Miocene Foraminifera of South-Central Chile

Miocene foraminifera from the
south-central coast of Chile

Kenneth L. Finger
University of California Museum of Paleontology
1101 Valley Life Sciences Building, Berkeley, CA 94720-4780 USA
email: kfinger@berkeley.edu
ABSTRACT: Foraminifera are abundant and diverse in the Neogene sedimentary units of south-central Chile (3443S),
but their age and depositional paleoenvironment have been points of contention for decades. The marine strata most often
referred to the Navidad Formation have been a focal point of study in the region by geoscientists in academia as well as the
Chilean oil industry. Most of the foraminiferal assemblages documented in this study are from three sedimentary basins
isolated from each other by more than 400km. From north to south, they are (1) the Navidad Formation in the vicinity of the
name-bearing town located southwest of Santiago, (2) the Ranquil Formation on the Arauco Peninsula southeast of Concepcon, and (3) the Lacui Formation on and near Chilo Island. Also included in this study are two assemblages from deposits
of uncertain affinity in San Sebastin (northeast of the Navidad area and just north of San Antonio), and one from the Santo
Domingo Formation in the vicinity of Valdivia (about midway between Arauco and Chilo Island). The 27 localities reported in this study span nearly 1000km of the south-central Chilean margin. Most of the samples are siltstones from coastal
bluffs and wave-cut platforms in which bedding is nearly horizontal, but often massive or indistinct.
Regional workers have disagreed on which subepochs these units represent. Suggested ages based on macro- and microfossils have ranged from Late Oligocene to Early Pliocene. The planktic foraminifera identified in this study suggest that most,
if not all, are Miocene. Excluding the two San Sebastin assemblages, which are slightly younger, one assemblage devoid
of planktic foraminifera, and three with only long-ranging planktic foraminifera, all of the remaining 21 concurrent ranges
begin or end in the Early Miocene. Nineteen of those ranges are restricted to that subepoch and two range into the early
Middle Miocene. Seventeen of 18 localities had an 87Sr/86Sr age ranging into or within the Early Miocene. The exception
yielded an isotopic age within the latest Oligocene. The majority of the analytical data presented in this report suggest that
most of the assemblages are of late Early Miocene (Burdigalian) age.
The most abundant macrofossils in the Navidad, Ranquil, Lacui, and Santo Domingo formations (collectively and informally referred to in this study as the Navidad group) are gastropods typical of neritic habitats, but specimens tend to be widely
scattered in the outcrops and they are associated with mixed-depth assemblages of foraminifera. I therefore concluded that
there had been significant downslope displacement. The upper depth limits of the deepest-dwelling benthic foraminifera in
each assemblage indicate that final deposition occurred at bathyal depths, well below the neritic zone.
All of the 358 taxa (336 benthics + 22 planktics) identified in this study are presented systematically and illustrated. New
species described are Astacolus novambiguus, Cornuspira libella, Cristellariopsis petersonae, Fissurina ambicarinata,
Globocassidulina chileensis, Karreriella biglobata, Percultazonaria encinasi, Percultazonaria obliquispinata, Pseudolingulina nielseni, and Pseudononion ranquilensis. In addition, I propose one new (substitute) name, Lenticulina neopolita, to
resolve a synonymy.
The presence of Neogene cosmopolitan deep-water benthic foraminifera in all of the assemblages supports the hypothesis
that deep water masses derived from the Antarctic Circumpolar Current have enabled many Neogene deep-water foraminifera to disperse widely in the global ocean. This compendium is a useful guide to identification of Neogene foraminifera in
subtropical/temperate regions in and beyond south-central Chile, and it also provides a regional dataset that can be incorporated into future studies on benthic foraminiferal biostratigraphy and biogeography.
INTRODUCTION
In comparison to most other regions, western South America is

represented by relatively few significant publications on fossil
foraminifera. This monograph fills a major void by documenting
the foraminiferal taxa in Miocene strata of south-central Chile.
This study focuses on the foraminifera of the El Peral beds
(Lo Abarca Formation?) and the Navidad group (an informal
name used here to refer collectively to the Navidad, Ranquil,
Santo Domingo, and Lacui formations). These units crop out in
south-central Chile between 33 and 44S (text-figs. 15). Age
interpretations of the units have ranged widely in the literature
(text-figs. 6 and 7). Molluscan and strontium isotope data indicate
a Late OligoceneMiddle Miocene age, and the gastropod fauna

is typical of neritic depths. In contrast, studies of microplankton
have yielded ages of Early, Middle, or Late Miocene, or Early
Pliocene, and deposition at bathyal depths. The Navidad group
was long thought of as Early Miocene shallow-marine deposits
on the basis of its molluscan fauna (Groves and Nielsen 2003;
Nielsen et al. 2004). In contrast, Finger et al. (2007) identified
benthic foraminifera interpreted as lower-bathyal indicators,
and sedimentary features typical of turbidites. Ichnofossils
supported that deep-water interpretation. Foremost among
their explanations for these discordancies were reworking and
downslope displacement by massive slumping and turbidity
currents. Since that time, however, these issues have remained
unsettled.
Micropaleontology, vol. 59, nos. 45, pp. 341492, text figures 118, tables 1-8, plates 1-24, 2013
341
Kenneth L. Finger: Miocene foraminifera from the south-central coast of Chile
TEXT-FIGURE 1
Map of south-central Chile showing primary areas of study and sample
localities not within any of the areas shown on text-figures 25.
The initial goal of this study was to produce a monograph on

the foraminiferal fauna of the Navidad group, but the conflicting
age and depth interpretations needed further investigation. In
addition to a large section on systematic taxonomy, this report
revisits these issues and presents extensive data sets and analyses
used in the current interpretation.
PREVIOUS STUDIES
Charles Darwin was first to explore the geology of this region,

where he visited from November 1834 to March 1835 (Darwin
1839). Darwin (1846) later named and described the older
Tertiary strata in the coastal bluffs near the town of Navidad,
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TEXT-FIGURE 2
Las Cruces area (Lo Abarca Formation) collecting sites. Neogene strata
are exposed in shaded areas.
~140km southwest of Santiago, as the Formation of Navidad,

noting that the unit had a rich gastropod fauna. The bluff on the
west side of Punta Perro (text-fig. 3) has generally been referred to
as the Navidad stratotype, although Martnez-Pardo and Osorio
(1964) noted that it does not match Darwins locality description.
Sowerby (1846) documented the gastropods collected by Darwin,
and Philippi (1887) subsequently provided a more thorough
monograph based on that and subsequent collections. Mricke
(1896) was the first to recognize the molluscan fauna as Miocene.
Micropaleontology, vol. 59, nos. 45, 2013
TEXT-FIGURE 4
Arauco area (Ranquil Formation) and Valdivia (Santo Domingo Formation) collecting sites. Neogene strata are exposed in shaded areas. Not
plotted is RQS, which yielded few foraminifers.
TEXT-FIGURE 3
Navidad area (Navidad Formation) collecting sites. Neogene strata are
exposed in shaded areas. Not plotted are CCQ, MAP, PPG, and PPS,
which yielded few foraminifers.
Tavera (1968, 1979) confined the Navidad Formation to the

Burdigalian (late Early Miocene) based on correlations with
Patagonian gastropods, and divided it into the Navidad,
Lincancheu, and Rapel members. The age of the basal Navidad
Member and related units, however, became a matter of debate
following micropaleontological studies that concluded different
ages (text-figs. 6, 7). Martnez-Pardo and Osorio 1964 studied
an assemblage of foraminifera and ostracodes from the marine
terrace on the west side of the Punta Perro peninsula and
suggested it was Late Miocene. Osorio (1978) later detailed the

ostracodes and used the stratigraphic distribution of some of the
species in the Caribbean region to support that interpretation.
Dremel (in Herm 1969) identified 17 species of planktic
foraminifera from the terrace that placed the unit in the Early
Miocene. Martnez-Pardo and Valenzuela (1979) later reported
that the Navidad Formation had discoasters indicative of the
Middle Miocene. As a result of these studies, Martnez-Pardo
(1990) referred to the unit as Early to Late Miocene. Ibaraki
(1992a) sampled the Navidad Formation about six miles south of
Punta Perro and identified planktic foraminifera that correlated
with the Late Miocene (also see Tsuchi et al. 1990; Tsuchi 2002;
Ibaraki 1992b).
343
Their study was the first on benthic foraminifera in the regional

Neogene, and they interpreted the assemblage as indicative of
bathyal deposition at 200500m.
Martnez and Pino (1979) ascribed the Santo Domingo
Formation to the Miocene based on the overall composition
of the foraminiferal assemblage, as all of their identifications
of planktic foraminifera were conferred (e.g., Globoquadrina
cf. dehiscens). Later, Marchant and Pineda (1988) recorded
Globigerina pachyderma from this unit, which first appears in
the Late Miocene.
Although Martnez (1990) synonymized the Navidad, Ranquil,
and Lacui formations as the Navidad Formation, but their
distinction is retained here in order to distinguish them as
discrete regional units and because their relationships remain
to be confirmed. Encinas et al. (2006) proposed elevating the
Navidad, Lincancheu, Rapel, and La Cueva members of the
Navidad Formation to the rank of formation because they are
separated by regional paraconformities. Because of these
proposed changes, and other referrals of different Miocene units
in the region to the Navidad Formation, and for the convenience
of communication in this report, those units and assemblages
investigated in the present study are collectively referred to as
Navidad.
Anticipating data that would be useful in my study, I anxiously
awaited the findings of the ODP (Offshore Drilling Program Leg
202) investigations off central Chile (text-fig. 8) at Site 1232 in
the Chile Basin (west of the trench) and Sites 12331235 on the
Chile margin (forearc). Unfortunately, drilling did not extend
below the very thick Pleistocene section (Tiedemann and Mix
2007).
TEXT-FIGURE 5
Chilo area (Lacui Formation) collecting sites. Neogene strata are exposed in shaded areas.
The Ranquil Formation crops out mostly along the Arauco

Peninsula, ~60km SSW of Concepcin (text-fig. 4). Tavera
(1942) and Garca (1968) ascribed the unit to the Miocene based
on molluscs and foraminifera, respectively. Nearly 400km to the
south of that area, outcrops at Carelmapu and on Chilo Island
(text-fig. 5) are of the Lacui Formation, which Valenzuela (1982)
referred tentatively to the Pliocene, based on its stratigraphic
position between Miocene volcanics and Pleistocene sediments.
He also recognized as Miocene the four species of molluscs that
Machado (1909, p. 5) had identified in a calcareous sandstone
at Carelmapu, but questioned their provenance. Sernageomin
(1998) later assigned the Lacui to the earliest Serravallian based
on foraminifera (Duhart, Muoz and Stern 2000).
Martnez and Parada (1968) used benthic foraminifera to place
the San Sebastin beds, referred to in this report as the El Peral
beds, in the Pliocene, as they did not recover any planktic species.
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Encinas et al. (2006, 2007) and Finger et al. (2007) analyzed

the sediments and fossils (i.e., trace fossils, gastropods,
foraminifera, ostracodes) of the Navidad, Ranquil, and Lacui
formations in south-central Chile (33-45S), as well as the
foraminifera and ostracodes from numerous wells drilled in
the region. These authors described the composite section as
having a basal thin, shallow-marine, conglomerate overlain
by a clastic succession with Bouma cycles, parallel-laminated
sandstones, synsedimentary breccias, slides, slumps, diamictites,
and massive siltstones and sandstones. Deposition was attributed
to gravity flows, primarily turbidity currents and sandy debris
flows. The presence of Zoophycos ichnofacies and certain deepwater species of benthic foraminifera, ostracodes, and gastropods
support a deep-water origin. The benthic foraminifera suggest
deposition on the lower middle to lower continental slope below
1500m. These authors concluded that these units were deposited
in the Late Miocene or Early Pliocene as slope aprons as the
result of rapid and major forearc subsidence that led to subduction
erosion (Encinas et al. 2008; Encinas, Finger and Buatois 2012).
Finger et al. (2007) mentioned, tabulated, or illustrated only
fossils they considered significant in their interpretation. The
present study provides the details of the complete foraminiferal
dataset compiled from the Navidad, Ranquil, Lacui, and Santo
Domingo formations, as well as the deposits referred to herein as
the El Peral beds.
Planktic foraminifera reported by Finger et al. (2007) and Encinas
et al. (2008) indicated deposition of the Navidad, Ranquil,
and Lacui formations occurred in the Late Miocene and Early
Pliocene. Their incongruent association with older Miocene
planktic foraminifera, Late Oligocene-early Middle Miocene
gastropods (DeVries and Frassinetti 2003), and late Oligocene-
TEXT-FIGURE 6
Previous interpretations of the geologic units referred to in this study. Publications listed are not necessarily the original source of the data or the
interpretations. Black bars are biostratigraphic; grey bars are chronostratigraphic.
345
schemes were modified and refined over the last four decades.
To enhance communication in this text, I refer to the N zones,
even though Berggren et al. (1995) constructed a transitional
(temperate) Mt zonation that is more appropriate for the Miocene
of south-central Chile. The comparison of zonations illustrated
in text-figure 9 shows the correlations between the zonation of
Berggren et al. (1995) and those of Gradstein, Ogg and Smith
(2004) and Wade et al. (2004), which enable realignment of
the Mt zones with the most recent revision of the low-latitude
scheme of Hilgen, Lourens and Van Dam (2012).
TEXT-FIGURE 7
Distribution of age determinations for the units derived from data in
text-figure 1.
Early Miocene shark teeth (Surez, Encinas and Ward 2006),

however, was problematic. Finger et al. (2007) suggested that
the older fossils were reworked by massive slumps from an older
stratigraphic unit that had yet to be identified. Although it seemed
to be a reasonable hypothesis, as giant continental slope failures
have been identified in this region (Geersen et al. 2011), there
are no known outcrops of that older age nor is there any direct
sedimentological evidence of massive slump blocks; hence, those
working on the Navidad continued to debate the age and, to a
lesser degree, the depositional environment. Recently, Gutirrez
et al. (2013) composited stratigraphic sections for the coastal
cliffs exposed between La Boca and Matanzas. They mostly
utilized published data (i.e., 40Ar/39Ar dates on pumice clasts, Sr87/
Sr86 dates on calcareous fossils, and molluscan and palynological
biostratigraphic correlations) to place the lower member of the
Navidad Formation in the Aquitanian Stage at 22.022.5 Ma,
which is early Early Miocene. Rather than questioning the
identifications of Late Miocene and Early Pliocene planktic
foraminifera in Finger et al. (2007), they concluded that those
species must have originated in the southeastern Pacific Ocean
during the Early Miocene. The following discussion delves
further into this issue.
Biostratigraphic framework
Micropaleontologists working the foraminifera-rich land-based

sections in the Caribbean region were the first to construct
widely accepted biostratigraphic zonations of Cenozoic planktic
foraminifera. Bolli (1957, 1966, 1970) applied species names to
zones based on the first or last occurrences of species, which
he used to define the lower and upper boundaries of each zone.
Banner and Blow (1965) and Blow (1969, 1979) developed a
similar scheme characterized by numbered P (Paleogene) and
N (Neogene) zones. Because the alphanumeric scheme was
more convenient to communicate, it became the more popular
of the two. In subsequent years, the two frameworks melded
and incorporated zonations of other widely used planktic
microfossils (e.g., calcareous nannoplankton), tying all into
polarity chrons and absolute ages (see Berggren et al. 1995, for
example). Jenkins (1966, 1967, 1971) created the first Cenozoic
planktic foraminiferal zonation for temperate regions based
on his work in New Zealand. Both the temperate and tropical
346
Deep-sea cores generally provide the most complete and

continuous successions of planktic foraminifera, and their
records can be invaluable guides to regional land-based
sequences. Several expeditions have drilled the Southeast Pacific
off southern Peru and Chile (text-fig. 8), but none have explored
the region between 23 and 36S. Contemporaneous with this
study, ODP Leg 202 drilled 14 coreholes at four sites off central
Chile. Site 1232 is in the Chile Basin, west of the Peru-Chile
Trench, 265km NE of Chilo Island. The other three sites are
on the Chilean margin (forearc): Site 1233 (41.00S, 74.45W) is
92km NE of Chilo Island, and Sites 1234 (36.22S, 73.68W)
and 1235 (36.16S, 73.57W) are 86km NE of Concepcon. A
comparative Neogene section was greatly anticipated, but none
of these coreholes penetrated below the thick Pleistocene section
(Tiedemann and Mix 2007). Also disappointing were IODP Leg
141 sites 859863 at the Chile Triple Junction (c. 46S, 75W),
none of which drilled deeper than the upper Pliocene (Spiegler
and Mller 1995).
The nearest Miocene deep-sea core sections were obtained from
the tropical zone off Peru (c. 921S) at the Nazca Plate, Nazca
Ridge, and East Pacific Rise (DSDP sites 319321, IODP Leg
142, and ODP sites 12351237). The stratigraphy at ODP Site
1237 (16.00S, 76.38W), 447km south of Lima on the eastern
end of the Nazca Ridge west of the trench, ranks among the
best hemipelagic and pelagic reference sections from the South
Pacific, having provided a complete pelagic Oligocene (~31 Ma)
to Holocene sequence that was relatively unmodified by burial
(Tiedemann and Mix 2007). The Shipboard Scientific Party
(2003) identified 23 planktic foraminifer datums in the Miocene
of Site 1237 (text-fig. 9). Incorporating that biostratigraphy into
studies beyond that region, however, could result in inaccurate
interpretations. For example, Hodell and Kennett (1986) found
many planktic foraminiferal datums markedly depart from
the correlation suggested by magnetostratigraphy between the
subtropics of the South Atlantic and Southwest Pacific, which
indicates that such datum levels can be unreliable for correlation
between these ocean basins. Also, Berggren et al. (1995) noted
that the LA of Globoquadrina dehiscens ranges from 5.8 Ma
in the tropics to 6.8 Ma in the subtropics, and Keller (1980)
correlated its LA in DSDP Hole 319 (131.0380S, 10131.4580W,
Bauer Deep, Nazca Plate) with the middle of N16 (~10 Ma).
This diachrony spans about 4 Myr and suggests that the species
disappeared even earlier off south-central Chile. Extrapolating
the datums recorded at Site 1237 to this project area could result
in errors on the order of several million years. This would not be
a significant problem if those particular datums were well within
or beyond the concurrent range determined for any spot sample.
PALEOBATHYMETRY
Depth zonation models
Two depth zonations have been widely used in foraminiferal

studies (table 1): one specific to the tectonic margin of the East
Pacific, championed by the California (Bandy) school (e.g., Ingle
TEXT-FIGURE 8
Map showing locations of ODP sites off Peru and Chile between 7S and 50 S relative to the present region of study.
347
TEXT-FIGURE 9
Neogene time scale with planktic foraminiferal zonations and datums. Species in bold were found in the Chilean samples. Abbreviations of genera: C.=Catapsydrax, F. = Fohsella, G.=Globigerina, Gd.=Globoquadrina, Glb.=Globigerinoides, Glr.=Globorotalia, Gq =Globoquadrina,
Gt.=Globoturborotalita, Gtl. = Globigerinatella, Ng.=Neogloboquadrina, O.=Orbulina, P.=Pulleniatina, Pg.=Paragloborotalia, Pr.=Praeorbulina. Abbreviations of zonal units: CRZ=Concurrent Range Zone, CRSZ=Concurrent Range Subzone, ISZ=Interval Subzone, IZ=Interval Zone,
PRZ=Primary Range Zone, RSZ=Range Subzone, TRZ=Transitional Range Zone. Datum symbols: p first occurrence; q last occurrence; open
triangle if not reported from ODP 1237.
348
1980), and the other presented by van Morkhoven, Berggren

and Edwards (1986) in their book on cosmopolitan deep-water
foraminifera. The latter authors provide no explanation of the
zonation they used, so the reader is left to assume that it has
global applicability. The data presented in their compilation,
however, are biased toward passive ocean margins, a notion
recently confirmed by one of its coauthors (W. A. Berggren,
pers. comm. 2011). The present study incorporates the Pacific
specific scheme developed by Ingle (1980) and applied by Ingle,
Keller and Kolpack (1980) to the benthic foraminifera off southcentral Chile.
ranges of modern species to the fossil record using the priniciple

of homeomorphy, which assumes that similar morphologies
reflect similar environments. Micropaleontologists have assumed
that this approach was fairly reliable for at least the last 25 Myr
because, as many authors have documented, the modern benthic
foraminiferal fauna has changed little since the Oligocene or
Eocene. The depth ranges of extant species enable us to use fossil
assemblages to interpret the depositional history of sedimentary
basins. The method has been applied to various aspects of
paleoenvironmental reconstruction, including sedimentation
rates, eustatic changes, and other processes.
Ingle (1980) based his foraminiferal depth zonation on the most

significant depth-related parameters in the middle latitudes,
which include light penetration, temperature, dissolved oxygen,
stratification of the water column, pressure, and CCD (carbonate
compensation depth). For example, the thermocline off
California is at 200m, the oxygen-minimum zone (with high P
and N) from 150 to 450m, nutrient-poor Antarctic Intermediate
Water from 400900m, another oxygen minimum zone from
9001700m and the CCD at 4000m. These depths are not all
coincident with water mass boundaries, which counters the
general concept that prevailed three to four decades ago, which
attributed bathymetric distributions of benthic foraminifera to
the water-mass stratification of the oceans (e.g., Douglas and
Heitman 1979; Pujos-Lamy 1973; Streeter 1973; Schnitker
1974; Streeter and Shackleton 1979). Subsequent studies (e.g.,
Mackensen et al. 1990; Schmiedl, Mackensen and Miller 1997;
Jorissen, Fontanier and Thomas 2007; Mojtahid et al., 2010;
Phipps et al., 2012) have clearly shown that major water-mass
boundaries do not necessarily coincide with faunal changes.
Natland and Kuenen (1951) and Phleger (1951) established

the utility of benthic foraminifera as tracers of displacement,
and Bandy (1961) substantiated the logical concept that the
percentage of displaced species should progressively increase
downslope. To avoid being misled by those species with wide
bathymetric ranges and any displaced from shallower depths,
only the upper depth limits (UDLs) of the deepest-dwelling
forms in the fossil assemblage should be used to estimate the
minimum depositional depth (Bandy 1961, 1967; Ingle 1967,
1980; Ingle and Keller 1980). The absence of species with deeper
UDLs does not eliminate the possibility that deposition occurred
deeper. Foraminiferal assemblages recovered from bathyal
depths along active margins are likely to be misinterpreted
if the most common species are considered indicative of the
depositional environment, as they may not be in situ.
It is important to recognize that all depth zonations are only

rough approximations because the physicochemical parameters
that control bathymetric distributions of foraminifera vary
both geographically and temporally. The most significant
differences between the two schemes are in the lower part of
the bathyal realm, where the zones are offset by as much as
2000m. Regardless of which zonation is used, workers should
consider the zones relative to one another rather than the precise
numerical depths assigned to them (e.g., middle bathyal depths
are undeniably well below neritic depths). This would avoid
potential conflicts with regional geologic concepts based on
modern scenarios.
Benthic foraminifera as paleodepth indicators
Benthic foraminifera can be valuable tools for deciphering

paleoecological parameters, reconstructing paleoenvironments,
and recognizing reworked or displaced sediments. As with most
other benthic marine organisms, their biofacies change with
depth; hence, depth zonations have been determined for modern
faunas in many regions, particularly in the oil-rich regions of
the northern Gulf of Mexico and southern California, where
the data enhanced stratigraphic correlations. The methodology
of paleodepth interpretation was developed in California,
beginning with Natlands (1933) classic dissertation study in
which he collected modern data from the Santa Monica basin
off Los Angeles to interpret a thick Pliocene-Pleistocene section
exposed about 70km WNW, in Hall Canyon just west of Ventura.
Of particular significance is Natlands recognition of mixeddepth assemblages, a phenomenon that Phleger (1951) later
reported in modern foraminiferal assemblages in the San Diego
Trough and Sigsbee Deep. Subsequent studies that refined the
methodology of interpreting paleodepths include Natland (1952,
1957), Bandy (1953a, 1953b, 1960), Bandy and Arnal (1957,
1960), and Ingle (1967, 1980). These workers applied the depth
The reliability of this method has been scrutinized and criticized

in numerous publications. Several authors have correlated largescale depth distributions with water masses (Lohman 1978;
Blake and Douglas 1980; Schnitker 1980; Culver and Buzas
1980, 1981a, b), which have migrated vertically (and laterally)
over time. Although the concept of isobathyal species has been
largely abandoned in recent years, benthic foraminifera tend to
display depth-related distributions within areas of considerable
size (Culver 1988). Nevertheless, it is prudent to apply
modern upper depth limits from the same region as that of the
paleontologic study.
What controls the depth distribution of benthic foraminifera?
Murray (2006) discusses the more commonly measured
parameters in modern foraminiferal studies: salinity, temperature,
dissolved oxygen, nutrients, tides, substrate, competition, space,
food supply, and controls on anoxia. It was long thought that
the marine temperature gradient was the primary controlling
factor, but it is now believed to play only a very minor role, if
any. Van der Zwaan et al. (1999) concluded that the interplay
between organic flux and dissolved oxygen control foraminiferal
depth distributions, noting that both tend to decrease with water
depth. They asserted that individual species are never good
paleodepth markers even though some typically deep-water
taxa are adapted to low organic flux, while others proliferate at
the shelf margin where there is high organic load. High organic
flux, as well as weak circulation, leads to oxygen depletion, and
the tolerance to dysoxia varies substantially among species of
foraminifera. In deep-water environments, organic matter flux
has a major influence on the spatial and depth distributions
of benthic foraminifera; thus local and broad distinctions can
be made between depth zones (Gooday 2003, and references
therein). Van der Zwaan, Jorissen and Verhallen (1990) proposed
using the P:B ratio as a more reliable indicator of paleodepth,
as it is independent of flux because both planktics and benthics
are dependent on it. Providing that the CCD is not reached, the
ratio increases with depth because depth typically increases with
distance offshore. The wide range of ratios found at intermediate
349
depths limits this method. For example, the %P vs. depth plot
shown in Van der Zwaan et al. (1999, fig. 12) has a range of
about 070% P at a depth of 100m, and 60100% at 1000m. On
that chart, 19% P (the average value found in the present study)
correlates with depths ranging 30-700m. These broad variations
suggest that the P:B ratio is more useful in determining the
temporal trend in depth from a stratigraphic sequence than it is
in assigning depositional depths to spot samples. In addition, and
most relevant to the present study, downslope displacement of
benthic foraminifera can significantly lower P:B values, and thus
result in inaccurate paleobathymetric determinations.
It is obvious from the above discussion that the use of foraminifera
as paleodepth indicators has its limitations, but the methodology
has often produced useful results in reconstructing the history
of Neogene coastal depocenters, such as the Gulf of Mexico
(Katz and Miller 1993a, b) and California (Ingle 1980; Lagoe
1985; Olson 1990). As with benthic biostratigraphy, the key is
temporal and geographic proximity of the past to the present.
Workers should be less concerned with specific depth ranges
or zones than with which modern counterparts of the fossil
assemblage characterize the shelf vs. the slope, particularly in
the given provincial area. Although benthic foraminiferal species
distributions in the deep sea may be dependent on organic flux
and oxygen, it is undeniable that many benthic genera and species
are typical of shallow, intermediate, deep, or very deep waters.
The south-central region of Chile, as delimited in this study, is
adjacent to two marine ecoregions. The Araucanian ecoregion,
at the southern end of the Warm Temperate Southeastern Pacific
Province, extends from Valparaiso to Punta Chocoi (east of
Puerto Montt). The Chiloense ecoregion, at the northern end of
the Magellanic Province, encompasses Chilo Island (Spalding
et al. 2007). For ease of discussion, the present study refers
to these two marine ecoregions as the South-central Chilean
Province; hence, their foraminiferal communities constitute the
modern provincial fauna.
Fieldwork and sample localities
TEXT-FIGURE 10
Results of linear regression analyses. A, number of specimens vs. species richness. B, number of specimens vs. Fisher a diversity index. C,
species richness vs. Fisher a diversity index. Locality number sequence
(1 to 27) is from north to south.
350
This study originally focused on 46 samples from 32 localities

along the central Chilean coast south of Valparaiso (~3300S),
from Las Cruces (~3430S) to Cucao (~4240S) (text-figs. 15).
Alfonso Encinas, Sven Nielsen, and I collected nearly all of the
outcrop samples during 20002003. During the course of this
study, I also examined hundreds of micropaleontological slides
representing 25 wells drilled by ENAP (Empresa Nacional del
Petrleo [the national oil company of Chile]) in this region:
Belavista #1, Cholchol #1 and #2, Colegual #1 and #2, Cuva #1,
Huilma #1 and #2, Labranza nos. 2, 48, and 10, Los Muermos
#1 and #2, Los Pinos #1, Navidad #1, Pozo B1, Puerto Montt
#1, Rahue #1 and #2, Rio Blanco #1, and Tegualda #1. One
assemblage from the Navidad #1 well is included in my dataset.
The Miocene sequences in these wells have the same benthic
fauna that I report here, in some cases overlying an Eocene
sequence with coal, or the Cretaceous crystalline basement.
Encinas provided additional slides from 24 outcrops in the
Central Valley, but most of the assemblages were weak and
very poorly preserved. My examination of these supplementary
assemblages observed only rare specimens of a few Miocene
species that are not among those documented from the primary
localities.
In the following, the 32 localities are arranged alphabetically
and each includes its description, latitude and longitude (from
Google Earth), name of collector(s) and year(s) of collection,
and equivalent UCMP (University of California Museum of
TEXT-FIGURE 11
Dendrogram from Q-mode cluster analysis using Wards method and a presence-absence matrix for all 27 benthic assemblages, based on the 52
dominant (common and abundant) species. Nodal points marked by a dot () denote clusters that only group samples from same geographic area.
Paleontology) microfossil locality number. Most of the samples

provided by Nielsen were matrix sediments from gastropod
samples. The five sample localities that are not in bold type
yielded few, if any, foraminifera and are excluded from the
statistical analyses. Photographs of localities CCQ, CUC, MAT,
PPP, PPS, and RAN are included in Finger et al. (2007).
CCQ: Navidad Formation. White coquina between granitic
basement and massive sandstone ~15m north of MOS,
approximately 6km north of the mouth of Estero Maitenlahue,
near Punta Toro, north of Mostazal and Candelero, San Antonio
Province, Valparaso Region, 334711S, 714742W. Collected
by A. Encinas, 2003. UCMP MF9003.
CHE: Lacui Formation. Yellowish sandstone from coastal bluffs,

south of Chepu, Isla Chilo (see Watters and Fleming 1972),
Chilo Province, Los Lagos Region, 420426S, 74 021W.
Collected by S. N. Nielsen, 2001. UCMP MF9030.
CHO: Lacui Formation. Grey silty mudstone, intertidal platform
along southern coast of Punta Chocoi, Llanquihue Province,
Los Lagos Region, 414443S, 735036W. Collected by S. N.
Nielsen, 2001. UCMP MF9027.
CPUP: Navidad Formation. Grey siltstones in a small roadcut
along Camino de Pupuya, approximately 2.5km due south of
La Boca, Cardinal Caro Province, Libertador General Bernardo
OHiggins Region, 340252S, 715036W. Collected by A.
Encinas, 2004. UCMP MF9016.
351
TEXT-FIGURE 12
Dendrogram from R-mode cluster analysis using Wards method and a presence-absence matrix for all 27 assemblages, based on same data matrix
as text-figure 11.
CUC: Lacui Formation. Grey sandstone, coastal bluff blockfall

south of Cucao, Isla Chilo, Chilo Province, Los Lagos Region,
424228S, 740810W. Collected by S. N. Nielsen, 2001.
UCMP MF9031.
FRA: Ranquil Formation. Brown sandstone, concordantly
superjacent to FRM, coastal bluff at Punta Fraile, west of Tubul,
Arauco Peninsula, Arauco Province, Biobo Region, 371213S,
732927W. Collected by S. N. Nielsen, 20002002. UCMP
MF9018.
FRM: Ranquil Formation. Grey mudstone to siltstone, basal
part of exposure concordantly subjacent to FRA, intertidal
352
platform, Punta Fraile, west of Tubul, Arauco Peninsula, Arauco

Province, Biobo Region, 371213S, 732927W. Collected by
S. N. Nielsen, 20002002. UCMP MF9019.
LBZ: Navidad Formation. Fossiliferous lenses of yellowish to
orange-brown sandstones with abundant Pinna semicostata
at top of 5-m thick interval of interbedded sandstone and
microconglomerate, basal part of coastal bluff located
approximately 1km north of Las Brisas, adjacent to a small
lagoon formed at the mouth of the Estero Navidad. south of Punta
Perro, between La Boca and Las Brisas, Cardinal Caro Province,
Libertador General Bernardo OHiggins Region, 335543S,
6km north of the mouth of Estero Maitenlahue, just south of

Punta Toro, north of Mostazal and south of Rocas de Santo
Domingo, San Antonio Province, Valparaso Region, 334649S,
714738W. Collected by A. Encinas, 2003. UCMP MF9002.
MPUP: Navidad Formation. Coarse grey sandstone lens ~15
stratigraphic feet above PPP coastal bluff, Punta Perro, Cardinal
Caro Province, Libertador General Bernardo OHiggins Region,
335439S, 715038W, Collected by S. N. Nielsen 2000 &
2001 and K. L. Finger 2003. UCMP MF9015.
TEXT-FIGURE 13
Dendrogram from Q-mode cluster analysis using Wards method and a
presence-absence matrix for the faunas in each of the five geologic units,
based on the 172 species that had a relative abundance of at least 1% in
one assemblage.
715046W. Collected by S. N. Nielsen, 2000 & 2001. UCMP

MF9010. (Corresponds to NV4 of Encinas et al. 2008)
LEB: Ranquil Formation. Greenish grey calcareous sandstone
cobbles derived from channel fill cutting through well-sorted
Eocene(?) sandstones, isolated bluff consisting of large filled
crevice in in well-sorted nonfossiliferous, presumably Eocene
sandstones, northern end of Lebu, Arauco Peninsula, Arauco
Province, Biobo Region, 373539S, 733816W. Collected by
S. N. 2000 & 2002; K. L. Finger 2003. UCMP MF9025.
LPER: El Peral beds (unnamed unit). Off the northeast end of
Laguna El Peral, on east side of road (Avda. El Peral) between
San Sebastin and Las Cruces, just below the bend (Laguna de
los Patos locality of Martnez and Parada 1968), San Antonio
Province, Valparaso Region, 333012S, 713626W. Portion
of Martnez and Paradas bulk sample obtained by A. Encinas
from the Museo Nacional de Historia Natural, Santiago.
(Locality recollected by K. L. Finger in 2003, but retrieved
assemblage was poorly preserved.) UCMP MF9001.
MAP: Navidad Formation. Grey siltstone with coarse sandstone
and shell lenses, intertidal platform about 1km north of
Matanzas, Cardinal Caro Province, Libertador General Bernardo
OHiggins Region, 335658S, 715205W. Collected by S. N.
Nielsen 2001 & 2002; and K. L. Finger 2003. UCMP MF9013.
MAT: Navidad Formation. Small lens of glauconitic sandstone
23m above base of coastal bluff and approximately 2m
stratigraphically above MAP, Los Goterones (about 1km north
of Matanzas), Cardinal Caro Province, Libertador General
Bernardo OHiggins Region, 335655S, 715159W. Collected
by S. N. Nielsen, 20002002. UCMP MF9012.
MIB: Ranquil Formation. Grey mudstones to siltstones, coastal
bluff on southern part of Caleta Ranquil, close to fault contact
with Millongue Formation, Arauco Peninsula, Arauco Province,
Biobo Region, 373228S, 733646W. Collected by K. L.
Finger, 2003. UCMP MF9024.
MOS: Navidad Formation. Seven-meter thick interval of grey
siltstone that overlies a 0.5-m thick microconglomerate capping
large granitic boulders at base of coastal bluff approximately
MS10: Ranquil Formation. Grey siltstone on Isla Quiriquina,

entrance to Baha de Concepcin, Biobo Region. Collected by
J. Tavera. Locality details not available (approximate center of
island is 363738, 733038). Sample obtained by A. Encinas
from the Museo Nacional de Historia Natural, Santiago. UCMP
MF9017.
NAV5: Navidad Formation. Navidad #5 well, Matanzas, Cardinal
335730S, 715223W. Foraminiferal residues from well
depths of 2430 and 3036m, loaned by ENAP. UCMP MF9014.
NLP: El Peral beds (unnamed unit). Shovel diggings from the
excavation of a water well to a depth of 7m, at the side of the
road (Avda. El Peral) between San Sebastin and Las Cruces,
just north of the western shoreline at the northern end of
Laguna El Peral, approximately 200m northwest of LPER, San
Antonio Province, Valparaso Region, 333004S, 713626W.
Collected by A. Encinas, 2004. UCMP MF9000.
PCB: Lacui Formation. Yellowish coquina containing
foraminifera and calcareous fragments of echinoderms, oysters,
pectinids, bryozoans and claystone clasts, coastal bluff west
of Playa Chaumn, Isla Chilo, Chilo Province, Los Lagos
Region, 415544S, 740139W. Collected by S. N. Nielsen,
2001. UCMP MF9028.
PNH: Lacui Formation. Dark gray siltstone from a small
(approximately 2m 1m) sedimentary pocket in the Ancud
Volcanic Complex near the pinguinera (penguin station) at
Puihuil, Isla Chilo, Chilo Province, Los Lagos Region,
415544S, 740139W. Collected by S. N. Nielsen, 2001.
UCMP MF9029.
PPG: Navidad Formation. Calcareous matrix between granitic
boulders on beach, slightly north of PPS, Punta Perro, Cardinal
335438S, 715041W. Collected by S. N. Nielsen, 2000
2002. UCMP MF9008.
PPN: Navidad Formation. Fossiliferous lenses of yellowish to
orange-brown sandstones in the coastal bluff at Punta Perro and
stratigraphically below PPP, Cardinal Caro Province, Libertador
General Bernardo OHiggins Region, 335423S, 715018W.
Collected by S. N. Nielsen 20002002. UCMP MF9007.
PPP: Navidad Formation. Grey siltstone, surface of intertidal
platform at the northwestern tip of Punta Perro, just south of
the mouth of Rio Rapel, Cardinal Caro Province, Libertador
General Bernardo OHiggins Region, 335415S, 715013W.
Collected by S. N. Nielsen, 20002002 and K. L. Finger, 2003.
UCMP MF9005.
PPS: Navidad Formation. Coarse grey sandstone lens, coastal
bluff, Punta Perro, Cardinal Caro Province, Libertador General
Bernardo OHiggins Region, 335439S, 715038W. Collected
353
TEXT-FIGURE 14
Principle coordinates plot from NMDS analysis showing the relationships between localities based on the 52 dominant (common and abundant) species occurrences.
by S. N. Nielsen 2000 & 2001 and K. L. Finger 2003. UCMP

MF9009.
335623S, 71514W. Collected by S. N. Nielsen 2002 and K.

L. Finger 2003. UCMP MF9011.
PPT: Navidad Formation. Top of grey siltstone interval, approximately 15m stratigraphically above PPP, coastal bluff, Punta
Perro, Cardinal Caro Province, Libertador General Bernardo
OHiggins Region, 335416S, 71509W. Collected by A.
Encinas, 2006. UCMP MF9006.
RAN: Ranquil Formation. Brown massive sandstones with

intermittent beds of glauconitic sandstone, overlying RQT and
transected by RQS, coastal bluff of Punta Huenteguapi, Ranquil,
733528W. Collected by S. N. Nielsen 20002002 and K. L.
Finger 2003. UCMP MF9023.
PTA: Navidad Formation. Fossiliferous lens of grey siltstone

similar to that of PPP and PPN, at top of a 20-m thick
siltstone interval that overlies a 30-m thick interval of massive
microconglomerates and medium- to coarse-grained sandstones,
coastal bluff almost below dirt road, Punta Alta, south of
Las Brisas, Libertador General Bernardo OHiggins Region,
354
RAP: Navidad Formation. Grey, reddish-brown and darkbrown sandstones in an undifferentiated blockfall from steep
cliffs along the coast north of Ro Rapel, San Antonio Province,
Valparaso Region, 335320S, 714934W. Collected by S. N.
Nielsen, 2000 & 2002. UCMP MF9004.
TEXT-FIGURE 15
Principle coordinates plot from NMDS analysis showing relationships between species based on same data matrix as text-figure 14.
RQK: Ranquil Formation. Highly fossiliferous sandstone

boulders restricted to the northernmost part of the beach, Punta
Huenteguapi, Arauco Peninsula, Arauco Province, Biobo
Region, 373020S, 733526W. Derived from the top of
the sequence that is no longer present on the top of the bluffs.
Collected by S. N. Nielsen 2001 & 2002 and K. L. Finger 2003.
UCMP MF9022.
RQS: Ranquil Formation. Gastropod-rich sandstone displaced
from upper part of adjacent bluffs and scattered on beach,
733524W. Collected by K. L. Finger 2003. UCMP9020.
RQT: Ranquil Formation. Grey mudstones to siltstones like
FRM, intertidal platform of Punta Huenteguapi, Ranquil,
733524W. Collected by S. N. Nielsen 2001 & 2002 and K. L.
Finger 2003. UCMP MF9021.
VAL: Santo Domingo Formation. Dark grey mudstone to
siltstone, bluff behind roadside house approximately 20km south
of Valdivia, Valdivia Province, Los Lagos Region, 395543S,
730732W. Collected by S. N. Nielsen, 2001. UCMP MF9026.
METHODS
Sample processing
Foraminifera were processed from 46 sedimentary rock samples

representing 32 localities by (1) soaking in water or hydrogen
peroxide until most of the sediment had disaggregated, (2)
washing the residue over a U.S. Standard 230-mesh (63-m
openings) sieve and (3) drying by funneling through fast-flow
filter paper, followed by oven-drying at 30C. Specimens were
then picked with a 000 sable hair brush and sorted by species
onto 63 60-grid micropaleontological slides, from which
primary types and hypotypes were selected and transferred
onto single-hole slides for reference and imaging. Five localities
were excluded from this study because they had poor yields of
foraminifera with no unique species, reducing the number of
localities to 27. After species identifications and counts were
made, assemblage data from sites that were sampled multiple
times were composited (i.e., one assemblage per site).
Both assemblage and individual species slides are in the
microfossil collection of the University of California Museum of
Paleontology (UCMP) in Berkeley. Primary types and hypotypes
were imaged with a succession of three environmental scanning
electron microscopes (ESEMs) at the UC Electron Microscope
Laboratory. A photomicroscope setup with the 30-year old
Infinite Focus system (Irvine Optical Corporation) was also
utilized, as in some cases it produced more revealing and useful,
albeit lower resolution, images. That system involves time-lapsed
photomicrography of a specimen on a motorized stage as it
slowly passes through a plane of illumination. Similar images
were subsequently obtained with a Leica IC80 HD microscope
camera, which is an integrated digital system that is considerably
more efficient, and the auto-blend (photostacking) feature of
Adobe Photoshop.
Taxonomic procedure
The primary resources initially used in this study for identifying

planktic foraminifera Kennett and Srinivasan (1983), Bolli and
Saunders (1985), Jenkins (1985), and Scott, Bishop and Burt
(1990). Also referred to were studies on planktic foraminifera
of Oligocene and Miocene deep-sea core sections, including
Brnnimann and Resig (1971), Spezzaferri and Premoli-Silva
(1992), Chaisson and Leckie (1993), Leckie, Farnham and
Schmidt (1993), and Majewski (2010). Identifications of benthic
species were based primarily on the type descriptions and figures
in the Catalogue of Foraminifera (Ellis and Messina 1940 et seq.),
revisions of the nine early European works on Tertiary to Recent
foraminifera listed in table 2, Atlas of Cosmopolitan Deep-water
Benthic Foraminifera (van Morkhoven, Berggren and Edwards
355
TEXT-FIGURE 16
Biostratigraphic correlation of Chilean samples based on concurrent ranges of planktic foraminifers that have first or last occurrences in the Miocene.
Arrow at top of bar indicate range continues post-Miocene; arrow at bottom of bar indicate species appears earlier in Tertiary. Species in bold and
preceded by a solid datum triangle are those found in this study. Light grey columns are samples that yielded no markers. Cross-hatched bars indicate
Sr ages from Nielsen and Glodny (2009) and Encinas (unpublished) and shaded grey where they overlap the biostratigraphic range; that for NAV5 and
PPT were obtained from tests of Paragloborotalia bella and Pg. zealandica, respectively; all others were derived from gastropod shells collected near
the microfossil locality they are associated with. See Figure 8 caption for abbreviation keys.
1986), Benthic Cenozoic foraminifera from Ecuador (Whittaker

1988), the recent tome on deep-water uniserial taxa (Hayward et
al. 2012), and the new Atlas of Benthic Foraminifera (Holbourn,
Henderson and MacLeod 2013). Literature on the modern
356
foraminifera of Chile was also perused for this purpose (see

following subsection).
Many of the benthic species identified in this study were originally described from the Pacific (OligoceneRecent), Caribbean
TEXT-FIGURE 17
Modern upper-depth limits of 76 foraminiferal genera represented in the Chilean Neogene that are common in the global fauna. Based on modern
global data from Murray (1991, 2006). Modern provincial data from Ingle, Keller and Kolpack (1980), Zapata and Moyano (1997), Zapata and Varela
(1975), and Figueroa et al. (2005, 2006). Key: Genera in bold = restricted to bathyal depths; dark grey cells = global common; light grey cells = global
infrequent; SCC = south-central Chile (provincial) UDL.
(Oligocene and Miocene), Mediterranean (MioceneRecent) Car

Nicobar Island in the Andaman Sea (Pliocene), and the Vienna
Basin of the Central Paratethys (OligoceneMiocene). A few
species were first described from the Atlantic and polar regions,
or from pre-Oligocene strata.
Previous taxonomic studies on the modern Chilean fauna
Marchant, Zapata and Hromic (2007) provide a thorough

bibliography of studies on the modern foraminifera of Chile.
Those most pertinent to the present study are discussed below.
The earliest report on the modern foraminifera off central Chile

describes species occurring in littoral sands from the coasts of
South America (i.e., Brazil to Ecuador; dOrbigny 1839c). Of the
81 new species described in that study, 56 were from the Atlantic
and 25 were from the Pacific. DOrbignys only sample taken off
Chile is from Baha de Valparaiso (33S), which yielded 12 new
species [brackets denote current generic assignment]: Rotalina
[Buccella] peruviana, Globigerina bulloides, Truncatulina
[Planulina] depressa, Truncatulina [Planulina] ornata,
Rosalina [Valvulineria] araucana, Valvulina [Nonionella]
357
TEXT-FIGURE 18
Modern slope profiles off south-central Chile. Horizontal bands show depth ranges where slopes level out. (Modified from Geersen et al. 2011).
auris, Valvulina [Cancris] inflata, Bulimina pulchella, Bulimina

[Praeglobobulimina] ovula, Bolivina plicata, Bolivina punctata,
and Quinqueloculina araucana. Of these, only Buccella
peruviana and Bulimina ovula occur in my samples. Nine other
species identified herein were described by dOrbigny (1839c)
from the southwest Atlantic one from Patagonia and eight
from the Falkland Islands.
Additional species were documented in H. B. Bradys (1884)
tome on the foraminifera collected by the Challenger Expedition
(18731876). That global venture included sample stations west
of Chilo and throughout the archipelago of southern Chile.
Egger (1893) subsequently worked on samples from the Gazelle
Expedition (18741876) that were collected off northern Chile
(north of Valparaiso). Bandy and Rodolfo (1964) studied 32
trawl and core samples from depths of 1796250m off Peru
and Chile, but only as far south as Valparaiso (32.3S). The
foraminifera off south-central Chile were included in studies by
Khusid (1971, 1974, 1977, 1979a, b) and Saidova (1969, 1971,
1975). Boltovskoy and Theyer (1970) analyzed 20 samples taken
at depths of 44260m off central Chile (29574216S). A few
years later, studies focused on specific locations off Chile began
appearing in South American journals (e.g., Zapata and Varela
1975). Resig (1981) analyzed 121 core-top samples taken from
depths of 822286m on the northern Nazca plate (027S) and
on the continental margin from 220S. More pertinent to the
present study is the analysis by Ingle, Keller and Kolpack (1980)
of bottom samples collected from depths of 1354500m along
three transects off central Chile (31.539.2S).
In the last two decades, several marine biologists in central
Chile have studied the modern foraminifera of the Chilean
margin (e.g., Zapata and Moyano 1997; Zapata and Cear 2004;
Figueroa et al. 2005, 2006; Marchant, Zapata and Hromic 2007).
Zapata (1999) studied benthic foraminifera in Cumberland
Bay (3341S, 7850W), Robinson Crusoe Island, in the Juan
Fernandez Archipelago ~670m west of the mainland at San
Antonio. His samples taken from depths down to 20m yielded
85 species, but he noted the degree of affinity with the Chilean
province was only 35% and, therefore, suggested that they were
different subprovinces. Zapata and Cear (2004) provided the
most thorough report on littoral foraminifera off the coast of
northern Chile (18283156S), having documented 151 species
358
from depths of 1170m. Nearly half of the species illustrated in

that study resemble those that occur in the Chilean Miocene, but
I retain the same binomina for only 20 of them. Most pertinent
among the studies by the Chileans is that of Figueroa et al.
(2005, 2006), who recorded 117 species of calcareous benthic
foraminifera from multicores taken at depths of 1253485m in
the south-central Chilean province (i.e., from Valparaiso to the
southern end of Chilo Island).
Taxonomic problems
Subjective synonymies are the nemesis of foraminiferal

taxonomy. Early workers were often unaware of publications
in foreign languages, as evidenced by the lack of comments
comparing their new species with previously described forms.
Also, as discussed by Lipps (2002), synonymies invaded the
foraminiferal literature in the 19th Century, partly because British
workers rejected dOrbignys concept of foraminiferal taxonomy.
Attitudes changed when H. B. Bradys (1884) Challenger tome
recognized many of dOrbignys genera.
In the first half of the 20th Century, J. A. Cushman pioneered
the application of benthic foraminifera in the North American
oil industry, and he soon became the most prolific authority on
their taxonomy. Unfortunately, he and his contemporaries tended
to view foraminifera as highly provincial and mostly ignored
species that had already been described in foreign languages.
This resulted in a multitude of synonyms that inundate the
topical literature. Murray (2007) estimates that for the modern
fauna as many as 25% of the species names are synonyms.
It has become increasingly evident that many species of benthic
foraminifera have much wider geologic and geographic ranges
than previously envisioned. In addition, it appears that many
named varieties, subspecies, and species that may have utility
in local biostratigraphic correlations are simply ecophenotypes
(i.e., invalid taxa). Wide geographic distributions are probably
due primarily to the dispersal of propagules by oceanic currents
(Alve and Goldstein 2003, 2010). To a lesser degree, testate
specimens are dispersed by water masses, detached algae, and
migrating marine animals (i.e., fish, birds, mammals). As colder,
denser water flows from high latitudes toward the equator,
the oceans become increasingly stratified. This phenomenon
enhances the cosmopolitan nature of the deep-water fauna,

TABLE 1
Comparison of bathymetric zonation schemes.
which preliminary genetic data supports (Pawlowski et al. 2007).

Neritic and marginal-marine foraminifera, on the other hand,
can be transported freely by surface currents or winds and their
geographic distribution can be assisted by adherence to floating
wood and algae or highly mobile marine animals (Murray 1991;
Culver and Buzas 2002). Ingestion by the latter vector is another
possibility, as there is some evidence that foraminifera can
survive passage through the digestive tracts of marine animals
(e.g., Brand and Lipps 1982).
It is both a blessing and a curse that foraminifera are so abundant
and diverse, and that they have received so much attention by
the scientific community. Their numerous applications are well
established in the earth sciences, especially in biostratigraphy,
paleoecology, paleoceanography, paleoclimatology, and
environmental science. Their great temporal and spatial diversity
of morphotypes, and the different opinions of taxonomists,
unfortunately have resulted in the conundrum of synonyms
previously noted. Boltovskoy (1965) expounded on this
taxonomic quagmire that may forever plague foraminiferology.
His pessimistic view would likely have been greater had he lived
long enough to learn that DNA sequencing has revealed several
cryptic species of benthic and planktic foraminifera (Gooday
and Jorissen 2012, and references therein). Early monographs
revised with better images of type specimens (table 2) certainly
have been a great asset in deciphering synonymies and detecting
misidentifications in the literature, but the taxonomic study of
the Foraminifera remains a formidable task. Those who have
already provided these valuable resources have sealed many
of the cracks in the foundation of foraminiferology and they
are commended for their extraordinary efforts. Nevertheless,
Linnaean taxonomy is typological, aligning species concepts
with primary type specimens. As concluded by Scott (2011)
in reference to planktic foraminifera, Typological practices
served well for the zonal biostratigraphic studies promoted by
Loeblich et al. (1957). That and allied research, which focused
on discovery of homotaxial stratigraphic markers, made little
demand on knowledge of populations. Its legacy is a host of
poorly described taxa. This certainly rings true for benthic
foraminifera as well.
Identifications made in this study
In the present study, I initially identified taxa by comparing

specimens with illustrations in notable papers on Oligocene to
Holocene faunas and modern publications bearing high-quality
images of contemporaneous specimens. Most useful among those
illustrating the modern Chilean fauna were those of Ingle, Keller
and Kolpack (1980) and Resig (1981). I utilized the Catalogue
of Foraminifera (Ellis and Messina 1940 et. seq.) extensively to
confirm species identifications, to construct synonymies, and to
select other comparative species worthy of mention.
TABLE 2
Early foraminiferal monographs and their latest revisions, and the number of their benthic species recognized in this study.
This compilation is the only extensive documentation of fossil

foraminifera from Chile and it serves as the definitive reference
to the Miocene fauna of this region. As with any study of similar
scope, further sampling may recover additional species, but the
large number of assemblages perused throughout the course of
this study suggests that they are most likely to be relatively rare
occurrences.
Applied statistics
Species diversity and assemblage similarity measurements used

a variety of applications provided by the PAST software package
of Hammer, Harper and Ryan (2001). I applied the Simpson
and Fisher diversity indice to each assemblage by using its
species richness and numbers of specimens. The Simpson index
indicates dominance and ranges from 0 (all taxa equally present)
to 1 (a monospecific assemblage). Fishers is a diversity index
defined by S = ln(1 + n/), where S is the number of taxa and
n is the number of specimens. To detect faunal similarities and
differences between areas and geologic units, I applied cluster
analyses and non-linear multidimensional scaling (NMDS) with
presence-absence matrices that were reduced in size first by the
exclusion of all species that do not account for at least 1% of
one assemblage), then by the exclusion of all species that do not
account for at least 5% of one assemblage. I used the similarity
coefficients of Jaccard and Simpson, as well as Wards method,
in the cluster analyses, and the correlation coefficient (Pearsons
r) for the NMDS.
RESULTS
The 27 assemblages analyzed in this study are represented by

more than 16,000 specimens that were picked and sorted on
60-grid micropaleontological assemblage slides (UCMP5043850499), from which I isolated representative specimens on
single-holed slides (UCMP50000-50437) for imaging and
reference. Table 3 shows the relative abundances of benthic
foraminifera present in each of the 27 assemblages. Table 4
simply indicates the presence/absence of planktic foraminiferal
species in each assemblage because many specimens were
diagenetically distorted or had obscured features. There was
also a wide range of transitional or variant forms. The recorded
fauna comprises 336 benthic and 24 planktic species. Table 5
presents the numerical calculations and diversity indices for each
assemblage. Compositing assemblages obtained by resampling
sites resulted in a wide range of specimen counts (1652133;
mean 610, median 471) and benthic species richness (20138;
mean 60, median 63). Planktic:benthic (P:B) ratios range 00.68
(mean 0.20, median 0.14).
Thirty-one benthic species occur at more than half the 27 localities.
The most widespread (number of localities in parentheses) are
Lenticulina subcultrata (26), Quinqueloculina akneriana (23),
Sphaeroidina bulloides (22), Glandulina laevigata (22), Bulimina
spicata (21), Cibicidoides compressus (20), Hoeglundina
359
TABLE 3
Relative abundance of benthic species vs. localities checklist. VR = 1 specimen; R 2 specimens if <1%, or 2 specimens if 1%; F = 15%; C = 525%;
A = 2550%; VA >33%.
360

TABLE 3
Continued.
361
TABLE 3
Continued.
362

TABLE 3
Continued.
363
TABLE 3
Continued.
364

TABLE 4
Planktic species presence vs. localities checklist. Relative abundances: VR = 1 specimen; R 2 specimens if <1%, or 2 specimens if 1%; F = 15%;
C = 525%; A = 2550%; VA >33%.
elegans (20), Dentalina striatissima (19), Hansenisca altiformis

(19), Laevidentalina elegans (19), Martinottiella communis
(19), Neugeborina longiscata (18), Saracenaria schencki (18),
Siphonodosaria lepidula (17), Globocassidulina chileensis (16),
Pseudononion novozealandicum (16), Pyrgo depressa (16), and
Zeaflorilus chiliensis (16).
Oligocene to Holocene. This is reflected in the distribution of the

type ages for the identified species (or comparative cf. species) in
Chile, 97% of which are Oligocene or younger (51% Quaternary,
38% Neogene and 8% Oligocene). This is also apparent in the
known ranges presented in the Systematic Taxonomy section,
nearly all of which are post-Eocene.
Only five species are relatively abundant (>25%) in any assemblage:

Cibicides umboniferus (LBZ), Ciperozoa basispinata (MIB),
Globocassidulina chileensis (PNH), Rectuvigerina transversa
(VAL), and Zeaflorilus chiliensis (PPN, CHE). Forty other
species are common (525%) at one or more localities. Of these
45 species that are at least 5% at one locality, 26 are restricted
to one locality, and two were recorded at multiple localities but
restricted to one formation; the remaining 17 are less constrained
(table 6).
Table 5 includes the results of the statistical measurements of

diversity, as well as the P:B values calculated for each of the
27 assemblages. Thirteen of the Fisher values are within the
normal range of 516 for open-marine environments (Murray,
1973), but the rest range higher, up to 32.66, and the 27
assemblages average 18.21. Considering that the normal range
is based on modern assemblages, and the values obtained from
fossil assemblages may have been lowered by post-mortem
disaggregation of weakly agglutinated species, the majority of
the Chilean values are abnormally high for fossil assemblages.
I performed linear regressions to examine the relationships
between the number of specimens, species richness, and Fisher
(text-figure 10). As more specimens are observed, the number
of species counted increases to a point where only very rare
components of the assemblage are likely to be found. Following
Phleger (1954), foraminiferologists have traditionally placed
that at 300 specimens the number derived from an analysis
of heavy mineral frequencies (Dryden 1931) and later extended
to zoological studies (Fisher, Corbett and Williams 1943). Even
though diversity indices are based on that logarithmic trend,
Fisher correlated much better with species richness than with
species number, possibly because the richness value is a factor of
the number, not vice versa.
Benthic species previously known from MioceneHolocene

deposits in the middle latitudes predominate in all assemblages
from Neogene outcrop and well samples examined during the
course of this study. Both benthic and planktic microfaunas
in temperate zones typically include taxa also known to
occur in subpolar and subtropical latitudes, and the samples
studied herein are no exception. Many benthic species or their
homeomorphs inhabiting the deep waters off Chile occur in the
provincial Neogene and some are members of the cosmopolitan
deep-water fauna documented by Morkhoven, Berggren and
Edwards (1986).
Boltovskoy (1980) claimed that less than 2.5% of Neogene bathyal
benthic foraminiferal taxa have provincial or regional utility as
guide fossils because the fauna has remained fairly stable from
365
TABLE 5
Numerical and statistical data tally for the 28 foraminiferal assemblages.
The Q-mode cluster analysis produced a dendrogram (text-fig.

