Taxonomic remarks on Cenozoic pseudolivid gastropods from South America

6 Pages·1997·3 MB·

THE VELIGER
© CMS, Inc., 1997
TheVeliger40(l):23-28 (January 2, 1997)
Taxonomic Remarks on Cenozoic Pseudolivid
Gastropods from South America
by
GEERAT VERMEIJ
J.
Department of Geology and Center for Population Biology, University of California at Davis,
Davis, California 95616, USA
AND
THOMAS DeVRIES
J.
Box 13061, Burton, Washington 98013, USA
Abstract. The neogastropod family Pseudolividae was represented in the Neogene ofPeru and Chile
by a single genus, which we name Testallium to replace Gastridium Sowerby, 1846, non Gastndium
Sowerby, 1842, or Gastridium Modeer, 1793. Besidesthetype species, Testalliumcepa (Sowerby, 1846),
from the Miocene of Peru and Chile, we recognize T. valuta (Olsson, 1932) from the early Miocene of
northern Peru, and T. escalonia, sp. nov. from the PlioceneofChile. Gastridium retusum Philippi, 1887,
is assigned to Buccinorbis Conrad, 1865.
INTRODUCTION BMNH,
Institutional abbreviationsareasfollows: Brit-
ish Museum of Natural History, London, England; CAS,
Thefamily Pseudolividae is apoorly known groupofLate CaliforniaAcademyofSciences,SanFrancisco,California,
Cretaceousto Recent gastropod mollusks that, on the basis USA; PRI, Paleontological Research Institute, Ithaca, New
of shell characters as well as internal anatomy, belongs to York, USA; SGO PI, Museo Nacional de Chile, Institute
theneogastropodsuperfamily Buccinoidea (see Cossmann, de Palontologia, Santiago, Chile; UCMP, Museum ofPa-
1901; Squires, 1989; Kantor, 1991). Although the bathyal
leontology, University of California, Berkeley, California,
genus Benthobia Dall, 1889, is widely distributed in the USA; YPM, Yale Peabody Museum, New Haven, Con-
AtlanticandsouthwesternPacificOceans, Recentshallow- necticut, USA.
water members of the Pseudolividae are restricted to sub-
tropicalandwarm-temperatewatersinwesternandsouth-
ern Africa, the Australian region, and the west coast of SYSTEMATIC PALEONTOLOGY
North America. During the Miocene and Pliocene, how-
ever, the family was also represented on the west coast of Class Gastropoda
South America by an extinct genus that has been referred Order Neogastropoda
to as Gastridium Sowerby, 1846. Unfortunately, Gastri-
BUCCINOIDEA
dium was preoccupied by Gastridium Sowerby, 1842, and Superfamily
Modeer, 1793. The distinctive South American Neogene
Family Pseudolividae Cossmann, 1901
pseudolivids therefore require a new generic name. In this
paper, we (1) propose the genus Testallium for three Discussion: Kantor (1991) has made cogent arguments
Miocene and Pliocene pseudolivid species from Peru and for recognizing the Pseudolividae as a family distinct from
Chile;(2)compare Testalliumtootherpseudolividgenera; such other families as the Buccinidae and Olividae with
(3) review the described species assigned or compared to which it had previously been associated as a subfamily. In
Gastridium Sowerby, 1846, by previous authors; and (4) fact, anatomical peculiarities of the group prompted him
describeTestallium.escalonia,sp.nov.fromstrataofprob- to isolate the Pseudolividae in its own suborder Pseudo-
able Pliocene age in Chile. livoidei. This taxon was characterized by the presence of
Page 24 The Veliger, Vol. 40, No. 1
a gland and valve ofLeiblein, and by having the proboscis "Buccini" rather than to the "Olivi." In fact, Sowerby
formed by the elongation of the buccal tube rather than assigned B. plumbeum to the buccinid genus Eburna La-
by the elongation of the dorsal wall of the buccal cavity marck, 1822 (non Lamarck, 1801), whose oldest valid
as in the suborder Muricoidei (Kantor, 1991). The most nameisBabylonia Schliiter, 1838 (seevan Regteren Altena
obvious shell character that distinguishes Pseudolividae & Gittenberger, 1981). Philippi also disliked the name
from the Buccinidae {sensu lato) is the presence of a basal Pseudolwa, and applied Gastridium Sowerby, 1846, to G.
