Paleocene Pareorine Turritellid Gastropods From The Pacific Slope of North America

THE NAUTILUS 121(1):1–16, 2007 Page 1
Paleocene pareorine turritellid gastropods from the Pacific slope

of North America
Richard L. Squires LouElla R. Saul
Department of Geological Sciences Invertebrate Paleontology Section
California State University Natural History Museum of Los
Northridge, CA 91330-8266 USA Angeles County
richard.squires@csun.edu 900 Exposition Boulevard
Los Angeles, CA 90007 USA
lousaul@earthlink.net
ABSTRACT (e.g., Zinsmeister, 1983) have generally placed both spe-

cies in Mesalia Gray, 1847, but some workers (e.g., Saul,
This paper presents the first detailed study of Mesalia martin- 1983a: fig. 2; Squires, 2003: table 2.4) have been hesitant
ezensis (Gabb, 1869) and Mesalia clarki (Dickerson, 1914a), the
to unequivocally use the genus name because of inad-
only two known pareorine (spout-bearing) turritellid gastro-
pods from the Pacific slope of North America. Both species are equate morphologic information about these species.
redescribed, in light of new morphologic information that also The shape of the aperture of the latter one was unknown
confirms their assignment to genus Mesalia Gray, 1847, which until now, and apertural information is critical in distin-
we believe to be congeneric with Sigmesalia Finlay and Mar- guishing parerorine gastropods from similar looking tur-
wick, 1937. New stratigraphic information allows for refine- ritellids (see “Systematic Paleontology” for morphological
ment of the chronologic range of each species. Mesalia mar- comparisons). There has also been the possibility that
tinezensis is of early late Paleocene (late Danian) to early late Mesalia macreadyi Waring, 1914, which has commonly
Paleocene (early Thanetian) age and ranges from northern been put into synonymy with M. martinezensis, could be
California to northern Baja California. Mesalia clarki is of late a distinct species.
middle to early late Paleocene age (late Selandian to early
We conclude that there are only two species, both
Thanetian) age and is known only from California; in southern
California it is restricted to a coralline-algal facies. Both species belonging to Mesalia, which we believe to be congeneric
have considerable variability in their spiral sculpture. with Sigmesalia Finlay and Marwick, 1937. In addition to
Mesalia originated during either the Late Cretaceous (Maas- providing new morphologic information about the study
trichtian) in northern Africa or the early Paleocene (Danian) in area Mesalia, we refine their geographical (Figure 1) and
northern Africa and western Iran. It became widespread during chronologic ranges (Figure 2). Both M. martinezensis
the warm (greenhouse) conditions of the Paleocene and and M. clarki have considerable variability in their spiral
Eocene but became geographically restricted during subse- sculpture.
quent cooler global conditions. Mesalia is an extant genus with Mesalia has long been reported (e.g., Cossmann, 1912)
possibly six species, and whose total geographic range is in as originating during the Late Cretaceous in the region
coastal waters in southern Portugal, southern Spain, Mediter-
between northern Africa and western Iran. Our review of
ranean Sea (primarily the western part), Canary Islands, and
the west coast of northern Africa. the literature shows that the genus most likely originated
during either the Late Cretaceous (Maastrichtian) or the
early Paleocene (Danian) in this Old World region.
Modern systematic studies of Mesalia are sorely lack-
INTRODUCTION ing as are detailed ecologic studies of the few extant
species. The classification scheme used here follows that
The object of this study was to rectify the identification of Bouchet et al. (2005: 249), and morphological terms
uncertainities concerning the only two known pareorine are taken from Cox (1960). Pacific slope of North
(spout-bearing) turritellid gastropod species from shal- America Turritella zones are taken from Saul (1983b).
low-marine rocks on the Pacific slope of North America. Institutional abbreviations used in the text are: ANSP:
They are Mesalia martinezensis (Gabb, 1869) and Mesa- Academy of Natural Sciences of Philadelphia; LACM:
lia clarki (Dickerson, 1914a). Most modern workers Natural History Museum of Los Angeles County, Mala-
cology Section; LACMIP: Natural History Museum of
Los Angeles County, Invertebrate Paleontology Section;
Corresponding author: Richard Squires UCMP: University of California, Berkeley, Museum of
email: richard.squires@csun.edu Paleontology.
Page 2 THE NAUTILUS, Vol. 121, No. 1
depths. These mollusks commonly include shallow-

marine mollusks such as naticid and buccinid gastropods,
as well as glycymerid and crassatellid bivalves. All are
indicative of normal marine salinities.
The type locality of Mesalia martinezensis has been
generally assigned (e.g., Keen and Bentson, 1944) to the
“Martinez Formation” in the vicinity of the city of Mar-
tinez, Contra Costa County, northern California (Figure
1). The history of how early workers referred to this
originally poorly defined “formation” has been summa-
rized by Mallory (1959). In this present study, we follow
the work of Weaver (1953), who refined the stratigraphy
of the Paleocene and Eocene formations in the vicinity of
the area where the “Martinez group” was first estab-
lished. He established new stratigraphic names, and the
rocks that pertain to the type locality of M. martinezensis
belong in his Paleocene Vine Hill Sandstone.
Mesalia clarki is only known from two locales: 1) its
type locality (UCMP loc. 1540, see “Appendix”) on the
north flank of Mount Diablo, Contra Costa County,
northern California, and 2) from the Santa Monica
Mountains, Los Angeles County, southern California
(Figure 1). Its type locality is near the site of Stewartville,
and approximately 24 km east-southeast of the city of
Martinez, and, according to Dickerson (1914a: 74), this
locality is “300 to 400 ft. above the base of the Martinez
Figure 1. Locales and latitudinal distribution of the study in hard, gray-green glauconitic sandstone.” Numerous
area gastropods. mollusks have been found at this locality (Dickerson,
1914a: 75). They consist of turritellid and buccinid gas-
tropods, crassatellid bivalves, and other shallow-marine
STRATIGRAPHY AND DEPOSITIONAL species, all indicative of normal marine salinities. Turri-
ENVIRONMENTS tella infragranulata pacheocensis Stanton, 1896, which is
found at this locality, is indicative of the upper middle
The geologic ages of the formations and most of the
Paleocene (upper Selandian) (Figure 2). On the geologic
depositional environments of the formations containing
map of Brabb et al. (1971), the locality plots within the
the two studied species are mentioned in Squires (1997).
Mesalia martinezensis is widespread in the study area glauconitic sandstone lower member of the “Martinez”
(Figure 1) and always found in siltstone or sandstone Formation. Megafossils are generally scarce in the “Mar-
beds that formed either as storm accumulations of mol- tinez” Formation in the vicinity of this type locality (E.
lusk-rich assemblages in shelfal-marine depths or as lo- Brabb, personal commun.), thus, it seems plausible that
calized displaced shallow-marine mollusks in deeper the fossils probably occur in storm-derived, isolated
lenses.
Mesalia clarki in the Santa Susana Formation in the
Palisades Highlands area of the Santa Monica Moun-
tains, southern California, is always found near outcrops
of coralline-algal limestone. Hoots (1931: 91–92, 133–
134, pl. 19B) reported that these limestones are resistant,
can be cliff forming, weather white, are nodular and ir-
regularly bedded, up to 35 m thick, up to 1200 m in
lateral extent, and commonly terminate in an abrupt wall.