11) that groups some assemblages from the same geologic units,
but there are many geologically and geographically incongruent
pairings. The associated R-mode clustering (text-fig. 12) resulted
in groupings that for the most part are paleobathymetrically
inconsistent, as expected. The relationships between the five
units are shown in another dendrogram (text-fig. 13), where the
Navidad and Ranquil formations are differentiated from the
other three, but the latters groupings are perplexing. The NMDS
plot (text-fig. 14) more clearly distinguishes the geologic units,
although those of the Navidad group show considerable overlap.
Each of the three primary units (Navidad, Ranquil, and Lacui
formations) overlaps the other two, perhaps related to similarities
in age and environment. The NMDS on the 52 common species
(text-fig. 15) yielded high stress values (S = 0.2399 for localities;
S = 0.4937 for species) that indicate poor ordinations. Similarly
poor values had been obtained prior, when the NMDS was
performed on the larger 172-species dataset that excluded all rare
species (which produced a very cluttered species plot).
DISCUSSION
Age of the units
In Peru, Navidad molluscs such as Miltha vidali, Acanthina

katzi, Olivancillaria claneophila, and Testallium cepa occur in
Late OligoceneMiddle Miocene sections (Nielsen et al. 2003),
but not in younger successions (DeVries and Frassinetti 2003).
366
Other Navidad species such as Ficus distans are restricted in

Peru to the EarlyMiddle Miocene, whereas Eucrassatella
ponderosa, Glycymeris ibariformis, and G. colchaguensis only
occur in the Late OligoceneEarly Miocene. Encinas (2006)
obtained Early Miocene ages of 24.70.4 and 20.40.5 Ma
from 87Sr/86Sr analyses of two O. claneophila specimens from
the Navidad Formation. Among these particular species in the
Navidad group, A. katzi (Finger et al. 2007, fig. 9) ranges the
youngest, to about 13 Ma. Encinas (2006) reported Sr isotope
dates in the range of 31.5 Ma (Early Oligocene) to 16.0 Ma
(early Middle Miocene) for 29 of 30 mollusc specimens from the
Navidad group, but only a few of the younger samples are from
the same localities as the microfossil assemblages reported here.
Nielsen and Glodny (2010) presented 87Sr/86Sr ages obtained
from molluscs collected at 14 of the Navidad group localities
in the general proximity of where the foraminiferal samples
were taken. Text-figure 16 shows the chronostratigraphic ages
(derived from both analytical data sets) that represent 18 of
the foraminiferal sample localities. Those ages range from
between 25.1 and 15.6 Myr Ma, or latest Oligocene (Chattian)
to early Middle Miocene (Langhian), but all but one (LEB) are
represented by at least one chronostratigraphic date extending
into or restricted to the Early Miocene. Evidence for being in this
lower, warmer interval of the Miocene may also be the presence
of benthic genera like Rectuvigerina, which in California has its
last occurrence at about 14 Ma (Finger 1992).

TABLE 6
Dominant (common and abundant) species in each assemblage.
Finger et al. (2007) previously addressed the discordant

interpretations that were the impetus for undertaking this study.
In that report, I identified several species of planktic foraminifera
in the Navidad, Ranquil, and Lacui formations that indicate Late
Miocene and Early Pliocene ages. Several colleagues agreed with
those identifications, and many of those index species had been
reported by others who previously worked in the region, some
having collected from the same localities examined in the present
study; hence, the identifications were thought to be accurate. In
2010, I had the opportunity to show my images of these species
to Martin Crundwell, who is intimately familiar with the mid-
latitude Miocene planktic fauna and the excellent stratigraphic

sections in New Zealand, from which many were first described.
Crundwell kindly provided his taxonomic opinions and argued
for an EarlyMiddle Miocene age. I subsequently followed
his suggestion that I peruse George Scotts publications on
the Miocene globorotaliids, and I also asked Scott to examine
the images. He confirmed the inaccuracy of some of my
identifications, but he was unable to assign several definitively
to species. It became obvious that some of the Chilean Miocene
taxa could not be reliably speciated because of their relative
rarity and stratigraphic isolation, which preclude a contextual
367
TABLE 7
Upper-depth limits of 383 foraminiferal taxa living off the central Chilean margin between 3344S. Data compiled from Ingle, Keller and Kolpack
(1980) and Figueroa (2005, 2006). Species found in the present study are indicated in bold; five of those species reported by Ingle, Keller and Kolpack
(I) are typically deep-water, but they were purported to occur much shallower by Figueroa (F).
368

TABLE 7
Continued.
369
understanding of where their particular morphologies fit among

the species used to subdivide the Miocene. I have carefully
reexamined my Chilean specimens in light of the comments from
both experts and Scotts pertinent publications. Although some
taxonomic uncertainties remain, the apparent indications better
agree with data from other sources, most notably molluscan
biostratigraphy and strontium isotope chronostratigraphy.
Nevertheless, unsettling discrepancies still challenge the
integrity of the planktic taxonomy and biostratigraphy of these
geologic units.
Other types of data mostly support the molluscan and isotopic
indications of ages in the Late OligoceneEarly Miocene
interval. Surez, Encinas and Ward (2006) identified the teeth of
various elasmobranch fishes in the Navidad Formation, including
Carcharoides totuserratus, an uncommon shark that has this
range (Surez and Marquardt 2001). Encinas (2006) dated six
volcanic scoria and pumice clasts in the Navidad Formation
by K/Ar and Ar/Ar analyses and found five were of Early to
Middle Miocene age (22.215.9 Ma); the exception yielded an
age of 11.060.19 Ma (earliest Late Miocene), which is 3.9 Myr
younger than the biostratigraphic range, and, therefore, assumed
to be unreliable.
The affinity of the El Peral beds with the Lo Abarca Formation
could only be postulated based on the regional sequence because
the former are silty mudstones with foraminifera, but no molluscs,
and the latter, described by Covacevich and Frassinetti (1990),
was primarily a limestone with molluscs but no foraminifera.
Covacevich and Frassinetti (1990) differentiated the Lo Abarca
Formation as younger than the Navidad Formation at Punta
Perro by comparing their molluscan faunas. Encinas et al. (2006,
2010) reported that the beds overlying the basal conglomerate of
the Lo Abarca stratotype yielded two diatom markers that have a
concurrent range of 12.211.3 Ma (Serravallian) in the equatorial
Pacific (Barron, 2003). That interval encompasses the Sr age
of 11.51.0 Ma obtained from an oyster shell collected at that
same level (Encinas, personal comm.). The two foraminiferal
assemblages from the Laguna el Peral area, 5km northwest of
the Lo Abarca stratotype, are also noticeably different from
those of the Navidad Formation. This is readily apparent in their
dominance by Neouvigerina hispida, which is not a dominant
constituent of any assemblage in the Navidad group (see table
3). The presence of Globorotalia miotumida in NLP and LPER
indicates a younger Miocene age of 15.0 7.3 Myr, but LPER
also has Globorotalia praemenardii, which has a more restricted
range of 14.211.6 Myr (Middle Miocene). This range overlaps
those obtained for the type Lo Abarca Formation, supporting the
notion that the El Peral beds belong to that unit, and dispelling
Martinez and Paradas interpretation of the LPER locality as
Pliocene, which they based on benthic foraminifera.
For the Navidad Formation in its type area along Punta Perro,
Martnez-Pardo and Osorio (1964), Cecioni (1970), Osorio
(1978), and Ibaraki (1992a, 1992b) suggested a Late Miocene
age. Ibaraki (1992a) was the first to apply modern planktic
foraminiferal biostratigraphy in her interpretation, and her
identification of Neogloboquadrina acostaensis (10.9 Ma FAD
in Berggren et al. 1995) placed the unit in the Tortonian. Shuto
(1990), Tsuchi et al. (1990), and Tsuchi (2002) also assigned
the Navidad at Punta Perro to the Late Miocene, presumably
based on Ibarakis report, even though it was associated with a
subtropical molluscan assemblage recorded by Covacevich and
Frassinetti (1980), which suggests that it preceded the global MidMiocene cooling event. Tsuchi (2002) correlated the molluscs
with one of the relative abundance spikes of warm-water planktic
370
foraminifera that he used to determine warm episodes in the

Pacific Neogene, notably that recognized at ~5.7 Ma in Japan,
Ecuador, Peru, and the Caleta Herradula de Mejillones section
near Antofagasta, northern Chile. Although Finger et al. (2007)
did not recognize Ng. acostaensis in their Punta Perro samples,
they reported the species from five other localities representing
the Navidad (NAV5), Ranquil (FRA, RQK), and Lacui (CHO,
CUC) formations. Their identifications of Globoturborotalia
apertura and Ng. pachyderma at another five localities (PTA,
MAT, MOS, RQT, MIB, PCB) also indicated a Late Miocene
age. The youngest index species they reported were Globorotalia
sphericomiozea (5.6 FAD in Berggren et al., 1995) at PTA and
Glr. puncticulata (4.6 Ma FAD in Berggren et al. 1995) at six
localities (PPP, PPT, PTA, FRA, RQK, CUC) that were therefore
referred to the Early Pliocene. They noted longer concurrent
range zones extending upward into the Late Miocene for 12
other planktic assemblages (LPER, NLP, CPUP, LBZ, MOS,
PPN, RAP, MS10, FRM, LEB, RAN, CHE). Finger et al. (2007)
concluded that faunal similarities among all of these localities
suggested similar ages within the Late Miocene to Early Pliocene
interval.
Of the 21 planktic foraminifer species with Miocene datums that
were recognized in ODP Site 1237, off southern Peru (text-fig.
9), I recognized only Catapsydrax dissimilis, Globigerinoides
primordius, Gln. trilobus, Globoquadrina dehiscens, and
Globorotalia praemenardii in the outcrop samples from central
Chile. All three species have datums in the Early Miocene.
Gutirrez et al. (2013) recently challenged the deduction by Finger
et al. (2007) that the Navidad Formation was a Late Miocene
Early Pliocene deep-water deposit, by insisting that the unit is an
Early to Middle Miocene shallow-water deposit. There are two
plausible explanations for the age disagreement: (1) reworking,
as proposed by Finger et al. (2007), and (2) misidentification of
index species by Finger et al. (2007). Gutirrez et al. (2012) did not
consider the latter possibility, but instead assumed the planktic
markers had to have evolved much earlier in the Southeast Pacific
than elsewhere. Modern microfossil biostratigraphy, honed by
several decades of deep-sea core studies, immediately dismisses
that hypothesis because the voluminous amount of global data
show that any regional differences in first appearance datums
are on are a much shorter time scale, and such diachronous
events certainly would have been detected by foraminiferal
biostratigraphers and paleoceanographers long ago.
It is now apparent to me that the younger age determinations
are incorrect, and the result of misidentifications. This can be
attributed partly to the preservational state of most specimens
and the absence of any extended or continuous stratigraphic
sequences that would put their morphologic variability into
temporal perspective. The most common planktic species in the
Navidad group are Globigerina venezuelana, Globigerinella
obesa, Globoquadrina dehiscens, and Globoturborotalita
woodi, all of which have long ranges in the Miocene. Of
these, only Gq. dehiscens has a Miocene datum, being its first
occurrence just above the base of the Miocene; hence, none
of these four species is useful in restricting an assemblage to
a single subepoch or age. Although less abundant, the most
informative species in the Navidad group are Catapsydrax
dissimilis (N6 LAD), Globigerinoides primordius (N4AN5
FAD), Globoquadrina dehiscens (N4 FAD), transitional forms
between Paragloborotalia nana (N6 LAD), Neogloboquadrina
continuosa, (N6 FAD), Pg. bella (N4N8), and the Pg. zealandica
group (N5N7). For the El Peral beds they are Orbulina
universa (N9 FAD), Globorotalia miotumida (N9 FAD), and

TABLE 8
Paleobathymetric interpretations based on 68 benthic foraminifera in the Chilean Miocene that have bathyal UDLs. Species are placed in depth zones
according to their upper depth limits off south-central Chile as recorded by Ingle, Keller and Kolpack (1980), with secondary consideration given
to global UDLs reported by van Morkhoven, Berggren and Edwards (1986) and Hayward et al. (2012). Figueroa et al. (2005, 2006) recorded the 12
species in shaded cells from the inner shelf; their exclusion would shift only four depth zone interpretations, and those would be from from lower to
lower middle bathyal (shaded cells in bottom row).
371
Glr. praemenardii (N10N13). Catapsydrax dissimilis is present

in seven of the Navidad group assemblages, often in association
with what were previously thought to be younger taxa, including
the orbulines. Whereas C. dissimilis is highly resistant to
dissolution (Kennett and Srinivasan 1983), reworking seemed
a logical explanation for its presence. All of the specimens
identified as Orbulina universa have since been reexamined, and
only those in the two El Peral assemblages tested positive for
calcium carbonate, revealing that the other porous, nonspinose
spheres (Pl. 16, Fig. 14) were the predominant radiolarians in the
washed sample residues representing 11 localities in the Navidad
and Ranquil formations.
Finger et al. (2007) erroneously reported five species of planktic
foraminifera in the Navidad group that have first appearance
datums in the Late Miocene or Early Pliocene. These were
Neogloboquadrina acostaensis (N16 FAD), Ng. pachyderma (N16
FAD), Globoturborotalia apertura (N16 FAD), Globorotalia
sphericomiozea (N16 LAD), and Globorotalia puncticulata (N19
FAD). As previously noted, these were identified on the basis of
their illustrations in Kennett and Srinivasan (1983), Bolli and
Saunders (1985), and Jenkins (1985). In addition, Ng. acostaensis
had been reported by Ibaraki (1992a) from Punta Perro and by
Osorio and Elgueta (1990) from the ENAP Labranza #1 well
drilled west of Temuco, where they also recorded Gt. apertura
and Ng. pachyderma. Marchant and Pineda (1988) and Marchant
(1990) also recorded Ng. pachyderma in the vicinity of Valdivia.
I have carefully reexamined these species in my assemblages,
with particular reference to the detailed descriptions, morphostatistical analyses, and excellent images provided by Scott
(1983, 2011), Scott, Bishop and Burt (1990), and Scott et al.
(2007). My revisions are as follows: (1) the rare specimens that
appear identical to Glt. apertura are large-apertured variants
within the Glt. woodi populations they are associated with;
(2) the specimens previously referred to Ng. acostaensis, Ng.
continuosa, and Ng. pachyderma are now ascribed to various
transitional forms in Pg. bella, Pg. nanaNg continuosa, and
Pg. nana, respectively; (4) the rare specimens identified as Glr.
sphericomiozea now confer with Glr. miozea; (5) what was
thought to be Glr. puncticulata are now recognized as juvenile
Pg. zealandica, and (6), as noted above, the specimens in the
Navidad group that had been ascribed to Orbulina universa are
actually radiolarians. The features that distinguish each of these
species are discussed in the Systematic Taxonomy section.
Excluding the assemblages devoid (VAL) and nearly devoid
(PNH) of planktic foraminifera, and two with only longranging species (MPUP, LEB), each of the 21 assemblages
from the Navidad group begin or end in the Early Miocene;
19 of those ranges are restricted to that subepoch, whereas
two others (FRM, CHE) range into the Middle Miocene (textfig. 16). I did not find any species with a Late Oligocene LAD.
Seventeen of 18 localities had a 87Sr/86Sr age coincident with the
Early Miocene (text-fig. 16); the exception yielded an isotopic
age considerably younger than the biostratigraphic range and
was therefore considered unreliable. One isotopic age (RQK)
ranges into the Middle Miocene, whereas two (MAT, FRM)
cross over the boundary into the latest Oligocene. The Sr age
obtained for MAT, however, is 0.5 My older than that indicated
by the foraminifera. Two Sr dates were obtained for VAL,
one Early Miocene and the other latest Oligocene. The only
other Sr age restricted to the latest Oligocene was from LEB,
which, as noted above, did not yield any planktic foraminifera
useful in constraining the biostratigraphic age within the latest
OligoceneMiocene interval. In summary, 16 localities yielded
Sr-isotope ages that at least partly overlap the concurrent range
372
indicated by planktic foraminifera. Overall, the analytical data

place nearly all of the material collected from the Navidad group
within the Burdigalian stage of the late Early Miocene.
Depositional paleoenvironment
Previously, Finger et al. (2007) attempted to end the disagreement

among regional workers about the depositional depth of the
units by determining which taxa are regionally restricted to
deep water according to the depths reported in Ingle, Keller and
Kolpack (1980) and van Morkhoven, Berggren and Edwards
(1986), and then identifying the deepest minimal depth zone
indicated within each assemblage. The findings led to two
conclusions that are repeated here. First, all samples yielded
mixed-depth assemblages of benthic foraminifera, indicating the
prevalence of downslope transport and contributing to the faunal
heterogeneity between sample sites. Ingle, Keller and Kolpack
(1980) previously documented this phenomenon in their study
of transects taken off Valparaiso (33S), Cabo Carranza (36S),
and Valdivia (39S), as did Resig (1990) in her foraminiferal
study of 21 drill sites along the Peru margin (DSDP Legs 18 and
112, respectively). Such downslope displacement of sediments
is a common phenomenon on tectonically active margins (e.g.,
Shipp, Weimer and Posamentier 2011; Slatt and Zavala 2012).
The second conclusion was that all of the sampled units were
deposited at deep bathyal depths.
If a fossil assemblage has undergone significant bathymetric
mixing, it may yield anomalously high values of species
richness and diversity that reflect the conglomeration of taxa
from different depth-related biofacies. Most of the benthic
foraminiferal assemblages in this study yielded numbers that
are unusually high for a modern in situ temperate assemblage,
despite any taphonomic loss that may have occurred. From
another perspective, the depths interpreted for the fossil
assemblages have a narrower range than those from which the
modern fauna was sampled, yet the total number of species in the
fossil fauna is not much less than that of the modern provincial
fauna. This similarity in species richness could be explained
by the paleobathymetric mixing that is evident in the fossil
assemblages.
Reworking is most readily recognized by the presence of
significantly older fossils that show a poorer state of preservation,
but the only reliable evidence of this phenomenon is a single
Cretaceous globotruncanid test recovered from the FRA locality
and the association of Praeorbulina with slightly younger species
in LPER. Each assemblage appears more likely to be the product
of mixing unconsolidated sediments that had accumulated along
a depth transect of downslope displacement.
The paleobathymetric study has been expanded to incorporate
data on the modern fauna reported in South American journals. Of
those, Figueroa et al. (2005, 2006) provide the most bathymetric
data for the provincial fauna, including many neritic occurrences
shallower than those reported by Ingle, Keller and Kolpack (1980).
Combined, these three reports total 108 genera and 374 species
(table 7).
On the generic level, the Chilean fauna is similar to those from
other Neogene locations in temperate and subtropical zones,
particularly the diverse and well-studied units of the Caribbean
and New Zealand, but many of the benthic species are recorded
only from one of these three regions. Regardless of unrealized
synonymies, this clearly indicates that many species did not have
wide geographic or temporal ranges.
Typical modern outer neritic and deeper benthic foraminiferal

assemblages have 30 to 60 species per thousand specimens (Murray
1973). The 27 Chilean assemblages have species:specimen ratios
that equate to 44 to 223 species per thousand specimens, with an
average of 122. Accordingly, species diversity indices range well
above the normal, as previously mentioned. These disparities are
also evident in the wide variation of species that dominate each
assemblage (table 6). Despite the mixing, the premise that the
Navidad, Ranquil, Santo Domingo, and Lacui formations have
similar faunas that may have been contemporaneous is borne out
by the multivariate statistical analyses (text-figs. 1115), which do
not clearly distinguish them from each other.
Text-figure 17 shows the modern provincial and global UDLs of
76 common genera, all of which are represented in the Chilean
Miocene. It suggests that most, if not all, of these genera have
been recorded from neritic depths; in the south-central Chilean
province, however, 14 of the genera are represented by species
found only at bathyal depths: Ammobaculites, Anomalinoides,
Bathysiphon, Chilostomella, Gaudryina, Globocassidulina,
Laticarinina, Osangularia, Pleurostomella, Quadrimorphina,
Rhabdammina, Robertina, Tritaxis, and Virgulinella. One or
more of these are represented in 23 of the 27 assemblages (the
exceptions are PPN, LBZ, MAT, and CHO); thus, despite any
inconsistency in species identifications between different workers,
and some seemingly anomalous UDLs, the evidence strongly
favors deposition at bathyal depths.
Modern benthic foraminifera off central and northern Chile were
first reported in geographically broader works by dOrbigny
(1839c), Gay (1854), and Brady (1884). The Southeast Pacific
margin was first isolated for study by Bandy and Rodolfo (1964),
who examined foraminifera in 32 trawl and core samples taken
from depths of 1796250m off Peru and Chile, but only as far
south as Valparaiso (32.3S). Ingle, Keller and Kolpack (1980)
analyzed the fauna in bottom samples collected from depths of
1354500m along three transects off central Chile (31.539.2S).
Resig (1981) analyzed 121 core-top samples taken from depths
of 822286m on the northern part of the Nazca plate (027S)
and on the continental margin off Guayaquil, Ecuador (2S)
to Iquique (20S), Chile. Recently, several marine biologists
have focused on the modern foraminiferal fauna off Chile (e.g.,
Zapata and Cear 2004; Zapata and Moyano 1997; Figueroa et al.
2005, 2006; Marchant, Zapata and Hromic 2007). A study by
Zapata (1999) of the benthic foraminifera down to 20m depth in
Cumberland Bay (3341S, 7850W), Robinson Crusoe Island,
Juan Fernandez Archipelago (~670m west of the mainland at
San Antonio), yielded 85 species but he noted the degree of
affinity with the Chilean province was only 35% and suggested
that they were different subprovinces. Zapata and Cear (2004)
provided the most thorough report on littoral foraminifera off
the coast of northern Chile (18283156S). They documented
151 species from depths of 1170m, but only 20 of those species
are recognized in the Miocene fauna, and about half appear to
be different species. From the south-central Chilean province
extending from Valparaiso to Chilo, Figueroa et al. (2005,
2006) recorded 117 species of calcareous benthic foraminifera
from multicores taken at depths of 1253485m. Hence, Ingle,
Keller and Kolpack (1980) and Figueroa et al. (2005, 2006)
are the source of the 374 provincial upper-depth limits (UDLs)
listed in table 7, which serves as the foundation for extrapolating
provincial modern bathyal UDLs into the regional fossil record
(table 8).
Considering that the average time range of a Cenozoic benthic
foraminiferal species is estimated to be 1525 million years
(Buzas and Culver 1984), it is not surprising that 63 (22%) of

the Chilean Miocene benthic species are provincially extant, nor
that about half of those have been provincially recovered only
from bathyal depths. Deep-water deposition is further supported
by 21 genera represented in the Chilean Miocene that have been
recorded only at bathyal depths off south-central Chile (text-fig.
17). Other species UDLs noted in the systematics section are
extrapolated from the modern cosmopolitan deep-water fauna
documented by van Morkhoven, Berggren and Edwards (1986),
Hayward et al. (2012), and Holbourn, Henderson and MacLeod
(2013).
Table 8 shows the distribution of 63 species assigned bathyal
UDLs in this study. The 27 assemblages range 433 bathyal
species with an average of 16; the numbers of middle or lower
bathyal indicators per assemblage range 121 and average 9.
Among these are the seemingly anomalous inner shelf records
of Cyclammina cancellata, Fissurina sp., Favulina hexagona,
Fontbotia wuellerstorfi, Laticarinina pauperata, Martinottiella
communis, Melonis pompilioides (f. spheroides), M. barleeanus,
Oridorsalis umbonatus, Pullenia bulloides, Pyrgo murrhina, and
Triloculina trigona. These 12 taxa are included in table 8 in their
otherwise bathyal depth zones, deepening the paleobathymetric
interpretations for three localities (MOS, RQK, CHE), from
lower middle bathyal to lower bathyal. Excluding those 12 species
from the set of 63 bathyal depth indicators would result in 4 upper
middle bathyal, 5 lower middle bathyal, and 18 lower bathyal
paleodepth zone interpretations. The 559 bathyal indications
(332 being middle and lower bathyal) in table 8 should erase
any lingering doubts about the deep-water interpretation for
the units, as it is unlikely that any significant number of the 63
species consistently had anomalously shallow occurrences. Their
association with neritic species is considered here to be evidence
of downslope displacement and bathymetric mixing with final
deposition on the continental slope, most likely at middle to lower
bathyal depths.
The paleobathymetric interpretation of the foraminifera fits
the modern depositional scenario off south-central Chile,
where earthquakes trigger slumps and debris flows that evolve
into turbidity currents and mudflows that rework and funnel
slope sediments through deep submarine canyons (Raitzsch,
Volker and Huebeck 2007). Displaced sediments accumulate
in topographic depressions and where the seafloor levels out;
in the latter case off south-central Chile, these depocenters are
at depths between 19002200m for normal slopes and between
28003600m for slope embayments (text-figure 18).
An argument can be made about the accuracy of the temporal
consistency of depth zones assigned to the Chilean assemblages,
as the UDLs are based on extrapolation from the Holocene to
the Early Miocene, a span of more than 16 Myr that includes the
late Middle Miocene global cooling event, and it often assumes
that similar congeneric morphotypes lived at similar depths. In
addition, UDLs vary geographically there are no isobathyal
species. With the exception of polar emergence, those geographic
differences should rarely exceed a few hundred meters or one
bathymetric zone. All of the assemblages, except LBZ, are
interpreted to have been deposited in the lower middle bathyal
(15002000m) or lower bathyal (20004000m) zone, but the
margin of error is unknown. All of the Chilean assemblages
have indications of deep-water deposition, well below 500m. It is
noteworthy that ODP Site 1237, drilled at a water depth of 3212m,
yielded a Neogene fauna characterized by Chrysalogonium
spp., Cibicidoides mundulus, Globocassidulina subglobosa,
Gyroidinoides soldanii, G. orbicularis, Laticarinina pauperata,
373
Melonis affinis, Oridorsalis umbonatus, Planulina wuellerstorfi,

Pullenia bulloides, Pyrgo murrhina, Rectuvigerina striata,
Siphonina tenuicarinata, Stilostomella abyssorum, Stilostomella
subspinosa, and Vulvulina spinosa (Shipboard Scientific Party
2003). Most of those species, as well as very similar taxa and
possible synonyms, are identified in the present study. Regardless
of purported or verified shallow-water occurrences, these species
and many of those identified in the present study are typically
found at bathyal depths.
Comparison with other American Cenozoic faunas
There are numerous well-documented Oligocene, Miocene,

and Pliocene benthic foraminiferal faunas in the lower-latitude
Americas. They have many genera, but comparatively fewer
species (including likely synonyms), in common with the
Chilean Early Miocene. These studies include the faunas of six
formations spanning the Middle Oligocene to Lower Miocene
of Puerto Rico (Galloway and Heminway 1941), the Oligocene
Cipero Marl in Trinidad and Tobago (Cushman and Stainforth
1945), the Early Miocene La Boca Formation of Panama
(Blacut and Kleinpell 1969), the lower Pliocene of southeastern
Mexico (Kohl 1985), the middle to Late Miocene Buff Bay
Formation of Jamaica (Robertson 1998), and the late middle to
Late Miocene Gatun Formation of Panama (Collins et al. 1996,
Coates et al. 2009). Of additional utility in comparing with the
Caribbean taxa is the compendium by Bolli, Beckmann and
Saunders (1994) on the Cretaceous to Miocene foraminiferal
biostratigraphy of Trinidad, Venezuela, and Barbados. The most
thorough study on a contemporaneous fauna from the Pacific
side of South America is Whittakers (1988) work on benthic
foraminifera from the Late Oligocene to Pliocene sequences in
Ecuador, which provides taxonomic and distributional data for
130 species. Many of the Ecuadorian Miocene taxa, particularly
the deep-water species, occur in the Chilean Neogene. Although
Natland et al. (1974) recognized 200 species of foraminifera
from Tertiary sequences in the Magallanes Basin in southern
Patagonia, they only provided information on those 25 species
determined to be of biostratigraphic utility in their study, and
none of them is recognized in the present study.
Biogeography
Biogeographic inferences are difficult to make for benthic

foraminifera because species identifications in the literature
reflect worker subjectivity on intraspecific variation and
provincial vs. cosmopolitan distributions, as well as their
experience and taxonomic skills, available resources (i.e.,
imaging, literature, collections, colleagues, time), and the
amount of time and effort devoted to identifying specimens.
Some relevant comments are included in the beginning of
the Systematics section of this report. Considering that ocean
currents and other vectors effectively transport tests and
propagules, I find it difficult to conceive any bona fide marine
microfossil species can be restricted to its type locality and
horizon, although it might appear that many are, especially if
they lack adequate type-figures and subsequent workers apply
other names without recognizing their synonymies, or if similar
coeval facies had not been studied elsewhere. Many workers
have been misled by Cushmans profuse contributions in which
he had a propensity to designate a new species if it was found
in a different region or epoch than a very similar or identical
morphotype already described. As expected, the vast majority of
species in the Chilean Miocene fauna were originally described
from the OligoceneHolocene, and their regional distributions
are widely scattered across the globe. Although many of the
modern offshore Chilean species or their homeomorphs are
374
recognized in the Chilean Neogene, the provincial foraminiferal

fauna was unknown until the second half of the 20th Century,
well after the vast majority of common Neogene species had
already been described elsewhere.
Many of the Neogene benthic foraminiferal species in Chile appear
to have wide geographic ranges, which suggests that oceanic
pathways connected these disparate regions. The Mediterranean
basin was not silled off from the North Atlantic in the Early
Neogene, and transoceanic migration, particularly via the deep
water masses, could have distributed benthic species. This might
explain why many of the same species are recognized in both the
Mediterranean and Caribbean regions. Deep-water straits across
Central America would have enabled the Caribbean foraminifera
to migrate to and from the subtropical Northeast Pacific. Coates
et al. (2009) reported that benthic foraminifera indicate that the
deepest parts of the Chucunaque-Tuira and Sambu basins in the
Darien province of Panama were at lower-bathyal depths during
the Middle Miocene, but the basins shallowed as the Panama arc
began colliding with South America, rising to neritic depths in
the Early Pliocene and emerging at 4.8 Ma. Similarities between
the Miocene foraminifera of Car Nicobar and Chile, on the other
hand, could be due to oceanic pathways of cold, deep water
masses emanating from the Southern Ocean.
Modern water masses of the Southeast Pacific are described
by Strub et al. (1998). Off central Chile today, cold, nutrientenriched subpolar water is transported northward by the PeruChile Current (PCC). The Coastal Current (CC) also flows
northward but is significantly affected by an admixture of lowsalinity waters from the Chile fjord region. In between them,
100300km offshore, the Peru-Chile Counter Current (PCCC)
transports subtropical surface water to the south. The polewardflowing Gunther Undercurrent underlies these surface-water
masses at depths of 100400m and transports relatively lowoxygen and high-salinity water masses southward along the
shelf edge. At depths of 4001000m is the northward-flowing
Antarctic Intermediate Water (AAIW), which is relatively high
in oxygen and low in salinity. It overlies the southward-flowing,
nutrient-rich Pacific Central Water (PCW). If a similar pattern
of stratification and circulation existed in the Oligo-Miocene, it
could have provided both northerly and southerly pathways for
potential foraminiferal migration.
The Chilean Neogene benthic foraminiferal fauna has relatively
few species in common with the welldocumented Neogene
deep-water basins of Japan and California. This suggests that
Oligocene and Early Miocene, foraminiferal migration across
the Equator may have been more difficult in the Pacific than in
the Atlantic. Although van Morkhoven, Berggren and Edwards
(1986) designated a select number of deepwater Neogene
foraminifera as cosmopolitan, they presented relatively few
data from the Southeast Pacific in their study. The present study
indicates that many other species may belong to the cosmopolitan
deep-water fauna.
CONCLUSIONS
The benthic foraminiferal faunas of the Navidad, Ranquil, Santo

Domingo, and Lacui formations (the Navidad group) cannot
readily be distinguished from each other due to similarities in
geologic age, depositional history, and species composition. Most
of the species (excluding very rare ones) occur in two or more
of these units. Only the northernmost strata in this study, the El
Peral beds, yielded assemblages that do not correlate with the
Navidad group, but instead may belong to the nearby Lo Abarca
Formation. I conclude that all of the Navidad group localities are
Early Miocene, based primarily on the presence of the planktic

species Catapsydrax dissimilis, Globigerinoides primordius,
Paragloborotalia bella, and Paragloborotalia zealandica s.l.,
as well as strontium-isotope chronostratigraphy and molluscan
biostratigraphy. The results of statistical analyses of the benthic
foraminiferal data weakly correlate with geography and geology
of the areas studied due to their faunal similarities. Foraminiferal
assemblages from the El Peral beds differ from the those of the
Navidad group by the dominance of benthic species not found in
the other units and by the presence of orbulines, Globorotalia
miotumida, and Glr. praemenardii, which indicate Middle and
Late Miocene ages.
Benthic foraminifera indicate that all of the samples were
deposited on the lower half of the continental slope between
1000 and 2500m. Downslope displacement and deep-water
deposition on the forearc of the Peru-Chile trench is supported
not just by the tectonic setting, but also by the recognition of
deep-dwelling (psychrospheric) ostracodes, partial Bouma
sequences, and the Zoophycos ichnofacies. All of these findings
imply that the well-studied gastropods of the Navidad group are
not in situ.
This report fulfills the need for a guide to the Neogene foraminifera
of south-central coastal Chile, and it is anticipated that it will
impact future studies on the stratigraphy, sedimentology,
and paleontology of the region. In addition, the abundance of
cosmopolitan deep-water species in this fauna extends its the
utility of this publication to Neogene foraminiferal studies
elsewhere in the mid-latitudes.
ACKNOWLEDGMENTS
I thank my colleagues Alfonso Encinas and Sven Nielsen for

inviting me to join them on a collaborative, interdisciplinary
project to study the geology and paleontology of south-central
Chile. They were indispensable as field partners and sources of
additional samples, regional literature, and pertinent data, and
together we coauthored several publications on various aspects of
the project. I also thank them for their comradeship and endless
support throughout the entire project, and for pre-reviewing the
manuscript. ENAP (Empresa Nacional del Petrleo) graciously
loaned many slides of unsorted foraminifera from numerous
wells drilled in the region. I am also very grateful to have had
the assistance of the late Dawn E. Peterson in sample processing
and scanning electron microscopy. Humberto Carvajal Chitty
(BioStrat International, Venezuela) and Lee Hsiang Low (Centre
for Ecological and Evolutionary Synthesis, University of Oslo)
kindly provided some sought after advice on the statistical
analyses. With my utmost appreciation, I acknowledge C.
Wylie Poag (USGS, Reston) for taking on, and completing, the
daunting task of reviewing the entire manuscript and providing
many useful comments, suggestions, and annotations. The
UCMP provided funds for my fieldwork in Chile and scanning
electron microscopy at the Robert D. Ogg Electron Microscope
Laboratory (EML), University of California, Berkeley.
SYSTEMATIC TAXONOMY
The 27 Chilean Neogene localities yielded a foraminiferal

fauna consisting of 336 benthic and 22 planktic species. All
assemblage and type specimen slides have been deposited in the
microfossil collections at the UCMP (University of California
Museum of Paleontology). The taxa identified in this study
represent 162 genera, and are systematically arranged according
to the supraspecific framework provided by Loeblich and
Tappan (1987) and subsequent revisions above the rank of family

(Lee 1990, 2000; Loeblich and Tappan 1992, 1993; Sen Gupta
1999; Cavalier-Smith 2002), including that for the suffix of
superfamilies (ICZN 4th Ed. 1999, Art. 29.2). The subdivisions of
the Foraminifera should be recognized as uncertain because they
are not fully consistent with molecular phylogenetic data (Adl et
al. 2005). Ten new species described are Karreriella biglobata,
Cornuspira libella, Pseudolingulina nielseni, Cristellariopsis
petersonae, Percultazonaria encinasi, Percultazonaria obliquispina, Astacolus novambiguus, Fissurina ambicarinata,
Globocassidulina chileensis, and Pseudononion ranquilensis.
One new (substitute) name, Lenticulina neopolita, is proposed
for an objective junior synonym. Among the benthic fauna are
19 species conferred (cf.) to another species, and an additional 46
left in open nomenclature because they are not represented by any
specimens that are distinct and well preserved enough to warrant
their establishment as new species.
Format of this section
For each species, reference to its original designation and

description is included. Many include a synonymy based on
comparison with published images.
Distinguishing features: Primary characteristics used to
distinguish (1) relatively new (post-Loeblich and Tappan 1987)
genera from similar genera represented in this study, and (2)
selected species that may not be readily distinguished from
others identified in this study.
Type age and locality: The general age and locality designated
for the holotype. If a type locality was not indicated, the first
reported localities are noted.
Stratigraphic range: For benthic species, this is a minimum
range based only on the type level, the present study, and ages
indicated in the global studies by van Morkhoven, Berggren and
Edwards (1986), Jones (1994), and Hayward et al. (2012). For
planktic species, the stratigraphic range is indicated by the age
range corresponding to the Paleogene (P) and Neogene (N) zones
of Blow (1969, 1979) that define the first and last appearances of
the species. Ranges of planktic species are derived from Kennett
and Srinivasan (1983) and Bolli and Saunders (1985), which vary
slightly from each other. Revised datums presented by Berggren
et al. (1995) are incorporated, especially if they specified their
relevance to the temperate zone. Age ranges of deep-dwelling
benthic species are based on van Morkhoven, Berggren and
Edwards (1986), Jones (1994), and Hayward et al. (2012). For
any other benthic taxon, the known age indicated is that between
its type age and its occurrence in the Chilean Miocene.
Upper depth limit: The shallowest depth zone in which the
species has been recorded. Derived primarily from bathymetric
ranges presented in Bandy and Rodolfo (1964), Hayward et
al. (2012), Ingle, Keller and Kolpack (1980), van Morkhoven,
Berggren and Edwards (1986), Figueroa et al. (2005, 2006),
Hayward et al. (2012), and Holbourn, Henderson and MacLeod
(2013). In a few cases, California UDLs recorded by Ingle
(1980) are incorporated, but only if shallower than the other
determinations, since they tend to be deeper in California than
in most other regions, or if data for a particular species was not
presented in any of the three primary references. If the UDL
is based on another species, that probable synonym, isomorph,
or comparable morphospecies is indicated. Numerical depths
are assigned to depth zones according to the Southeast Pacific
scheme of Ingle, Keller and Kolpack(1980).
375
Remarks: Distinguishing features or comments about the Chilean

specimens, but excluding comparisons with other species, which
are the next category.
Upper depth limit: Lower bathyal; based on Bathysiphon sp.

recorded off Chile by Bandy and Rodolfo (1964) and Ingle,
Keller and Kolpack (1980).
Comparative species: Features that distinguish the Chilean

species from similar taxa, which are referred to by their original
binomen with author and date, age, and location; uncertain
synonymies are noted.
Remarks: The illustrated specimen recovered is white and

aplitic. Electron microscopy and spectrum analysis reveal that it
is completely covered by flat spiraling rosettes of gypsum.
Occurrence: Refers to the geologic units and their localities

where the species is recorded in this study (see text-figures 15).
Maximum relative abundance: Based on the following
percentages calculated for each assemblage: 1 specimen = VR
(very rare), 2 specimens or <1% = R (rare), 1% to <5% = F (few),
5% to <25% = C (common), 25% = A (abundant).
Superkingdom Eukaryota Chatton 1925
Kingdom Rhizaria Cavalier-Smith 2002
Superphylum RETARIA Cavalier-Smith 2002
Phylum FORAMINIFERA (dOrbigny 1826) Lee 1990
Class Polythalamea Ehrenberg 1838
Order ASTRORHIZIDA Brady 1881
Family BATHYSIPHONIDAE Avnimelech 1952
Bathysiphon M. Sars, in G.O. Sars 1872
Type species: Bathysiphon filiformis M. Sars, in G.O. Sars
1872.
Bathysiphon giganteus Cushman
Plate 1, Figure 1
Bathysiphon falvidus var. giganteus Cushman 1917a, p. 651;

type-figure in Cushman 1921, pl. 2, fig. 4.
Type age and locality: Recent, Caribbean, 1997m.
Comparative species: Bathysiphon filiformis Sars 1872 (Recent,

Norway) has a smoother outline and transverse wrinkles.
Occurrence: El Peral beds (NLP), Navidad Fm. (PPP).
Maximum relative abundance: Rare.
Bathysiphon sp.
Plate 1, Figures 2, 3
Upper depth limit: Bathyal; based on its type occurrence and
Bathysiphon sp. recorded off Chile by Bandy and Rodolfo (1964)
and Ingle, Keller and Kolpack (1980).
Remarks: All specimens are short, collapsed segments of finegrained tubes.
Occurrence: Navidad Fm. (MPUP, PPP, PPT, RAP), Ranquil
Fm. (FRA, FRM, RAN), Santo Domingo Fm. (VAL), Lacui Fm.
(PCB).
Maximum relative abundance: Few.
Rhabdammina M. Sars, in Carpenter 1869
Type species: Rhabdammina abyssorum M. Sars, in Carpenter

1869.
Rhabdammina abyssorum M. Sars, in Carpenter 1869
PLATE 1
Figures 46, 15, 17, and 22 are photomicrographs; all other images are SEMs. Scale bars in m.
1 Bathysiphon giganteus Cushman, UCMP50000, RAP.

2, 3 B
athysiphon sp.: 2, UCMP50001, PCB; 3, UCMP50002,
FRA
4, 5 Rhabdammina abyssorum Sars, PPP: 4, UCMP50003. 5,
UCMP50004.
6 Ammodiscus incertus (dOrbigny), UCMP50005, FRM.
7 Reophax agglutinatus Cushman, UCMP50006, FRA.
13, 14 H
aplophragmoides sp.: 13, UCMP50012, PPP. 14, UCMP
50013, CUC.
1518 Cyclammina cancellata (Brady): 15, UCMP50014, CHO.
16, UCMP50015, MAT. 17, UCMP50016, CHO. 18, typ
ical large specimen covered with matrix, UCMP50017,
MAT.
8 Reophax sp., UCMP50007, VAL.
19 A
mmobaculites agglutinans (dOrbigny), UCMP50018,
PPP.
9 H
aplophragmoides impressus Voloshinova, UCMP50008,
MOS
20 A
mmobaculites exilis Cushman and Brnnimann, UCMP
50019, MS10.
10 H
aplophragmoides mexicanus Kornfeld, UCMP50009,
PTA.
11 Haplophragmoides pulicosus Saidova, UCMP50010, FRA.
376
12 A
lveophragmium orbiculatum Shchedrina, UCMP50011,
PCB.
21 Vulvulina pacifica Cushman, UCMP50020, RQT.

22 T
ritaxis challengeri (Hedley, Hurdle and Burdett), UCMP
50021, PPP.
Kenneth L. Finger
Micropaleontology, vol. 59, nos. 45
Plate 1
377
Rhabdammina abyssorum M. Sars, in Carpenter 1869, p. 60,

type-figure not given. BOLTOVSKOY and THEYER 1970, p.
355, pl. 4, fig. 26; pl. 5, figs. 3, 4. INGLE, KELLER and KOL
PACK 1980, pl. 9, fig. 8. JONES 1994, p. 32, pl. 21, figs. 1 8,
1013.
Occurrence: Navidad Fm. (PPP).

Order LITUOLIDA Lankester 1885
Superfamily AMMODISCOIDEA Reuss 1862
Family AMMODISCIDAE Reuss 1862
Subfamily AMMOVOLUMMININAE Chernykh 1967
Ammodiscus Reuss 1862; emend. Rhumbler 1895 and Loeb
lich and Tappan 1954.
Type species: Ammodiscus infimus L. G. Bornemann 1874.
Ammodiscus incertus (dOrbigny 1839a)
Plate 1, Figure 6
Operculina incerta DORBIGNY 1839a, p. 49, pl. 8, figs. 16, 17.

Ammodiscus incertus (dOrbigny). INGLE, KELLER and KOLPACK
1980, pl. 9, fig. 9.
Remarks: Smaller specimens tend to be circular, while larger

ones often become ovate.
Comparative species: The distinction between dOrbignys
species and others is blurred by growth rate, growth stage,
micro
spheric vs. megalospheric dimorphism, and test com
position derived from the local substrate. Possible synonyms
of dOrbignys species include Ammodiscus miocenicus Karrer
1877 (Miocene, central Europe), A. tenuis (= Trochammina
(Ammodiscus) tenuis Brady 1884; Recent, multiple localities),
and A. discoideus (A. incertus var. discoideus Cushman 1917a;
Recent, East Indies, 3060m).
Upper depth limit: Upper middle bathyal; based on A. tenuis and
A. incertus pacificus, which have the shallowest UDLs for the
genus off Chile (table 8).
Occurrence: Ranquil Fm. (FRA).

Maximum relative abundance: Very rare.
Reophax sp.
Plate 1, Figure 8
Distinguishing features: 45 subspherical chambers, rapidly
increasing in size.
Occurrence: Santo Domingo Fm. (VAL).
Superfamily LITUOLOIDEA de Blainville 1827
Family HAPLOPHRAGMOIDIDAE Maync 1952
Haplophragmoides Cushman 1910a
Type species: Nonionina canariensis dOrbigny 1839b.
Haplophragmoides impressus Voloshinova, in Voloshinova and
Budasheva 1961
Plate 1, Figure 9
Haplophragmoides impressus Voloshinova, in Voloshinova and

Budasheva 1961, p. 192, pl. 5, figs. 57.
Type age and locality: Late Miocene, Sakhalin Island (NW

Pacific).
Upper depth limit: Upper bathyal; conservatively based on most
of the extant congeneric species.
Comparative species: This Chilean form is coarser grained
than Haplophragmoides scitulum (= Lituola (Haplophragmium)
scitulum Brady 1881; Recent, Faroe Channel, Scotland).
Occurrence: Navidad Fm. (MOS, PPP), Ranquil Fm. (MS10),
Lacui Fm. (CUC).
Haplophragmoides mexicanus Kornfeld 1931
Plate 1, Figure 10
Remarks: Large specimens tend to be less flat and less circular

than smaller ones.
Haplophragmoides canariensis var. mexicana Kornfeld 1931, p.

83, pl. 13, fig. 4.
Occurrence: Navidad Fm. (CPUP, MPUP, PPP), Ranquil Fm.

(FRA, FRM), Santo Domingo Fm. (VAL), Lacui Fm. (CHO).
Type age and locality: Recent, Louisiana; neritic.
Superfamily HORMOSINOIDEA Haeckel 1894

Family HORMOSINIDAE Haeckel 1894
Subfamily REOPHACINAE Cushman 1910a
REOPHAX de Montfort 1808
Type species: Reophax scorpiurus de Montfort 1808.
Reophax agglutinatus Cushman 1913a
Plate 1, Figure 7
Reophax agglutinatus CUSHMAN 1913a, p. 637, pl. 79, fig. 6.
Type age and locality: Recent, Philippines, 732m.

Upper depth limit: Upper middle bathyal; based on type occurrence.
Remarks: Represented by a single, small specimen.
378
Remarks: Specimens match the holotype by being very

compressed, six-chambered forms.
Occurrence: El Peral beds (NLP), Navidad Fm. (MOS, PPP,
PPT, PTA), Ranquil Fm. (MS10), Valdivia (VAL).
Maximum relative abundance: Common (MOS).
Haplophragmoides pulicosus Saidova 1970
Plate 1, Figure 11
Haplophragmoides pulicosus Saidova 1970, p. 150, pl. 4, fig. 4.
Upper depth limit: Lower bathyal, based on Saidovas (1970)

report of the species at depths ranging from 2611m in the
Southern Ocean to 7320m in the North Pacific.
Remarks: The Chilean specimens have four (or possibly five)
chambers rapidly increasing in size and distinctly depressed
sutures. The type specimen is finer grained and described as

having five chambers, although its fifth chamber is barely visible.
Comparative species: This species bears some resemblance
to the 45 chambered Haplophragmoides globigerinoides (=
Trochammina globigerinoides Haeusler 1882, emend. Osterle
1968), which is a less lobulate form from the Late Jurassic.
Type age and locality: Recent, Kurile Islands, 5095m.
Haplophragmoides spp.
Remarks: These are rather nondescript internal molds of

somewhat inflated planispiral forms, many of which may be
juvenile Cyclammina cancellata.
Comparative species: Ammobaculites salsus Cushman and

Brnnimann 1948 (Recent, Trinidad, mangrove swamp) has
sutures on its uniserial portion that dip toward the coil and its
penultimate chamber accounts for half of the test size.
Occurrence: Ranquil Fm. (MS10).
Superfamily LOFTUSIOIDEA Brady 1884
Family CYCLAMMINIDAE Marie 1941
Subfamily CYCLAMMININAE Marie 1941
Alveophragmium Shchedrina 1936
Type species: Alveolophragmium orbiculatum Shchedrina
1936.
Alveolophragmium orbiculatum Shchedrina 1936
Plate 1, Figure 12
Alveolophragmium orbiculatum SHCHEDRINA 1936, p. 315, text-fig. 1.
Occurrence: Navidad Fm. (MOS, RAP, PPP, PPT, PTA, MPUP,

CPUP), Ranquil Fm. (FRA, FRM, MS10, RAN, RQK, RQT),
Santo Domingo Fm. (VAL), Lacui Fm. (CUC, PCB).
Type age and locality: Recent, Japan Sea, 57200m.
Maximum relative abundance: Common (MPUP, CUC).
Upper depth limit: Outer neritic.
Family LITUOLIDAE de Blainville 1827

Subfamily AMMOMARGINULININAE Podobina 1978
Ammobaculites Cushman 1910a; emend. Hglund 1947
Type species: Spirolina agglutinans dOrbigny 1846.
Occurrence: Navidad Fm. (LBZ), Lacui Fm. (PCB).
Ammobaculites agglutinans (dOrbigny 1846); emend. Barten

stein 1952
Plate 1, Figure 19
Spirolina agglutinans DORBIGNY 1846, p. 137, pl. 7, figs. 1012.

Haplophragmium agglutinans (dOrbigny). BRADY 1884, pl. 32, figs.
19, 20, 2426.
Ammobaculites midwayensis PLUMMER 1933, p. 63, pl. 5, figs. 711.
? Ammobaculites subagglutinans BANDY 1949a, p. 27, pl. 3, figs. 5a, b.
Ammobaculites agglutinans (dOrbigny). BARTENSTEIN 1952, p.
318, pl. 1, fig. 1; pl. 2, figs. 1016. PAPP and SCHMID 1985, p.
54, pl. 45, figs. 69. JONES 1994, p. 39, pl. 32, figs. 19, 20, 24
26. LOEBLICH and TAPPAN 1987, p. 74, pl. 58, figs. 34.
KAMINSKI and GRADSTEIN 2005, p. 324, fig. 70, 13. HOL
BOURN, HENDERSON and MACLEOD 2013, p. 2627.
Type age and locality: Middle Miocene (Badenian), Vienna

Basin, Austria.
Stratigraphic range: Late Cretaceous to Recent.
Upper depth limit: Inner neritic.
Occurrence: Navidad Fm. (PPP), Ranquil Fm. (MS10), Santo
Domingo Fm. (VAL).
Ammobaculites exilis Cushman and Brnnimann 1948
Plate 1, Figure 20
Ammobaculites exilis Cushman and Brnnimann 1948, p. 39,

pl. 7, fig. 9.
Type age and locality: Recent, Trinidad, 012m.

Upper depth limit: Inner neritic; based on type locality.
Stratigraphic range: Early Miocene to Recent.

Subfamily CYCLAMMININAE Marie 1941
Cyclammina Brady 1879a
Type species: Cyclammina cancellata Brady 1879a.
Cyclammina cancellata Brady 1879a
Plate 1, Figures 1518
Cyclammina cancellata BRADY 1879a, p. 62, pl. 37, figs. 816.

BRADY 1884, p. 351, pl. 37, figs. 816. GRADSTEIN and
BERGGREN 1981, p. 254, pl. 7, figs. 13. BOLTOVSKOY and
THEYER 1970, p. 323, pl. 2, fig. 11. KOHL 1985, p. 28, pl. 1,
fig. 4. LOEBLICH and TAPPAN 1987, pl. 107, figs. 26.
JONES, p. 43, pl. 37, figs. 816. ZAPATA and CEAR 2004, p. 4,
fig. 12. HOLBOURN, HENDERSON and MACLEOD 2013, p.
222223.
Type age and locality: Recent; widespread localities, none

shallower than 640m.
Upper depth limit: Upper middle bathyal (Theyer, 1971; Ingle,
Keller and Kolpack 1980).
Remarks: Most of the larger Chilean specimens resemble
Haplophragmoides grandiformis Cushman 1910b (Recent,
East Indies, 204m), as they are relatively huge and appear to
be coarsely agglutinated because a granular, often glauconitic,
veneer obscures the very fine-grained, many-chambered
tests typical of Cyclammina. The original test material is not
preserved, and most specimens appear to be casts; apertures
were not observed, but many specimens have regular patterns of
black grains that reflect the labyrinthic interior, which was clearly
observed on some broken fragments. Many smaller specimens of
similar composition were recovered, some showing the numerous
chambers, but most had indecipherable sutures. Attempts to split
them into discrete morphotypes based on dimensions were futile.
Whereas no other forms were preserved as casts, I assume they
379
are of the same species. Similarly obscure specimens have been

referred to this species by Kaminski and Gradstein (2005).
At the Peru-Chile Trench, Theyer (1971) found C. cancellata
increases in size from 0.6mm at 463m to 6.3mm at 2120m. The
largest Chilean Miocene specimens range 45mm, matching
what Theyer recorded from depths between 1000 and 4000m.
Comparative species: In his redescription of Haplophragmium
incisum Stache 1865 (Late Tertiary, New Zealand), Hornibrook
(1971) examined topotypes and found them to have a labryinthic
internal architecture, but none had the porous apertural face
characteristic of Cyclammina; nevertheless, he transferred
the species to that genus stating The possibility that true
Cyclammina has a considerable depth range and produces a
porous apertural face more consistently at greater depths seems
worthy of consideration. He noted that the species otherwise
bears close resemblance both externally and internally to
Cyclammina cancellata. If Staches species has the apertural
pores, it would match Cyclammina bradyi Cushman 1910a
(Recent; localities given in N Pacific at 2358 and 5266m),
which is probably synonymous with C. cancellata. If that were
the case, C. incisa would have priority. The Chilean Miocene
specimens are assigned to C. cancellata to conform with the
modern provincial literature and because they similarly attain
much larger sizes and more chambers in the outer whorl than
indicated in the type descriptions of the other two species.
Tritaxis challengeri (Hedley, Hurdle and Burdett 1964)
Plate 1, Figure 22
Trochammina challengeri Hedley, Hurdle and Burdett 1964,

p. 425; type-figure in Brady 1884, pl. 41, fig. 3 (as Trochammina
squamata Jones and Parker). JONES 1994, p. 46, pl. 41, fig. 3.
Type age and locality: Recent, Virgin Islands, 2340m.

Upper depth limit: Outer neritic (Jones 1994).
Trochamminopsis Brnnimann 1976
Type species: Trochammina quadriloba Hglund 1948.
Trochamminopsis quadriloba Hglund 1948
Plate 2, Figure 1
Trochammina quadriloba Hglund 1948, p. 46 (nom. nov. pro T.
pusilla Hglund 1947, p. 201, pl. 17, fig. 4).
Type age and locality: Recent, Skagerrak (S of Sweden), 366m.
Occurrence: Navidad Fm. (CPUP, MAT, MPUP, NAV5, PPP,

PPT, PTA), Ranquil Fm. (FRM, MS10, RAN, RQT), Santo
Domingo Fm. (VAL), Lacui Fm. (CHO, CUC, PCB).
Maximum relative abundance: Abundant.
Occurrence: Ranquil Fm. (FRA), Santo Domingo Fm. (VAL).
Superfamily SPIROPLECTAMMINOIDEA Cushman 1927b

Family SPIROPLECTAMMINIDAE Cushman 1927b
Subfamily VULVULININAE Saidova 1981
Vulvulina dOrbigny 1826
Type species: Vulvulina capreolus Defrance, in dOrbigny
1826.
Vulvulina pacifica Cushman 1932b
Plate 1, Figure 21
Vulvulina pacifica Cushman 1932b, p. 78, pl. 10, figs. 8, 9.

