Table Of ContentB.M. Landau, F. Frydman&CM. DaSilva Novapex 8 (3-4): 111-121.10 décembre 2007
Morum (Oniscidia) domingense from the Caribbean Neogene
vs. the Récent M. (O.) lindae: One taxon or two?
Bernard M. LANDAU
International Health Centres, Av. Infante D. Henrique 7, 8200 Albufeira, Portugal. Centra de
Geologia. Universidade de Lisboa. C6. Campo Grande.
1749-016 LISBOA. Portugal.
[email protected]
FRYDMAN
Franck
3, Rue Charles François Dupuis, 75003 Paris, France.
Carlos M. DA SILVA
Departamento e Centra de Geologia. Universidade de Lisboa. C6. Campo Grande.
1749-016 LISBOA. Portugal.
KEYWORDS. Gastropoda, Harpidae, Morum. Caribbean. Neogene.
ABSTRACT. The Morum chipolanum-group ofMorum (Oniscidia) species has a long géologie
history in Tropical America, although their spécimens are never abundant. Most of the original
descriptions were based on very few shells. The lack of information concerning intraspecific
variability makes classification sometimes difficult within this group of species with somewhat
conservative shell design. This study attempts to distinguish several Caribbean Neogene taxa on
the basis of combined shell morphometry and qualitative shell characteristics, paying spécial
attention to the distinction between the shells ofM. (O.) domingense from the Upper Miocene-
Lovver Pliocène ofthe Dominican Republic and those ofthe Récent M. (O.) lindae from the coasts
of Colombia and Venezuela, in order to ascertain if they should be assigned to the same or to
distinct taxa. The largest number of spécimens yet considered is assembled, drawn from
adequately illustrated spécimens in the literature, collections and high quality digital photographs
posted on the Internet. Analysis shows that by morphometry alone thèse two taxa cannot be
reliably distinguished, but when the morphometric data is considered in combination with other
observable shell characteristics the two sets ofshells are most likely distinct, actually representing
two différent taxa. This conclusion is furtherstrengthened by palaeobiogeographical data.
INTRODUCTION 1959) and the Récent Brazilian M. (O.) matthewsi
Emerson, 1967.
The genus Morum Rôding, 1798, now placed in the A handful ofspecies belonging to the M. chipolanum-
family Harpidae Bronn. 1849 (Hughes & Emerson, group hâve been described from the Caribbean
1987) superfamily Muricoidea Rafinesque. 1815 Neogene and Récent faunas.
(Bouchet & Rocroi, 2005), comprises a small group of The earliest Tropical American records are for the
species ofcarnivorous marine gastropods (J. Smith in Peruvian M. peruvianum Olsson, 1931 from the
Beesley et al., 1998), occurring in tropical to Eocene Chira Formation (Olsson. 1931) and for M.
subtropical seas Worldwide. They are found in shallow (O.) harpula (Conrad, 1848) from the Lower
to fairly deep waters, but are always uncommon to Oligocène Vicksburg group of Mississippi. USA
very rare (J. Smith in Beesley et ai. 1998). The (MacNeil & Dockery, 1984).
subgenus Oniscidia Môrsch, 1852 (= Cancellomorum From the Lower Miocène the following species hâve
Emerson & Old, 1963, see Vokes, 1998) comprises a been reported: Morum chipolanum Dali, 1925 from
group of Morum species with a cancellate sculpture; the Chipola Formation of Florida, USA and Baitoa
for full discussion on the taxonomic problems Formation of the Dominican Republic (see Vokes,
regarding the name Oniscidia we refer to Beu (1976) 1998), and M. (Onicidia)jungi Landau, 1996 from the
and Vokes (1998). In this paper we will only discuss Cantaure Formation ofVenezuela (see Landau. 1996).