Additional geologic and/or paleontologic details concern-
ing these limestones are mentioned in Strathearn et al.
(1988), Colburn (1996), Squires and Saul (1998), Squires
and Kennedy (1998), and Squires and Saul (2001).
At LACMIP loc. 10508, in the Palisades Highlands
area, abundant specimens of M. clarki are found in a thin
Figure 2. Chronostratigraphic position of the study area taxa. coralline-algal-rich micaceous muddy siltstone bed about
Ages of stage boundaries from Gradstein et al. (2004). Turri- 1 m stratigraphically below a blocky, coralline-algal-
tella zones from Saul (1983b). limestone interval approximately 24 m thick. Also found
R. L. Squires and L. R. Saul, 2007 Page 3
in this bed is the large neritid gastropod Corsania (Jan- onymized J. Müller’s specimens and G. Müller’s speci-
uncia) rhoga Saul and Squires, 1997, as well as the bi- mens with their Kassenberg quarry specimens. We be-
valves Plicatula lapidicina Squires and Saul, 1998, and lieve, however, that Kiel and Bandel’s Cenomanian
Plicatula trailerensis Squires and Saul, 1998. Occurring specimens have nothing to do with T. multilineata and
in nearby beds in close association with the coralline- represent, at best, a very questionable Mesalia. More
algal deposits are the gastropods Terebralia susana specimens of this possible Mesalia from the Cenomanian
Squires and Kennedy, 1998, and Campanile greenellum of Germany are needed in order to determine its generic
Hanna and Hertlein, 1939. All of these aforementioned assignment.
mollusks indicate very nearshore, tropical to subtropical Cossmann (1912: 126) reported the chronologic range
conditions (Squires and Saul, 1997; Squires and of Mesalia to be Late Cretaceous (Turonian) to Recent,
Kennedy, 1998). The latter workers concluded that the as did Wenz (1939), who apparently simply reiterated
coralline-algal limestones, like those at locality 10508, Cossmann’s findings. We were unable, however, to cor-
were deposited in a protected bay (no deeper than 40 to roborate any of Cossmann’s Cretaceous occurrences. He
70 m) with warm-algal-limestone buildups associated reported Mesalia gazellensis Whitfield (1891: 424, pl. 9,
with shoals on the bay floor. These limestone buildups fig. 10) as being from the Turonian of Syria, but the
are very similar in lithology and sedimentologic/tectonic aperture of this species is unknown. In addition, the
setting to limestones in the Paleocene Sepultura and Ba- sculpture is obsolete, which is unlike Mesalia.
hia Ballenas formations in northern Baja California (Ab- Cossmann (1912: 126) listed five Mesalia species of
bott et al., 1995), as well as similar to limestones in the Late Cretaceous (Senonian) age, and these are discussed
upper Paleocene to lower middle Eocene Sierra Blanca in the following sentences. Arcotia indica Stoliczka
Limestone in Santa Barbara County, southern California. (1868: 215, 469, pl. 16, figs. 12, 12a; pl. 19, fig 6) from
These limestones were deposited when tectonic plate- southern India is not a Mesalia. This species is also dis-
edge strain in the fore-arc basin caused local basement cussed later under “Systematic Paleontology.” Specimens
highs to form within the otherwise deeper marine envi- of Turritella ventricosa Forbes (1846: 123, pl. 13, fig. 3;
ronment (Whidden et al., 1995; Abbott et al., 1995). It is Stoliczka, 1868: 227, pl. 17, fig. 15) from southern India
likely that the Santa Susana Formation coralline-algal are missing the aperture. Turritella martinezensis Gabb
limestones formed under similar conditions. (1869: 159, pl. 28, fig. 51) from California is not of Cre-
Although Mesalia clarki and Mesalia martinezensis taceous age. Mesalia nettoana White (1887: 164–165, p.
both occur in the Santa Susana Formation in the Santa 18, figs. 3, 4) from the Maria Farinha beds in Brazil is
Monica Mountains, southern California, they never oc- Paleocene age. Mesalia hebe White (1887: 165, pl. 18, fig.
cur together in the same beds. Mesalia martinezensis is 5), also from Brazil, looks like a juvenile specimen of M.
not associated with the coralline-algal facies there or any- nettoana.
where else in the study area. Cossmann (1912: 126) also listed two Late Cretaceous
(Maastrichtian) species. The first one is Mesalia jovisam-
monis (Quaas, 1902: 256, pl. 26, figs. 18–20), which
PALEOBIOGEOGRAPHIC IMPLICATIONS Quaas reported, in a very generalized way, to be associ-
ated with the Exogyra overwegi biozone at Ammonite
Kiel and Bandel (2004: 120, fig. 7I) reported two speci- Hill in the Great Sand Sea in western Egypt. This bio-
mens of Mesalia cf. multilineata (J. Müller, 1851) from zone can also be recognized in the Maastrichtian (but not
Cenomanian strata at the Kasssenberg quarry in Ger- latest Maastrichtian) part of the Ammonite Hill Member
many. If these specimens actually belong to Mesalia, they of the Campanian to Paleocene Dakhla Formation in
would be the geologically oldest. The conical-turriculate western Egypt (Barthel and Herrmann-Degen, 1981).
shell with convex whorls bearing strong spiral ribs does Tantawy et al. (2001) assigned this member an early to
resemble Mesalia, but there are no specimens with an early late Maastrichtian age, based on planktic forami-
intact aperture nor with a protoconch. Turritella multi- nifera, calcareous nannofossils, and macrofossils. They
lineata J. Müller (1851: 29, pl. 4, figs. 4, 6) was originally also determined, however, that the entire formation
reported from the lower Campanian Aachen strata of ranges in age from early Maastrichtian to early Danian.
Germany, thus, it is considerably younger than the Kass- Immediately above the widespread K/T disconformity in
senberg quarry material. Turritella multilineata was also the region, a sedimentologically complex sequence
figured by G. Müller (1898: 97, pl. 13, figs. 4, 5), who marks the lower Danian Bir Abu Minquar horizon, which
reported it from middle Santonian to lower Campanian contains a mixture of Maastrichtian (reworked) and Da-
strata at Braunschweig/Ilsede, Germany. There is also a nian fossils, including both microfossil and macrofossil
mention of T. multilineata in Kollmann and Odin (2001: species (e.g., including some ammonites.). Unfortu-
441), and they also consider this Campanian species to nately, Quaas did not provide any information as to
belong to genus Turritella. The pleural angle of J. where exactly in the stratigraphic section he collected the
Müller’s figure is much narrower than that of Kiel and specimens of M. jovisammonis. His specimens were lost,
Bandel (2004), and in our opinion, Kiel and Bandel’s M. so it is not possible to match their rock matrix to actual
cf. M. multilineata (J. Müller) does not look like J. outcrops. Recollecting of this gastropod is necessary to
Müller’s species. Kiel and Bandel (2004) tentatively syn- decipher its exact geologic age.
Figures 3–15. Type species of Mesalia and Sigmesalia, plus comparative pictures of Mesalia solida (Deshayes, 1861). Specimens
coated with ammonium chloride. 3–9. Mesalia mesal (Adanson, 1757), Baie de Hann, Senegal, West Africa (Recent). 3–7.
Hypotype LACM 173163. 3. Apertural view, height 45 mm, diameter 15 mm. 4. Tip of specimen shown in previous figure, height
14 mm, diameter 5.5 mm. 5. Protoconch and earliest spire whorls, apertural/ right-lateral view, height 1 mm, diameter 0.7 mm. 6.
