j^s-^ 
Vol. 100(3) July 30, 1986 THE NAUTILUS     85 
NOTES ON THE MORPHOLOGY OF ADMETE VIRIDULA 
(GASTROPODA: CANCELLARIIDAE) 
M. G. Harasewych and Richard E. Petit' 
Department of Invertebrate Zoology 
National Museum of Natural History 
Smithsonian Institution 
Washington, DC 20560 
ABSTRACT 
The anatomy and shell morphology of Admete viridula (Fabricius, 1780), a 
boreal cancellariid, are described. This species, the type o/Admete Kroyer, 18U2, 
differs from members of the Cancellariinae and Trigonostominae in having a 
thinner shell composed of a single aragonitic layer, a reduced kidney and jaw, and 
a long, expanded prostate gland, as well as in lacking a raduLa, separate sperm 
ingesting gland and dorsally recurved albumen gland. Based on cladistic analysis 
of 20 shell and anatomical characters, we suggest that the family Admetinae is an 
early offshoot from primitive en,icellariid stock, that the subfamily Trigono- 
stominae contains the most primitive cancellariids studded anatomically to date, 
and that Cancellariinae coinprise a comparatively recent radiation with 
specialized chemosensory capabilities. 
The composition and taxonomic history of the 
superfamily Cancellariacea have been briefly 
discussed in previous papers (Harasewych & 
Petit, 1982,1984). The relationship of this group 
to other prosobranchs is still uncertain, as is the 
phylogenetic arrangement of the nearly 100 
nominal supraspecific taxa proposed almost ex- 
clusively on the basis of shell characters. 
Recent supraspecific taxa have, in general, 
been clustered around three morphological 
types, represented by the genera Cancellaria, 
Trigonostoma and Admete, that have been given 
subfamily status by Cossmann (1899). Members 
of the Cancellariinae and Trigonostominae in- 
habit tropical and temperate waters while 
Admetinae is generally limited to polar regions 
with some species inhabiting deeper waters of 
the temperate zone. The gross anatomy of 
Cancellaria reticulata (Linn?, 1767), the type of 
Cancellariinae, and Olssonella smithii (Dall, 
1888) a trigonostomine, have previously been 
described (Harasewych and Petit, 1982, 1984). 
No general anatomical studies of any adme- 
tine are known to us, although brief descriptions 
of the animal of Admete virid^ula (Fabricius, 
1780) were given by M?ller (1842: 88, as A. 
'Research Associate 
29582. 
P.O. Box 30, North Myrtle Beach, SC 
crispa M?ller) and Jeffreys (1876; 322). Simple 
drawings of the living animal were published by 
H. & A. Adams (1853: pi. 29, fig. 5; copied by 
Tryon, 1885: pi. 7, fig. 32), Troschel (1865: pi. 4, 
fig. 14) and Morse (1921: pi. 7, fig. 43). Troschel 
(1865: pi. 4, figs. 16, 17) figured the jaws of A. 
viridula and the synonymous A. crispa, refer- 
ring to these structures as "hard apparatus" and 
suggesting that they may be toxoglossan 
radular teeth. The latter figure was copied by 
Fischer (1883: 595, fig. 359) and labelled as a 
radular tooth. Tryon (1885: pi. 7, fig. 33) copied 
Fischer's figure without comment. Thiele (1904: 
172) identified Troschel's figures as jaws and 
reported the absence of a radula in A. viridula. 
The austral species A. magellanica Strebel, 
1905, and Waipaoa marwicki Dell, 1956, were 
shown to lack radulae, respectively, by Powell 
(1951: 167) and Beu and Climo (1974: 327). Not 
all admetines are without radulae as evidenced 
by the figures of the radula of Nothoadmete 
t?mida Oliver, 1982, that accompanied the 
description of that species. The gross anatomy 
and shell morphology of Admete viridula 
(Fabricius, 1780), the type species of Admete 
M?ller, 1842 [as A. crispa M?ller, 1842] are here 
described and compared to other cancellariid 
taxa. 
The checkered nomenclatural history of this 
?}' 0 J U 1 
86     THE NAUTILUS July 30, 1986 Vol. 100 (3) 
species is detailed by Bouchet & Waren (1985: 
257), who point out that North American 
authors have used the junior synonym Admete 
couthouyi (Jay, 1839) for the species, based on 
incorrect statements by Ball (1887: 298; 1918: 
318, 328) contending that the type of A. viridula 
was a turrid. Although this was refuted by 
Pilsbry (1938), Macpherson (1971: 107), citing 
Ball, followed other New World malacologists in 
using A. couthouyi. 