Vulvulina jarvisi Cushman 1932b (Eocene, Trinidad). ROBERTSON
1998, p. 16, pl. 1, fig. 3 (Miocene, Jamaica).

Upper depth limit: Upper middle bathyal; based on its type
locality.
Comparative species: Vulvulina pennatula has raised sutures
and deeply excavated chambers, and lacks a spinose periphery.
Upper depth limit: Upper bathyal; based on its type locality.
Order TEXTULARIINA Ehrenberg 1839

Superfamily VERNEUILINOIDEA Cushman 1911
Family PROLIXOPLECTIDAE Loeblich and Tappan 1985
Karrerulina Finlay 1940
Type species: Karrerulina apicularis Cushman 1911.
Karrerulina apicularis (Cushman 1911)
Plate 2, Figure 3
Gaudryina apicularis Cushman 1911, p. 69, tf. 110.

Karrerulina apicularis (Cushman). Loeblich and Tappan 1987,
pl. 139, figs. 79.
Type age and locality: Recent, type locality not designated;

localities noted in North Pacific at 38407224m.
Upper depth limit: Lower bathyal, based on Loeblich and Tappan
(1987) for genus.
Occurrence: Ranquil Fm. (RAN, RQT).
Occurrence: Navidad Fm. (MPUP, PPP), Ranquil Fm. (MS10),

Santo Domingo Fm. (VAL).
Order TROCHAMMINIDA Schwager 1877

Superfamily TROCHAMMINOIDEA Schwager 1877
Family TROCHAMMINIDAE Schwager 1877
Subfamily TROCHAMMININAE Schwager 1877
Tritaxis Schubert 1921
Type species: Rotalina fusca Williamson 1858.
Family VERNEUILINIDAE Cushman 1911

Subfamily VERNEUILININAE Cushman 1911
Gaudryina dOrbigny 1840
Type species: Gaudryina rugosa dOrbigny 1840.
380

Gaudryina sp.
Plate 2, Figure 13
Remarks: The single specimen from the Chilean Neogene is
somewhat similar to Gaudryina frankei Brotzen 1936 (Cretaceous,
Sweden), which is less robust. Gaudryina trinitatensis Nuttall
1928 (Late Oligocene or Early Miocene, Trinidad) and G.
pliocenica Cushman, Stewart and Stewart 1949 (Pliocene,
Washington) similarly have a distinct triserial stage but those
species are acutely triangular in cross section and have more
numerous biserial chambers that are not robust.
Occurrence: Ranquil Fm. (FRM).
Family TEXTULARIELLIDAE Grnhagen and Luterbacher 1966
Guppyella Brnnimann 1951
Type species: Goesella miocenica Cushman 1936a.
Guppyella crassa (Cushman and Renz 1941)
Plate 2, Figure 4
Liebusella pozonensis var. crassa Cushman and Renz 1941, p. 10,

pl. 2, figs. 3, 4a.
Type age and locality: Late Oligocene, Venezuela.

Stratigraphic range: Late Oligocene to Early Miocene.
Occurrence: Navidad Fm. (CPUP, MOS, PPP, PPT, PTA, RAP),
Ranquil Fm. (FRM, MS10, RQT), Santo Domingo Fm. (VAL),
Lacui Fm. (CHO).
Verneuilina bradyi Cushman 1911, p. 54, text-fig. 87.

Eggerella bradyi (Cushman). KOHL 1985, p. 32, pl. 3, fig. 3. LOE
BLICH and TAPPAN 1987, pl. 189, figs. 1, 2. JONES 1994, pl.
47, figs. 47. ROBERTSON 1998, p. 24, pl. 3, fig. 4. HAY
WARD, GRENFELL, SABAA, NEIL and BUZAS 2010, p. 144, pl.
5, figs. 23, 24. HOLBOURN, HENDERSON and MACLEOD
2013, p. 232233.
Type age and locality: Recent, North Pacific, >1829m.

Stratigraphic range: Late Eocene to Recent.
Upper depth limit: Upper middle bathyal based on New Zealand
records (Hayward et al. 2010) and middle bathyal assignment in
van Morkhoven, Berggren and Edwards (1986).
Occurrence: Ranquil Fm. (FRA, RAN).
Eggerelloides Haynes 1973
Type species: Verneuilina bradyi Cushman 1911.
Eggerelloides scabrus (Williamson 1858)
Plate 2, Figure 7
Bulimina scabra Williamson 1858, p. 65, pl. 5, figs. 136, 137.

Eggerelloides scabrus (Williamson). LOEBLICH and TAPPAN 1987,
pl. 189, figs. 57.
Eggerelloides scaber (Williamson). JONES 1994, p. 52, figs. 1517.
Type age and locality: Recent, type locality not designated, but
localities given in United Kingdom, depths not indicated.
Superfamily TEXTULARIOIDEA Ehrenberg 1838

Family EGGERELLIDAE Cushman 1937
Subfamily DOROTHIINAE Balakhmatova 1972
Dorothia Plummer 1931; emend. Desai and Banner 1987
Type species: Gaudryina bulletta Carsey 1926.
Type species: Gaudryina siphonella Reuss 1851b.
Dorothia cylindrica (Nuttall 1932)
Karreriella biglobata Finger, n. sp.
Gaudryina cylindrica Nuttall 1932, p. 7, pl. 2, fig. 7.
Description: Test robust with initial triserial stage followed by an

elongate biserial stage comprised of at least two pairs of inflated
rectilinear chambers succeeded by two abruptly larger and
nearly globular chambers; periphery rounded and increasingly
lobulate; aperture slightly intercameral, elongate, very narrow,
with slight lip.
Plate 2, Figure 5
Type age and locality: Oligocene, Mexico.

Stratigraphic range: Oligocene to Early Miocene.
Upper depth limit: Upper bathyal; based on Dorothia spp.
recorded as bathyal off Chile by Bandy and Rodolfo (1964); also
upper bathyal in Southwest Pacific (Hayward et al. 2012).
Comparative species: Karreriella cylindrica Finlay 1940 (Late
Miocene, New Zealand) has an areal aperture.
Occurrence: Navidad Fm. (MPUP, PPP, RAP), Ranquil Fm.
(FRM, RAN, RQT), Lacui Fm. (CHO, PCB).
Maximum relative abundance: Common (FRM).
Subfamily EGGERELLINAE Cushman 1937
EGGERELLA Cushman 1933b
Type species: Verneuilina bradyi Cushman 1911.
Eggerella bradyi (Cushman 1911)
Plate 2, Figure 6

Karreriella Cushman 1933b
Plate 2, Figure 2
Remarks: Although represented by a single specimen, this form

is quite distinct from all known species of Karreriella. The
populations of K. bradyi show little variation to suggest this is an
aberrant individual of that species. Some species of Gaudryina,
however, are characterized by a similar dramatic inflation of the
last two chambers.
Occurrence: Ranquil Fm. (FRM) type locality.
Holotype: UCMP50023.
Type age and locality: Early Miocene; Ranquil Formation in
coastal bluff level below intertidal platform of Punta Fraile,
coastal central Chile.
381
Etymology: Morphodescriptor derived from the Latin bi +

globatus, meaning two spheres, in reference to the shape of the
final chambers.
Karreriella bradyi (Cushman 1911)
Plate 2, Figure 8
Gaudryina bradyi Cushman 1911, p. 67; text-fig. 107.

Karreriella bradyi (Cushman). KOHL 1985, p. 32, figs. 4, 5.
JONES 1994, p. 50, pl. 46, figs. 14. ROBERTSON 1998, p.
26, pl. 4, fig. 1. HAYWARD, GRENFELL, SABAA, NEIL and
BUZAS 2010, p. 145, pl. 5, figs. 2527. HOLBOURN, HEN
DERSON and MACLEOD 2013, p. 318319.
Type age and locality: Recent; localities given have an average

depth of 1829m off Hawaiian Islands, Galpagos Islands, Guam,
and Japan.
Upper depth limit: Middle bathyal; based on occurrences off
New Zealand (Hayward et al. 2010); Bandy and Rodolfo (1964)
report it as bathyal.
Occurrence: Ranquil Fm. (FRA, FRM, MIB), Lacui Fm. (PCB).
Karreriella subcylindrica (Nuttall 1932)
Gaudryina subcylindrica Nuttall 1932, p. 76, pl. 3, figs. 17, 18.
Type age and locality: Early Miocene, Trinidad.

Stratigraphic range: Early Miocene.
Upper depth limit: Middle bathyal; based on Karreriella bradyi
in New Zealand (Hayward et al. 2010).
Remarks: Many of the Chilean specimens of K. bradyi are
relatively large, including robustly fusiform juveniles.
Comparative species: Karreriella caribaea (= K. subcylindrica
var. caribaea Acosta 1940; Recent, Cuba, 1481m) is more
slender with higher chambers.
Occurrence: El Peral beds (LPER, NLP), Navidad Fm. (MPUP,

PTA), Ranquil Fm. (FRA, FRM, MIB, RAN, RQK, RQT).
Maximum relative abundance: Common (MIB).
Martinottiella Cushman 1933b
Type species: Clavulina communis dOrbigny 1826.
Martinottiella communis (dOrbigny 1846)
Plate 2, Figure 12
Clavulina communis DOrbigny 1846, p. 196, pl. 12, figs. 1, 2.

Martinottiella communis (dOrbigny). INGLE, KELLER and KOL
PACK 1980, pl. 4, fig. 14, 15. PAPP and SCHMID 1985, p. 74,
pl. 66, figs. 18. KOHL 1985, p. 33, pl. 4, fig. 2. LOEBLICH
and TAPPAN 1987, pl. 190, figs. 3, 4. JONES 1994, p. 52, pl. 66,
figs. 18. HOLBOURN, HENDERSON and MACLEOD 2013,
p. 350351.

Basin, Austria.
Upper depth limit: Upper bathyal; based on its occurrence off
Chile recorded by Ingle, Keller and Kolpack (1980). Its UDL off
New Zealand is the same (Hayward et al. 2010).
Remarks: The neritic specimen identified as this species in
Zapata and Cear (2004) is much coarser and is more similar to M.
juncea. Multifidella nodosa is a very similar form distinguished
by multiple apertures.
Occurrence: El Peral beds (LPER, NLP), Navidad Fm. (CPUP,
MOS, MPUP, NAV5, PPP, PPT, PTA), Ranquil Fm. (FRM,
MS10, RAN, RQK, RQT), Santo Domingo Fm. (VAL), Lacui
Fm. (CHE, CUC, PCB, PNH).
Maximum relative abundance: Common (MS10)
Martinottiella juncea (Cushman 1936a)
Plate 2, Figure 11
Pseudoclavulina juncea Cushman 1936a, p. 19, pl. 3, fig. 8.
PLATE 2
Figures 7 and 15 are photomicrographs; all other images are SEMs. Scale bars in m.
1 T
rochamminopsis quadriloba (Hglund), UCMP50022,
FRA.
2 K
arreriella biglobata Finger, n. sp., holotype UCMP50023,
FRM.
382
8 Karreriella bradyi (Cushman), UCMP50029, FRA.

9, 10 K
arreriella subcylindrica (Nuttall): 9, without bise
rial stage UCMP50030, PTA. 10, with biserial stage,
UCMP50031, FRA.
3 Karrerulina apicularis Cushman, UCMP50024, MS10.
11 Martinottiella juncea (Cushman), UCMP50032, FRA.
4 Guppyella crassa (Cushman and Renz), UCMP50025, PPP.
12 Martinottiella communis (dOrbigny), UCMP50033, PPP.
5 Dorothia cylindrica (Nuttall), UCMP50026, RAP.
13 Gaudryina sp., UCMP50034, FRM.
6 Eggerella bradyi (Cushman), UCMP50027, RAN.
14 Textularia lythostrota (Schwager), UCMP50035, FRA.
7 E
ggerelloides scabrus (Williamson), crushed specimen,
UCMP50028, PPP.
15 Martinottiella pallida (Cushman), UCMP50036, PPP.

16 Textularia miozea Finlay, UCMP50037, NAV5.
Kenneth L. Finger
Plate 2
383
Upper depth limit: Middle bathyal; based on its type occurrence.

Comparative species: Morphology is very similar to that of
Martinottiella perrosa (= Listerella perrosa Cushman and
Bermdez 1937; Eocene, Cuba).
Type age and locality: Recent, Caribbean, depth not recorded.

Occurrence: Navidad Fm. (PPT), Ranquil Fm. (FRA).
Occurrence: El Peral beds (NLP), Navidad Fm. (CPUP, MPUP,

PPP, PPT, PTA), Ranquil Fm. (RAN), Lacui Fm. (CHO).
Martinottiella pallida (Cushman 1927f)
Textularia lythostrota (Schwager 1866)
Clavulina communis var. pallida Cushman 1927f, p. 138, pl. 2, fig.

1.
Plecanium lithostrotum Schwager 1866, p. 194, pl. 4, fig. 4.

Textularia lythostrota (Schwager). KOHL 1985, p. 30, pl. 2, fig, 2.
Type age and locality: Recent, off California at 850m.
Type age and locality: Early to Middle Pliocene, Car Nicobar.
Remarks: Differs from Martinottiella communis in having a

white test that is much smaller in size and apparently weaker in
construction, as it often diagenetically compressed.
Occurrence: Navidad Fm. (NAV5, PPP, PPT, PTA), Ranquil Fm.

(FRA), Lacui Fm. (PCB).
Plate 2, Figure 15
Occurrence: Navidad Fm. (PPP, RAP), Ranquil Fm. (RQT),

Santo Domingo Fm. (VAL).
Family TEXTULARIIDAE Ehrenberg 1838
Subfamily TEXTULARIINAE Ehrenberg 1838
Textularia Defrance, in de Blainville 1824
Type species: Textularia sagittula Defrance, in de Blainville
1824.
Remarks: Although Textularia spp. in Ingle, Keller and Kolpack
(1980) were not found above the upper middle bathyal zone, the
genus is common in neritic assemblages elsewhere.
Textularia agglutinans dOrbigny 1839a
Textularia agglutinans DOrbigny 1839a, p. 144, pl. 1, fig. 17, 18,

32, 34. JONES 1994, p. 48, pl. 43, figs. 13.
Plate 2, Figure 14

Textularia miozea Finlay 1939c
Plate 2, Figure 16
Textularia cf. T. miozea Finlay 1939c, p. 326, pl. 129, figs. 159161.
Type age and locality: Middle Miocene, New Zealand.

Occurrence: Navidad Fm. (CPUP, NAV5, PPT), Ranquil Fm.
(RQT).
Textularia schencki Cushman and Valentine 1930
Plate 3, Figure 3
Textularia schencki Cushman and Valentine 1930, p. 8, pl. 1,

fig. 3.
Type age and locality: Recent, California, 537m.
PLATE 3
1, 2 Textularia agglutinans dOrbigny: 1, UCMP50038, CHO.

2, UCMP50039, PTA.
3 Textularia schencki Cushman and Valentine, UCMP50040,
PCB.
4 Textularia sp. A, UCMP50041, PNH.
384
10 Cornuspira planorbis Schultze, UCMP50047, FRA.

11 Cornuspiroides foliaceus (Philippi), UCMP50048, RAP.
12 Cornuspira veleronis (McCulloch), UCMP50049, PPP.
13 Nummulopyrgo globulus (Bornemann), UCMP50050, NLP.
5 Textularia sp. B., UCMP50042, NLP.
14 Spiroloculina robusta Brady, UCMP50051, NLP.
6 Textularia sp. C, UCMP50043, RQT.
15 Spiroloculina incisa Cushman, UCMP50052, FRA.
7 Textularia? sp., UCMP50044, PTA.
16 Quinqueloculina akneriana dOrbigny, UCMP50053, MIB.
8 Pseudoclavulina mexicana (Cushman), UCMP50045, FRA.
17 Quinqueloculina badenensis dOrbigny, UCMP50054, RAP.
9 C
ornuspira libella Finger, n. sp., holotype UCMP50046,
FRA.
18 Q
uinqueloculina cf. Q. benwestonensis McCulloch, UCMP50055, FRA.
Kenneth L. Finger
Plate 3
385
Upper depth limit: Inner neritic; based on type locality and this
form identified as T. deltoidea by Zapata and Cear (2004).
Comparative species: Textularia secasensis Lalicker and
McCulloch 1940 (Recent, Pacific off Mexico) and T. candeiana
dOrbigny 1839a (Recent, Caribbean) have a similar flaring
test shape but more inflated chambers. Gaudryina subglabrata
Cushman and McCulloch 1939 (southern California, Recent,
13m) is nearly identical to T. schencki, but has a very early
triserial stage.
Occurrence: Navidad Fm. (PPP), Ranquil Fm. (RAN), Lacui
Fm. (PCB).
Comparative species: Clavulina serventyi Chapman and Parr

1935 (Recent, off southern Australia, 128m) differs primarily
in the shape of the apertural face. Clavulina pacifica Cushman
1924 (Recent, off Samoa, 46m) is triangular throughout the
length of its test.
Occurrence: El Peral beds (LPER, NLP), Navidad Fm. (MOS,
PPP, PTA), Ranquil Fm. (FRA, FRM, LEB, MS10, RAN, RQT),
Lacui Fm. (PCB).
Plate 3, Figure 4
Order MILIOLIDA Lankester 1885

Superfamily CORNUSPIROIDEA von Schultze 1854
Family CORNUSPIRIDAE von Schultze 1854
Subfamily CORNUSPIRINAE von Schultze 1854
Cornuspira von Schultze 1854
Type species: Orbis foliaceus Philippi 1844.
Occurrence: Ranquil Fm. (RAN), Lacui Fm. (PNH).
Cornuspira libella Finger, n. sp.

Textularia sp. A

Textularia sp. B
Plate 3, Figure 5
Occurrence: El Peral beds (NLP).
Textularia sp. C
Plate 3, Figure 6
Occurrence: Ranquil Fm. (RQT).
Textularia? sp.
Plate 3, Figure 7
Remarks: The single specimen recovered is a slightly twisted
form consisting mostly of a biserial stage followed by what
appears to be a short uniserial stage, but its aperture is not
discernible and the test does not resemble the early stage of
any biserial-to-uniserial species in the fauna; hence, the generic
placement of this specimen is uncertain.
Occurrence: Lacui Fm. (CHO).
Family PSEUDOGAUDRYINIDAE Loeblich and Tappan 1985
Subfamily PSEUDOGAUDRYININAE Loeblich and Tappan 1985
Pseudoclavulina Cushman 1936
Type species: Clavulina clavata Cushman 1926a.
Pseudoclavulina mexicana (Cushman 1922a)
Plate 3, Figure 8
Clavulina humilus Brady var. mexicana Cushman 1922a, p. 83;

type-figure not given, but pl. 16, figs. 13 are paratypes.
Clavulina mexicana Cushman. KOHL 1985, p. 33, pl. 4, fig. 1.
ROBERTSON 1998, p. 32, pl. 7, fig. 1.
Type age and locality: Recent, Gulf of Mexico, 384m.

Upper depth limit: Upper bathyal; based on type locality.
386
Plate 3, Figure 9
Description: Test an opaque, flat planispiral of 3+ whorls;

peripheral edge subrounded; ornamented with nodes separated
by a smooth interspiral ridge. Whorls progressively increase
their number of nodes from 24 to 42, accompanied by a slight
increase in widths of nodes and interspiral ridge. Nodal pattern
resembles a coiled string of beads, and is identical on both sides
of test.
Remarks: This form is unusual for the genus in possessing
ornamentation, and therefore may represent a new genus. Though
only one specimen was recovered, its exquisite ornamentation
clearly distinguishes it as a unique species worthy of description.
The ornamentation of this form is more akin to spirillinids,
which are typically vitreous and relatively minute.
Comparative species: The ornate pattern of Cornuspira libella
resembles the interspirally grooved side of Turrispirillina
denticulogranulata (= Spirillina denticulo-granulata Chapman
1907; off Australia, depth not indicated). Another similar
pattern of ornamentation is that of S. seymourensis McCulloch
1977 (Galpagos, 24m), which has the spiral nodes but lacks the
spiral ridge. Spirillina nodosa Terquem 1880 (Recent, northern
France) has 67 whorls and the nodules tend to coalesce into
radial ridges.
Type age and locality: Early Miocene, Ranquil Formation
exposed in coastal bluffs at Punta El Fraile (Ranquil Fm. locality
FRA), west of Tubul.
Etymology: Morphodescriptor from the Latin libella, meaning
small silver coin.
Cornuspira planorbis von Schultze 1854
Plate 3, Figure 10
Cornuspira planorbis VON SCHULTZE 1854, p. 40, pl. 2, fig. 21.
Type age and locality: Recent, off Mozambique, depth not

indicated.
Comparative species: Cornuspira involvens (= Operculina

involvens Reuss 1850; Tertiary, Germany) has an acute periphery
and its outer whorl is several times wider than the previous whorl.
Upper depth limit: Neritic (for C. involvens in Zapata and Cear
2004).
Occurrence: El Peral beds (NLP), Navidad Fm. (MOS, PTA),
Ranquil Fm. (FRA, MIB, MS10, RAN, RQT), Lacui Fm. (CHO).
Cornuspira veleronis (McCulloch 1981)

Spiroloculina dOrbigny 1826
Type species: Spiroloculina depressa dOrbigny 1826.
Spiroloculina incisa Cushman 1921
Plate 3, Figure 15
Spiroloculina grateloupi var. incisa Cushman 1921, p. 397, pl. 78,
fig. 5.
Spiroloculina alveata Cushman and TODD 1944, p. 28, pl. 4, figs.
29, 30. (Miocene, Jamaica)
Plate 3, Figure 12
Cyclogyra veleronis McCULLOCH 1981, p. 31, pl. 270, fig. 31.
Upper depth limit: Outer neritic; based on type locality (shelf

edge for Spiroloculina sp. in Ingle, Keller and Kolpack 1980).
Type age and locality: Recent, Gulf of Venezuela, 86m.

Remarks: The figured Chilean specimen has a robust test with
concave spiral face textured with very faint oblique furrows, and
subacute periphery with flat edge. Juvenile specimens lack the
furrows and have a more acute periphery. McCulloch described
small specimens without mention of furrows, which appear to be
juveniles.
Comparative species: The type figure of Cornuspira involvens
(= Operculina involvens Reuss 1850; Tertiary, Germany) has an
acute periphery and its outer whorl is several times wider than the
previous whorl.
Occurrence: Navidad Fm. (PPP), Ranquil Fm. (FRA).
Cornuspiroides Cushman 1928
Type species: Orbis foliaceus Philippi 1844.
Cornuspiroides foliaceus (Philippi 1844)
Plate 3, Figure 11
Orbis foliaceus Philippi 1844, p. 147, pl. 24, fig. 26.

Cornuspiroides foliaceus (Philippi). JONES 1994, p. 27, pl. 11, figs. 5, 6.
Cornuspiroides yabei ASANO 1951, p. 2, text-figs. 3, 4. (Japan, Plio
cene)
Type age and locality: Recent and Tertiary, Sicily; depth not
indicated for Recent specimens.
Occurrence: El Peral beds (NLP), Navidad Fm. (CPUP, PPP,
RAP), Lacui Fm. (CUC).
Superfamily MIOLIOLOIDEA Ehrenberg 1839
Family SPIROLOCULINIDAE Wiesner 1920
Nummulopyrgo Hofker 1983
Type species: Nummulopyrgo globulus (Bornemann 1855).
Comparative species: Spiroloculina depressa dOrbigny 1826

(fossil, Italy and Recent, Mediterranean) might be the same
species, but the holotype is figured as opposite sides of a broken
specimen and it lacks an original description. Spiroloculina
badenensis dOrbigny 1846 (Middle Miocene, Vienna Basin)
does not have deeply depressed sutures.
Occurrence: El Peral beds (LPER), Navidad Fm. (PPP, PPT,
RAP), Ranquil Fm. (FRA, FRM, MIB, RAN, RQK, RQT),
Lacui Fm. (CHO).
Spiroloculina robusta Brady 1884
Plate 3, Figure 14
Spiroloculina robusta BRADY 1884, p. 150, pl. 9, figs. 7, 8.

Distinguishing features: Rapid increase in inflation of
progressive chambers, edges relatively thick, deeply concave;
later chambers are progressively more enveloping, so that only
34 chambers are visible.
Comparative species: Spiroloculina incisa is not nearly as robust
and concave, showing a more gradual increase in the inflation of
progressive chambers that are also much less enveloping. The
overall shape of S. robusta resembles Pyrgo lucernulinoides
Ujii 1990, but differs in coiling mode.
Occurrence: El Peral beds (NLP), Navidad (PPP, PPT)
Family HAUERINIDAE Schwager 1876
Subfamily HAUERININAE Schwager 1876
Quinqueloculina dOrbigny 1826
Type species: Serpula seminulum Linnaeus 1758.
Biloculina globulus Bornemann 1855, p. 349, pl. 19, fig. 3.
Upper depth limit: Quinqueloculina is generally considered as

having its upper depth limit in the inner neritic zone. Off Chile,
several species occur at neritic depths (Zapata and Cear 2004),
and the genus is not represented at bathyal depths along the PeruChile Trench (Ingle, Keller and Kolpack 1980).
Type age and locality: Oligocene, Germany.
Quinqueloculina akneriana dOrbigny 1846
Nummulopyrgo globulus (Bornemann 1855)
Plate 3, Figure 13
Occurrence: El Peral beds (NLP), Navidad Fm. (PPP).
Plate 3, Figure 16
Quinqueloculina akneriana dOrbigny 1846, p. 100, p. 290, pl. 18, figs.

1621. PAPP and SCHMID 1985, pl. 95, figs. 15.
387

Basin, Austria.
Remarks: This common form resembles what many workers have
identified as Quinqueloculina seminula (= Serpula seminulum
Linnaeus 1758), even though the type-figure of that species
(Plancus 1743, fig. 1) displays a more sinuous outline on one side
and a larger aperture.
MAT, MPUP, MOS, NAV5, PPP, PPT, PTA, RAP), Ranquil Fm.
(FRA, FRM, MIB, MS10, RAN, RQK, RQT), Santo Domingo
Fm. (VAL), Lacui Fm. (CHE, CHO, CUC, PNH).
Maximum relative abundance: Common (MOS, RAP, MS10).
Type age and locality: Recent, off southern California, 320m.

Distinguishing features: One edge broad and flat, the other
subacute.
Occurrence: Navidad Fm. (NAV5, PTA), Ranquil Fm. (FRA,
RQT).
Quinqueloculina boueana dOrbigny 1846
Plate 4, Figure 1
Quinqueloculina boueana DORBIGNY 1846, p. 293, pl. 19, figs. 79.

PAPP and SCHMID 1985, p. 101, pl. 96, figs. 8, 9. JONES
1994, p. 23, pl. 7, fig. 13.
Quinqueloculina badenensis dOrbigny 1846
Plate 3, Figure 17

Basin, Austria.
Quinqueloculina badenensis dOrbigny 1846, p. 299, pl. 20, figs.

1012. PAPP and SCHMID 1985, p. 105, pl. 101, figs. 610.
Distinguishing features: Distinctly costate; subacute edges,

slightly concave sides.

Basin, Austria.
Comparative species: Quinqueloculina bicostata dOrbigny
1839a (Recent, Cuba) has more-pronounced double ridges along
its periphery and its outline reflects the quinqueloculine pattern,
making it less ovate (see Li Qianyu 1977). Quinqueloculina
arctica Cushman 1933a (Recent, off Greenland) does not have
concave surfaces.
Occurrence: Navidad Fm. (NAV5, RAP), Lacui Fm. (CHO).
Maximum relative abundance: Common (RAP)
Quinqueloculina cf. Q. benwestonensis McCulloch 1977
Plate 3, Figure 18
Quinqueloculina benwestonensis McCulloch 1977, p. 482, pl. 211,

fig. 1.
Occurrence: Navidad Fm. (PPN, RAP).

Maximum relative abundance: Common (RAP).
Quinqueloculina cf. Q. flexuosa dOrbigny 1839c
Plate 4, Figure 2
Quinqueloculina flexuosa DORBIGNY 1839c, p. 73, pl. 4, figs. 46.

(Recent, Chile)
Remarks: This is a fairly robust form similar to Triloculina cf. T.

brochita (Pl. 5, Fig. 5), but the display of four chambers suggests
a quinqueloculine arrangement.
Comparative species: Quinqueloculina flexuosa has a sinuous
outline and fine costae.
Occurrence: Ranquil Fm. (MIB, RAN), Lacui Fm. (PCB).
PLATE 4
1 Quinqueloculina boueana dOrbigny, UCMP50056, RAP.
8 Quinqueloculina sp. C, UCMP50063, RAP.
2 Q
uinqueloculina cf. Q. flexuosa dOrbigny, UCMP50057,
RAN.
9 Biloculinella labiata (Schlumberger), UCMP50064, PPP.
3 Q
uinqueloculina magellanica dOrbigny, UCMP50058,
CUC.
uinqueloculina opulenta McCulloch, UCMP50059,
4 Q
NAV5.
5 Q
uinqueloculina suborbicularis dOrbigny, UCMP50060,
MIB.
6 Quinqueloculina sp. A, UCMP50061, MIB.
7 Quinqueloculina sp. B, UCMP50062, RAN.
388
10 C
ribromiliolinella subvalvularis (Parr), UCMP50065,
MIB.
11, 12 Miliolinella suborbicularis (dOrbigny): 11, UCMP50066,
RAP. 12, UCMP50067, CUC.
13 P
seudotriloculina cf. P. cyclostoma (Reuss), UCMP50068, MIB.
14 Pyrgo depressa (dOrbigny), UCMP50069, RAN.
15 Pyrgo lunula (dOrbigny), UCMP50070, RQT.
Kenneth L. Finger
Plate 4
389
Quinqueloculina magellanica dOrbigny 1839c
Quinqueloculina sp. C
Quinqueloculina magellanica DORBIGNY 1839c, p. 77, pl. 9, figs.

1921.
Occurrence: Navidad Fm. (RAP), Ranquil Fm. (RQT).
Plate 4, Figure 3
Type age and locality: Recent, Falkland Islands.

Distinguishing features: Outline slightly higher than wide;
apertural view has triangular outline with subrounded to
subacute edges.
Occurrence: Navidad Fm. (RAP), Lacui Fm. (CHO, CUC).
Maximum relative abundance: Common (CHO).
Quinqueloculina opulenta McCulloch 1977
Plate 4, fig. 8

Subfamily MILIOLINELLINAE Vella 1957
Biloculinella Wiesner 1931
Type species: Biloculina labiata Schlumberger 1891.
Biloculinella labiata (Schlumberger 1891)
Plate 4, Figure 9
Biloculina labiata Schlumberger 1891, p. 556, pl. 9, figs. 6062;

p. 556 tf. 13; pl. 557 tf. 14.
Plate 4, Figure 4
Type age and locality: Recent, Mediterranean Sea, 555m.
Quinqueloculina opulenta McCulloch 1977, p. 500, pl. 216, figs. 2,

3, 6.
Comparative species: Biloculina isabelleana dOrbigny 1839c

(Recent, Falkland Islands) has a rounded edge.
Type age and locality: Recent, Galpagos Islands, 29m.
Occurrence: Navidad Fm. (MOS, PPP).
Distinguishing feature: Flat peripheral edge.
Occurrence: NAV5.
CRIBROMILIOLINELLA Saidova 1981

Type species: Triloculina subvalvularis Parr 1950.

Quinqueloculina suborbicularis dOrbigny 1905
Plate 4, Figure 5
Quinqueloculina suborbicularis DORBIGNY 1826, p. 302, modle

no. 29 (nomen nudem).
Quinqueloculina suborbicularis dOrbigny in FORNASINI 1905, p.
67, pl. 4, fig. 3. HAYWARD, GRENFELL, SABAA, NEIL and
BUZAS 2010, p. 155, pl. 8, figs. 23 25.
Occurrence: Navidad Fm. (CPUP, RAP), Ranquil Fm. (MIB,

RAN).
Quinqueloculina sp. A
Plate 4, Figure 6
Distinguishing features: Nearly triloculine, rounded edges,

short flap-like tooth extends halfway into aperture.
Remarks: This form might be an aberration of Triloculinella
bornemanni (Pl. 5, Fig. 10).
Occurrence: Navidad Fm. (LBZ, MAT), Ranquil Fm. (LEB,
MIB).
Quinqueloculina sp. B
Plate 4, Figure 7
Comparative species: Quinqueloculina inaequalis dOrbigny

1839b (Recent, Canary Islands) has a sinuous profile in apertural
view.
Occurrence: Ranquil Fm. (FRA), Santo Domingo (VAL),
390
Cribromiliolinella subvalvularis (Parr 1950)
Plate 4, Figure 10
Triloculina subvalvularis PARR 1950, p. 296, pl. 7, fig. 4.

Cribromiliolinella subvalvularis (Parr). LOEBLICH and TAPPAN
1987, pl. 348, figs. 1618.
Type age and locality: Recent, off Antarctica, 193m.

Occurrences: Navidad Fm. (CPUP), Ranquil Fm. (MIB, RQT).
Miliolinella Wiesner 1931
Type species: Vermiculum subrotundum Montagu 1803.
Distinguishing features: Early stage quinqueloculine, later
planispiral with 47 chambers visible.
Miliolinella suborbicularis (dOrbigny 1839a)
Triloculina suborbicularis DORBIGNY 1839a, p. 177, type figure in

v. 8, pl. 10, figs. 911.
Type age and locality: Recent, Caribbean.

Distinguishing features: Compressed, rounded edge, fine costae.
Occurrence: Navidad Fm. (MOS, PPP, RAP), Lacui Fm. (CUC).
PSEUDOTRILOCULINA Cherif 1970
Type species: Triloculina lecalvezae Kaaschieter 1961.
Pseudotriloculina cf. P. cyclostoma Reuss 1850
Plate 4, Figure 13

Biloculina cyclostoma REUSS 1850, p. 382, pl. 49, fig. 6.
Pseudotriloculina cyclostoma (Reuss). LOEBLICH and TAPPAN
1987, p. 342, pl. 352, figs. 612.

Basin, Austria.
Distinguishing features: Only two chambers visible.
Remarks: Differs from P. cyclostoma by being less inflated.
Comparative species: Triloculina inornata dOrbigny 1846
(Middle Miocene, Vienna Basin) has a similar shape, but
displays a third chamber.
Occurrence: Navidad Fm. (MPUP), Ranquil Fm. (MIB).
Pyrgo Defrance, in de Blainville 1824
Type species: Pyrgo laevis Defrance, in de Blainville 1824.
Pyrgo clypeata (dOrbigny 1846)
Biloculina clypeata DOrbigny 1846, p. 266, pl. 15, figs. 1921.

Pyrgo clypeata (dOrbigny). PAPP and SCHMID 1985, p. 89, pl. 82,
figs. 46. HAYWARD, GRENFELL, SABAA, NEIL and BU
ZAS 2010, p. 150, pl. 7, figs. 710.

Basin, Austria.
Comparative species: DOrbigny named numerous species
of ovate Pyrgo that can difficult to consistently differentiate.
He described the Recent Biloculina oblonga (off Cuba), B.
patagonica (off Argentina), and B. peruviana (off Peru) in 1839,
and, from the Middle Miocene of Austria in 1846, B. affinis, B.
clypeata, B. inornata, and B. simplex. The primary differences
seen in his type figures are width, chamber inflation, angle and
orientation of the ultimate chambers inward ledge and the edgeview shape of its suture, but gradations among these features
blur the differentiation of species. On the basis of SEMs in Papp
and Schmid 1985, Pyrgo clypeata most closely resembles the
specimens from the Chilean Miocene.
Upper depth limit: Neritic (this species as P. ringens in Zapata
and Cear 2004).
PTA, RAP), Ranquil Fm. (FRA, FRM, LEB, MIB, RAN, RQK,
RQT), Santo Domingo Fm. (VAL), Lacui Fm. (PNH).
Distinguishing features: Circular outline in side view, subcircular

in edge view, blunt keel.
Comparative species: See remarks under Pyrgo murrhina below.
Pyrgo scutella (= Biloculina scutella Karrer 1868; Miocene,
Romania) has a wider and thinner keel and is more compressed.
Occurrence: El Peral beds (NLP), Navidad Fm. (CPUP, LBZ,
MPUP, MOS, NAV5, PPP, PTA, RAP), Ranquil Fm. (FRA,
FRM, MS10, RAN, RQT), Lacui Fm. (CHO, CUC, PCB).
Pyrgo lunula (dOrbigny 1846)
Plate 4, Figure 15
Biloculina lunula DORBIGNY 1846, p. 264, pl. 15, figs. 2224.

Biloculina scutella KARRER 1868, p. 134, pl. 1, fig. 7.
Pyrgo lunula (dOrbigny). PAPP and SCHMID 1985, p. 89, pl. 82, fig.
79.
Type age and locality: Miocene, Romania.

Comparative species: Pyrgo depressa (Pl. 4, Fig. 14) has a wider
aperture that is rimmed on all sides, similar to P. murrhina.
Pyrgo murrhina (Pl. 5, Fig. 4) has a more compressed test with a
subrounded to round aperture, and a more acute peripheral edge.
Occurrence: Navidad Fm. (CPUP, MPUP, PTA); Ranquil Fm.
(RQT).
Maximum relative abundance: Few
Pyrgo murrhina Schwager 1866
Plate 5, Figure 4
Pyrgo murrhina Schwager 1866, p. 203, pl. 4, fig. 15. KOHL

1985, p. 35, pl. 5, fig. 1. van Morkhoven, BERGGREN and
EDWARDS 1986, p. 50, pl. 15, figs. 1, 2. JONES 1994, p. 18, pl.
2, figs. 10, 11, 15. HAYWARD, GRENFELL, SABAA, NEIL
and BUZAS 2010, p. 151, pl. 7, figs. 20123.
Biloculina bradyi Fornasini 1886, p. 251, type-figure not given
(Pliocene, Italy)

Upper depth limit: Inner neritic, but most often middle bathyal
(van Morkhoven, Berggren and Edwards 1986).
Distinguishing features: Test diameter considerably greater than
thickness; relatively thin, moderately broad, carina, often with
aboral sinus; rimmed aperture ontogenetically transitions from
circular to ovate-quadrate.
Pyrgo depressa (dOrbigny 1826)
Occurrence: El Peral beds (LPER), Navidad Fm. (MOS, RAP),

Ranquil Fm. (FRA, MIB, RQK, RQT).
Biloculina depressa DORBIGNY 1826, p. 298, modle no. 91.

Pyrgo depressa (dOrbigny). JONES 1994, p. 19, pl. 2, figs. 12, 16, 17.
HAYWARD, GRENFELL, SABAA, NEIL and BUZAS 2010, p.
150, pl. 7, figs. 13, 14.
Type age and locality: Fossil and Recent, Italy.
Pyrgoella sphaera (dOrbigny 1839c)
Plate 4, Figure 14
Upper depth limit: Upper middle bathyal; based on occurrence

recorded off Chile by Bandy and Rodolfo (1964).
Pyrgoella Cushman and White 1936

Type species: Biloculina sphaera dOrbigny 1839c.
Plate 5, Figure 3
Biloculina sphaera DOrbigny 1839c, p. 66, pl. 8, figs. 1316.

Pyrgoella sphaera (dOrbigny). JONES 1994, p. 18, pl. 2, fig. 4.
391
152, pl. 8, figs. 911.

Pyrgoella(?) generalis McCulloch 1977, p. 522, pl. 242, figs. 918.
(Recent, Pacific off Mexico)
Type age and locality: Late Pleistocene, Ryuku Islands.
Type age and locality: Recent, Falkland Islands, depth not

recorded.
Upper depth limit: Bathyal; based on type-description.
Stratigraphic range: Early Miocene to late Pleistocene.
Upper depth limit: Outer neritic; based on records off New

Zealand (Hayward et al. 2010).
Remarks: The type description notes that although adult

specimens externally show two chambers, juveniles show a third
chamber. The Chilean forms show three chambers throughout
an ontogenetic series, although very little of the third chamber is
visible in adults.
Comparative species: Pyrgoella dokici (= Biloculina dokici

Pavlovi 898; Tertiary, Yugoslavia) is not as spherical and its
apertural end protrudes outward.
Comparative species: Its shape resembles that of Spiroloculina

robusta Brady 1884, but T. lucernuloides has a sinuous outline
and a different mode of coiling.
Occurrence: El Peral beds (NLP), Navidad Fm. (CPUP, MOS,

PPP), Ranquil Fm. (FRA, RQT), Santo Domingo Fm. (VAL).
Occurrence: Navidad Fm. (CPUP, MAT, MOS, NAV5, PPP,

PTA), Ranquil Fm. (MS10. RQT), Santo Domingo Fm. (VAL).
Triloculina dOrbigny 1826; emend. uczkowska 1972

Type species: Miliola (Miliolites) trigonula Lamarck 1804.
Triloculina oblonga (Montagu 1803)
Triloculina cf. T. brochita Carter 1964
Vermiculum oblongum Montagu 1803, p. 522, pl. 19, fig. 9.
Plate 5, Figure 5
Triloculina brochita CARTER 1964, p. 59, pl. 1, figs. 3, 4.
Type age and locality: Lower Miocene, Victoria, Australia.
Plate 5, Figure 6
Type age and locality: Recent, England, depth not recorded.

Occurrence: Lacui Fm. (CUC).
Comparative species: This form is similar to Triloculina oblonga

dOrbigny 1839a (Recent, Cuba) and T. chemnitziana dOrbigny
1839b (Recent, Canary Islands). Montagus type description and
figure are inadequate for comparison and dOrbigny does not
comment on Montagus species. Assuming they are synonymous,
Montagus nomen has seniority.
Occurrence: Navidad Fm. (NAV5), Ranquil Fm. (FRA).
Triloculina lucernuloides (Ujii 1990)
Remarks: In apertural view, Triloculina brochita has a

subtriangular outline. The apertural tooth characteristic of the
genus is not preserved.
Plate 5, Figure 7
Pyrgo lucernuloides Ujii 1990, p. 15, pl. 3 figs. 7, 8.
Triloculina striatotrigonula Parker and Jones 1941
Plate 5, Figure 8
PLATE 5
1, 2 Pyrgo clypeata (dOrbigny): 13, UCMP50071, MIB. 14,

UCMP50072, FRA.
3 Pyrgoella sphaera (dOrbigny), UCMP50073, FRA.
4 Pyrgo murrhina Schwager, UCMP50074, FRA.
5 Triloculina cf. T. brochita Carter, UCMP50075, CUC.
6 Triloculina oblonga (Montagu), UCMP50076, FRA.
7 Triloculina lucernuloides (Ujii), UCMP50077, PPP.
8 T
riloculina striatotrigonula Parker and Jones, orally distort
ed specimen, UCMP50078, FRA.
392
9 Triloculina trigonula (Lamarck), UCMP50079, CHO.

10 Triloculinella bornemanni (Bosquet), UCMP50080, RAP.
11 Triloculina sp., UCMP50081, RAP.
12 Triloculinella striata (Fischer), UCMP50082, NLP.
13 Triloculinella sp. A, UCMP50083, VAL.
14 Triloculinella sp. B, UCMP50084, VAL.
15 Sigmopyrgo vespertilio (Schlumberger), UCMP50085, PPP.
Kenneth L. Finger
Plate 5
393
Triloculina striatotrigonula PARKER and JONES, in Parr 1941, p.

305; type-figure is that of Miliolina insignis Brady in Brady 1884,
pl. 4, fig. 10 (non fig. 8).
Triloculina trigonula var. striatotrigonula Parr. JONES 1994, p. 21, pl.
4, fig. 10.
Type age and locality: Recent, Australia, 6973m.

Upper depth limit: Neritic.
Remarks: The single specimen recovered has poorly defined
striae and an obscured aperture.
Occurrence: El Peral beds (NLP), Ranquil Fm. (FRA).
Triloculina trigonula (Lamarck 1804)
Plate 5, Figure 9
Miliolites trigonula Lamarck 1804, p. 351; type-figure in LA

MARCK 1807, pl. 17, fig. 4.
Triloculina trigonula (Lamarck). LOEBLICH and TAPPAN 1987, pl.
351, figs. 19-21. JONES 1994, p. 20, pl. 3, figs. 15, 16.
Type age and locality: Eocene, France.

Stratigraphic range: Eocene to Recent.
Occurrence: Navidad Fm. (CPUP, PPP, RAP), Ranquil Fm.
(FRA, RAN, RQT), Lacui Fm. (CHO).
Triloculina sp.
Plate 5, Figure 11
Remarks: In apertural view, it appears subrounded-subtriangular,
as all three sides are convex and the corners are well rounded.
Occurrence: (RAP)
Triloculinella Riccio 1950
Type species: Triloculinella obliquinodus Riccio 1950.

Triloculinella striata (Fischer 1927)
Plate 5, Figure 12
Miliolina valvularis var. striata Fischer 1927, p. , pl. 217, fig. 128.
Type age and locality: Pliocene, Moluka Islands (Moluccas),

Indonesia.
Triloculinella sp. A
Plate 5, Figure 13
Remarks: Juveniles display the broad apertural flap.

Comparative species: Similar in subglobose form to Triloculina
flavescens dOrbigny (in Fornasini 1905; Recent, France), which
has the apertural tooth characteristic of that genus.
Occurrence: El Peral beds (NLP, LPER), Navidad Fm. (PPP),
Ranquil Fm. (FRA), Santo Domingo Fm. (VAL), Lacui Fm.
(PNH).
Triloculinella sp. B
Plate 5, Figure 14
Distinguishing features: Test very inflated, broadly ovate in

side view, width exceeding height, broadly subovate and widest
across ultimate chamber in apertural view; 3 chambers visible;
aperture broadly U-shaped, wider than high.
Remarks: None of the specimens preserved the apertural flap,
but the very robust shape is more typical of Triloculinella than
Triloculina.
Occurrence: Navidad Fm. (RAP), Ranquil Fm. (FRA, RAN,
RQT), Santo Domingo Fm. (VAL),
Remarks: Triloculinella is differentiated from Miliolinella by its

more inflated test, a cryptoquinqueloculine to quinqueloculine
arrangement, and final three to five chambers externally visible.
It differs from Triloculina by having a more rounded outline in
latitudinal section and a more rounded aperture with a flap-like
tooth.
Subfamily SIGMOILINITINAE uczkowska 1974

Nummoloculina Steinmann 1881
Type species: Biloculina contraria dOrbigny 1846.
Triloculinella bornemanni (Bosquet 1859)
Biloculina contraria dOrbigny 1846, p. 266, pl. 16, figs. 46.

Nummuloloculina contraria (dOrbigny). PAPP and SCHMID 1985, p.
90, pl. 83, figs. 79. LOEBLICH and TAPPAN 1987, pl. 355, figs.
1723. HAYWARD, GRENFELL, SABAA, NEIL and BUZAS
2010, p. 150, pl. 7, figs. 46. HOLBOURN, HENDERSON and
MACLEOD 2013, p. 376377.
Plate 5, Figure 10
Triloculina bornemanni Bosquet 1859, p. 25, pl. 2, fig. 12.
Type age and locality: Tertiary, The Netherlands.

Distinguishing features: Broadly ovate in side view, subtriangular
with rounded edges in apertural view; 3 chambers visible;
aperture V-shaped with rounded angle and similarly shaped flap.
Comparative species: Triloculinella selene (= Triloculina selene
Karrer 1868; Miocene, Mediterranean) is probably synonymous.
Occurrence: El Peral beds (NLP), Navidad Fm. (RAP), Santo
Domingo Fm. (VAL).
394
Nummoloculina contraria (dOrbigny 1846)

Plate 6, Figure 3

Basin, Austria.
Distinguishing features: Three to four chambers visible in the
outer whorl.

Sigmoilinita Seiglie 1965
Type species: Quinqueloculina tenuis Cjek 1848.
Sigmoilinita tenuis (Cjek 1848)
Plate 6, Figure 1
Quinqueloculina tenuis Cjek 1848, p. 149, pl. 13, figs. 3134.

Sigmoilina miocenica Cushman 1946 (Miocene, Florida).
Spirosigmoilina tenuis (Cjek). KOHL 1985, p. 36, pl. 5, fig. 5.
LOEBLICH and TAPPAN 1987, pl. 356, figs. 17, 18. FINGER
1992, p. 68, pl. 1, fig. 30. JONES 1994, p. 26, pl. 10, figs. 7, 8, 11.
157, pl. 9, figs. 21, 22.
Type age and locality: Tertiary, Austria.

Upper depth limit: Outer neritic; based on its placement in
California by Ingle (1980).
Remarks: The very flat, fusiform shape of Cjeks species is
uncharacteristic of Spirosigmoilina Parr 1942, to which it often
has been assigned.
Comparative species: Sigmoilinita concinna (= Quinqueloculina
concinna Reuss 1850; Tertiary, Austria) shows a great difference
in the number of chambers between opposite sides, whereas
Sigmoilinita tenuis does not.
Occurrence: Navidad Fm. (CPUP, MOS, PPP, PTA), Ranquil
Fm. (RQK, RQT), Lacui Fm. (CHO).
Sigmopyrgo Hofker 1983
Type species: Biloculina vespertilio Schlumberger 1891.
Sigmopyrgo vespertilio (Schlumberger 1891)
Sigmoilina schlumbergeri A. Silvestri 1904a, p. 267, 269; type-fig

ures in Schlumberger 1887, pl. 7, figs. 1214; p. 481, tf. 6; p.
482, tf. 7.
Sigmoilopsis schlumbergi (A. Silvestri). KOHL 1985, p. 36, pl. 5, fig.
6. LOEBLICH and TAPPAN 1987, pl. 356, figs. 8-13. JONES
1994, p. 23, pl. 8, figs. 14. HAYWARD, GRENFELL, SABAA,
NEIL and BUZAS 2010, p. 157, pl. 9, figs. 15, 16.
Type age and locality: Recent, Gulf of Gascogne (Bay of Biscay;

NE Atlantic), France, 6001200m.
Upper depth limit: Neritic, but in greatest abundance at bathyal
depths (van Morkhoven, Berggren and Edwards 1986; Hayward
et al. 2010).
Order LAGENIDA Lankester 1885
Superfamily NODOSARIOIDEA Ehrenberg 1838
Family CHRYSALOGONIIDAE Mikhalevich 1993
Anastomosa Hayward, in Hayward et al. 2012
Type species: Nodosaria gomphiformis Schwager 1866.
Anastomosa brevilocula (Cushman and Jarvis 1934)
Plate 6, Figure 4
Chrysalogonium brevilocolum CUSHMAN and JARVIS 1934, p. 74,

pl. 10, fig. 13.
Anastomosa brevilocula (Cushman and Jarvis). HAYWARD,
Kawagata, Sabaa, Grenfell, Van Kerckhoven,
Johnson and Thomas 2012, p. 115, pl. 2, figs. 2025.
Type age and locality: Early Miocene, Trinidad.

Stratigraphic range: Late Eocene to Early Miocene.
Plate 5, Figure 13
Upper depth limit: Unknown.
Biloculina vespertilio Schlumberger 1891, p. 561, pl. 10, figs.

7476; p. 561, tfs. 2021; p. 562, tf. 22.
Pyrgo vespertilio (Schlumberger). BOLTOVSKOY and THEYER
1970, p. 353, pl. 4, fig. 9.
Sigmopyrgo vespertilio (Schlumberger). LOEBLICH and TAPPAN
1987, pl. 257, figs. 1418. JONES 1994, p. 18, pl. 2, fig. 8.
Occurrence: El Peral beds (NLP)
Type age and locality: Recent, Golfe de Gascogne (Bay of

Biscay; NE Atlantic), 1850m.
Comparative species: In edge view, Sigmopyrgo calostoma
(= Biloculina bulloides var. calostoma Karrer 1868; Miocene,
Romania) shows a sinuous, not planar, contact between the two
visible chambers.
Subfamily SIGMOILOPSINAE Vella 1957
Sigmoilopsis Finlay 1947
Type species: Sigmoilina schlumbergeri A. Silvestri 1904a.
Sigmoilopsis schlumbergeri (A. Silvestri 1904a)
Plate 6, Figure 2

Anastomosa lamellata (Cushman and Stainforth 1945)
Nodosaria lamellata CUSHMAN and STAINFORTH 1945, p. 24,

pl. 3, figs. 23, 24 (nom. subst. pro Dentalina carinata Neugeboren
1856). ROBERTSON 1998, p. 44, pl. 15, fig. 4.
Anastomosa lamellata (Cushman and Stainforth). HAYWARD,
Type age and locality: Early to Middle Miocene, Venezuela.

Stratigraphic range: Late Cretaceous to early Pleistocene.
Upper depth limit: Lower middle bathyal (1100m).
Occurrence: Navidad Fm. (PPP)
Chrysalogonium Schubert 1908
Type species: Nodosaria polystoma Schwager 1866.
395
Remarks: The genus is distinguished by having a finely cribrate

aperture without a toothplate. None of the specimens assigned
here to Chrysalogonium have a preserved aperture; hence,
the generic assignment is based solely on chamber shape and
ornamentation, which, together, appear unique to the species
indicated.
Occurrence: Navidad (MPUP), Ranquil Fm. (FRM), Lacui Fm.

(CUC).
Chrysalogonium deceptorium (Schwager 1866)
Plate 6, Figure 12
Nodosaria deceptorium SCHWAGER 1866, p. 231, 5, pl. 6, fig. 66.

Chrysalogonium deceptorium (Schwager). HAYWARD, Kawaga
ta, Sabaa, Grenfell, Van Kerckhoven, Johnson and
Thomas 2012, p. 118, pl. 4, figs. 916.

Stratigraphic range: Late Cretaceous to Early Pliocene.
Upper depth limit: Upper bathyal, 500m (Hayward et al. 2012).
Comparative species: Dentalina barnesi Rankin (in Cushman

and Kleinpell 1934; Miocene, California) has about half as many
costae.
Chrysalogonium equisetiformis (Schwager 1866)
Plate 6, Figure 13
Nodosaria equisetiformis Schwager 1866, p. 231, pl. 6, fig. 66.

Chrysalogonium equisetiformis (Schwager). HAYWARD, Kawaga
Thomas 2012, p. 119, pl. 4, figs. 1722.
Nodosaria spirostriolata Cushman 1917b, p. 656, pl. 38, fig. 4.
(Recent, Philippines, 903m)

Stratigraphic range: Late Cretaceous to early Pleistocene.
PLATE 6
Figures 2a, 3a,b, 46, 15, 18, 19, 24b, 34, and 37 are photomicrographs;
all other images are SEMs. Scale bars in m.
1 Sigmoilinita tenuis (Cjek), UCMP50086, RQT.

2 S igmoilopsis schlumbergeri (A. Silvestri), UCMP50087,
NLP.
3 Nummoloculina contraria (dOrbigny), UCMP50088, PPP.
4 A
nastomosa brevilocula (Cushman and Jarvis), UCMP
50089, NLP.
5, 6 A
nastomosa lamellata (Cushman and Stainforth), PPP:
5. megalospheric, UCMP50090. 6, microspheric, UCMP
50091.
7, 8 L
otostomoides pyrulus (dOrbigny): 7, UCMP50092, PTA.
8, UCMP50093, PPP.
20 Dentalina mutsui Hada, UCMP50105, FRA.

21 Dentalina obliquecostata (Stache), UCMP50106, RAP.
22, 23 D
entalina striatissima Stache, FRM: 22, UCMP50107.
23, UCMP50108.
24 Enantiodentalina muraii Uchio, UCMP50109, PPP.
25 Fingerina weaveri (Finger and Lipps), UCMP50110, PPP.
26, 27 P
yramidulina acuminata (Hantken): 26, microspher
ic form, UCMP50111, FRM. 27, megalospheric form,
UCMP50112, MIB.
28 Dentalina flintii (Cushman), UCMP50113, PPP.
9 Neugeborina longiscata (dOrbigny), UCMP50094, FRA.
29 Laevidentalina cf. L. planata (Blake), UCMP50114, PPN.
10, 11 L otostomoides asperulum (Neugeboren): 10, UCMP

50095, PTA. 11, UCMP50096, NAV5.
30 L aevidentalina roemeri (Neugeboren), UCMP50115, MIB.
12 C
hrysalogonium deceptorium (Schwager), UCMP50097,
FRM.
13 C
hrysalogonium equisetiformis (Schwager), UCMP50098,
RAN.
14 C
hrysalogonium rudis (dOrbigny), UCMP50099, FRA.
15 O
rthomorphina jedlitschkai (Thalmann), UCMP50100,
PPP.
16 Orthomorphina perversa (Schwager), UCMP50101, PTA.
17, 18 D
entalina aciculata dOrbigny, PPP: 17, UCMP50102. 18,
UCMP50103.
19 Dentalina albatrossi (Cushman), UCMP50104, MPUP.
396
31 Laevidentalina sp. A, UCMP50116, PPP.