what Vokes (1998) called the M. chipolanum-group of M. harrisi Maury. 1925. from the Lower Miocène
species, characterized by an ornamentation of Pirabas Limestone of Brazil. is based on internai
elongated pustules on the pariétal shield as opposed to moulds and its apertural features are unknown
coarse rugae, as seen in Morum ("Oniscidia'") sp. from (Maury, 1925). Morum (O.) coxi (Trechmann. 1935)
the Middle Eocene Gatuncillo Formation (Woodring, occurs in the lower Middle Miocène (Robinson &
111
H M. I WDAl ,F. FRYDMAN&< M DaSILVA Morum (Oniscidia) domingense
June. 1972)Grand Baj FormationofCamacou(Jung, 1, Figs 1-2) rather than of M. domingense." (Vokes,
ll)7l). In the Upper Miocène and Louer Pliocène 1998,p. 20).
assemblages M. (O). domingense (Sowerby, 1850) In ail Caribbean assemblages in which Morum occurs
OCCUTS in the Cercado, Gurabo and Mao Formations of it is uncommon to very rare and at the time when
the Dominican Republic (Vokes. 1998). many of the above taxa were described, they where
Morum (O.) meganae Raymond, 1997 was recently known from one or two spécimens only. As larger
described from the Middle Pliocène Pinecrest Beds of numbers of shells hâve become available and the
I lorida and M. (O.) macgintyi Smith. 1937 [M. (O.) intraspecific variability better represented, the
obrienae Olsson &MPetit. 1964. is almost certainly a characteristics separating some of the taxa hâve
junior synonym of (O.) macgintyi (Emerson, 1967; become less clear.
Vokes. 1998)] occurs in the Upper Pliocène This paper started as a discussion over the validity of
Caloosahatcb.ee Formation ofFlorida (Petuch. 1994). M. (O.) lindae as a distinct taxon from the Mio-
The living M. (O.) dennisoni (Reeve, 1842), found Pliocene M. (O.) domingense. For each argument put
from the coasts of Louisiana, USA south to the forward supporting their séparation, a Récent shell
Caribbean, is immediately distinguished from this refuting their isolation could be found. Therefore, in
group by the larger size of fully adult spécimens, by the light of the greater amount of material available,
its weaker spiral sculpture, with cords of unequal we attempt to clarify whether M. (O.) lindae and M.
strenght making it diffîcult often to distinguish cords (O.) domingense are indeed distinct taxa by means of
ofprimary or secondary strength. This is not the case a morphometric comparison. We hâve added
with any of the other species of the M. chipolanum- morphometric data on the shells of two further
group in which ail the cords are more strongly undisputed extinct species; M. (O.) chipolanum and
elevated and of roughly equal strength. The subsutral M. (O.) jungi and the living M. (O.) dennisoni in
ramp of the last whorl in M. (O.) dennisoni is much order to investigate the interspecific variability within
vvider than in any ofthe other members of the group this group of gastropods with very similar shells.
and there is a marked thickening ofthe inner aspect of Although the number of shells used in this study is
the outer lip mid-height in M. (O.) dennisoni not less than that usually used for morphometric studies,
présent in the other group members. Finally the we stress that for ail species this is the largest number
pustules tend to be coarser in the shells of M. (O.) ofspécimens so faravailable.
dennisoni. Rios (1994) suggested M. (O.) lindae was a
synonym of M. (O.) dennisoni. At the time of MATERIAL AND METHODS
publication probably only a couple of spécimens of
the latterwere known, and Rios probably considered it Data for the fossil shells were obtained from
a subadult spécimen of M. (O.) dennisoni. However, measurements, rib and cord counts as well as
as more material of M. (O.) lindae has become observations of shell characteristics of the type
available it is clearthèse différences are consistMent and material clearly illustrated in Landau (1996) and
not gerontic characters, as the smallest shell of (O.) Vokes (1998) and further spécimens in the Bernard
dennisoni we hâve found is within the maximum size Landau collection (M. (O.) chipolanum: 10 spécimens,
range of M. (O.) lindae and still shows thèse 3 from locality TU (Tulane University locality
consistent différences. number) 458, 3 from TU 546, 5 from TU 826; M. (O.)
