Notes on the genus Anadema H. and A. Adams, 1854 (Gastropoda : Colloniidae)

2008·7.2 MB·English

by  James H Mclean

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SociedadEspañoladeMalacologta Iberus, 26 (1): 53-63, 2008
Notes on the genusAnadema H. and A. Adams, 1854 (Gastropoda:
Colloniidae)
Notas sobre el género Anadema H. yA. Adams, 1854 (Gastropoda:
Colloniidae)
James H. MCLEAN* and Serge GOFAS**
Recibidoel15-1-2008.Aceptadoel23-IV-2008
ABSTRACT
Shell morphology and characters of the living animal ofthe poorly known, Atlantic Moroc-
can species Anadema mocandrewii (Mórch, 1864) are described and illustrated, based
on beach collected specimens and a single live-collected specimen. The genus ¡s mono-
typic and is assigned to the Colloniidae rather than Turbinidae because of the dome-
shaped profile of the shell, open umbilicus, symmetrical tooth rows of the radula, lack of
cephalic lappets, and the non-bicarinate ¡uvenile shell. Within the Colloniidae, it unusual
for its relatively large mature shell, ¡uvenile shell with a keeled profile, and the lack of the
secondary flap above the rachidian tooth. The species is regarded as sexually dimorphic,
with the female shell having a raised periumbilical rim comparable to that of other tro-
choideans modified for brooding by means ofan enlarged umbilical cavity.
RESUMEN
Se describe e ilustra la morfología de la concha y del animal vivo de Anadema macan-
drewii (Mórch, 1864), una especie poco conocida de la costa atlántica de Marruecos. El
género es monotípico y se asigna a la familia Colloniidae, en lugar de a los Turbinidae
por la forma abombada de la concha, el ombligo abierto, las filas de dientes radulares
simétricas, la ausencia de lóbulos cefálicos y por su concha juvenil no bicarenada. Entre
los Colloniidae, la especie es insólita por el tamaño relativamente grande de la concha
adulta, el perfil de la concha juvenil con una quilla y la ausencia de un repliegue secun-
dario sobre el diente raquídeo. Se considera que existe dimorfismo sexual en esta
especie, pues la concha de la hembra tiene un reborde periumbilical elevado comparable
con el de otros trocoideos modificados para incubar en una cavidad umbilical ampliada.
KEYWORDS:Trochoidea, Morocco,brooding,endemism.
PALABRAS CLAVE:Trochoidea,Marruecos, incubación,endemismo.
INTRODUCTION
The poorly known genus Anadema been regarded as a trochid, a liotiid, a
H. and A. Adams, 1854 has at times colloniid, and a turbinid. Here we
*NaturalHistoryMuseumofLosAngelesCounty, California90007,USA.
**DepartamentodeBiologíaAnimal,FacultaddeCiencias, UniversidaddeMálaga,E-29071 Málaga, Spain.
53
Iberus, 26 (1), 2008
review the taxonomy and update what figured and the repository of the oper-
is now known about the type and only culate specimen examined by Morch is
knownspeciesofthisgenus. unknown.
Anadema was proposed for the com- Pilsbry (1888) ignored the replace-
bination Omphalius (Anadema) caelata A. ment ñame and called it Leptothyra
Adams, 1854, an Atlantic Moroccan caelata, a not unreasonable choice, as it
species then thought to be a trochid in hardly resembles a species of Turbo. Lep-
the absence of data on a calcified oper- tothyraisnowassignedtoColloniidae.
culum. The species was not to be illus- Wenz (1938: 340) recognized the
trated until 19 years later but it was genus Anadema and placed it in the
described in sufficient detail that its turbinid subfamily Liotiinae, under a
identity has never been in question. In broad definition in which he also
his remarks that followed the Latin included genera with calcareous oper-
description of the species, Adams (1855: cularelatedtoHomalopoma.
39) wrote: "The character of this shell is Keen (1960: 270) placed Anadema in
so peculiar, on account of the internal the more restricted Homalopomatinae, a
spiral callus of the umbilicus, and the group with fully calcified operculum,
absence of characters which constitute now subsumed under the turbinid sub-
alliedforms,thatIproposetoconsiderit family Colloniinae in the classification
a subgenus of Omphalius under the ofHickmanandMcLean(1990).