32 Laevidentalina sp. B, UCMP50117, FRM.
33 Laevidentalina advena (Cushman), UCMP50118, PPP.
34, 35 L aevidentalina communis (dOrbigny): 34, UCMP50119,
PPP. 35, UCMP50120, PTA.
36, 37 L
aevidentalina elegans (dOrbigny): 36, UCMP50121,
FRA. 37, UCMP50122, PPP.
38 Laevidentalina inflexa (Reuss), UCMP50123, PPP.
39 L aevidentalina inornata (dOrbigny), UCMP50124, FRA.
Kenneth L. Finger
Plate 6
397

Distinguishing features: Approximately 30 striae continue
across sutures of barrel-shaped chambers.
Remarks: Generic placement is based on Hayward et al. (2012).
Comparative species: Dentalina obliqua (= Nautilus obliquus
Linnaeus 1758; Recent, Adriatic Sea) has oblique sutures and
fewer striae that do not continue across sutures. Dentalina
strigosa (= Nodosaria (Dentalina) strigosa Costa 1856; fossil,
Italy) has a less lobulate profile. Dentalina striatissima Stache
1865 (Late Tertiary, New Zealand) has about half as many
striae. Dentalina subbullata (= Orthocerina subbullata Costa,
in Fornasini 1894; Tertiary, Sicily) is similar.
Occurrence: Ranquil Fm. (FRA, RAN).
Chrysalogonium rudis (dOrbigny 1846)
Plate 6, Figure 14
Nodosaria rudis dOrbigny 1846, p. 33, pl. 1, figs. 1719. PAPP

and SCHMID 1985, p. 24, pl. 4, fig. 5.
Siphonodosaria setosa SCHWAGER 1866, p. 219, pl. 5, fig. 40. (Plio
cene, Car Nicobar)
Chrysalogonium rudis (dOrbigny). HAYWARD, Kawagata,
Sabaa, Grenfell, Van Kerckhoven, Johnson and
Thomas 2012, p. 121, pl. 5, figs. 613.
Type age and locality: Middle Miocene, Austria.

Upper depth limit: Lower middle bathyal, 1100m (Hayward et
al. 2012).
Occurrence: Ranquil Fm. (FRA, FRM, MIB, RAN, MIB),
Lacui Fm. (PCB).
Lotostomoides Hayward and Kawagata, in Hayward et al. 2012
Type species: Nodosaria asperula Neugeboren 1852.
Distinguishing features: Aperture coarsely reticulate, slightly
domed to conical mesh, usually on a neck and often divided into
pillars or tubes.
Lotostomoides asperulum (Neugeboren 1852)
Nodosaria asperula NEUGEBOREN 1852, p. 54, , pl. 1, figs. 40, 41.

Nodosaria holoserica Schwager 1866, p. 398, text-fig. 9. (Pliocene,
Car Nicobar)
Nodosaria aspera O. Silvestri 1872, p. 76, pl. 8, figs. 191200.
(Pliocene, Italy)
Lotostomoides asperulum (Neugeboren). HAYWARD, Kawagata,
Thomas 2012, p. 125, pl. 6, figs. 2023.
Type age and locality: Tertiary, Romania.

Stratigraphic range: Early Miocene to Pliocene.
Upper depth limit: Upper middle bathyal, 900m (Hayward et
al. 2012).
Distinguishing features: Long neck, spherical chambers, without
apical spine. In addition to being hispid, some of the Chilean
specimens have very faint longitudinal striae.
398
Comparative species: Nodosaria insecta Schwager 1866

(Pliocene, Car Nicobar) has a radiate aperture and more ovate
chambers. Single-chamber segments could be mistakenly
identified as N. fichteliana Neugeboren 1852 (Tertiary,
Romania). Chrysalogonium rudis (= Nodosaria rudis dOrbigny
1846; Middle Miocene, Austria) is also densely hispid, but its
chambers are connected by long narrow necks and its aperture
is cribrate.
Occurrence: El Peral beds (NLP), Navidad Fm. (MPUP, MOS,
NAV5, PPP, PPT, PTA), Ranquil Fm. (MIB, MS10, RAN, RQT).
Lotostomoides pyrulus (dOrbigny 1826)
Nodosaria pyrula DORBIGNY 1826, p. 255; first described and

figured in Parker, Jones and Brady 1871, p. 253, no. 6.
KOHL 1985, p. 43, pl. 6, fig. 4. PAPP and SCHMID, p. 1985, pl.
4, figs. 8, 10.
Nodosaria? pyrula dOrbigny. ROBERTSON 1998, p. 46, pl. 15, fig. 8.
Grigelis pyrulus (dOrbigny). LOEBLICH and TAPPAN 1987, pl. 441, figs. 4, 5.
Type age and locality: Fossil, Italy.

Remarks: Most of the Chilean forms have subspherical-lacriform
chambers, but can be widest above or below the middle.
Comparative species: Lotostomoides lageniferus (= Nodosaria
lagenifera Neugeboren 1852; Tertiary, Romania) has more
slender chambers with shorter interchamber sections, but
those might be intraspecific variations. Dentalina guttifera
dOrbigny 1846; Middle Miocene, Austria) lacks the rod-like
extensions between most chambers. Lotostomoides orectus (=
Grigelis orectus Loeblich and Tappan 1994; Timor Sea, 263m) is
distinguished by its relatively long tubular necks between slender
pyriform chambers.
Occurrence: Navidad Fm. (MAT, MPUP, MOS, NAV5, PPP,
PPT, PTA), Ranquil Fm. (FRA, FRM, MIB, RAN, RQK, RQT)
Family GLANDULONODOSARIIDAE A. Silvestri 1901.
Neugeborina Podescu, in Cicha, Rgl, Rupp and Ctyroka
1998
Neugeborina longiscata (dOrbigny 1846)
Plate 6, Figure 9
Nodosaria longiscata DORBIGNY 1846, p. 32, pl. 1, figs. 1012.

KOHL 1985, p. 43, pl. 6, fig. 3. PAPP and SCHMID 1985, p. 23, pl.
3, figs. 15.
Ellipsonodosaria longiscata (dOrbigny). CUSHMAN and CEDER
STROM 1945, p. 32, pl. 4, figs. 1921.
Ellipsonodosaria cf. E. longiscata (dOrbigny). CUSHMAN 1948, p. 239,
pl. 19, figs. 12, 13.
Nodosaria? longiscata dOrbigny. ROBERTSON 1998, p. 44, pl. 15, fig. 5.
Neugeborina longiscata (dOrbigny). KAWAGATA, HAYWARD and
GUPTA 2006, p. 237, pl. fig. X. HAYWARD, Kawagata, Sa
baa, Grenfell, Van Kerckhoven, Johnson and Thomas
2012, p. 134, pl. 8, figs. 1923.

Basin, Austria.
Stratigraphic range: Late Cretaceous to middle Pleistocene.
Remarks: Some of the Chilean specimens display very faint

striae.
Type species: Nodosaria (les Dentalines) cuvieri dOrbigny

1826.
Comparative species: Similar morphotypes are Neugeborina

ewaldi (= Nodosaria ewaldi Reuss 1851b; Eocene, Germany)
and Neugeborina arundinea (Nodosaria arundinea Schwager
1866; Pliocene, Car Nicobar). In addition, Neugeboren (1852)
described several variants of this form from the Tertiary
of Romania, including Nodosaria ackneriana, N. exilis, N.
gracillis, and N. roemeriana. Slightly less elongate forms, such
as N. farcimen A. Silvestri 1872 and N. elongata dOrbigny 1902
(in Fornasini 1902; nom. nov. pro N. ovicula dOrbigny 1826),
may also be variants of Neugeborina longiscata.
Distinguishing features: Elongate arcuate test, radiate aperture,

longitudinal costae.

MPUP, MOS, NAV5, PPP, PPT, PTA), Ranquil Fm. (FRA, FRM,
MIB, MS10, RAN, RQK, RQT), Santo Domingo Fm. (VAL),
Lacui Fm. (PCB, PNH).
Maximum relative abundance: Common (MIB).
ORTHOMORPHINA Stainforth 1952
Type species: Nodogenerina havanensis Cushman and Bermdez 1937.
Orthomorphina jedlitschkai (Thalmann 1937)
Plate 6, Figure 15
Nodogenerina jedlitschkai Thalmann 1937, p. 341; type-figure in

BRADY 1884, pl. 62, figs. 1, 2.
Orthomorphina jedlitschkai (Thalmann). JONES 1994, p. 73, pl. 62,
figs. 1, 2. JONES 1994, p. 73, pl. 62, figs. 1, 2; suppl. pl. 2, figs.
6, 7. HAYWARD, Kawagata, Sabaa, Grenfell, Van
Kerckhoven, Johnson and Thomas 2012, p. 136, pl. 8,
figs. 2427.
Type age and locality: Recent, SE Pacific, 773m.

Upper depth limit: Upper middle bathyal in the Southwest Pacific
(based on type locality and Hayward 2002).
Occurrence: Navidad Fm. (PPP), Ranquil Fm. (MS10), Lacui
Fm. (PCB).
Dentalina aciculata (dOrbigny 1826)
Nodosaria (Dentaline) aciculata DOrbigny 1826, p. 255; first

described and figured in Parker, Jones and Brady 1871, p.
160, pl. 9, fig. 52.
Type age and locality: Recent, Adriatic Sea, depth not recorded.
Distinguishing features: Large, tapering test with ovate to
spherical chambers, surface smooth or finely hispid with short
intercameral costae; spherical chambers ornamented with fine
short spinules.
Comparative species: The overall shape of Nodosaria ambigua
Costa 1856 (age not given but probably Pleistocene, Italy)
suggests it is synonymous with Dentalina aciculata, but its type
figure shows none of the details needed to confirm this.
Occurrence: Navidad Fm. (CPUP, MAT, MPUP. PPP, PTA),
Santo Domingo Fm. (VAL), Lacui Fm. (PCB).
Dentalina albatrossi (Cushman 1923)
Plate 6, Figure 19
Nodosaria vertebralis var. albatrossi Cushman 1923, p. 312, pl. 57,

fig. 5.
Dentalina albatrossi (Cushman). JONES 1994, p. 76, pl. 63, fig. 35; pl.
64, figs. 1114.

Occurrence: Navidad Fm. (MPUP).
Orthomorphina perversa (Schwager 1866)
Dentalina flintii (Cushman 1923)
Nodosaria perversa Schwager 1866, p. 212, pl. 5, fig. 29.

Orthomorphina jedlitschkai (Thalmann). JONES 1994, p. 73, pl. 62,
figs. 1, 2. HAYWARD, Kawagata, Sabaa, Grenfell,
Van Kerckhoven, Johnson and Thomas 2012, p. 137, pl.
8, figs. 3538; pl. 9, figs. 14.
Nodosaria flintii Cushman 1923, p. 85, pl. 14, fig. 23.

Dentalina flintii (Cushman). JONES 1994, p. 76, pl. 64, figs. 2022.
Plate 6, Figure 16

Upper depth limit: Upper bathyal, 500m (Hayward et al. 2012)
Occurrence: Navidad Fm. (PTA), Lacui Fm. (PCB).
Family NODOSARIIDAE Ehrenberg 1838
Subfamily NODOSARIINAE Ehrenberg 1838
Dentalina Risso 1826
Plate 6, Figure 28
Type age and locality: Recent, off NE United States, 1811m.

Upper depth limit: Lower middle bathyal based on type
occurrence; middle bathyal in Southwest Pacific (Hayward 2002).
Remarks: Unlike the holotype, most of the Chilean specimens
have costae that do not extend across the ultimate chamber.
Comparative species: Dentalina albatrossi (= Nodosaria verte
bralis var. albatrossi Cushman 1923; Recent, Gulf of Mexico)
has a profile that is barely lobulate. Dentalina mutabilis (=
Nodosaria mutabilis Costa 1855; Tertiary, Italy), a junior
subjective synonym of D. mutabilis Bailey 1851, has oblique
sutures and more robust costae.
399
Occurrence: Navidad Fm. (PPP), Ranquil Fm. (MIB, RAN,

RQT).
Dentalina mutsui Hada 1931
Plate 6, Figure 20
Dentalina mutsui Hada 1931, p. 97, pl. 97, text-fig. 50.
Type age and locality: Recent, Japan, 2746m.

Occurrence: El Peral beds (NLP), Navidad Fm. (all except RAP

and PPN), Ranquil Fm. (FRA, FRM, LEB, RAN, RQT), Santo
Domingo Fm. (VAL), Lacui Fm. (CHE, PCB, PNH).
Enantiodentalina Marie 1941
Type species: Enantiodentalina communis Marie 1941.
Distinguishing features: Early stage biserial; surface smooth.
Enantiodentalina muraii Uchio 1953
Plate 6, Figure 24
Upper depth limit: Inner neritic, based on type occurrence.
Enantiodentalina muraii UCHIO 1953, p. 152, pl. 14, fig. 2.
Comparative species: This species is very similar to Dentalina

semicostata (= Nodosaria (Dentalina) pungens var. semicostata
Reuss 1870; Late Oligocene, Germany), the type figure of
which shows the chambers comprising the latter half of the
test as smooth and nearly rectilinear. Dentalina raristriata
(= Nodosaria raristriata Chapman 1893; Lower Cretaceous,
England) is a similar morphotype.
Distinguishing features: Early chambers of all specimens

are biserial and slightly compressed, some tests are apiculate;
sutures oblique, depressed.
Occurrence: El Peral beds (LPER, NLP), Navidad Fm. (CPUP),

Ranquil Fm. (RQT).
Dentalina obliquecostata (Stache 1865)
Plate 6, Figure 21
Nodosaria obliquecostata Stache 1865, p. 197, pl. 22, fig. 24.
Type age and locality: Late Tertiary, New Zealand.

Stratigraphic range: Miocene.
Remarks: The holotype is a four-chambered segment; Hornibrook
(1971) illustrates three topotype partial tests that he assigns to this
species. The Chilean form matches his plate 7, figure 11, but not
the others, which may not be the same species.
Comparative species: Dentalina pseudoinvolvens Cushman
and McGlamery 1939 (Early Oligocene, Alabama) has longer
chambers and more depressed sutures.
Occurrence: Navidad Fm. (PPN, PPP, RAP), Ranquil Fm.
(RAN), Lacui Fm. (PNH).
Maximum relative abundance: Common (RAP).
Dentalina striatissima Stache 1865
Dentalina striatissima Stache 1865, p. 208, pl. 22, fig. 38.

Stratigraphic range: Miocene to Pliocene(?).
Comparative species: Dentalina sulcata Nilsson 1826 (Cre
taceous, Sweden) is similar except its costae extend to the
aperture. Dentalina pseudoinvolvens Cushman and McGlamery
1939 (Early Oligocene, Alabama) has a less lobulate profile with
finer and more numerous oblique costae.
Occurrence: Navidad Fm. (CPUP, PPP, PTA), Ranquil Fm.

(RQT), Lacui Fm. (CHO).
Fingerina Hayward, in Hayward et al. 2012
Type species: Nodosaria weaveri Finger and Lipps, in Finger et
al. 1990.
Distinguishing feature: Aperture comprised of radiate bars
separated by triangular gaps, forming a distinct terminal
protrusion.
Fingerina weaveri Finger and Lipps 1990
Plate 6, Figure 25
Nodosaria weaveri Finger and Lipps, in Finger, Weaver, Lipps

and Miller 1990, p. 28, pl. 1, figs. 19, 20. HAYWARD,
Type age and locality: Early Miocene, California.

Stratigraphic range: Early Miocene to early Pleistocene.
al. 2012).
Comparative species: Orthomorphina ambigua (= Nodosaria
ambigua Neugeboren 1856; Neogene, Romania) has a round
aperture with everted lip, more constricted sutures, and no
apical spine (see Hayward 2002). Stilostomella consobrina (=
Nodosaria consobrina dOrbigny 1846; Middle Miocene, Austria)
has chambers that are not compressed, as well as an aperture
characteristic of its genus.
Occurrence: Navidad Fm. (CPUP, MOS, PPP, PPT, PTA).
Laevidentalina Loeblich and Tappan 1986b
Type species: Laevidentalina aphelis Loeblich and Tappan 1986b.
Distinguishing features: Elongate arcuate test with radiate
aperture and longitudinal costae.
Laevidentalina advena (Cushman 1923)
Plate 6, Figure 33
Nodosaria advena Cushman 1923, p. 79, pl. 14, fig. 12.
400

Dentalina advena (Cushman). JONES 1994, p. 74, pl. 63, fig. 1.
Laevidentalina subemaciata (Parr). ZAPATA and CEAR 2004, p. 28,
pl. 9, fig. 8,
Type age and locality: Recent, off NE United States, 960m.

Occurrence: Navidad Fm. (MPUP, NAV5, PPP, PPT, RAP),
Ranquil Fm. FRM).
Laevidentalina communis (dOrbigny 1826)
Occurrence: Navidad Fm. (PPP, PPT), Ranquil Fm. (FRM,

MIB, MS10).
Laevidentalina inornata (dOrbigny 1846)
Plate 6, Figure 39
Nodosaria inornata DORBIGNY 1846, p. 28, p. 2, pl. 1, fig. 5.

PAPP and SCHMID 1985, pl. 9, figs. 58. HAYWARD,
GRENFELL, SABAA, NEIL and BUZAS 2010, p. 171, pl. 12, figs.
48520.

Basin, Austria.
Nodosaria (Dentaline) communis DORBIGNY 1826, p. 254; no

type-figure or description given, but earliest are in dOrbigny 1840, p.
13, pl. 1, fig. 4.
Dentalina communis dOrbigny. FINGER 1992, p. 68, pl. 2, fig. 9.
Upper depth limit: Upper middle bathyal; based on California

occurrence (Ingle 1980).
Type age and locality: Recent, Adriatic Sea.

Comparative species: Laevidentalina filiformis (= Dentalina
filiformis Terquem 1878; Late Pliocene, Greece) has barrelshaped chambers. Laevidentalina inflexa (= Nodosaria
(Dentalina) inflexa Reuss 1866; middle Tertiary, Germany) has
greater sutural constriction.
Occurrence: Navidad Fm. (CPUP, PPP, PTA, RAP), Ranquil
Fm. (MIB, MS10), Lacui Fm. (CHE)
Laevidentalina elegans (dOrbigny 1846)
Dentalina elegans DORBIGNY 1846, p. 28, , p. 45, pl. 1, figs. 5256.

PAPP and SCHMID 1985, p. 28, pl. 10, figs. 18.

Basin, Austria.
Upper depth limit: Outer neritic; based on 122155m range
reported by Parr (1950).
Comparative species: Similar forms have often been referred

to Dentalina communis (= Nodosaria (Dentaline) communis
dOrbigny 1826; Recent, Adriatic Sea) for which subsequent
figures in dOrbigny (1840) and Parker, Jones and Brady (1871)
look quite dissimilar, the former having very oblique chambers
twice as broad as high, whereas those in the latter have more
rectilinear chambers that are twice as high as broad. Dentalina
gomphoides (= Nodosaria (Dentalina) gomphoides Costa 1856;
fossil, Italy) differs by the chamber width increasing more
rapidly and a more lobulate outline.
NAV5, PPP, PPT, PTA), Ranquil Fm. (FRA, FRM, MIB, MS10,
RAN), Santo Domingo Fm. (VAL).
Laevidentalina cf. L. planata (Blake 1876)
Plate 6, Figure 29
Dentalina planata Blake 1876, p. 458, pl. 18, fig. 22.

Type age and locality: Lower Jurassic, England.
Remarks: Although the single Chilean specimen is remarkably
similar to holotype of Laevidentalina planata, the wide
difference in age suggests that they are not the same species.
Occurrence: Navidad Fm. (PPN).
Occurrence: El Peral beds (NLP), Navidad Fm. (CPUP, MAT,

MOS, MPUP, NAV5, PPN, PPP, PPT, PTA), Ranquil Fm. (MS10,
RAN), Santo Domingo Fm. (VAL), Lacui Fm. (CHO, CUC,
PCB, PNH).
Dentalina roemeri Neugeboren 1856, p. 82, pl. 2, figs. 1317.

FINGER 1992, p. 69, pl. 2, figs 1922.
Laevidentalina inflexa (Reuss 1866)
Laevidentalina roemeri (Neugeboren 1856)
Plate 6, Figure 30
Plate 6, Figure 38
Type age and locality: Miocene to Pliocene, Romania.
Nodosaria (Dentalina) inflexa Reuss 1866, p. 131, pl. 2, fig. 1.

Dentalina inflexa Reuss. JONES 1994, p. 73, pl. 62, fig. 9.
Type age and locality: Middle Oligocene, Germany.

Stratigraphic range: Middle Oligocene to Recent.
Distinguishing feature: Asymmetrically ovate chambers.
Comparative species: Laevidentalina obliqua (= Nodosaria

(Dentaline) obliqua dOrbigny 1826; Recent, Adriatic Sea)
may be a senior synonym, but its type figure (Parker, Brady
and Jones 1871) is a simple sketch of a more curved specimen
with highly oblique chambers half as high as broad and a
lobulate periphery. Laevidentalina neugeboreni (= Nodosaria
401
neugeboreni Schwager 1866; Pliocene, Car Nicobar) is another

possible synonym.
Nodosaria acuminata VON Hantken 1876, p. 28, pl. 2, fig. 9; pl. 13,
fig. 5.
Occurrence: Ranquil Fm. (MIB), Lacui Fm. (CHO).
Type age and locality: Early Oligocene, Hungary.
Stratigraphic range: Early Oligocene to Recent.
Laevidentalina sp. A
Upper depth limit: Lower bathyal in the Southwest Pacific

(Hayward 2002).
Distinguishing features: Large, robust, nearly straight test with

slightly oblique sutures and nearly equidimensional chambers.
Remarks: This species resembles Nodosaria affinis dOrbigny

1846 and N. bacillum dOrbigny 1846 (both Middle Miocene,
Austria), which Papp and Schmid (1985) synonymized with
Nodosaria raphanistrum (OD = Nautilus raphanistrum
Linnaeus 1758; Recent, Mediterranean Sea). The basis for that
synonymy is not evident, as Linnaeus did not designate or figure
a type, nor did he indicate its depository. Vnec-Peyr (2005)
could not locate any type specimens of N. affinis in dOrbignys
collection at the Musum National dHistoire Naturelle, Paris.
The Chilean form is, therefore, aligned with the newer nomen,
Pyramidulina acuminata. Following their examination of
topotypes, Hayward et al. (2012) realized that the form assigned
to N. acuminata in Hayward (2002) was the smaller, deeperwater species, Anastomosa lamellata (= Nodosaria lamellata
Cushman and Stainforth 1945; new name for Dentalina carinata
Neugeboren 1856).
Plate 6, Figure 31
Comparative species: Laevidentalina advena (= Nodosaria

advena Cushman 1923; Recent, off NE United States, 960m) has
an arched test with more oblique sutures. The Chilean specimens
resemble the one illustrated by Papp and Schmid (1985, pl.
13, figs. 8, 9) as Dentalina inornata dOrbigny 1846 (Middle
Miocene, Austria), but the type figure and their other images (pl.
9, figs. 6, 7) of that species show it to be a slender, arcuate form
with very oblique sutures.
Occurrence: Ranquil Fm. (NAV5, PPP).
Laevidentalina sp. B
Plate 6, Figure 32
Remarks: This very rare form might be an aberrant L. inornata.

Comparative species: Hemirobulina yabei (= Dentalina yabei
Asano 1936a; Pliocene, Japan) has more compressed chambers
and a pupate form. Nodosaria fustiformis Schwager 1866
(Pliocene, Car Nicobar) is very similar but its sutures are not
oblique.
Occurrence: Navidad Fm. (FRM).
Nodosaria splendidula Schwager 1878
Plate 7, Figure 1
Nodosaria splendidula Schwager 1878, p. 521, pl. 1, fig. 5.
Type age and locality: Miocene, Sicily.

Comparative species: Nodosaria parexilis Cushman and K. C.
Stewart 1930 (Pliocene, California) lacks the apical spine.
Occurrence: El Peral beds (LPER), Navidad Fm. (MOS),
Ranquil Fm. (FRA, MIB, MS10, RAN, RQT), Santo Domingo
Fm. (VAL), Lacui Fm. (PCB, PNH).
Pyramidulina Fornasini 1894
Type species: Pyramidulina eptagona Fornasini 1894.

Distinguishing features: Same as Nodosaria but with distinct
longitudinal costae.
Pyramidulina acuminata (von Hantken 1876)
402
Comparative species: Pyramidulina raphana (= Nautilus

raphanus Linnaeus 1758; Recent, Mediterranean Sea) and P.
latejugata (= Nodosaria latejugata von Gmbel 1868; Late
Eocene, Germany) have two to three times more costae than P.
acuminata.
Occurrence: El Peral beds (NLP), Navidad Fm. (PPP, RAP),
Ranquil Fm. (FRA, FRM, MIB), Lacui Fm. (PCB).
Pandaglandulina Loeblich and Tappan 1955b
Type species: Pandaglandulina dinapolii Loeblich and Tappan

1955b.
Distinguishing features: Early stage slightly arcuate with
chambers increasing rapidly in size, followed by rectilinear
series of nearly equidimensional chambers.
Pandaglandulina obliquesuturata (Stache 1865)
Dentalina obliquesuturata Stache 1865, p. 207, pl. 22, fig. 6.

Cristellaria (Marginulina) angistoma Stache 1865, p. 207, pl. 22, fig.
46.
Marginulina subcrassa Schwager 1866, p. 240, pl. 6, fig. 82. (Plio
cene, Car Nicobar)

Remarks: Populations of this species in the Tertiary of Chile
vary widely in chamber size and sutural angles.
Comparative species: Stache (1865) distinguished more elongated
specimens as Cristellaria (Marginulina) angi-stoma, but Horni
brooks (1971) topotypes and the Chilean specimens form a
gradational series between the two species. Pandaglandulina
dinapolii Loeblich and Tappan 1955b (Early Pliocene, Italy) might
also be synonymous.
Occurrence: Navidad Fm. (CPUP, MOS, MPUP, NAV5, PPP,

PPT, PTA), Ranquil Fm. (MS10, FRA, RAN, RQK, RQT), Santo
Domingo Fm. (VAL).
Pseudonodosaria Boomgaart 1949
Type species: Glandulina discreta Reuss 1850.
Distinguishing features: Elongate, smooth, cylindrical test,
tapered or broadly rounded base, flush latitudinal sutures.
Pseudonodosaria aequalis (Reuss 1863b)
Plate 7, Figure 4
Glandulina aequalis Reuss 1863b, p. 48, pl. 1, fig. 28.

Pseudoglandulina aequalis (Reuss). JONES 1994, p. 73, pl. 61, fig. 32.
Glandulina? koreana McCulloch 1977, p. 12, pl. 49, fig. 37; pl. 96,
fig. 9. (Recent, off Korea, 42m)

Occurrence: El Peral beds (NLP), Navidad Fm. (PPP, PPT),
Ranquil Fm. (RAN, RQT), Lacui Fm. (PCB).
Pseudonodosaria brevis (dOrbigny 1846)
Dentalina brevis DORBIGNY 1846, p. 48, pl. 2, figs. 9, 10. PAPP

and SCHMID 1985, p. 30, pl. 12, figs. 811.
Nodosaria radicula var. glanduliniformis Dervieux 1894, p. 599,
pl. 5, figs. 37. (Miocene, Italy)
Glandulina aperta STACHE 1865, p. 188, pl. 22, fig. 11. (Late Tertiary,
New Zealand)

Basin, Austria.
Remarks: The type description does not detail the aperture, but
in every other respect the Chilean form matches the type figure.
Should examination of the type specimen reveal a different kind
of aperture, this form would represent an undescribed species.
Occurrence: Ranquil Fm. (MS10), Navidad Fm. (CPUP, MAT,
PPP), Ranquil Fm. (RQT).
Pseudonodosaria comatula (Cushman 1923)
Plate 7, Figures 7

Subfamily LINGULININAE Loeblich and Tappan 1961
Lingulina dOrbigny 1826; emend. Sellier de Civrieux and
Dessauvagie 1965; emend. Norling 1972.
Type species: Lingulina carinata dOrbigny 1826.
Lingulina sirakawaensis Nakamura 1942
Plate 7, Figure 8
Lingulina sirakawaensis Nakamura 1942, p. 92, pl. 6, fig. 5.
Type age and locality: Late Tertiary, Taiwan.

Stratigraphic range: Miocene to Pliocene(?).
Comparative species: Lingulina carinata dOrbigny 1839b
(Recent, Canary Islands) is similar but noncarinate.
Pseudolingulina McCulloch,1977
Type species: Pseudolingulina advena McCulloch 1977.
Pseudolingulina digitata (dOrbigny 1826)
Plate 7, Figure 9
Frondicularia digitata DORBIGNY 1826, p. 256, type-figure not

given; see Parker, Jones and Brady 1871, pl. 10, fig. 65.
Type age and locality: Pliocene, Italy.

Comparative species: Pseudolingulina linearis (= Frondicularia
linearis Philippi 1843; Tertiary, Germany) has slightly narrower
chambers and a less lobulate periphery; it could be synonymous.
Pseudolingulina gordabankensis McCulloch 1977 (Recent, Sea
of Cortez, 118m) has gently curved sutures and an arcuate test.
Pseudolingulina sacculus (= Frondicularia sacculus Terquem
1866; Early Jurassic) and P. spathulata (= Frondicularia
spathulata Brady 1879b; Recent, near New Guinea) lack the
medial depression.
Occurrence: Navidad Fm. Formation (CPUP).
Pseudolingulina nielseni Finger, n. sp.
Plate 7, Figure 10
Description: An elongate, lanceolate, compressed uniseries

of approximately 15 chambers separated by slightly concaveupward, flush sutures trending approximately 25 upward from
test periphery to longitudinal median, forming a stack of inverted
obtuse chevrons that terminate in a protruding radiate aperture;
periphery smooth, edges rounded; surface smooth, devoid of any
ornamentation.
Occurrence: Navidad Fm. (CPUP)
Upper depth limit: Upper bathyal; based on type occurrence.
Maximum relative abundance: Very rare (one specimen).
Occurrence: Navidad Fm. (CPUP, MOS, NAV5, PPP, PPT,

PTA), Ranquil Fm. (FRA, RQT), Lacui Fm. (PCB).
Nodosaria comatula Cushman 1923, p. 83, pl. 14, fig. 5.

Pseudoglandulina comatula (Cushman). KOHL 1985, p. 44, pl. 7, fig.
10. JONES 1994, p. 76, pl. 64, figs. 15.
Pseudonodosaria comatula (Cushman). ROBERTSON 1998, p. 50, pl.
17, fig. 1, 2.
403
Type age and locality: Early Miocene; roadcut exposure of

Navidad Formation along Camino de Pupuya (locality CPUP),
approximately 2.5km due south of La Boca.
Etymology: Named for paleontologist Sven N. Nielsen
(Universidad Austral de Chile, Valdivia).
Tollmannia Sellier de Civrieux and Dessauvagie 1965
Type species: Lingulina costata dOrbigny 1846.
Tollmannia costata (dOrbigny 1846)
Plate 7, Figure 11
Lingulina costata DORBIGNY 1846, p. 62, pl. 3, figs. 15. PAPP

and SCHMID, pl. 20, figs. 15.
Tollmannia costata (dOrbigny). LOEBLICH and TAPPAN 1987, pl.
442, figs. 1624.

Thomas 2012, p. 143, fig. 63, pl. 9, figs. 2225.
Type age and locality: Oligocene, Trinidad.

Upper depth limit: Probably upper bathyal based on M. compressa.
Comparative species: Mucronina morreyae (= Plectofrondicularia morreyae Cushman 1929; Tertiary, Ecuador) lacks a
medial ridge.
Occurrence: Navidad Fm. (PPP), Ranquil Fm. (MIB).

Basin, Austria.
Mucronina compressa (Costa 1855)
Frondicularia compressa COSTA 1855, p. 372, pl. 3, fig. 2.

Frondicularia foliacea SCHWAGER 1866, p. 236, pl. 6, fig. 76. (Plio
cene, Car Nicobar).
Frondicularia advena CUSHMAN 1923, p. 141, pl. 20, figs, 1, 2.
(Recent, off NE USA, 1962m).
Plectofrondicularia jarvisi Cushman and Todd 1945, p. 38, pl. 6,
fig. 4. (Miocene, Jamaica).
Mucronina compressa (Costa). HAYWARD, Kawagata, Sabaa,
Grenfell, Van Kerckhoven, Johnson and Thomas
2012, p. 144, fig. 63, pl. 9, figs. 3032; pl. 10, figs. 16.
Occurrence: Navidad Fm. (PPP, PPT).

Family Plectofrondiculariidae Montanaro-Gallitelli 1957
Mucronina Ehrenberg 1839
Type species: Nodosaria (les Mucronines) hasta dOrbigny
1826.
Mucronina acuta (Cushman and Stainforth 1945)
Plate 7, Figure 13
Plate 7, Figure 12
Plectofrondicularia nuttalli var. acuta Cushman and Stain

forth 1945, p. 39, pl. 5. fig. 24.
Mucronina acuta (Cushman and Stainforth). HAYWARD, Kawaga
PLATE 7
Figures 4, 10, 14, 17, 18, 20, 22a, and 23b are photomicrographs; all other images are SEMs. Scale bars in m.
1 Nodosaria splendidula Schwager, UCMP50125, FRA.

2, 3 P
andaglandulina obliquesuturata (Stache): 2, UCMP50126,
RQT. 3, UCMP50127, MS10.
4 Pseudonodosaria aequalis (Reuss), UCMP50128, RAN.
13 Mucronina compressa (Costa), UCMP50137, FRA.

14 Mucronina spatulata (Costa), UCMP50138, MOS.
1517 M
ucronina striata (dOrbigny): 15, UCMP50139, PPP. 16,
UCMP50140, CHO. 17, aberrant form, UCMP50141, PPP.
5, 6 P
seudonodosaria brevis (dOrbigny), PPP: 5, UCMP50129.
6, UCMP50130.
18 P
lectofrondicularia californica Cushman and Stewart,
UCMP50142, CPUP.
7 Pseudonodosaria comatula (Cushman), UCMP50131, FRA.
19, 20 P
lectofrondicularia digitalis (Neugeboren): 19, UCMP50143, FRA. 20, UCMP50144, PPP.
8 Lingulina sirakawaensis Nakamura, UCMP50132, PPP.

9 Pseudolingulina digitata (dOrbigny), UCMP50133, CPUP.
10 P
seudolingulina nielseni Finger, n. sp., holotype UCMP50134, CPUP.
11 Tollmannia costata (dOrbigny), UCMP50135, PPP.
12 Mucronina acuta (Cushman and Stainforth), UCMP50136, MIB.
404
ristellariopsis petersonae Finger, n. sp., holotype

21 C
UCMP50145, MAT.
22 Lenticulina calcar (Linn), UCMP50146, PPP.
23 Lenticulina douglasi Finger, UCMP50147, CUC.
Kenneth L. Finger
Plate 7
405
Comparative species: Plectofrondicularia vaughani Cushman

1927d (Eocene, Mexico) lacks the short costae at the apical end.
MOS, PPP, PPT, ), Ranquil Fm. (FRA, FRM, MIB, MS10)
156, figs. 64, 65; pl. 14, figs. 18.

Plectofrondicularia floridana CUSHMAN 1930a, p. 41, pl. 8, fig. 1
(Miocene, Florida). KOHL 1985, p. 45, pl. 9, figs. 6, 7. ROB
ERTSON 1998, p. 56, pl. 18, fig. 7. LOEBLICH and TAPPAN
1987, pl. 443, figs. 1, 2.
Type age and locality: Pliocene, California.
Mucronina spatulata (Costa 1855)
Stratigraphic range: Oligocene to earliest middle Pleistocene.
Plate 7, Figure 14
Frondicularia spatulata COSTA 1855, p. 372, pl. 2, fig. 19.

Mucronina spatulata (Costa). HAYWARD, Kawagata, Sabaa,
2012, p.151, fig. 63; pl. 12, figs. 1522.

Stratigraphic range: Late Eocene to middle Pleistocene.
Upper depth limit: Upper bathyal (400m).
Mucronina striata (dOrbigny 1826)
Frondicularia striata DORBIGNY 1826, p. 256, pl. 10, fig. 67.

Mucronina striata (dOrbigny). HAYWARD, Kawagata, Sabaa,
2012, p. 152, fig. 62; pl. 13, figs. 38.
Type age and locality: Fossil (age unknown), Italy.

Stratigraphic range: Late Eocene to early Pleistocene.
Upper depth limit: Not determined.
Comparative species: Ornamentation is similar to that of
Mucronina whitei (= Plectofrondicularia whitei Martin 1943;
Eocene, California), but that species has an asymmetrical outline
and the costae are discontinuous at the sutures. Mucronina
awamoana (= Plectofrondicularia awamoana Finlay 1939;
Middle Miocene, New Zealand) is a more slender form in which
the central ribs are the most prominent. Mucronina japonica
(= Parafrondicularia japonica Asano 1938; Pliocene, Japan) is
broadest toward the apertural end, and its surface is ornamented
with more numerous and finer longitudinal striations. The
Chilean species varies in position of maximum breadth and
continuity and extent of costae, but most specimens are widest
at midlength. The Chilean species may be the same form that
Robertson (1998) identified as Frondicularia sp. A.
Occurrence: El Peral beds (LPER, NLP), Ranquil Fm. (MS10),
Navidad Fm. (CPUP, MAT, MPUP, MOS, PPP, PPT, PTA),
Ranquil Fm. (MIB, MS10), Lacui Fm. (CHO, PCB).

al. 2012).
Occurrence: El Peral beds (LPER, NLP), Navidad Fm. (CPUP).
Plectofrondicularia digitalis (Neugeboren 1850)
Frondicularia digitalis NEUGEBOREN 1850, p. 120, pl. 3, fig. 3.

Plectofrondicularia parri FINLAY 1939b, p. 516, pl. 68, fig. 4. (Mid
dle Miocene, New Zealand)
Plectofrondicularia digitalis (Neugeboren). HAYWARD, Kawaga
Thomas 2012, p. 157, figs. 64, 65; pl. 14, figs. 1420.

Stratigraphic range: Late Eocene to Pliocene.
Upper depth limit: Outer neritic (for Plectofrondicularia parri
in van Morkhoven, Berggren and Edwards 1986; not found in
deep-water cores studied by Hayward et al. 2012).
Distinguishing features: Few costae, lobate chambers.
Comparative species: Plectofrondicularia semicostata (=
Frondicularia semicostata Neugeboren 1850; Tertiary, Romania)
has prominent lateral ridges.
Occurrence: Navidad Fm. (CPUP, MAT, MOS, NAV5, PPN, PPP,
PPT, PTA, RAP), Ranquil Fm. (FRA, LEB, MS10, RAN, RQT),
Lacui Fm. (CHO).
Family Vaginulinidae Reuss 1860
Remarks: This family is represented in a great majority of
foraminiferal studies, but there has been little consistency in
the assignments of some of its commonly represented elongate
forms (i.e., those with a uniserial stage). They are differentiated
in this study as follows:
Early stage distinctly coiled................................... LENTICULININAE
Initially keeled or angled periphery, uniseries circular in section.......
.................................................................................... Marginulinopsis
Maximum relative abundance: Common (MS10).
Compressed with prominent ornamentation...........Percultazonaria
Plectofrondicularia Liebus 1902

Type species: Plectofrondicularia concava Liebus 1902.
Compressed without ornamentation.........................Cristellariopsis
Plectofrondicularia californica Cushman and Stewart 1926
Plate 7, Figure 18
Plectofrondicularia californica Cushman and STEWART 1926, p.

39, pl. 6, figs. 18. HAYWARD, Kawagata, Sabaa, Gren
fell, Van Kerckhoven, Johnson and Thomas 2012, p.
406
Early stage indistinctly coiled.............................. MARGINULININAE

Flattened, numerous broad, low, oblique chambers.......... Astacolus
Circular in section with prominent costae....................Marginulina
Circular in section without ornamentation................. Hemirobulina

Slightly compressed ornamentation....................... Vaginulinopsis
Early stage not coiled............................................... VAGINULININAE
Compressed blade-like to palmate.................................. Vaginulina
Subfamily LENTICULININAE Chapman, Parr and Collins

1934
Cristellariopsis Rzehak 1895
Type species: Cristellariopsis punctata Rzehak 1895.
Cristellariopsis petersonae Finger, n. sp.
Plate 7, Figure 21
Description: Test compressed, initially involute planispiral,

later uncoiling; surface smooth; uniserial stage with relatively
wide rectilinear chambers; periphery of coil and dorsal side of
uniserial stage with a broad thin carina; apertural face ovate with
rounded edges; aperture radiate, protruding at dorsal angle.
Remarks: Whereas other uncoiling lenticulinine taxa are given
generic status, I am inclined to disagree with the synonymy
of this genus with Lenticulina noted by Loeblich and Tappan
(1987: 64).
Occurrence: El Peral beds (NLP), Navidad Fm. (LBZ, MAT,

MPUP, MOS, PPP, PPT PTA, RAP), Ranquil Fm. (MS10, RAN,
RQT), Lacui Fm. (CHO, CUC).
Maximum relative abundance: Common (MOS).
Lenticulina douglasi Finger 1992
Plate 7, Figure 23
Lenticulina douglasi Finger 1992, p. 70, pl. 4, figs. 3239.
Type age and locality: Miocene, California.

Diagnostic features: Subcircular-subpolygonal periphery with
narrow, rounded carina; usually 6 chambers, with slightly
depressed sutures.
Comparative species: Differs from Lenticulina subcultrata (=
Robulus subcultratus dOrbigny 1839c; Recent, Falkland Islands
and Canary Islands) by its lateral outline.
Occurrence: Navidad Fm. (MAT).
Occurrence: Ranquil Fm. (MS10), Navidad Fm. (MOS, MPUP,

PPP, PTA), Ranquil Fm. (FRM, MS10, RQT), Lacui Fm. (CHE,
CUC).
Maximum relative abundance: Rare (3 specimens).
Type specimens: Holotype UCMP50145; paratype UCMP50430

(from MAT).
Lenticulina foliata (Stache 1865)
Type age and locality: Early Miocene; Navidad Fm. Formation

in coastal bluff at Los Goterones (Navidad Fm. locality MAT),
about 1km north of Matanzas.
Cristellaria (Robulina) foliata Stache 1865, p. 245, pl. 23, fig. 24.
Etymology: Named for the late Dawn E. Peterson, who was

a great assistance in processing the Chilean samples for
microfaunal study.
Distinguishing features: Circular outline, moderately inflated,

smooth surface, with very narrow, sharp keel; 5 chambers in
final whorl, each shaped somewhat like a curved gourd, or a
stretched comma with tail flaring toward margin; sutures flush,
asymmetrically curved.
Lenticulina Lamarck 1804; emend. Marie 1941

Type species: Lenticulites rotulata Lamarck 1804.
Lenticulina calcar (Linnaeus 1758)
Plate 7, Figure 22
Nautilus calcar LinnAEUS 1758, p. 709; type-figure in Plancus

1743, pl. 1, figs. 3gI, 4ln; also Gualtieri 1742, pl. 19, figs. B, C.
Lenticulina calcar (Linnaeus). KOHL 1985, p. 47, pl. 10, figs. 4, 5.
WHITTAKER 1988, p. 47, pl. 5, figs. 3, 4. JONES 1994, p. 81, pl.
70, figs. 912. ROBERTSON 1998, p. 62, pl. 20, fig. 4. HAY
14, figs. 9, 10; pl. 15, figs. 1, 2. ZAPATA and CEAR 2004, p. 30,
pl. 10, fig. 10.

Upper depth limit: Neritic (Zapata and Cear 2004).
Distinguishing features: Circular outline, moderately inflated,
smooth surface, narrow carina; outer whorl of 56 chambers,
some with peripheral spines at midpoint of chamber margin.
Comparative species: Lenticulina douglasi Finger 1992;
Miocene, California) lacks peripheral spines, is more
compressed. and has a subcircular-subpolygonal outline.
Plate 8, Figure 1
Comparative species: Lenticulina orbicularis (= Cristellaria

vortex var. orbicularis A. Silvestri 1898; Pliocene, Italy) displays
7 chambers, but might be synonymous.
Occurrence: Navidad Fm. (PPT), Ranquil Fm. (FRA, MIB),
Lacui Fm. (CHE).
Lenticulina cf. L. gibba (dOrbigny 1839a)
Plate 8, Figure 2
Cristellaria gibba DORBIGNY 1839a, p. 40; type-fig. in v. 8, pl. 7,

figs. 20, 21.
Lenticulina gibba (dOrbigny). JONES 1994, p. 81, pl. 69, figs. 8, 9.
Type age and locality: Recent, Caribbean and Mediterranean.

Distinguishing features: Moderately compressed, slightly elongate,
noncarinate, translucent; shows five tightly coiled chambers with
moderatley arched, slightly depressed to flush sutures.
Comparative species: The type-figure of Lenticulina gibba has a
depressed umbilicus and is slightly carinate.
407
Occurrence: Navidad Fm. (NAV5), Ranquil Fm. (LEB, RQT),

Lacui Fm. (CHE, PNH).
Lenticulina glaucina (Stache 1865)
Plate 8, Figure 3
Cristellaria (Cristellaria) glaucina Stache 1865, p. 241, pl. 23, fig. 20.

Distinguishing features: Circular outline with wide, sharp carina;
910 chambers in last whorl; very oblique, curved sutures.
Comparative species: Lenticulina subcarinata (= Cristellaria
orbicularis var. subcarinata Cushman 1917b; Recent, East
Indies) has a narrower and more robust keel, with less tangential
sutures.
Occurrence: Ranquil Fm. (FRA, RQT).

Remarks: Hornibrook (1971) indicated its stratigraphic range in
New Zealand as Oligocene to possibly lower Miocene.
Occurrence: Navidad Fm. (CPUP, PPP, PTA, RAP), Ranquil
Fm. (FRA, FRM), Lacui Fm. (CHO, CUC, PNH).
Lenticulina miyagiensis (Asano 1937)
Plate 8, Figure 6
Robulus miyagiensis ASANO 1937, p. 32, text-fig. 3.
Type age and locality: Miocene, Japan.

Distinguishing features: Circular outline; sharp, wide keel; 10
12 chambers in last whorl, with moderately oblique sutures,
Plate 8, Figure 4
Comparative species: The sutures of Lenticulina iota (=

Cristellaria iota Cushman 1923; Recent, Gulf of Mexico, 359m)
are nearly radial. Lenticulina etigoensis (= Robulus etigoensis
Asano 1938 (Pliocene, Japan) has a narrower keel and its
chambers are longer and narrower.
Robulina grandis BORNEMANN 1860, p. 156, pl. 6, fig. 1.
Occurrence: Ranquil Fm. (MIB, RQT).
Type age and locality: Early Oligocene, Germany.
Stratigraphic range: Early Oligocene to Early Miocene.
Lenticulina neopolita Finger, new name

Lenticulina grandis (Bornemann 1860)
Occurrence: Navidad Fm. (PPN).

Lenticulina halophora (Stache 1865)
Plate 8, Figure 5
Planularia halophora Stache 1865, p. 248, pl. 23, fig. 28.

Cristellaria expansa Cushman 1917b, p. 658, pl. 46, fig. 2. (Recent,
China Sea, 878m)
Plate 8, Figure 7
Nomen substitutum pro Cristellaria polita Schwager 1866, p. 245,

pl. 7, fig. 89. (Not Cristellaria polita Reuss 1856)

Distinguishing features: Carina wide with each successive
section distinctly delineated.
PLATE 8
Figures 1b, 2b, 3b, 4a, 6b, 7b, 9b, 10a, 11a, 12b, 13a, and 14c are photomicrographs; all other images are SEMs. Scale bars in m.
1 Lenticulina foliata (Stache), UCMP50148, FRA.

2 Lenticulina cf. L. gibba (dOrbigny), UCMP50149, LEB.
3 Lenticulina glaucina (Stache), UCMP50150, FRA.
4 Lenticulina grandis (Bornemann), UCMP50151, PPN.
408
8 L
enticulina nuttalli (Cushman and Renz), UCMP50155,
MS10.
9 Lenticulina stellata (Seguenza), UCMP50156, RQT.
enticulina subcultrata (dOrbigny): 10, carinate, UCMP10, 11 L
50157, FRA. 11, noncarinate, UCMP50158, RQT.
5 Lenticulina halophora (Stache), UCMP50152, CPUP.
12 Lenticulina tangens (LeRoy), UCMP50159, MIB.
6 Lenticulina miyagiensis (Asano), UCMP50153, MIB.
13 Lenticulina thalmanni (Hessland), UCMP50160, NAV5.
7 L
enticulina neopolita Finger, new name, UCMP50154,
MIB.
14 Lenticulina variabilis (Reuss), UCMP50161, PPP.
Kenneth L. Finger
Plate 8
409
Remarks: The Chilean form matches well with the type figure
of Cristellaria polita Schwager 1866; Pliocene, Car Nicobar),
which is a junior homonym of Lenticulina polita (= Cristellaria
polita Reuss 1856). Srinivasan and Sharma (1980) reassigned
Schwagers species to Lenticulina (= Robulus cushmani
Galloway and Wissler 1927; Pleistocene, California), but that
species has a much narrower and slightly thicker keel.
Occurrence: Ranquil Fm. (MIB, RQT).
Etymology: Addition of suffix neo (new) to polita distinguishes
the new name from the old one.
Lenticulina nuttalli (Cushman and Renz 1941)
Plate 8, Figure 8
Robulus nuttalli Cushman and Renz 1941, p. 11, pl. 2, fig. 10.
Type age and locality: Late Oligocene to Early Miocene,

Venezuela.
Distinguishing features: Compressed, slightly uncoiled, slightly
lobulate periphery; sutures initially limbate, but later depressed;
final whorl has 67 chambers with short inner margins that rise
around a plug-like central boss.
Lenticulina stellata (Seguenza 1880)
Plate 8, Figure 9
Robulus stellata Seguenza 1880, p. 144, pl. 13, fig. 29.
Type age and locality: Late Miocene, Italy.

Distinguishing features: Very compressed, evolute coil with
about 911 narrow chambers in the final whorl and fringed with
a wide and thin keel.
Comparative species: Lenticulina iota (= Cristellaria iota
Cushman 1923; Recent, Gulf of Mexico, 359m) has nearly radial
sutures and a narrower keel. Lenticulina etigoensis (= Robulus
etigoensis Asano 1938; Pliocene, Japan) also has a narrow keel.
Occurrence: El Peral beds (NLP), Ranquil Fm. (RQT), Santo
Domingo (VAL).
Lenticulina subcultrata (dOrbigny 1839c)
Robulina subcultrata DORBIGNY 1839c, p. 98, pl. 5, figs. 19, 20.

Lenticulina clericii (Fornasini). WHITTAKER 1988, pl. 5, figs. 5, 6
(MiocenePliocene, Ecuador).
Type age and locality: Recent; localities in Falkland Islands and

Canary Islands.
Distinguishing features: Circular with smooth surface and
moderate umbo, varies from noncarinate to moderately keeled;
usually 810 chambers in final whorl, sutures slightly curved,
ranging from oblique to nearly radial.
410
Remarks: It is difficult to consistently differentiate dOrbignys

species from several others that were subsequently described
(see below), as the differences shown by the type figures may be
intraspecific.
Comparative species: Lenticulina subcultrata is very similar to,
and possibly a junior synonym of, L. nikobarensis (= Cristel
laria nikobarensis Schwager 1866; Pliocene, Car Nicobar).
Cristellaria clericii Fornasini 1895 (Lower Pliocene, Italy) is
noncarinate and subacute in edge view, and the inner half of its
sutures are strongly curved. Some Chilean specimens have the
wider keel of Lenticulina similis (= Robulina similis dOrbigny
1846; Middle Miocene, Austria), but L. similis lacks an umbo and
appears to have limbate sutures. Lenticulina iota (= Cristellaria
iota Cushman 1923; Recent, Gulf of Mexico) has 1315 chambers
in the final whorl. Lenticulina cushmani (= Robulus cushmani
Galloway and Wissler 1927; Pleistocene, California) has limbate
sutures, but otherwise matches the keeled form. Lenticulina
smileyi (= Robulus smileyi Kleinpell 1938; Miocene, California)
has a prominent boss and slightly limbate sutures. The plasticity
of this common form suggests that it is represented by multiple
synonyms.
Occurrence: All localities except MIB.
Maximum relative abundance: Common (MOS, RAP, MPUP,
RAN, CHO).
Lenticulina tangens (LeRoy 1939)
Plate 8, Figure 12
Robulus tangens LeRoy 1939, p. 233, pl. 8, figs. 12, 13.
Type age and locality: Miocene?, Indonesia.

Distinguishing features: Circular outline with thin keel, 57
chambers in final whorl, sutures very oblique, and umbilical
area, wide, glassy, and flush with surrounding surface.
Comparative species: Lenticulina nigrisepta (= Cristellaria
nigrisepta Koch 1926; middle Tertiary, Borneo) has 8 chambers
with broader sutures. Lenticulina gyroscalpra (= Cristellaria
(Cristellaria) gyroscalprum Stache 1865 (late Tertiary, New
Zealand) and L. taettowata (= C. (Robulina) taettowata Stache
1865; late Tertiary, New Zealand) have a more prominent boss.
Lenticulina americana (= Cristellaria americana Cushman
1918b; Miocene, Florida) has raised sutures extending from a
raised umbo to a raised narrow keel.
Occurrence: Ranquil Fm. (MIB).
Lenticulina thalmanni (Hessland 1943)
Plate 8, Figure 13
Robulus thalmanni Hessland 1943, p. 265; type-figure in Brady

1884, pl. 69, fig. 13.
Lenticulina (Robulus) americanus (Cushman). WHITTAKER, 1988,
pl. 5, figs. 1, 2 (late Cenozoic, Ecuador).
Lenticulina thalmanni (Hessland). JONES 1994, p. 81, pl. 69, fig. 13.
Type age and locality: Recent, West Indies.

Distinguishing features: Moderately inflated test with circular

outline; smooth surface; very narrow, blunt keel; ~9 chambers in
final whorl, slightly curved sutures.
Comparative species: Lenticulina americana (= Cristellaria
americana Cushman 1918b; Miocene, Florida) has 67
chambers in the final whorl with thick sutures that coalesce
into a prominent umbo. Lenticulina subcarinata (Cristellaria
orbicularis var. subcarinata Cushman 1917b; Recent, East
Indies) has a robust keel and a prominent boss.
Lenticulina sp. C
Plate 9, Figure 3
Distinguishing features: Sublacrimate outline, slightly com

pressed; outer whorl comprised of four chambersm, slight keel;
surface smooth, translucent.
Occurrence: Navidad Fm. (LBZ, MOS, NAV5, PPP), Ranquil

Fm. (RQT).
Lenticulina sp. D
Maximum relative abundance: Common (LBZ).
Distinguishing features: Outline slightly elongate, noncarinate.

moderately compressed in edge view; 5 tightly coiled, translucent
chambers; moderatley arched, slightly depressed.
Lenticulina variabilis (Reuss 1850)
Plate 8, Figure 14
Cristellaria variabilis Reuss 1850, p. 369, pl. 46, figs. 15, 16.
Lenticulina variabilis (Reuss). JONES 1994, p. 80, pl. 69, figs. 1116.
Lenticulina variabilis Romanova, in GLAZUNOVA et al. 1960, p. 73,
pl. 12, figs. 38.
Type age and locality: Tertiary, Germany.

Distinguishing features: Ovate outline, carinate; very
compressed in edge view; final whorl with 4 chambers that
increase rapidly in size; sutures curved, slightly depressed.
Comparative species: Lenticulina intumescens (= Cristellaria
(Robulina) pauperata var. intumescens Reuss 1866; middle
Tertiary, Germany) is much less compressed.
Occurrence: Ranquil Fm. (MS10), Navidad Fm. (PPP), Santo
Domingo Fm. (VAL).
Lenticulina sp. A
Plate 9, Figure 4
Occurrence: El Peral beds (LPER), Navidad Fm. (NAV5),

Ranquil Fm. (FRA, LEB).
Neolenticulina McCulloch 1977
Type species: Neolenticulina chathamensis McCulloch 1977.
Neolenticulina peregrina (Schwager 1866)
Cristellaria peregrina SCHWAGER 1866, p. 245, pl. 7, fig. 89.

Lenticulina peregrina (Schwager). Srinivasan and Sharma
1980, p. 34, pl. 6, fig. 26. KOHL 1985, p. 48, pl. 10, figs. 10, 11;
pl. 11, fig. 1. VAN MORKHOVEN, BERGGREN and ED
WARDS 1986, p. 92, pl. 27, figs. 1, 2.
Neolenticulina peregrina (Schwager). HOLBOURN, HENDER
SON and MACLEOD 2013, p. 368369.
Type age and locality: Middle Pliocene, Car Nicobar.

Plate 9, Figure 1
Lenticulina convergens (Bornemann). HOLBOURN, HENDERSON

and MACLEOD 2013, p. 332333.

PTA), Ranquil Fm. (FRA, FRM, RQT).
Distinguishing features: Outline rounded, but final two

chambers have nearly straight outer edges that adjoin at 135;
carinate, fusiform in edge view; 57 translucent chambers
in final whorl; sutures slightly curved, flush; umbo large, flush,
opaque; surface smooth.

Percultazonaria Loeblich and Tappan 1986
Type species: Cristellaria subaculeata Cushman 1923.
Remarks: Differs from Vaginulinopsis by its more prominent

coil and elevated ornamentation.
Percultazonaria encinasi Finger, n. sp.
Lenticulina sp. B
Plate 9, Figure 2
Distinguishing features: Ovate outline; inflated, with wide keel;

some nodules along sutures.
Comparative species: Lenticulina expansa (= Cristellaria
expansa Cushman 1917b; Recent, Philippines) is nearly identical
in lateral view, but it is highly compressed in edge view.
Plate 9, Figure 7
Description: Test elongate and moderately compressed, initial

stage planispiral and involute, later uncoiling and rectilinear,
chambers broad and low, sutures moderately oblique and slightly
curved, coil with thin, narrow keel bearing short spines, later
subacute on ventral side and slightly flattened on ventral side,
radiate aperture protruding from dorsal angle on neck, rows of
nodules along sutures tend to be less prominent between later
chambers.
Comparative species: Percultazonaria encinasi differs from P.
vaughani by its smaller size, more elongated and hook-shaped
test, more inflated later chambers, and weak sutural nodules.
Cristellaria spinulosa Costa (in (Fornasini) 1894 (Tertiary, Italy)
411
has ornamentation similar to the Chilean species, but it does not

appear to uncoil.
however, neither resemble the planispiral stage of the uncoiling

Percultazonaria species they are associated with.
Occurrence: Navidad Fm. (CPUP, LBZ, MAT, MOS, NAV5,

PPN, PPP, PPT, PTA, RAP), Ranquil Fm. (FRA, RAN, RQT),
Lacui Fm. (CHO, CUC).
Comparative species: Lenticulina (Robulus) vicksburgensis

(Cushman) of Whittaker 1988 (pl. 5, figs. 9, 10; late Cenozoic,
Ecuador) may be synonymous. The type figure of Lenticulina
vicksburgensis (= Cristellaria vicksburgensis Cushman 1922b,
Early Oligocene, Mississippi), however, has finer nodes clustered
very close to the periphery.