Admete viridula is exceedingly variable as at- 
tested by the synonymy listed by Bouchet & 
Waren (1985: 258), to which additional nomina 
will eventually be added. While the Canadian 
specimens used for this study are not "typical" of 
the species, in conchological characters, they do 
fall within the range of variation accepted by 
Bouchet & Waren (1985: figs. 683-689) who ex- 
amined thousands of North Atlantic specimens. 
Materials and Methods 
Anatomical data is based on 20- and 1? 
specimens from the Saguenay River, Quebec 
[48?24'48"N, 70?44'-70?48'W] Voucher Material- 
National Museum of Natural Sciences, Ottawa 
43946. 
Shell fragments for ultrastructural studies 
were broken from the outer lip with pliers. 
Specimens were then immersed in 10% hydro- 
chloric acid (HCl) until the shells dissolved. Soft 
parts were rinsed in distilled water and returned 
to 70% ethanol for dissection. Portions of the 
reproductive systems and anterior proboscis 
were sectioned at a thickness of 6 ^m, and the 
sections stained in hematoxylin and eosin. Bried 
shell fragments were powdered in a mortar and 
pestle, and the X-ray diffraction pattern deter- 
mined on a Philips APB 3600 Automated X-ray 
Powder Biffractimeter. 
Shell Morphology 
External: Shell, reaching 20 mm in length, 
thin, ovate with conical spire and rounded 
anterior (Fig. 1). Protoconch (Figs. 3, 4) pauci- 
spiral, consisting of 1 whorl, with a smooth, 
chalky surface incised by fine, spiral lines. Tran- 
sition to teleoconch marked by beginning of 
axial growth striae and an increase in shell 
thickness. Teleoconch with up to 5V2 convex 
whorls. Early whorls (Fig. 2) strongly shoul- 
dered, becoming more rounded with increase in 
shell size. Suture deeply impressed. Spiral 
sculpture of 16-21 shallow, rounded cords on 
body whorl and 7-10 on the penultimate whorl. 
Axial sculpture of 14-24 ribs per whorl, increas- 
ing in number but decreasing in prominence as 
the shell becomes larger. Aperture elliptical, 
deflected from coiling axis by 20-28?. Outer lip 
smooth to slightly corrugated. Siphonal canal 
short but pronounced in juveniles (Fig. 2), com- 
monly indiscernible in large adults (Fig. 1). 
Inner lip with 2 weak columellar folds and a 
siphonal fold. Color white to ivory, uniform 
within and without. 
Internal: Fractured shells revealed the inter- 
nal surfaces to be smooth and continuous, and to 
lack the apertural lirae and periodic increases in 
prominence of columellar folds found in other 
cancellariids (Harasewych and Petit, 1982, 
1984). 
Ultrastructure: A thin, brownish, lamellose 
periostracum overlies the shell, which appears 
to consist of a single layer of crossed-lamellar 
crystals (Fig. 5), oriented with lamellar planes 
parallel to the outer lip. X-ray diffraction 
analysis of powdered shell revealed it to consist 
primarily (>95%) of aragonite. 
Soft-parts Morphology 
External features: The soft-parts comprise 3 
to 3V3 whorls, of which the digestive gland (Fig. 
7, dg) occupies about 2V4 whorls, the kidney 
(Fig. 7, k) less than Vs whorl and the mantle cavi- 
ty just over V2 whorl. Preserved animals were 
orange tan in color, lacked any discernible 
markings and were retracted at most 74 whorl 
into the aperture. The foot is broad and ovate, 
lacking an operculum. Tentacles (Fig. 7, t) are 
long, tubular and symmetrical. The mantle edge 
is smooth, the siphon (Fig. 7, s) short but 
distinct. 
Mantle cavity: The mantle cavity is shorter 
and broader than in other cancellariids dis- 
sected, with palliai organs situated as in other 
higher prosobranchs. The osphradium (Fig. 7, 
os) is broad anteriorly (L/W = 3), tapers 
posteriorly, and consists of about 35 leaflets per 
side. Adjacent is the ctendium (Fig. 7, ct), twice 
as long and slightly narrower than the 
osphradium, composed of about 70 triangular 
leaflets. This organ and the voluminous, trans- 
versely pleated hypobranchial gland (Fig. 7, hg) 
Vol. 100 (3) July 30, 1986 THE NAUTILUS     87 
FIGS. 1-6. Features of the shells and jaw oi Admete vtridula {Fiibvicms). I, Apertural and right side views of specimen 
collected in the Sag-uenay River, Quebec. [48?24'48"N, 70?44'-70?48'W] sta. 62 30 (NMNS 43946) 3.0 x . 2, Scanning elec- 
tron micrograph of juvenile specimen dredged in 673 m, off Martha's Vineyard, Massachussetts, U.S. Fish. Comm. sta. 
