HGLC: 11.5 by 12.2 by 12.0; MNHN (Sta.
DW11): 15.7 by 17.7 by 9.2 (valve); MNHN (Sta.
CP889): 19.7 by 17.0 by 9.1 (valve).
South and Central Indo-
Pacific in 146–805 m depth.
Paratypes of S. ericia: AUSTRA-
LIA; South Cape Wiles, 174–183 m, 35°39
Ј S, 136°40Ј E,
AMS 032068, 1 left, 1 right valves (Zoological Results of
the F.I.S. E
, 28 Aug. 1909).
Holotype of S. japonica:
JAPAN. ANSP 49639, 1 shell. MNHN. SW PACIFIC.
Loyaute Islands, 16 lots [122 v]. TONGA IS. 12 lots [59
v]. GUAM. Marianas Islands, 3 lots [15 v]. AUSTRALIA.
South Cape Wiles, 1 lot [6 v]. NEW CALEDONIA.
South, 3 lots [7 specimens]; Banc Esponge, 2 lots [3
specimens]; Chesterfield Plateau, 1 specimen. PHILIP-
PINES. Aliguri Is. 2 lots [1 specimen and 3 v]; Bohol
Sea, Off Balicasag Island, 1 lot [1 v]. FIJI. 1 specimen.
MYANMAR (BURMA). 1 lot [5 v] Preparis North Chan-
nel, 1 lot [4 v]; N.W. of Tavoy I., 1 lot [11 v]. ANDA-
MANS SEA. 1 lot [1 v]. THAILAND. Phuket I., 1 lot [11
v]; Andaman Sea, 1 lot [1 v] (Details in Simone and
Cunha, in press.)
Spinosipella costeminens (Poutiers, 1981)
(Figures 60–65, 68–71, 83–92, 103–108)
4, figs 1–4, text fig 5).
143, 158 (figs. 1–2).
Shell with 16–17 tall radial ribs, those
more posterior to middle surface very taller, normally
possessing blade-like projections along tip; 3–4 more
posterior abruptly lower, preceded by a very tall, carina-
gree of convexity (width/length) in each valve approxi-
mately 0.50. Outer surface prickly, with somewhat cha-
otic organization (Figures 60, 61, 68–70). Sculptured by
strong, uniform, arched, radial ribs, from 16 to 17 in each
valve (Figures 60, 61); ribs increasing from region ante-
rior to umbo to region between middle and posterior
thirds, last ribs in this region taller and more separated
from each other, last one on a weak carina (Figure 70);
larger ribs normally possessing blade-like, projection
THE NAUTILUS, Vol. 122, No. 2
lar to those of anterior region; blade like projection ab-
sent in some specimens (Figures 68–71). Posterior edge
about twice broader than anterior edge. Between umbos
and anterior edge a concavity bearing transversal ribs
similar to ribs of remaining region (Figures 64, 65); pre-
umbonal region narrow, smooth about 10% of shell
length (Figures 64, 65, 69). Anterior, ventral, and poste-
rior edges forming zigzag (Figures 62, 63, 71, 103), with
tips longer and narrower projected in those middle and
larger ribs. Hinge with a large cardinal tooth in right
valve, stubby, tall (about 10% of valve width), broadly
extracted from shell, posterior view, showing siphonal area. 85. Whole right view, some portions of right mantle lobe extracted,
particularly regions ventral to septum, and ventral and dorsal to renal fold (rf) to expose inner surface; cerebral ganglion (ce) seen
by transparency. 86. Same, ventral-slightly right view. Scale bar = 5 mm.
L. R. L. Simone and C. M. Cunha, 2008
dent socket in left valve shallow, restrict to dorsal sur-
face; this socket flanked by small posterior tooth, with
insertion of anterior valve edge approximately in middle
region of this tooth (Figure 62), anterior tooth absent
Additional details for this species see Poutiers (1981),
Poutiers and Bernard (1995).
to those in preceding species, differing in being propor-
tionally shorter and wider (Figures 104–105). Length
of hinge length, and about 1.5 times wider
similar to those in preceding species. Anterior adductor
muscle about 20% dorso-ventrally longer (Figures 83, 85).