Type specimens: Holotype UCMP50168; paratypes UCMP50434
and UCMP50435 (both from CUC).
Type age and locality: Early Miocene; coastal bluff blockfall

south of Cucao (Lacui Fm. locality CUC).
Etymology: Named for geologist Alfonso Encinas (Universidad

de Concepcon).
Plate 9, Figure 9
Percultazonaria cf. P. mamilligera (Karrer 1865)
Plate 9, Figure 8
Cristellaria (Cristellaria) mamilligera Karrer 1865, p. 76, pl. 16,

fig. 5.
Lenticulina mamilligera (Karrer). HORNIBROOK 1971, p. 17, pl. 2,
fig. 21.

Distinguishing features: Surface ornamented with numerous
large tubercles; irregularly serrated keel narrows in later
chambers, becoming nonserrate with short spines.
Remarks: The type-figure of P. mamilligera shows a wider keel
whose edge consists of arcuate segments that adjoin as short,
oblique projections in line with the sutures. Hornibrooks (1971)
illustration of a topotype that is slightly uncoiled and noncarinate
might not be the same species. Both the Chilean form and
Karrers species appear to be juveniles that have yet to uncoil;
Percultazonaria obliquispinata Finger, n. sp.
Description: Test stout and compressed, initial stage planispiral

and involute, later uncoiling and rectilinear, chambers broad and
low, sutures slightly depressed, with rows of tubercles along base
of succeeding chambers, periphery of coil indistinctly carinate
with laterally asymmetrical spines, apertural face slightly
flattened, smooth, radiate aperture protruding from dorsal angle
on short neck.
Comparative species: Percultazonaria alazanensis (= Cristellaria alazanensis Cushman 1927d; Eocene, Mexico) and its
junior subjective synonym (= Cristellaria calcar var. alazanensis
Nuttall 1932; Oligocene, Mexico) lack sutural nodules and bear
spines that are more symmetrical than on P. obliquispinata.
Occurrence: Navidad Fm. (CPUP, NAV5), Ranquil Fm. (FRA,
FRM, MIB, RAN, RQK, RQT), Lacui Fm. (PNH, CUC).
(from FRM) and UCMP50437 (from CPUP).
PLATE 9
Figures 1a, 3b, 4b, 5b, 11a, 15b, 16b, 17b, and 18a are photomicrographs; all other images are SEMs. Scale bars in m.
1 Lenticulina sp. A, UCMP50162, MIB.
10 Percultazonaria vaughani (Cushman), UCMP50171, PPP.
2 Lenticulina sp. B, UCMP50163, FRM.
11, 12 Saracenaria schencki Cushman and Hobson, PPP: 11,

UCMP50172. 12, UCMP50173.
3 Lenticulina sp. C, UCMP50164, MIB.

4 Lenticulina sp. D, UCMP50165, PNH.
5, 6 N
eolenticulina peregrina (Schwager): 5, UCMP50166,
FRA; 6, juvenile form, UCMP50167, PPP.
ercultazonaria encinasi Finger, n. sp., holotype UCMP7 P
50168, CUC.
8 P
ercultazonaria cf. P. mamilligera (Karrer), UCMP50169,
FRM.
9 P
ercultazonaria obliquispina Finger, n. sp., holotype
UCMP50170, RQT.
412
13 Amphicoryna badenensis (dOrbigny), UCMP50174, RQT.

14 Amphicoryna cf. A. scalaris (Batsch), UCMP50175, PPP.
15 Astacolus crepidulus (Montfort), UCMP50176, LEB.
16 Astacolus cymboides (dOrbigny), UCMP50177, FRA.
17 Astacolus jordai (Colom), UCMP50178, FRM.
stacolus cf. A. mayi (Cushman and Parker), UCMP50179,
18 A
RAN.
19 Astacolus mexicanus (Nuttall), UCMP50180, FRM.
Kenneth L. Finger
Plate 9
413
Type age and locality: Early Miocene; Punta Huenteguapi

intertidal platform (Ranquil Fm. locality RQT).
Etymology: Morphodescriptor derived from the Latin obliquus
+ spinata, meaning obliquely spined.
Percultazonaria vaughani (Cushman 1918a)
Plate 9, Figure 10
Cristellaria vaughani Cushman 1918a, p. 61, pl. 22, fig. 3.

Lenticulina (Robulus) cristi SKINNER 1963, p. 150, text-figs. 1, 2.
(Miocene, Louisiana)
Type age and locality: Miocene, Panama.

Remarks: Some specimens have short marginal spine. The
degrees of uncoiling and of sutural nodosity vary within popu
lations.
Comparative species: Percultazonaria hanseni (= Lenticulina
hanseni Garrett 1942; Miocene, Texas) is a similar form, but has
a more inflated spiral stage and a tendency for sutural tubercles
coalesce into ridges. In the Chilean Neogene, P. obliquispinata is
a smaller and less compressed with more-inflated later chambers.
Occurrence: El Peral beds (NLP), Navidad Fm. (MPUP, PPP,
PTA).
Maximum relative abundance: Common.
Saracenaria Defrance, in de Blainville and 1824
Type species: Saracenaria italica Defrance, in de Blainville
1824.
Saracenaria schencki Cushman and Hobson 1935
Saracenaria schencki Cushman and Hobson 1935, p. 57, pl. 8, fig. 11.
Lenticulina (Saracenaria) carapitana Franklin. WHITTAKER 1988,
p. 51, pl. 5, figs. 19, 20 (Middle Miocene, Ecuador).
Subfamily MARGINULININAE Wedekind 1937

Amphicoryina Schlumberger, in Milne-Edwards 1881
Type species: Marginulina falx Jones and Parker 1860.
Amphicoryna badenensis (dOrbigny 1846)
Plate 9, Figure 13
Nodosaria badenensis DORBIGNY 1846, p. 38, pl. 1, figs. 34, 35.

PAPP and SCHMID 1985, p. 27, pl. 7, figs. 18.

Basin, Austria.
Comparative species: Amphicoryna scalaris (= Nautilus
scalaris Batsch 1791; probably Recent, Italy) and Amphicoryna
striaticollis (= Nodosaria striaticollis dOrbigny 1839b; Recent,
Canary Islands) have much more densely costate chambers that
increase more rapidly in size. The chambers of Amphicoryna
ehrenbergiana (= Nodosaria ehrenbergiana Neugeboren 1852;
Tertiary, Romania) are more fusiform. Amphicoryna protumida
(= Nodosaria protumida Schwager 1866; Pliocene, Car Nicobar)
is curved, much more tapered, more densely costate and with a
long, smooth, lipped neck. A few specimens of the Chilean A.
badenensis have the last two chambers separated by a long tube,
as in Amphicoryna separans (= N. scalaris var. separans Brady
1884; Recent, off New Zealand, 503m), which is probably a
variant of A. badenensis. Amphicoryna scalaroides (= Nodosaria
scalaroides ten Dam and Reinhold 1942; Middle Miocene, The
Netherlands) also has more costae. Amphicoryna catesbyi (=
Nodosaria catesbyi dOrbigny 1939; Recent, Cuba) has fewer
and more-pronounced costae (also see Le Calvez 1977).
Occurrence: Navidad Fm. (CPUP, PPN, PPP), Ranquil Fm.
(RQT), Chiloe (PCB, CHE).
Amphicoryna cf. A. scalaris (Batsch 1791)
Plate 9, Figure 14
Type age and locality: Eocene (Oligocene?), California.
Nautilus (Orthoceras) scalaris Batsch 1791, p. 1, 4, pl. 2, fig. 4.
Stratigraphic range: Eocene (Oligocene?)Middle Miocene.
Type age and locality: Oligocene, Hungary.
Upper depth limit: Outer neritic (for Saracenaria spp. occurrences

off California (Ingle 1980) and New Zealand (Hayward et al.
2010).
Remarks: I recovered a single microspheric specimen with

an arcuate early stage composed of narrow, oblique chambers
followed by a single chamber with bladed costae.
Comparative species: Saracenaria carapitana (= S. italica

var. carapitana Franklin 1944; Early Oligocene, Venezuela)
has less-oblique sutures that remain limbate to the peripheral
keel. The apertural face of S. italica DeFrance 1824 (fossil,
Italy) is nearly an equilateral triangle and the test is not arcuate.
Saracenaria senni Hedberg 1937 (middle Tertiary, Venezuela)
is a small form that in edge view shows early chambers that are
very narrow (compressed) and an ultimate chamber that is very
inflated. Saracenaria latifrons (= Cristellaria latifrons Brady
1884; Recent, localities off New Zealand and West Indies) looks
somewhat like an unusually inflated Astacolus.
Comparative species: Astacolus tunicata (= Cristellaria

(Marginulina) tunicata von Hantken 1868; Oligocene, Hungary)
is narrower and devoid of costae.
Occurrence: Navidad Fm. (CPUP, MAT, MOS, MPUP, NAV5,

PPP, PPT, PTA), Ranquil Fm. (FRA, FRM, MS10, RAN, RQK,
RQT), Lacui Fm. (CHO, PCB, PNH, CUC).
Plate 9, Figure 15
414

Astacolus de Montfort 1808
Type species: Astacolus crepidulatus de Montfort 1808.
Astacolus crepidulus (Fichtel and Moll 1798)
Nautilus crepidula Fichtel and Moll 1798, p. 107, pl. 19, figs. gi.
Astacolus crepidulus (Fichtel and Moll). Rgl and Hansen 1984,
p. 66, textfig 27, pl. 26, figs. 1, 2. FINGER 1992, p. 72, pl. 6, figs.
2, 3. JONES 1994, p. 79, pl. 67, fig. 20; pl. 68, figs. 1, 2. HAY
14, figs. 1, 2.
Type age and locality: Recent, Italy.
Astacolus mexicanus (Nuttall 1932)
Upper depth limit: Neritic; based on occurrences off New

Zealand (Hayward et al. 2010).
Occurrence: Navidad Fm. (MAT, PPT), Ranquil Fm. (LEB),
Lacui Fm. (PCB).
Astacolus cymboides (dOrbigny 1846)
Plate 9, Figure 19
Vaginulina elegans var. mexicana NUTTALL 1932, p. 16, pl. 3, figs.

12, 16.
Type age and locality: Early Oligocene, Mexico.

Stratigraphic range: Early Oligocene to Early Miocene or
possibly younger.
Plate 9, Figure 16
Occurrence: Navidad Fm. (CPUP), Ranquil Fm. (FRA, FRM).
Cristellaria cymboides DOrbigny 1846, p. 85, pl. 3, figs. 30, 31.

Lenticulina cymboides (dOrbigny). PAPP and SCHMID 1985, p. 39,
pl. 24, figs. 13.

Basin, Austria.
Comparative species: Astacolus mayi (= Robulus mayi Cushman
and Parker 1931; Miocene, California) is more strongly coiled
in its early stage and has an acute periphery. Cristellaria
berthelotiana dOrbigny 1839b (Recent, Canary Islands) is
flatter, and successive chambers increase more gradually in size.
Occurrence: Ranquil Fm. (FRA, FRM), Lacui Fm. (CHO).

Astacolus jordai (Colom 1943)
Plate 9, Figure 17
Lenticulina sublituus jordai Colom 1943, p. 238, pl. 21, figs. 15, 14,
15, pl. 24, figs. 7476.
Lenticulina (Astacolus) nuttalli (Todd and Kniker). Whittaker
1988, pl. 5, figs. 13, 14 (Middle Miocene, Ecuador).
Type age and locality: Miocene, Spain.

Comparative species: Astacolus nuttalli (= Marginulina nuttalli
Todd and Kniker 1952, nom. nov. pro L. sublituus Nuttall 1932;
Early Oligocene, Mexico) has narrow and less-oblique sutures,
and its chambers increase more rapidly in width. Astacolus
compressus (= Cristellaria compressus Stache 1865; early
Tertiary, New Zealand) has more chambers and thinner sutures.
Astacolus insolita (= Cristellaria insolita Schwager 1866;
Pliocene, Car Nicobar) has an arcuate test and slightly sinuous
thin sutures. Astacolus reniformis (= Cristellaria reniformis
dOrbigny 1846; Middle Miocene, Austria) is distinctly carinate.
Occurrence: Navidad Fm. (PPP, CPUP), Ranquil Fm. (FRM).
Astacolus cf. A. mayi (Cushman and Parker 1931)
Plate 9, Figure 18
Robulus mayi Cushman and Parker 1931, p. 2, pl. 1, figs. 35.
Astacolus multicameratus (Cushman and Stainforth 1945)
Plate 10, Figure 1
Marginulina sublituus var. multicamerata Cushman and Stain

forth 1945, p. 23, pl. 3, figs. 6, 7.
Type age and locality: Probably Late Oligocene, Trinidad.

Occurrence: Navidad Fm. (LBZ), Ranquil Fm. (FRA), Lacui
Fm. (PNH).
Astacolus novambiguus Finger, n. sp.
Plate 10, Figure 2
Description: Test very strongly compressed, narrow, elongate,

slightly arcuate, slightly lobulate; early stage slightly curved, later
chambers obliquely rectilinear, width/height ratio progressively
decreasing through series, sutures depressed; surface smooth;
aperture radiate at dorsal angle.
Comparative species: This species closely resembles Marginulina ambigua Schwager (in Waagen 1866; Late Jurassic,
Germany), which is a much older species with a slightly more
pronounced early stage and arcuate test, which appears to have a
narrow carina along its outer edge.
Occurrence: Navidad Fm. (CPUP, PPT, PTA), Lacui Fm. (PNH).
Type specimens: Holotype UCMP50182; paratype UCMP50429
(from CPUP).
Type age and locality: Early Miocene, Navidad Fm. Formation
in coastal bluff of Punta Alta (Navidad Fm. locality PTA).
Etymology: Derived by adding the Latin novus, meaning new or
young, as the prefix to the existing nomen ambiguus.
Astacolus sp. A
Plate 10, Figure 3
Type age and locality: Miocene, central California.
Remarks: Lenticulina mayi is more compressed and strongly

spiralled, with an acute periphery.
Occurrence: Navidad Fm. (PPP), Ranquil Fm. (RAN), Lacui

Fm. (PCB).
Plate 10, Figure 4
Astacolus sp. B

415

Hemirobulina dOrbigny 1826
Type species: Cristellaria arcuatula Stache 1865.
Comparative species: Hemirobulina glabra (= Marginulina

glabra dOrbigny 1826; Pliocene, Italy) has more chambers and
transverse sutures.
Hemirobulina pedum (dOrbigny 1846)
Occurrence: El Peral beds (NLP), Navidad Fm. (CPUP, NAV5,

PTA).
Marginulina pedum DORBIGNY 1846, p. 68, pl. 3, figs. 13, 14.

Vaginulinopsis pedum (dOrbigny). PAPP and SCHMID 1985, p. 37,
pl. 21, figs. 6, 7.
Pseudodimorphina galapagosensis McCulloch 1977, p. 9, pl. 76,
fig. 14; pl. 94, figs. 18-21, 23. (Recent, Galpagos)
Plate 10, Figure 5

Occurrence: Ranquil Fm. (FRA, FRM, MIB, RAN), Santo

Domingo Fm. (VAL)
Hemirobulina similis (dOrbigny 1846)
Hemirobulina yabei (Asano 1936a)
Plate 10, Figure 8
Dentalina yabei Asano 1936a, p. 329, pl. 36, figs. 1012.
Type age and locality: Pliocene, Japan.

Remarks: This form is nearly uniserial with slightly diagonal
sutures.
Occurrence: Navidad Fm. (CPUP, MOS, RAP).
Marginulina similis DOrbigny 1846, p. 69, pl. 3, figs. 15, 16.

PAPP and SCHMID 1985, p. 37, pl. 21. figs. 1012.
Marginulina subbullata Hantken. FINGER 1992, p. 69, pl. 3, figs. 15,
16. PAPP and SCHMID 1985, p. 73, pl. 6. figs. 32, 33.
Marginulina obesa (Cushman). HOLBOURN, HENDERSON and
MACLEOD 2013, p. 346347.
Marginulina dOrbigny 1826

Type species: Marginulina raphanus dOrbigny 1826.

Basin, Austria.
Marginulina cubana Palmer 1940
Marginulina cubana Palmer 1940, p. 278, pl. 52, fig. 18.
PLATE 10
Figures 1b, 3b, 4b, 16, 18a, 19a, 20a, and 22b are photomicrographs; all other images are SEMs. Scale bars in m
1 A
stacolus multicameratus (Cushman and Stainforth),
UCMP50181, LBZ.
2 A
stacolus novambiguus Finger, n. sp., holotype UCMP
50182, PTA.
15 Vaginulinopsis sp. A, UCMP50195, VAL.

16 Vaginulinopsis sp. B, UCMP50196, NLP.
3 Astacolus sp. A, UCMP50183, FRA.
17 Vaginulinopsis sp. C, UCMP50197, MS10.
4 Astacolus sp. B, UCMP50184, RQT.
18 Planularia cassis (Fichtel and Moll), UCMP50198, PPP.
5 Hemirobulina pedum (dOrbigny), UCMP50185, FRA.
19 P
rismatomorphia tricarinata (dOrbigny), UCMP50199,
NLP.
6, 7 H
emirobulina similis (dOrbigny): 6, UCMP50186, CPUP.
7, juvenile, UCMP50187, MS10.
8 Hemirobulina yabei (Asano), UCMP50188, MOS.
9, 10 Marginulina cubana Palmer, PPP: 9, megalospheric,
UCMP50189. 10, microspheric, UCMP50190.
11 Vaginulinopsis cf. V. chetae (Basov), UCMP50191, PCB
12 Vaginulinopsis costatus (Batsch), UCMP50192, PNH
13 V
aginulinopsis lueneburgensis (Clodius), UCMP50193,
PTA.
416
14 Vaginulinopsis subelegans? (Parr), UCMP50194, VAL.
20 Vaginulina alazanensis Nuttall, UCMP50200, PPP.

21 Vaginulina cf. V. spinata (Schubert), UCMP50201, CUC.
22 Vaginulina tenuis (J. G. Bornemann), UCMP50202, LBZ.
23 Lagena cf. L. alcocki White, UCMP50203, LEB.
24 Lagena alternans Terquem, UCMP50204, NAV5.
Kenneth L. Finger
Plate 10
417
Type age and locality: Late Oligocene, Cuba.
Vaginulinopsis costatus (Batsch 1791)
Plate 10, Figure 12
Stratigraphic range: Late Oligocene to Early Miocene.
Nautilus (Orthoceras) costatus Batsch 1791, p. 1, 4, pl. 1, fig. 1.
Remarks: This appears to be a dimorphic species, as some

specimens have an incomplete early spiral stage.
Type age and locality: Neither designated; probably Recent,

Adriatic coast of Italy.
Diagnostic species: The form without the early spiral stage

resembles Marginulina costata var. coarctata A. Silvestri 1896
(Early Pliocene, Italy), but the costae of that species are fewer in
number, and continuous across all sutures.
Distinguishing featuress: Slightly compressed test with 16

strong, continuous costae, but missing its later chambers.
Comparative species: Marginulina crepinae (= Marginulinopsis

crespinae Ludbrook 1966; Cretaceous, South Australia) has about
six chambers in a small spiral before uncoiling and its costae
are more parallel to the uniseries axis. Marginulina costata (=
Nautilus (Orthoceras) costatus Batsch 1791; probably Recent,
Italy) has fewer, but more prominent, costae that continue up to
the aperture, and they are indented across sutures. Marginulina
utsunomiyensis Uchio 1951 (Late Miocene, Japan) has slightly
more costae and a central aperture.
Occurrence: Navidad Fm. (MOS, PPP, PTA), Ranquil Fm.

(MS10).
Maximum relative abundance: Common (MS10)
Prismatomorphia Loeblich and Tappan 1986
Type species: Vaginulina tricarinata dOrbigny 1826.
Prismatomorphia tricarinata (dOrbigny 1826)
Occurrence: Lacui Fm. (PNH).
Vaginulinopsis lueneburgensis (Clodius 1922)
Plate 10, Figure 13
Cristellaria lneburgensis Clodius 1922, p. 118, pl. 1, fig. 6.
Type age and locality: Late Miocene, Germany.

Remarks: The single Chilean specimen differs from the type
figure of Vaginulinopsis lueneburgensis by having a fairly
straight, slightly less lobulate uniserial stage, and broader
peripheral spines.
Occurrence: Navidad Fm. (PPT, PTA), Ranquil Fm. (MS10),
Lacui Fm. (PNH).
Plate 10, Figure 19
Vaginulinopsis subelegans? (Parr 1950)
Vaginulina tricarinata DOrbigny 1826, p. 258; type-figure in

PARKER, JONES, and BRADY 1865, pl. 1, fig. 34 (after dOrbig
nys modle no. 4).
Vaginulina subelegans Parr 1950, p. 326, pl. 11, fig. 20.
Type age and locality: Recent, Adriatic Sea, depth not given.
Remarks: Represented by a single worn specimen with three
sides, each of which looks identical to an unornamented Plecto
frondicularia.
Vaginulinopsis A. Silvestri 1904a
Type species: Vaginulina soluta var. carinata A. Silvestri 1898.
Vaginulinopsis cf. V. chetae (Basov 1964)
Plate 10, Figure 14
Type age and locality: Recent, Southern Ocean (Indian Ocean

sector).
Occurrence: Navidad Fm. (MPUP), Santo Domingo Fm. (VAL),
Lacui Fm. (CHE, CUC, PCB, PNH).
Vaginulinopsis sp. A
Plate 10, Figure 15

Plate 10, Figure 11
Vaginulinopsis sp. B
Lenticulina (Marginulinopsis) chetae Basov 1964, p. 80, 81, pl. 1,

fig. 7.
Plate 10, Figure 16
Type age and locality: Late Jurassic, Siberia.
Remarks: This robust form is very similar to Basovs Jurassic

species.
Vaginulinopsis sp. C
Comparative species: Resembles Vaginulinopsis sp. A (Pl. 10.

Fig. 15) in lateral view but has an acute, not rounded, periphery.
Occurrence: Quiriquina Island, off Ranquil Fm. (MS10).
Occurrence: Lacui Fm. (PCB).

418
Plate 10, Figure 17

Planularia Defrance, in de Blainville 1824
Type species: Peneroplis auris Defrance, in de Blainville 1824.

Planularia cassis (Fichtel and Moll 1798)
Maximum relative abundance: Common (CHO).
Nautilus cassis Fichtel and Moll 1798, p. 95, figured vars. , ,

, ,
FAMILY LAGENIDAE Reuss 1862

Lagena Walker and Boys 1784; emend. A. Silvestri 1902
Type species: Serpula (Lagena) sulcata Walker and Jacob, in
Kanmacher 1798.
Plate 10, Figure 18

Remarks: Rgl and Hansen (1984) provide a thorough review
of Fichtel and Molls specimens as well as excellent images of
lectotypes that match well with the Chilean specimens.
Comparative species: Planularia kubinyi (= Cristellaria (Robu
lina) kubinyi von Hantken 1868; Oligocene, Hungary) has more
curved sutures and is devoid of tubercles.
Subfamily VAGINULININAE Reuss 1860
Vaginulina dOrbigny 1826; emend. Reuss 1862, 1874 and
Marie 1941
Type species: Nautilus legumen Linnaeus 1758.
Vaginulina alazanensis Nuttall 1932
Lagena cf. L. alcocki White 1956
Plate 10, Figure 23
Lagena alcocki WHITE 1956, p. 246, pl. 27, fig. 7.
Type age and locality: Pliocene, California.

Remarks: This single specimen closely resembles Lagena alcocki,
but the worn apertural area is not inflated, nor is it reticulate.
Occurrence: Ranquil Fm. (LEB).
Lagena alternans Terquem 1875
Plate 10, Figure 24
Lagena alternans Terquem 1875, p. 425, pl. 1, fig. 4.
Plate 10, Figure 20
Type age and locality: Recent, off northern France.
Vaginulina alazanensis Nuttall 1932, p. 17, pl. 1, figs. 11, 15.
Distinguishing features: Subspherical with numerous striae that

extend from base and alternate between those that reach neck
and those that stop about of the way.
Type age and locality: Early Oligocene, Mexico.

Occurrence: Navidad Fm. (CPUP, LBZ, PPP, PTA), Ranquil
Fm. (MS10, RQT), Santo Domingo Fm. (VAL), Lacui Fm.
(PCB, CUC).
Vaginulina cf. V. spinata (Schubert 1901)
Plate 10, Figure 21
Cristellaria cumulicostata var. spinata SChubert 1901, p. 24, pl. 1,

fig. 34.
Type age and locality: Early Oligocene, Italy.

Remarks: The Chilean form has a more slender and rectilinear shape
and its costae are not as extensive as those shown in the type figure of
Vaginulina spinata.
Occurrence: Navidad Fm. (RAP), Lacui Fm. (CUC).
Vaginulina tenuis (J. G. Bornemann 1855)
Plate 10, Figure 22
Marginulina tenuis J. G. Bornemann 1855, p. 25, pl. 13, fig. 14.

Vaginulinopsis tenuis (J. G. Bornemann). JONES 1994, p. 78, pl. 66, figs. 2123.

Occurrence: Navidad Fm. (LBZ, MAT, PPP, PTA, RAP),
Ranquil Fm. (FRA, RAN), Lacui Fm. (CHO, CUC).
Comparative species: Neither the Chilean specimens or the

holotype have the neck intact so it is not possible to confirm
whether or not Lagena dorseyae McLean 1956 (Late Miocene,
Virginia) is synonymous.
Occurrence: Navidad Fm. (MPUP, NAV5), Ranquil Fm. (FRA,
FRM, MIB, RAN, RQK, RQT), Lacui Fm. (CUC).
Lagena bassensis Collins 1974
Plate 11, Figure 1
Lagena bassensis COLLINS 1974, p. 22, pl. 1, fig. 10.
Type age and locality: Recent, Australia.

Distinguishing features: Subfusiform with thick costae and
smooth, conical collar.
Comparative species: Lagena barkeri Hofker 1978 (Recent,
South Pacific) is finely porous and the basal edge of its collar is
scalloped. Lagena alcocki White 1956 (Pliocene, California) is
a similar except for having a reticulate neck instead of a smooth
collar.
Occurrence: Navidad Fm. (NAV5), Ranquil Fm. (LEB), Lacui
Fm. (CHO, PCB).
Lagena filicosta Reuss 1863a
Plate 11, Figure 2
Lagena filicosta REUSS 1863a, p. 328, pl. 4, figs, 50, 51.
419
Type age and locality: Not designated; levels given in Recent

and Pliocene.
Occurrence: Navidad Fm. (PPP, PPT, PTA).
Lagena perlucida (Montagu 1803)
Vermiculum perlucidum Montagu 1803, p. 525, pl. 14, fig. 3.
Type age and locality: Recent, England, depth not given.

Diagnostic features: The test is widest about of the way from
base to neck, with 814 bladed costae and a long neck with
costae coiled at 45 (Pl. 11, Fig. 4).
Lagena semistriata Williamson 1848
Plate 11, Figure 5
Lagena striata var. semistriata var. Williamson 1848, p. 14, pl.
1, figs. 9, 10.
Lagena semistriata Williamson. JONES 1994, p. 64, pl. 57, figs. 14,
16.

Diagnostic features: Flask-shaped with 24 striae of extending
1/3 to nearly the full distance from the aboral end to the base of
the neck.
Remarks: Lagena striatula (= Oolina striatula Egger 1857;
Miocene, Germany) is represented by type figures of three distinct
morphotypes; the specimen shown in his figures 5 and 6 are
similar to L. semistriata.
Comparative species: Lagena incerta (= Phialina incerta
Seguenza 1862, Tertiary, Sicily) is of similar form but its striae
extend about up from the base.
Occurrence: Ranquil Fm. (FRA), Lacui Fm. (CHO, PCB).
Lagena striata (dOrbigny 1839c)
PLATE 11
1 Lagena bassensis Collins, UCMP50205, NAV5.
20 Procerolagena? sp., UCMP50224, PTA.
2 Lagena filicostata Reuss, UCMP50206, PPP.
21 P ygmaeoseistron asperoides (Galloway and Morrey),

UCMP50225, FRA.
3, 4 L
agena perlucida (Montagu), LEB: 3, UCMP50207. 4,
UCMP50208.
5 Lagena semistriata Williamson, UCMP50209, CHO.
6, 7 L agena striata (dOrbigny): 6, UCMP50210, RAN. 7,
UCMP50211, FRA.
8 Lagena cf. L. striata (dOrbigny), UCMP50212, MAT.
9 Lagena striatula (Egger), UCMP50213, FRA.
10 Lagena substriata Williamson, UCMP50214, RQS.
11 Lagena sulcata Walker and Jacob, UCMP50215, FRA.
12 Lagena vilardeboana (dOrbigny), UCMP50216, PTA.
13 Lagena sp. A, UCMP50217, PTA.
14 Lagena sp. B, UCMP50218, PCB.
23 Pygmaeoseistron globulohispidum (McLean), UCMP50227, FRA.

24 Pygmaeoseistron hispidum (Reuss), UCMP50228, PPP.
25 P
ygmaeoseistron parvuliporum (Bandy), UCMP50229,
FRA.
26 Reussoolina cf. R. apiculata (Reuss), UCMP50230, PPP.
27 Allanhancockia? sp., UCMP50231, RAN.
28 Globulina pirula Egger, UCMP50232, PNH.
29 Polymorphina fistulosa Williamson, UCMP50233, RAN.
30 Polymorphina sp., UCMP50234, RAN.
15 Lagena sp. C, UCMP50219, PTA.
31 P
seudopolymorphina atlantica Cushman and Ozawa,
UCMP50235, PPN.
16 Lagena sp. D, UCMP50220, RAN.
32 Pseudopolymorphina sp. A, UCMP50236, NLP.
17 P
rocerolagena distomum (Parker and Jones), UCMP50221,
CHO.
33 Pseudopolymorphina sp. B, UCMP50237, MAT.
18 Procerolagena ingens (Buchner), UCMP50222, FRA.

19 P
rocerolagena multilatera (McCulloch), UCMP50223,
RAN.
420
22 Pygmaeoseistron gibberum (Buchner), UCMP50226, FRA.
34 S igmomorphina trinitatensis Cushman and Ozawa,

UCMP50238, VAL.
Kenneth L. Finger
Plate 11
421
Oolina striata DORBIGNY 1839c, p. 21, pl. 5, fig. 12.

Lagena striata (dOrbigny). BOLTOVSKOY and THEYER 1970, p.
342, pl. 3, figs. 17, 23. INGLE, KELLER and KOLPACK 1980,
pl. 4, fig. 13. ZAPATA and CEAR 2004, p. 29, pl. 10, fig. 5.
Distinguishing features: Test ovate, with ~40 extensive striations;

neck reticulate in lower part, progressively coarsening until they
succeeded in upper part by a single stria that gently twists around
the next.
Type age and locality: Recent, Falkland Islands, depth not given.
Remarks: Neck ornamentation on Lagena can be difficult to

ascertain under a stereomicroscope, and most type descriptions
of Lagena species do not mention it. Some species appear to vary
in this repect, so until its taxonomic significance is demonstrated,
I refrain from questioning those species assignments where
neck ornamentation is apparent on the Chilean form, but not on
the type figure or in the type description of the species I have
ascribed it to.

Distinguishing features: Subsperical with 2030 well-developed
longitudinal costae, some slightly shorter than others; long neck has
reticulation that varies between specimens.
Occurrence: El Peral beds (NLP), Navidad Fm. (CPUP, NAV5, PPP,
PPT, PTA), Ranquil Fm. (FRA, FRM, MIB, RAN, RQK), Lacui
Fm. (CHO).
Lagena cf. L. striata (dOrbigny 1839c)
Plate 11, Figure 8
Oolina striata DORBIGNY 1839c, p. 21, pl. 5, fig. 12.
Type age and locality: Recent, Falkland Islands, depth not given
Distinguishing features: Globular with ~26 striations.
Remarks: The striae are much less pronounced than those on L.
striata (Pl. 11, Figs. 7, 8), but they are not nearly as faint as those of
Lagena sp. A (Pl. 11, Fig. 14). All recovered specimens are missing
the pper part of neck.
Comparative species: Lagena haidingeri (= Oolina haidingeri
Cjek 1848; Tertiary, Austria) has nearly twice as many striae.
Lagena ampliformis McCulloch 1977 (Recent, Galpagos Islands)
has three types of costae and a more pronounced collar.
Occurrence: Navidad Fm. (MAT, PPP), Ranquil Fm. (RAN, RQT).
Lagena striatula (Egger 1857)
Plate 11, Figure 9
Occurrence: Navidad Fm. (MAT, PPT), Ranquil Fm. (RAN,

RQK, RQT).
Lagena sulcata Walker and Jacob 1798
Plate 11, Figure 11
Serpula (Lagena) sulcata Walker and Jacob, in Kanmacher 1798,

p. 634, pl. 14, fig. 5.
Lagena sulcata Walker and Jacob. LOEBLICH and TAPPAN 1987, pl.
pl. 455, figs. 12, 13. JONES 1994, p. 64, pl. 57, figs. 23, 2527,
33, 34. HOLBOURN, HENDERSON and MACLEOD 2013, p.
324325.

Stratigraphic range: Early Cretaceous to Recent.
Distinguishing features: All costae extend at least to neck; those
that extend to aperture give the neck a triangular, rather than
tubular, outline in apertural view.
Comparative species: Lagena vilardeboana (see next entry)
is slightly more striate, as is L. isabelleana dOrbigny 1939
(Recent, Falkland Islands), which has a slightly less-rounded
aboral end. Those species might be variants of L. sulcata.
Oolina striatula Egger 1857, p. 269, pl. 5, figs. 38.
Occurrence: Navidad Fm. (CPUP), Ranquil Fm. (FRA, RAN,

RQT).
Type age and locality: Miocene, Germany.
Distinguishing features: Spherical, with about 14 striae extending

one-third to more than two-thirds up from the aboral end. Upper part
of neck missing on all recovered specimens.
Lagena vilardeboana (dOrbigny 1839c)
Remarks: The type figures of L. striatula show three distinct

morphotypes. The Chilean form matches Eggers first illustrated
specimen (his figs. 3 and 4).
Plate 11, Figure 12
Oolina vilardeboana DOrbigny 1839c, p. 19, pl. 5, figs. 4, 5.

Lagena vilardeboana (dOrbigny). BOLTOVSKOY and THEYER
1970, p. 343, pl. 3, fig. 8.
Distinguishing features: Somewhat flask-shaped, with long

costae of variable lengths, some extending onto elongate neck.
Lagena substriata Williamson 1848
Comparative species: See Lagena sulcata (Pl. 11, Fig. 11).
Plate 11, Figure 10
Lagena substriata Williamson 1848, p. 15, pl. 2, fig. 12. JONES

1994, pl. 57, fig. 19.
Type age and locality: Recent, Great Britain.
422
Occurrence: Navidad Fm. (PTA), Ranquil Fm. (RAN).

Lagena sp. A
Plate 11, Figure 13
Diagnostic features: Globular; ~20 faint striae; short, smooth

collar around base of neck.
Comparative species: Hyalinonetrion ingens (= Lagena distoma

var. ingens Buchner 1940; fossil, Italy) is more inflated.
Occurrence: Navidad Fm. (PPP, PTA), Lacui Fm. (PCB).
Occurrence: El Peral beds (NLP), Navidad Fm. (PPP, PTA),

Ranquil Fm. (FRA, RQK), Lacui Fm. (CHO, PCB).

Lagena sp. B
Maximum relative abundance: Rare
Plate 11, Figure 14
Procerolagena ingens (Buchner 1940)
Distinguishing features: Test widest 1/3 from base, upper 2/3

of chamber with nearly straight margins converging toward
aperture; ornamented with ~16 strongly bladed costae.
Lagena distoma var. ingens Buchner 1940, p. 425, pl. 2, fig. 22.
Remarks: Neck and aperture of single specimen missing, base

eroded.
Occurrence: Lacui Fm. (PCB)
Lagena sp. C
Plate 11, Figure 15

Distinguishing features: Lower half of test with 18 bladed costae.
Remarks: Single specimen recovered has neck broken near its
base, and eroded basal area.
Plate 11, Figure 18
Type age and locality: Fossil, Italy.

Remarks: See remarks for Hyalinonetrion distomum.
Occurrence: El Peral beds (NLP), Navidad Fm. (PPP, RAP),
Ranquil Fm. (FRA), Lacui Fm. (CHO, PCB).
Procerolagena multilatera (McCulloch 1977)
Plate 11, Figure 19
Lagena multilatera McCulloch 1977, p. 40, pl. 50, fig. 36.
Type age and locality: Recent, off Bikini, 480641m.
Comparative species: Lagena sp. B (Pl. 11, Fig. 15) has much
more prominent and extensive costae. The fusiform Lagena
sulcata (= Lagenulina sulcata Terquem 1876; Recent, France)
and L. florida Terquem 1882 (Eocene, France), and the spherical
L. exsculpta Brady 1881 (Recent, Indo-Pacific), have grooved,
not costate, lower halves.
Upper depth limit: Lower bathyal.
Occurrence: Ranquil Fm. (PTA).
Plate 11, Figure 20

Lagena sp. D
Plate 11, Figure 16

Distinguishing features: Subovate outline; smooth surface.
Remarks: Single specimen recovered is missing most of its neck.
It may be a worn Lagena semistriata (Pl. 11, Fig. 5).
Occurrence: Ranquil Fm. (RAN).
Procerolagena Puri 1954
Type species: Lagena gracilis Williamson 1848.
Procerolagena distomum (Parker and Jones 1864)
Plate 11, Figure 17
Lagena distoma Parker and Jones, in Brady 1864, p. 467, pl. 48,
fig. 6. INGLE, KELLER and KOLPACK 1980, pl. 4, fig. 12.
ZAPATA and CEAR 2004, p. 28, pl. 9, fig. 11.

Procerolagena? sp.
Distinguishing features: Slightly asymmetrical, narrow, and

fusiform with approximately 16 striae; long neck.
Remarks: This chamber is atypically inflated for the genus
Procerolagena. Whereas the aperture and base are eroded, the
two recovered specimens may be the ultimate chamber of a
uniserial species unrecognized in this study. If uniserial, generic
placement would be questionable.
Occurrence: Navidad Fm. (PPP, PTA).
Pygmaeoseistron Patterson and Richardson 1988
Type species: Lagena hispidula Cushman 1913b.
Pygmaeoseistron asperoides (Galloway and Morrey 1929)
Plate 11, Figure 21
Lagena asperoides Galloway and Morrey 1929, p. 16, pl. 16,

fig. 1. ROBERTSON 1998, p. 86, pl. 30, fig. 4.
Type age and locality: Probably Late Eocene, Ecuador.
Type age and locality: Not designated; lectotype from Recent,

coast of Norway.
Distinguishing features: Spherical with wide, lipped neck;

covered with coarse spinules.
Comparative species: Lagena aspera Reuss 1862 (Late Cre

taceous, The Netherlands) is coarsely tuberculate and its type
figure shows no indication of a neck.
Distinguishing features: Cylindrical with tapered ends; finely

striate.
423
Occurrence: Navidad Fm. (CPUP, NAV5), Ranquil Fm. (FRA,

MIB, RAN).
Pygmaeoseistron gibberum (Buchner 1940)
Plate 11, Figure 22
Lagena gibbera Buchner 1940, p. 423, pl. 3, figs. 4850. JONES

1994, p. 63, pl. 57, figs. 8, 9. ZAPATA 1999, fig. 18.
Type age and locality: Recent, Mediterranean, 50m and fossil,

Italy.
Upper depth limit: Neritic; based on type occurrence.
Diagnosis: Spherical with short apical protuberance; thick neck
ornamented with spinules.
Remarks: The recovered specimens appear weathered, but retain
the apical protuberance and some stubby remnants of the neck
spinules.
Pygmaeoseistron globulohispidum (McLean 1956)
elliptica Reuss 1863b and L. orbignyana var. elliptica Cushman

1923).
Lagena hispida (Reuss). JONES 1994, p. 63, pl. 57, fig. 1.
Type age and locality: Middle Eocene, Alabama

Upper depth limit: Lower bathyal; based on type occurrence of
Lagena hispidula Cushman 1913b (Recent, W Pacific, 3805m).
Distinguishing features: Ellipsoid; narrow neck; chamber and
neck hispid.
Comparative species: Pygmaeoseistron hispidula (= Lagena
hispidula Cushman 1913b) is flask-shaped, while P. hispidum
(Pl. 11, Fig. 24) is spherical.
REUSSOOLINA Colom 1956
Type species: Oolina apiculata Reuss 1851a.
Reussoolina cf. R. apiculata (Reuss 1851a)
Plate 11, Figure 26
Oolina apiculata REUSS 1851a, p. 22, pl. 2, fig. 1.
Plate 11, Figure 23
Type age and locality: Upper Cretaceous, Poland.
Lagena globulohispida McLean 1956, p. 331, pl. 39, fig. 7.
Remarks: This form has the deep radial grooves around the
aperture, but lacks the apical protrusion of R. apiculata.
Type age and locality: Late Miocene, Maryland.

Upper depth limit: Neritic (this species identified as Lagena
hispida in Zapata and Cear 2004).
Comparative species: Pygmaeoseistron globulohispidum is
intermediate between P. asperoides (Pl. 11, Fig. 19) and P. parvulipora (Pl. 11, Fig. 22) in shape and ornamentation.
Pygmaeoseistron hispidum (Reuss 1863a)
Plate 11, Figure 24
Lagena hispida REUSS 1863a, p. 335, pl. 6, figs. 7779.

Lagena aspera (Reuss). BOLTOVSKOY and THEYER 1970, p. 338,
pl. 3, fig. 15.

Distinguishing features: Spherical with long, narrow neck and
moderate-sized spinules.
Comparative species: Ornamentation is coarser than P.
parvuliporum (Pl. 11, Fig. 22), although not nearly as coarse as
P. asperoides (Pl. 11, Fig. 19).

Family POLYMORPHINIDAE dOrbigny 1839a
Subfamily POLYMORPHININAE dOrbigny 1839a
Allanhancockia McCulloch 1977
Type species: Allanhancockia lucenta McCulloch 1977.
Allanhancockia? sp.
Plate 11, Figure 27
Comparative species: This form looks identical to Allanhancockia

inculenta McCulloch 1977 (Recent, East Pacific), but that genus
is distinguished by an entosolenian tube, which is not apparent
in the Chilean specimens.
Occurrence: Navidad Fm. (MOS), Ranquil Fm. (RAN), Lacui
Fm. (CHE).
Globulina dOrbigny 1839a
Type species: Polymorphina (les Globulines) gibba dOrbigny
1826.
Globulina pirula Egger 1857
Plate 11, Figure 28
Polymorphina (Globulina) gibba var. pirula Egger 1857, p. 290, pl.

13, figs. 11, 12.
Pygmaeoseistron parvuliporum (Bandy 1949b)
Type age and locality: Miocene, Germany.
Plate 11, Figure 25
Lagena parvulipora Bandy 1949b, nom. subst. pro Lagena elliptica

BANDY 1949a, p. 55, pl. 7, figs. 16 (homonym of L. apiculata var.
424
Occurrence: Lacui Fm. (PNH).
Polymorphina dOrbigny 1826

Type species: Polymorphina burdigalensis dOrbigny 1826.
Pseudopolymorphina sp. B
Remarks: Numerous genera have been differentiated among

forms traditionally assigned to Polymorphina, but they are based
on internal chamber arrangements determined by dissection or
thin-sectioning, tasks that are impractical in most studies, so I
have retained the traditonally broad concept of Polymorphina
here.
Occurrence: Navidad (MAT), Lacui Fm.(PNH).
Polymorphina fistulosa Williamson 1858
Plate 11, Figure 29
Polymorphina lactea var. fistulosa Williamson 1858, p. 72, pl. 6, fig.

150.

Remarks: A single specimen was recovered in this study.
Polymorphina sp.
Plate 11, Figure 30

Distinguishing features: Slightly compressed, fusiform to slightly
pyriform shape; very slightly depressed sutures.
Remarks: All recovered tests are opaque and no single specimen
reveals the chamber arrangement clearly enough to enable
species recognition.
Occurrence: Ranquil Fm. (LEB, RAN).
Pseudopolymorphina Cushman and Ozawa 1928
Type species: Pseudopolymorphina hanzawai Cushman and
Ozawa 1928.
Pseudopolymorphina atlantica Cushman and Ozawa 1930
Plate 11, Figure 31
Pseudopolymorphina atlantica Cushman and Ozawa 1930, p. 94,

pl. 24, fig. 2.
Type age and locality: Recent, off eastern U.S. (1618050m).
Plate 11, Figure 33
Maximum relative abundance: Common (PNH).

Sigmomorphina Cushman and Ozawa 1928
Type species: Sigmomorphina (Sigmomorphina) yokoyamai
Cushman and Ozawa 1928.
Sigmomorphina trinitatensis Cushman and Ozawa 1930
Plate 11, Figure 34
Sigmomorphina trinitatensis Cushman and Ozawa 1930, p. 134,

pl. 36, figs. 1, 2.
Type age and locality: Eocene, Trinidad.

Comparative species: Sigmomorphina pernaeformis (= Polymorphina pernaeformis Stache 1865; late Tertiary, New
Zealand) has more chambers and its longitudinal suture is curved
and extends to the aperture. Sigmomorphina vaughani Cushman
and Ozawa 1930 (Eocene, South Carolina) has nearly the same
chamber arrangement, but is almost rhombic with depressed
sutures. Sigmomorphina problema (= Polymorphina (Guttulina)
problema dOrbigny 1826; fossil, Italy) is not fusiform and its
chambers are more inflated and arranged differently.
Occurrence: Navidad Fm. (MAT), Santo Domingo Fm. (VAL),
Lacui Fm. (CHE, PCB, PNH).
Maximum relative abundance: Common (MAT, PCB).
Subfamily RAMULININAE Brady 1884
Ramulina T.R. Jones, in J. Wright 1875; emend. Dakin 1906
Type species: Ramulina laevis T.R. Jones, in J. Wright 1875.
Ramulina globulifera Brady
Plate 12, Figure 1
Ramulina globulifera Brady 1879b, p. 272, pl. 8, figs. 32, 33.

KOHL 1985, p. 54, pl. 14, fig. 1. JONES 1994, p. 88, pl. 77, figs.
2228. ROBERTSON 1998, p. 98, pl. 36, figs. 3, 4. HAY
13, fig. 9.
Type age and locality: Recent, type locality not given; localities
in North Atlantic or South Pacific, 2651097m.
Comparative species: Polymorphina incavata Stache 1865

(Early Oligocene, New Zealand) is subrectangular in apertural
view and its chambers are more consistent in shape. Guttulina
yabei Cushman and Ozawa 1929 (Late Pliocene, Japan) is less
flaring and more rounded in cross-section.
Upper depth limit: Neritic; based on occurrence off New Zealand

(Hayward et al. 2010).
Occurrence: Navidad Fm. (LBZ, MAT, MPUP, PPN, PPP,

RAP), Ranquil Fm. (LEB, RAN, RQT), Lacui Fm. (PCB, PNH).
Occurrence: Navidad Fm. (PPP, PTA).
Maximum relative abundance: Common (PPN).

Pseudopolymorphina sp. A
Plate 11, Figure 32
Occurrence: El Peral beds (NLP), Navidad Fm. (PPP), Ranquil

Fm. (MIB).
Distinguishing features: Spherical chamber with many tubular

branches.
Ramulina pulchra Bermdez 1949
Plate 12, Figure 2
Ramulina globulifera var. pulchra Bermdez 1949, p. 164, pl. 11, fig.
12.
Type age and locality: Middle Miocene, Dominican Republic.

425
Distinguishing features: Subglobular chamber with numerous

long, robust tubercles.
Comparative species: Ramulina globulifera Brady 1879b
(Recent, N Atlantic and S Pacific) has a more globose chamber
with more tubercles.
Occurrence: Navidad Fm. (PPP, PPT), Ranquil Fm. (FRA, RAN).
Family ELLIPSOLAGENIDAE A. Silvestri 1923
Subfamily OOLININAE Loeblich and Tappan 1961
Favulina Patterson and Richardson 1988
Type species: Entosolenia squamosa var. hexagona Williamson
1848.
Favulina favosopunctata (Brady 1881)
Plate 12, Figure 3
Lagena favoso-punctata Brady 1881, p. 62; type-figures in Brady

1884, pl. 58, fig. 35; pl. 59, fig. 4; pl. 61, fig. 2.
Oolina favosopunctata (Brady). JONES 1994, p. 66, pl. 58, fig. 35.
Type age and locality: Recent, localities given in Indopacific

region, 930 and 102m.
Remarks: Brady figures three morphotypes that might not
conspecific. The Chilean specimen matches Bradys specimen
from the Torres Strait, differing only in not having a multiringed neck, which is not seen on his other types.
Occurrence: Lacui Fm. (CHO).
Favulina hexagona (Williamson 1848)
Plate 12, Figure 4
Entosolenia squamosa var. hexagona Williamson 1848, p. 20, pl.

2, fig. 23.
Lagena hexagona (Williamson). HOWE and WALLACE 1932, p. 28,
pl. 6, fig. 14. CUSHMAN 1935, p. 23, pl. 9, fig. 10.
Oolina hexagona (Williamson). BOLTOVSKOY and THEYER 1970,
p. 350, pl. 4, fig. 15. JONES 1994, p. 66, pl. 58, fig. 33. ROB
ERTSON 1998, p. 100, pl. 37, fig. 2. ZAPATA and CEAR 2004,
p. 31, pl. 11, fig. 12. HOLBOURN, HENDERSON and MA
CLEOD 2013, p. 382383.
Favulina hexagona (Williamson). LOEBLICH and TAPPAN 1987, pl.
463, figs. 1, 2.
Type age and locality: Recent, United Kingdom; depth not given.
Upper depth limit: Neritic (Zapata and Cear 2004), but typically
bathyal (Holbour, Andrews and McLeod 2013).
Remarks: A few Favulina specimens were found with hexagonal
ornamentation, but they vary in shape from globular to lacrimate,
and by the coarseness and regularity of the ornamentation. The
form shown here may be a variant of F. hexagona or possibly
another species. Favulina species are very minor components
of assemblages, which makes it difficult to ascertain their
intraspecific variations.
Occurrence: Navidad Fm. (PPP, PTA), Santo Domingo Fm.
(VAL).
Favulina melo (dOrbigny 1839c)
Plate 12, Figure 5
Oolina melo DOrbigny 1839c, p. 20, pl. 5, fig. 9. BOLTOVSKOY

and THEYER 1970, p. 350, pl. 5, fig. 21. ROBERTSON 1998, p.
100, pl. 37, fig. 3.
Type age and locality: Recent, Falkland Islands, depth not

recorded.
PLATE 12
Figures 1, 10, and 13 are photomicrographs; all other images are SEMs. Scale bars in m.
1 Ramulina globulifera Brady, UCMP50239, PPP.

2 Ramulina pulchra Bermdez, UCMP50240, FRA.
3 Favulina favosopunctata (Brady), UCMP50241, CHO.
4 Favulina hexagona (Williamson), UCMP50242, PPP.
5 Favulina melo (dOrbigny), UCMP50243, LEB.
6 Favulina squamosa (Montagu), UCMP50244, FRA.
7 Homalohedra sp., UCMP50245, PCB.
Finger,
n.
sp.,
holotype
12 Fissurina cuculatta Silvestri, UCMP50250, FRA.

13, 14 Fissurina marginata Seguenza, UCMP50251, PPP.
15 Fissurina cf. F. marginata Seguenza, UCMP50252, MIB.
16 Fissurina cf. F. obvia Seguenza, UCMP50253, CPUP.
17 Fissurina sp., UCMP50254, PPP.
8 Lagnea cf. L. enderbiensis (Chapman), UCMP50246, PCB.
seudoolina cf. P. fissurinea R. W. Jones, UCMP50255,

18 P
VAL.
9 Lagnea sp., UCMP50247, RAN.
19 Pseudoolina sp., UCMP50256, PPP.
10 Oolina laevigata dOrbigny, UCMP50248, RQK.
426
11 F
issurina ambicarinata
UCMP50249, RQT.
20 Parafissurina inermis (Buchner), UCMP50257, FRA.
Kenneth L. Finger
Plate 12
427
Remarks: The type figure shows a specimen that lacks a neck

and has a rounded aboral end; therefore, its outline is more
subovate or sublacrimate than the Chilean form illustrated here.
It has been customary to assign unilocular forms with similar
ornamentation to this species.
Comparative species: Both Favulina meloformis (= Oolina
meloformis McCulloch 1977, southern California, Recent, 20m)
and F. melosquamosa (O. melosquamosa McCulloch 1977;
northern California, Recent, 234m) have high-arched costae that
define intracostal areas that are higher than wide.
Occurrence: Ranquil Fm. (LEB), Lacui Fm. (CHO).
Favulina squamosa (Montagu 1803)
Plate 12, Figure 6
Vermiculum squamosum MONTAGU 1803, p. 526, pl. 14, fig. 2.

Oolina squamosa (Montagu). JONES 1994, p. 66, pl. 58, figs. 2832.
Type age and locality: Recent, England.

Remarks: This species is ornamented with a densely reticulated
pattern of aligned, arched areolae that transition just above
midpoint into hexagons.
Comparative species: Favulina melo (= Oolina melo dOrbigny
1839c; Recent, Falkland Islands) has a reticulation of quad
rangular areolae. The areolae of Favulina hexagona (Pl. 12, Fig.
4) are all hexagonal and generally larger and more distinct.
Occurrence: Ranquil Fm. (FRA, RAN), Lacui Fm. (PCB).
HOMALOHEDRA Patterson and Richardson 1988
Type Species. Lagena guntheri Earland 1934.
Homalohedra sp.
Plate 12, Figure 7

Distinguishing features: Subspherical, tapering from below
midpoint to pronounced aperture; 14 prominent costae extend
from base to of the way to the aperture.
Lagnea Popescu 1983
Type species: Fissurina radiata Seguenza 1862.

Lagnea sp.
Plate 12, Figure 9

Distinguishing features; Lacrimate; no peripheral ridge.
Remarks: Represented by a single poorly preserved specimen.
Oolina dOrbigny 1839c
Type species: Oolina laevigata dOrbigny 1839c.
Oolina laevigata dOrbigny 1839c
Plate 12, Figure 10
Oolina laevigata DORBIGNY 1839c, p. 19, pl. 5, fig. 3. LOE

BLICH and TAPPAN 1987, pl. 463, figs. 8, 9.
Type age and locality: Recent, Falkland Islands (depth not

indicated).
Distinguishing features: Slightly lacrimate outline, circular
in apertural view, with small, blunt apical spine; aperture
nonradiate, with entosolenian tube.
Comparative species: Reussoolina apiculata (= Oolina apicu
lata Reuss 1851a; late Cretaceous, Poland) has a narrower outline
and lacks an entosolenian tube.
Occurrence: El Peral beds (NLP), Navidad Fm. (NAV5),
Ranquil Fm. (LEB, RQK, RQT), Santo Domingo (VAL).
Subfamily ELLIPSOLAGENINAE A. Silvestri 1923
Fissurina Reuss 1850
Type species: Fissurina laevigata Reuss 1850.
Fissurina ambicarinata Finger, n. sp.
Plate 12, Figure 11
Description: Test lacrimate in outline with circular chamber

and broad carina; moderately compresed; chamber moderately
convex, smooth; aperture a marginal slit in thickened, protruding
area that creates a discontinuity in the keel; entosolenian tube
extends along one face about the length of the chamber.
Lagena enderbiensis CHAPMAN 1909, p. 339, pl. 16, fig. 1.
Remarks: This Chilean form most closely resembles Lagena

lateralis forma carinata Buchner (1940; Recent, Italy), which
differs only by having a narrower carina; the name is unavailable
for elevation to the species-group, as Fissurina carinata Reuss
1863a has priority.
Type age and locality: Recent, southern New Zealand (155m).
Distinguishing features; Flat with fusiform outline and ridged

perimeter.
Maximum relative abundance: Rare (4 specimens).
Lagnea cf. L. enderbiensis (Chapman 1909)
Plate 12, Figure 8
Remarks: Represented by a single poorly preserved specimen

with its aperture obscured by secondary mineralization.
Comparative species: Lagnea selseyensis (= Lagena selseyensis
Heron-Allen and Earland 1909; fossil, England) has a slightly
inflated test.
428
Type specimens: Holotype UCMP50249, paratype UCMP50431

(from RQT).
Type age and locality: Early Miocene; Ranquil Formation
forming intertidal platform of Punta Huenteguapi (Ranquil Fm.,
locality RQT).
Etymology: Morphodescriptor derived from the Latin ambi

(circumference) + carinata (carinate).
Fissurina cuculatta A. Silvestri 1902
Plate 12, Figure 12
Fissurina cuculatta A. Silvestri 1902, p. 146, figs. 2325.
Type age and locality: Recent, Tyrrenian Sea.

Comparative species: Fissurina palliolata Earland 1934
(Recent, Falkland Islands) has a narrower profile.