994-97. (USNM 43232) 20.0 x. 3, Protoconch of specimen in figure 2, lateral view, scale bar = 200 fjm. 4, Protoconch of 
specimen in figure 2, axial view, scale bar = 200 f^m. 5, Fracture surface. Plane of fracture parallel to outer lip, scale bar 
= 100 fjm. 6, Lateral view of jaw, scale bar = 50 (jm. 
88     THE NAUTILUS July 30, 1986 Vol. 100(3) 
rmc 
i'^IGS. 7-10. Anatomical features oi Admelt i<iridida. 7, Female specimen removed from shell, partially un- 
coiled and mantle cavity opened mid-dorsally to display contents. 8, Dissection of anterior portion of proboscis, 
opened mid-dorsally. 9, Diag-rammatic representation of the alimentary system. 10, Male reproductive system, 
a, anus; ag, albumen gland; asg, accessory salivary gland; be, bursa copulatrix; bg, buccal ganglia; bm, buccal 
mass; eg, capsule gland; ct, ctenidium; dg, digestive gland; hg, hypobranchial gland; int, intestine; ,j, jaw; k, 
kidney; m, mouth; me, mid-esophagus; ng, nephridial gland; nr, nerve ring; opm, opening to mantle cavity; os, 
osphradium; ot, oral tube; pap, papilla; pc, pericardium; pen, penis; pr, prostate gland; r, rectum; rmc, rear of 
mantle cavity; s, siphon; sg, salivary gland; sto, stomach; sv, seminal vesicle; t, tentacle; tes, testes; vd, vas 
deferens; vl, valve of Leiblein. 
span the roof of the mantle cavity. The palliai 
gonoducts and rectum (Fig. 7, r) are located 
along the right side of the mantle cavity, while 
its rear is formed by the pericardium (Fig. 7, pc) 
and kidney (Fig. 7, k). The kidney is small, being 
only twice the size of the pericardium. The 
nephridial gland (Fig. 7, ng) occupies nearly half 
the kidney along the pericardium. 
Alimentary system: The broad, tubular pleu- 
rombolic proboscis can be extended nearly the 
length of the shell aperture, and is the same 
color as the foot. Broad retractor muscles ex- 
tend from the columellar muscle, while thinner 
strands originate from the body wall. A short 
oral tube (Fig. 8, ot) extends from the mouth 
(Fig. 8, m) to the cuticularized, tubular portion 
of the jaw (Fig. 8, j), which expands posteriorly 
to envelope the lateral surfaces of the minute 
buccal mass (Fig.  8, bm). The longitudinally 
grooved buccal mass lacks a radula and has only 
a vestigial subradular membrane. A pair of 
short, extremely thin accessory salivary glands 
(Fig. 8, asg) lie alongside the buccal mass, 
emptying into the posterior portion of the oral 
tube through fine ducts. The wider and much 
longer salivary glands (Fig. 8, sg) join the buccal 
mass laterally, via short ducts, just anterior to 
the esophageal opening. A small valve of 
Leiblein (Fig. 8, vl) and a pair of buccal ganglia 
(Fig. 8, bg) are situated at the rear of the bucal 
mass. The narrow mid-esophagus (Figs. 8, 9, 
me) runs posteriorly from the valve of Leiblein, 
passing through the nerve ring (Fig. 9, nr) 
before expanding and becoming sacular. The 
esophagus narrows at the rear of the cephalic 
sinus and leads to a simple U-shaped stomach 
(Fig. 7, 9, sto) that is embedded in the anterior 
face of the digestive gland. Longitudinal folds 
Vol. 100(3) July 30, 1986 THE NAUTILUS     89 
TABLE 1. Admete viridula (Fabricius), measurements of 
shell characters. Linear measurements in millimeters, (n = 
10). 
Standard 
Character Mean       Deviation Range 
shell length 16.87 1.91 13.82-19.02 
shell width 9.90 1.13 8.18-10.97 
aperture length 10.21 1.42 8.13-11.58 
aperture length. 0.604 0.023 0.567-0.641 
shell length 
#whorls, teleoconch 4.78 0.35 4.33-5.33 
spire angle 55.2? 2.9? 45.5?-58.0? 
line the stomach and pass into the intestine (Fig. 
9, int), which runs through the kidney and along 
the right wall of the mantle cavity, expanding 
slightly to form the rectum (Figs. 7, 9, r). 