external spiral groove, which ends in a more or less prom- cepa and two other species, G. opimum (Hupe, 1854) and
inent tooth or spine at the edge ofthe outer lip (Cossman, G. retusum (Philippi, 1887). The objective synonymy of
1901). Other shell features that collectively set the Pseud- Gastridium Sowerby, 1842,andPseudolwaSwainson, 1840,
olividae apart from other neogastropod groups include the was recognized by most subsequent authors, including
following: notchorsinusatposteriorendofouterlip;outer Sowerby (1859), Cossmann (1901), Melvill (1903), and
lip lying in a plane; spiral sculpture consisting ofcords or Wenz (1938-1944). Moreover, the name Gastridium, as
threads that are strongest near base and weakest near used by Sowerby (1842, 1846), was preoccupied by Gas-
upperpartofwhorl;aperturenarrowlyelongate;columella tridium Modeer, 1793 (p. 106), a well-known Indo-Pacific
moreorlessconcaveonupperpart, sm^ooth; siphonal canal Recent member of the toxoglossan gastropod family Con-
short to almost absent. Genera are distinguished on the idae. Gray's (1847, p. 136) introduction of Gastndia re-
presenceor absence ofapertural lirae (spiral riblets on the solved the homonymy of Gastridium, but his name falls as
inner side of the outer lip), axial sculpture, an umbilicus, an objective synonym of Pseudolwa because it was based
and a parietal rib, as well as on shell shape. A general on the same type species, Buccinum plumbeum. Sowerby
review of the family by Vermeij is in progress. (1846), Philippi (1887), and Olsson (1932) all regarded
Gastridium inSowerby's (1846)senseasdistinctfromPseu-
Genus Testallium Vermeij & DeVries, gen. nov. dolwa, but none of these authors offered distinguishing
characters for the two groups. We agree that Sowerby's
Gastridium Sowerby, 1846: p. 261; non Gastridium Sowerby, (1846) Gastridium refers to a distinct group, and propose
c1i8e4s,2:Bup.cc3i1n2um(=plPusemubdeoulmwaCShweami.n,so=n,Bu1c8c4i0n)u(mtycpreassspuem- the new genus Testallium for it.
Gmelin, 1791); non Gastridium Modeer, 1793: p. 106 Testallium differs from Pseudoliva in shape, sculpture,
(type species: Conus geographus Linnaeus, 1758; Gas- and the development of callus. Whereas Testallium, is
tropoda: Toxoglossa: Conidae). constricted (that is, distinctly concave in profile)just above
Type species: Gastridium cepa Sowerby, 1846 (pi. 1, figs, ttoheablamsoestofstthreaisgihpth-osniadledc.anTalh,ePsfeauscdioollweaoifs wTeeasktlayllciounmveixs
a, b)
prominent, marked by a keel-like posterior angulation.
Diagnosis: Pseudolivid characterized by deep basal exter- That of Pseudolwa is indistinct and rounded. Testallium.
nal groove terminating at outer lip in blunt labral spine; hastwotoeightspiral cords below the basal groove, which
last whorl basally constricted above siphonal fasciole; fas- conspicuously crenulate the edge ofthe outer lip; and nu-
cicle adapically keeled; sculpture consisting of fine spiral merous spiral threads between the groove and the suture.
threads over whole surface of shell above groove, and of Earlier whorls carry fewer, somewhat stronger cords on
two to eight strong spiral cords below groove; outer lip the upper portion of the whorl. Pseudolwa's shell, by con-
forming notch at adapical end of aperture; inner side of trast,isessentially smooth except forthe basal groove. The
outer lip smooth; columellar callus of small extent; um- columellarcallusofTestallium.islimitedinextent,where-
bilical slit narrow or absent; parietal rib at adapical end as that in Pseudolwa is broad, thick, and heavy, obscuring
of inner lip distinct. the basal groove on the ventral half of the last whorl.