and disposition similar to those in S. deshayesiana. Byssal
gland relatively deep, running immersed in ventral re-
gion of pedal musculature at about half of byssal furrow
topology of some structures also shown. Scale bar = 5 mm. 88. Scheme of layers of tissue in indicated region of stomach. Scale bar
= 0.5 mm. 89. Fore- and midgut opened longitudinally for exposing inner surface (same scale of Figure 87). 90. Foot, ventral-slightly
posterior view, sectioned transversally in two levels to show inner layer of tissues. Scale bar = 1 mm.
at same level as pericardium (MNHN sta. CP767, Mallory, 5
m). Scale bar = 2 mm. 92. Same, detail of posterior region of mantle
border. Scale bar = 1 mm. 93–102. S. deshayesiana. 93. Detail of hinge region of left valve with lithodesma (lt) still attached, right
view. Scale bar = 2 mm. 94. Detail of posterior region of supraseptal chamber, right view, right mantle lobe removed (MNHN sta.
DW1499). Scale bar = 1mm. 95. Infraseptal chamber roof, ventral view, right mantle lobe removed (MNHN sta. CP767). Scale bar
= 2 mm. 96–99. Lithodesma (MNHN sta. DW739). Scale bar = 1 mm. 96. Ventral view. 97. Dorsal view. 98. Posterior-slightly dorsal
view. 99. Posterior view. 100. Same specimen, empty shell, ventral view, valves slightly open, lithodesma still in situ. Scale bar = 2
mm. 101. Same, detail of hinge and lithodesma. 102. Same, ventral-slightly anterior view. 103–108. S. costeminens. 103. Shell,
ventral view, valves open, lithodesma still attached to left valve (MNHN sta. CP1460). Scale bar = 2 mm. 104–105. Lithodesma, same
lot (other specimen), dorsal and ventral views respectively. Scale bar = 1 mm. 106. Detail of anterior region, right view, integument
removed, mainly showing right cerebral ganglion (ce) (same lot). Scale bar = 1 mm. 107. Infraseptal chamber roof, ventral view, right
mantle lobe removed (MNHN CP767). Scale bar = 1 mm. 108. Detail of posterior (siphonal) region, posterior view (MNHN
CP1460). Scale bar = 2 mm.
layer surrounding a nucleus of conective tissue (Figure
to those in preceding species, with following distinctive
characters. Pair of secondary tentacles positioned be-
tween incurrent and excurrent siphons (Figures 84, 108);
remaining tentacles similar in size and position. Ventral
pair of tentacles of incurrent siphon generally symmetri-
cal. Zigzag formed by mantle edge having secondary
folds positioned in more distal tips, possibly elated to
taller radial shell ribs (Figure 108). Radial mantle gland
(Figure 92) similar to S. deshayesiana.
similar to those in preceding species, except for wider
platform between posterior region of gills as part of sep-
tum (Figure 107).
to those in preceding species, differing mainly by wider
region separating pair of renal folds in supraseptal cham-
ber (Figure 85).
91): Pericardium and heart with characters similar to
those in S. deshayesiana (Figure 91). Kidneys of similar
features, differing mainly by enlargement of pair of renal
folds (Figures 85–86, rf), taller and wider, almost divid-
ing supraseptal chamber in two—internal and external—
halves. Height of renal fold about 80% of that of su-
praseptal chamber height. In addition to an enlargement,
both renal folds still have posterior end in more anterior
position and wider separation between folds and visceral
mass (Figure 85).
lar to those in preceding species. Esophagus with about
dicular to posterior surface of anterior adductor muscle
(Figure 87, es). Stomach main chamber with longer re-
gion as a blind-sac projected posteriorly. Gastric wall
constituted by external layer of weak connective tissue
(Figure 88, cj), two thick muscular layers of similar size,
with outer layer of longitudinal muscle and inner layer of
circular muscle (Figure 88, lo and cm). Inner surface of
stomach (Figure 89) with posterior end of esophageal
folds clearly more evident that together form a flat fold.
Another ventral fold surrounding apertures to digestive
diverticula. Gastric style narrower (about
width); internally a pair of tall folds separating intestinal
from style sac components (Figure 88, ss, in).
ceding species. Separated masculine and feminine com-
ponents of gonad shown through histological sections in
Figures 91(ts, ov).
ganglia with similar localization and size to those of pre-
Measurements (respectively length, height, width
MNHN (Sta. 1361): 22.0 by 28.1 by 12.5
(valve); MNHN (Sta. CC996): 20.0 by 24.3 by 14.3
(valve); MNHN (Sta. CP992): 19.6 by 23.3 by 12.6
Tropical West Pacific.