Base, diameter 15.6 mm. 7. Abapertural view, height 45 mm, diameter 15 mm. 8–9. Hypotype LACM 173164. 8. Oblique apertural
view, height 51 mm, diameter 16.3 mm. 9. Close-up of abapertural view of last whorl, diameter 17.2 mm. 10–13. Mesalia koeneni
(Le Renard, 1994), LACMIP hypotype 13397, Grignon, Paris Basin, France (middle Eocene, Lutetian Stage), height 45.8 mm,
diameter 20.3 mm. 10. Apertural view. 11. Close-up of aperture. 12. Right-lateral view (outer lip broken). 13. Base. 14–15. Mesalia
solida (Deshayes, 1861), hypotype LACMIP 13398, Le Guépelle, Paris Basin, France (late Eocene). 14. Apertural view, height 21.6
mm, diameter 8.1 mm. 15. Protoconch and earliest spire whorls, apertural to slightly right-lateral view, height 1 mm, diameter 0.7
mm.
Abbass (1963: 39–40, pl. 2, figs. 20–22) illustrated M. and Turner, 1942: card 110); Mesalia hardemanensis
jovisammonis from eastern Egypt and referred to it as (Gabb, 1860: 392, pl. 68, fig. 15; Stenzel and Turner,
Mesalia (Woodsalia) jovisammonis of Maastrichtian age. 1942: card 116); and Mesalia pumila (Gabb, 1860: 392,
He did not provide, however, any discussion as to how pl. 68, fig. 14; Stenzel and Turner, 1942: card 118).
this age was determined. In summary, our search of the literature revealed that
Mesalia cf. M. multisulcata (Lamarck 1804), reported Mesalia most likely originated during either the Maas-
by Serra (1937: 313–315, pl. 16, figs. 12, 12a, 13) from trichtian in northern Africa or the early Paleocene (Da-
near Tripoli, Libya, looks like it might be conspecific nian) in northern Africa and western Iran. During the
with Mesalia jovisammonis. Serra provided no detailed Danian it spread quickly to the Gulf Coast of the United
stratigraphic or geologic age information. States by means of westward-flowing ocean currents
The second species that Cossmann (1912) listed from emanating from the western Tethyan region. These cur-
the Maastrichtian is Mesalia fasciata (Lamarck, 1804) rents, which existed during the Late Cretaceous (Gor-
from Iran. Cossmann based this occurrence on work by don, 1973; Johnson, 1999) and continued into the Paleo-
Douvillé (1904: 329–330, pl. 47, figs. 23–27), who re- cene and Eocene (Saul, 1986; Squires, 1987), were part
ported M. fasciata from the “Couches à Cérithes” beds in of a circumglobal-tropical current that contributed to a
the Luristan region in west-central Iran. Douvillé (1904) widespread dispersal of marine biota (Haq, 1981). By the
believed that these Iranian specimens of M. fasciata, late Danian, it reached California and northern Baja
whose type locality is in middle Eocene (Lutetian Stage) California, Mexico, as well as Belgium (Cox, 1930; Glib-
strata at Grignon in Paris Basin, France (Eames, 1952: ert, 1973). By middle Paleocene, it reached Greenland
34), are of Maastrichtian age, but the “Couches a (Kollmann and Peel, 1983), and by the late Paleocene, it
reached Nigeria (Adegoke, 1977). During the Paleocene
Cérithes” beds contain the bivalve “Cardita” beaumonti
and Eocene, Mesalia reached its peak diversity and be-
d’Archiac and Haime, 1854, which is diagnostic of earli-
came most widespread, with occurrences mainly in the
est Danian age in Iran and Pakistan (Douvillé, 1928;
Old World western Tethys Sea region. We did not de-
Eames, 1952; Davies, 1975). Mesalia fasciata is long-
tect, however, any reported occurrences in Australia,
ranged geologically (early Paleocene to late Eocene) and New Zealand, Japan, or Antarctica. The Paris Basin of
widespread geographically (western Europe to Pakistan) France (see Cossmann and Pissarro, 1910–1913), south-
(Eames, 1952). western Nigeria (Adegoke, 1977), and the Gulf Coast of
Another Mesalia that needs investigation as to its pre- the United States (Stenzel and Turner, 1940, 1942;
cise stratigraphic position and geologic age is Mesalia Palmer and Brann, 1966) are the principal areas in which
foucheri Pervinquière (1912: 44, pl. 3, figs. 6–15), from numerous species of Mesalia have been recognized.
the Ghadames (Garat Temblili) region in Tunisia, north- Some species became very widespread. For example, as
ern Africa. Pervinquière (1912: 336) reported the species mentioned earlier, Mesalia fasciata ranged from the
as being of Maastricthian age, but no critical geologic Paris Basin, France to Pakistan (Eames, 1952). After the
details are provided. He did differentiate between Maas- warm greenhouse conditions that existed during the
trichtian and Danian fossils; thus, like in nearby Egypt Eocene, Mesalia was much reduced in its distribution
and Libya, the stratigraphic section containing M. and mainly occurred in what is now the Mediterranean
foucheri and other macrofossils in Tunisia, also spans the Sea region (Cossmann, 1912).
K/T boundary.
Two species of so-called Woodsalia Olsson, 1929, from
Upper Cretaceous (Campanian?) strata in northwestern MODERN MESALIA
Peru (Olsson, 1944) might eventually be placed in Me-
salia, once their apertures become known. They are: Mesalia is extant and review of the scant literature, as
Woodsalia paitana Olsson (1944: 69–70, pl. 11, fig. 5) well as use of the internet (note: <http://www.alboranshells
and Woodsalia paitana robusta Olsson (1944: 70, pl. 11, .com/turritellidae> was particularly helpful), revealed as
figs. 3, 9). many as possibly six species. The they are the following:
The so-called Mesalia (Mesalia) mauryae Allison M. mesal (Adanson, 1757), M. brevialis Lamarck, 1822;
(1955: 414–415, pl. 41, fig. 3; Perrilliat, 1989: 149, fig. M. varia Kiener, 1843; M. opalina Adams and Reeve,
51h) from the upper Aptian upper member of the Alisi- 1850; M. freytagi von Maltzan, 1884; and M. flammifera
tos Formation, Punta China region, Baja California, Locard, 1897, which includes the subspecies M. flam-
Mexico, is, according to Squires and Saul (2006), Turri- mifera flammifera Locard, 1897 and M. flammifera sim-
tella seriatimgranulata Roemer, 1849. plex Locard, 1897. There is much confusion as to exactly
In addition to the above-mentioned Old World Danian how many species there are, and potential synonyms
species of Mesalia, three New World Danian species are need to be resolved. For example, some workers (e.g.,
known from the Gulf Coast of the southeastern United Smith, 1915; Bowles, 1939) equated M. mesal with M.
States. They are from the Clayton Formation (Palmer brevialis, but other workers (e.g., Advovini and Cos-
and Brann, 1966), which is of earliest Danian age (Dock- signani, 2004) separated them. Bowles (1939) gave a
ery, 1986). The three species are: Mesalia allentonensis thorough review of the nomenclatural history of Mesalia
(Aldrich, 1894: 246–247, pl. 13, figs. 4a, 4b, 6; Stenzel brevialis.