PSEUDOOLina R. W. Jones 1984
Type species: Pseudoolina fissurinea R. W. Jones 1984.
Pseudoolina cf. P. fissurinea R. W. Jones 1984.
Plate 12, Figure 18
Pseudoolina fissurinea R. W. JONES 1984, p. 119, pl. 4, figs. 19, 20.
Comparative species: Pseudoolina fissurinea is less elongate

with a more pronounced apertural area and longer apertural
slit.
Fissurina marginata Seguenza 1862
Fissurina marginata Seguenza 1862, p. 66, pl. 2, figs. 27, 28.

LOEBLICH and TAPPAN 1987, p. 465, figs. 57.
Pseudoolina sp.
Plate 12, Figure 19
Type age and locality: Late Miocene, Sicily.
Distinguishing features: Ovate outline, circular in apertural

view; aperture a thickly lipped slit.
Occurrence: Navidad Fm. (PPP), Ranquil Fm. (RQK).
Remarks: The type figure is ovate and widest at midline, whereas

the Chilean form is widest below midline.

Fissurina cf. F. marginata Seguenza 1862
Subfamily ParafissurinINAE R.W. Jones 1984

Parafissurina Parr 1947
Type species: Lagena ventricosa A. Silvestri 1904b.
Parafissurina inermis (Buchner 1940)
Plate 12, Figure 20
Fissurina cf. F. marginata Seguenza 1862, p. 66, pl. 2, figs. 27, 28.
Lagena staphyllearia forma inermis Buchner 1940, p. 523, pl. 25,

fig. 522.
Type age and locality: Recent, Mediterranean.
Remarks: Differs from Fissurina marginata by being more

lacrimate and compressed, and the worn carina appears to be
ridged.
Comparative species: Fissurina marginata (Pl. 12, Figs. 13, 14)

appears to be a Parafissurina of similar outline, but possesses
a double margin. The aperture of P. sublata Parr 1950 (Recent,
off Tasmania) is a short, straight slit in a slight protrusion of the
test margin.

Fissurina cf. F. obvia Seguenza 1862
Plate 12, Figure 16
Fissurina (Fissurine) obvia Seguenza 1862, p. 60, pl. 2, fig. 1.

Distinguishing features: Lacrimate outline, inflated fusiform in oral
view, carinate, with apical protrusion; apertural area pronounced.
Occurrence: Navidad Fm. (CPUP).
Fissurina sp.
Plate 12, Figure 17

Comparative species: Similar to Fissurina ambicarinata (Pl.
12, Fig. 11), but with a narrower carina and a more pronounced
apertural region.

Family GLANDULINIDAE Reuss 1860
Subfamily GLANDULININAE Reuss 1860
Glandulina dOrbigny 1839a
Type species: Nodosaria (les Glandulines) laevigata dOrbigny
1826, subsequent designation by Cushman 1927f, p. 189.
Glandulina dentalinoides A. Silvestri 1903
Plate 13, Figure 1
Glandulina laevigata var. dentalinoides A. Silvestri 1903, p. 212,

text-fig. 3 (p. 212).
Type age and locality: Tertiary, Italy.

Distinguishing features: Spindle-shaped with blunt aboral end.
Occurrence: Navidad Fm. (MOS). Lacui Fm. (CHO).
429
Maximum relative abundance: Common (RQT).
Glandulina laevigata dOrbigny 1826
Glandulina simplex Hussey 1949
Nodosaria (Glanduline) laevigata DOrbigny 1826, p. 252, pl. 10,

figs. 13.
Glandulina ovula DORBIGNY 1846, p. 29, pl. 1, figs. 4, 5.
Glandulina laevigata dOrbigny. LOEBLICH and TAPPAN 1987, pl.
468, figs. 14. ROBERTSON 1998, p. 112, pl. 43, fig. 4.
Glandulina simplex Hussey 1949, p. 130, pl. 26, fig. 22.
Type age and locality: Fossil, Italy; Recent, Adriatic, depth not
recorded.
Remarks: The holotype shows the ultimate chamber comprising
about half of the test length, but this chamber overlap varies
considerably within populations, as does the degree of inflation.
Papp and Schmid (1985) noted the gradation between the two
species described by dOrbigny (1846), Glandulina laevigata
and G. ovula, and united them as G. ovula, even though G.
laevigata was numbered, described, and figured first. The very
short apical spine is characteristic of this form, but not always
present.
Comparative species: The stout form shown in Plate 13, Figure
3 matches Glandulina symmetrica Stache 1865 (late Tertiary,
New Zealand) (see illustration of topotype in Hornibrook 1971,
pl. 10, fig. 172). Glandulina hantkeni Franzenau 1894 (Neogene,
Hungary) also resembles this morphotype, but it has a short
apical spine. Type figures of G. simulans A. Silvestri 1903 (late
Tertiary, Italy) and G. haidingerina Neugeboren 1850 (Tertiary,
Romania) show more chamber overlap than most of the Chilean
specimens. Any of these morphotypes could be within the
interspecific variation of G. laevigata.
MOS, MPUP, NAV5, PPP, PPT, PTA, RAP), Ranquil Fm. (all
localities), Santo Domingo Fm. (VAL), Lacui Fm. (CHO, PCB,
PNH).
Plate 13, Figure 6
Type age and locality: Eocene, Louisiana.

Distinguishing features: Narrow fusiform shape with very rapid
increase in chamber size and flush sutures.
Comparative species: Glandulina elongata J. G. Bornemann
1855 (Oligocene, Germany) is more fusiform and slightly more
inflated.
Occurrence: Navidad Fm. (CPUP), Lacui Fm. (CHO).
Order ROBERTINIDA Mikhalevich 1980
Superfamily CERATOBULIMINOIDEA Cushman 1927c
Family CERATOBULIMINIDAE Cushman 1927c
Subfamily CERATOBULIMININAE Cushman 1927c
Ceratobulimina Toula 1915
Type species: Rotalina contraria Reuss 1851b.
Ceratobulimina jonesiana (Brady 1881)
Plate 13, Figure 8
Cassidulina jonesiana Brady 1881, p. 21: 59; type-figure in BRADY

1884, pl. 54, fig. 18.
Ceratobulimina alazanensis Cushman and Harris 1927 (Eocene,
Mexico). WHITTAKER 1988, p. 109, pl. 14, fig. 1416. ROB
ERTSON 1998, p. 114, pl. 44, fig. 1.
Ceratobulimina pacifica CUSHMAN and HARRIS 1927, p. 176, pl.
29, fig. 9. (Recent, Philippines, 903m)
Ceratobulimina jonesiana (Brady) JONES 1994, p. 60, pl. 54, fig.
18.
PLATE 13
Figures 1, 35, 9c, and 16 are photomicrographs; all other images are SEMs. Scale bars in m.
1 Glandulina dentalinoides Silvestri, UCMP50258, MOS.

landulina laevigata dOrbigny: 2, UCMP50259, FRA. 3,
25 G
UCMP50260, FRM. 4, UCMP50261, PPP. 5, UCMP50262,
NLP.
6 Glandulina simplex Hussey, UCMP50263, CHO.
7 Robertina subteres (Brady), UCMP50264, MS10.
8 Ceratobulimina jonesiana (Brady), UCMP50265, FRA.
9 Hoeglundina elegans (dOrbigny), UCMP50266, FRA.
430
10, 11 G
lobigerina concinna Reuss: 10, UCMP50267, PPT. 11,
juvenile, UCMP50268, PPP.
12 Globigerina praebulloides Blow, UCMP50269, FRA.
13 Globigerina venezuelana Hedberg, UCMP50270, FRA.
14 Globorotalia miotumida Walters, UCMP50271, NLP.
15 Globorotalia cf. Glr. miozea Walters, UCMP50272, PTA.
16 G
loborotalia praemenardii Cushman and Stainforth,
UCMP50273, NLP.
Kenneth L. Finger
Plate 13
431
Type age and locality: Recent, Indonesia (1061m).
Order GLOBIGERINIDA Carpenter, Parker and Jones 1862

Superfamily GLOBOROTALIOIDEA Cushman 1927c
Family GLOBOROTALIIDAE Cushman 1927c
Globorotalia Cushman 1927c
Type species: Pulvinulina menardii var. tumida Brady 1877.
Comparative species: Ceratobulimina haueri (= Rotalina

hauerii dOrbigny 1846; Miocene, Austria) is more compressed,
subacute in edge view, and has twice as many chambers visible.
Occurrence: El Peral beds (NLP), Navidad Fm. (CPUP, MOS,
PPP, PPT, PTA, RAP), Ranquil Fm. (all except LEB), Lacui Fm.
(CHE).
Family EPISTOMINIDAE Wedekind 1937
Subfamily EPISTOMININAE Wedekind 1937
Hoeglundina Brotzen 1948
Type species: Rotalia elegans dOrbigny, 1826.
Hoeglundina elegans (dOrbigny 1826)
Plate 13, Figure 9
Rotalia (Turbuline) elegans dOrbigny 1826, p. 276, type-figure not

given.
Hoeglundina elegans (dOrbigny). INGLE, KELLER and KOLPACK
1980, pl. 2, fig. 11. KOHL 1985, p. 59, pl. 14, figs. 4, 5. PAPP
and SCHMID 1985, pl. 49, figs. 16. VAN MORKHOVEN,
BERGGREN and EDWARDS 1986, p. 97, pl. 29, figs. 1a, b, 2a,
b. WHITTAKER 1988, p. 110, pl. 14, figs. 1719. JONES
1994, p. 104, pl. 105, figs. 36. ROBERTSON 1998, p. 114, pl. 44,
fig. 2. ZAPATA and CEAR 2004, p. 28, pl. 9, fig. 57. HAY
16, figs. 1621. HOLBOURN, HENDERSON and MACLEOD
2013, p. 298299.
Type age and locality: Not given.

Diagnostic feature: Number of chambers in final whorl is 610.
Occurrence: El Peral beds (LPER, NLP), Ranquil Fm. (MS10),
Navidad Fm. (CPUP, MAT, MOS, MPUP, MS10, NAV5, PPN,
PPP, PTA), Ranquil Fm. (all), Santo Domingo Fm. (VAL).
Superfamily ROBERTINOIDEA Reuss 1850
Family ROBERTINIDAE Reuss 1850
Subfamily ROBERTININAE Reuss 1850
Robertina dOrbigny 1846; emend. Hglund 1947
Type species: Robertina arctica dOrbigny 1846.
Robertina subteres (Brady 1881)
Plate 13, Figure 7
Bulimina subteres Brady 1881, p. 55; type-figure in Brady 1884,

pl. 50, figs. 17, 18.
Robertina bradyi Cushman and Parker 1936, p. 99, pl. 16, fig. 9.
(Recent, Caribbean Sea, 699m).
Type age and locality: Recent, Caribbean (713m) and off Fiji
Islands (1116m).
Comparative species: Robertina arctica dOrbigny 1846 (Middle
Pliocene, Austria) emend. Hglund 1947 has broader and more
numerous chambers.
432
Globorotalia miotumida Jenkins 1960
Plate 13, Figure 14
Globorotalia menardii miotumida Jenkins 1960, p. 362, pl. 4, figs.

9ac.
Globorotalia miotumida Jenkins. Jenkins 1985, p. 278, figs. 7.12,
7.13 SCOTT, BISHOP and BURT 1990, p. 39, figs. 2628.
Stratigraphic range: Middle Miocene to Early Pliocene (Jenkins

1985); base of N9N18/19.
Remarks: Kennett and Srinivasan (1983) note that Globorotalia
conoidea Walters 1965 may be a thick-walled form of this
species.
Occurrence: El Peral beds (NLP, LPER).
Globorotalia cf. Glr. miozea Finlay 1939
Plate 13, Figure 15
Globorotalia miozea Finlay 1939, p. 326, pl. 29, figs. 159161.

Stratigraphic range: Globorotalia miozea ranges from late Early
through Middle Pliocene (N6N13).
Remarks: Of the four specimens found in this study, three have
four chambers in the last whorl, whereas the one figured has five.
In axial view, these forms differ from Glr. miozea s.s. by having
a more rounded peripheral edge, nearly planar spiral side, and
broader penultimate chamber.
Occurrence: Navidad Fm. (PTA).
Globorotalia praemenardii Cushman and Stainforth 1945
Plate 13, Figure 16
Globorotalia praemenardii CUSHMAN and STAINFORTH 1945, p.

70, pl. 13, fig. 14. KENNETT and SRINIVASAN 1983, p. 122,
pl. 28, figs. 68. BOLLI and SAUNDERS 1985, fig. 32.7.
Stratigraphic range: Late Middle Miocene Zones N10N13.

Globorotalia praescitula Blow 1959
Plate 14, Figure
Globorotalia praescitula Blow 1959, p. 221, pl. 19. figs. 128ac.

KENNETT and SRINIVASAN 1983, p. 108, pl. 24, fig. 1; pl. 25,
figs. 46. SCOTT, BISHOP and BURT 1990, p. 27, figs. 19, 30.
CHAISSON and LECKIE 1993, p. 162, pl. 4, figs. 711.
Globorotalia scitula praescitula Blow. BOLLI and SAUNDERS 1985,
fig. 31.6.
Globorotalia praescitula Blow plexus. MAJEWSKI 2010, p. 4, figs.
3, 5.
Stratigraphic range: Late Early Miocene Zone N6 to Recent.

Occurrence: Navidad Fm. (PPP, PPT).
Remarks: In axial view, this form is more equally biconvex and
narrower than the other globorotalids observed in this study.
Paragloborotalia Cifelli 1982
Type species: Globorotalia opima opima Bolli 1957.

Remarks: The Paragloborotalia specimens recovered in this
study are ascribed to four plexuses that encompass the vast
majority of forms that slightly differ from the primary types or
the hypotypes shown in other publications. This morphologic
diversity can be attributed intraspecific variation, evolutionary
transition, and diagenetic alteration. It is my opinion that the
biostratigraphic range of each named species can be applied to its
plexus without introducing a significant error when determining
the relative age interval of an assemblage, especially when that
age is derived from a concurrent species range.
Paragloborotalia bella (Jenkins 1967) plexus
Plate 14, Figure 2
Globorotalia bella Jenkins 1967: 1069, fig. 3, nos. 16. KEN

NETT and SRINIVASAN 1983, p. 174, pl. 43, figs. 13.
Paragloborotalia bella (Jenkins). SCOTT, BISHOP and BURT 1990,
p. 124, figs. 81, 82.
Stratigraphic range: Early to late Middle Miocene, N4B to N8

(Kennett and Srinivasan 1983).
Remarks: Based on primarily on shape and numbers of chambers, I
originally split these populations into a Late Miocene association
comprised of Neogloboquadrina acostaensis (= Globorotalia
acostaensis Blow 1959), Ng. continuosa (= Globorotalia opima
Bolli subsp. continuosa Blow 1959), and Ng. pachyderma (=
Aristerospira pachyderma Ehrenberg 1861). Reports of Ng.
acostaensis at Punta Perro (Ibaraki 1992b) and Ng. pachyderma
in the vicinity of the Santo Domingo Fm. (Marchant and Pineda
1988; Marchant 1990) seemed to support this interpretation, and
larger populations (e.g., FRA) include forms resembling those
species.
Jenkins (1960) reported Globorotalia acostaensis in New
Zealand, but in 1967 he no longer recognized its presence in
that region, having realized that his specimens were older and
smaller, with radial sutures, less-inflated chambers, and lessdeveloped apertural lips. He thereupon described his late Middle
Pliocene 4.5- to 5-chambered form as Glr. mayeri nympha,
and also distinguished it from Glr. pachyderma, which has
four chambers and recurved sutures. In the same publication,
Jenkins described a similar form from the Early Miocene as
Glr. bella, which has a quinquelobate equatorial periphery. Its
spiral sutures also are more radial than curved, and in edge view
the ultimate chamber appears vertically compressed and wider
than those at the opposite edge. Kennett and Srinivasan (1983:
174) stated Morphological similarity between Gr. (J.) bella and
Gt. (J.) mayeri, especially with regard to the axially depressed
final chamber in peripheral view, suggests a close phylogenetic
relationship.
I now view the specimens I previously aligned with the
Ng. continuosa-acostaensis-pachyderma group as bettermatched with the Paragloborotalia bella-nympha group. The
distinction between this plexus and that of the Pg. nana-Ng.
continuosa and Pg. mayeri groups, however, are not clear.

In his description of Ng. continuosa, Blow noted that forms
transitional with Globorotalia opima nana Blow 1959 occur in
the Early Miocene. Bolli and Saunders (1982) considered Ng.
continuosa to be a four-chambered variant of Glr. mayeri, but
in their 1985 publication they differentiated them and gave the
Glr. opima nana-continuosa transitional forms a P22N6 range.
Bolli (1957) described Glr. opima nana as a 4- to 5-chambered
Oligocene species with radial sutures. Jenkins differentiated
Glr. bella from Glr. nana only by 5 vs. 4 chambers in the outer
whorl despite his remark that a few of his Glr. bella have 4.5
chambers, as does the holotype of Glr. nana. Obviously, neither
the number of chambers or the shape of the sutures can be relied
upon to separate the two species, which apparently overlap in
the Early Miocene. Regardless of whether they are assigned to
the Pg. nana-Ng. continuosa or Pg. bella-nympha lineage, the
morphologies represented by the Chilean populations indicate a
late Early Miocene (Burdigalian) age.
Comparative species: Paragloborotalia mayeri s.l. (see next
entry) are more robust with a high arched aperture, curved
sutures, and 45 chambers in the outer whorl.
Occurrence: Navidad Fm. (NAV5), Ranquil Fm. (FRA, RQT),
Lacui Fm. (CHO, PCB).
Paragloborotalia mayeri (Cushman and Ellisor 1939) plexus
Plate 14, Figure 3
Globorotalia mayeri Cushman and Ellisor 1939, p. 11, pl. 2, fig.

4. KENNETT and SRINIVASAN 1983, p. 174, pl. 43, figs. 46.
BOLLI and SAUNDERS 1985, p. 201, figs. 26.3126.43.
Paragloborotalia mayeri (Cushman and Ellisor). SCOTT, BISHOP
and BURT 1990, p. 127, figs. 83, 84. CHAISSON and LECKIE
1993, p. 164, pl. 8, figs. 1620.
Stratigraphic range: Late Oligocene Zone P22 (N4A) to late

Middle Miocene Zone N14 (Kennett and Srinivasan 1983).
Remarks: The holotype of Pr. mayeri has 6 chambers in the
outer whorl, whereas all of the Chilean specimens have 45.
Comparative species: Paragloborotalia bella by has curved
sutures, a lower aperture, and an ultimate chamber that is axially
compressed.
Occurrence: Navidad Fm. (PPN, MAT), Ranquil Fm. (MIB.
RAN, RQK, RQT).
Paragloborotalia nana (Bolli 1957) plexus
Plate 14, Figure 4
Globorotalia opima nana BOLLI 1957, p. 118, pl. 28, fig. 3.

Globorotalia nana (Bolli). KENNETT and SRINIVASAN 1983, p.
106, pl. 24, figs. 35.
Praegloborotalia nana (Bolli). CHAISSON and LECKIE 1993, p.
165, pl. 8, figs. 10, 11.
Stratigraphic range: Eocene to Middle Miocene Zone N12.

Remarks: The last appearance datum reported for this species
ranges from latest Oligocene Zone P22 in Bolli and Saunders
(1985) to Early Miocene Zone N5 in Srinivasan and Kenneth
(1983), and to Middle Miocene Zone N12 in Chaisson and Leckie
(1993). This may reflect the difficulty in differentiating the forms
433
in the Pg. bella-nympha and Pg. nana-Ng. continuosa lineages,

as well as in distinguishing the two lineages from each other.
Occurrence: Lacui Fm. (PCB)
Paragloborotalia zealandica (Hornibrook 1958) plexus
Globorotalia zealandica Hornibrook 1958, p. 667, figs. 18 19, 30.

KENNETT and SRINIVASAN 1983, p. 108, pl. 25, figs. 13.
Jenkins 1985, p. 279, fig. 7.6.
Paragloborotalia zealandica (Hornibrook). SCOTT, BISHOP and
BURT 1990, p. 115, fig. 76.
Globorotalia zealandica (Hornibrook) plexus. MAJEWSKI 2010, p. 4,
figs. 3, 4.
Stratigraphic range: Paragloborotalia zealandica ranges N5

N7 (Early Miocene).
Remarks: The specimens are mostly small, 4-chambered,
subplanoconvex tests that have an outer whorl comprising
four elongate-arcuate chambers, the earlier of which is often
encrusted, a bluntly rounded to subacute peripheral edge, and a
moderately to highly arched aperture with a moderately thick lip.
Comparative species: This species closely resembles
Globorotalia amuria Scott, Bishop and Burt 1990 (N9N15),
which Majewski (2010) included in his Glr. zealandica plexus.
Some populations (e.g., PTA) are more subconical in edge view
and include 5-chambered forms that approach the form of Glr.
miozea Walters.
Occurrence: Navidad Fm. (MOS, PPP, PPT, PTA), Ranquil Fm.
(RQK, RQT), Lacui Fm. (CUC).
Family CATAPSYDRACIDAE Bolli, Loeblich and Tappan
1957
Catapsydrax Bolli, Loeblich and Tappan 1957
Type species: Globigerina dissimilis Cushman and Bermdez
1937.
Catapsydrax dissimilis (Cushman and Bermdez 1937)
Globigerina dissimilis Cushman and Bermdez 1937, p. 25,

pl. 3, figs. 46.
Catapsydrax dissimilis (Cushman and Bermdez). KENNETT
and SRINIVASAN 1983, p. 22, pl. 2, figs. 38. BOLLI and
SAUNDERS 1985, p. 186, figs. 17.117.4. Spezzaferri and
Premoli-Silva 1992, pl. 1, fig. 1. LECKIE, FARNHAM and
SCHMIDT 1993, p. 123, pl. 3, figs. 16, 17.
Globigerinita dissimilis (Cushman and Bermdez). Brnnimann
and Resig 1971, p. 1303, pl. 25, figs. 7, 8.
Stratigraphic range: Middle Eocene P13 through Early Miocene

N6 (Kennett and Srinivasan 1983); in low latitudes, its last
occurrence is in lower N5 (Bolli and Saunders 1985).
Occurrence: Navidad Fm. (PPP), Ranquil Fm. (MS10), Ranquil
Fm. (MIB, RAN, RQK, RQT), Lacui Fm. (PCB).
Globoquadrina Finlay 1947
Type species: Globoquadrina dehiscens Chapman, Parr and Collins
1934.
Globoquadrina dehiscens (Chapman, Parr and Collins 1934)
Plate 14, Figure 10
Globorotalia dehiscens Chapman, Parr and Collins 1934, p. 569,

pl. 11, fig. 36.
Globoquadrina dehiscens (Chapman, Parr and Collins). KENNETT and
SRINIVASAN 1983, p. 184, pl. 45, figs. 79. BOLLI and SAUN
DERS 1985, figs. 15.415.7 CHAISSON and LECKIE 1993, p. 159,
pl. 9, figs. 1416. LECKIE, FARNHAM and SCHMIDT 1993, p.
124, pl. 8, figs. 17, 18.
Stratigraphic range: Early to Late Miocene. Kennett and

Srinivasan (1983) have it ranging from N4B through N18 and
note that its FAD and LAD are widely accepted as lower and
upper boundaries of the Miocene in most areas. Berggren et al.
(1995) indicate that its FAD is 5.8 Ma in the tropics and 6.8 Ma
in the subtropics, and its LAD is diachronous within N17 (Late
Miocene), Keller (1980) had placed its LAD in Southwest Pacific
DSDP Hole 590 in the middle of N16 at ~10 Ma.
Occurrence: Navidad Fm. (all localities except MPUP), Ranquil
Fm. (FRA, FRM, MIB, RAN, RQT), Lacui Fm. (CHE, CHO,
PCB).
PLATE 14
Figures 6ac are photomicrographs; all other images are SEMs. Scale bars in m.
globorotalia zealandica-Globorotalia miozea transitional,

UCMP50280, PTA.
1 Globorotalia praescitula Blow, UCMP50274, PPP.

2 Paragloborotalia bella (Jenkins), UCMP50275, FRA.
3 P
aragloborotalia mayeri (Cushman and Ellisor) s.l.,
UCMP50276, PPT.
4 Paragloborotalia nana (Bolli), UCMP50277, NAV5.
58 P
aragloborotalia zealandica (Hornibrook) plexus: 5, 6,
UCMP50278, PPP. 7, juvenile, UCMP50279, PPP. 8, Para-
434
9 G
loboquadrina praedehiscens Blow and Banner, UCMP50281, RAN.
10
Globoquadrina dehiscens (Chapman, Parr and Collins),

UCMP50282, FRA.
Kenneth L. Finger
Plate 14
435
Globoquadrina praedehiscens Blow and Banner 1962
Plate 14, Figure 9
Globoquadrina dehiscens praedehiscens Blow and Banner 1962,

p. 116, pl. 15, figs. QS.
Globoquadrina praedehiscens Blow and Banner. KENNETT and
DERS 1985, p. 304, fig. 5.11
Stratigraphic range: Latest Oligocene Zone P22 to Early

Miocene Zone N6.
Occurrence: Navidad Fm. (MOS, PPP), Ranquil Fm. (FRM,
RAN).
Superfamily GLOBIGERINOIDEA Carpenter, Parker and
Jones 1862
Family GLOBIGERINIDAE Carpenter, Parker and Jones 1862
Subfamily GLOBIGERININAE Carpenter, Parker and Jones
1862
Globigerina Delage and Hrouard 1896
Type species: Globigerina bulloides dOrbigny 1826.
Globigerina concinna Reuss 1850
Globigerina concinna REUSS 1850, p. 373, pl. 47, fig. 8.
Stratigraphic range: Upper N6N12, late Early Miocene to late

Middle Pliocene.
Comparative species: Globigerina concinna Reuss 1850 was
described from the Late Miocene as having five chambers per
whorl. A similar form was described from the lower Oligocene
as G. ciperoensis Bolli 1954 (P19N4B, possibly N5), but
it was distinguished by its smaller test with lower spire and a
less rapid increase in the size of successive chambers, and by
having only four chambers in its penultimate whorl. Globigerina
praebulloides pseudociperoensis Blow 1969 (N7N12) was later
named for another 5-chambered form occurring in the Middle
Pliocene, but Blow compared it only with G. ciperoensis, perhaps
because the type material of G. concinna was lost. It is likely that
these two species are synonymous.
Occurrence: Navidad Fm. (NAV5, PPP, PPT, PTA), Ranquil Fm.
(MIB), Lacui Fm. (PCB).
Globigerina praebulloides Blow 1959; emend. Blow and Banner
1962
Plate 13, Figure 12
Globigerina praebulloides BLOW 1959, p. 180, pl. 8, fig. 47; pl. 9, fig.
48. BLOW and BANNER 1962, p. 92, pl. 9., figs. O-Q. KEN
NETT and SRINIVASAN 1983, p. 38, pl. 6, figs. 13. BOLLI
and SAUNDERS 1985, p. 181, fig. 13.14. Spezzaferri and
Premoli-Silva 1992, pl. 6, figs. 3, 4, 6. CHAISSON and
LECKIE 1993, p. 156, pl. 1, figs. 17, 18. LECKIE, FARNHAM
and SCHMIDT 1993, pl. 9, figs. 13, 14.
Stratigraphic range: N7N12, Late Early to late Middle

Pliocene.
Comparative species: Globoturborotalita woodi (P21N21)
differs by its apertural lip and strongly pitted, cancellate texture
(see Pl. 15, Fig. 8)
436
Occurrence: Navidad Fm. (MOS, PPP, PTA, MAT, NAV5,

CPUP), Ranquil Fm. (all except MS10), Santo Domingo Fm.
(VAL), Lacui Fm. (CHE, CHO, PCB).
Globigerina venezuelana Hedberg 1937
Plate 13, Figure 13
Globigerina venezuelana Hedberg 1937, p. 681, pl. 92, fig. 72b.

Brnnimann and Resig 1971, p. 1302, pl. 5, fig. 7; text-figs. 13,
14. BOLLI and SAUNDERS 1985, p. 180, fig. 13.20.
Globoquadrina venezuelana (Hedberg). KENNETT and SRINI
VASAN 1983, p. 180, pl. 44, figs. 57.
Globigerina venezuelana Hedberg. Spezzaferri and Premo
li-Silva 1992, pl. 7, figs. 2, 4.
Stratigraphic range: Middle Eocene to Early Pliocene Zone N19.

Remarks: The species has often been assigned to Globoquadrina.
Occurrence: El Peral beds (NLP, LPER), Navidad Fm. (CPUP,
MAT, MOS, NAV5, PPN, PPP, PPT, PTA, RAP), Ranquil Fm. (FRA,
FRM, LEB, MIB, RAN, RQK, RQT), Lacui Fm. (CHE, PCB).
Globigerinella Cushman 1927c
Type species: Globigerina aequilateralis Brady 1879b.
Globigerinella obesa (Bolli 1957)
Globorotalia obesa Bolli 1957, p. 119, pl. 29, figs. 2, 3. Brn

nimann and Resig 1971, p. 1314, pl. 50, figs. 7, 8; text-fig. 19.
BOLLI and SAUNDERS 1985, fig. 26.44.
Globigerinella obesa (Bolli). CHAISSON and LECKIE 1993, p. 157, pl.
1, figs. 3, 4. LECKIE, FARNHAM and SCHMIDT 1993, p. 124, pl.
7, figs. 15, 16. KENNETT and SRINIVASAN 1983, p. 234, pl. 59,
figs. 35.
Stratigraphic range: Late Oligocene Zone P22 to Recent

(Kennett and Srinivasan 1982; for the lower latitudes, Bolli and
Saunders (1985) have the range from Early Miocene N5 through
Middle Miocene N15.
Remarks: Bolli and Saunders (1985) do not mention Globi
gerinella pseudobesa (= Turborotalia pseudobesa Salvatorini
1967), a closely related form that Kennett and Srinivasan (1983)
range from Middle Miocene Zone N13 to Early Pliocene Zone
N19. They note that Glla. pseudobesa is characterized by its
interiomarginal, umbilical-extraumbilical aperture, and they
differentiate it from Glla. obesa by the wide range of variation
in apertural position from extraumbilical to nearly umbilical as
in Globigerina, and that the aperture commonly extends onto
the spiral side of the test. It is apparent from these comments,
the disparate age datums, and the diagenetic distortion common
among the Chilean specimens, that Glla. obesa and Glla.
pseudobesa may not be significantly different enough to warrant
their distinction as separate species.
Occurrence: El Peral beds (NLP), Navidad Fm. (CPUP, MAT,
MOS, NAV5, PPN, PPP, PTA, RAP), Ranquil Fm. (MS10),
Ranquil Fm. (FRA, FRM, LEB, RAN, RQK, RQT), Lacui Fm.
(CHE, CHO, PCB).
Globigerinoides Cushman 1927c; emend. Blow 1979
Type species: Globigerina rubra dOrbigny 1839a.
Stratigraphic range: Late Oligocene to Early Miocene, Zones

P22 (N4A) and N5.
Jenkins 1964; Early Miocene, N4N7). I have reidentified very

large-apertured specimens that I had referred to Gt. apertura (=
Globigerina apertura Cushman 1918b; N16 FAD) as Gt. woodi
primarily because of the Early Miocene age indicated by the
planktic assemblages. It is interesting that the Chilean forms
occur much earlier than the Middle Miocene intergradation
of very large-apertured Gt. woodi with Gt. apertura noted by
Kennett and Srinivasan (1983). Chaproniere (1988) restudied the
type section of Gt. woodi and noted an apparent grade in overall
test morphology with G. bulloides.
Remarks: The supplementary aperture on the spiral side of

this species is quite small and not discernible on most of the
recovered specimens due to adhered matrix material.
Occurrence: El Peral beds (NLP), Navidad Fm. (all localities),

Ranquil Fm. (all localities except LBZ), Lacui Fm. (CHE, CHO,
CUC, PCB).
Occurrence: El Peral beds (NLP, LPER), Navidad Fm. (CPUP,

MAT, LBZ, MS10, PPN, PPP, PPT, PTA), Ranquil Fm. (MIB,
MS10, RQK).
Globigerinoides primordius Blow and Banner 1962
Plate 15, Figure 6
Globigerinoides quadrilobatus primordius BLOW and BANNER

1962, p. 115, pl. 9, figs. DdFf; p. 138, text-fig. 14: figs. DdFf.
Globigerinoides primordius Blow and Banner. KENNETT and
DERS 1985, p. 195, fig. 20.16.
Subfamily ORBULININAE von Schultze 1854

Orbulina dOrbigny 1839a
Type species: Orbulina universa dOrbigny 1839a.
Globigerinoides trilobus (Reuss 1850)
Orbulina universa dOrbigny 1839a
Globigerina triloba REUSS 1850, p. 374, pl. 447, fig. 11.

Globigerinoides trilobus (Reuss). KENNETT and SRINIVASAN
1983, p. 62, pl. 13, figs. 13.
Orbulina universa DORBIGNY 1839a, p. 106; v. 8, pl. 1, fig. 1.

KENNETT and SRINIVASAN 1983, p. 86, pl. 20, figs. 46.
JONES 1994, p. 90, pl. 82, figs. 13. BOLLI and SAUNDERS
1985, p. 201, figs. 23.1, 24.2.
Plate 15, Figure 7
Stratigraphic range: Early Miocene Zone N4 to Pleistocene

Zone N22.
Occurrence. Navidad Fm. (CPUP, MAT, MOS, NAV5, PPN, PPP,
PTA, RAP), Ranquil Fm. (MIB, RAN, RQK), Lacui Fm. (CHE,
UC, PCB).
Plate 15, Figure 12
Stratigraphic range: Early Middle Miocene Zone N9 to Recent

(Kennett and Srinivasan 1983).
Occurrence: El Peral beds (LPER)
Globoturborotalita Hofker 1976

Type species: Globigerina rubescens Hofker 1956.
Praeorbulina Olsson 1964

Type species: Globigerinoides glomerosa subsp. glomerosa
Blow 1956.
Globoturborotalita brazieri (Jenkins 1966)
Praeorbulina glomerosa Blow 1956, s.l.
Globigerina brazieri JENKINS 1966, p. 1098, fig. 6, nos. 4351.
Globigerinoides glomerosa glomerosa Blow 1956, p. 64. tf. 1, nos.

1519; tf. 2, nos. 1, 2. KENNETT and SRINIVASAN 1983, p.
82, pl. 18, figs. 58. BOLLI and SAUNDERS 1985, p. 199, figs.
23.4, 24.5.
Plate 15, Figure 8
Stratigraphic range: Latest Oligocene Zone P23 to Early

Miocene Zone N8.
Occurrence: Navidad Fm. (MAT, MOS, PPN, PPP, PPT, RAP),
Ranquil Fm. (MIB, RAN, RQT), Lacui Fm. (CHO).
Maximum relative abundance: Common
Globoturborotalita woodi (Jenkins 1960)
Globigerina woodi Jenkins 1960, p. 352, pl. 2, fig. 2. KENNETT

and SRINIVASAN 1983, p. 43, pl. 7, figs. 46. Jenkins 1985,
p. 275, fig. 6.21 CHAISSON and LECKIE 1993, pl. 1, figs. 17, 18.
Globoturborotalita woodi (Jenkins). MAJEWSKI 2010, p. 6, figs. 7, 8.
Stratigraphic range: Late Oligocene Globorotalia kugleri Zone

to Late Pliocene Glr. tosaensis Zone (Kennett and Srinivasan
1983).
Remarks: Chilean populations of this species are vary greatly in
aperture size. I could not consistently differentiate morphologies
suggestive of Gt. connecta (= Globigerina woodi connecta
Plate 15, Figure 13
Stratigraphic range: Late Early Miocene Zone N8 to Middle

Miocene Zone N9 (Kennett and Srinivasan 1983).
Remarks: The single specimen recovered shows the early spiral
slightly protruding from the final chamber, but the supplementary
apertures are not clearly defined; hence, subspecies identification
is not possible.
Order BULIMINIDA Fursenko 1958
Superfamily BOLIVINOIDEA Glaessner 1937
Family BOLIVINIDAE Glaessner 1937
Bolivina dOrbigny 1839c
Type species: Bolivina plicata dOrbigny 1839c.
Remarks: Most species of Bolivina have their upper depth limit
in the outer neritic or upper bathyal zone. Specimens of this
437
genus are surprisingly rare in the Early Miocene assemblages

from Chile.
Occurrence: El Peral beds (LPER, NLP), Navidad Fm. (PPP,

PPT).
Bolivina advena Cushman 1925
Plate 16, Figure 1
Bolivina advena Cushman 1925, p. 29, pl. 5, fig. 1.
Bolivina alazanensis Cushman 1926b
Plate 16, Figure 4
Bolivina alazanensis Cushman 1926b, p. 82, pl. 12, fig. 1.
Occurrence: Navidad Fm. (NAV5), Lacui Fm. (PCB).
Type age and locality: Oligocene, Mexico.
Remarks: The illustrated specimen closely resembles that of

Bolli, Beckmann and Saunders (1994, fig. 53.3).
Bolivina aenariensis Costa 1856; emend. Sgarrella 1992
Bolivina aenariensis Costa 1856, p. 297, pl. 15, fig. 1; emend.

Sgarrella 1992, p. 318, 320, 322, pl. 1, figs. 113, pl. 2, figs.
111.
Bolivina subaenariensis CUSHMAN 1922a , p. 46, pl. 7, fig. 6. (Re
cent, Atlantic); see Sgarrella (1992).
Type age and locality: Age not indicated, Italy; Sgarrellas

paralectotypes are from the Pleistocene of Italy.
Distinguishing features: Lanceolate test moderately curved,
limbate sutures and costae.
Remarks: The Chilean populations vary from unornamented to
having as many as 14 fine to moderate costae extending from the
proloculus to at least midpoint of the test, and in some specimens
nearly to almost reaching the last pair of chambers. None of the
recovered specimens have the apical spine seen on most of those
shown by Sgarrella (1992).
Comparative species: Bolivina multicostata (= B. aenariensis
var. multicostata Cushman 1918b; Miocene, Florida) has
nearly twice as many costae, which extend almost the full
length of the test. Bolivina imbricata Cushman 1925 (Miocene,
California) has distinctly limbate sutures and its costae extend
less than halfway toward the aperture. These two forms may be
synonymous with B. aenariensis.
Comparative species: The form recognized as Bolivina serrata

(= B. subadvena var. serrata Natland 1938; Recent, California)
by Whittaker (1988: pl. 13, figs. 13) in the Late Miocene
and Pliocene of Ecuador may be synonymous. Bolivina perca
Garrett 1938 (middle Tertiary, Texas) is medially flat, more
limbate and coarsely perforate. Bolivina barbata Phleger and
Parker 1951 (Recent, Gulf of Mexico) is narrower with strongly
curved sutures. Bolivina alazanensis is coarsely punctate with a
distinct keel.
Occurrence: Navidad Fm. (PPP), Ranquil Fm. (RQK).
Bolivina arta MacFadyen 1930
Plate 16, Figure 5
Bolivina arta MacFadyen 1930, p. 58, pl. 4, fig. 21.
Comparative species: Bolivina acerosa Cushman 1936b

(Miocene, Dominican Republic) is nearly identical to this
species, but it has very fine longitudinal striae on the lower half
of its test.
Type age and locality: Miocene, Sinai Peninsula.
Occurrence: Navidad Fm. (PPP, PTA), Ranquil Fm. (MS10).
PLATE 15
Figures 13c, d are photomicrographs; all other images are SEMs. Scale bars in m.
13 C
atapsydrax dissimilis (Cushman and Bermdez): 1,
UCMP50283, RQT. 2, UCMP50284, RQT. 3, UCMP50285,
MIB.
4, 5 G
lobigerinella obesa (Bolli): 4, UCMP50286, PPP. 5,
UCMP50287, FRA.
6 G
lobigerinoides primordius Blow and Banner, UCMP50288,
PPP.
7 Globigerinoides trilobus (Reuss), UCMP50289, PTA.
438
8 Globoturborotalita brazieri (Jenkins), UCMP50290, PPP.

9, 10 G
loboturborotalita woodi (Jenkins): 9, UCMP50291, FRA.
10, UCMP50292, PPP.
11 Radiolarian, UCMP50293, RQK.
12 Orbulina universa dOrbigny, UCMP50294, LPER.
13 Praeorbulina glomerosa (Blow), UCMP50295, NLP.
Kenneth L. Finger
Plate 15
439
Bolivina pukeuriensis Hornibrook 1961
Plate 16, Figure 6
Bolivina pukeuriensis HORNIBROOK 1961, p. 76, pl. 9, figs. 172, 173.
Bramlette 1951, p. 61, pl. 22, fig. 7.
Type age and locality: Late Miocene, California.
Occurrence: Concepcin (MS10), Navidad (PTA).
Comparative species: Similarly small species that are possible

synonyms are Cassidulinoides yamagaensis Asano and Murata
(in Murata 1961; Early Miocene, Japan) and C. mekranense
Haque 1970 (Miocene, Pakistan).
Maximum Relative Abundance: Few.
Occurrence: Navidad Fm. (PTA).
Bolivina tumida Cushman 1925
Type age and locality: Early Miocene, New Zealand.
Plate 16, Figure 7
Bolivina tumida Cushman 1925, p. 32, pl. 5, fig. 9.

Superfamily CASSIDULINOIDEA dOrbigny 1839a
Family CASSIDULINIDAE dOrbigny 1839a
Subfamily CASSIDULININAE dOrbigny 1839a
Cassidulinoides Cushman 1927c
Type species: Cassidulina parkeriana Brady 1881.
Cassidulinoides californiensis Bramlette, in Woodring and
Bramlette 1951
Plate 16, Figure 8
Cassidulinoides californiensis Bramlette, in Woodring and
Cassidulinoides porrectus (Heron-Allen and Earland 1932)
Cassidulina crassa var. porrecta HERON-ALLEN and EARLAND

1932, p. 358, pl. 9, figs. 3437.
Cassidulinoides porrectus (Heron-Allen and Earland). NOMURA,
1999,
Type age and locality: Recent, between Punta Arenas and East
Falkland Islands.
Globocassidulina Voloshinova 1960
Type species: Cassidulina globosa von Hantken 1876.

Remarks: Most Globocassidulina have their upper depth limits
in the upper middle bathyal zone (Ingle 1980).
PLATE 16
Figures 5b, 9, 10, 11b, 12b, 13c, 14a, 15b, and 25 are photomicrographs; all other images are SEMs. Scale bars in m.
1 Bolivina advena Cushman, UCMP50296, PCB.

2, 3 B
olivina aenariensis Costa: 2, UCMP50297, NLP. 3,
UCMP50298, PPT.
4 Bolivina alazanensis Cushman, UCMP50299, PPP.
5 B
olivina arta MacFadyen, opposite side views, UCMP50300, PPP.
6 Bolivina pukeuriensis Hornibrook, UCMP50301, MS10.
7 Bolivina tumida Cushman, UCMP50302, MIB.
14 G
lobocassidulina subglobosa (Brady), UCMP50309,
PCB.
15 P
lanocassidulina curvicamerata Voloshinova, UCMP50310, PCB.
16 Ehrenbergina fyfei Finlay, UCMP50311, PPP.
17 S tainforthia cf. S. complanata (Egger), UCMP50312,
MS10.
18 Rectuvigerina transversa (Cushman), UCMP50313, LEB.
19 Bulimina alazanensis Cushman, UCMP50314, CPUP.
8 C
assidulinoides californiensis Bramlette, UCMP50303,
PTA.
20 Bulimina spicata Phleger and Parker, UCMP50315, FRA.
9, 10 C
assidulinoides porrectus (Heron-Allen and Earland),
PCB: 9, UCMP50304. 10, UCMP50305.
21, 22 G
lobobulimina pacifica Cushman, PPP: 21, UCMP50316;
22, UCMP50317.
11 G
lobocassidulina chileensis Finger, n. sp., holotype UCMP50306, PNH.
23 Praeglobobulimina ovata (dOrbigny), UCMP50318,

RQK.
12 Paracassidulina lobatula (Cushman), UCMP50307, MAT.
24 P
raeglobobulimina
50319, MIB.
13 G
lobocassidulina quadrata (Cushman and Hughes),
UCMP50308, FRA.
440
ovula
(dOrbigny),
UCMP-
25 P
raeglobobulimina socialis (Bornemann): UCMP50320,
RQK.
Kenneth L. Finger
Plate 16
441
Globocassidulina chileensis Finger, n. sp.
Plate 16, Figure 11
Description: Test large, subcircular outline in side view,

compressed sublenticular to subovate in edge view; peripheral
edge subacute to subrounded, slightly lobulate; chambers
broadly elongate, four pairs in final whorl; slightly inflated;
umbilical region closed and not depressed; aperture a commashaped opening along peripheral edge of ultimate chamber and
bending almost perpendicular to the suture, with small cristate
tooth; surface smooth, polished, opaque; wall structure granular.
Occurrence: Lacui Fm. (CHE, PCB PNH).
Maximum relative abundance: Very abundant (PNH).
and UCMP50433 (all from PNH).
Type locality: A small sedimentary pocket of the Lacui
Formation in volcanic rocks near the pinguinera (penguin
station) at Punihuil (locality PNH).
Cassidulina subglobosa Brady 1881, p. 60; type-figure in BRADY

1884, pl. 54, fig. 17. BANDY 1949a, p. 140, pl. 26, figs. 7a, b.
Cassidulina globosa Hantken. CUSHMAN 1935, p. 49, pl. 20, figs.
12a, b. CUSHMAN 1948, p. 241, pl. 20, fig. 2.
Globocassidulina subglobosa (Brady). JONES 1983, p. 27, pl. 14,
fig. 12. INGLE, KELLER and KOLPACK 1980, pl. 1, figs. 14,
15. RESIG 1981, pl. 7, fig. 7. KOHL 1985, p. 88, pl. 30, fig. 2.
LOEBLICH and TAPPAN 1987, pl. 557, figs. 1823. JONES
1994, p. 60, pl. 54, fig. 17. WHITTAKER 1988, p. 107, pl. 14,
figs. 10, 11. HOLBOURN, HENDERSON and MACLEOD 2013,
p. 264265.
Type age and locality: Recent, off Brazil, 1143m.

Upper depth limit: Outer neritic (Ingle, Keller and Kolpack
1980).
Occurrence: Navidad Fm. (CPUP, MPUP, MOS, NAV5, PPP),
Ranquil Fm. (LEB, RAN), Santo Domingo Fm. (VAL), Lacui
Fm. (CHE, PCB, PNH).
Maximum relative abundance: Common (PNH).
Etymology. Named for the country from which it is described.
Paracassidulina Nomura 1983a

Type species: Cassidulina delicata Cushman 1927a.
Type age and locality: Early Miocene, Chile.
Paracassidulina lobatula (Kaiho 1984)
Comparative species: The subacute to subrounded edge and

slightly lobulate periphery of G. chiliensis resembles that
of Islandiella japonica (= Cassidulina japonica Asano and
Nakamura 1937; Recent, Japan), which has sutures that appear
radial in lateral view and an aperture and wall structure
characteristic of Islandiella (Nomura 1983a, b). Cassidulina
miocenica Voloshinova (in Voloshinova and Borovleva 1952;
Miocene, Sakhalin Island, Russia) has a rounded periphery
and aperture parallel to the margin of the ultimate chamber.
Cassidulina imamurai Tai 1959 (Miocene, Japan) is not as
compressed and its chambers are not as rectilinear. Cassidulina
brocha Poag 1966 (Miocene, Mississippi) has a more acute edge,
curved sutures, and an aperture parallel to the periphery. The type
figure of Cassidulina laevigata dOrbigny 1826 (description, age
and locality not given) displays an acute periphery and a more
lobulate test. Cassidulina miocenica Voloshinova and Borovleva
1952 (Miocene, Sakhalin Island, Russia) has the same lateral
appearance of chambers, but it is nonlobulate and the aperture
parallels the penultimate chamber margin.
Occurrence: Navidad Fm. (NAV5), Lacui Fm. (CHE, PCB
PNH).
Plate 16, Figure 12
Cassidulina lobatula Kaiho 1984, p. 126, pl. 10, figs. 4, 5.
Type age and locality: Late Eocene, Japan.

Upper depth limit: Possibly upper middle bathyal.
Remarks: Kaiho (1984) also reported this species from the
Oligocene. The Chilean occurrences indicate that this species
ranges into the Early Miocene.
Comparative species: Takayanagia delicata (= Cassidulina
delicata Cushman 1927a; Recent, Pacific off Panama, 783m)
has a more delicate and more compressed test with a much
wider aperture and a radial wall structure. Cassidulina minuta
Cushman 1933c (Recent, off Paumotu Islands, tropical Pacific,
1544m) is only moderately compressed, with bulbous chambers
and deeply depressed sutures.
Occurrence: Navidad Fm. (MAT, NAV5), Ranquil Fm. (RQK).
Maximum relative abundance: Very abundant.
PLANOCASSIDULINA Gudina 1966; emend. Nomura 1999.

Type species: Cassidulina norcrossi Cushman 1933c.
Globocassidulina quadrata (Cushman and Hughes 1925)
Planocassidulina curvicamerata (Voloshinova 1952)
Cassidulina subglobosa var. quadrata Cushman and Hughes

1925, p. 15, pl. 2, figs. 7. RESIG 1981, pl. 4, fig. 14.
Globocassidulina subglobosa (Brady) (horizontalis-type). WHIT
TAKER 1988, p. 107, pl. 14, figs. 8, 9.
Cassidulina curvicamerata Voloshinova 1952, p. 92, pl. 2, fig. 9.
Plate 16, Figure 13
Type age and locality: Pleistocene, California.

Occurrence: Ranquil Fm. (FRA, FRM, RQT).
Globocassidulina subglobosa (Brady 1881)
Plate 16, Figure 14

442
Plate 16, Figure 15
Type locality and age: Miocene, Sakhalin Island, Russia.

Occurrence: Chiloe (PCB).
Subfamily EHRENBERGININAE Cushman 1927c
Ehrenbergina Reuss 1850
Type species: Ehrenbergina serrata Reuss 1850.
Ehrenbergina fyfei Finlay 1939c, p. 323, pl. 28, figs. 119, 121, 122.
Remarks: Although designated as the type for the genus

Transversigerina by Mathews (1945), that nomen is considered
to be a subgenus of Rectuvigerina (van Morkhoven, Berggren
and Edwards 1986).
Comparative species: Ehrenbergina compressa Cushman

(1927a; Recent, Pacific off Panama, 783m) is of similar
dimensions but the edges of its chamber are bluntly angled.
Occurrence: Navidad Fm. (MAT), Ranquil Fm. (FRA, FRM,

LEB, RQK, RQT), Santo Domingo Fm. (VAL), Lacui Fm.
(PCB).
Type age and locality: Late Miocene, New Zealand.
Maximum relative abundance: Common (MAT).
Occurrence: Navidad Fm. (CPUP, MOS, NAV5, PPP, PPT,

PTA), Ranquil Fm. (MS10, RQT), Lacui Fm. (CHE, PCB).
Family BULIMINIDAE T. R. Jones, in J. Wright 1875

Bulimina dOrbigny 1826
Type species: Bulimina marginata dOrbigny 1826.\
Ehrenbergina fyfei Finlay 1939c
Plate 16, Figure 16

Superfamily TURRILINOIDEA Cushman 1927c
Family STAINFORTHIIDAE Reiss 1963
Stainforthia Hofker 1956
Type species: Virgulina concava Hglund 1947.
Stainforthia cf. S. complanata (Egger)
Plate 16, Figure 17
Virgulina schreibersiana var. complanata Egger 1893, p. 292, pl. 8,

figs. 91, 92.
Type age and locality: Recent, Coral Sea.

Remarks: Differs from the type figure of Stainforthia complanata
by having less depressed sutures, a less lobulate outline, and a
wider early half.
Comparative species: Cassidella pacifica Hofker 1951 (Recent,
Indonesia) could be synonymous, as its author implied, but
he did not indicate a toothplate or compare it with Stainforthia
complanata.
Order BULIMINIDA Fursenko 1958
Superfamily BULIMINOIDEA T. R. Jones, in J. Wright 1875
Family SIPHOGENERINOIDIDAE Saidova 1981.
Subfamily SIPHOGENERININAE Saidova 1981
Rectuvigerina Mathews 1945
Type species: Siphogenerina multicostata Cushman and Jarvis
1929.
Rectuvigerina transversa (Cushman 1918a)
Plate 16, Figure 18
Siphogenerina raphanus var. transversus Cushman 1918a, p. 64, pl.

22, fig. 8.
Transversigerina transversa (Cushman). LOEBLICH and TAPPAN
1987, pl. 570, figs. 15, 16. WHITTAKER 1988, p. 73, pl. 9, figs.
810.
Rectuvigerina transversa (Cushman). FINGER 1992, p. 82, pl. 21 , figs
16.
Type age and locality: Miocene, Panama.

Stratigraphic range: Latest Oligocene to Middle Miocene Zones
P22 (N4A) to N11 (van Morkhoven, Berggren and Edwards
1986: 172).
Upper depth limit: Upper bathyal (van Morkhoven, Berggren
and Edwards 1986: 172).
Bulimina alazanensis Cushman
Plate 16, Figure 19
Bulimina alazanensis Cushman 1927e, p. 161, pl. 25, fig. 4.

KOHL 1985, p. 66, pl. 20, fig. 2. ROBERTSON 1998, p. 148, pl.
56, fig. 5.
Bulimina rostrata Brady 1884 (Recent, type locality not designated).
MARTNEZ and PARADA 1968, pl. 1, figs. 1, 2. INGLE,
KELLER and KOLPACK 1980, pl. 9, fig. 2. WHITTAKER 1988,
pl. 7, figs. 8, 9. JONES 1994, p. 56, pl. 51, figs. 1415. HOL
Type age and locality: Late Oligocene, Mexico.

Upper depth limit: Upper middle bathyal (1061m) in Jones 1994;
lower middle bathyal in Ingle, Keller and Kolpack 1980).
Comparative species: Bradys type figure of Bulimina rostrata
shows a fusiform test that tapers at both ends and is finely
pointed at its base. Bulimina truncana von Gmbel 1868 (Late
Eocene; localities given in Germany) is noticeably stouter and
with shorter costae. Bulimina truncanella Finlay 1940 (Middle
Eocene, New Zealand) was found by its author in beds as young
as Late Miocene, but it has many more costae than B. alazanensis.
Bulimina fossa Cushman and Parker 1938 (Pliocene, California)
also has more costae, which are somewhat irregular.
Occurrence: Navidad Fm. (CPUP).
Bulimina spicata Phleger and Parker 1951
Plate 16, Figure 20
Bulimina spicata Phleger and Parker 1951, p. 16, pl. 7, figs. 25,
31. KOHL 1985, p. 66, pl. 20, fig. 5.
Bulimina mexicana var. striata Cushman. INGLE, KELLER and
KOLPACK 1980, pl. 2, fig 4. RESIG 1981, pl. 1, fig. 12.
Bulimina pagoda Cushman. WHITTAKER 1988, p. 54, pl. 7, fig. 10
(Pliocene, Ecuador)
Type age and locality: Recent, 228m, northern Gulf of Mexico.

Upper depth limit: Upper bathyal (for B. striata mexicana in
Ingle, Keller and Kolpack 1980, and B. costata in van Morkhoven,
Berggren and Edwards 1986: 11).
Distinguishing features: Moderate number of bladed costae,
which terminate as short, downward-projecting marginal spines.
Comparative species: Bulimina pagoda Cushman 1927g
(Recent, Pacific off Panama) has coarse spines projecting
outward at about 45 rather than bladed costae. Several Neogene
443
species are differentiated from B. spicata by the nature of

their costae: B. inflata Seguenza 1862 (Pleistocene, Italy) has
more-spinose costae; B. inflata var. mexicana Cushman 1922a
(Recent, southern Gulf of Mexico) has better-defined, more
numerous ridges; B. subacuminata Cushman and Stewart 1930
emend. Haller 1980 (Pliocene, northern California) has lessspinose costae that extend part way up the chambers of the last
whorl; and B. pagoda var. hebespinata Stewart and Stewart 1930
(Early Miocene, southern California) has indistinct, irregularly
arranged costae, and larger, blunt spines that extend outward.
Variations within populations suggest that some, if not all, of
these forms are conspecific.
Occurrence: El Peral beds (LPER, NLP), Navidad Fm. (all
except PPN and LBZ), Ranquil Fm. (all), Lacui Fm. (CHE)
Globobulimina Cushman 1927c; emend. Hglund 1947
Type species: Globobulimina pacifica Cushman 1927c.
Remarks: Most Globobulimina have their upper depth limits in
the upper middle bathyal zone (Ingle 1980).
Globobulimina pacifica Cushman 1927g
Globobulimina pacifica CUSHMAN 1927g, p. 67, pl. 14, fig. 12.

KOHL 1985, p. 67, pl. 21, fig. 1. ROBERTSON 1998, p. 150, pl.
57, fig. 1.
Bulimina galliheri Kleinpell 1938, p. 253, pl. 17, figs 2, 3. (Middle
Miocene, California)
Bulimina pacifica (Cushman). BOLTOVSKOY and THEYER 1971, p.
334, pl. 2, fig. 11.
Type age and locality: Recent, East Pacific.

Remarks: The last whorl covers nearly all (Fig. 22) or all (Fig.
23) of the previous chambers.
Occurrence: Navidad Fm. (CPUP, MAT, MPUP, MOS, NAV5,
PPP, PPT, PTA), Ranquil Fm. (FRM, MIB, MS10, RAN, RQT).
Lacui Fm. (CHO).
Praeglobobulimina Hofker 1951
Type species: Bulimina pyrula dOrbigny var. spinescens Brady
1884.
Praeglobobulimina ovata (dOrbigny 1846)
Plate 16, Figure 23
Bulimina ovata DOrbigny 1846, p. 85, pl. 11, figs. 13, 14.
Praeglobobulimina ovata (dOrbigny), emend. HAYNES 1954, p. 150,
text-figs. 912; pl. 35, figs. 2, 3. (Paleocene, England)]
Bulimina pyrula (dOrbigny). PAPP and SCHMID 1985, pl. 62, figs.
24.
Praeglobobulimina ovata (dOrbigny). KOHL 1985, p. 67, pl. 21, fig. 3.