Petuch (1981) assigned a spécimen collected from 11 jungi: 5 spécimens from TU 1269; M. (O.)
meters depth off the Goajira Peninsula, Colombia to domingense: 34 spécimens, 2 Cercado Formation from
M. domingense, noting at the time that the spécimens NMB (Naturhistorisches Muséum Basel locality
of the species known from the fossil record had 12 number 16832, 32 Gurabo Formation, 2 from TU
axial ridges, whereas the Récent shell had 16. 1354, 14 from TU 1215, 13 from TU 1219,3 from TU
Subsequently, he reassigned the Récent spécimen to 1373).
Morum (Cancellomorum) lindae Petuch, 1987 (p. 95, The shells ofsome Morum (Oniscidia) species display
pi. 23, Figs 1-2). HMe considered M. domingense "the a considérable variation in size between fully adult
direct ancestor" of lindae and noted that the fossil spécimens, herein interpreted as shells with a fully
M
species: "(...) has fewer axial ribs, is broader and expanded pariétal shield and thickened outer lip. In
more angled, has a smaller pariétal shield, and is far chipolanum, for example, adult shells range from 22.2
less sculptured and less squamose" (Petuch, 1987, p. mm to 38.8 mm, M. M(O.) domingense from 20.5 mm to
95). 35.8 mm, adults of (O.)jungi seem more uniform
Vokes (1998) accepted the distinction between the in size. For the purpose of this study we hâve not
mm
fossil and récent taxa, specifying that the number of included any adult spécimen smaller than 23.5 in
axial ribs in the fossil shells of M. (O.) domingense height.
varied from 10-12 and added that: "(...) the nature of
the low stepped spire, with its incised suture and the Data for Récent shells of M. (O.) lindae were
numerous fine axial lamellae, suggests that M. lindae collected from three spécimens in the Franck Frydman
is the linear descendent ofthe more recently described collection and a further 13 shells illustrated on the
Cantaure Formation M.jungi Landau (1996, p. 53, pi. Internet (Fémorale, 7 spécimens; Jaxshells, 3
112
.
B.M. Landau, F. Frydman &CM. Da Silva Novapex 8 (3-4): 111-121, 10 décembre 2007
spécimens; worldwideconchology. 1 spécimen; internet data was collected from the following sites:
mineralislamacla. 1 spécimen). M. (O.) dennisoni data Guest Shells (1996). Frank (1998). Coltro
was collected from the Internet (Fémorale 8). Only (2006); worldwideconchology.com (2006).
spécimens illustrated with both ventral and dorsal Measurements were taken as illustrated in Figure 1
views and with the correct standard orientation were Measurements of Internet photographs were taken
considered. Whilst collecting data from images on the with the measuring tool after copying the pictures into
web may not usually be idéal, we suggest that for rare Photoshop, thereafter measurements calculated based
spécimens such as this (the type material consists of on the original length given for each spécimen. To
the holotype alone) it is a useful way to consider the minimize observer error ail measurements were taken
maximum number ofspécimens known. Forthis study by the first author.
Figure 1. Morphometric measurements taken from a Morum shell.
W
H = total height. = width ofshell, Hs = height ofspire. Ha = height ofaperture, Wsh = maximum width of
pariétal shield.
RESULTS Caribbean non-planktotrophic development is the
norm. This coincides with the type of development
Several spécimens of the fossil taxa M. (O.) observed in the living western Atlantic species Morum
domingense and M. (O.) chipolanum, and one Récent oniscus (Linnaeus. 1767) (type species of Morum).
shell of M. (O.) lindae had their protoconchs well which lays its eggs beneath small slabs ofcoral at low
preserved. In ail three the protoconch was small. tide level. The eggs undergo direct development and
consisting of 1.25-1.5 smooth whorls with a small the gastropods hatch as crawling young (Work. 1969).
nucleus. This small paucispiral type of protoconch is Protoconch morphology was not, therefore, useful in
highly suggestive of non-planktotrophic larval distinguishing between species within the M.
development. Bouchet (2002) reviewed data on Indo- chipolanum-group.