A comprehensive malacological study of the modern whereas the modern occurrences have contracted pri-
species of Mesalia is greatly needed. Because of the un- marily to the Iberian Peninsula, Alboran Sea, and north-
certainties stemming from the poorly known systematics, western Africa.
it is confusing to try to determine which species is found
where. We were able to establish with certainty (see SYSTEMATIC PALEONTOLOGY
references below), however, that modern Mesalia is only
Superfamily Cerithioidea Fleming, 1822
found in the Atlantic coastal areas of southern Portugal,
Family Turritellidae Lovén, 1847
southwestern Spain, Morocco, Canary Islands, Western
Sahara, Mauritania, Senegal, and Guinea, as well as in Discussion: Allmon (1996: 9–12, table 1) thoroughly
the westernmost Mediterranean Sea, particularly in the reviewed the history of the classification of turritellid
Alboran Sea (i.e., Strait of Gibraltar to southern Spain on gastropods and listed the five subfamilies and all the
the north and Morocco on the south) and the Aegean Sea genera/subgenera within each subfamily. These subfami-
region of western Turkey. lies are: Turritellinae Lóven, 1847; Protominae Marwick,
Mesalia mesal and M. brevialis have the widest distri- 1957; Pareorinae Finlay and Marwick, 1937; Vermicu-
bution. Mesalia mesal has been reported from the Al- lariinae Lamarck, 1799; and Turritellopsinae Marwick,
garve region of southern Portugal, the Algeciras region of 1957. Bouchet et al. (2005) included the first four of
southwestern Spain, and the Alboran Sea (Poppe and these subfamilies, but removed Turritellopsinae. Instead,
Goto, 1991), the Canary Islands (Macedo and Borges, they included subfamily Orectospirinae Habe, 1955.
1999), Senegal (Bouchet, 1977; Ardovini and Cossignani, Subfamily Pareorinae Finlay and Marwick, 1937
2004), and western Turkey (Demir, 2003). Mesalia bre- Discussion: Pareorine turritellids are characterized
vialis has been reported from the Algarve region of from the other subfamilies of family Turritellidae by hav-
southern Portugal (Afonso et al., 2000; Alves et al., 2003), ing an aperture obliquely effuse over the anterior end of
southwestern Spain and the Alboran Sea (Hidalgo, the columella and forming a sinus (short spout), with the
1917), Senegal (Ardovini and Cossignani, 2004), and adapical margin of the sinus usually making a spiral ridge
Guinea (Pasteur-Humbert, 1962). Mesalia opalina has on the columella (Marwick, 1957).
been reported from the Canary Islands and Morocco Mesalia can be readily identified if its aperture is in-
(Poppe and Goto, 1991), as well as from Mauritania (Ar- tact, but when it is missing, workers have commonly
dovini and Cossignani, 2004). The other modern species/ misassigned it to the similar looking genus “Turritella”
subspecies of Mesalia are apparently restricted to the Lamarck, 1799, sensu lato, a group comprising at least 35
northwestern coast of Africa (Ardovini and Cossignani, genera and subgenera names (Allmon, 1996), all of which
2004). are turritellines whose apertures do not have a sinus
Mesalia melanioides Reeve, 1849, was reported (short spout) at the anterior end of the aperture nor have
(Smith, 1915) to be from West Australia, but this species a spiral ridge on the columella. In addition, according to
is now the type species of Neodiastoma Cotton, 1932, Smith (1915), the corneous operculum of Mesalia is pau-
which differs from Mesalia by having axial sculpture on cispiral and not multispiral, as in “Turritella,” but this
the early spire. Marwick (1957) summarized the system- later distinction is not useful when studying fossil spe-
atics of Neodiastoma and classified it as a pareorine. cies.
Mesalia is found today on both muddy and sandy bot- Ten pareorine genera were listed by Marwick (1957),
toms in coastal waters ranging in depth from lower in- who also provided illustrations of the growth-line traces
tertidal to 20 m (Hidalgo, 1917; Pasteur-Humbert, 1962; of some of these genera. Comparative information about
Poppe and Goto, 1991; Afonso et al., 2000; Demir, 2003; the stratigraphic range, growth-line details, whorl profile,
Alves et al., 2003). Bouchet (1977) reported that M. me- and protoconch shape of most of these genera was given
sal, although not common there, can be found in the by Allmon (1996: table 1).
seaward part of mangrove-swamp systems along the coast Genus Mesalia Gray (nomen nudum, 1840), 1847
of Senegal. Specimen LACM 17316 (Figures 3–7) of M.
mesal was collected in approximately 5 m depth, on sand Type Species: Cerithium mesal Adanson, 1757 [=Tur-
and rubble in Senegal. ritella mesal Deshayes, 1843], by original designation;
Large numbers of M. mesal have been reported Recent, southern Portugal, southwestern Spain, Alboran
(Afonso et al., 2000) as almost always being partially in- Sea, Canary Islands, Senegal, and western Turkey.
faunal (with their apices pointed upward) when found on Description: Small to large (up to approximately 95
exposed low-tide mud flats on the inner lagoon sides of mm shell height), turritelliform, slender to conical ro-
islands within the Rio Formosa coastal-lagoon system of tund. Pleural angle ranges from 15° to 41°. Protoconch
southern Portugal. conical, small, smooth, and approximately two whorls.
The modern ecological parameters mentioned above Protoconch to teleoconch transition gradual. Teleoconch
are not totally reliable for fossil Mesalia because prefer- whorls up to 16, whorl sides convex to flattish/concave.
ences for substrate and depth of water might have pos- Sculpture on early juvenile teleoconch whorls variable,
sibly changed over time. In addition, the fossil occur- ranging from nearly smooth or with very fine, unicostate,
rences of Mesalia had a pan-Tethyan distribution, bicostate, or tricostate spiral lirae; sculpture on adult
whorls highly variable, ranging from smooth to numer- species of Mesalia are illustrated in Figures 3–9, and
ous, weak to moderately strong closely spaced spiral ribs, various views of a representative specimen of the type
but less commonly with fewer and more prominent spiral species of Sigmesalia are illustrated in Figures 10–13. Its
ribs. Growth lines parasigmoidal on last whorl (including type species, Sigmesalia koeneni Le Renard, 1994 [new
base); lateral sinus variable in amount of concavity (flex- name for Turritella sulcata Lamarck, 1804 (original des-
ure). Aperture with shallow effuse spout, ranging from ignation), non Bosc, 1801], is of middle Eocene (Lu-
somewhat constrained to broad. Adapical edge of spout tetian) age and from Paris Basin, France. Finlay and
usually forms weak spiral ridge that continues onto col- Marwick (1937) stated that the aperture and growth lines
umella. of the type species of Mesalia seem to be generically
different than those of the Paris Basin shells, but they did
Discussion: Mesaliopsis Thiele, 1929 [type species: not provide any details. Davies (1971: 312, figs. 677a,
Mesalia opalina (Adams and Reeve, 1850)], Recent, was 677b) mentioned that the growth lines of Sigmesalia
reported by Wenz (1939) to be a subgenus of Mesalia, have a more flexed outer lip sinus than does Mesalia.
but future work might show it to be synonymous with According to Marwick (1957: 163), Sigmesalia differs
Mesalia. from Mesalia by usually having a wider pleural angle.