,
ñame of Anadema." The generic ñame Nordseick (1968: 33) overlooked
Omphalius Philippi, 1847 now pertains Mórch (1864) and placed itin Liotiinae:
to the Tegulinae, as defined by "Deckel spiralig und mit Kalkbesatz,
HickmanandMcLean(1990). nicht verdickt", apparently having
Ten years after the introduction of assumed thatitmusthave the opercular
Anadema Mórch (1864: 46) reported definition of the now understood Lioti-
,
that a living specimen had been col- idae, in which the operculum is multi-
lected at the type locality. Morch stated spiral with calcareous beads on the
that: "Omphalius (Anadema) caelata is outersurface.
provided with a calcareous operculum, Resolution of the uncertainty is here
which proves that this species may be provided by a preserved immature
removed to Turbo. As there is already a specimen with operculum, which was
T. caelata, L., I propose for this species collectedbythe secondauthorin 1991 at
Turbo macandrewii." Morch's placement Essaouira (formerly Mogador), the type
of the species in Turbo predated most locality. We therefore take this opportu-
work on the genera of turbiniform gas- nity to illustrate the species and confirm
tropods, and was based on the assump- itsplacement.
tion that any turbiniform species with a
calcareous operculum could only be a
Turbo. Thiswas unfair toArthurAdams, MATERIALS
but the ICZN rules about secondary
homonymy forcé us to abandon the This report is based on material col-
original ñame and to use the replace- lected by the second author, which is
ment ñame. This ñame honors the re- now in the malacology collection of the
discoverer Robert McAndrew, who Muséum National d'Histoire Naturelle,
dredged a living specimen at Mogador Paris (MNHN). The external features of
(now Essaouira), the type locality of the the single live-collected specimen were
species. drawn with the animal fully extended.
The first shell figure known to us Subsequent preservation of the speci-
was provided still later by P. Fischer men resulted in retraction within the
(1873), who used the replacement ñame shell. It was later critical-point dried for
Turbo macandrewii; this is a drawing that SEM examination, after which it was
has been copied by many subsequent rehydrated for extraction of the radula
authors. The operculum has never been forSEManalysis.
54
McLeanAND GOFAS: Notes on thegenusAnademaH. andA. Adams, 1854
SYSTEMATICS
SuperfamilyTrochoideaRafinesque, 1815
Hickman and McLean (1990) we here treat the Liotiidae, Colloniidae,
divided the family Turbinidae into a and Turbinidae at the family level
number of subfamilies, including Lioti- within the superfamily Trochoidea,
inae, Colloniinae, and Turbininae. which simply raises the ranking in the
However, Williamsand Ozawa (2007) existing classification. This has already
have reported that their data toward a been adopted by McLean and Kiel
molecular phylogeny of the family (2007). Additionally, and without dis-
Turbinidae indicates that there are two cussion,Warénand Bouchetin Bouchet
well-supported groups withintheprevi- and Rocroi (2005: 245) have separated
ously defined Turbinidae, which there- Turbinoidea and Trochoidea at the
fore preeludes the placement of all of superfamily level, which also indicates
the subfamilies in the same family. In that the higher classification for trochi-
view of the ongoing reconsideration of form vetigastropods is currently unset-
the relationships among these groups. tled.
FamilyColloniidae Cossmann, 1916
SubfamilyColloniinae Cossmann, 1916
Hickman and McLean (1990) pro- loniidae, based on opercular morphol-
vided an extensive treatment ofthe Col- ogy: the basal and mostly extinct
loniidae (then as Colloniinae), distin- Petropomatinae Cox in Knight et al.,
guishing them from Turbinidae (then as 1960, having a calcified operculum that
Turbininae) ontheirsmallersize,non-bi- is conical on the inner surface, with a
carinate juvenile shell, lack of cephalic fully multispiral pattern, and the Col-
lappets,symmetricaltoothrowsandinner loniinae, in which the operculum is fíat
lateralteeththatarenotgreatlyenlarged. on the inner surface with a multispiral
This is in contrast with the Turbinidae, pattern that changes to broadly pau-
characterizedbylargersize,bicarinateju- cispiral on the final volution. There is
venileshell,asymmetricaltoothrowsand one living genus (Liotipoma McLean and
enlargedinnerlateralteethoftheradula. Kiel, 2007) of Petropomatinae, whereas
McLean and Kiel (2007) distin- there are numerous living and fossil
guished two subfamilies withinthe Col- generaofColloniinae.