Holotype; Additional material
(MNHN): SW PACIFIC. 4 lots [32 v, 11 specimens];
Wallis Is., 6 lots [15 v]; Banc Combe, 5 Lots [28 v];
Fortuna Is., 5 lots [18 v]; Banc Waterwitch, 2 lots [3 v];
Banc Tuscarora, 29 lots [63 v]; South Vanuatu - Monts
Gemini, 4 lots [4 v, 1 specimen]; TONGA IS. 8 lots [52
v]; Eua Is. 6 lots [12 v]; Seamount, 6 lots [29 v]; South of
Nomuka group, 1 lot [25 v]; Ha
Јapai Group, 2 lots [4 v];
N Ha’apai group, 3 lots [ 6 v]; NW Tongatapu, 3 lots [16
v]; SW Tongatapu, 5 lots [22 v]; Tongatapu, 6 lots [8 v];
S. Nomuka group, 2 lots [6 v]; Vava’ group, 1 lot [2 v];
NEW CALEDONIA. 5 lots [5 v, 5 specimens]; Lord
Howe, 1 lot [1 v]; Banc Nova, 2 lot [8 v, 1 specimen];
North New Caledonia, 10 lots [tota 20 v]; South New
Geographic distribution of Spinosipella spp.
[98 v]; Banc Esponge, 11 lots [144 v]; Banc Kaimon-
Maru, 9 lots [38 v]; Banc Antigonia, 1 lot [1 v]; Banc
Jumeau-West, 4 lots [17 v]; Banc Introuvable, 7 lots [16
v]; Banc Stylaster, 1 lot [1 v] ; Volcans Hunter and Mat-
thew, 2 lots [2 v]; S.E. New Caledonia, 2 lots [2 v]; East
New Caledonia, 6 lots [30 v] Banc Capel, 1 lot [lota 12 v];
Banc Kelso, 1 lot [6 v]; I. Loyaute, 22 lots [44 v]. FIJI.
South of Viti Levu, 42 lots [328 v]; Southeast of Viti
Levu, 17 lots [57 v]; Bohol/Sulu Seas, 2 lots [5 v]; Bohol
Sea - Balicasag Island, 3 lots [5 v]; Bordau, 1 specimen;
TAIWAN. Bashi channel, 2 lots [3 v]; South China Sea,
1 lot [2 v]; East Taiwan, 2 lots [5 v]. (Details in Simone
and Cunha, in press.)
Despite their larger size, the prickly outer surface of the
ates Spinosipella from the remaining verticordiids, this
taxon has traditionally been considered a subgenus of the
genus Verticordia. This set of characters is sufficient in
my opinion to allocate Spinosipella as a separate genus.
This view was previously defended by the author of the
genus (Iredale, 1930) and by Poutiers and Bernard
(1995). Other distinctive characters are the spiral um-
bones (Figures 5, 7, 21, 22, 33, 54, 53), the tall, some-
what uniform radial sculpture, triangular in section; and
the obesity of the valves. The spiral umbones and the
obesity of Spinosipella are quite similar to those in the
fossil genus Pecchiolia Savi and Meneghini in Murchison,
1850 [type-species (by monotypy): Pecchiolia argentea
Savi and Meneghini in Murchison, 1850 (= Chama ari-
etina Brocchi, 1814) middle Tertiary, Europe] (Keen,
1969: 857), from which Spinosipella differs in having
well-developed ribs and zigzag edges.
The full genus status of Spinosipella is based on the
differences with the typical Verticordia sensu stricto
[type species (by monotypy) Verticordia cardiiformis
Sowerby, 1844], such as the higher size and obesity of the
valves; the additional development of the prickly surface
(which also covers the radial ribs, whereas in Verticordia,
when a prickly surface is present, it does not cover the
radial ribs), the absence of lunule; the spiral fashion of
both valves; and the similarity among the radial ribs (rep-
resentatives of Verticordia usually have an unusually
larger rib or space between ribs). The same set of char-
acters also differentiates Spinosipella from Trigonulina
d’Orbigny, 1842 [type species (by monotypy) T. ornata
d’Orbigny, 1842] in the sense of Jung (1996: 46–47).