Mesalia somewhat resembles Lithotrochus Conrad, The following paragraphs deal with our observations con-
1855, of Jurassic age from Chile, South America. Coss- cerning these proposed diagnostic features of Sigmesalia.
mann (1912: 125) reported Lithotrochus to be a junior Inspection of representative specimens of several of
synonym of Mesalia, but Wenz (1938: 280, fig. 596) and the Eocene Paris Basin species, including the type spe-
Cox (1960: I248–I249, fig. 159,11) believed Lithotrochus
cies of Sigmesalia, stored in the LACMIP collection, as
to be a trochid. It is an extraordinarly large gastropod
well as inspection of photographs of 17 Paris Basin spe-
(height 125 mm) with a wide pleural angle, domed upper
cies (see Cossmann and Pissarro, 1910–1913: pl. 21, figs.
spire, turritelliform shape, anteriorly carinate whorls, and
relatively few spiral ribs. Details of its aperture are un- 126–1 to 126–15), revealed variability in the shape of the
known. aperture of Sigmesalia. For example, the aperture of Sig-
Cossmann (1912: 125) also reported Lithotrochus to mesalia incerta (Deshayes, 1832; Cossmann and Pissarro,
be a junior synonym of Arcotia Stoliczka, 1868, whose 1910–1913: pl. 21: fig. 126–4, two views) is similar to that
type species, Arcotia indica Stoliczka (1868: 215, 469, pl. of M. mesal, in that the spout is broad and not well
16, figs. 12, 12a; pl. 19, fig. 6) is from Upper Cretaceous constrained. The aperture of Sigmesalia koeneni how-
(Trichinopoly Group) strata near the town of Alundan- ever, is better developed (Cossmann and Pissarro, 1910–
apooram, southern India. According to Sundaram et al. 1913: pl. 21, fig. 126–15).
(2001: fig. 3), this town’s name is also referred to as The amount of flexure of the outer lip sinus is basically
Alundalippur and, from information in their map, this similar in all the Eocene Paris Basin species, although
town is underlain by the Kulakkalnattam Formation of Mesalia solida (Deshayes, 1861) does show some vari-
Turonian age. Wenz (1939) synonymized Arcotia with ability. The amount of flexure of this feature is variable in
Mesalia. Finlay and Marwick (1937) reviewed the mor- M. mesal and can be similar to the amount seen on
phology of Arcotia and reported that, based on its Eocene Paris Basin species. Variability in the amount of
straight growth lines and open umbilicus, this genus is flexure for both the Eocene and modern Mesalia shells
not a synonym of Mesalia. They reported, furthermore, can also occur in proximity of growth checks and break-
that Aroctia appears to be a mathildid. Bandel (2000) ages of the outer lip incurred during the life of the gas-
came to the same conclusion. tropod.
Mesalia is similar to the pareorine Woodsalia Olsson The pleural angle of the Eocene Paris Basin shells is
1929, whose type species, Woodsalia negritosensis Ols- quite variable, ranging from 21° to 41°, but the low end
son (1929: 13–15, pl. 4, figs. 5, 6) is from lower Eocene of this range [e.g., Mesalia ecki (Laubrière, 1881; Coss-
rocks in northwestern Peru. Woods (1922: 78–79, pl. 7, mann and Pissarro, 1910–1913: pl. 21, fig. 126–2)] is very
figs. 5–7; pl. 8, figs. 1–3) and Wenz (1939: 651, fig. 1852, close to the value (16° to 18°) on M. mesal. Sigmesalia
two views) also illustrated this species. The full shape of koeneni has one of widest pleural angles (41°; see Figures
the aperture of this Peruvian gastropod, however, is not 10 and 12). For comparative purposes, an illustration
known. (Figure 14) is provided for Sigmesalia solida. It has a
Genus Sigmesalia Finlay and Marwick, 1937, was relatively narrow pleural angle of 25°, more like that
originally erected to accommodate a group of Eocene found on M. mesal (Figures 3, 4, and 7).
gastropods from the Paris Basin, France that were pre- Other morphologic features that are variable on the
viously identified as Mesalia. There has been no consen- Eocene Paris Basin shells are strength and number of
sus as to whether or not Sigmesalia is a distinct genus. spiral ribs, pattern of development of sculpture on the
Marwick (1957) reported it to be a separate genus, as did early juvenile teleoconch whorls, and degree of indenta-
Le Renard (1994). Eames (1952) reported it to be a tion of the suture. Even the strength of the spiral ridge
subgenus of Mesalia, and Palmer and Brann (1966) re- (Figure 10) on the columella is variable. Mesalia mesal pos-
ported Sigmesalia to be synonymous with Mesalia. sesses a spiral ridge on the columella, as do most specimens
Various views of representative specimens of the type of Sigmesalia koeneni (compare Figures 8 and 11).
A few species of Sigmesalia are similar to Mesalia in quite prominent) on lower spire. Last whorl with three
having a relatively narrow pleural angle and bicostate primaries, both posterior and anterior to carina. Base
sculpture on the early juvenile whorls but not on the with three secondaries, anteriormost one weak; ribs ob-
adult whorls. They are the following: Sigmesalia instabi- solete on short neck. Aperture relatively small, D-
lis (Briart and Cornet, 1873: 86, pl. 12, figs. 9a–9b) of shaped; columella relatively broad, smooth; spout effuse
early Paleocene (Danian) age from Belgium; Sigmesalia and short with anterior end projecting slightly; growth-
salvani (Adegoke, 1977: 86–88, pl. 14, figs. 10–16) of line trace of last whorl (including base) parasigmoidal,
Paleocene age from Nigeria; and Sigmesalia fasciata with lateral sinus flexure strongest in vicinity of carina.
(Lamarck, 1804: 217) from Eocene strata in France, Bel-
gium, and Pakistan (Cossmann and Pisarrro, 1910–1913; Holotype: ANSP 4344, height 57 mm, diameter 23
Cox, 1930; Eames, 1952); Sigmesalia pagoda (Cox, 1930: mm.
160–161, pl. 18, figs. 6a–b, 7a–b) from Eocene strata in Type Locality: Martinez, northern California (details
Pakistan; Sigmesalia biplicata (Bowles, 1939: 328, pl. 34, not given).
figs. 6, 8) from Paleocene strata in Alabama; and Sigme-
salia gomin (Bowles, 1939: 326–327, pl. 33, fig. 9) from Geologic Range: Late early Paleocene to early late
Paleocene strata in South Carolina. Paleocene (near the Danian-Selandian boundary to early
The protoconchs of Mesalia mesal and Sigmesalia Thanetian).
solida are very similar (compare Figures 5 and 15); both
are small, smooth, have essentially the same shape, and Distribution: DANIAN = Turritella peninsularis qua-
the transition to the teleoconch is gradual. leyi Zone: lower San Francisquito Formation, Warm
In summary, we found that the morphologic features Springs Mountain, Los Angeles County, southern Cali-
of the Eocene Paris Basin shells are variable. We could fornia (new stratigraphic occurrence, LACMIP loc.