Neither an anal gland nor identifiable gut con- 
tents were detected in any of the specimens ex- 
amined. 
Female reproductive system: As the only 
female specimen available to us was poorly pre- 
served, only the palliai portion of the oviduct is 
described. The oviduct passes through the 
kidney and expands to form a broad, laterally 
compressed albumen gland (Fig. 7, ag), which 
expands further to form the long, blunt, capsule 
gland (Fig. 7, eg). A small, ovate, bursa copula- 
trix (Fig. 7, be) is situated below the anterior 
margin of the capsule gland. An ingesting gland 
was not found. Egg capsules o? Admete viridula 
are figured by Bouchet and Waren (1985: fig. 
687). 
Male reproductive system: The testes (Fig. 10, 
te) consist of a yellowish mass of ascinous 
tubules embedded in the right side of the diges- 
tive gland and extend nearly a full whorl from 
its anterior edge. These tubules converge to 
form a testicular duct that immediately forms a 
serpentine seminal vesicle (Fig. 10, sv). The duct 
straightens at the anterior limit of the digestive 
gland, passes through the kidney and enters the 
mantle cavity ventral to the rectum. It greatly 
expands to form the prostate gland (Fig. 10, pr), 
which extends along the posterior third of the 
mantle cavity and communicates with it by a 
long ventral slit (Fig. 10, opm). From the 
anterior portion of the prostate gland, a tubular 
vas deferens (Fig. 10, vd) leads to the base of the 
penis (Fig. 10, pen), which is ovate in cross- 
section, extends approximately % the length of 
the mantle cavity and has a terminal papilla 
(Fig. 10, pap). 
Nervous system: Admete viridula has a typical 
cancellariid nervous system (Bouvier, 1887: 
Harasewych and Petit, 1982, 1984), with the 
cerebral, pedal and pleural and subesophageal 
ganglia fused into a nerve ring, and the buccal 
ganglia (Fig. 8, bg) at the rear of the buccal 
mass. 
Discussion 
Cladistic analysis of the 20 shell and 
anatomical characters listed in Table 2 produced 
the cladogram in Figure 11. Despite the lack of 
agreement on the relationships of supraspecific 
taxa, nearly all classifications proposed to date 
separate the Admetinae from the remaining 
Cancellariidae (H. and A. Adams, 1853: 
Troschel, 1865; Fischer, 1883; Tryon, 1885; 
Cossmann, 1899; Thiele, 1929; Wenz, 1943), and 
this arrangement is supported by the present 
study. 
Of the characters used to distinguish the 
Admetinae from other cancellariids, most con- 
TABLE 2. Shell and anatomical characters used to distin- 
guish between Admete viridula, Co.ncellaria reticuLato, and 
Olssonella smilhii. Primitive states denoted by (0), derived 
states by (1). 
1) Shell: (0) thick; (1) thin. 
2) Internal varices: (0) pronounced; (1) reduced or absent. 
3) Shell ultrastructure of: (0) 2 orthogonal layers; (1) 1 
layer. 
4) Protoconch of: (0) 2 whorls; (1) 1 whorl. 
5) Tentacles: (0) symmetrical, unmodified; (1) asymmetri- 
cal, left tentacle modified. 
6) Osphradium with approximately: (0) 35 leaflets/side; (1) 
70 leaflets/side. 
7) Ctenidium with: (0) < 125 leaflets; (1) > 200 leaflets. 
8) Distance between osphradium and ctenidium: (0) nor- 
mal; (1) large relative to either organ. 
9) Kidney occupying: (0) > 'A whorl; (1) < Vs whorl. 
10) Proboscis: (0) tubular; (1) ventrally flattened, papillose. 
11) Jaws with: (0) short posterior lobes; (1) long posterior 
lobes. 
12) Buccal mass: (0) small; (1) large, filling retracted pro- 
boscis. 
13) Radula: (0) present; (1) absent. 
14) Outer cusps of radular teeth: (0) simple, smooth; (1) 
with secondary dentition. 
15) Accessory salivary gland: (0) shorter than; (1) longer 
than; salivary gland. 
16) Mid-esophagus posterior to nerve ring: (0) sacular; (1) 
convoluted tube. 
17) Anal gland: (0) present; (1) absent. 
18) Sperm ingesting gland: (0) present; (1) absent. 
19) Albumen gland: (0) dorsally recurved; (1) laterally com- 
pressed. 
20) Prostate  gland:  (0) long,  expanded;  (1) short,  con- 
voluted. 
90     THE NAUTILUS July 30, 1986 Vol. 100(3) 
3 
> 
< 
re 
TO 
3 ? 
