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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 33º and 44ºS (text-figs. 1–5). Age interpretations of the units have ranged widely in the literature (text-figs. 6 and 7). Molluscan and strontium isotope data indicate

a Late Oligocene–Middle 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.

Miocene foraminifera from thesouth-central coast of Chile

Kenneth L. Finger

University of California Museum of Paleontology 1101 Valley Life Sciences Building, Berkeley, CA 94720-4780 USA

email: [email protected]

ABSTRACT: Foraminifera are abundant and diverse in the Neogene sedimentary units of south-central Chile (34–43ºS), 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 Concep-cíon, and (3) the Lacui Formation on and near Chiloé Island. Also included in this study are two assemblages from deposits of uncertain affinity in San Sebastián (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 Chiloé Island). The 27 localities report-ed 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 microfos-sils 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 Sebastián 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 informal-ly 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, Pseudolin-gulina 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 foramin-ifera 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 incorpo-rated into future studies on benthic foraminiferal biostratigraphy and biogeography.

Micropaleontology, vol. 59, nos. 4–5, pp. 341–492, text figures 1–18, tables 1-8, plates 1-24, 2013

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Kenneth L. Finger: Miocene foraminifera from the south-central coast of Chile

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,

~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 Martínez-Pardo and Osorio (1964) noted that it does not match Darwin’s 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. Möricke (1896) was the first to recognize the molluscan fauna as Miocene.

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 2–5.

TEXT-FIGURE 2 Las Cruces area (Lo Abarca Formation) collecting sites. Neogene strata are exposed in shaded areas.

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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). Martínez-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. Martínez-Pardo and Valenzuela (1979) later reported that the Navidad Formation had discoasters indicative of the Middle Miocene. As a result of these studies, Martínez-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).

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.

TEXT-FIGURE 4 Arauco area (Ranquil Formation) and Valdivia (Santo Domingo Forma-tion) collecting sites. Neogene strata are exposed in shaded areas. Not plotted is RQS, which yielded few foraminifers.

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The Ranquil Formation crops out mostly along the Arauco Peninsula, ~60km SSW of Concepción (text-fig. 4). Tavera (1942) and García (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 Chiloé 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, Muñoz and Stern 2000).

Martínez and Parada (1968) used benthic foraminifera to place the San Sebastián beds, referred to in this report as the El Peral beds, in the Pliocene, as they did not recover any planktic species.

Their study was the first on benthic foraminifera in the regional Neogene, and they interpreted the assemblage as indicative of bathyal deposition at 200–500m.

Martínez 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 Martínez (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 1233–1235 on the Chile margin (forearc). Unfortunately, drilling did not extend below the very thick Pleistocene section (Tiedemann and Mix 2007).

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 (33-45°S), 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 deep-water 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 5 Chiloé area (Lacui Formation) collecting sites. Neogene strata are ex-posed in shaded areas.

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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.

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Early Miocene shark teeth (Suárez, 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, Gutiérrez 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 22.0–22.5 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 frameworkMicropaleontologists 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

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).

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 36°S. 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 Chiloé Island. The other three sites are on the Chilean margin (forearc): Site 1233 (41.00°S, 74.45°W) is 92km NE of Chiloé Island, and Sites 1234 (36.22°S, 73.68°W) and 1235 (36.16°S, 73.57°W) are 86km NE of Concepcíon. 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 859–863 at the Chile Triple Junction (c. 46°S, 75°W), none of which drilled deeper than the upper Pliocene (Spiegler and Müller 1995).

The nearest Miocene deep-sea core sections were obtained from the tropical zone off Peru (c. 9–21°S) at the Nazca Plate, Nazca Ridge, and East Pacific Rise (DSDP sites 319–321, IODP Leg 142, and ODP sites 1235–1237). The stratigraphy at ODP Site 1237 (16.00°S, 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 (13°1.0380’S, 101°31.4580’W, 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 modelsTwo 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 7 Distribution of age determinations for the units derived from data in text-figure 1.

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TEXT-FIGURE 8 Map showing locations of ODP sites off Peru and Chile between 7ºS and 50º S relative to the present region of study.

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TEXT-FIGURE 9 Neogene time scale with planktic foraminiferal zonations and datums. Species in bold were found in the Chilean samples. Abbreviations of gen-era: 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.=Praeorbu-lina. 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.

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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 south-central Chile.

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 400–900m, another oxygen minimum zone from 900–1700m 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.