Etymology: testa (Latin: shell) and allium (Latin: onion), AnothersimilargenusisBuccinorbis Conrad, 1865, spe-
referring to the specific name cepa (Latin: onion) of the cies of which are known from the late Paleocene (Thane-
tian) to the late Eocene (Priabonian). Like the Miocene
type species.
to Recent West and South African Pseudolwa, it has ex-
Discussion: Sowerby (1842, p. 312) named thegenus Gas- tensive columellar callus, and the base is not constricted;
tridium to include Buccinum plumbeum Chem. (= Buccin- but thegroove is located relatively higheronthelast whorl
umcrassum Gmelin, 1791). In 1846 (p. 261),heintroduced (see also Squires, 1989). Three to five strong spiral cords
the name Gastridium again as a new genus, this time for below the groove form well-defined crenulations on the
Gastridium cepa Sowerby, 1846, a Miocene species col- edgeoftheouterlipbelowthelabralspine(seealsoPalmer,
lected by Charles Darwin in 1833 at Navidad on the 1937). Buccinorbis differs from Testallium by the absence
central coastofChile. In his 1846 paper, Sowerby (p. 261) of basal constriction, the large, heavy columellar callus,
noted that G. cepa was similar to Buccinum plumbeum, and the absence ofa parietal rib at the adapical end ofthe
and that Swainson (1840, p. 82, p. 306) had made the inner lip.
latter species the type o[Pseudolwa Swainson, 1840. Sow- The western North American early Miocene to Recent
erby (1846) considered the name Pseudolwa "absurd" be- genus Macron H. & A. Adams, 1853, differs from Tes-
cause B. plumbeum in his mind clearly belonged to the tallium. chiefly by possessing lirae on the inner side ofthe
G. Vermeij & T. DeVries, 1997 Page 25
J. J.
outerlip. Thetypespecies,BuccinumaethiopsReeve, 1847,
isoftenextremely prominently sculptured with two heavy,
rounded cords below the groove and up to six spiral cords
abovethegroove. Otherspecies, however, such as theearly
Miocene M. hartmanni Hertlein & Jordan, 1827 and the
Pleistocene to Recent M. lividus A. Adams, 1855, and M.
orcuttiDall, 1918, havefinespiralthreadsonthelastwhorl
between the groove and the suture, and in this respect
strongly resemble species of Testallium.
Testallium is strongly convergent on the Miocene to
Recentwestern SouthAmericanocenebrinemuricidgenus
Chorus Gray, 1847. Both genera have a broadly fusiform,
basally constricted shell with a basal spiral groove ending
in a labral spine at the edge ofthe outer lip. Chorus differs
from Testallium by the absence ofa posterior sinus in the
outer lip, by lacking a parietal rib, and by having an
indistinct, rounded siphonal fasciole rather than a mark-
Figure
edly angulated one. The basal groove is also shallowerand 1
less conspicuous in Chorus than in Testallium. Moreover, Testallium cepa. Holotype (BMNH G 26399); a, b. lateral and
Chorushasspiralsculptureconsistingofsixtosevenwidely dorsal views. The shell is 5.5 cm in height.
separated primary spiral cords on the last whorl, which
do not increase in prominence basally. blainvillei (d'Orbigny, 1842) from strata inferred to have
been of early Pliocene age at Chepu, on Isla Chiloe, in
southern Chile. They favored a Pliocene age in part be-
Testallium cepa (Sowerby, 1846)
cause Chorus and several other genera in the fauna were
(Figure la, b) not known from the Miocene at that time. The presence
of distinctive Miocene fossils persuades us, however, that
MGaosntorciedriousmlcaebpiaaleSoHwuepreb,y,1815844:6,ppp.. 216919,-2p0i0..4, figs 68, 69. the Chepu beds are of late early Miocene age, and that
Monoceros opimum Hupe, 1854, p. 200. they are correlated with the similarly aged strata of the
Fusus labialis Hupe, 1854, pi. 3, fig. 3a. Navidad Formation, from which T. cepa was originally
Fusus opimus Hupe, 1854, pi. 2, figs. 6, 6a. collected (DeVries & Vermeij, in preparation). Fleming
Gastridium cepa Sowerby, 1846: Philippi, 1887, pp 59-60, noted that the spiral sculpture of his specimen was weak,
pi. 6, fig. 2. that the spire was higher, and that the shell was narrower
Monoceros labialis Hupe, 1854; Philippi, 1887, pi. 5 fig. 1; than that of Chorus blainvillei. Moreover, his specimen
not pi. 5, fig. 6 (indeterminate species).