GenusAnadema H. andA. Adams, 1854
Anadema H. and A. Adams, 1854: 430 [as subgenus of Omphalius Philippi, 1847]. Type species
(monotypy): Omphalius (Anadema)caelataA.Adamsin H. andA.Adams, 1854 [= Turbomacan-
drewiiMorch,1864;notTurbocaelataLinnaeus,1758].
Anadema macandrewii (Morch, 1864) (Figs. 1-23)
Omphalius(Anadema)caelataA.Adams,inHandA.Adams,1854:430 [astypeofnewsubgenus].
A.Adams,1855:39 [moredetaileddescription].
Turbo macandrewii Morch, 1864: 46 [new ñame for secondary homonym Omphalius (Anadema)
caelata,notTurbocaelataLinnaeus,1758].
Turbo macandrewi. Fischer, 1873: 98, pl. 29, fig. 3 [firstillustration]. Pasteur-Humbert, 1962: 132
[listed].
Leptothyracoelata [sic].Pilsbry,1888:255,pl.48,fig.38 [figureafterFischer],
55
y
Iberus, 26 (1), 2008
Turbo(Anadema)macandrewi.Pallary, 1920:63.
Anadema coelata [sic]. Wenz, 1938: 340, fig. 795 [figure after Fischer]. Nordesick, 1968: 33, fig.
17.00
Anadema caelata. Keen, 1960: 270 [no figure]. Trew, 1992: 19 [listing ofspecies describedby H.&
A.Adams].
Material examined: BMNH, 2 syntypes, BMNH 1968183, Mogador (now Essaouira), Atlantic
Morocco(height12.3mm,diameter16.6mm;height12.1mm,diameter17.3mm).MNHN,Essaouira
(formerlyMogador),AtlanticMorocco (31° 31' N, 9o47' W), 1 live-collected male specimenand
severalbeach-wornshells,leg.Gofas,23September1991.MNHN,ElJadida,AtlanticMorocco(33°
16' N, 8o 29' W),beachwornshells, leg. Gofas, 26September, 1991. MNHN,Mohammedia (for-
merlyFedala),AtlanticMorocco(33°43'N,7o21'W),10beach-wornshells,leg.Gofas,1970-71.
Description: Because the genus is developed in interspaces but not
monotypic, the description that follows expressed on surface ofnodularbeads;
appliesbothtothegenusand species. umbilicusopeninjuvenileshell,closedin
Shellcomposedof5whorlswithalow male shell at basal diameter of 9 mm;
spire;profiledome-shaped,broaderthan remainingopeninfemaleshell;umbilicus
high,suturenotimpressed;whorlsweakly ofmaturefemaleshellborderedbyraised,
rounded, periphery of immature shell unbeaded peripheral cord thatpartially
stronglyprojecting,formingakeel;periph- obstructsfinalquarterwhorlandconnects
ery spinose in early stages, but losing directly at base of aperture; aperture
spination atmaturity; mature shellwith oblique, thickened within, descending
asubangulatebase;axialsculptureoffine, slightlyonfinalwhorl;interiornacreous,
raised lamellae; spiral sculpture oflow, inner wall of aperture of female shell
strongly beaded cords; cords between smooth,edgewithU-shaped sinus; shell
suture andbeadedbasal cord increasing colourbrick-red.
from three injuvenile to 6 at maturity; Shell dimensions. Mature female
base slightlyconvex,basalcords ofcom- shell (Figs. 1-3): height 13.0, diameter
parable strengthandbeading to those of 17.1 mm; immature male shell (Figs. 4-
bodywhorl,increasingfromthreeinjuve- 6): height 4.9, diameter 9.1 mm;
nile to 8 at maturity; spiral sculpture of máximum dimensions possible for male
bothbodywhorl andbase separatedby shell unknown; immature female shell
narrow interspaces; axial lamellae well (Figs. 7-9),height5.9, diameter9.1 mm.