O 
E 
FIG. U. Cladogram of phylogenetic relationships of cancel- 
lariid taxa. Single slashes across tree branches represent 
transformations of the corresponding character from the 
primitive (0) to the derived (1) state. Double slashes indicate 
transformations that occur more than once. 
spicuous are their thin shells lacking pro- 
nounced surface sculpture and apertural lirae. 
This is likely a consequence of their polar 
habitat, as Graus (1974) has shown that de- 
creased calcification in high latitudes is due to 
reduced availability of calcium carbonate in 
colder waters. Admetines also lack such anti- 
predatory features as internal varices that are 
found in tropical and temperate cancellariids. 
The predominance of aragonite in the shell is 
plesiomorphic, as calcite is more easily formed 
(Wilbur, 1964) and more stable (Lowenstam, 
1954) at low temperatures. 
In terms of anatomical organization, Admete 
viridula differs from ancestral cancellariid mor- 
phology, most nearly approximated by Olsso- 
nella smithii of the taxa studied to date, in hav- 
ing a shallower mantle cavity, a reduced kidney, 
and in lacking color pattern. At least some of 
these may also be adaptations to cold water. 
Cancellariines and Trigonostomines have uni- 
serial "tricusped" radular teeth with elaborate 
secondary dentition on the outer cusps and a 
comparatively simple ventrally recurved central 
cusp (Harasewych and Petit, 1982, 1984; Petit 
and Harasewych, 1986). The radula of Notho- 
admete t?mida, the only known radulate 
admetine, has barbed central cusps and simple 
bulbous outer cusps (Oliver, 1982: figs. 3, 5). 
Presence of central cusp barbs in Cancellaria 
atopodonta Petit and Harasewych (1986: figs. 
15, 16) suggests that this feature is primitive 
and occurred in the ancestor of all Recent 
cancellariids. The jaws of admetines lack the 
long posterior lobes found in cancellariines and 
trigonostomines. Although the diet of cancel- 
lariids remains unknown, we have suggested 
that they are piercing suctorial feeders based on 
the functional morphology of their alimentary 
systems (Harasewych and Petit, 1982, 1984; 
Petit and Harasewych, 1986). The loss of radula 
and reduction in the size of the buccal mass in 
most admetines suggests that they feed suc- 
torially, but on a different group of prey 
organisms than other cancellariids. 
The lack of an anal gland in Admete viridula 
and Cancellaria reticulata is likely a com- 
paratively recent convergence, as this organ has 
been reported in Cancellaria cancellata 
(Graham, 1966). 
Additional differences between admetines and 
other cancellariids are found in the reproductive 
systems. The prostate gland of Admete viridula 
is long and greatly expanded, while in Olssonella 
smithii and Cancellaria reticulata it is short and 
convoluted. Admete viridula also lacks the 
separate sperm ingesting gland and dorsally 
recurved albumen gland found in cancellariines 
and trigonostomines. 
The phylogenetic arrangement in Figure 11 is 
supported by the fossil record, as several 
"admetines" have been reported from the Upper 
Cretaceous, and Waipaoa, known to be 
aradulate in the Recent fauna, dates back to the 
Oligoc?ne. Cancellaria, sensu stricto, first ap- 
pears in the Miocene and is characterized by 
modifications to the left cephalic tentacle, pro- 
boscis, osphradium and ctendium, organs in- 
volved in tactile and distance chemoreception. 
Acknowledgments 
We thank Jane B. Topping, Invertebrate 
Zoology Division, National Museum of Natural 
Sciences, Ottawa, for making available the 
preserved specimens used in this study. Some of 
this work was done at the Smithsonian Marine 
Station at Link Port, Florida, and we thank Dr. 
Vol. 100(3) July 30, 1986 THE NAUTILUS     91 
Mary Rice for making these facilities available. 
This is contribution number 158 of the Smith- 
sonian Marine Station at Link Port. Critical 
review of the manuscript by Dr. R. S. Houbrick, 
National Museum of Natural History, Smith- 
sonian Institution, is gratefully acknowledged. 
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BOOK REVIEW 
Washington Public Shore Guide: Marine Waters 
by James W. Scott and Melly A. Reuling. 
1986. 348 pp., numerous photos, maps and 
habitat sketches. Cloth, $25.00; paperback, 
$14.95. 
If you are planning on studying or visiting the 
shorelines of the State of Washington, this is a 
great bargain with many useful facts about the 
2,400-mile saltwater shoreline of 14 counties. It 
largely ignores mollusks, a fact which offers a 
challenge to conchologists.-?. T. Abbott.