Gastridium opimum (Hupe, 1854): Philippi, 1887, p. 60, pi. lacked the nodes characteristic of C. blainvillei, a Pliocene
57, fig. 7. species. Fleming'sspecimenrepresentsamoderatelyshoul-
Chorusaff.Cblainvillei(d'Orbigny, 1842):Walters&Flem- dered form of Testallium. cepa.
ing, 1972: p. 398, pi. 28, fig. 6u; non Chorus blainvillei Two probably unrelated gastropods are strongly con-
(d'Orbigny, 1842) (Muricidae). vergentinform,aswell asintheexpression andvariability
Gastridum cepa Sowerby, 1846: Tavera Jerez, 1979. p. 97, of the shoulder, to Testallium. cepa. The Recent Panamic
pi. 20, figs. 74, 75.
rapaninemuricid Vasulamelones (Duclos, 1832)oftenforms
Discussion: Testallium cepa is a highly variable species, akeel-likeridgeattheshoulderinitsfinalstagesofgrowth,
a fact that accounts for its long synonymy. The most vari- whereas younger shells are weakly shouldered or rounded
ablecharacteristheshoulder. Theposterior(oranal)notch on the upper part of the last whorl. Every gradation be-
is strongly produced in the type specimen of T. cepa, and tween strongly shoulderedand roundedshellscan befound
the shoulder is correspondingly well developed. Hupe's within populations throughout the range of the species.
(1854) Monoceros opimum (labeled Fusus on his plate, and The "buccinid" Triumphisdistorta (Wood, 1828), also from
assigned to Gastridium by Philippi, 1887) represents an thePanamicProvince,alsoshowsakeeled,ridgelikeshoul-
unshouldered shell in which the posterior notch is not der in the adult stage and a rounded profile of the upper
produced. Monoceros labiale Hupe, 1854 (labeled Fususon part ofthe body whorl in younger shells. It is striking that
Hupe'splate,andassignedtoMonocerosbyPhilippi, 1887) both Vasula melones and Triumphis distorta occur in rocky
is intermediate between the cepa and opimum phenotypes. areas with intermixed sand, a habitat similar to that of
Philippi's (1887) illustration ofGastridium opimum (Hupe, the pseudolividsPseudolivacrassa (Gmelin, 1791) andLui-
1854) (pi. 5,fig.7)isindistinguishablefromhisillustration ziazebrina (A. Adams, 1855) inAngola(S. Gofas,personal
of Monoceros labialis. communication to G. Vermeij, July, 1995).
Fleming (inWatters & Fleming, 1972) figured a poorly TestalliumoccupiedasimilarhabitatinPeruandChile.
preserved specimen of Testallium cepa as Chorus aff. C. In Chile, T. cepa is found in pebbly sandstones ofprobable
Page 26 The Veliger, Vol. 40, No. 1
early Miocene age (DeVries & Vermeij, in preparation) Testallium.. Apertural features are obscured by matrix.
near basement rocks at Punta Ahuenco on the east coast Nelson (1870, p. 199) suggested that Clavella solida is
of Chiloe (Watters & Fleming, 1972); in massive sand- relatedto Tnumphisdistorta (Wood, 1828). We tentatively
stones at Islas Ipun and Stokes, south of Chiloe (DeVries assign it to the genus Nicema Woodring, 1964, which is
et al., 1984); and in fine to medium-grained sandstone in usuallyincluded with Tnumphis Gray, 1857, inthefamily
the Navidad basin (Tavera Jerez, 1979), sometimes in Buccinidae (see Woodring, 1964).