(Rightpage) Figures 1-15.Anadema macandrewii(Morch, 1864). 1-3: Mature, beach-worn female
shell, from Essaouira, Morocco (MNHN), 3views, height 13.0 mm, diameter 17.1 mm; 4-6: live-
collected, immature male specimen with operculum in place, same locality (MNHN), 3 views,
height 4.9 mm, diameter 9.1 mm; 7-9: immature, beach-worn female shell, from Mohammedia,
Morocco (MNHN), 3views, height 5.9 mm, diameter9.1 mm; 10, 11:juvenile beach-worn shell,
from Essaouira, Morocco (MNHN), 2 views, height 2.1 mm, diameter 5.0 mm; 12,13: juvenile
shell with protoconch showing in umbilical view, from Essaouira, Morocco (MNHN), 2 views,
diameter2.7 mm; 14, 15: operculum ofspecimen shown in Figures4-6 and 16, exteriorand inte-
riorviews, máximumdiameter2.1 mm.
(Página derecha) Figuras 1-15. Anadema macandrewii (Morch, 1864). 1-3: Concha adulta,
explayada, de una hembra, deEssaouira, Marruecos (MNHN), 3 vistas, altura 13,0 mm, diámetro
17,1 mm; 4-6: ejemplarmacho inmaduro, recolectado vivo, con elopérculo cerrando la concha; misma
localidad(MNHN), 3 vistas, altura4,9mm, diámetro9,1 mm; 7-9: concha inmadura, explayada, de
una hembra, deMohammedia, Marruecos (MNHN), 3 vistas, altura5,9mm, diámetro9,1 mm; 10,
11: conchajuvenilexplayada, deEssaouira, Marruecos (MNHN), 2 vistas, altura 2,1 mm, diámetro
5,0 mm; 12, 13: conchajuvenilcon laprotoconcha visible desde elombligo, deEssaouira, Marruecos
(MNHN), 2vistas, diámetro2,7mm; 14, 15: opérculodelejemplarrepresentado en lasFiguras 4-6
16, vistasexterioreinterior, diámetromáximo2,1 mm.
56
McLeanAND GOFAS: Notes on thegenusAnademaH. andA. Adams, 1854
57
Iberus, 26 (1), 2008
Figure 16.Anademamacandrewii(Morch, 1864). DrawingoflivingspecimenshowninFigures4-6.
Figura 16. Anademamacandrewii (Morch, 1864). Dibujodelanimalvivo, delmismoejemplardelas
Figuras4-6.
Juvenile shell (Figs. 10-13) not with the upper part, which bears strong pus-
raised axial lamellae, of low profile, tules in the región adjacent to the col-
exposing protoconch and early whorls umella; outer edge with a narrow
in basal view; profile not equally bicari- groove; inner surface becoming broadly
nate, the upper carination spinose and paucispiral in final volution. This oper-
more strongly projecting than lower, culum is chipped or somehow reduced
non-spinose carination; protoconch and at its inner edge, to the extent that the
first teleoconch whorl positioned outline is not oval; the chitinous layer of
slightlybelowlevelofsecondwhorl. theinnersurfacethatshows atthelower
Operculum (Figs. 14-15) calcareous, left of Figure 15 is missing at the upper
thick,withaconcavityatthebasewhere left.
the white outer surface is coloured light Flead-foot (Figs. 16-19). Head pro-
reddish-brown. There is a strong ridge portionally large, provided anteriorly
and groove of similar width leading to with a snout terminating in a broad.
(Right page) Figures 17-23. Anadema macandrewii (Morch, 1864). 17-19. Scanning electrón
micrographs ofcritical-point dried specimen, same as shown in Figure 16. 17: general view of
head-foot; 18: close-up ofsnout; note the absence ofcephalic lappets; 19: close-up ofright ante-
rior bundle ofepipodial tentacle. 20-23. Scanningelectrón micrographs ofthe radula, same speci-
men as Figure 16. 20: complete radula; 21: close-up ofseveral complete rows; 22: detail ofcentral
and lateral teeth; 23: detail ofmarginal teeth. Abbreviations, ct: cephalic tentacle; et: epipodial
tentacles; f: foot (pointingtopropodium); sn: snout.