Basin, Austria.
Occurrence: Ranquil Fm. (MIB), Lacui Fm. (PCB).
Stratigraphic range: Paleocene to Middle Miocene.
444
Praeglobobulimina ovula (dOrbigny)
Plate 16, Figure 24
Bulimina ovula DOrbigny 1839c, p. 51, pl. 1, figs. 10, 11. PAPP
and SCHMID 1985, RESIG 1981, pl. 1, fig. 10.
Globobulimina ovula (dOrbigny). BOLTOVSKOY and THEYER
1971, p. 334, pl. 2, fig. 11. INGLE, KELLER and KOLPACK
1980, pl. 2, figs. 5, 6.
Type age and locality: Recent, type locality not given; localities
recorded off Ecuador, Peru, and Chile.
Distinguishing features: Nearly parallel sides; depressed and
highly oblique sutures; small, pointed initial end; last whorl
overlapping most previous whorls.
Comparative species: Praeglobobulimina ovata (= Bulimina
ovata dOrbigny 1846; Middle Pliocene, Austria) is broadest
at midlength and has considerably less overlap of chambers.
Praeglobobulimina (= B. pseudotorta Cushman 1926d; Late
Miocene, central California) is broadest near its apertural end
and has less oblique sutures.
Occurrence: Ranquil Fm. (MIB, RQK).
Praeglobobulimina socialis (J. G. Bornemann 1855)
Plate 16, Figure 25
Bulimina socialis J. G. Bornemann 1855, p. 342, pl. 16, fig. 10.

Bulimina affinis (dOrbigny). INGLE, KELLER and KOLPACK 1980,
pl. 4, figs. 10, 11.

Distinguishing features: Widest at midlength, where the width
is about half the length; moderately to weakly depressed sutures.
Comparative species: Praeglobobulimina affinis (= Bulimina
affinis dOrbigny; Recent, Cuba, beach) is spindle-shaped
and tapers aborally to a point. Praeglobobulimina subaffinis
(= Bulimina subaffinis Cushman 1921; Recent, Philippines,
1013m) is fusiform with a sharp apical end and smooth outline,
apparently due to flush sutures.
NAV5, PPP), Ranquil Fm. (FRA, FRM, MIB, RAN, RQK).
Maximum relative abundance: Common (LPER, FRM).
Praeglobobulimina spinescens (Brady 1884)
Plate 17, Figure 1
Bulimina pyrula var. spinescens Brady 1884, p. 400, pl. 50, figs. 11,
12.
Praeglobobulimina spinescens (Brady). KOHL 1985, p. 67, pl. 21,
fig. 3. LOEBLICH and TAPPAN 1987, pl. 571, figs. 1316.
JONES 1994, p. 54, pl. 50, figs. 11, 12.
Type age and locality: Recent, Indo-Pacific.

Comparative species: Praeglobobulimina spinifera (= Bulimina
spinifera Cushman 1927g; Recent, Pacific Ocean) is probably
synonymous.
Occurrence: Navidad Fm. (NAV5, PTA).
Protoglobobulimina Hofker 1951

Type species: Bulimina pupoides dOrbigny 1846.
Family UVIGERINIDAE Haeckel 1894

Subfamily UVIGERININAE Haeckel 1894
Ciperozea Vella 1961
Type species: Siphogenerina ongleyi Finlay 1939c.
Protoglobobulimina pupoides (dOrbigny 1846)
Plate 17, Figure 2
Bulimina pupoides DOrbigny 1846, p. 185, pl. 11, figs. 11, 12.
Bulimina pyrula (dOrbigny). PAPP and SCHMID 19785, p. 69, pl. 62,
figs. 24.
Praeglobobulimina pupoides (dOrbigny). LOEBLICH and TAPPAN
1987, pl. 572, figs. 16. JONES 1994, p. 55, pl. 50, figs. 14, 15.
192, pl. 17, figs. 35, 36.

Basin, Austria.
Remarks: DOrbigny named three morphotypes from the Vienna
Basin: Bulimina pyrula, B. pupoides, and B. ovata that can be
differentiated by the degree to which the later chambers envelope
the earlier part of the test. On the basis of metric measurements
of dOrbignys specimens, Papp and Schmid (1985) suggested
they are conspecific and should be synonymized as B. pyrula,
although B. ovata would have priority. I retain those taxonomic
divisions here because the end members of that plexus look
drastically different and a definitive intergradation of that extent
is not evident in the Chilean fauna.
Comparative species: Protoglobobulimina pupula (= Bulimina
pupula Stache 1865; late Tertiary, New Zealand) might be
synonymous (see illustration of neotype in Hornibrook 1971, pl.
10, fig. 180).
Ciperozea basispinata (Cushman and Jarvis 1929)
Plate 17, Figure 4
Siphogenerina basispinata Cushman and Jarvis 1929, p. 23, pl. 3,

figs. 4, 5.
Rectuvigerina basispinata (Cushman and Jarvis). KOHL 1985, p. 69,
pl. 22, fig. 3.
Uvigerina basispinata (Cushman and Jarvis). WHITTAKER 1988, p.
64, pl. 8, figs. 12, 13.
Type age and locality: Late Middle Miocene, Trinidad.

Stratigraphic range: Late Oligocene Zone P21 to Middle
Miocene Zone N12.
Upper depth limit: Probably bathyal based on C. multicostata.
Remarks: Whittaker (1988) notes that Lamb and Miller (1984)
synonymized this species as Uvigerina gallowayi, which they
considered to be a gerontic stage, but they did not realize that
Siphogenerina basispinata was erected earlier in the same year.
In the Chilean Miocene, I found the form referred to U. gallowayi
at only one locality (PPP), while Ciperozoa basispinata occurs
at seven (including PPP). Although U. gallowayi and C.
basispinata may be synonymous, their typical disassociation in
my samples prompts me to retain them as separate species.
Occurrence: Navidad Fm. (MOS, PPP), Ranquil Fm. (all except
LEB), Lacui Fm. (PCB).
Occurrence: Navidad Fm. (PPP, PTA), Ranquil Fm. (FRA,

MIB, MS10, RQT) Lacui Fm. (PNH).
Maximum relative abundance: Common (RAN).
Plate 17, Figure 5
Family BULIMINELLIDAE Hofker 1951

EUBuliminella Revets 1993
Type species: Eubulimina elegans var. exilis Brady 1884.
Eubuliminella bassendorfensis (Cushman and Parker 1937)
Plate 17, Figure 3
Buliminella bassendorfensis Cushman and Parker 1937, p. 40,

pl. 4, fig. 13.
Buliminella curta Cushman. WHITTAKER 1988, p. 61, pl. 7, figs. 16,
17.
Eubuliminella bassendorfensis (Cushman and Parker). REVETS 1993,
p. 142, pl. 2, figs. 3, 4.
Type age and locality: Oligocene, Oregon.

Remarks: Although the rare specimens found in the Chilean
material are poorly preserved, they match the holotype.
Comparative species: Eubuliminella curta (= Buliminella curta
Cushman 1925; Middle Miocene, California) is a stout form
with about 4 whorls and 56 chambers per whorl. Eubuliminella
subfusiformis (= Buliminella subfusiformis Cushman 1925;
Middle Miocene, California) is an elongate form with a straight
axis and about 7 whorls and 34 chambers per whorl.
Occurrence: Ranquil Fm. (RQK).
Ciperozea multicostata (Cushman and Jarvis 1929)

Siphogenerina multicostata Cushman and Jarvis 1929, p. 14, pl.
3, fig. 6.
Rectuvigerina multicostata (Cushman and Jarvis). VAN MORK
HOVEN, BERGGREN and EDWARDS 1986, p. 115, pl. 36, figs.
14. WHITTAKER 1988, p. 72, pl. 9, figs. 12, 13. ROBERT
SON 1998, p. 144, pl. 55, figs. 6, 7.
Ciperozoa multicostata (Cushman and Jarvis). LOEBLICH and TAP
PAN 1987, pl. 569, figs. 28, 29.
Type age and locality: Probably Miocene, Trinidad.

Stratigraphic range: Late Oligocene Zone P21 to Early Pliocene
Zone N20.
Upper depth limit: Upper bathyal; based on van Morkhoven,
Berggren and Edwards (1986: 116), who refer to it as bathyal.
Remarks: The costae on most of the Chilean Miocene specimens
show a slight tendency toward spinosity.
Comparative species: The type figure of Cushman and Jarvis
shows a nonserrate form, whereas that of Ciperozoa mayi (=
Siphogenerina mayi Cushman and Parker 1931; Miocene,
California) displays serration at the base of its chambers.
Although S. mayi is not mentioned as a subjective synonym by
van Morkhoven, Berggren and Edwards 1986), their illustrations
and the type description suggest that these two forms are end
445
members of a morphological grade; hence, Ciperozea mayi may

be a synonym of C. multicostata.
Occurrence: Navidad Fm. (CPUP, NAV5).
Ciperozea ongleyi (Finlay 1939c)
Plate 17, Figure 6
Siphogenerina ongleyi Finlay 1939c, p. 111, pl. 13, figs. 42, 43.
Ciperozea ongleyi (Finlay). LOEBLICH and TAPPAN 1987, pl. 573,
fig. 3.
Type age and locality: Early Miocene, New Zealand.

Upper depth limit: Probably bathyal based on Ciperozoa multi
costata.
Distinguishing features: Multiserial stage 1/3 1/2 of test length;
low costae tend to be somewhat continuous across sutures.
Remarks: The type-figures of this species are of specimens with
more irregular and tapering outlines than those from Chilean.
The type-description states that the costae terminate at sutures
as short, blunt spines, but that is not readily apparent in the typefigures, and should not be considered a definitive feature of the
species.
Comparative species: Ciperozoa mayi (= Siphogenerina mayi
Cushman and Parker; Miocene, California) has slightly abrupt
lower chamber margins that impart a more serrate appearance.
Ciperozoa costostriata (= Siphogenerina costostriata Galloway
and Heminway 1941; Late OligoceneEarly Miocene, Puerto
Rico) has a less lobulate periphery, a minimal uniserial stage and
discontinuous, and finer ornamentation. Ciperozoa hubbardi
(= Siphogenerina hubbardi Galloway and Heminway 1941;
Late OligoceneEarly Miocene, Puerto Rico) has a much more
extensive uniserial stage and its costae are continuous across the
sutures. Ciperozoa stonei (= Siphogenerina stonei Bermdez
1949 (Middle Oligocene, Dominican Republic) has a short apical
spine and its costae twist near the aperture.
Occurrence: Navidad Fm. (PPP), Ranquil Fm. (RQK), Lacui
Fm. (PCB).
Maximum relative abundance: Common (RQK).
Neouvigerina Hofker 1951
Type species: Uvigerina asperula var. ampullacea Brady 1884
(designated by Thalmann 1952).
PLATE 17
Figures 2, 3, 25, and 32a are photomicrographs; all other images are SEMs. Scale bars in m.
1 P
raeglobobulimina spinescens (Brady), UCMP50321,
NAV5.
18 Fursenkoina vicksburgensis (Cushman), UCMP50338,

RQT.
2 Protoglobobulimina pupoides (dOrbigny), UCMP50322,

RQT.
19 E
llipsopolymorphina zuberi (Grzybowski), UCMP50339,
FRA.
3 E
ubuliminella bassendorfensis (Cushman and Parker),
UCMP50323, RQK.
20 Virgulinella pertusa (Reuss), UCMP50340, CHO.
4 C
iperozea basispinata (Cushman and Jarvis), UCMP50324, FRA.
5 Ciperozea multicostata (Cushman and Jarvis), UCMP50325, CPUP.
6 Ciperozea ongleyi (Finlay), UCMP50326, PPP.
7 Neouvigerina auberiana (dOrbigny), UCMP50327, MIB
8, 9 N
eouvigerina hispida (Schwager): 8, UCMP50328, MIB.
9, UCMP50329, PPP.
10 Neouvigerina gallowayi (Cushman), UCMP50330, PPP.
11 Neouvigerina schwageri (Brady), UCMP50331, NAV5.
12 U
vigerina hispidocostata Cushman and Todd, UCMP50332, LEB.
13, 14 U
vigerina kernensis Barbat and Estorff, PTA: 13,
UCMP50333; 14, UCMP50334.
15 Uvigerina peregrina Cushman, UCMP50335, RQK.
16, 17 T
rifarina angulosa (Williamson): 16, UCMP50336,
NAV5. 17, UCMP50337, LEB.
446
21 O
besopleurostomella brevis (Schwager), UCMP50341,
MIB.
22 Pleurostomella alternans Schwager, UCMP50342, FRA.
23, 24 Siphonodosaria insecta (Schwager): 23, UCMP50343,
RAN. 24, UCMP50344, MIB.
25 Siphonodosaria pomuligera (Stache), UCMP50345, PPT.
26 Strictocostella pupa (Karrer), UCMP50346, MOS.
27 S iphonodosaria sentifera
UCMP50347, LEB.
(Cushman
and
Parker),
28, 29 S iphonodosaria lepidula (Schwager): 28, UCMP50348,

PPP. 29, UCMP50349, FRA.
30, 31 T
oddostomella hochstetteri (Schwager): 30, microspher
ic, UCMP50350, FRA. 31, megalospheric, UCMP50351,
NLP.
32, 33 C
ancris auriculus (Fichtel and Moll): 32, UCMP50352,
NLP. 33, UCMP50353, PCB.
Kenneth L. Finger
Plate 17
447
Neouvigerina auberiana (dOrbigny 1839a)
Upper depth limit: Upper middle bathyal.
Uvigerina auberiana DORBIGNY 1839a, p. 106, pl. 2, figs. 23,

24. LE CALVEZ 1977, p. 126, fig. 1 INGLE, KELLER and
KOLPACK 1980, pl. 6, fig. 1. WHITTAKER 1988, p. 63, pl. 8,
fig. 14. JONES 1994, p. 86, pl. 75, figs. 69.
Siphouvigerina auberiana (dOrbigny). KOHL 1985, p. 70, pl. 22, figs.
7, 8; pl. 23, fig. 1.
Comparative species: Neouvigerina rugosa (= Uvigerina rugosa

dOrbigny; Tertiary, Italy) has less dense, slightly coarser, and
more blunt spinosity. Neovigerina rustica (= Uvigerina rustica
Cushman and Edwards 1938 (Oligocene?, Venezuela) also has
less dense spinosity, but its spines are even coarser and more
blunt than those of N. rugosa.
Plate 17, Figure 7

localities noted in Cuba, Jamaica, and Martinique.
Upper depth limit: Upper bathyal; based on Uvigerina probo
scidea in van Morkhoven, Berggren and Edwards (1986: 30),
refer to it as bathyal.
Occurrence: El Peral beds (LPER, NLP), Navidad Fm. (MIB,

MPUP, MOS, PPP, PPT, PTA), Ranquil Fm. (FRA, FRM, MS10,
RAN, RQK, RQT), Lacui Fm. (PCB)
Maximum relative abundance: Common (LPER, NLP).
Neouvigerina schwageri (Brady 1884)
Plate 17, Figure 11
Comparative species: Neouvigerina proboscidea (= Uvigerina

proboscidea Schwager 1866; Pliocene, Car Nicobar) may be
synonymous. Neouvigerina senticosa (= Uvigerina senticosa
Cushman 1927; Recent, Pacific, 2542m) has a densely spinulose
texture.
Uvigerina schwageri Brady 1884, p. 575, pl. 74, figs. 810.

WHITTAKER 1988, p. 71, pl. 8, fig. 11.
Occurrence: Navidad Fm. (MOS, MPUP, PPP, PTA), Ranquil

Fm. (MIB, RQK).
Distinguishing features: Costae poorly defined, partial, few in

number.

Neouvigerina gallowayi (Cushman 1929)
Plate 17, Figure 10
Uvigerina gallowayi CUSHMAN 1929, p. 94, pl. 13, figs. 33, 34.
Type age and locality: Middle Pliocene, Ecuador.

Stratigraphic range: Late Oligocene Zone P21 to Middle
Miocene Zone N12, based on possible synonymy with Ciperozoa
basispinata.
Upper depth limit: Probably bathyal based on possible synonymy
noted above.
Type age and locality: Recent, Fiji Islands, 384m.

Comparative species: Neouvigerina carapitana (= Uvigerina

carapitana Hedberg 1937; Early Miocene, Venezuela) has fewer,
if any, costae, which are faint.
Occurrence: Navidad Fm. (MPUP, NAV5), Lacui Fm. (CHE,
CHO).
Maximum relative abundance: Common (NAV5).
Uvigerina dOrbigny 1826
Type species: Uvigerina pygmaea dOrbigny 1826.
Uvigerina hispidocostata Cushman and Todd 1945
Plate 17, Figure 12
Uvigerina hispido-costata Cushman and Todd 1945, p. 51, pl. 7,

figs. 27, 31. ROBERTSON 1998, p. 156, pl. 58, figs. 4, 5.
Uvigerina peregrina var. dirupta Todd. Cushman and McCull
och 1948, p. 267, pl. 34, fig. 3. (Recent, off southern California,
896m)
Uvigerina peregrina Cushman. WHITTAKER 1988, pl. 8, fig. 1.
(Recent, West Atlantic)
Maximum abundance: Few.
Type age and locality: Miocene, Jamaica.
Neouvigerina hispida (Schwager 1866)
Stratigraphic range: Miocene
Distinguishing features: Costae widely spaced, nearly linear,

continuous.
Remarks: See remarks for Ciperozoa basispinata.
Uvigerina hispida Schwager 1866, p. 249, pl. 7, fig. 95.

MARTNEZ and PARADA 1968, pl. 1, figs. 5, 6. INGLE,
KELLER and KOLPACK 1980, pl. 8, fig. 8. RESIG 1981, fig.
4. KOHL 1985, p. 72, pl. 24, fig. 3. VAN MORKHOVEN,
BERGGREN and EDWARDS 1986, p. 62, pl. 20, figs. 14. ROB
ERTSON 1998, p. 154, pl. 58, fig. 3. HAYWARD, GRENFELL,
SABAA, NEIL and BUZAS 2010, p. 203, pl. 21, figs. 4, 5.
Uvigerina rugosa dOrbigny. WHITTAKER 1988, p. 70, pl. 8, figs. 19,
20.

Stratigraphic range: Late Oligocene Zone P21 through
Pleistocene Zone N23.
448
Distinguishing features: Costate until last few chambers, which

are hispid.
Comparative species: Uvigerina peregrina s.s is fully costate.
The last-formed chambers of U. subperegrina (Cushman and
Kleinpell 1934; Middle Miocene, California) and U. hannai
Kleinpell 1938 (Late Miocene, California) are relatively smooth.
Occurrence: Navidad Fm. (CPUP, NAV5), Ranquil Fm. (LEB,
MS10).
Uvigerina kernensis Barbat and Estorff 1933

Uvigerina kernensis Barbat and Estorff 1933, p. 172, pl. 23, fig.
13.
Uvigerina hispida Schwager gr. WHITTAKER 1988 , p. 71, pl. 8, fig.
17.
Type age and locality: Early Miocene, California.

Upper depth limit: Upper middle bathyal; based on the
Uvigerina peregrina dirupta group recorded off Chile by Bandy
and Rodolfo (1964).
Distinguishing features: Relatively small test interspersed fine
costae and spinules, the proportion of which varies between
specimens.
Occurrence: Navidad Fm. (PPP, PPT, PTA).
Uvigerina peregrina Cushman 1923
and Earland 1932; Recent, subantarctic) infers that it may be an

ecophenotype of T. angulosa.
Occurrence: Navidad Fm. (CPUP, NAV5), Ranquil Fm. (LEB),
Santo Domingo Fm. (VAL), Lacui Fm. (PCB).
Superfamily FURSENKOINOIDEA Loeblich and Tappan 1961
Family FURSENKOINIDAE Loeblich and Tappan 1961
Fursenkoina Loeblich and Tappan 1961
Type species: Virgulina squammosa dOrbigny 1826.
Fursenkoina vicksburgensis (Cushman 1936b)
Plate 17, Figure 18
Virginulina vicksburgensis Cushman 1936b, p. 48, pl. 7, fig. 6.
Type age and locality: Oligocene, Mississippi.
Plate 17, Figure 15
Uvigerina peregrina Cushman 1923, p. 166, pl. 42, figs. 710.

MARTNEZ and PARADA 1968, pl. 1, figs. 10, 11. WHITTAK
ER 1988, pl. 8, fig. 3. LOEBLICH and TAPPAN 1987, pl. 573,
figs. 2428.
Type age and locality: Recent, NW Atlantic, 2136m.

Upper depth limit: Upper bathyal; based on occurrence off
California reported by Ingle (1980).
Remarks: The neritic form identified as this species in Zapata
and Cear (2004) is another species devoid of costae on its
ultimate chamber.
Comparative species: Uvigerina bifurcata dOrbigny 1839c
(Recent, Falkland Islands), as the name implies, has bifurcating
costae. Uvigerina hollicki Thalmann 1957 (nom. nov. pro U.
peregrina var. bradyana Cushman (1923; Recent, off NE USA)
has a coarsely perforate test with nearly twice as many costae.
Occurrence: Ranquil Fm. (RQK), Lacui Fm. (CUC).
Family VIRGULINELLIDAE Loeblich and Tappan 1984

Virgulinella Cushman 1932a
Type species: Virgulina pertusa Reuss 1861.
Virgulinella pertusa (Reuss 1861)
Plate 17, Figure 20
Virgulina pertusa Reuss 1861, p. 362, pl. 2, fig. 16. LOEBLICH

and TAPPAN 1994, pl. 579, figs. 2022.
Virgulina miocenica CUSHMAN and PONTON 1931, p. 32, pl. 4, figs.
1416. (Middle Miocene, Florida)
Virgulinella pertusa (Reuss). FINGER 1992, pl. 23, figs. 37, 38.
Type age and locality: Pliocene, Belgium.

Stratigraphic range: Early Miocene to Middle Pliocene.
Upper depth limit: Neritic?
Trifarina Cushman 1923

Type species: Trifarina bradyi Cushman 1923.
Comparative species: Although the type description of Virgulinella miocenica (= Virgulina miocenica Cushman and
Ponton 1931; Miocene, Florida) distinguishes it as being much
more slender and tapering, as well as smaller than V. pertusa
Reuss of Europe, I view them as synonyms.
Trifarina angulosa (Williamson 1858)
Occurrence: Ranquil Fm. (RQK), Lacui Fm. (CHO).
Uvigerina angulosa Williamson 1858, p. 67, pl. 5, fig. 140.

Angulogerina angulosa (Williamson). BOLTOVSKOY and THEYER
1970, p. 300, pl. 1, fig. 3.
Trifarina angulosa (Williamson). JONES 1994, p. 86, pl. 74, figs. 19,
20. LOEBLICH and TAPPAN 1987, pl. 574, figs. 59. HAY
21, figs. 1215.
Superfamily PLEUROSTOMELLOIDEA Reuss 1860

Family ELLIPSOIDINIDAE A. Silvestri 1923
Ellipsopolymorphina A. Silvestri 1901
Type species: Ellipsopolymorphina fornasinii Galloway 1933,
nom. subst. pro Ellipsopolymorphina deformis (Fornasini) =
Dimorphina deformis (part) Fornasini 1890 (non Glandulina
deformis Costa 1856).
Ellipsopolymorphina zuberi (Grzybowski 1896)
Upper depth limit: Outer neritic (Ingle 1980).

Remarks: All recovered specimens are relatively small.
Comparative species: The type-description of Trifarina
pauperata (= Uvigerina angulosa var. pauperata Heron-Allen
Plate 17, Figure 19
Pleurostomella zuberi Grzybowski 1896, p. 291, pl. 9, figs. 32, 33.

Ellipsopleurostomella schlichti A. SILVESTRI 1904b , p. 5, text-figs.
1, 2.
Ellipsoglandulina labiata (Schwager). HAYWARD 2002, p. 301, pl.
2, fig. 15.
Ellipsopolymorphina zuberi (Grzybowski). HAYWARD, Kawaga
449

Thomas 2012, p. 208, pl. 29, figs. 17.
Type age and locality: Early Oligocene, Poland.

Stratigraphic range: Early Oligocene to Early Miocene.
Upper depth limit: Upper middle bathyal, 900m (Hayward et al.
2012).
Remarks: Hayward (2002) differentiated three genera that are
in contention here: Ellipsoglandulina (uniserial throughout;
semilunate aperture), Ellipsopleurostomella (initially biserial;
triseriate aperture with hood on one side) and Ellipsopolymorphina
(initially biserial; lunate aperture). The Chilean specimens match
the type figure of Ellipsopleurostomella schlichti (A. Silvestri
1904b), which Hayward (2002) referred to Ellipsopolymorphina,
but he did not include an image; however, in the same publication,
Hayward illustrated this form as Ellipsoglandulina labiata (=
Polymorphina labiata Schwager 1866). The type figure of that
species is a considerably less-inflated polymorphine-like form
with a much less-enveloping ultimate chamber. Adding to the
confusion is that the E. labiata neotype selected by Srinivasan
and Sharma (1980: 59, pl. 8, figs. 9, 10) is a robust, uniserial form
with transverse sutures that does not resemble either species type
figure. Haywards form is not quite biserial in the early stage, but
the sutures are oblique and its aperture can be described as lunate;
thus, it appears more likely to be a variant of E. schlichti, which
Hayward et al. (2012) synonymized with E. zuberi.
Pleurostomella alternans Schwager 1866
Plate 17, Figure 22
Pleurostomella alternans SCHWAGER 1866, p. 238, fig. 79; emend.

Galloway and Heminway 1931. ROBERTSON 1998, p. . pl. 64,
fig. 6. HAYWARD, Kawagata, Sabaa, Grenfell, Van
Kerckhoven, Johnson and Thomas 2012, p. 228, pl. 36,
figs. 1015.

Stratigraphic range: Late Cretaceousmiddle Pleistocene
(Hayward et al. 2012)
Upper depth limit: Lower bathyal off Chile (Bandy and Rodolfo
1964); upper bathyal (400m) elsewhere (Hayward et al. 2012).
Remarks: Finger (2007) identified the figured specimen as
Pleurostomella elliptica Galloway and Heminway 1941 (Late
Oligocene and Early Miocene, Puerto Rico), but Hayward et al.
(2012) reassigned it as P. alternans Schwager 1866 (Pliocene,
Car Nicobar).
Comparative species: Pleurostomella elliptica Galloway and
Heminway 1941 (Late Oligocene and Early Miocene, Puerto
Rico) is more compressed, P. tenuis Hantken 1883 (Eocene,
France) has more elongate chambers, and P. acuminata Cushman
1922a (Recent, Caribbean) has a semicircular, smaller aperture.
Occurrence: Ranquil Fm. (FRA, FRM, RAN, RQT).
Occurrence: Ranquil Fm. (FRA, FRM, RQT).
Family STILOSTOMELLIDAE Finlay 1947

Siphonodosaria A. Silvestri 1924
Type species: Nodosaria abyssorum Brady 1881.
Family Pleurostomellidae Reuss 1860

Obesopleurostomella Hayward, in Hayward et al.
2012
Type species: Pleurostomella bierigi Palmer and Bermdez
1936.
Obesopleurostomella brevis (Schwager)
Plate 17, Figure 21
Pleurostomella brevis Schwager 1866, p. 239, pl. 6, figs. 21.

JONES 1994, p. 56, pl. 51, fig. 20.
Obesopleurostomella brevis (Schwager). HAYWARD, Kawagata,
Thomas 2012, p. 221, pl. 33, figs. 38.

al. 2012).
Comparative species: Obesopleurostomella recens (= Pleurostomella rapa var. recens Dervieux 1899 (Late Tertiary, Italy) has
numerous early chambers that come to a sharp aboral tip, so the
test looks like a spinning top. Nevertheless, Hayward et al. 2012
synonymizes O. recens with O. brevis.
Occurrence: Ranquil Fm. (FRA, MIB, RQT).
Pleurostomella Reuss 1860
Type species: Pleurostomella subnodosa (Reuss 1860).
450
Remarks: Siphonodosaria differs from other uniserial genera by

the aperture having a crenulate lip and tooth. Although those
features are frequently eroded or obscured in fossil specimens,
most species can be recognized by other morphologic criteria
(i.e., test shape, chamber shapes, sutures, ornamentation).
Generic assignment in this study is based on those few specimens
that show the details of the aperture, and by overall morphologic
comparison with the species illustrated in Hayward et al. (2012).
The upper depth limit of Siphonodosaria generally is considered
to be middle bathyal.
Siphonodosaria insecta (Schwager 1866)
Nodosaria insecta SCHWAGER 1866, p. 224, pl. 5, figs. 53, 54.

Nodosaria (?) abyssorum Brady 1881, p. 63; type-figures in
BRADY 1884, pl. 63, figs. 8, 9.
Stilostomella abyssorum (Brady). JONES 1994, p. 74, pl. 63, figs. 8, 9;
suppl. pl. 2, figs. 8, 9.
Siphonodosaria insecta (Schwager). HAYWARD, Kawagata,
Thomas 2012, p. 171, pl. 17, figs. 2429.

Stratigraphic range: Oligocene to Recent.
Upper depth limit: Upper middle bathyal (see Remarks below).
Remarks: All of the Chilean specimens are short segments and
those with the initial chamber have a very short apical spine.
Hayward et al. (2012) consider Siphonodosaria abyssorum (=
Nodosaria abyssorum Brady 1881; Recent, S. Pacific, 3338m)
and S. pacifica (= Nodogenerina pacifica Boomgaart 1949;

Pliocene, Java; Recent, Philippines, 903m) as synonyms of S.
insecta; however, they indicated a UDL of 2000m (top of lower
bathyal) despite the type occurrence of S. pacifica at an upper
middle-bathyal depth.
Siphonodosaria lepidula (Schwager 1866)
Nodosaria lepidula SCHWAGER 1866, p. 210, pl. 5, figs. 27, 28.

Nodosaria sagrinensis BAGG 1912, p. 58, pl. 16, fig. 4.
Nodogenerina sagrinensis (Bagg). FINGER 1990, p. 174, p. 175 figs.
17. FINGER, WEAVER, LIPPS and MILLER 1990, pl. 1, figs.
4345. FINGER 1992, p. 84, pl. 24, figs. 1016. FINGER,
NIELSEN, DEVRIES, ENCINAS and PETERSON 2007, fig, 13F.
Siphonodosaria matanzana (Palmer and Bermdez). ROBERTSON
1998, p. 180, pl. 67, fig. 2.
Siphonodosaria subspinosa (Cushman). ROBERTSON 1998, p. 180,
pl. 67, fig. 3.
Siphonodosaria lepidula (Schwager). HAYWARD, Kawagata,
Thomas 2012, p. 174, pl. 18, figs. 620.

Remarks: Hayward (2012: 175) noted As the most common and
morphotypically variable species of Siphonodosaria, S. lepidula
has been recorded and described under many names. They
reassigned many of those other names to different genera based
on details of the aperture, which is often missing and had not been
thoroughly investigated until recently. The extensive synonymy
presented by Hayward for S. lepidula includes the specimens
imaged here. That shown in Plate 16, Figure 29 matches very
well with S. hispidula (Cushman; Recent Philippines, 903m)
in Hayward (2002, pl. 3, fig. 16), which Hayward (2012) later
synonymized with S. lepidula. Most of the Chilean specimens
are more like that of Plate 16, Figure 28 , which was previously
included in Finger (2007) as S. sagrinensis (Bagg 1912; Early
Miocene, California), but Hayward (2012) also synonymized
that species with S. lepidula. Chilean specimens typically have
subspherical chambers fringed below centerline with very small
spines, which in later chambers tend to transform into fine striae,
and most specimens also have an apical spine.
Comparative species: Nodosaria insecta var. spinifera LeRoy
1941 (MiocenePliocene, Sumatra) is not mentioned in Hayward
(2012). It has similarly shaped chambers with small spines on the
lower part of chambers and its type figure has an apical spine; these
are features seen within Chilean populations of S. lepidula and if
LeRoys variety has the aperture of Siphonodosaria, S. spinifera
should be added to the synonymy of S. lepidula. Siphonodosaria
antillea (= Nodosaria antillea Cushman 1923; Recent, off east
coast of United States. 307m) and S. bradyi (= Nodogenerina
bradyi Cushman 1927b; Recent, South Pacific) have campanulate
chambers with more prominent spines at the basal shoulder
of each chamber; Hayward (2002) synonymized both with
S. lepidula, but Hayward (2012) retained the latter species as
distinct. The chambers of Strictocostella pupa (Pl. 17, fig. 26)
and Strictocostella advena (= Nodogenerina advena Cushman
and Laiming 1931; Early Miocene, California) are rectilinear

and smooth. Siphonodosaria pomuligera (Pl. 17, Fig. 25) has the
apical spine and subspherical chambers of S. sagrinensis, but lacks
the fine ornamentation. Siphonodosaria sagrinensis, therefore,
appears to be morphologically intermediate between S. lepidula
and S. pomuligera.
Occurrence: Navidad Fm. (CPUP, MOS, MPUP, NAV5, PPP,
PPT, PTA), Ranquil Fm. (FRA, FRM, LEB, MIB, MS10, RAN,
RQK, RQT), Lacui Fm. (CUC, PCB).
Maximum relative abundance: Common (PPN).
Siphonodosaria pomuligera (Stache 1865)
Plate 17, Figure 25
Dentalina pomuligera Stache 1865, p. 204, pl. 22, fig. 31.

Siphonodosaria pomuligera (Stache). HAYWARD, Kawagata,
Thomas 2012, p. 178, pl. 19, figs. 412.
Type age and locality: Early Oligocene, New Zealand.

Upper depth limit: Upper bathyal, 400m (Hayward et al. 2012)
Distinguishing features: Chambers subspherical and smooth;
offset apical spine.
Remarks: Generic assignment based on Hayward (2012), as the
single Chilean specimen is missing the ultimate chamber with
aperture.
Occurrence: El Peral beds (LPER, NLP), Navidad Fm. (MAT,
PPT, PTA), Lacui Fm. (CHE, PCB).
Siphonodosaria sentifera (Cushman and Parker 1931)
Plate 17, Figure 27
Nodosaria parexilis var. sentifera Cushman and Parker 1931, p.

6, pl. 1, fig. 16.
Siphonodosaria sentifera (Cushman and Parker). HAYWARD,

Stratigraphic range: Late Cretaceous to Miocene.
al. 2012).
Remarks: As is the case for the holotype, the Chilean specimens
are short segments consisting of spiny, ovate chambers.
Comparative species: The test of Nodosaria aculeata dOrbigny
1846 (Middle Pliocene, Austria) is considerably stouter with less
constricted sutures.
Strictocostella Patterson 1987
Type species: Ellipsonodosaria modesta var. prolata Cushman
and Bermdez 1937.
451
Strictocostella pupa (Karrer 1878)
Plate 17, Figure 26
Nodosaria pupa Karrer 1878, p. 89, pl. 5, fig. 9.

Nodogenerina advena CUSHMAN and LAIMING 1931, p. 106, pl. 11,
fig. 19. (Early Miocene, California)
Ellipsonodosaria oinomikadoi ISHIZAKI 1943, p. 685, figs. 710.
(Early Pliocene, Japan).
Type age and locality: Late Tertiary, Philippines, depth not

recorded.
Stratigraphic range: Early Miocene to Early Miocene.
Upper depth limit: Middle bathyal (see above Remarks for
genus).
Remarks: Karrer did not indicate a depository for his holotype,
nor did he mention or illustrate apertural details, but his type
figure is a form otherwise identical to Siphonodosaria (=
Nodogenerina advena Cushman and Laiming 1931; Early
Miocene, California), whose holotype is missing the early
chambers. Hayward et al. (2012) did not mention Karrers species
anywhere in their book, but they reassigned Cushman and
Laimings species to Strictocostella because of its denticulate
aperture, which had been shown in more recent publications
(e.g., Finger et al. 1990). It is assumed here that the species are
synonymous, but a study of S. pupa topotypes may be the only
way to confirm this.
Comparative species: Siphonodosaria paucistriata (=
Nodosarella paucistriata Galloway and Morrey 1929; probably
Late Eocene, Ecuador) has a curved test with short striae
crossing sutures. A similar form is Nodosaria glabra dOrbigny
1826 (fossil, Italy), which was based on synonymy with figures in
Soldani (17891799), but a primary type was never described or
figured; Parker, Jones and Brady (1865) subsequently examined
Soldanis figures and stated that the chambers are globular (their
figure shows subglobular chambers).
Occurrence: El Peral beds (LPER, NLP), Navidad Fm. (MOS,
PTA), Ranquil Fm. (MIB).
TODDOstomella Hayward, in Hayward et al. 2012
Type species: Siphonodosaria chileana Todd and Kniker 1952.
Toddostomella hochstetteri (Schwager 1866)
Nodosaria hochstetteri Schwager 1866, p. 349, text-fig. 5 (p. 351).

Nodosaria? hochstetteri Schwager. ROBERTSON 1998, p. 44, pl. 15,
fig. 3.
Toddostomella hochstetteri (Schwager). HAYWARD, Kawagata,
Thomas 2012, p. 195, pl. 24, figs. 810.

Stratigraphic range: Cretaceous to early Pleistocene.
Upper depth limit: Middle bathyal, 1500m (Hayward et al. 2012).
Remarks: None of the Chilean specimens preserve the
Y-shaped tooth and crenulated lip, but their chamber shape and
ornamentation are also diagnostic features of Toddostomella
(Hayward et al. 2012).
452
Comparative species: Toddostomella spinosa (= Dentalina

spinosa dOrbigny 1846; Middle Pliocene, Austria) is a slender
form that does not show a tendency toward rounder chambers.
Occurrence: El Peral beds (NLP), Navidad Fm. (CPUP),
Ranquil Fm. (FRA).
Order ROTALIIDA Ehrenberg 1838
Superfamily DISCORBOIDEA Ehrenberg 1838
Family BAGGINIDAE Cushman 1927c
Subfamily BAGGININAE Cushman 1927c
Cancris de Montfort 1808
Type species: Cancris auriculatus de Montfort 1808 = Nautilus
auriculus Fichtel and Moll 1798.
Cancris auriculus (Fichtel and Moll 1798)
Nautilus auricula Fichtel and Moll 1798, p. 108, pl. 20, figs. af.
Rotalia brongniartii DORBIGNY 1826, p. 273; type fig. not given.
(fossil, Italy)
Rotalia sagra DORBIGNY 1839a, p. 77, pl. 5, figs. 1315. (Recent,
Cuba and Jamaica)
Pulvulina oblonga (Williamson). BRADY 1884, p. 688, pl. 106, fig. 4.
Cancris sagra var. communis Cushman and Todd 1942, p. 79, pl. 19,
figs. 810; pl. 20, fig. 1 (Middle Pliocene, Florida).
Cancris intermedius Cushman and Todd 1942, p. 88, pl. 22, figs. 11, 12
(Miocene, Australia).
Cancris ovatus Cushman and Todd 1942, p. 89, pl. 22, fig. 1 (Oligo
cene, Australia).
Cancris auriculus (Fichtel and Moll). PAPP and SCHMID 1985,
pl. 52, figs. 7-13. LOEBLICH and TAPPAN 1987, pl. 591, figs.
13. JONES 1994, p. 105, pl. 106, fig. 4. ZAPATA and CEAR
2004, p. 19, pl. 3, fig. 1. HOLBOURN, HENDERSON and MA
CLEOD 2013, p. 134135.
Cancris sagra (dOrbigny). WHITTAKER 1988, p. 115, pl. 15, figs.
1315 (Pliocene, Ecuador).
Type age and locality: Age not designated, Mediterranean Sea.

Stratigraphic age: Oligocene to Recent.
Comparative species: Cushman and Todd (1942) distinguished
C. sagra var. communis Cushman and Todd 1942 (Middle
Pliocene, Florida), C. intermedius Cushman and Todd 1942
(Miocene, Australia) and C. ovatus (Oligocene, Australia), all of
which fall into the range of variation exhibited by the types of
Fichtel and Molls species illustrated in Le Calvez (1977) and
Rgl and Hansen (1984).
Occurrence: El Peral beds (LPER, NLP), Navidad Fm. (PPP,
PTA), Lacui (PCB).
Valvulineria Cushman 1926d
Type species: Valvulineria californica Cushman 1926d.
Valvulineria ecuadorana Cushman and Stevenson 1948
Plate 18, Figure 1
Valvulineria ecuadorana Cushman and STEVENSON 1948, p. 64,

pl. 10, fig. 12.
Type age and locality: Early Miocene, Ecuador.
Stratigraphic age: Early Miocene.
Upper depth limit: Neritic (for V. araucana and V. inflata in

Zapata and Cear 2004).
Orbitina Sellier de Civrieux 1977

Type species: Orbitina carinata Sellier de Civrieux 1977.
Occurrence: Navidad Fm. (CPUP, MOS, PPP), Ranquil Fm.

(MS10).
Orbitina parri (Collins 1974)
Rosalina parri Collins 1974, p. 46, 47, pl. 3, fig. 36 (plesiotype);

holotype figured in Parr 1945, pl. 10, fig. 3.
Family DISCORBIDAE Ehrenberg 1838

EPONIDES Montfort 1808
Type species: Eponides repandus (Fichtel and Moll 1798)
Plate 18, Figure 5
Type age and locality: Holocene, off southeastern Australia

(depth not given).
Eponides orientalis Asano 1937
Eponides orientalis ASANO 1937, p. 117, pl. 16, fig. 8.
Remarks: Differs from holotype in having much finer radial

grooves on the umbilical side.
Type age and locality: Recent, Japan
Occurrence: Ranquil Fm. (LEB), Lacui Fm. (PCB).
Family ROSALINIDAE Reiss 1963

Gavelinopsis Hofker 1951
Type species: Discorbina praegeri Heron-Allen and Earland
1913.
Plate 18, Figure 2
Distinguishing features: Sutures on spiral side are thick, flush,

and oblique, whereas those on umbilical side are thick, raised,
and radial.
Occurrence: Lacui Fm. (CHO)
Eponides ouachitaensis Howe and Wallace 1932
Gavelinopsis alhamensis Gonzlez-Donoso 1968
Plate 18, figs. 6
Gavelinopsis alhamensis Gonzlez-Donoso 1968, p. 74, 76, pl.

1, fig. 7.
Plate 18, Figure 3
Type age and locality: Late Miocene, Spain.
Eponides ouachitaensis HOWE and WALLACE 1932, p. 69, pl. 13,

fig. 8.
Stratigraphic range: Early to Late Miocene.
Stratigraphic range: Eocene to Early Miocene
Occurrence: Navidad Fm. (NAV5), Ranquil Fm. (LEB), Santo

Domingo Fm. (VAL).
Upper depth limit: Neritic
Occurrence: Ranquil Fm. (RQK).
Rosalina dOrbigny 1826

Type species: Rosalina globularis dOrbigny 1826.

Eponides parantillarum (Galloway and Heminway 1941)
Plate 18, Figure 4
Eponides parantillarum Galloway and Heminway 1941, p. 374,

pl. 18, fig. 1.
Neoeponides parantillarum (Galloway and Heminway). KOHL 1985,
p. 76, pl. 25, figs. 7, 8. ROBERTSON 1998, p. 194, pl. 74, fig. 2.
Type age and locality: Early Miocene, Puerto Rico.

Comparative species: Eponides antillarum (= Rotalina antillarum dOrbigny 1839a; Recent, Cuba) has depressed sutures
and a lobulate periphery.
Occurrence: El Peral beds (LPER), Navidad Fm. (CPUP, MOS,
NAV5, PPN, PPP, RAP), Ranquil Fm. (FRA, RAN, RQK, RQT),
Lacui Fm. (CHE, PCB, PNH).
Rosalina rugosa dOrbigny 1839c
Plate 18, Figure 7
Rosalina rugosa DORBIGNY 1839c, p. 42, pl. 2, figs. 1214.
Type age and locality: Recent, coast of Patagonia, Argentina

(depth not given).
Comparative species: Rosalina candeina dOrbigny 1839a
(Recent, Cuba) has more rounded chambers and a more rounded
periphery. Rosalina natlandi Church 1968 (Recent, off Baja
California) has a rounded edge, as well as six chambers in the
final whorl. Rosalina californica Finger and Lipps 1990 (in
Finger et al. 1990; Miocene, California) has chambers that are
less inflated on the umbilical side, the sutures are more curved
and oblique, and the periphery is only slightly lobulate.
Occurrence: Navidad Fm. (MOS, PTA).
453

Family SPHAEROIDINIDAE Cushman 1927c
Sphaeroidina dOrbigny 1826
Type species: Sphaeroidina bulloides dOrbigny 1826.
Sphaeroidina bulloides dOrbigny 1826
Plate 19, Figure 8
Sphaeroidina bulloides DORBIGNY 1826, p. 257, modle no. 265.

INGLE, KELLER and KOLPACK 1980, pl. 9, fig. 16. KOHL
1985, p. 59, pl. 14, fig. 6. PAPP and SCHMID 1985, pl. 90, figs.
712. VAN MORKHOVEN, BERGGREN and EDWARDS
1986, p. 80, pl. 24, figs. 1, 2. LOEBLICH and TAPPAN 1987, pl.
617, figs. 16. WHITTAKER 1988, p. 107, pl. 14, figs. 12, 13.
JONES 1994, p. 91, pl. 84, figs. 15. ROBERTSON 1998, p. 196,
pl. 74, fig. 4. HAYWARD, GRENFELL, SABAA, NEIL and
BUZAS 2010, p. 231, pl. 32, figs. 12, 13.
Type age and locality: Fossil and Recent, Italy.

Stratigraphic range: Early Oligocene Zone P19 through
Pleistocene Zone N23 (van Morkhoven, Berggren and Edwards
1986).
Upper depth limit: Middle neritic, but most abundant at upper
and middle bathyal depths; size increases with depth so that
upper bathyal forms are about one third the size of deeper ones
(van Morkhoven, Berggren and Edwards 1986: 82). Modern
UDL off Chile is lower bathyal (table 9). Chilean Miocene
specimens tend to be large; hence, they are assigned a UDL of
upper middle bathyal.
Remarks: Specimens of this species are relatively large and
abundant in the Chilean Neogene.
Occurrence: El Peral beds (NLP, LPER), Navidad Fm. (all except
LBZ), Ranquil Fm. (all except LEB), Santo Domingo Fm. (VAL),
Lacui Fm. (CHE, PCB).
Maximum relative abundance: Common (MAT, MPUP, FRM,
RAN, MIB, VAL).
Superfamily DISCORBINELLOIDEA Sigal 1952
Family PARRELLOIDIDAE Hofker 1956
Cibicidoides Thalmann 1939
Type species: Truncatulina mundula Brady, Parker and Jones
1888.
Cibicidoides bradyi (Trauth 1918)
Truncatulina bradyi Trauth 1918, p. 235; type-figure not designat

ed.
Cibicidoides bradyi (Trauth). INGLE, KELLER and KOLPACK 1980,
pl. 6, figs. 11, 12. WHITTAKER 1988, p. 141, pl. 22, figs. 1315.
VAN MORKHOVEN, BERGGREN and EDWARDS 1986, p.
100, pl. 30, figs. 1, 2. HAYWARD, GRENFELL, SABAA, NEIL
and BUZAS 2010, p. 208, pl. 22, figs. 13.
Cibicidoides robertsonianus (Brady). KOHL 1985, p. 98, pl. 35, fig. 4.
Cibicidoides ferasini (Conato). ROBERTSON 1998, p. 202, pl. 78, fig.
4.

noted localities include the Atlantic, SW Pacific and Coral
Sea.
Stratigraphic range: Late Early Eocene Zone P9 through Pleistocene Zone N23 (van Morkhoven, Berggren and Edwards 1986).
Upper depth limit: Upper bathyal based on Gulf of Mexico
(Pflum and Frerichs 1976). Van Morkhoven, Berggren and
Edwards (1986) note that Egger (1893) recorded this form as
Truncatulina dutemplei from outer neritic depths, it is more
often found at bathyal depths.
Remarks: Van Morkhoven, Berggren and Edwards (1986: 100)
note that Cibicidoides bradyi has 610 chambers in the final
whorl and a broadly rounded axial periphery and it grades into
C. robertsonianus (Brady 1881), which has 1314 chambers in
the final whorl and an angular to carinate periphery. The Chilean
Neogene specimens show the full range of edge variations but
only 56 chambers in the outer whorl. They are lumped here
as the older nomen, C. bradyi, because it was not possible to
consistently different them by test and chamber shapes, or
coarseness of porosity. Although this species has been recorded
as shallow as outer neritic by Egger (1893), it is most often found
in the bathyal realm (see van Morkhoven, Berggren and Edwards
1986: 102). Cibicidoides bradyi has an upper bathyal UDL in
the Gulf of Mexico (Pflum and Frerichs 1976), where UDLs
tend to be shallower than in the East Pacific; this suggests that
the Chilean Neogene UDL is probably deeper, which is what
Ingle, Keller and Kolpacks (1980) data on the Peru-Chile trench
indicate.
PLATE 18
Figures 2, 3, 4c, 5c, and 6c are photomicrographs; all other images are SEMs. Scale bars in m.
1 V
alvulineria ecuadorana Cushman and Stevenson, UCMP50354, MS10.
6 G
avelinopsis alhamensis Gonzlez-Donoso, UCMP50359, VAL.
2 Eponides orientalis Asano, UCMP50355, CHO.
7 Rosalina rugosa dOrbigny, UCMP50360, PTA.
3 Eponides ouachitaensis (Howe and Wallace), UCMP50356, RQK.
810 C
ibicidoides bradyi (Trauth): 8, UCMP50361, PTA. 9,
UCMP50362, NLP. 10, UCMP50363, FRA.
4 E
ponides parantillarum Galloway and Heminway, UCMP50357, RAP.
ibicidoides havanensis (Cushman and Bermdez), NLP:

11, 12 C
11, UCMP50364. 12, UCMP50365.
5 Orbitina parri (Collins), UCMP50358, LEB.
454
Kenneth L. Finger
Plate 18
455
Comparative species: Cibicidoides robertsonianus (= Planorbulina

(Truncatulina) robertsonianus Brady 1881; Caribbean, 2340m)
has an acute, noncarinate edge and 1314 chambers in the outer
whorl. Cibicidoides spiralis (= Cibicides spiralis Natland 1938;
Lower Pliocene, California) has nearly equidimensional chambers, a
subrounded periphery, and 10 chambers in its outer whorl.
Occurrence: El Peral beds (NLP, LPER), Navidad Fm. (PTA),
Ranquil Fm. (FRA, MIB, RQK, RQT), Santo Domingo Fm.
(VAL), Lacui Fm. (PCB).
Cibicidoides compressus (Cushman and Renz 1941)
Cibicides floridanus var. compressa Cushman and Renz 1941, p.

24, pl. 6, fig. 9.
Cibicidoides compressus (Cushman and Renz). VAN MORKHOVEN,
BERGGREN and EDWARDS 1986, p. 137, pl. 44, figs. 1, 2.
HOLBOURN, HENDERSON and MACLEOD 2013, p. 166167.
Cibicidoides floridanus (Cushman). WHITTAKER 1988, p. 143, pl.
20, figs. 46. KOHL 1985, p. 97, pl. 35, fig. 2. ROBERTSON
1998, p. 202, pl. 79, fig. 2.
Type age and locality: Early to Middle Pliocene, Venezuela.

Stratigraphic range: Early Miocene Zone N5 through Late
Miocene Zone N16 (van Morkhoven, Berggren and Edwards
1986).
Upper depth limit: Upper bathyal; primarily upper middle
bathyal (van Morkhoven, Berggren and Edwards 1986).
Distinguishing features: Outline subcircular, periphery weakly
lobulate; compressed and biconvex to nearly planoconvex in
edge view; spiral side shows relatively wide outer whorl and
prominent boss, more so in megalospheric specimens; few
chambers of earlier whorl are discernible; chambers of outer
whorl with 1016 chambers that are typically 24 times longer
than wide, nearly doubling in width as they flare outward; sutures
arcuate, limbate on both dorsal and ventral sides.
Remarks: This is one of the most abundant and commonly
occurring species in the Chilean fauna. Although the type-figure
of Cibicides floridanus var. compressa shows it to be more
ovate and with much longer later chambers, van Morkhoven,
Berggren and Edwards (1986) noted that the species shows much
morphological variation, and their figures closely match the form
shown here as Plate 18, Figure 3.
Comparative species: Whittaker (1988: pl. 20, figs. 412) identified
similar forms in the Early and Middle Pliocene of Ecuador as
Cibicidoides floridanus (= Truncatulina floridana Cushman
1918b; Miocene, Florida), C. crebbsi (= Eponides crebbsi Hedberg
1937; OligoceneMiocene, Venezuela), and C. mckannai (=
Cibicides mckannai Galloway and Wissler 1927; Pleistocene,
California). His figured specimen of C. floridanus most closely
matches that shown here as Plate 18, Figure 3. The type figures
of those three species, however, are noticeably different from his
illustrated specimens. Cibicidoides floridanus has the umbilical
side showing raised sutures coalescing at a distinct umbo, C.
mckannai is more inflated ventrally, and C. crebbsi is more
inflated and with very sinuous ventral sutures. There are many
other species that resemble this form in one aspect or another, and
they can be difficult to distinguish from ecophenotypic variants.
Cibicidoides falconensis (= Cibicides falconensis Renz 1948; Late
OligoceneEarly Miocene, Venezuela, Mexico, and Trinidad)
456
has a sharp edge and small umbo on the spiral side. Cibicidoides
miocenica (= Cibicides floridanus miocenica Colom 1946;
Miocene, Spain) has a small, clear umbilical plug. Cibicidoides
umbonatus (= Cibicides umbonatus Phleger and Parker 1951;
Pleistocene, Gulf of Mexico) clearly displays its whorls.
Cibicidoides pseudoungeriana (= Truncatulina pseudoungeriana
Cushman 1922a; Oligocene, Mississippi) has a very lobulate
periphery. Cibicidoides pachyderma (= Truncatulina pachyderma
Rzehak 1886; early Middle Pliocene, Czechoslovakia) is not as
compressed, particularly along the periphery, and has a large
umbilicus. Cibicidoides terryi (= Cibicides terryi Coryell and
Mossman 1942; Pliocene, Pacific coast of Panama) has a small,
flat umbo on the spiral side. Cibicidoides mecapetecensis (=
Anomalina mecapetecensis Nuttall 1932; Early Oligocene,
Mexico) has a lobulate periphery and a narrow, beaded periphery.
Cibicidoides kullenbergi (= Cibicides kullenbergi Parker, in
Phleger, Parker and Person 1953; Holocene, North Atlantic) has
an umbilical side that is much more convex; van Morkhoven,
Berggren and Edwards (1986) consider C. kullenbergi a junior
synonym of C. mundulus (= Truncatulina mundulus Brady, Parker
and Jones 1888; Recent, off Brazil, 475m).
PPP, PPT, PTA), Ranquil Fm. (all except MS10), Lacui Fm.
(CHE, CHO, PCB, PNH).
Maximum relative abundance: Abundant (LEB).
Cibicidoides havanensis (Cushman and Bermdez 1937)
Cibicides havanensis Cushman and Bermdez 1937, p. 28, pl. 3,

figs. 13.
Cibicidoides havanensis (Cushman and Bermdez). VAN MORK
HOVEN, BERGGREN and EDWARDS 1986, p. 189, pl. 64A, figs.
14; pl. 64B, figs. 1, 2. ROBERTSON 1998, p. 204, pl. 80, fig. 1.
Type age and locality: Eocene, Cuba.

Stratigraphic range: Early Eocene Zone P7 through the Middle
Miocene Zone N10.
Upper depth limit: Lower middle bathyal (van Morkhoven,
Berggren and Edwards 1986).
Remarks: Van Morkhoven, Berggren and Edwards (1986)
recognized this species as one of the cosmopolitan deep-water
species. Resig (1976) recorded it at DSDP Sites 320 and 321,
on the eastern margin of the Nazca Ridge, as ranging to early
Middle Miocene Zone N8.
Occurrence: El Peral beds (NLP, LPER).
Cibicidoides renzi (Cushman and Stainforth 1945)
Planulina renzi Cushman and Stainforth 1945, p. 72, pl. 15,

fig. 1. VAN MORKHOVEN, BERRGREN and EDWARDS 1986,
p. 133, pl. 43A, figs. 15; pl. 43B, figs. 1, 2. ROBERTSON 1998,
p. 214, pl. 85, fig. 2. HOLBOURN, HENDERSON and MA
CLEOD 2013, p. 410411.
Cibicidoides renzi (Cushman and Stainforth). WHITTAKER 1988, p.
147, pl. 21, figs. 16.
Type age and locality: Middle Miocene, Trinidad.
Stratigraphic range: Early Oligocene through Late Miocene.