Representatives of Spinosipella also resemble those of
the genera Haliris Dall, 1886, and Euciroa Dall, 1881, by
their larger size, convexity, and prickly shell surface. Spi-
higher degree of convexity, reflected in more obese
shells in its species; in the much more developed and
taller radial ribs; higher degree of spiralization of the
valves; and in the expansion of the ribs beyond the shell
phylogeny can be found in the literature (e.g., Pelseneer,
1888; Salvini-Plawén and Haszprunar, 1982; Bieler and
COMPARISON BETWEEN THE
The differentiation between the five species of Spino-
Table 1. The degree of differentiation in the samples of
each species examined allows for specific separations.
The number of radial ribs is the most notable feature;
despite certain a small amount of intraspecific variation,
the number of radial ribs is somewhat constant in each
species, at least in specimens of larger size. The fossil S.
acuticostata is the species with fewest ribs, 12–13 (Fig-
ures 36, 38, 39), while S. deshayesiana has the largest
number of ribs, 16–19 (Figures 46, 48, 49, 54, 53). The
other species possess an intermediary number of ribs.
The species of Spinosipella usually have radial ribs of
relatively uniform size; the single exception is S. costem-
posterior shell slope, however, the ribs abruptly reduce
in size, although in some specimens, particularly in the
young ones, this character is not so clear, i.e., the ribs are
somewhat uniform-sized. The shell inflation is well de-
veloped in most Spinosipella species, but this is clearer in
Tropical W. Atlantic;
Caribbean; to SE Brazil
South and Central
Tropical West Pacific
Prickly ribs outer
Number of Ribs
L. R. L. Simone and C. M. Cunha, 2008
considerably flatter (Figures 41–45). The prickly outer
shell surface is an outstanding character of the Spino-
sipella species; however, this character is conservative
among the five species; the single exception is the rela-
tively chaotic arrangement in S. agnes (Figure 8) and S.
costeminens, while in the remaining species a radial ar-
rangement is apparent (parallel to the radial ribs) (Figure
32). The Pacific species S. deshayesiana has much larger,
crispy prickles along the tip of the ribs (Figures 42, 45,
46, 48, 49). This is lacking in the remaining species,
except in some very young specimens (e.g., USNM
810889, S. agnes, 6 mm), where the prickles, however,
are not fully developed. The prickly surface is strongly
damaged in eroded specimens (Figure 55), becoming
almost completely smooth. Spinosipella deshayesiana,
perhaps because of this character, has the distal tips of
the zigzag edges of the shell even longer and more pro-
jected (Figures 41, 44, 47, 50, 51, 59, 67). The series of
radial ribs is interrupted in the region between the um-
bos, where a triangular smooth area appears. This area is
particularly large in S. agnes (Figures 7, 9), but is prac-
tically absent in S. tinga (Figures 21, 22); it is narrow in
the remaining three species. The size of the specimens
appears to be another distinctive feature, as S. tinga is
small (around 10 mm), whereas the remaining species
are larger (20–30 mm). The hinge does not vary much
between the Spinosipella species; however, some par-
ticularities exist. The posterior tooth of the left valve is
well developed in S. agnes [Figures 4, 10, 12, 27, 28
(arrow)], very low in S. acuticostata (Figures 35, 39), and
practically absent in remaining species (Figures 20, 25,
50). The tall and pointed cardinal tooth of the right valve
is more developed in S. agnes, in such it is also sharply
pointed and curved (Figures 3, 10, 12). In the remaining
species this tooth is weakly shorter and more rounded
(Figures 26, 34, 47, 51).
The geographic and stratigraphic distribution are
somewhat mutually exclusive for most of the species
(Fig. 72); Spinosipella acuticostata is the only Mediter-
ranean species, S. agnes occurs from Florida to Rio de
Janeiro, S. tinga is found from Rio de Janeiro to Rio
Grande do Sul, along the Brazilian coast. The fine-
resolution distribution of the Indo-Pacific species is still
unclear, but S. deshayesiana and S. costeminens, appear
to be sympatric. Spinosipella acuticostata is a fossil spe-
cies, occurring in Pliocene strata, while the remaining
species are found in the Recent. Apparently no Recent
All samples of Spinosipella from the Atlantic and
Mediterranean have previously been accepted as belong-
ing to the single species S. acuticostata (e.g., Abbott,
1974; Abbott and Dance, 1983; Rios, 1994). However,
analyses of the conchological, geographic, and strati-
graphic differences, show that the separation into three
species is warranted. As the shape changes considerably
during ontogeny, a specimen of S. agnes at same size as
the holotype of S. tinga was chosen to show the differ-