Pacific Morum species and concluded that
planktotrophy was the standard method of Morphometric results on the teleoconch show:
development in thèse species. He also noted that ail
the Caribbean species; M. lindae, M. domingense, and 1. In relation to their overall shape, there is overlap
M. dennisoni had paucispiral protoconchs. It therefore between the spécimens of four species when
seems that unlike the Indo-Pacific species. in the comparing width in relation to height. although
13
»
B M. l \\l)\l , F. FKYDMAN& CM. I)\ Su \ \ Morum (Oniscidia) domingense
\l il).) domingense tonds to hâve a narrower, more pack having a relatively narrower shell in relation to
elongated shell compared to the one of M. (O.) height (Fig. 2).
lindae. \l (O.) dennisoni can be separated from the
(Width height) height
/ /
0,70- iO. xx *
.c °A^ *° X°X
A A
0)
£
0,60- 4 +
"D
£ X
0,50-
1 1 1 mm
20 30 40 50 60
Height
O A
Figure 2. Morphometric plot of(width/height)/height. - M. domingense; - M. chipolanum; - M.jungi;
X - M. lindae; + - M. dennisoni.
2. None ofthe species, except M. dennisoni, can be apertural height (Figs 3,4).
distinguished on the basis ofrelative spire height or
(Aperture height height) height
/ /
*—
96" o X
O) °-
.c +
^
0,92- # +
.Oc) oX**Ox x +
S 0,88- a^x^ xk + +
AA a? x <^*
<D +
3 x
C 0,84- O A*X
X
<Q. X +
X
0,80-
1 1 1 mm
20 30 40 50 60
Height
O A
Figure 3. Morphometric plot of(aperture height/height)/height. - M. domingense; - M. chipolanum;
M.jungi; X - M. lindae; +- M. dennisoni.
14
B.M. Landau, F. Frydman& CM. Da Silva Novapex 8 (3-4): 111-121, 10 décembre 2007
(Spire height height) height
/ /
X
0,16-
D5 Qdo
+
+
0,12-
D)
*
.CCD \* x + +
X X
CD 0,08- + +
i_
+
c/)
0,04-
I 1 1 mm
20 30 40 50 60
Height
O A
Figure4. Morphometric plot of(spire height/height)/height. - M. domingense; - M. chipolanum; - M.
M
jungi; X - lindae; + - M. dennisoni.
3. M. (O.) chipolanum has the broadest pariétal shield with M. (O.) domingense and M. (O.) lindae, there is
ofthe five species, although there is some overlap no overlap with M. (O.)jungi (Fig. 5).
(Shield width height) height
/ /
.c .•ô
O) / o o
CD O o +
-c 0,30- +
X. .*° °q,/ +
+
•-.Q----X-.. x.y +
+
+
-o 0,20-
cd
le
C/) X
0,10-
I I I mm
20 30 40 50 60
Height
O A M
Figure 5. Morphometric plot of(shield width/height)/height. - M. domingense; - M. chipolanum: -
M
jungi; X - lindae; + - M. dennisoni.
M
4. (O.)jungi has consistently feweraxial ribs in fewerribs than M. (O.) lindae. M. (O.) dennisonican
M
relation to height than lindae. Although there is also be separated in having fewerribs in relation to
some overlap, M. (O.) domingensealso tends to hâve size (Fig. 6).
115
Ii.\l I U1DAU,F.FRYDMAN&CM.DASlLVA \4orum (Oniscidia) domingense
Number of ribs height
/
O
x"--.
16- >«X<
>
x xj
xy
'•*X €X
.Q — >S< . +'
E3 12- <H Q'+ +
p «o + +
-H-.;
A0<» A A
.
,AA A +
R- .
1 i i mm
20 30 40 50 60
Height
Figure 6. Morphometric plot ofnumberofaxial ribs/height. - M. domingense; O - M. chipolanum; A - M.
jungi; X - A/, /inclue: + - M. c/ennisoni.