find no reliable, constant morphologic characters to dis- 21581). NEAR THE DANIAN-SELANDIAN BOUND-
tinguish Mesalia from Sigmesalia; hence, we regard them ARY = Turritella peninsularis qualeyi Zone transitional
as congeneric. with Turritella peninsularis Zone: Martinez Formation,
Herndon Creek east of Lower Lake, Lake County, north-
Mesalia martinezensis (Gabb, 1869) ern California (Stanton, 1896 [faunal list]; Dickerson,
(Figures 16–23) 1914a; Merriam, 1941); upper Las Virgenes Sandstone,
Simi Hills, Ventura County, southern California (Waring,
Turritella martinezensis Gabb, 1869: 169–170, 228, pl. 28, fig. 1917; Nelson, 1925 [faunal list]; Merriam, 1941; Zins-
51; Dickerson, 1914a: pl. 13, fig. 10; Waring, 1917: 100, pl. meister, 1983; Saul, 1983a). PROBABLY NEAR THE
14, fig. 5. DANIAN-SELANDIAN BOUNDARY: Reworked
Turritella maccreadyi Waring, 1914: 783; Waring, 1915: fig. 15 specimens in Santa Susana Formation, Poison Oak Can-
[not fig. 14]; Waring 1917: 87–88, pl. 12, fig. 10. yon, north side Simi Valley, Los Angeles County, south-
Mesalia maccreadyi (Waring).—Paredes-Mejia, 1989: 176–177, ern California (new stratigraphic occurrence, LACMIP
pl. 3, figs. 3–6. loc. 21554); Reworked specimens in Stokes Canyon
Mesalia martinezensis (Gabb).—Cossmann, 1912: 126; Stew-
art, 1927: 353–354, pl. 25, fig. 1; Schenck and Keen, 1940:
Breccia Member of the middle Miocene Calabasas For-
pl. 20, fig. 5; Merriam, 1941: 127–128, pl. 39, figs. 1–5, 7; mation, Stokes Canyon, Santa Monica Mountains, Ven-
Zinsmeister, 1974: 118–119, pl. 12, figs. 5, 9; Zinsmeister, tura County (new stratigraphic occurrence, LACMIP
1983: pl. 2, fig. 14; Paredes-Mejia, 1989: 173–176, pl. 3, loc. 25281). SELANDIAN = Turritella peninsularis
figs. 7–10; Saul, 1983a: text-fig. 2, pl. 1, fig. 2. Zone: Lower Vine Hill Sandstone, Martinez area, Contra
Mesalia clarki (Dickerson).—Zinsmeister, 1983: pl. 2, fig. 13. Costa County, northern California (Weaver, 1953 [faunal
list]); lower San Francisquito Formation, Pinyon Ridge
Description: Large (up to approximately 95 mm east of Big Rock Creek, Valymero area, Antelope Valley,
height). Turritelliform. Pleural angle approximately 20°. Los Angeles County, southern California (Dickerson,
Protoconch unknown. Teleoconch up to 12 whorls, in- 1914b [faunal list]; Merriam, 1941; Kooser, 1980 [faunal
creasing rapidly in size from the apex. Suture slightly list]); lower Santa Susana Formation (= “Martinez ma-
impressed. Sculpture consisting only of spiral ribs of dif- rine member” of Nelson, 1925 [faunal list]), Simi Hills,
fering strength but dominated by carina located anteri- Ventura County, southern California (Kew, 1923 [faunal
orly; ribs generally becoming stronger with growth; spiral list]; Nelson, 1925 [faunal list]; Zinsmeister, 1983; Saul,
threads on interspaces and on carina surface. Carina usu- 1983a). LOWER THANETIAN = Turritella infragranu-
ally strongly angulate but can be rounded or even sub- lata Zone: Upper Vine Hill Sandstone, Martinez area,
dued. Posterior to carina, several widely spaced spiral Contra Costa County, northern California (Weaver, 1953
ribs of variable strength occur, ranging from tertiaries to [faunal list]); upper Santa Susana Formation, Palisades
primaries: two ribs on uppermost spire, three to four on Highlands, Santa Monica Mountains, Los Angeles
upper spire, and one to three on lower spire. Anterior to County, southern California (new stratigraphic occur-
carina, several spiral ribs of variable strength occur, rang- rence, LACMIP locs. 7060 and 11717); Sepultura For-
ing from tertiaries to primaries: approximately five ribs mation, Mesa San Carlos, northern Baja California,
on upper spire and one to two ribs (both occasionally Mexico (Paredes-Mejia, 1989).
Figures 16–23. Mesalia martinezensis (Gabb, 1869). Specimens coated with ammonium chloride. 16–18. Hypotype LACMIP
13399, height 58 mm, diameter 25.4 mm. 16. Apertural view. 17. Oblique apertural view. 18. Abapertural view. 19. Hypotype
LACMIP 13400, LACMIP loc. 22557, apertural view, height 36.3 mm, diameter 14.1 mm. 20. Hypotype LACMIP 13401, LACMIP
loc. 21607, abapertural view, height 38.3 mm, diameter 19.3 mm. 21. Hypotype LACMIP 13402, LACMIP loc. 22698, abapertural
view, height 33.9 mm, diameter 21.6 mm. 22. Hypotype LACMIP 13403, LACMIP loc. 26897, apertural view, height 10.6 mm,
diameter 6.5 mm. 23. Hypotype LACMIP 13404, LACMIP loc. 22330, base, diameter 17.2 mm.
Discussion: The largest specimens of this species oc- shells misidentified by some workers as Turritella ma-
cur in the lower San Francisquito Formation, Pinyon creadyi Waring, 1914.
Ridge east of Big Rock Creek, Valymero area, Antelope The overall teleoconch morphology of the 10 mm-high
Valley, Los Angeles County, southern California. tip of Mesalia martinezensis superficially resembles that
There is considerable variability in the strength of the of the 15-mm high mathildid Carinathilda diminuata
spiral ribs on M. martinezensis. Most specimens are cari- (Perrilliat, Vega, and Corona, 2004) illustrated by Kiel et
nate on all whorls, including the last whorl. On some al. (2002: 329–330, fig. 2.4) from the lower Maastrichtian
specimens, however, the carina becomes weaker on the of the Mexcala Formation, Guerrero, southern Mexico.
later whorls as the other spiral ribs become stronger, Carinathilda diminuata is definitely a mathildid because
giving these whorls a convex shape (Figures 19–21), like it has a heterostrophic protoconch. The resemblance be-
tween these two gastropods, nevertheless, provides evi- strength primaries (anteriorly located ribs can be some-
dence that the Late Cretaceous mathildids and lower what angulate); spiral threads on all interspaces; poste-
Paleogene turritellids can have similar looking adult riormost rib part of broad band; ribs on anterior part of
shells. whorl tend to be slightly stronger than posteriorly located
Mesalia martinezensis resembles “Mesalia” virginiae ribs; base of last whorl with three ribs. Angulate whorl
Stilwell et al. (2004: 29–30, pl. 5, figs. 6–10) from lower shape: upper spire with one secondary on posterior part
Paleocene (Danian) rocks on Seymour Island, Antarctic and two (bicostate), well developed, flat-topped prima-
Peninsula, but M. martinezensis has a subtle effuse spout ries on anterior part; lower spire and last whorl with
rather than the longer and more distinct, twisted narrow three primaries on posterior part and two stronger pri-
anterior canal that “M.” virginiae possesses. In addition, maries, with one secondary in between each, on anterior
M. martinezensis has stronger ribs and a parasigmoidal part; spiral threads on all interspaces. Base (including
growth line, rather than an opisthocyrt one on the last short neck) of last whorl with approximately seven,
whorl. In our opinion, the aperture of “M.” virginiae is evenly spaced ribs; interspaces and ribs covered by spiral
unlike that of Mesalia. threads. Aperture small; columella narrow with thin cal-
Gabb (1869) mentioned that the broadly expanding lus, occasionally with single, weak fold, slight twist on
whorl of martinezensis approaches that seen on Turri- anterior end of columella. Spout effuse, short, and nar-
tella robusta Gabb (1864: 135, pl. 21, fig. 74; not = T. row. Growth-line trace on last whorl (including base)
(Haustator) robusta Grzybowski, 1899), but Merriam parasigmoidal, with lateral sinus flexure strongest medi-
(1941: 128) reported that the Late Cretaceous T. ro- ally.