stratawith intercalations and lenses ofgravel. Inthe Pisco SpecimensofT. voluta fromnothern Peruarefilledwith
Basinofsouthern Peru,specimensofTestalliumarefound acoarse-grainedbioclasticmatrix. Olsson (1932) proposed
in coarse-grained sandstones at numerous localities. At with some uncertainty that they were collected at the base
Callejon de Cerro de Piedra, near Nazca (Rivera, 1957), ofshales fromthe Heath Formation, overlying Punta Bra-
specimens of T. cepa occur with numerous specimens of vo "grits." Olsson (1932) believed the Heath Formation
Turritella woodsi Lisson, 1925, in probably early Miocene to be of Oligocene age, but subsequent work by Zuiiiga
conglomeratesandcoarse-grainedsandstonesdirectlyover- & Cruzado (1979) indicates its age to be early Miocene.
lying basement crystalline rocks. In the Lomas Chilcatay,
east of Bahia de la Independencia, specimens of T. cepa
arefoundtogetherwith adiverse assemblageofearlyearly Testallium escalonia Vermeij & DeVries, sp. nov.
Miocene mollusks in basal sandstones of the Chilcatay
(Figure 2a, b)
Formation and in bioclastic, balanid-rich sandstones high-
er in the same section that have been dated as late early Diagnosis: Testallium with a relatively high spire (spire
Miocene (Dunbar et al., 1990). one-fourth to one-third shell height), 17 to 23 fine spiral
Distibution: Early Miocene {Rocella gelida Zone to Tri- threads on last whorl, and narrow umbilical slit.
ceratiumpileus Zone, about 24-18 Ma; H. Schrader, writ- Description: Shell relatively high-spired, consistingoffive
tencommunication, 1987),southernPerutosouthern Chile. teleoconchwhorlsseparatedbyshallow,appressedsutures;
Testallium valuta (Olsson, 1932) spire one-fourth to one-third total shell height; sculpture
consisting of two cords below basal groove, and 17 to 23
Acanthiza {Chorus) valuta Olsson, 1932: pp. 184-185, pi. 19, fine spiral threads between the groove and the suture on
figs. 3, 6, 7. the last whorl, increasing in strength toward the base; last
Discussion: Acanthiza Fischer is probably a misspelling quarter ofbody whorl marked with closely spaced growth
of Acanthma Fischer de Waldheim, 1807, another genus lines; groove terminates in small blunt labral tooth at edge
with a labral spine from western South America. Chorus ofouterlip; fasciole prominent,sharplyangledposteriorly,
Gray, 1847, is yet another genus with a labral spine from rounded below; narrow umbilical slit on inner side, edge
the same region. Olsson (1932) compares T. valuta with faintlycrenulated inaccordancewithexternalspiralsculp-
Monoceros laevis Philippi, 1887 (= Chorus laevis). Speci- ture; columella bearing five or six low oblique ridges on
mens of T. voluta are distinguished from those of both lower half; weak anal notch where outer lipjoins penul-
Acanthina and Chorus by their strong axial folds on the timate whorl posteriorly.
earliest whorls and anal notch. Holotype: CAS 66806.01, height 34.0 mm, diameter 24.2
Differences between T. voluta and T. cepa are more
mm, aperture height 25.4 mm.
subtle. Specimens of both species show strong spiral cords
on the earliest whorls, but only T. voluta shows any axial Paratype 1: CAS 66806.02, height, 28.9 mm, diameter
sculpture. What most distinguish specimens of T. voluta 19.7 mm, aperture height 21.4 mm.
alreengtah)shaonrdt,siinphaodnualltscpaencailme(nwsel,lauwnedaekrthoalmfodtehreaatpeerstuulrcauls P15a.r8atmymp,e a2p:erUtuCrMe Pheig3h9t88200,.5hemimg.ht 25.4 mm, diameter
on the body whorl that lies anterior to a moderately an-
gulate,tabulateshoulder. ShoulderedspecimensofT. cepa, Type locality: El Ganso (ganso = goose), 34°13'S, west
in contrast, have sloping shoulders, no sulci so low on the of Fundo Las Damas, also known as the La Cueva Lo-
body whorls, and moderately produced siphonal canals cality. La Cueva Formation ofprobable late Pliocene age,
(equal to half the apertural length). Chile.
