Genera of American strombid gastropods (Gastropoda : Strombidae) and remarks on their phylogeny

2007·6 MB·English

by  Gijs C Kronenberg

THE VELIGER
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TheVeliger49(4):256-264(December 11, 2007)
Genera of American Strombid Gastropods (Gastropoda: Strombidae) and
Remarks on Their Pnylogeny
KRONENBERG
GIJS C.
Milieu Educatie Centrum, P.O. Box 435, NL-5600 AK Eindhoven, The Netherlands
(e-mail: [email protected])
HARRY G. LEE
4132 Ortega Forest Dr., Jacksonville, FL 32210, USA
(e-mail; [email protected])
Abstract. Persististrombus gen. nov. is created to accommodate a number offossil and one Recent species which
formadistinct lineage startingin EarlyOligocene ofEurope with Strombusradix Brongniart, 1823, via the Oligocene
to early Miocene S. bonelliisensu stricto Brongniart, 1823 and a number ofCaribbean extinct species to the Recent
Panamic faunal province Strombus- grauulatiis. The genus level name Lobatus Iredale, 1921 (type species
bitiiberciilatiis Lamark, 1822) is available. Possible relationships between Persististrombus gen. nov. and other
Caribbean and Panamic Strombidae are discussed but remain uncertain as the phylogeny ofthese species is not fully
elucidated. Putative evolutionary scenarios are briefly considered.
Gastropoda, Strombidae, new genus, Persististrombus, Lobatus, phylogeny
INTRODUCTION Pacific Lentigo lentiginosusand L. pipus. More recently
The discovery ofaberrant specimens ofthe well known it has been demonstrated, based on anatomical
characters (Simone, 2004) and molecular sequencedata
Strombus grcmulatus Swainson, 1821 from the Islas
(Latiolais, 2003 and Latiolais et al., 2006), that the
&GalLaepeago(s20a04n)d IcsolambdeilneCdocwoitdhiscaunsseedarblyierKrpoanpeenrberbyg genus Strombus sensu Abbott is not monophyletic.
Lozouet & Maestrati (1986) compelled the authors to Sacco (1893:12) was the first to recognize a hneage
further investigate the relationships ofthis species and fBrroonmgntihaerfto,ss1il82S3.,rfaodrixwhBircohngSntiraortm,bu1s82n3odaonsdusS.(Bboornselolni,i
the fossil record ofits relatives. 1820) might be an earlier name, see Sacco (1893:4) of
Strombus granulatus was assigned to Lentigo Jous- the Recent S. granulatus, an opinion followed by
seaume, 1886 by Abbott (1960), and this allocation was Lozouet and Maestrati (1986)]. This was acknowledged
followed by subsequent workers like Walls (1980), by Jung & Heitz (2001), who described a number of
Kronenberg and Berkhout (1984), and DeTurck et al. fossil species, allocating those to Lentigo. They also
(1999). included the Recent Strombus raninus Gmelin, 1791,
As indicated before (Kronenberg & Vermeij, 2002), a species previously assigned to Tricornis Jousseaume,
the lines between subgenera as recognized by Abbott 1886 or, more recently, to Lobatus Iredale, 1921
(1960) are rather blurred and arbitrary. Lentigo, to (Petuch, 1994).
which Abbott (1960) assigned tlve species, viz. Strom- Consistent with reasoning ofKronenberg & Vermeij
bus lentiginosus Linnaeus, 1758 (type species TS); S. (2002) and data presented by Latiolais (2003), Simone
pipus (Roding, 1798); 5. fasciatus Bom, 1778; S. latus (2004), and Latiolais etal. (2006), and to accommodate
Gmelin, 1791; and S. granulatus, is an example ofthis the lineage of the Early Oligocene species of Europe
problem. On the basis of shell characters there are at and a number of fossil species described by Jung &
least three supraspecific taxa included in this group. Heitz (2001) to the Recent S. granulatus, a newgenus is
Moolenbeek& Dekker(1993) have already allocated S. described herein. As Jung & Heitz (2001) argued that
fasciatus to Conomurex Fischer, 1884 based on shell the name Lobatus Iredale, 1921 is unavailable, we
morphology and characters ofthe radula. Subsequent- discuss the nomenclatorial status of that taxon. A
ly, DeTurck et al. (1999) replaced it in Lentigo without review of the literature for possible relationships
comment. Kronenberg & Vermeij (2002) indicated that between the new genus described herein and other
both Strombus granulatus and S. latus differed in strombid genera revealed that there is a number of
a number of conchological characters from the Indo- possible relationships between the Recent Western
G. C. Kronenberg & H. G. Lee, 2006 Page 257
Atlantic and Panamic Province fauna and fossil species
known from the Early Oligocene to Early Miocene of
Europe. These are discussed briefly.