busta, which is represented by a single poorly preserved Holotype: UCMP 11936, height 25 mm, diameter
type specimen, is probably not a Mesalia. This type speci- 16.5 mm.
men has an umbilicus, therefore it is not a turritellid. It
is from the Redding area, northern California, and not Type Locality: UCMP loc. 1540.
from Tuscan Springs, as erroneously reported by Mer-
riam (1941). Jones et al. (1978: pl. 1, fig. 19) identified Geologic Range: Late Paleocene = Turritella infra-
this specimen, which is of Turonian age, to be Glauco- granulata Zone.
nia? robusta (Gabb, 1864).
Distribution: “Martinez” Formation, northeast side of
Merriam (1941: 10, 116) stated that mainly in profile
Mount Diablo, Contra Costa County, northern Califor-
the Pacific slope Miocene Turritella temblorenesis
nia (Dickerson, 1914a; Merriam, 1941; Zinsmeister and
Wiedey, 1928, might readily be confused with Mesalia
Paredes-Mejia, 1988 [faunal list]; upper Santa Susana
martinezensis. The latter also resembles the Pacific slope
Formation, Trailer and Quarry canyons, Los Angeles
Miocene Turritella temblorensis tritschi Hertlein, 1928, County, Santa Monica Mountains, southern California
and Turritella ocoyana Conrad, 1857. The latter, how- (Strathearn et al., 1988 [faunal list]; Squires and Saul,
ever, has a different growth line. In addition, T. martin- 1998: 1025).
ezensis strongly resembles Turritella fredeai Hodson,
1926, of Miocene age from northern Colombia and Discussion: Mesalia clarki is abundant in the upper
northern Venezuela. None of these above-mentioned Santa Susana Formation at LACMIP loc. 10508, in the
Miocene species, however, has the effuse spout of Me- Santa Monica Mountains, Los Angeles County, southern
salia. California. The anterior ends of the shells are very frag-
ile, and nearly all the specimens have incomplete aper-
Mesalia clarki (Dickerson, 1914a) tures. None of the specimens has retained their proto-
(Figures 24–32) conch, and most specimens are missing their upper spire.
Growth lines are hard to discern, usually visible only on
a single whorl (typically the penultimate whorl), and
Turitella [sic] clarki Dickerson, 1914a: 142–143, pl. 13, fig. 8. were rarely preserved on the base of the last whorl. Some
Mesalia clarki (Dickerson).—Merriam, 1941: 128, pl. 39, fig. 6; of the specimens appear to have a wider pleural angle
Zinsmeister, 1983: table 1, pl. 2, fig. 14.
(23°) than normal, but these particular specimens have
been crushed. A few of the specimens (five percent) have
Description: Medium small (up to approximately 31 naticid boreholes, and a few other specimens are en-
mm height). Turritelliform. Pleural angle approximately crusted, in part, by bryozoans. Rare specimens are
21 to 22°. Protoconch unknown. Teleoconch up to 12 coated by calcareous algae.
whorls, consisting of two whorl shapes: flatish rounded All previous workers assigned Mesalia clarki to various
and anteriorly angulate. Sutural area indented. Sculpture genera without knowledge of the shape of the aperture.
consisting only of spiral ribs, variable in number, Our cleaning of representative specimens of Dickerson’s
strength, and spacing. Flattish to rounded whorl shape: species revealed it to have a short, shallow effuse spout
upper spire with one or two secondaries on posterior part (Figures 24–25) and bicostate sculpture on the juvenile
and two (bicostate) primaries on anterior part; lower whorls (Figure 30). There is considerable variation in the
spire and last whorl with seven to eight nearly equal sculpture and the shape of the whorls on M. clarki. Some
Figures 24–32. Mesalia clarki (Dickerson, 1914). Specimens coated with ammonium chloride. All from LACMIP loc. 10508. 24.
Hypotype LACMIP 13405, apertural view, height 21.5 mm, diameter, 19.6 mm. 25. Hypotype LACMIP 13406, slightly oblique
apertural view, height 23.1 mm, diameter 10.8 mm. 26–27. Hypotype LACMIP 13407, height 22.6 mm, diameter 9.4 mm. 26.
Abapertural view. 27. Oblique apertural view. 28. Hypotype LACMIP 13408, apertural view, height 23.2 mm, diameter 9.2 mm. 29.
Hypotype LACMIP 13409, abapertural view, height 21.5 mm, diameter 11 mm. 30. Hypotype LACMIP 13410, abapertural view,
height 23.6 mm, diameter 9.7 mm. 31. Hypotype LACMIP 13411, base, diameter 9.5 mm. 32. Hypotype LACMIP 13408, base of
same specimen shown in Figure 28, diameter 8.9 mm.
specimens have nearly uniform sculpture and flattish Zinsmeister (1983: pl. 2, fig. 14), Zinsmeister and
whorls (Figure 24), others have carinate whorls (Figure Paredes-Mejia (1988: table 1), and Paredes-Mejia (1989:
26), whereas others have uniform sculpture with convex table 3) reported M. clarki from the Santa Susana For-
whorls (Figure 29). mation in the Simi Hills, southern California. These re-
ports, however, were based on the misidentification of a Aldrich, T. H. 1894. The (Midway) Clayton Tertiary section
specimen of Mesalia martinezensis that happens to lack a and its fossils. In: E. Smith, J. Allen, C. Lawrence and
strong anterior carina on the otherwise convex whorls. D. W. Landgon, Jr., Report on the Geology of the Coastal
Mesalia clarki resembles Motyris aralica Plain of Alabama. Alabama Geological Survey, 759 pp.
Allison, E. C. 1955. Middle Cretaceous Gastropoda from Punta
(Michailovski, 1912; Wenz, 1939: 652, fig. 1856) from China, Baja California, Mexico. Journal of Paleontology
upper Eocene rocks in the Aral Sea region. Motyris 29: 400–432.
Eames, 1952, was formerly known as Tomyris Allmon, W. D. 1996. Systematics and evolution of Cenozoic
Michailovski, 1912. See Marwick (1957: 162–163) for American Turritellidae (Mollusca: Gastropoda) I: Paleo-
more taxonomic information about Motyris. Mesalia cene and Eocene coastal plain species related to “Turri-
clarki differs from M. aralica by not having tabulate tella mortoni Conrad” and “Turritella humerosa Conrad.”
whorls with strongly indented sutures. The full aperture Palaeontographica Americana 59: 1–134.
of M. aralica is unknown, and details about its apical Alves, F. L. Chícharo, A. Nogueira and J. Regala. 2003.
whorl development are wanting. The only other species Changes in benthic community structure due to clam
of Motyris that we are aware of is Motyris pseudoaralica dredging on the Algarve coast and the importance of sea-
sonal analysis. Journal of the Marine Biological Association
Eames (1952: 30–31, pl. 1, fig. 15; pl. 2, figs. 58a, b) from of the United Kingdom 83: 719–729.