M
5. M. (O.) lindae has consistently fewer spiral cords in hâve fewercords than (O.) domingense and M. (O.)
M
relation to height than (O.) chipolanum. Although jungi(Fig. 7).
M
there is some overlap, (O.) lindae also tends to
Number of cords height
/
12-
"oE /OO**** €>
o
10- IMMW) O
JtÉ»^.. :x + X"\ + + +
E
8- ,X «A>9S< X >ï( ,/
••-, X x '"'
fi-
1 1 1 mm
20 30 40 50 60
Height
O A M
Figure 7. Morphometric plot ofnumberofspiral cords/height. M. domingense; - M. chipolanum; -
jungi; X - M. lindae; + - M. dennisoni.
DISCUSSION than M. (O.) lindae, and M. (O.) lindae has
consistently fewer spiral cords than M. (O.)
M
It is clear from the results above that by this chipolanum while tending to hâve fewercords than
morphometric assessment alone it is not possible to (O.) domingense and M. (O.)jungi.
distinguish groups of spécimens easily within this Two further shell features are important in
assemblage of taxa with highly conservative shell distinguishing the four groups ofshells, which are not
form. Ourplots show that M. (O.) domingense tends to amenable to morphometric analysis: Thèse are the
M
hâve a narrower shell than (O.) lindae, M. (O.) shape of the pariétal shield and the degree of
chipolanum tends to hâve the broadest pariétal shield, development of secondary scabrous or squamous
that M. (O.) jungi has consistently fewer axial ribs sculpture formed by the axial growth lines. Morum
116
B.M. Landau, F. Frydman &CM. Da Silva Novapex 8 (3-4): 111-121, 10 décembre 2007
(Oniscidia) chipolamim and M. (O.)jungi both hâve a found in the West Indian and Colombian-Venezuelan-
shield which is rounded adapically, whereas in M. (O.) Trinidad Subprovince share very few taxa in common.
domingense and M. (O.) lindae the shield is somewhat Despite extensive collecting in rich Pliocène
pointed in the area ofthe posterior canal (see Plate 1). sédiments in the area, there are no reported
This character does not seem too variable occurrences ofM. (O.) domingense in the fossil record
intraspecifically. ofthe Colombian-Venezuelan-Trinidad Subprovince.
The second, the surface scabrosity, is difficult to Morum (O.) lindae occurs exclusively off the
quantify. There is no doubt that MM. (O.) chipolamim Caribbean coast of Colombia and Venezuela. As in
has the most scabrous surface and (O.) domingense the Neogene, the fauna ofthe northern coasts ofSouth
the least. Whilst some intraspecific variability exists, America contains a high number ofendémie éléments
the most scabrous spécimen of M. (O.) domingense (Petuch, 1987; Diaz, 1995). Indeed, Landau et al. (in
(Plate 1, Figure 4) is still less scabrous than the print PPP) argued that the Colombian-Venezuelan-
smoothest M. (O.) chipolamim. Morum (O.) lindae Trinidad Subprovince was probably in place since at
usually has a more scabrous surface than M. (O.) least the Early Miocène and has continued as a distinct
domingense, although in the occasional spécimen palaeobiogeographical unit to the présent day.
(Plate 1, Figure 4) there is little différence between Although Petuch (1982) argued that this area had
them. acted as a refugium for Gatunian species unchanged
Vokes (1998) discussed a further character, the spire since the Pliocène (a primary relict pocket, according
and considered M. (O.) lindae to hâve a more stepped to his nomenclature), this does not stand up to
spire than M. (O.) domingense. As can be seen from taxonomic scrutiny. Petuch (1988) laterconsidered it a
the spécimens illustrated (Plate 1) and the plot of secondary relict pocket, i.e. containing species which
(apertural height/height)/height (Figure 3) there is no closely resemble their Pliocène ancestors. There is no
différence in relative spire height between the three fossil record of M. (O.) lindae in the southern
species and only M. (O.) jungi has a significantly Caribbean assemblages. Only one single fragment of
more stepped spire, i.e. the infrasutural platform is Morum (Oniscidia) sp. from the Lower Pliocène Punta
more horizontal. Gavilân Formation, Falcôn Province, mainland
Venezuela, is known (BL collection), and it is too
PALAEOBIOGEOGRAPHIC
incomplète to identify specifically.