Upper depth limit: Upper bathyal (referred to as bathyal by van
Morkhoven, Berggren and Edwards 1986).
Cibicidoides sp.
Plate 19, Figure 7

Distinguishing features: Compressed, coarsely punctate, 910
chambers in last whorl, oblique and slightly curved limbate
sutures connected to bluntly rounded peripheral carina.
Occurrence: Lacui Fm. (CHE, CHO, CUC).
Family PSEUDOPARRELLIDAE Voloshinova, in Voloshinova
and Dain 1952
Subfamily PSEUDOPARRELLINAE Voloshinova, in Voloshi
nova and Dain 1952
Pseudoparrella Cushman and ten Dam 1948
Type species: Pulvinulinella subperuviana Cushman 1926d.
Pseudoparrella naraensis Kuwano 1950
Plate 19, Figure 10
Pseudoparrella naraensis Kuwano 1950, p. 317, text-fig. 6 on p. 313.

noted localities are in the Atlantic and Indian Oceans.
Stratigraphic range: Early Oligocene Zone P19 through Pleistocene Zone N23 (van Morkhoven, Berggren and Edwards 1986).
Upper depth limit: Upper middle bathyal (Bandy and Rodolfo
1964).
Remarks: Van Morkhoven, Berggren and Edwards (1986: 90)
refer to this species as a bathyal-abyssal taxon, noting that in
the Gulf of Mexico it is approximately twice as abundant in
the lower bathyal zone than at middle and upper bathyal depths
(Parker 1954; Pflum and Frerichs 1976). Its upper depth limit
off California is lower middle bathyal (Bandy 1953a).
Comparable species: Laticarinina bullbrooki Cushman and
Todd 1942 (Miocene, Trinidad) is probably synonymous.
Superfamily PLANORBULINOIDEA Schwager 1877
Family PLANULINIDAE Bermdez 1952
PLANULINA dOrbigny 1826
Type species: Planulina ariminensis dOrbigny 1826.
Planulina sp.
Plate 19, Figure 11
Planulina sp. 3 of van Morkhoven, EDWARDS and BERG

GREN 1986, p. 55, pl. 17, figs. 13.
Stratigraphic range: Early Miocene through Pleistocene.
Distinguishing features: 56 chambers in outer whorl, nearly

smooth periphery, rounded edges.
Upper depth limit: Bathyal.
Comparative species: Pseudoparrella exigua (= Pulvinulina

exigua Brady 1884; Recent, 12 localities given in Atlantic,
Pacific, and Southern oceans, 1175011m) has a lobulate,
subacute edge. Pulvinulinella subperuviana Cushman 1926d
(Miocene, California) has 1011 chambers in its outer whorl and
a subacute edge.
Occurrence: Navidad Fm. (MAT, MOS, NAV5).
Family DISCORBINELLIDAE Sigal 1952
Subfamily DISCORBINELLINAE Sigal 1952
Laticarinina Galloway and Wissler 1927
Type species: Laticarinina carinata Galloway and Wissler
1927.
Laticarinina pauperata (Parker and Jones 1865)
Plate 19, Figure 9
Pulvinulina repanda var. menardii subvar. pauperata Parker and

Jones 1865, p. 395, pl. 16, figs. 50, 51.
Laticarinina pauperata (Parker and Jones). INGLE, KELLER and
KOLPACK 1980, pl. 9, fig. 12. RESIG 1981, pl. 6, fig. 8.
KOHL 1985, p. 77, pl. 26, fig. 1. VAN MORKHOVEN, BERG
GREN and EDWARDS 1986, p. 89, pl. 26, fig. 1. LOEBLICH
and TAPPAN 1987, pl. 631, figs. 14. WHITTAKER 1988, p.
128, pl. 17, figs. 1921. ROBERTSON 1998, p. 210, pl. 84, figs. 1,
2. HAYWARD, GRENFELL, SABAA, NEIL and BUZAS 2010,
p. 213, pl. 24, figs. 1921. HOLBOURN, HENDERSON and
MACLEOD 2013, p. 328329.
Comparative species: Planulina caribaea Cushman 1921

(Recent, Jamaica) shows ~2.5 whorls and has curved, not
subangular, sutures. Planulina antillea Drooger 1953 (Miocene,
Aruba) is not as compressed and lacks a keel. Planulina
ecuadorana Cushman and Stevenson 1948 (Early Miocene,
Ecuador) has depressed and gently curved sutures. Planulina
retia Belford 1966 (Middle Pliocene, Papua New Guinea) has
similarly subangularly reflexed sutures but its test distinctly
planoconvex. Planulina marialana Hadley 1934 (Oligocene,
Cuba) has strongly arched sutures but its chambers are narrower
and its dorsal center is pustulose.
Occurrence: Ranquil Fm. (RAN), El Peral beds (NLP).
Family CIBICIDIDAE Cushman 1927c
Subfamily CIBICIDINAE Cushman 1927c
Cibicides de Montfort 1808
Type species: Cibicides refulgens de Montfort 1808.
Cibicides cicatricosus (Schwager 1866)
Anomalina cicatricosa Schwager 1866, p. 260, pl. 7, figs. 4, 108.

Truncatulina akneriana (dOrbigny). BRADY 1884, pl. 94, fig. 8.
Cibicidoides cicatricosa (Schwager). RESIG 1981, fig. 8, figs. 13, 14.
VAN MORKHOVEN, BERGGREN and EDWARDS 1986, p.
63, pl. 16, fig. 1. JONES 1994, p. 98, pl. 94, fig. 8.
Cibicides cicatricosus (Schwager). WHITTAKER 1988, p. 142, pl. 21,
figs. 79.
457
Cibicidoides cicatricosus (Schwager). ROBERTSON 1998, p. 200, pl.

78, fig. 1. HOLBOURN, HENDERSON and MACLEOD 2013,
p. 164165.

Stratigraphic range: Late Oligocene (P21b) through Pleistocene
Zone N23).
Upper depth limit: Upper bathyal (van Morkhoven, Berggren
and Edwards 1986: 55).
Remarks: Younger specimens have a slight, subacute keel on
early chambers of their last whorl, whereas the peripheral edge
of adults is smoothly rounded. Typical specimens have thick,
slightly raised sutures on the early chambers of the last whorl, in
contrast to the depressed sutures of succeeding chambers.
LBZ, PTA, RAP), Ranquil Fm. (all except MS10), Lacui Fm.
(PCB, PNH, CHE).
Stratigraphic range: Early to Middle Pliocene.

Remarks: Populations of this species include forms ranging from
slightly concavo-convex to biconvex, with 69 chambers in the
outer whorl.
Comparative species: Cibicides miocenica (= C. floridanus
miocenica Colom 1946; Miocene, Spain) has more chambers
in the outer whorl and a more distinct inner whorl, but that
morphotype could be conspecific with C. mediocris.
Occurrence: El Peral beds (NLP, LPER), Navidad Fm. (MAT,
PPN, RAP), Ranquil Fm. (FRA, MS10, RQK), Santo Domingo
Fm. (VAL), Lacui Fm. (CHE, CHO, CUC, PCB).
Maximum relative abundance: Common (PPN, MS10, CHO,
CHE)
Cibicides umboniferus (Howchin and Parr 1938)
Maximum relative abundance: Common (LBZ, PCB)
Operculina (?) umbonifera Howchin and Parr 1938, p. 309, pl. 18,
figs. 3, 4, 6, 13, 14.
Cibicides lobatulus? (Walker and Jacob 1798)
Type age and locality: Miocene, Australia.
Nautilus spiralis lobatulus Walker and Jacob, in KANMACH

ER 1798, p. 642, pl. 14, fig. 36.
Distinguishing features: Moderately and equally to unequally

biconvex to nearly planoconvex with subacute to rounded
periphery, opaque umbo visible on both sides, very large on
spiral side, less so on umbilical side.
Plate 20, Figure 5
Type age and locality: Recent shore sand, England.

Remarks: The rare Chilean form lacking the slight keel of the
holotype.
Occurrence: Navidad Fm. (MOS, NAV5, PPN), Ranquil Fm.
(RAN), Lacui Fm. (CHE, PCB)
Cibicides mediocris Finlay 1940

Occurrence: Navidad Fm. (MOS, RAP, PPN, PTA), Ranquil Fm.
(FRA, LEB, MS10, RQK), Lacui Fm. (CHO, CHE, CUC, PCB).
Maximum relative abundance: Very abundant (PPN).
Cibicides sp.
Plate 21, Figure 1
Cibicides mediocris Finlay 1940, p. 464, pl. 67, figs. 198, 199.
Remarks: Distinguished by its planoconvex test, this does not

appear to be a juvenile form of any other Cibicides species
recognized in the Chilean Miocene.
Type age and locality: Middle Miocene, New Zealand.
Occurrence: Ranquil Fm. (LEB, RAN), Lacui Fm. (CHE, PCB)
PLATE 19
Figs. 3a, 5a, 9b, and 11a are photomicrographs; all other images are SEMs.
14 C
ibicidoides compressus (Cushman and Renz): 1, UCMP50366, MIB. 2, UCMP50367, FRA. 3, UCMP50368, FRA. 4,
rare form with unusually thick sutures. UCMP50369, PTA.
5, 6 Cibicidoides renzi (Cushman and Stainforth), PCB: 5,
UCMP50370. 6, UCMP50371.
7 Cibicidoides sp., UCMP50372, CHE.
458
8 Sphaeroidina bulloides dOrbigny, UCMP50373, FRA.

9 L
aticarinina pauperata (Parker and Jones), UCMP50374,
FRA.
10 Pseudoparrella naraensis Kuwano, UCMP50375, NAV5.
11 Planulina sp., UCMP50376, NLP.
Kenneth L. Finger
Plate 19
459
Falsocibicides Poignant 1958

Type species: Falsocibicides aquitanicus Poignant 1958.
Superfamily NONIONOIDEA von Schultze 1854

Family NONIONIDAE von Schultze 1854
Subfamily Nonioninae von Schultze 1854
Nonionella Cushman 1926c
Type species: Nonionella miocenica Cushman 1926c.
Falsocibicides sp.
Plate 21, Figure 2
Upper depth limit: Cibicidids that conform to a hard substrate

are generally restricted to inner-neritic depths.
Nonionella miocenica Cushman 1926c
Occurrence: El Peral beds (NLP), Navidad Fm. (LBZ, MOS),

Ranquil Fm. (LEB, RAN), Lacui Fm. (CUC).
Nonionella miocenica Cushman 1926c, p. 64; type-figure not given.

INGLE, KELLER and KOLPACK 1981, pl. 2, figs. 1518.
Saunders and Mller-Merz 1982, p. 270, pl. 2, figs. 913.
LOEBLICH and TAPPAN 1987, pl. 689, figs. 1821. WHIT
TAKER 1988, p. 166, pl. 24, figs. 1618.
Nonionella pulchella HADA 1931, p. 120, text-fig. 79. (Recent, Japan,
760m).
Nonionella auris (dOrbigny 1839c). BOLTOVSKOY and THEYER
1970, p. 323, pl. 4, fig. 8. (Recent, Chile, 88 & 120m).

Fontbotia Gonzlez-Donoso and Linares 1970
Type species: Anomalina wuellerstorfi Schwager 1866.
Fontbotia wuellerstorfi (Schwager 1866)
Plate 21, Figure 3
Anomalina wuellerstorfi Schwager 1866, p. 258, pl. 7, figs. 105,

107.
Truncatulina wuellerstorfi (Schwager). BRADY 1884, pl. 93, figs. 8, 9.
Cibicidoides wuellerstorfi (Schwager) RESIG 1981, pl. 8, figs. 16, 17.
JONES 1994, p. 98, pl. 93, figs. 8, 9. WHITTAKER 1988, p.
153, pl. 21, figs. 1012, 1621.
Planulina wuellerstorfi (Schwager). VAN MORKHOVEN, BERG
GREN and EDWARDS 1986, p. 48, pl. 14, figs. 1, 2. ROBERT
SON 1998, p. 216, pl. 86, fig. 2. HOLBOURN, HENDERSON
and MACLEOD 2013, p. 416417.
Fontbotia wuellerstorfi (Schwager). LOEBLICH and TAPPAN 1987,
pl. 634, figs. 1012.
Cibicides wuellerstorfi (Schwager). HAYWARD, GRENFELL, SA
BAA, NEIL and BUZAS 2010, p. 210, pl. 23, figs. 1722.

Stratigraphic range: Early Miocene through Pleistocene.
Upper depth limit: Usually lower bathyal; based on van Morkhoven, Berggren and Edwards (1986).
Remarks: The neritic form identified as Planulina wuellerstorfi
in Zapata and Cear (2004) appears to be a flatter and less ovate
species.
Occurrence: El Peral beds (LPER, NLP), Navidad Fm. (PPP),
Ranquil Fm. (FRA, RAN), Lacui Fm. (CUC).

Upper depth limit: Neritic (this species identified as Nonionella
auris in Zapata and Cear, 2004).
Occurrence: Navidad Fm. (CPUP, MAT, MOS, MPUP, NAV5),
Ranquil Fm. (RAN).
Nonionella stella Cushman and Moyer 1930
Plate 21, Figure 6
Nonionella miocenica var. stella Cushman and MOYER 1930, p.

56, pl. 7, fig. 17.
Type age and locality: Recent, California, 64166m.

Upper depth limit: Inner neritic.
Remarks: Cushman and Moyer (1930) believed the fingerlike processes over the umbilicus was a varietal character of
Nonionella miocenica, but I have never found it in association
with N. miocenica in the Miocene of California and Chile.
Occurrence: Navidad Fm. (PPP), Ranquil Fm. (RQK, RQT),
Lacui Fm. (CHO).
PLATE 20
All images are SEMs. Scale bars in m.
14 C
ibicides cicatricosus (Schwager): 1, UCMP50377, RAN. 2,
UCMP50378, FRA. 3, UCMP50379, FRA. 4, UCMP50380,
FRM.
5 C
ibicides lobatulus? (Walker and Jacob), UCMP50381,
PCB.
460
68 C
ibicides mediocris Finlay: 6, UCMP50382, PTA; 7,
UCMP50383, CHO; 8, UCMP50384, PTA.
9, 10 C
ibicides umboniferus (Howchin and Parr): 8, UCMP50385, PCB. 9, UCMP50386, RAN.
Kenneth L. Finger
Plate 20
461

Pseudononion Asano 1936b
Type species: Pseudononion japonicum Asano 1936b.
Pseudononion communis (dOrbigny 1846)
Plate 21, Figure 7
Nonionina communis DOrbigny 1846, p. 106, pl. 5, figs. 7, 8.

Nonion pizarrensis BERRY 1928, p. 269, text-figs. 13. (Recent, Peru,
48m)
Nonionina pizarrense var. basispinata Cushman and Moyer
1930, p. 54, pl. 7, fig. 18. (Recent, California, 6491m).
Nonion commune (dOrbigny). PAPP and SCHMID 1985, p. 45, pl. 34,
figs. 15.
Pseudononion pizarrensis (Berry). WHITTAKER 1988, p. 171, pl. 24,
figs. 13, 911.

Basin, Austria.
Stratigraphic range: Early to Late Miocene.
Remarks: Although its type figure shows an acutely edged
form without sutural pustules, Papp and Schmid (1985, pl.
34, figs. 25) illustrate specimens from dOrbignys sample
that show and edge progressing from subacute to rounded and
pustules filling depressed sutures. Whittaker (1988) notes that
syntypes of Nonion pizarrensis Berry 1928 (Recent, Peru,
48m) have the pustules characteristic of Pseudononion, which
invalidates the differentiation of Pseudononion basispinata (=
Nonionina pizarrense var. basispinata Cushman and Moyer
1930; Recent, California, 210300m); it is likely synonymous
with Pseudononion communis, as is P. incisum (= Nonionina
incisa Cushman 1926c; Miocene, California). Pseudononion
oinomikadoi Matsunaga 1963 (Late Miocene?Pliocene, Japan)
has a more ovate outline, limbate sutures and slightly more
compressed test.
Occurrence: El Peral beds (LPER, NLP), Navidad Fm. (PPT).
Pseudononion cuevasensis Saunders and Mller-Merz 1982
Plate 21, Figure 8
Pseudononion cuevasensis Saunders and Mller-Merz 1982,

p. 267, pl. 4, figs. 3438.
Pseudononion atlanticum (Cushman 1947). Saunders and

Mller-Merz 1982, p. 264, 268, pl. 3, figs. 1925.
Florilus grateloupi (dOrbigny). KOHL 1985, p. 92, pl. 32, fig. 3.
Pseudononion grateloupi (dOrbigny). ROBERTSON 1998, p. 224, pl.
90, fig. 6.
Type age and locality: Recent, Trinidad and Tobago, 18m; fossil
occurrence noted as ranging down to upper Miocene.
Upper depth limit: Inner neritic; based on type occurrence.
Remarks: The type description and figure of Nonionina
grateloupii dOrbigny 1839a (Recent, Caribbean) do not
indicate the presence of extra-umbilical pustules characteristic
of Pseudononion, nor do the photomicrograph and description
of its lectotype (Le Calvez 1977). Saunders and Mller-Merz
(1982) recognized the species collected off Trinidad as having a
small, deep umbilicus with a rim of pustules around it and with
no pustules centrally placed. They distinguished P. cuevasensis,
also collected off Trinidad, as having pustules along the spiral
suture and extending a short distance along the intercameral
sutures, and did not observe any forms intermediate between the
two species.
Comparative species: Nonionina elongata dOrbigny 1852
(Tertiary, France) is slightly more elongate, but not described as
having pustules; it could be synonymous with either Nonionina
grateloupi or Pseudononion cuevasensis. Pseudononion
japonicum Asano 1936b (type level not designated; recorded as
PlioceneRecent, Japan) has a less elongate test that is acutely
edged in the early part of the ultimate whorl, and its later
chambers are lobulate.
Occurrence: Navidad Fm. (NAV5, PPP, RAP), Ranquil Fm.
(FRA, RQK), Lacui Fm. (CHO, CUC, PCB).
Pseudononion novozealandicum (Cushman 1936a)
Nonionella novo-zealandica CUSHMAN 1936a, p. 88, pl. 13, fig. 16.
Type age and locality: Miocene, New Zealand.

PLATE 21
All images are SEMs. Scale bars in m.
1 Cibicides sp., UCMP50387, RAN.
6 Nonionella stella Cushman and Moyer, UCMP50392, CHO.
2 Falsocibicides sp., UCMP50388, LEB.
7 Pseudononion communis (dOrbigny), UCMP50393, NLP.
3 Fontbotia wuellerstorfi (Schwager), UCMP50389, FRA.
8 P
seudononion cuevasensis Saunders and Mller-Merz,
UCMP50394, RAP.
4, 5 N
onionella miocenica Cushman, NAV5: 4, UCMP50390. 5,
UCMP50391.
462
9, 10 P
seudononion novozealandicum (Cushman): 9, UCMP50395, PPP. 10, UCMP50396, CHO.
Kenneth L. Finger
Plate 21
463
Comparative species: Pseudononion papillatum Galloway and

Heminway 1941 (Late Oligocene, Puerto Rico) displays the
earlier whorl on the nonpapillate spiral side.
PPP, PPT, PTA), Ranquil Fm. (FRA, FRM, MIB, MS10, RAN,
RQT), Lacui Fm. (CHO, CUC, PNH).
Maximum relative abundance: Common (MPUP, NAV5)
Pseudononion ranquilensis Finger, n. sp.
acute periphery and the width of the outer whorl increases more
rapidly. Zeaflorilus pizarrensis (= Nonion pizarrensis Berry
1928; Recent, Peru) has fewer chambers, a rounded periphery,
depressed sutures, and a significantly greater increase in width
of its outer whorl. Other comparable forms have the granular
umbilical area that characterizes Pseudononion.
Occurrence: Navidad Fm. (LBZ, MAT, NAV5, PPN, PPP, PTA,
RAP), Ranquil Fm. (FRA, LEB, MS10, RQK, RQT), Lacui Fm.
(CHE, CHO, CUC).
Plate 22, Figure 1
Maximum relative abundance: Very abundant (CHE, PPN).
Description: Test slightly trochospiral, nearly planispiral,

moderately inflated, periphery slightly lobulate, outer whorl
consisting of six chambers, sutures depressed; surface smooth
but very finely papillate in sutures as they extend out from
the umbilicus and on base of apertural face; aperture an
interiomarginal slit.
Subfamily ASTRONONIONINAE Saidova 1981

Fijinonion Hornibrook 1964
Type species: Astrononion fijiensis Cushman and Edwards
1937.
Remarks: The other Pseudononion species in the Chilean

fauna are larger, more compressed forms with subacute edges
and considerably more chambers. Although only one specimen
was found, it is described as a new species because it is well
preserved and appears to be a unique form.
Type age and locality: Early Miocene; Ranquil Formation
locality MIB, exposed on coastal bluff in southern part of Caleta
Ranquil, close to fault contact with Millongue, coastal area of
central Chile.
Etymology: Name derived from that of the lithostratigraphic unit
in which it occurs.
Zeaflorilus Vella 1962
Type species: Nonionella parri Cushman 1936a.
Zeaflorilus chiliensis (Cushman and Kellett 1929)
Nonionella chiliensis Cushman and KELLETT 1929, p. 6, pl. 12,

fig. 4. BOLTOVSKOY and THEYER 1970, p. 348, pl. 4, fig.
6. ZAPATA, ZAPATA and GUTIRREZ 1995, p. 27, pl. 6, figs.
13.
Nonionella parri Cushman 1936a, p. 89, pl. 13, fig. 17. (Recent
shore sand, New Zealand)
Fijinonion obesum (Carter 1964)
Plate 22, Figure 5
Astrononion obesum CARTER 1964, p. 112, pl. 10, figs. 205, 206.
Type age and locality: Early Miocene, Victoria, Australia.

Upper depth limit: Probably neritic.
Distinguishing features; Involute planispiral, becoming subplanispiral in adults, with slightly lobulate outline, shows 10 or
more chambers; sutures flush to slightly depressed, umbilical flaps
fused.
Remarks: The Chilean form differs from the holotype of F. obesum
only by being more compressed. The sutures are occasionally
thickened as on Melonis affinis. Smaller specimens display
supplementary apertures, not seen on larger specimens due to the
more extensive fusion of the umbilical flaps. The fused area is not
thick or raised, nor does it have the stellate shape characteristic of
Astrononion. Planispiral specimens resemble Melonis barleeanus
(Pl. 22, Fig. 8), while the very low trochospiral ones are similar to
Anomalinoides salinasensis (Pl. 23, Fig. 6).
Comparative species: Melonis simplex (= Nonionina simplex
Karrer 1865; Miocene New Zealand) is more compressed and
slightly flaring in edge view, and lacks distinct umbilical flaps.
Fijinonion sphaeroides Saidova 1975 (Recent, Pacific, 1235
1350m ) is more robust. The holotype of Astrononion fijiensis
Cushman and Edwards 1937 (Recent, off Fiji) has a more
prominent, fused umbilical ring.
Type age and locality: Recent, off Chile.
Occurrence: El Peral beds (NLP), Lacui Fm. (PCB).
Upper depth limit: Inner neritic (based on Zapata, Zapata and

Gutirrez 1995).
Subfamily PULLENINAE Schwager 1877

Melonis de Montfort 1808
Type species: Melonis etruscus de Montfort 1808 = Nautilus
pompiliodes Fichtel and Moll 1798.
Remarks: Most of the Chilean Miocene specimens have 1418

chambers in the outer whorl as described for Nonionella parri,
but they show an ontogenetic progression in which the test goes
from fairly compressed and smooth and to less compressed with
very thick raised sutures. Populations include intermediate forms
that suggest the two species are conspecific variants.
Comparative species: Zeaflorilus boueanus (= Nonionina
boueana dOrbigny 1846; Middle Miocene, Vienna Basin) has an
464
Melonis affinis (Reuss 1851b)
Plate 22, Figure 7
Nonionina affinis Reuss 1851b, p. 72, pl. 5, figs. 32.

Nonion affine (Reuss). BOLTOVSKOY and THEYER 1970, p. 346, pl.
3, fig. 11.
Melonis affinis (Reuss) INGLE, KELLER and KOLPACK 1980, pl. 5,

figs. 1, 2. RESIG 1981, pl. 4, figs. 8, 9. WHITTAKER 1988,
pl. 24, figs. 2527. KOHL 1985, p. 100, pl. 36, fig. 4. ROB
ERTSON 1998, p. 226, pl. 91, fig. 3. JONES 1994, p. 107, pl.
109, figs. 8, 9. HAYWARD, GRENFELL, SABAA, NEIL and
BUZAS 2010, p. 222, pl. 28, figs. 17, 18.
Melonis sphaeroides Voloshinova. FIGUEROA et al. 2005, p. 349.
Type age and locality: Eocene, Germany.

Occurrence: Navidad Fm. (NAV5), Ranquil Fm. (FRM), Lacui
Fm. (PCB).
Melonis barleeanum (Williamson 1858)
Plate 22, Figure 8
Nonionina barleeana Williamson 1858, p. 32, pl. 3, figs. 68, 69.

Melonis barleeanus (Williamson). INGLE, KELLER and KOLPACK
1980, pl. 7, figs. 14, 15.
Melonis barleeanum (Williamson). LOEBLICH and TAPPAN 1987,
pl. 696, figs. 5, 6. HOLBOURN, HENDERSON and MACLEOD
2013, p. 354355.
Type age and locality: Recent, British Isles.

Upper depth limit: Upper middle bathyal (Ingle, Keller and
Kolpack 1980).
Occurrence: Navidad Fm. (NAV5, PPN, PPT, PTA, RAP),
Ranquil Fm. (FRA, RQT), Lacui Fm. (CUC).
Melonis pompilioides (Fichtel and Moll 1798), including f.
sphaeroides Voloshinova 1958
Plate 22, Figure 6
Nautilus pompilioides FICHTEL and MOLL 1798, p. 31, pl. 2, figs.

ac.
Nonionina soldanii DORBIGNY 1846, p. 109, pl. 5, figs. 15, 16.
Brady 1884, p. 727, pl. 109, figs. 10, 11. (Middle Pliocene,
Austria)
Nonionina pompilioides (Fichtel and Moll). Brady 1884, p. 727, pl.
109, figs. 10, 11. (Recent, North Atlantic).
Melonis pompilioides (Fichtel and Moll). MARTNEZ and PARADA
1968, pl. 1, figs. 31, 32. INGLE, KELLER and KOLPACK 1980,
pl. 9, fig. 14. RGL and HANSEN 1984, p. 30, pl. 2, figs. 1, 2;
pl. 3, fig. 1. VAN MORKHOVEN, BERGGREN and EDWARDS
1986, p. 72, pl. 23C, fig. 1. LOEBLICH and TAPPAN 1987,
pl. 696, figs. 7, 8. ROBERTSON 1998, p. 228, pl. 91, fig. 4.
JONES 1994, p. 108, pl. 109, figs. 10, 11. HAYWARD, GREN
FELL, SABAA, NEIL and BUZAS 2010, p. 223, pl. 28, figs. 19, 20.
Melonis sphaeroides Voloshinova 1958, pl. 3, figs. 8, 9.
WHITTAKER 1988, p. 165, pl. 24, figs. 28, 29. ROBERTSON
1998, p. 228, pl. 91, fig. 5. HOLBOURN, HENDERSON and
MACLEOD 2013, p. 358359.
species is most typical of deep-water associations. Lower bathyal

based on M. pompilioides recorded by Bandy and Rodolfo (1964)
and Ingle, Keller and Kolpack (1980) off Chile. Van Morkhoven,
Berggren and Edwards (1986) place the UDL in the middle bathyal
zone. The form reported by Figueroa (2005) as M. spheroides [sic]
appears to be M. affinis.
Remarks: Voloshinova (1958) recognized that Fichtel and Molls
holotype was from a Pliocene neritic deposit and differed from
deep-water forms that had been recovered from the North Atlantic.
In naming the latter form Melonis sphaeroides, she chose Bradys
figured specimen from 2840m in the Northeast Atlantic as the
holotype. Van Morkhoven, Berggren and Edwards (1986) viewed
M. pompilioides and M. sphaeroides as conspecific, but retained
the sphaeroides nomen as a forma because of its ecological
significance. The M. sphaeroides ecophenotype differs by its
smaller size, pronouncedly involute coiling, deeper umbilicus,
more rapid increase in chamber size, coarser perforations, and thin
to virtually unrecognizable flush sutures. The Chilean Miocene
populations of M. pompilioides vary inconsistently with regard to
these features; thus, it is difficult to clearly differentiate the two
primary ecophenotypes, but overall they trend toward the deepwater morphology. The compressed specimen shown in Holbourn,
HENDERSON and MACLEOD (2013, p. 356) as M. pompilioides
is M. affinis.
Occurrence: Navidad Fm. (CPUP, MOS, NAV5, PPP, PPT, PTA),
Ranquil Fm. (FRA, FRM, MIB, MS10, RAN, RQK, RQT), Santo
Domingo Fm. (VAL), Lacui Fm. (CHE).
Maximum relative abundance: Common (MS10).
Pullenia Parker and Jones, in Carpenter, Parker and Jones
1862
Type species: Nonionina bulloides dOrbigny 1846.
Pullenia bulloides (dOrbigny 1846)
Plate. 21, Figure 9
Nonionina bulloides DORBIGNY 1846, p. 107, pl. 5, figs. 9, 10.

Pullenia bulloides (dOrbigny). INGLE, KELLER and KOLPACK
1980, pl. 5, fig. 7. RESIG 1981, pl. 7, fig. 13. PAPP and
SCHMID 1985, p. 45, pl. 34, figs. 69. WHITTAKER 1988, p.
173, pl. 24, figs. 2032. JONES 1994, p. 92, pl. 84, figs. 12, 13.
225, pl. 29, figs. 16, 17.

Basin, Austria.
Upper depth limit: Shelf break (van Morkhoven, Berggren and
Edwards 1986, fig. 8).
Occurrence: El Peral beds (NLP), Navidad Fm. (NAV5, RAP),
Ranquil Fm. (FRA, MIB, RAN, RQT), Santo Domingo Fm.
(VAL).
Type age and locality: Pliocene, Italy and Recent, Mediterranean

Sea.
Pullenia subcarinata (dOrbigny 1839c)
Stratigraphic range: Late Oligocene through Pleistocene; Early

Miocene to Pliocene for f. sphaeroides.
Nonionina subcarinata DORBIGNY 1839c, p. 28, pl. 5, figs. 23, 24.

Nonionina quinqueloba REUSS 1851b, p. 71, pl. 5, fig. 31. (Eocene,
Germany)
Pullenia compressa Seguenza 1880, p. 307, pl. 17, fig. 14. (Plio
cene, Italy)
Pullenia salisburyi Stewart and Stewart 1930, p. 72, pl. 8, fig.
Upper depth limit: Globally neritic for Melonis pompilioides s.s.,

middle bathyal for f. sphaeroides (see remarks below), but the
465
2. (Pliocene, California).
Pullenia quinqueloba JONES 1994, p. 92, pl. 84, figs. 14, 15.
Pullenia subcarinata (dOrbigny) ZAPATA 1999, fig. 27.
BOLTOVSKOY and THEYER 1970, p. 352, pl. 5, fig. 18. WHIT
TAKER 1988, p. 173, pl. 24, figs. 3338.

Upper depth limit: Neritic (Zapata 1999).
Remarks: Although this species was described as a 6-chambered
form with a subacute peripheral edge, the range is 46 chambers,
with 5 chambers most common.
Comparative species: Pullenia inglei Finger and Lipps (in
Finger et al. 1990; Miocene, California) has 56 chambers in
the outer whorl and a more rounded peripheral edge. Pullenia
malkinae Coryell and Mossman 1942 (Pliocene, Pacific coast of
Panama) has 78 chambers in the outer whorl.
Remarks: The recovered Chile specimens are relatively large.

Occurrence: Navidad Fm. (PTA), Ranquil Fm. (FRA, FRM).
Chilostomella Reuss, in Cjek 1849
Type species: Chilostomella ovoidea Reuss 1850.
Chilostomella ovoidea Reuss 1850
Plate 23, Figure 1
Chilostomella ovoidea Reuss 1850, p. 380, pl. 48, fig. 12. LOE
BLICH and TAPPAN 1987, pl. 701, figs. 6, 7. ROBERTSON
1998, p. 234, pl. 93, fig. 3. FINGER 1992, p. 88, pl. 34, figs,
1928. JONES 1994, p. 61, pl. 55, figs. 1516, 1923. HAY
21, figs. 3640.
Chilostomella oolina Schwager 1878. INGLE, KELLER and KOL
PACK 1981, pl. 5, figs. 9, 10. ZAPATA and CEAR 2004, p. 20,
fig. 1. HOLBOURN, HENDERSON and MACLEOD 2013, p.
148149.
Occurrence: El Peral beds (NLP), Navidad Fm. (PPP), Ranquil

Fm. (FRA, FRM, LEB, MIB, RAN, RQK, RQT), Santo Domingo
Fm. (VAL), Lacui Fm. (PCB).
Type age and locality: Tertiary, Poland.
Superfamily CHILOSTOMELLOIDEA Brady 1881

Family CHILOSTOMELLIDAE Brady 1881
Subfamily CHILOSTOMELLINAE Brady 1881
Allomorphina Reuss, in Cjek 1849
Type species: Allomorphina trigona Reuss 1850 (first species
named).
Occurrence: Navidad Fm. (CPUP, MOS, PPP, PPT, PTA),

Ranquil Fm. (MIB, RQK, RQT).
Allomorphina pacifica Cushman and Todd 1949
Plate 22, Figure 12
Allomorphina pacifica Cushman and Todd 1949, p. 68, pl. 12, figs.
69. JONES 1994, p. 61, pl. 55, figs. 2426.
Type age and locality: Pliocene, Fiji Islands.

Stratigraphic range: Miocene or earlier to Recent.

HIDINA Gheorghian, Iva and Gheorghian 1968
Type species: Hidina variabilis Gheorghian, Iva and Gheorghian
1968.
Hidina variabilis Gheorghian, Iva and Gheorghian 1968
Plate 23, Figure 2
Hidina variabilae Gheorghian, Iva and Gheorghian 1968, p.

195, pl. figs. 14.
Hidina variabilis Gheorghian, Iva and Gheorghian. LOEBLICH and
TAPPAN 1987, pl. 701, figs. 1114.
PLATE 22
Figures 3a and 5 are photomicrographs; all other images are SEMs. Scale bars in m.
1 P
seudononion ranquilensis Finger, n. sp., holotype UCMP50397, MIB.
24 Z
eaflorilus chiliensis (Cushman and Kellett): 2, UCMP50398, PPN. 3, UCMP50399, LEB; 4, UCMP50400, RQS.
5 Fijinonion obesum (Carter), UCMP50401, NLP.
6 M
elonis pompilioides (Fichtel and Moll), UCMP50402,
NAV5.
7 Melonis affinis (Reuss), UCMP50403, PCB.
466
8 Melonis barleeanus (Williamson), UCMP50404, NAV5.

9 Pullenia bulloides (dOrbigny), UCMP50405, FRA.
10, 11 P
ullenia subcarinata (dOrbigny): 10, 4-chambered
variant (f. quinqueloba), UCMP50406, FRM. 11,
UCMP50407, FRA.
12 A
llomorphina pacifica Cushman and Todd, UCMP50408,
FRM.
Kenneth L. Finger
Plate 22
467
Type age and locality: Miocene, Romania.

Distinguishing features: Robust, subspherical test showing 34
chambers and pronounced flange-like apertural lip.
Comparative species: Hidina hidensis Popescu 1975 (Early
Miocene, Romania) and Hidina cubensis (= Allomorphina
cubensis Palmer and Bermdez 1936; Early Oligocene, Cuba)
are more ovate forms. Hidina globata (= Chilostomella globata
Galloway and Heminway 1941; Late Oligocene and Early
Miocene, Puerto Rico) is similarly globose, but the final whorl
comprises only two chambers.
Occurrence: Navidad Fm. (MAT, PPP, PTA), Ranquil Fm.
(FRM, MS10, RQT).
Family QUADRIMORPHINIDAE Saidova 1981
Quadrimorphina Finlay 1939b
Type species: Valvulina allomorphinoides Reuss 1860.
Quadrimorphina glabra (Cushman 1927g)
Plate 23, Figure 3
Valvulineria vilardeboana var. glabra Cushman 1927g, p. 161, pl.

4, figs. 5, 6.
Quadrimophina glabra (Cushman). KOHL 1985, p. 90, pl. 31, fig. 8.
Type age and locality: Recent, Pacific off Central America.

Occurrence: Navidad Fm. (PTA), Ranquil Fm. (RQT).
Family OSANGULARIIDAE Loeblich and Tappan 1964
Osangularia Brotzen 1940
Type species: Osangularia lens Brotzen 1940.
Osangularia culter (Parker and Jones 1865)
Plate 23, Figure 4
Planorbulina farcta var. ungeriana subvar. culter Parker and

Jones 1865, p. 382, 421, pl. 19, fig. 1.
Anomalina bengalensis Schwager 1866, p. 259, pl. 7, fig. 111.
Osangularia bengalensis (Schwager). LOEBLICH and TAPPAN 1987,
pl. 708, figs. 4, 5. WHITTAKER 1988, p. 134, pl. 18, figs. 16.
JONES 1994, p. 100, pl. 96, fig. 3.
Osangularia culter (Parker and Jones). ROBERTSON 1998, p. 238,
pl. 95, figs. 2, 3. HAYWARD, GRENFELL, SABAA, NEIL and
BUZAS 2010, p. 227, pl. 30, figs. 79. HOLBOURN, HENDER
SON and MACLEOD 2013, p. 386387.
Type age and locality: Recent, Northeast Pacific, 1482m.

Stratigraphic range: Late Oligocene to Recent.
Upper depth limit: Lower bathyal; based on modern UDL of
genus off Chile.
Remarks: Although the type figure of Anomalina bengalensis
Schwager 1866 (Pliocene, Car Nicobar) has 1.5X the number of
chambers in the outer whorl than Osangularia culter, I suspect
they are synonymous.
468
Occurrence: El Peral beds (NLP), Ranquil Fm. (FRA, MIB,

RAN, RQK, RQT), Lacui (PNH).
Family ORIDORSALIDAE Loeblich and Tappan 1984
Oridorsalis Andersen 1961
Type species: Oridorsalis westi Andersen 1961.
Oridorsalis umbonatus (Reuss 1851a)
Plate 23, Figure 5
Rotalina umbonata REUSS 1851b, p. 75, pl. 5, fig. 35.

Truncatulina tenera Brady 1884, p. 665, pl. 95, fig. 11.
Oridorsalis umbonatus (Reuss) RESIG 1981, pl. 8, fig. 8. KOHL
1985, p. 95, pl. 33, fig. 6. JONES 1994, p. 99, pl. 95, fig. 11; p. 104,
pl. 95, fig. 2. WHITTAKER 1988, p. 137, pl. 19, figs. 13. ZAPA
TA and CEAR 2004, p. 32, pl. 12, fig. 5. HAYWARD, GREN
FELL, SABAA, NEIL and BUZAS 2010, p. 227, pl. 30, figs. 36.
Oridorsalis tener (Brady) INGLE, KELLER and KOLPACK 1980, pl.
5, figs. 5, 6.
Oridorsalis tenera (Brady) LOEBLICH and TAPPAN 1987, pl. 708,
figs. 911.
Type age and locality: Eocene, Germany.

Occurrence: El Peral beds (NLP), Navidad Fm. (CPUP, NAV5,
PPP, PPT, PTA), Ranquil Fm. (FRA, FRM, MS10, RAN, RQK,
RQT), Santo Domingo Fm. (VAL), Lacui Fm. (PCB).
Family HETEROLEPIDAE Gonzles-Donoso 1969
Anomalinoides Brotzen 1942
Type species: Anomalinoides plummerae Brotzen 1942.
Anomalinoides salinasensis (Kleinpell 1938)
Plate 23, Figure 6
Anomalina salinasensis Kleinpell 1938, p. 347, pl. 13, fig. 1.

Anomalina ornata (Costa). MARTNEZ and PARADA 1968, pl. 1,
figs. 1517.
Anomalinoides salinasensis (Kleinpell). WHITTAKER 1988, p. 140,
pl. 23, figs. 1012. FINGER 1992, p. 88, pl. 35, figs. 3035.
Type age and locality: Middle Miocene, California.

Stratigraphic range: Early to Middle Pliocene.
Upper depth limit: Middle bathyal, as reported in California by
Ingle (1980).
Remarks: The Chilean form is identical to specimens from the
California Miocene, as noted in Kleinpells type description, it
is somewhat variable. The outer whorl of this form has 1011
chambers that maintain a fairly consistent height:width ratio.
The coil ranges from evolute to involute. Specimens with a nearly
involute low trochospire resemble Melonis barleeanus.
Comparative species: Anomalinoides globosa (= Anomalina
globulosa Chapman and Parr 1937; Recent, Southern Ocean)
is coarsely perforate and characterized by a depressed
trochospiral with about 7 chambers in the outer whorl; the type
figure, however, shows only the involute umbilical side and 9

chambers.
PPP, NAV5, MPUP, CPUP), Ranquil Fm. (LEB), Santo Domingo
Fm. (VAL), Lacui Fm. (PCB, PNH).
Family GAVELINELLIDAE Hofker 1956
Subfamily GAVELINELLINAE Hofker 1956
Gyroidina dOrbigny 1826
Type species: Gyroidina orbicularis dOrbigny 1826.
Gyroidina laevigata dOrbigny 1826
Plate 23, Figure 7
Gyroidina laevigata DOrbigny 1826, p. 278; figure in Parker,

Jones and Brady 1871, pl. 12, fig. 130; also see text-fig. of For
nasini 1898, p. 52.

Comparative species: Gyroidinoides nitida (= Rotalina nitida

Reuss 1844; Cretaceous, Europe) may be synonymous, but the
type figures are too small to confirm this, and the age discrepancy
suggests otherwise.
Occurrence: Navidad Fm. (PPP, NAV5), Ranquil Fm. (MIB,
RQK).
Hansenisca Loeblich and Tappan 1987
Type species: Gyroidina soldanii dOrbigny 1826.
Hansenisca altiformis (R. E. and K. C. Stewart 1930)
Plate 23, Figure 10
Gyroidina soldanii var. altiformis R. E. and K. C. Stewart 1930, p.

67, pl. 9, fig. 2.
Gyroidina neosoldanii BROTZEN 1936, p. 158, pl. 107, figs. 6, 7
(Recent; N and S Pacific Ocean).
Gyroidina altiformis R. E. and K. C. Stewart. INGLE, KELLER and
KOLPACK 1980, pl. 6, fig. 1. WHITTAKER 1988, p. 130, pl. 18,
figs. 1618.
Gyroidinoides altiformis (R.E. and K.C. Stewart). KOHL 1985, p. 95,
pl. 34, fig. 3. ROBERTSON 1998, p. 242, pl. 98, fig. 1.
Remarks: The Chilean forms are small with a circular outline

and about 10 slightly inflated, nearly equidimensional chambers
separated by slightly limbate, curved sutures.
Type age and locality: Early Pliocene, California.
Comparative species: Gyroidina laevis (= Rotalia laevis

dOrbigny 1852; Tertiary, localities given in Italy) has curved
sutures with chambers that are about twice as high as broad.
Gyroidina orbicularis dOrbigny 1826 (Recent, Adriatic Sea)
has 6 outer chambers that are 23 times wider than high. The
outer whorl of Hansenisca soldanii (Pl. 22, Fig. 11) has a deeper
umbilicus and chambers on the spiral side are much broader than
high, and separated by depressed sutures.
Upper depth limit: Generally outer neritic.
Occurrence: El Peral beds (NLP), Ranquil Fm. (FRA, MIB,

RAN), Lacui Fm. (PCB, PNH).
Gyroidina sp.
Stratigraphic range: Late Oligocene to Recent.

Distinguishing features: Partly concave spiral surface with
backward-tilted dorsal faces of chambers, and oblique, limbate
sutures.
Occurrence: Navidad Fm. (all except RAP and PPN), Ranquil
Fm. (all except FRA), Santo Domingo Fm. (VAL), Lacui Fm.
(CHO, CUC).
Maximum relative abundance: Common (NAV5, VAL).
Hansenisca soldanii (dOrbigny 1826)
Plate 23, Figure 11
Remarks: Test differs from other Gyroidina by its slightly

concave spiral side with depressed umbilicus and very slightly
depressed, nonlimbate sutures. It might be an aberrant G.
laevigata, as the number of chambers and test size are similar.
Gyroidina soldanii DORBIGNY 1826, p. 278; type-figure not desig

nated. INGLE, KELLER and KOLPACK 1980, pl. 7, figs. 12, 13.
Gyroidinoides soldanii (dOrbigny). MARTNEZ and PARADA 1968,
pl. 1, figs. 1, 2. JONES 1994, p. 106, pl. 107, figs. 6, 7. HOL
Hansenisca soldanii (dOrbigny). LOEBLICH and TAPPAN 1987, p.
106, pl. 719, figs. 59.
Gyroidinoides Brotzen 1942

Type species: Rotalina nitida Reuss 1844
Plate 22, Figure 8
Gyroidinoides umbonatus (A. Silvestri 1898)
Plate 23, Figure 9
Rotalia soldanii var. umbonata A. SILVESTRI 1898 , p. 329, pl. 6, fig.

14.
Gyroidina nipponica Ishizaki 1944, p. 102, pl. 3, fig. 3. (Early
Pliocene, Japan)
Type age and locality: Early Pliocene, Italy.

Stratigraphic range: Early Miocene to Early Pliocene.
Remarks: DOrbignys earliest representation of this species is

modle 36, which he based upon the figure of Soldani (1789),
and which Parker, Jones and Brady (1871) later illustrated as a
planoconvex form with an ultimate whorl of 8 chambers on the
spiral side (vs. 9 on the umbilical side) with a sutural progression
from oblique to radial. DOrbignys (1846) publication on the
Middle Miocene foraminifera of the Vienna Basin shows the
species with 9 chambers in both spiral and umbilical views,
but the spiral sutures are radial. There is enough variability in
sutural obliquity among the Chilean specimens to indicate that
this is not a reliable criterion to distinguish H. soldanii from
469
related species; on the spiral side, the diagnostic features are the
rounded surface of the outer whorl and the progressive elongation
of its chambers.
Comparative species: Whittaker (1988: pl. 15, figs. 1618)
identified a biconvex form, lenticular in edge view, with nearly
radial sutures in the Late OligocenePliocene of Ecuador as
Gyroidina soldani (dOrbigny). Hansensica nitidula (= Rotalia
nitidula Schwager 1866; Pliocene, Car Nicobar) is probably
synonymous.
Occurrence: El Peral beds (LPER), Ranquil Fm. (FRA, FRM,
RAN, RQK, RQT), Lacui Fm. (CHE).
Hanzawaia Asano 1944
Type species: Hanzawaia nipponica Asano 1944.
Hanzawaia cf. H. nipponica Asano 1944
Hanzawaia nipponica Asano 1944, p. 98, pl. 4, figs. 1, 2.

Upper depth limit: Inner neritic (for H. basiloba in Ingle 1980).
Remarks: This form differs from Hanzawaia nipponica Asano
by having less broadly curved sutures and by not having the
earlier spiral completely obscured by coalesced umbilical flaps.
Comparable species: Hanzawaia mantaensis (= Anomalina
mantaensis Galloway and Morrey 1929; probably Late Eocene,
Ecuador) is inflated and has limbate sutures. Hanzawaia lucida
Saidova 1975 (Recent, off New Guinea) is even more inflated
with a thickness half the diameter, and the sutures are thicker
and more broadly curved.
Hanzawaia strattoni (Applin 1925)
Plate 24, Figure 3
Truncatulina americana var. strattoni Applin, in Applin, Ellisor

and Kniker 1925, p. 99, pl. 3, fig. 3.
Hanzawaia concentrica (Cushman). KOHL 1985, p. 98, pl. 36, fig. 1.
Type age and locality: Miocene, Louisiana.

Stratigraphic range: MioceneRecent.
Upper depth limit: Inner neritic (for H. basiloba in Ingle 1980).
Occurrence: Navidad Fm. (LBZ, MAT, NAV5, PPN, RAP),
Ranquil Fm. (FRA, LEB, MIB), Santo Domingo Fm. (VAL),
Lacui Fm. (CHO, CUC, PCB, PNH).
Maximum relative abundance: Common (RAP, PPN, CHO).
Family TRICHOHYALIDAE Saidova 1981
Buccella Andersen 1961
Type species: Eponides hannai Phleger and Parker 1951.
Buccella peruviana (dOrbigny 1839c)
Plate 24, Figures 4, 5.
Rotalina peruviana DORBIGNY 1839c, p. p. 41, pl. 1, figs. 1214.

Buccella peruviana (dOrbigny). BOLTOVSKOY and THEYER 1970,
p. 310, pl. 1, fig. 19. ZAPATA and CEAR 2004, p. 18, pl. 2, fig. 7.
Calvo-Marcilese and Langer 2012, figs. 2A2O.
Type age and locality: Recent; localities noted off Ecuador,

Peru, and Chile.
Upper depth limit: Inner neritic (Zapata and Cear 2004).
Remarks: The species is highly variable in morphology (CalvcoMarcilese and Langer 2012).
Comparative species: Buccella frigida (Cushman) emend.
Andersen 1952 is noncarinate and ovate in edge view, but
considered a synonym by Calvo-Marcilese and Langer (2012).
Buccella sinulata McCulloch 1977 (Recent, East Pacific) has
9 chambers in the last whorl, and the umbilical sutures are
curved. Buccella tenerrima (= Rotalia tenerrima Bandy 1950;
PLATE 23
Figs. 4a, 6c, 7c, 10c, and 11c are photomicrographs; all other images are SEMs. Scale bars in m.
1 Chilostomella ovoidea Reuss, UCMP50409, RQT.
7 Gyroidina laevigata dOrbigny, UCMP50415, FRA.
2 Hidina variabilis Gheorghian, Iva and Gheorghian,

UCMP50410, MIB.
8 Gyroidina sp., UCMP50416, MIB.
3 Quadrimorphina glabra (Cushman), UCMP50411, RQT.

4 Osangularia culter (Parker and Jones), UCMP50412, FRA.
10 H
ansenisca altiformis (R. E. and K. C. Stewart), UCMP50418,
PPP.
5 Oridorsalis umbonatus (Reuss), UCMP50413, FRA.
11 Hansenisca soldanii (dOrbigny), UCMP50419, FRA.
6 Anomalinoides salinasensis (Kleinpell), UCMP50414, CPUP.
470
9 Gyroidinoides umbonatus (A. Silvestri), UCMP50417, NAV5.
Kenneth L. Finger
Plate 23
471
Pleistocene, Oregon) has 810 chambers in the last whorl, and

peripheral ends of the sutures are conspicuously tuberculate.
Occurrence: El Peral beds (NLP, LPER), Navidad Fm. (LBZ,
MAT, NAV5, PPN, PPP, PTA, RAP), Ranquil Fm. (FRA, RAN,
RQK, RQK), Lacui Fm. (CHE, CUC).
Maximum relative abundance: Common (RQK).
Superfamily ROTALIOIDEA Ehrenberg 1839
Family ELPHIDIIDAE Galloway 1933
Subfamily ELPHIDIINAE Galloway 1933
Cribroelphidium Cushman and Brnnimann 1948
Type species: Cribroelphidium vadescens Cushman and Brn
nimann 1948.
Cribroelphidium hauerinum (dOrbigny 1846)
Plate 24, Figure 7
Polystomella hauerina DORBIGNY 1846, p. 122, pl. 6, figs. 1, 2.

Elphidium hauerinum (dOrbigny). PAPP and SCHMID 1985, p. 49,
pl. 38, figs. 510.

Basin, Austria.
Comparative species: Cribroelphidium trinitatensis Cushman
and Brnnimann 1948 (Recent, Trinidad and Tobago, mangrove
swamp) and C. irrasum McCulloch 1977 (Recent, East Pacific)
are probably synonymous. Cribroelphidium incertum var.
clavatum Cushman 1930b (Recent, off Maine) emend. Loeblich
and Tappan 1953 is more biconvex with curved sutures.
Occurrence: Lacui Fm. (CHE).
Elphidium de Montfort 1808
Type species: Nautilus macellus var. Fichtel and Moll 1798.
Elphidium macellum (Fichtel and Moll 1798)
Plate 24, Figure 9
Nautilus macellus var. Fichtel and Moll 1798, p. 68, pl. 10,
figs. h, i, k; type-figures also in RGL and HANSEN 1984, pl. 14,
fig. 2, pl. 15, figs. 1, 2; p. 51 tf. 18B.

Elphidium granti KLEINPELL 1938, p. 238, pl. 19, figs. 1, 11. (Mio
cene, California)
Elphidium pilasense MCCULLOCH 1977, p. 223, pl. 98, fig. 3. (Re
cent, N Pacific)
Type age and locality: Recent, Mediterranean.

Occurrence: Navidad Fm. (NAV5), Lacui Fm. (CHE).
Subfamily ROTALIINAE Ehrenberg 1839
Rotalinoides Saidova 1975
Type species: Rotalia papillosa Brady 1884.
Rotalinoides margaritifera (Brady 1881)
Plate 24, Figures 6, 8, 10
Planorbulina (Truncatulina) margaritifera BRADY 1881, p. 66; typefig. in BRADY 1884, pL. 96, fig. 2 (as Truncatulina margaritifera).
Rotalia papillosa var. compressiuscula Brady 1884, p. 708, pl. 107,
fig. 1, pl. 108, fig. 1. (Recent, multiple localities given in Indopacific
ranging 911448m)
Rotalinoides compressiusculus (Brady). JONES 1994, p. 106, pl. 107,
figs. 1, 3.
Type age and locality: Recent, off the Philippines, 174183m.

Remarks: Bradys two species synonymized here were both
described from the Recent Indopacific region and appear to be
variants of the same species. The Chilean forms include beaded
and nonbeaded varieties.
Comparative species: Rotalinoides crassimura (= Notorotalia
crassimura Carter 1958; Late Oligocene, Australia) has a very
prominent umbilical boss.
Occurrence: Navidad Fm. (PPN), Ranquil Fm. (LEB), Santo
Domingo Fm. (VAL).
PLATE 24
1, 2 Hanzawaia cf. H. nipponica Asano, UCMP50420, PPP.

3 Hanzawaia strattoni (Applin), UCMP50421, RAP.
4, 5 B
uccella peruviana (dOrbigny): 4, UCMP50422, PPN.
5, UCMP50423, PTA.
6, 8, 10 R
otalinoides margaritifera (Brady): 6, UCMP50424, PPN. 8, UCMP50425, VAL. 10, UCMP50426,
LEB.
472
7 C
ribroelphidium hauerinum (dOrbigny), UCMP50427,
CHE.
9 Elphidium macellum (Fichtel and Moll), UCMP50428,
CHE.
Kenneth L. Finger
Plate 24
473
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Miocene Foraminifera of South-Central Chile

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Miocene Foraminifera of South-Central Chile

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Miocene foraminifera from the

south-central coast of Chile

In comparison to most other regions, western South America is

a Late OligoceneMiddle Miocene age, and the gastropod fauna

Kenneth L. Finger: Miocene foraminifera from the south-central coast of Chile

The initial goal of this study was to produce a monograph on

Charles Darwin was first to explore the geology of this region,

~140km southwest of Santiago, as the Formation of Navidad,

Micropaleontology, vol. 59, nos. 45, 2013

Tavera (1968, 1979) confined the Navidad Formation to the

suggested it was Late Miocene. Osorio (1978) later detailed the

Kenneth L. Finger: Miocene foraminifera from the south-central coast of Chile

Their study was the first on benthic foraminifera in the regional

The Ranquil Formation crops out mostly along the Arauco

Encinas et al. (2006, 2007) and Finger et al. (2007) analyzed

Micropaleontology, vol. 59, nos. 45, 2013

Kenneth L. Finger: Miocene foraminifera from the south-central coast of Chile

Early Miocene shark teeth (Surez, Encinas and Ward 2006),

Micropaleontologists working the foraminifera-rich land-based

Deep-sea cores generally provide the most complete and

Two depth zonations have been widely used in foraminiferal

Micropaleontology, vol. 59, nos. 45, 2013

Kenneth L. Finger: Miocene foraminifera from the south-central coast of Chile

Micropaleontology, vol. 59, nos. 45, 2013

1980), and the other presented by van Morkhoven, Berggren

ranges of modern species to the fossil record using the priniciple

Ingle (1980) based his foraminiferal depth zonation on the most

Natland and Kuenen (1951) and Phleger (1951) established

It is important to recognize that all depth zonations are only

Benthic foraminifera can be valuable tools for deciphering

The reliability of this method has been scrutinized and criticized

Kenneth L. Finger: Miocene foraminifera from the south-central coast of Chile

This study originally focused on 46 samples from 32 localities

Micropaleontology, vol. 59, nos. 45, 2013

Paleontology) microfossil locality number. Most of the samples

CHE: Lacui Formation. Yellowish sandstone from coastal bluffs,

Kenneth L. Finger: Miocene foraminifera from the south-central coast of Chile

CUC: Lacui Formation. Grey sandstone, coastal bluff blockfall

platform, Punta Fraile, west of Tubul, Arauco Peninsula, Arauco

Micropaleontology, vol. 59, nos. 45, 2013

6km north of the mouth of Estero Maitenlahue, just south of

715046W. Collected by S. N. Nielsen, 2000 & 2001. UCMP

MS10: Ranquil Formation. Grey siltstone on Isla Quiriquina,

Kenneth L. Finger: Miocene foraminifera from the south-central coast of Chile

by S. N. Nielsen 2000 & 2001 and K. L. Finger 2003. UCMP

335623S, 71514W. Collected by S. N. Nielsen 2002 and K.

RAN: Ranquil Formation. Brown massive sandstones with

PTA: Navidad Formation. Fossiliferous lens of grey siltstone

Micropaleontology, vol. 59, nos. 45, 2013

RQK: Ranquil Formation. Highly fossiliferous sandstone

Foraminifera were processed from 46 sedimentary rock samples

The primary resources initially used in this study for identifying

Kenneth L. Finger: Miocene foraminifera from the south-central coast of Chile

1986), Benthic Cenozoic foraminifera from Ecuador (Whittaker

foraminifera of Chile was also perused for this purpose (see

Micropaleontology, vol. 59, nos. 45, 2013

(Oligocene and Miocene), Mediterranean (MioceneRecent) Car

Marchant, Zapata and Hromic (2007) provide a thorough

The earliest report on the modern foraminifera off central Chile

Kenneth L. Finger: Miocene foraminifera from the south-central coast of Chile

auris, Valvulina [Cancris] inflata, Bulimina pulchella, Bulimina

from depths of 1170m. Nearly half of the species illustrated in

1 Bathysiphon giganteus Cushman, UCMP50000, RAP.