Strombusalbirupianus Dall, 1890, described from the
Late Eocene (Jackson) white limestone overlying the
Claiborne sands, Claiborne Bluff, Alabama, does not
appear to be closely related to this lineagejudging from
the description and figures by Dall (1890:174-175, pi.
12 figs. 2, 10) and is not considered here.
Likewise, two other American fossil strombid
species, S. liocyclus Dall, 1915 from the Miocene
Tampa silex beds. Ballast Point, Tampa Bay (Florida,
USA) \fide Boss et al., 1968] and 5. leurus Woodring, Figure 1. Persististrombus aldriclu (Dall, 1890). U.S.A.,
1928 from the Pliocene Bowden Formation ofJamaica Florida, Calhoun Co. Chipola Riverjust above Farley Creek
Chipola Formation, Early Miocene. Leg. C. Hertweck,
arenotdiscussed here as they apparently left no Recent collection H. G. Lee. Actual size47.9 mm. A. Apertural view,
descendants in the Americas. Strombus liocyclus was B. Dorsal view. Photos H. G. Lee.
allocated to Canarium Schumacher, 1817 by Abbott
(1960:63). Strombus leurus was not discussed by
Derivation: Derived from the Latin persisteiis (persis-
Abbott, but the species bears a strong resemblance to
tent) combined with Strombus, as the general shell
species allocated to Dolomena Wenz, 1940 [Dolomena
Iredale, 1931 is not available; see Kronenberg & morphology of species assigned to this genus has
remained almost unchanged from the Early Oligocene
Dharma, 2005 and references therein] (Woodring, (Lozouet & Maestrati, 1986) to Recent. The late
1928:326-327; pi. 24. figs. 3-5).
Eocene record by Lozouet and Maestrati (1986) for
Another possible clade. consisting of the genus
Orthaulax Gabb, 1873 (TS O. mornatus Gabb, 1873), 5. radix may be erroneous (personal communication
known from the lowermost middle Eocene ofItaly (O. Lozouet to GCK, January 2007).
dainelliiSavazzi, 1989)with a number ofOligocene and Other species assigned to Persististrombus gen.nov.
Miocene species in America (see Yokes & Yokes, 1968 are: Strombus aldriclu Dall, 1890 from the early
for a review and discussion) is not discussed herein as Miocene of the Chipola Beds, Florida, U.S.A. (Fig-
we think that Orthaulax is not closely related to the ure 1); S. baltrae Garcia-Talavera, 1993 from the PHo-
Recent American species. Pleistocene of Isla Baltra, Islas Galapagos; S. barrigo-
Further, a number of fossil strombid species are nensis Jung & Heitz, 2001 from the Cubagua Forma-
known from South America, dating as far back as the tion, early Pliocene of Yenezuela; S. bonellii Brong-
Eocene. These have been allocated to various (sub)- niart, 1823 from the Early Miocene of France
genera, like Oostrombus Sacco, 1893 (type species (Figures 2, 3) [this may be a junior synonym of S.
Strombus problematicus Michelotti, 1861) but are nodosus Borson, \?>20,fide Sacco (1893:4)]; S. insulanus
probably not closely related to the Recent species and Jung & Heitz, 2001 from the Escudo de Yeraguas
may belong to anotherclade which has become extinct. Formation, middle Pliocene of Panama; S. mardieae
Petuch, 2004 from the early Miocene of the Chipola
beds, Florida, U.S.A.; S. obliteratus Hanna, 1926 from
SYSTEMATICS
the Pliocene ofImperial County, California, U.S.A.; S.