(Página derecha) Figuras 17-23. Anadema macandrewii (Morch, 1864). 17-19. Micrografías elec-
trónicasde barrido delejemplardelaFigura 16deshidratadoporpunto crítico. 17: vistageneraldela
cabezay delpie; 18: vista aumentada delhocico; nótese la ausencia de lóbulos cefálicos; 19: vista
aumentada de un haz anteriorderecho de tentáculos epipodiales. 20-23. Micrografías electrónicas de
barrido de la rádula, ejemplarde la Figura 16. 20: radula completa; 21: vista aumentada dealgunas
filascompletas;22:detalledelosdientescentralesylaterales;23:detalledelosdientesmarginales.Abre-
viaturas, ct: tentáculocefálico;et:tentáculosepipodiales;f:pie(señaladoelpropodio);sn:hocico.
58
McLeanAND GOFAS: Notes on the genusAnademaH. andA. Adams, 1854
59
Iberusy 26 (1), 2008
flattened area surrounding the mouth, projectingrimofmaturefemale shells; it
laterally with two prominent, smooth is therefore identified as a male speci-
bulges containing rather large, black men. There are immature beach-worn
mm
eyes and, next to these on the anterior shells of about 10 in diameter (Figs.
side, two slender, villose cephalic tenta- 7-9), which are considered to be female
cles; cephalic lappets lacking. Neck shells,havingthebeginningofaproject-
tobes present, rather symmetrical, undi- ing rim that will form the strong peri-
vided. Foot rather small, less than half umbilical angulation of the large female
the diametre of the shell when shell (Fig. 2). It is evident that male
extended; each side of the epipodium shells are much less frequent among the
provided in its anterior part with a beach-worn shells. Because the male
cluster of three tentacles, the foremost shell with an operculum is larger than
on hardly more than a rounded bulge, any of the comparable beach-worn
the next two tapering and villose, shells considered to be male shells, and
hardly one-tenth of the cephalic tenta- because the lip is immature, there is no
cles in size; in the posterior part with indication from the material at hand as
two more tentacles similar in size and to the possible size reached by male
shapetothelatter. shells. It may be that male shells can
Radula (Figs. 20-23) strikingly sym- reach a size similar to that of female
metrical, with broad rachidian having shells. If so, they would differ from
lateral extensions, and moderately long, female shells in having a rounded
tapered overhanging cusps; with four umbilical wall, rather than the project-
pairs of similar lateral teeth, which are ingumbilicalrimoffemaleshells.
elbowed like the rachidian and extend Both male and female shells seal the
above the shaft of the next tooth; shaft umbilicus upon attaining a half-grown
of fifth lateral tooth not in cióse contact diameter of about 10 mm. The female
with shaft of fourth lateral tooth; this shell then proceeds to form a secondary
tooth broad on both sides, with a less umbilicus for the brooding cavity. The
prominent cusp than those of the inner initial sealing ofthe umbilicus mayhelp
lateral teeth or any of the marginal toprotecttheshellfromexposuredue to
teeth; innermost pair of marginal teeth erosión of the apical whorls, which is
with short overhanging cusps, followed also avoided in most marine gastropods
by four pairs of marginal teeth with by shell deposition of a plug from
longer cusps; outer marginal teeth with within.
longershafts, overhangingcuspsshorter There is variation in the peripheral
anddeeplyserrateonsidesofthecusps. spination of immature stages. The
The live-collected specimen (Figs. 4- spination of the live-collected male
6, 16) reported here (diameter 9.1 mm) specimen (Figs. 4-6) is strongerthanthat
has anearlyclosedumbilicus and a gen- of any of the beach-worn shells of
erally unmodified base, with no indica- similar size, whether identified as male
tion of the incipient formation of the orfemale.
DISCUSSION
Knowledge of the genus Anadema single live-collected specimen reported
has been slow to develop because the here from the intertidal zone is an
speciesA. macandrewiilivesinthe sublit- unusualrecordforthespecies.
toral zone on rocky bottoms exposed to Systematic position: There is now no
strong surf, where there have appar- doubt that Anadema should be assigned
ently been few efforts at collecting by to the familyColloniidae, onthebasis of
diving due to the exposure, low visibil- shell and opercular characters, charac-
ity, and extensive muddy bottoms off- ters of the external anatomy, and the
shore in relatively shallow water. The radula. The fine lamellar sculpture had
60
McLeanAND GOFAS: Notes on the genusAnademaH. andA.Adams, 1854
brought to mind a comparison with cal, and the inner margináis are not
Liotiidae, but that is ruled out by the greatly enlarged. In turbinids, the asym-
calcareousoperculum. metrical tooth row causes the tooth
There are many possibilities for alignment to be skewed, because the
opercular morphology in the Colloni- large inner laterals must altérnate in
idae, just as there are in the Turbinidae zipperfashionwhen the radula is longi-
(see Vermeij and Williams, 2007). tudinally folded (as detailed by
There is no comparative work on the HickmanandMcLean, 1990).