Pakistan, but its aperture is unknown. We believe that d’Archiac, A. and H. Haime. 1854. Description des animaux
when the great variability of Mesalia is taken into ac- fossiles du groupe nummulitique de l’Inde. Guide et J.
count, Motyris will prove to be congeneric. Baury, Paris, pp. 225–373.
Ardovini, R. and T. Cossignani. 2004. West African seashells
(including Azores, Madeira and Canary Is.). Volume 2.
ACKNOWLEDGMENTS L’Informatore Piceno, Ancona, Italy, 319 pp.
Earl Brabb (U. S. Geological Survey, Menlo Park) pro- Bandel, K. 2000. Some gastropods from the Trichinopoly
Group Tamil Nadu, India and their relation to those from
vided very useful information regarding the stratigraphy
the American Gulf Coast. Geological Society of India,
of the beds in the vicinity of the type locality of Mesalia Memoir 46: 65–111.
clarki. Lindsey T. Groves (LACM, Malacology Section) Barthel, K.W. and W. Herrmann-Degen. 1981. Late Creta-
kindly provided key literature dealing with the ecology of ceous and early Tertiary stratigraphy in the Great Sand
modern Mesalia. The manuscript benefited from the re- Sea and its SE margins (Farafra and Dakhla oases), SW
views by Warren D. Allmon (Paleontological Research In- Desert, Egypt. Mitteilungen der Bayerischen Staats-
stitute, Ithaca, New York) and Steffen Kiel (University of sammlung für Paläontologie und historische Geologie 21:
Leeds, England and Department of Paleobiology, Smith- 141–182.
sonian Institution’s National Museum of Natural History). Bosc, L. A. G. 1801. Histoire naturelle des coquilles. In:
Steffen Kiel also provided us with an important hard-to- G. L. L. de Buffon (ed.) Histoire naturelle de Buffon, etc.,
nouvelle édition. Mollusca: Volume 4. Déterville, Paris,
find reference and very useful stratigraphic information.
280 pp.
Bouchet, P. 1977. Distribution des mollusques dans les man-
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Zinsmeister, W. J. and L. M. Paredes-Mejia. 1988. Paleocene “Meier Canyon,” Santa Susana Quadrangle (7.5 minute,
biogeography of the west coast of North America: A look 1951, photorevised 1969), south side of Simi Valley, Simi
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gene Symposium, Volume 58, pp. 9–22. 22557. Sandstone bed below small waterfall [dry] west of road
going south through Barclay Ranch, 10,162 ft. south and
5660 ft. wet of junction of Southern Pacific railroad and
APPENDIX Los Angeles Ave. about 0.25 mi. east of Santa Susana,
Santa Susana Quadrangle (7.5 minute, 1951, photorevised
1969), Simi Hills, Ventura County, southern California.
LOCALITIES CITED Paleocene. Santa Susana Formation. Coll.: M. Murphy,
Localities are LACMIP, unless otherwise noted. All spring, 1950.
quadrangle maps are U. S. Geological Survey maps. 22698. On first large ridge; trending southwest to west of ridge
trending south of hill 2150. Bearing from the northwest
7060. Elevation 1427 ft., on ridge between Temesal and Santa corner of the Calabasas Quadrangle is S14°E; distance
Ynez canyons at edge of fire road on top of ridge, Topanga 12,210 ft., Calabasas Quadrangle (7.5 minute, 1952, pho-
Canyon Quadrangle (7.5 minute, 1952, photorevised torevised 1967), Simi Hills, Ventura County, southern
1981), Los Angeles County, southern California. Pale- California. Paleocene. Santa Susana Formation. Coll.: J. H.
ocene. Santa Susana Formation. Coll.: H. D. B. Wilson, Fantozzi, June 1, 1951.
June 1, 1941. 25281. Sandstone at elevation of 1000 ft., about 400 ft. south
10508. North slope of Trailer Canyon near top of ridge be- and 1000 ft. west of northeast corner of section 5, T. 1 S,
tween Quarry and Trailer canyons at approximately 1325 R. 17 W, Malibu Beach Quadrangle (7.5 minute, 1950,
ft. elevation and just west of saddle, just below coralline- photorevised 1967), on west side of northern tributary to
algal beds in limy siltstone west of small fault, road cut Stokes Canyon, western Santa Monica Mountains, Los An-
north side of unpaved road 5600 ft. north of San Vicente geles County, southern California. Reworked Paleocene
y Santa Monica Grant boundary, 10,400 ft. east of Los (Selandian) fossils in middle Miocene Calabasas Forma-
Angeles City boundary, Topanga Quadrangle (7.5 minute, tion, Stokes Canyon Breccia Member. Coll.: J. Stark and
1952, photorevised 1981), east of Santa Ynez Canyon, Pali- T. Susuki family, May 5, 1965.
sades Highlands, Santa Monica Mountains, Los Angeles 26897. Gully west side of Temesal Canyon opposite 2nd ‘e’ of
County, southern California. Lower upper Paleocene Temesal at about 1475 ft. elevation; approximately 1082 m
(lower Thanetian). Santa Susana Formation. Coll.: G. (3550 ft.) south; 533m (1750 ft.) east of hill 22036; San
Strathearn and others, fall, 1982. Vincente and Santa Monica Grant, Topanga Quadrangle
11717. Float at about 1600 ft. elevation in bottom of south- (7.5 minute, 1952, photorevised 1967), Santa Monica
flowing gully joining Quarry Canyon at about 1410 ft. el- Mountains, Los Angeles County, southern California.
evation; 1500 ft. SW of hill 2036, Topanga Quadrangle (7.5 Middle upper Paleocene (middle Thanetian). Santa Su-
minute, 1952, photorevised 1981), Los Angeles County, sana Formation. Coll.: J. M. Alderson, March 9, 1980.
southern California. Paleocene. Santa Susana Formation. UCMP 1540. Elevation 1000 ft., 1 mi. south of Stewartville
Coll.: J. M. Alderson, November 11, 1980. (site), northeast corner of NW 1/4, section 15, T. 1 N,
21581. Black nodular shale and conglomerate on road 1.1 mi. R. 1 E, Antioch South Quadrangle (7.5 minute, 1973, pho-
east from Cienaga Camp at Fish Canyon forks toward torevised), 300 ft. south of basal Tejon conglomerate and
Warm Springs summit; on northwest side of ravine; north 600 ft. north of Chico-Martinez contact, northeast side of
side East Fork Fish Canyon, T. 6 N, R. 16 W, approximately Mount Diablo, Contra Costa County, northern California.
2050 ft. north, 750 ft. east of bench mark 2205, Warm Upper middle Paleocene (Selandian) = Turritella infra-
Springs Mountain Quadrangle (7.5 minute, 1958, photo- granulata pachecoensis Zone. “Martinez” Formation,
revised 1974), Los Angeles County, southern California. Pa- lower member. Coll.: R. E. Dickerson, circa 1912.

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Paleocene Pareorine Turritellid Gastropods From The Pacific Slope of North America

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THE NAUTILUS 121(1):1–16, 2007 Page 1

Paleocene pareorine turritellid gastropods from the Pacific slope

ABSTRACT (e.g., Zinsmeister, 1983) have generally placed both spe-

depths. These mollusks commonly include shallow-

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