Family Strombidae Rafinesque, 1815 radix Brongniart, 1823 from the Early Oligocene of
Europe (Figure 4); S. toroensis Jung & Heitz, 2001
Persististrombus Kronenberg and Lee gen. nov. from the Cayo Agua Formation, early Pliocene of
Panama; a radiation ofthe Middle Miocene ofEurope
Type species: Strombus grantilatus Swainson, 1822 (spHeacrizehsaurseepror&tedKrboyneJnubnegrg&inHperietpz.)(a2n00d1)a nwuhmicbheraroef
Pliocene to Recent
identified by means ofopen nomenclature or by letters.
Description: Shell ofmoderate size for family, fusiform, Powell (1988:17) also listed a - possibly new - species
shoulder knobs distinct on body whorl, slightly which he allocated to Strombus (Lentigo), but speci-
expanded outer lip with sharp, unglazed rim and no mens ofit haven't been examined by us. This species is
extensions, regularly divided callus on columella, not unlikely a Persististrombus as well. We agree with
anterior canal short, posterior canal or groove absent Jung & Heitz (2001:28) that S. granulatus cortezianus
or obsolete. Protoconch elongate and conical with four Durham, 1962 [new name for S. granulatus acutus
to five smooth whorls. Adaxial side of outer lip Durham, 1950 non G. Perry, 1811] is a synonym of5.
smooth, plicate, or granulate. granulatus. For an overview of species allocated to
Page 258 The Veliger, Vol. 49, No. 4
Figure 4. Persististvonibus radix (Brongniart, 1823). France,
dept. Landes, Espibos, Gaas. Chattian, lateOligocene. Leg. B.
Landau, collection B. Landau. Actual size 64.0 mm. A.
Figure 2. Persististrombiis honel/ii (Brongniart, 1823). Apertural view, B. Apical view, C. Dorsal view. Photos
France, Dept. Gironde, Le Peloua. Burdigalian, Early B. Landau.
Miocene. Leg. B. Landau, collection B. Landau. Actual size
86.8 mm. A. Apertural view, B. Apical view, C. Dorsal view.
Photos B. Landau. a more distinct posterior canal, a number of small
triangular extensions at the abapical side of the outer
Persististrombiis gen. nov. through time and space, see lip on the flange between the stromboid notch and the
Table 1 and Figure 5. anteriorcanal, very often ratherworn in L. lentiginosiis,
Although Persististrombusgen. nov. has a numberof and a columellar callus which does not reach the base
characters in common with Lentigo Jousseaume, 1886 of the columella, but is thickened at its abapical part,
(TS by monotypy: Stroinbiis lentiginosiis Linnaeus, but not forming a distinct pad as in some species of
1758), there are conspicuous differences: in Lentigo the Euprotonnis. Species assigned to Persististrombus gen.
adapical part ofthe outerlip has two notches, resulting nov. also have a relatively higher spire than do species
in two lobes, ofwhich the most adaxial one is attached of Lentigo, but within Persististrombus gen. nov.
to the spire ofthe shell; species assignedto Lentigohave species with a low spire do occur (Harzhauser &
Kronenberg, in prep.). Lentigo is here considered to be
restricted to the Indo-Pacific; see also Kronenberg &
Vermeij (2002). Persististrombus gen. nov. has many
characters in common with Strombus Linnaeus, 1758
(type species by SD Montfort, 1810: Strombus pugilis
Linnaeus, 1758), especially the spire. Differences
between Persististrombus gen. nov. and Strombus are
more difficult to quantify, and are more qualitive. In
Strombus the tips of the shoulder knobs are pointed
whereas in Persististrombus gen. nov. these tips are
usually rounded. But the population of P. granulatus
from the Islas Galapagos has the tips of the shoulder
knobs, better referred to as spines, pointed. In
Strombus there is no sculpture in the form of knob-
like structures abapical of the row of these shoulder
knobs. In Persististrombus gen. nov. there usually are
one or sometimes two of such rows, but in the middle
FFriagnucree, 3.DepPte.rsiGsitriosntdroem.lniLse hPoneelloluiai. (BBurrodniggnailairatn,, 1E8a2r3l).y Miocene of Europe there is at least one species of
Miocene. Leg. B. Landau, collection B. Landau. Actual size Persististrombus gen. nov. which only has the shoulder
83.0 mm. A. Apertural view, B. Dorsal view. Photos knobs present, and again the population of P.