CONSIDERATIONS
CONCLUSIONS
The présent Caribbean région in the Miocène and
Pliocène was part ofthe larger palaeobiogeographical Using a combination of quantitative shell parameters
Gatunian Province, which straddled the présent day and qualitative shell characters of shield shape and
région of the Isthmus of Panama and included the scabrous surface one can conclude that M. (O.)
modem Caribbean région and Tropical American chipolanum is most clearly distinguished from its
Pacific (Woodring, 1974; Vermeij, 2005). The closure congeners by the round shape of its shield and
of the Central American Seaway (CAS) divided the strongly scabrous surface. Morum (O.) jungi is
Gatunian Sea, causing a significant change in the characterized again by its abapically rounded shield,
faunal composition on the Atlantic side. After the total although less expansive than in the previous species
closure of the CAS the Atlantic portion of the and in having consistently fewer axial ribs than its
Neogene Gatunian Province gave rise to the Récent congeners. Morum (O.) domingense and M. (O.)
Caribbean Province. The Neogene Gatunian Province lindae are less easily distinguished; M. (O.)
is characterized by a relative stability of gastropod domingense tends to hâve a more elongated shell,
taxa at generic level, but a very rapid turnover at fewer axial ribs as well as a greater numberofprimary
spécifie level (Landau et al., in print PPP). Thèse high spiral cords and also tends to hâve a smoother surface
rates of extinction and local disappearance are although there is some overlap with ail thèse features
accompanied by high rates of speciation (Allmon et between the two taxa. Apertural height, spire height
al., 1993; Jackson et ai, 1993). At species level it is and shield width are not useful to distinguish the
not unusual to hâve an extinction rate since the Early shells ofthe two species.
Pliocène to présent times of80-85% (Woodring, 1928; On the basis of protoconch morphology ail members
Jung, 1969, Landau et al., in press), with about half ofthe M. chipolanum-group, in which the protoconch
the long-lived taxa belonging to the most notoriously is known, hâve a small paucispiral protoconch, which
cosmopolitan group of gastropods, the tonnoideans strongly suggests a non planktotrophic larval
(Beu, in prep). development. This type ofdevelopment is commonly
Within the Gatunian Province, Morum (Oniscidia) associated with species having a shorter geological
domingensis is found exclusively in the West Indian longevity and a more restricted geographical
Subprovince of Woodring (1974). The northern distribution (Jablonski & Lutz, 1980; Scheltema,
Caribbean coasts of South America formed the 1989; Gili & Martinell, 1994), which might support
Colombian-Venezuelan-Trinidad Subprovince of the séparation ofthe two taxa. However, this does not
Woodring (1974). At spécifie level, the assemblages seem to be universally true across ail gastropod
117
B.M. l \\D\i . F. Frydm \\ &C.M.Da Sii.va Morum (Oniscidia) domingensi
groups, as Jackson et al. (Ic)%) l'ound no corrélation Melbourne. Part A, i-xvi, 1-563; Part B, i-viii,
between species longevity and inferred developmental 565-1234.
modes in Neogene Tropical American strombinids. Beu, A. G. 1976. Revision ofthe southwest Pacific
Whilst M. (O.) domingense and M. (().) lindae show species ofMorum (Oniscidia) (Gastropoda:
very similar shells, on the basis of our présent Cassidae).Journalojthe MalacologicalSocietyof
knowledge of the Caribbean faunal turnover and the Australia, 3(3-4): 223-321.