colloniid operculum, but the operculum Anadema is highly unusual within
ofAnadema is within the range ofpossi- Colloniidae for its large size and its
ble expression for the family. Immature keeled early stage. With its máximum
shells of the Mediterranean turbinid shell diameter of 17 mm, it may well be
species Bolma rugosa (Linnaeus, 1767) the largest known colloniid. All of the
have a resemblance to Anadema macan- colloniid radulae illustrated by
drewii in having a somewhat similar Hickman and McLean (1990) have a
operculumandaspinoseperipheralcar- secondary flap that projects above the
ination, but mature specimens of Bolma rachidian tooth; the radula ofAnadema
are much larger, have a higher profile is unusual in not having the secondary
and an impressed suture, a closed flap. In addition, the morphology of the
umbilicus and an expansive columellar fifth lateral ofAnadema seems also to be
callus that forms a columellar shield unusualforthefamily.
nearly as broad as the aperture. For a Larval brooding: Brooding of larvae
review of Bolma see Beu and Ponder within the umbilical cavity has been
,
(1979).Anadema macandrewiidiffersfrom broadly reported among the Trochoidea
all illustrated species ofBolma in its low, (Hickman, 1992: 254). In the Trochidae
dome-shaped profile and open umbili- it is known in Margantes vorticiferus, as
cusatmaturity. shown by Lindberg and Doberteen
The keeled earlyjuvenile shells indi- (1981). In Liotiidae, it has been reported
cate that early sculpture is not evenly for Arene socorroensis by Shasky (1968)
bicarinate, which is a defining feature and again by Hertz (1998), and for
for Turbinidae, according to the "Munditia" subquadrata it has been
restricted definition provided by reported by Burn (1976). In the Colloni-
Hickman and McLean (1990: 55) at the idae, it is known in the recently
previously recognized subfamily level. described genus Liotipoma, as reported
Most colloniids have the juvenile shell by McLean and Kiel (2007). Its occur-
with even spiral cords, but there are rence inAnadema is therefore the second
exceptions to that generalization. The known example. In each of these cases,
keeled early stage brings to mind the the umbilical rim of the female shell is
recently described Liotipoma McLean raised to increase the volume of the
and Kiel, 2007, in the basal subfamily umbilical cavity. Such a modification
Petropomatinae, whichalsohas anearly can be the only explanation for the
keel. strongly raised umbilical rim of what
Thelackofcephaliclappetsisconsis- we interpret as the female shells of
tentwith the assignment to Colloniidae, Anademamacandrewii.
in which lappets are lacking (Hickman The size reached by the shells of
and McLean, 1990). This is in contrast males remains to be discovered; it is
to the Turbinidae, in which the lappets possible that mature male shells are
are well-developed (Hickman and smaller thanfemale shells. Insupportof
McLean, 1990), albeit they are small in that possibility, the operculum illus-
theMediterraneanBolmarugosa. trated here shows the expansión of the
The radular morphology ofAnadema final volution that is characteristic of a
provides convincing evidence that it is mature operculum. However, this shell
colloniid rather than turbinid because is somewhat immaturebecause the final
the tooth rows are perfectly symmetri- lip is not thickened. All beach-worn
61
Iberus, 26 (1), 2008
male shells in the material on hand thefirstauthor)werepreparedinPhoto-
seem to be immature. The immature shop by Michelle Schwengel (formerly
male shells are represented by fewer LACM) and píate preparation for the
specimens than the larger shells attrib- shells was completed by Ángel Valdés
utedto,forreasonsunknown. (formerly LACM). Critical-point dried
preparation of the live-collected speci-
men and the SEM views of the head-
ACKNOWLEDGMENTS foot, as well as the SEM illustrations of
the radula, were provided by Daniel L.
We thank Philippe Bouchet and Vir- Geiger of the Santa Barbara Museum of
ginie Heros of the MNHN for arranging Natural FFistory. We thankthereviewers
the loan of the Anadema specimens. for their suggestions, which led to
Illustrations of shells (from photos by improvementsinthepaper.
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