B. Landau. granulatus from the Islas Galapagos does not have
G. C. Kronenberg & H. G. Lee, 2006 Page 259
Table
1
Distribution in time and space ofPersististrombiis gen.nov. species.
Species Age Locality
P. radix Oligocene Mediterranean region
P. bonellii early Miocene Mediterranean region
P. aldrichi early Miocene Florida, U.S.A.
P. mardiae early Miocene Florida, U.S.A.
P. spp. Vienna basin early middle Miocene Vienna Basin, Austria
P. cf. insulanus early middle Miocene Grenadines
P. sp. C (Jung and Heitz) middle - late Miocene Venezuela
P. sp. A (Jung and Heitz) late Miocene Venezuela
P. barrigonensis early Pliocene Venezuela
P. toroensis early Pliocene Panama
P. sp. E (Jung and Heitz) early Pliocene Jamaica
P. insulanus middle Pliocene Panama
P. obliteratus Pliocene California, U.S.A.
P. sp. B (Jung and Heitz) Pliocene Panama
P. bahrae Plio-Pleistocene Islas Galapagos
P. granulatus late Pliocene - Recent Panamic fauna province
P. sp. D (Jung and Heitz) Pleistocene Panama
a second or third row ofknobs present in all specimens body whorl is never smooth as in Strombus. In
(Kronenberg & Lee, 2005). Fossil species of Strombus Strombus the outer lip is more widely expanded than
s.s. may have spiral sculpture (e.g., Strombus lindae inPersististrombusgen. nov., and the adapical aspect of
Petuch, 1991; see Petuch 1994: pi. 21, fig. A) as grooves the outer lip (wing) is more or less pointed (not very
on the body whorl. But also the ^'nicaragueusis" form evident in all specimens of 5'. alatus). This outer lip
ofStrombuspugilis (see Clench & Abbott, 1941: pi. 6) expansion is particularly evident in the apical view: In
has this kind of spiral sculpture on (part of) the body Strombusthelabrumarchesventrally fromits posterior
whorl. In P. aldrichi this sculpture, though less origin at the suture so as to form a large open sinus
conspicuous, is present, but the abapical part of the with its free margin directed abaxially, whereas that
Figure 5. Geographic and Stratigraphic Distribution ofPersististrombus gen. nov. species.
Page 260 The Veliger, Vol. 49, No. 4
Table 2
Summary ofdistribution and characters ofLeuligo; Persististrombus gen. nov.; Stronibus; and Lobatus.
Lentigo Persististrombus Strombus Lobatus
Zoogeographical province Indo-Pacific Parathetys ~ Caribbean - Panamic ? Parathetys - Caribbean -
Caribbean - Panamic
Panamic
Tip ofshoulder knobs rounded rounded pointed rounded
Knobs abapical ofshoulder knobs present usually present absent usually absent
Spiral grooves in adult species absent absent absent or present present
Outer lip slightly slightly expanded clearly expanded clearly expanded
expanded
Transition lateral part outer lip to rounded rounded pointed; not always Variable, from rounded to
adapical part outer lip clear in S. alatus pointed, but when pointed
formingadistinctgroovefrom
tip ofpoint into the aperture
Abapical part ofouter lip bilobed simple simple simple
Triangular projections between present absent absent absent
strombid notch and anterior cana
1
Columellar callus thickened not thickened not thickened not thickened
abapical
aspect of Persististrombus shows a narrower sinus larvae originating from the continental population. It
which tends to curve adaxially at its free margin. seems that there is a genetic factor involved in the
Juveniles of P. granulatus (see e.g., Emerson & Old, various insular populations' phenotypy. There may
1963:8fig. 7)haveadistinctspiralsculpture, acharacter even be significant segregation of genomes among
which they share with Strombus. Strombus is restricted populations ofP. granulatus (s.l.) by island or group of
toAmerica,anditmayhavebeenderivedfroma species islands in the Galapagos. The islands may be the
ofPersististrombus gen. nov. metaphorical battlefield for the (genetic) independence
Persististrombus gen. nov. differs from other Amer- of the earlier immigrant waifs [as in the Isla Santa Fe
ican species, here assigned to Lobatus Iredale, 1921, in morphs, see illustrations in Kronenberg & Lee (2004)].
