ences between those species. Figures 28–31 illustrate
these differences. Spinosipella agnes has fewer, taller,
and more widely spaced ribs than S. tinga (Figures 19,
29). The shape of the shell edge is much more uniform
in S. tinga than in S. agnes, in that the tips of the ribs are
more expanded, extending longer beyond the shell mar-
gin (Figures 20, 24, 28). The posterior cardinal tooth in
the hinge of the left valve is present in S. agnes, in ani-
mals larger than 5–6 mm, while this tooth is never
present in S. tinga (Figures 20, 25, 27–28, arrow). The
degree of convexity is higher in S. agnes and in S. tinga
(Figures 30, 31); S. agnes has a degree of convexity
(width/length) in each valve of about 0.57, while S. tinga
it is 0.47.
The comparison of the previously valid species Spino-
sipella ericia, including paratypes (Figures 41–44), and S.
deshayesiana, does not reveal any distinction between
them. Normally, specimens of smaller size were identi-
fied as S. ericia, and the large ones as S. deshayesiana.
But examination of shell features along a growth series
show a complete gradient linking the two taxa. The same
lack of distinction is found in the literature for both spe-
cies, including the original descriptions. For these rea-
sons, despite the fact that S. ericia is the type species of
the genus, the older name S. deshayesiana should be
used. Furthermore, a type specimen of S. japonica was
also examined (Figure 45), confirming the synonymy of
this species with S. deshayesiana.
The distinction between the Pacific species Spino-
sipella deshayesiana and S. costeminens is not always
easy. With the large quantity of specimens kindly pro-
vided by the MNHN (Paris), it was possible to analyze
the degree of variation of both species. Spinosipella
outstandingly large, carina-like spiral ridge between the
middle and posterior thirds of the shell, but sometimes
this ridge is not so different from the others, and the
animal become more rounded, similar to S. deshayesi-
ana. The distinction is based mainly on the presence of at
least a weak carina in the region between middle and
posterior thirds, and also by the more robust ridges of S.
costeminens specimens (Figures 60, 61), while those of S.
deshayesiana lack any clear radial carina and the ridges
are more delicate, uniform and apparently close from
each other (Figures 46, 48).
The lot USNM 63200 includes 3 valves (2 left and 1
right), collected in Barbados, the known geographic dis-
tribution of Spinosipella agnes. However, the right valve
has the characters of S. deshayesiana, instead of those of
S. agnes. In addition, is looks different in the state of
conservation, color and associated sediment, from the
other 2 valves of the same sample.
DISCUSSION ON ANATOMY
More in-depth anatomical descriptions and discussions
on verticordiids are provided by Allen and Turner
THE NAUTILUS, Vol. 122, No. 2
ever, no information on the anatomy of the genus Spi-
information is available here only for two of the five
species of the genus (of course one of them is a Pliocene
fossil), some systematic inferences can be made based on
the scenario given in the literature the Verticordiidae and
related families (Allen and Turner, 1974, and others, e.g.,
Fisher, 1860, 1862b; Pelseneer, 1888; Nakazima, 1967;
Allen and Morgan, 1981). Besides the conchological
characters discussed above, some anatomical features are
possibly restricted to Spinosipella, such as: the wide lith-
odesma (Figures 93, 96–104, lt); the simplified siphonal
tentacles (Figure 108), which normally have secondary
papillae; the papilla on the roof of the excurrent chamber
(Figures 78–80, 85, 94: pi); the absence of incurrent
valve in infraseptal chamber. However, wide lithodesma
have been reported for Policordia lisbetae Knudsen,
1970 (fig. 90), which has very different shell and pallial
tentacular characters. The study on the incurrent sipho-
nal structures is of particular importance in septibranchs,
as the modified incurrent siphon is the main structure
used in prey capture (Morton, 1987).
On the other hand, some features appear to be char-
acteristic of Verticordiidae, such as: elongation of lateral
region of kidneys; the muscular stomach (see also Pur-
chon, 1956, 1963); the separation between testis and
ovary. By the proximity of the esophagus from anterior
adductor muscle, by the lack of incurrent valve, and by
the simplified buccal structures, e.g., lack of buccal cavity
and tongue, it is possible to suggest that Spinosipella is a
basal taxon inside Verticordiidae. Unfortunately, no
member of the genus was analyzed in the recent com-
parative studies on anomalodesmatans (Harper et al,
the material for this study: Winston Ponder and Ian Loch
(AMS) for types of Spinosipella ericia; Néstor E. Ardila
(MHNMC) and Emilio Garcia (EGC) for S. agnes (Co-
lombia) and S. deshayesiana (Philippines); Harry G. Lee
for S. agnes (Florida) (MHNMC) for a large lot of S.
and Philippe Maestrati, MNHN, for the loan of a huge
quantity of lots coming from several places of the world,
mostly from the Indo-Pacific. For thorough comments
and additional information about S. acuticostata we
thank Rafael La Perna. For material of Haliris fisheriana
we thank to Daniel Mansur Pimpão (PPG-BAN,
UFRGS). For help with SEM procedures, we thank Lara
Guimarães (MZSP). For Rachel Collin, Smithsonian In-
stitution at Panama, we thank for the help in the text and
language. We thank also both referees and the Editor for
the thoughtful correction on the manuscript. This project
is supported by FAPESP (Fundação de Amparo a Pes-
quisa do Estado de São Paulo), procs. no. 03/05860-6,
04/02333-8, and a “Treinamento Técnico 3” grant, under
the supervision of Antonia Cecília Z. Amaral and Luiz
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