T. volutawasconsideredbyOlsson (1932)tobeancestral Etymology: Latin noun for small onion.
to Clavella solida Nelson, 1870, from the late Miocene
Cardalitos Beds ofnorthern Peru (Olsson's 1932 figure of Discussion: Testallium, escalonia differs from the older
T. voluta is excellent, and the reader is referred to it for T. cepa chiefly in having a higher spire (spire one-fourth
illustration). Olsson (1932) assigned Nelson's species to to one-third as compared to one-seventh of shell height as
what he called Acanthiza (Chorus). Our examination of in T. cepa) and in being less constricted at the base of the
type material of Clavella solida (Yale Peabody Museum siphonal canal, which is relatively shorter. Herm (1969)
YPM
00507) reveals that C. solida has strong axial ribs did not record this or any other species ofGastridium (our
on early teleoconch whorls, as does Testallium voluta, but Testallium) from the Pliocene of Chile. Oddly enough,
it lacks the basal groove of T. voluta and other species of the holotype and Paratype 1 were collected by Herm. He
&
G. Vermeij T. DeVries, 1997 Page 27
J. J.
Figure 2
Testallium escalonia. Vermeij & De Vries, sp. nov. a, b. holotype (CAS 66806.01), ventral and dorsal views; c,
d. paratype 1 (CAS 66806.02), ventral and dorsal views. The holotype (a, b) is 4 cm in height; the paratype (c, d)
is 3.5 cm in height.
mayhavemistakenthemforwornspecimensofthemuricid 1922, from the early Eocene of northern Peru, is a sub-
genusChorus,whichsuperficially resemblesspeciesofTes- jectivesynonymof5. retusa. Wehave been unabletolocate
tallium. Woods's holotype to confirm this suspicion. According to
The precise age of the La Cueva fauna is not entirely Olsson (1928), B. pannasensis occurs in the Salina, Ne-
clear, but it appears to be Pliocene (Herm, 1969). gritos, and Parinas Formations of early to middle Eocene
Testallium escalonia is the youngest known species of age. The extent of callus formation is variable in this
itsgenus. Along with many other genera and species, Tes- species, as is the strength and extent ofspiral sculpture on
tallium evidently became extinct near the end ofthe Plio- the last whorl between the spiral groove and the suture.
cene in western South America. The specimen described from the middle Eocene of Co-
lombia as Pseudoliva {Buccinorbis) cf. P. (B.) pannasensis
Genus Buccinorbis Conrad, 1865 by Clark & Durham (1946) differs from B. retusum and
Type species. Buccinum vetustum, Conrad, 1833 Woods's P. pannasensis from Peru by possessing an um-
bilicus. This feature may well have been variable in B.
Buccinorbis retusa (Philippi, 1887) retusa, but we prefer to keep Clark & Durham's (1946)
Co\omhianpannasensisformoutsidethelimitsof5. retusa.
Gastridium retusum Philippi, 1887. p. 59, pi. 6, fig. 3, 3b. Olsson's (1928) variety samanica from the late Eocene
Pseudoliua pannasensis Woods, 1922, p. 93-94, pi 12, figs. TalaraandSamanbeds(=Verdun Formation)ofnorthern
4-6.