biogeography of the Neogene Atlantic Gatunian Beu, A. G. (in prep). Neogene Paleontology in the
faunas it would be very unlikely for an exclusively Northern Dominican Republic XX. Neogene
Upper Miocene-Lower Pliocène Gatunian West Indian tonnoidean gastropods ofthe Central American
species to occur today within the Colombian- région, with a revision ofthe Neogene and Récent
Venezuelan-Trinidad Subprovince. Bursidae, Personidae and Ranellidae ofCentral
Taking ail the above arguments into considération it is and South America.
most likely thaï M. (().) domingense and M. (().) Bouchet, P. 2002. Protoconchs, dispersai, and tectonic
lindae actually represent two distinct taxa. There is plates biogeography: new Pacific species of
insufficient évidence to suggest a direct lineage Morum (Gastropoda: Harpidae). Journalof
between the two based on their shell morphology and Conchology 37(5):533-549.
palaeobiogeographic distribution. Further information Bouchet, P. & Rocroi, J.P. 2005. Classification and
on the Morum (Oniseidia) species présent in the nomenclature ofgastropod families. Malacologia
Lower Pliocène Punta Gavilân Formation of 47(1-2): 1-397.
Venezuela might help to clarify the relationship Coltro M. 2006. Fémorale. Colombia - Harpidae.
between thèse members of the M. chipolanum-group Available at
in the southern Caribbean. http://www.femorale.com.br/shellphotos/
local spec2.asp?localidade=Colombia&family=H
ACKNOWLEDGEMENTS ARPIDAE&nav=2 (cited 20/12/2006).
Conchology, The Art and Science ofNature.
Our thanks to Marcus Coltro (Fémorale), Brazil, for
M Available from
sending us high quality photographs of lindae, and
http://www.worldwideconchology.com. Date of
giving permission to reproduce some of thèse access: February-March, 2007
photographs.
Diaz, J. 1995. Zoogeography ofmarine gastropods in
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Allmon, W. D.. Rosenberg, G, Portell, R. W. & 2): 104-121.
Schindler, K. S. 1993. Diversity ofAtlantic Emerson, W. K. 1967. A new species ofMorum from
coastal plain mollusks since the Pliocène. Science Brazil, with reamarks on related species
260: 1626-1629. (Gastropoda : Tonnacea). Veliger9(3): 289-292.
Beesley, P. L., Ross, G. J. B. & Wells, A. (Editors) Frank, W. 1998. Shells, Shell collecting, Nature.
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Australia, volume 5. Australian Biological http://www.jaxshells.org/ 913b.htm (cited
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Figures 8-23
8-9. Morum (Oniscidia) chipolanum Dali, 1925. Loc. TU 546. Chipola River, Calhoun County, Florida, USA,
mm
Lower Miocène, Chipola Formation. Height 38.6 (B. L. coll).
10-11. Morum (Oniscidia) domingense(Sowerby, 1850). Loc. TU 1219. Amina River, Dominican Republic,
mm
Lower Miocène, Chipola Formation. Height 34.9 (B. L. coll).
12-13. Morum (Oniscidia) domingense (Sowerby, 1850). Loc. TU 1215. Gurabo River, Dominican Republic,
Lower Miocène, Chipola Formation. Height 35.6 mm (B. L. coll).
14-15. Morum (Oniscidia) dennisoni (Reeve, 842). Vitoria, Espirito Santo State, Brasil, Récent. Height48.8
1
mm
(photo: Fémorale).
16-17. Morum (Oniscidia)jungi Landau, 1996. Loc. TU 1269. Casa Cantaure, Paraguanâ Peninsula, Falcôn
mm
State, Venezuela, Lower Miocène, Cantaure Formation. Height 25.0 (B. L. coll).
18-19. Morum (Oniscidia) lindae Petuch, 1988. Cabo de la Vêla, Colombia, Récent. Height 36.0 mm (photo:
Fémorale).
20-21. Morum (Oniscidia) lindae Petuch, 1988. Cabo de la Vêla, Colombia, Récent. Height 31.5 mm (photo:
Fémorale).
22-23. Morum (Oniscidia) lindae Petuch, 1988. Cabo de la Vêla, Colombia, Récent. Height 39.4 mm (photo:
Fémorale).
118
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