rate ofexpansion of the outer lip and sculpture ofthe However, at present we see no constant difference in
bodywhorl. Severalgenuslevel taxa fortheseAmerican characters of the shells to make a clear conchological
species are available, viz. Aliger Thiele, 1929 (TS separationamongpopulations. Weareaware thatthere
Strombus gallus Linnaeus, 1758); Eustrombus Wenz, is no proof for this hypothesis, and therefore this is
1940 (TS Strombusgigas Linnaeus, 1758); Macrostrom- highly speculative at present. However, this hypothesis
bus Petuch, 1994 (TS Strombus co.status Gmelin, 1791); can probably be tested by molecular analysis.
and Titanostrombus Petuch, 1994 (TS Strombus goliath A NOTE ON LOBATUS IREDALE, 1921
Schroter, 1805 [not 1905 (Petuch, 1994:261), an
apparent lapsus calami]. The relationships within these
Jung and Heitz (2001:48-50, fig. 26) described
American species are still unclear, and whether these Strombusfetus from the Escudo de Veraguas Forma-
should be regarded as subgenera ofLobatus is beyond tion (late Pliocene) ofPanama and assigned this species
the scope ofthe present paper; see table 2. to the subgenus Lentigo. They based their description
Persististrombusgranulatus occurs from the northern on only one specimen and stated that this species is not
end of the Gulf of California (Sea of Cortez) to relatedto anyofthe speciestheystudied, statingthatS.
Ecuador (Keen, 1971:421). Skoglund (2002:55) added fetus resembles S. raninus Gmelin. 1791 only superfi-
some records to the known distribution, including Isla cially. They mentioned only one difference, viz. the size
Gorgona (Colombia) and Islas del Coco (Costa Rica) of the knobs on the shoulder of the body whorl.
southward to Zorritos, Peru. Finet (1994) listed Islas Indeed, most specimens of the Recent S. raninus have
Galapagos and provided other references in support. two large, spine-like knobs on the shoulder ofthe body
Given the rather aberrant shape of some of the whorl, which character is expressed in the nomen S.
specimens originating from the Islas Galapagos in bituberculatus Lamarck, 1822, a synonym. Yet this is
comparison to the continental specimens of P. granu- not always the case. In the privatecollection ofthe first
latus, we believe a process of speciation of the author there are two specimens, viz. one from Aruba,
Galapagos population is emerging. This speciation Paardenbaai, inside reef in seaweed field, leg. Jan
may, however, be frustrated by an infrequent influx of Berkhout, 1967 (GCK 5419); one from Aruba, Secoedi
&
G. C. Kronenberg H. G. Lee, 2006 Page 261
Palma, leg. Jan Berkhout, 1968 (GCK 5423) in which Scenario 1)
the two last knobs on the shoulder ofthe body whorl
are only slightly enlarged and one specimen from All recent American species are descendants of
Panama, Isla Bastimentos, Bocas del Toro, found dead a species which is also ancestral to the recent P.
in surf zone, leg. Monika Forner, 30 November 1997 granulatus. This may have happened in one single wave
(GCK 5858) in which the development ofthe shoulder (that is that one species ofPersististrombus is ancestral
knobs is not different from S. fetus as illustrated by to all western Atlantic and Panamic Recent species) or
Jung & Heitz. After studying the description and in two (or more) waves in which one of these waves
illustrations, we regard Strombusfetusas a synonym of ended up in S. pugilis, S. alatus and S. gracilior; the
S. raninus, which may also be true for S. praercminus other(s) resulted in all other species. Based on the
Kronenberg & Dekker, 2000 [new name for Strombus molecular data as presented by Latiolais et al. (2006)
this scenario oftwo waves seems most likely.
wilsonorum Petuch, 1994 non Abbott, 1967] and
Strombus magolecciai Macsotay & Villaroel, 2001.
