, C.S. & LICÍNIO, M.V. The genus Ampithoe (Crustacea, Amphipoda, Ampithoidae) from the Brazilian
coast.41
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