Peru ismore heavily callused, larger, and less rotund than
Pseudolivapannasensis Woods: Olsson, 1928, p. 123. isB. retusa, and has a narrower anterior end. Forms ques-
Discussion: Philippi (1887) recognized three species of tionably assigned to this variety were described from the
Gastridium: G. cepa Sowerby, G. opimum (Hupe), and G. late Eocene of Colombia (Clark & Durham, 1946) and
retusum,Philippi, 1887. Ofthese,thefirsttwoarereferable Curasao(Jung, 1974). Olsson's(1928)varietymancorensis
toasinglespeciesofTestallium(seeabove),butG.retusum hasa laterally flattened ratherthan an evenly rounded last
is distinct. A cast of Philippi's holotype (SGO PI 765), as whorl and a massive callus in the columellar and parietal
well as Philippi's figures, shows that the species belongs regions of the shell. The final stages of growth are char-
to Buccinorbis Conrad, 1865. Like other species of Buc- acterized by adapical migration ofthe outer lip and by the
cinorbis, B. retusa has the spiral groove at a relatively high increasing prominence of the shoulder. This variety was
position on the last whorl, and lacks basal constriction. described from the Chira, Mancora, and Heath Forma-
There is a strongly developed columellar and parietal cal- tions, all regarded as Oligocene by Olsson (1928). The
lus.Noneoftheknownspecimenshastheapertureexposed Mancora and Chira Formations, however, are of latest
or the outer lip intact. Philippi (1887) thought B. retusa Eoceneage(Zuniga & Cruzado, 1979),andtheoccurrence
to be of Cretaceous age, but strata at the type locality ofthe mancorensis form in the Heath Formation, which is
(Algarrobo, Chile) are Eocene (Tavera Jerez, 1979). early Miocene, is questionable. We tentatively retain the
Judgingfromthefigures,PseudolivapannasensisWoods, varieties samanica and mancorensis of Olsson, 1928, as
Page 28 The Veliger, Vol. 40, No. 1
stratigraphic subspecies oiB. retusa, but further work and Nelson, E. T. 1870. On the moUuscan fauna of the later
additional material may reveal that all are members of a Tertiary ofPeru. Transactions ofthe Connecticut Academy
single, long-ranging species in which whorl profile, callus ofArts and Sciences 2:186-206.
development, late-stage shouldering, expression of spiral Olsson,a.a. 1928. ContributionstotheTertiarypaleontology
ofnorthernPeru: Part 1,EoceneMolluscaandBrachiopoda.
sculpture, and size are all highly variable traits. Pseudoliva Bulletins of American Paleontology 14 (52):47-154.
(Buccinorbis) vientoensis Clark & Durham, 1946, fromthe Olsson,A.A. 1932. ContributionstotheTertiarypaleontology
Eocene of Colombia, may also prove to part of this vari- of northern Peru: Part 5, the Peruvian Miocene. Bulletins
ation. ofAmerican Paleontology 19 (68):l-272.
Palmer, K. W. V. 1937. The Claibornian Scaphopoda, Gas-
ACKNOWLEDGMENTS tropodaanddibranchiateCephalopodaofthesouthernUnit-
ed States. Bulletins ofAmerican Paleontology 7:548 pp.
We thank Gary Rosenberg of the Philadelphia Academy Philippl R. a. 1887, Die Tertiaren und Quartaren Verstei-
of Natural Sciences for his help in locating important nerungen Chiles. F. A. Brockhaus: Leipzig: 266 pp.
literature. We also thank Paul Jeffery of the Natural RegterenAltena,C.O.van&E.Gittenberger. 1981. The
genus Babylonia (Prosobranchia, Buccinidae). Zoologische
History Museum in London for photographing the ho- Verhandelingen 188:1-57.
lotype of Testallium cepa, and Mary Graziose at UC Rivera, R. 1957. Moluscos fosiles de la Formacion Paracas,
Davis for additional photographic work. Janice Cooper departamento de lea. Boletin de la Sociedad Geologica del
provided valuable technical assistance. The research was Peru 32:165-220.
funded by a grant (NSF EAR-94-05537) from the Na- SOWERBY, G. B. 1842. A Conchological Manual. 2d ed., Con-
siderably Enlarged and Improved. H. G. Bohn: London.
tional Science Foundation to Vermeij.
SowERBY, G. B. 1846. Descriptions of Tertiary fossil shells
LITERATURE CITED fromSouthAmerica. Pp. 249-264inC. Darwin,Geological
Observations on South America, Being the Third Part of
Clark. B. L. & J. W. Durham. 1946. Eocene faunas from tmhaenGdeoolfogCyaptof.tFhietzVrooyy,agRe.No.f,thDeuBreianggle,theUnYdeearrsth1e8C32omt-o
the Department ofBolivar, Colombia. Geological Society of
1836. Smith, Elder, and Company: London.
.America Memoir 16:1-126.