Scenario 2)
Petuch (1994) named a number of fossil Caribbean
strombid taxa, both at the genus and species level. For Sacco (1893:12) postulated that the late Miocene to
Strombus ranimis, Petuch used the subgenus name late Pliocene Strombus coronatus De France, 1827
Lobatus Iredale, 1921. Jung and Heitz (2001:40) (from the Tortonian through Piacenzian of Europe) is
criticized the use of the name Lobatus by Petuch ancestral to both the Recent West African Strombus
because the introduction of Lobatus was an historical latus (as Strombus bubonius Lamarck, 1822) and the
accident, referring to Abbott (1960:53). However, the Recent Strombus costatus (as Strombus accipitrinus
introduction o^LobatusbyIredale (1921:208) meetsthe Lamarck, 1822). All threeofthesespecies areextremely
requirements of the ICZN (Article 12.2.5), and variable (see for Strombus (s.l.) coronatus e.g., Sacco,
therefore the name Lobatus Iredale, 1921 (TS Strombus 1893, pi. 1, for Strombus (s.l.) latus DeTurck et al. pis
bituberculatus by monotypy) is available. 102, 103 and for Strombus (s.l.) costatus DeTurck et al.
pis 43, 44), and a large number ofnames are available,
especially for S. coronatus; see Sacco (1893).
POSSIBLE PHYLOGENIES OF RECENT Beneventi & Piccoli (1969), based on a number of
AMERICAN STROMBIDS
fossil species, elaborated on this scenario and conclud-
Kronenberg & Vermeij (2002:53) argued that the ed that many Recent species descend from a lineage
Recent Western Atlantic and Panamic strombids (in- started by StrombusfortisiBrongniart, 1823 through S.
cluding the West African Strombus latus Gmelin, 1791, radix and S. (s.l.) coronatus, to Recent Western
Atlantic and Panamic province species, but also to
and excluding the Indo-Pacific Gibberulus gibbosus
Recent Indo-Pacific species assigned to Euprotomus
(Roding, 1798) which was reported by Mienis (1978)
and Lentigo. [The evolution of Persististrombus in
from the Islas Galapagos [as Strombus (Gibberulus)
Europe, except for P. radix and P. bonellii will be
gibberulus gibbosus]), are monophyletic. This may be discussed elswhere (Harzhauser & Kronenberg. in
true because we have not been able to trace any fossil &
prep.)]. In their tree (Beneventi Piccolo, 1969:17)
evidence ofany Strombus (s.l.) in the Western Atlantic Persististrombusgranulatusalso descendedfrom Strom-
fossil record before Persististrombus gen.nov. made its bus coronatus, contrary to the results as shown by Jung
appearance there. & Heitz (2001) and our view. Also, the loss of the
Within this possibleclade a number ofgroups (based extremely dilated outer lip, as in Dilatilabrum, in
on overall shell characters, and for which genus-level Persististrombus radix and the subsequently regaining
names are available; see above) can be discerned, viz.: such a wing as present in many ofthe western Atlantic
S. costatus; S. gigas; S. ranimis + S. peruvianus + S. and Panamic fauna province species, seems unlikely.
gallus; S. goliath + S. galeatus. It should be noted here However, a derivation of some Indo-Pacific species
that in the analysis of stromboidean genus-level taxa, from a strombus radix-Hke species is not that unlikely.
based principally on anatomical characters, Simone There are some morphological characters which link
(2004) puts Strombus ranimis (allocated to Tricornis by bernielandaui Harzhauser, 2007, from the Oligocene
Simone) apart from the other American strombid late Chattian Warak formation, Gebel Madrakah,
species, i.e., branching off before Lambis, while all Oman, S. gijskronenbergi Harzhauser, 2007 from the
otherAmerican strombid species(asfarasexamined by Miocene Aquitanian Gubbarah formation, Gebel
Simone) branch offafter Lambis. Madrakah, Oman, S. quilonensis Dey, 1961 from the
Some possible scenarios for a phylogeny of the ?late Miocene of southern India and Strombus pre-
American speciescan be taken into consideration. With occupatus Finlay, 1927 from the early to late Miocene
no pretense to an exhaustive presentation, we mention ofJava and Borneo (Indonesia) although the knobs are
a few such, which can be tested using molecular data. more strongly developed and much more spine-like.
Page 262 The Veliger, Vol. 49, No. 4
reminiscent ofStronihiis coronatus and Stronihiis latus, modern fauna, and disappeared during the extinction
and to some degree resembling Persististrombus. wave at the end ofthe Eocene leaving only D. roegli to
Stvombiispreoccupatus was first assigned to Lentigo persist into the Oligocene, when it perished.
(as a subgenus) by Abbott (1960:123) but later Savazzi {fide Harzhauser, 2001:58) suggests that the
(1965:402) transferred to Dolomena Wenz, 1940 (also strombid notch evolved twice. Given the presence of
as a subgenus). [Dolomena Iredale, 1931:212 is a nomen the strombid notch in some other grades or clades of
nudwn, see Kronenberg & Dharma, 2005]. Stroinhus Stromboidea (Pugnellidae; see remark by Kronenberg
sedanensis Martin, 1899 of the early Miocene of & Burger, 2002:43) and Rimella-Mke species, Ectinochi-
Indonesia and Pakistan was assigned to Dolomena (as lus, Varicospira and other genera (Clark & Palmer,
a subgenus) by Abbott (1960:102). This was followed 1923; Burger& Kronenberg, 2006), we suggest that this
by Raven (2002:13, pi. 5 fig. 26a, 26b) with Dolomena character may have arisen even more than twice.
as a genus. In general shape Strombus sedanensis also Simone (2005: 248) used the absence of a strombid
reminds one ofcertain forms of S. coronatus, S. latus, notch in Recent Lobatus goliath as a reversion in his
and, to a lesser extent, as the outer lip ofthat species is listing of characters he used in his analysis. This
clearly more dilated, S. costatus. Without providing an strombid notch, however, is present in some specimens
allocation for both these species, we reject the of L. goliath, and the absence of the notch in certain
assignment of S. sedanensis to Lentigo (see remarks specimens is here considered not to be a reversion as
under description of Persististrombus gen. nov.) or argued by Simone (2005) but part of the intraspecific
Dolomena, because the structure ofthe outer lip clearly variation ofthis species.
differs from species of that latter genus. Other species The position of Strombiconus Marks, 1951 (TS
from the Miocene of Indonesia, such as S. injlatus Strombiconus ecuadorensis Marks, 1951, which is the
Martin, 1879, S. herklotsi Martin, 1880, S. tuberosus only species ever assigned to this genus) from the Early
Martin, 1883, S. tjilonganensis Martin 1899, as well as Miocene of Ecuador is enigmatic. The two known
S. mekranicus Vredenburg, 1928 from the Miocene of specimens on which both the description ofthe species
Pakistan (all allocated to Tricornisby Abbott, 1960:61- andthegenuswerebased arejuvenileswithworn apices
62) should be critically re-examined in teiTns of their (Marks, 1951:141-142, pi. 9 figs 10-11), and the
genericposition andpossible relationto theRecent and systematic position ofthis genus and species, although
fossil Indo-Pacific species mentioned above. This task probably strombid, cannot be determined. The same is
is beyond the scope ofthe present report. true for the more recently described Austrombus
Nielsen, 2005 (TS Conus medinae Philippi, 1887) from
Scenario 3) the Miocene of Chile, as already acknowledged by
Nielsen (2005:1122).