COSSMANN, M. 1901. Essaisdepaleoconchologiecomparee. M. SoWERBY, G. B. 1859. Monograph of the genus Pseudoliva,
Swains. Thesaurus Conchyliorum 3:73-75.
De \Croisessm,aTn.n:J.P,arLi.s.M2.93Stpop.tt & VV. J. Zinsmeister. 1984. Squirtehse,LRo.weL.r T1e9r8t9i.arAy onfewNoprstehudAomlievriincea.gaJsoturronpoadlgoefnuPsalferoonm-
Neogeneforearcbasinsinsouthern Chile. AntarcticJournal
tology 63:38-47.
Dunboafrt,heR.UnBi.t,eRd.SCtatMesar1t9:y12&-13P.. .\. Baker. 1990. Cenozoic SwaiCnlsaosnsi,fiWc.atio1n840o.f SAheTlrlesataisnedonShMelallfaicsho.logLyo;ngormatnhe,NaOtrumrea,l
mPaalraienoegseeodgirmaepnhtya,tiPoanleionctlhiemaSteoclhougry,aaPnadlaPeioseccoolBoagsiyns7,7:P2e3r5u-. Brown, Green, and Longmans: London: 419 pp.
261. Tavera Jerez, J. 1979. Estratigrafia y paleontologia de la
formacion Navidad, Provincia de Colchagua, Chile (Lat
Gray, J. E. 1847. .\ list of genera of Recent Mollusca, their 30°50'-34°S). Boletin del Museo Nacional de Historia Nat-
synonyms and types. Proceedings of the Zoological Society
ural 36(5-10): 5-176, and 21 plates.
Hermof,LDo.nd1o9n691.5:M1a29r-i2n1e9s.Pliozan und Pleistozaninnord- und Watters, W. a. & C. A. Fleming. 1972. Contributions to
thegeologyandpaleontologyofChiloeIsland,southernChile.
mittel-Chile unter besonderer Beriicksichtigung der En-
Philosophical Transactions of the Royal Society of London
twicklung der MoUusken-Faunen. Zitteliana 2:1-159.
(B) 263:369-408.
HuPEoH,giisHcta.o)r.ic1Ma8a5uF4i.lsdiecMaaelytacRPooellniootguiic:aaPdyaercioCsnh.qiul4ie4l.i9oVlopolpg..ia8.ainndC.atGlaasy(Z(eodo.l)-, Wenzo,brWan.chi1a9.38H-a4n4d.buGcahstdreorpoPdaala.oTzeoiolloI:giAeUg6.emBoerinnterraeugnerd:PBreors--
Jung, P. 1974. Eocene mollusks from Curasao, West Indies. Woodlrini.ng, W. p. 1964. Geology and paleontology of Canal
VerhandlungenderNaturforschendenGesellschaftBasel84:
ZoneandadjoiningpartsofPanama: descriptionofTertiary
483-500.
Kantor, Y. I. 1991. On the morphology and relationships of Gmeoolllougsikcsal(GSausrtvreoypodPrso:feCsosliuonmablelPlaipdeare 3t0o6V-oCl:u2t4id1a-e2)9.7.U. S.
some oliviform gastropods. Ruthenica 1:17-52. Woods, H. 1922. MolluscafromEoceneandMiocenedeposits
LiSSON, C. I. 1925. Algunos fosiles de Peru. Boletin de la of Peru. Pp. 51-113 in T. O. Bosworth (ed.). Geology of
Sociedad Geologica del Peru 1:23-30.
Melvill,J.C. 1903. ThegeneraPseudolivaandMacron.Jour- otfhePeTreur.tiMaraycmainldlaQnuaatnedrnCaormypaPenryi:odsLoinndotnhe.Northern Part
Modenealr,ofaC.onc1h79o3l.ogyIn1l0e:d3n2i0n-g3t3i0l.kunskapen om maskkraken i ZuSlPGeAr,uaF.no&.JB.olCertiunzadedolaCS.oci1e9d7a9d. GBeioolsotgriactiagrdaelfiPaedreuln6o0r:o2e1s9t-e
allmamhet.Kongl.vetenskapsAcademiensnyahandligar 14:
232.
83-112.