The genus Dilatilabrum Cossmann, 1904 [TS Strom- The overriding problem of convergence in the
busfortisi Brongniart, 1823 from the Lutetian (middle Strombidae impedes a morphological analysis of
Eocene) ofItaly] may have been ancestral to (some oO evolutionary relationships. An example is the resem-
the broad winged species. Species of Dilatilabrum are blance between Lobatus gallus and Tricornis tricornis
characterized by a widely-dilated outer lip (wing) and and the above-mentioned close resemblance of Dilati-
a large, narrow keel on the shoulder of the dorsal side labrum with some of the fossil and Recent Caribbean
ofthe body whorl. Based on the general shape and the and Panamic species. What appears to emerge is
presence of a keel-like ridge on the dorsum, both S. a mosaic pattern of characters that appear, disappear
dominator Pilsbry & Johnson, 1917 from the Miocene and reappear between lineages, but also within one
of Santo Domingo, Dominican Republic and S. single lineage, see also Landau etal. (in prep.) Weagree
dominator delabechei Rutsch, 1931, known from the with Petuch (1994:258) that none of the American
Pliocene Bowden Formation of Jamaica, could be species should be assigned to Tricornis Jousseaume,
assigned to DUatilabrum, thus linking the Eocene 5. 1886, whichonly superficially resemblescertainofthese
fortisi to the Recent broad winged species. But S. American species.
dominators.s. andS. dominatordelabechihaveashallow
strombid notch and a more or less clearly developed Acknowledgments. We thank Dr. Alan G. Beu, Institute of
spiral sculptureon thelatterpart ofthelastwhorl, both Geological and Nuclear Sciences, Lower Hutt, New Zealand,
absent in D. fortisi. The large D. roegli (Harzhauser, formaking a copy ofthe paper by Garcia-Talavera available;
2001) [as Strombus (Dilatilabrum) roegli] from the Dr. Eugene V. Coan, Palo Alto, California, U.S.A., for
Oligocene of Greece and Iran does have a strombid information on Strombus obliteratus; Dr. Yves Finet for
information on Islas Galapagos references; William Frank of
notch, but does not have the spiral sculpture present in
Jacksonville, Florida, U.S.A. for technical assistance in
S. dominator s.l.. Based on the consensus tree as constructing the distribution map; Richard E. Petit, North
presented by Latiolais et at. (2006:440) we do not think Myrtle Beach, South Carolina, U.S.A.. forsharinghisopinion
that the genus Dilatilabrum is closely related to the on the status of Lobatus; Han Raven, the Hague, the
&
G. C. Kronenberg H. G. Lee, 2006 Page 263
Netherlands for the translation from Spanish: Dr. Geerat J. Islands: a documented faunal list. Ed. Museum d'Histoire
Vermeij, University of California at Davis, for critically naturelle: Geneve. Pp. 1-180.
reading the first draft ofthe manuscript and making valuable Garcia-Talavera, F. 1993. Resultados cientificos del
suggestions. Dr. Mathias Harzhauser, Naturhistorisches proyecto Galapagos: patrimonio de la humanidad.
Museum in Wien, read the second draft of the manuscript No. 3. Los moluscos marinos fosiles. TFMC: Tenerife,
and gave useful tips. An anonymous reviewer critically read Spain. Pp. 1-61, 9 pis.
the manuscript and made valuable remarks and suggestions. Grant, U. S. & H. R. Gale. 1931. Catalogue ofthe Marine
Bernard Landau, Albvifeira, Portugal, isgreatlyacknowledged Pliocene and Pleisocene Mollusca of California and
for making images of specimens of P. radix and P. honellii adjacent regions with notes on their morphology,
available. The first author also wants to thank Adri W. classification, and nomenclature and a special treatment
Burger, Heerhugowaard, the Netherlands and Ron P.A. of the Pectinidae and the Turridae (including a few
Voskuil, Hulst, the Netherlands for stimulating discussions. Miocene and Recent species) together with a summary of
Dr. Mathias Harzhauser, Naturhistorisches Museum in Wien thestratigraphicrelations ofthe formations involved. San
forkindhospitalityand access to thecollection in hiscustody, Diego Society ofNatural History, Memoir 1, 1-1036.
and finally Marianne Matthijssen for abiding support. Hanna, G. D. 1926. Paleontology of Coyote Mountain,
Imperial County, California. Proceedings ofthe Califor-
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