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 indicatorsBenthic 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 Natland’s (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 Natland’s recognition of mixed-depth 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

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.

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.

The reliability of this method has been scrutinized and criticized in numerous publications. Several authors have correlated large-scale 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

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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 0–70% P at a depth of 100m, and 60–100% 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 Chiloé 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 localitiesThis study originally focused on 46 samples from 32 localities along the central Chilean coast south of Valparaiso (~33°00’S), from Las Cruces (~34°30’S) to Cucao (~42°40’S) (text-figs. 1−5). Alfonso Encinas, Sven Nielsen, and I collected nearly all of the outcrop samples during 2000–2003. During the course of this study, I also examined hundreds of micropaleontological slides representing 25 wells drilled by ENAP (Empresa Nacional del Petróleo [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, 4−8, 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 10 Results of linear regression analyses. A, number of specimens vs. spe-cies 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.

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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, Valparaíso Region, 33°47’11”S, 71°47’42”W. Collected by A. Encinas, 2003. UCMP MF9003.

CHE: Lacui Formation. Yellowish sandstone from coastal bluffs, south of Chepu, Isla Chiloé (see Watters and Fleming 1972), Chiloé Province, Los Lagos Region, 42°04’26”S, 74° 02’1”W. 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, 41°44’43”S, 73°50’36”W. 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 O’Higgins Region, 34°02’52”S, 71°50’36”W. Collected by A. Encinas, 2004. UCMP MF9016.

TEXT-FIGURE 11 Dendrogram from Q-mode cluster analysis using Ward’s 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.

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CUC: Lacui Formation. Grey sandstone, coastal bluff blockfall south of Cucao, Isla Chiloé, Chiloé Province, Los Lagos Region, 42°42’28”S, 74°08’10”W. 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, Biobío Region, 37°12’13”S, 73°29’27”W. Collected by S. N. Nielsen, 2000–2002. UCMP MF9018.

FRM: Ranquil Formation. Grey mudstone to siltstone, basal part of exposure concordantly subjacent to FRA, intertidal

platform, Punta Fraile, west of Tubul, Arauco Peninsula, Arauco Province, Biobío Region, 37°12’13”S, 73°29’27”W. Collected by S. N. Nielsen, 2000–2002. 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 O’Higgins Region, 33°55’43”S,

TEXT-FIGURE 12 Dendrogram from R-mode cluster analysis using Ward’s method and a presence-absence matrix for all 27 assemblages, based on same data matrix as text-figure 11.

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71°50’46”W. 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, Biobío Region, 37°35’39”S, 73°38’16”W. 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 Sebastián and Las Cruces, just below the bend (Laguna de los Patos locality of Martínez and Parada 1968), San Antonio Province, Valparaíso Region, 33º30’12’’S, 71º36’26”W. Portion of Martínez and Parada’s 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 O’Higgins Region, 33°56’58”S, 71°52’05”W. Collected by S. N. Nielsen 2001 & 2002; and K. L. Finger 2003. UCMP MF9013.

MAT: Navidad Formation. Small lens of glauconitic sandstone 2–3m above base of coastal bluff and approximately 2m stratigraphically above MAP, Los Goterones (about 1km north of Matanzas), Cardinal Caro Province, Libertador General Bernardo O’Higgins Region, 33°56’55”S, 71°51’59”W. Collected by S. N. Nielsen, 2000–2002. 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, Biobío Region, 37°32’28”S, 73°36’46”W. 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

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, Valparaíso Region, 33°46’49”S, 71°47’38”W. 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 O’Higgins Region, 33°54’39”S, 71°50’38”W, Collected by S. N. Nielsen 2000 & 2001 and K. L. Finger 2003. UCMP MF9015.

MS10: Ranquil Formation. Grey siltstone on Isla Quiriquina, entrance to Bahía de Concepción, Biobío Region. Collected by J. Tavera. Locality details not available (approximate center of island is 36°37’38”, 73°30’38”). Sample obtained by A. Encinas from the Museo Nacional de Historia Natural, Santiago. UCMP MF9017.

NAV5: Navidad Formation. Navidad #5 well, Matanzas, Cardinal Caro Province, Libertador General Bernardo O’Higgins Region, 33°57’30”S, 71°52’23”W. Foraminiferal residues from well depths of 24–30 and 30–36m, 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 Sebastián 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, Valparaíso Region, 33°30’04”S, 71°36’26”W. 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 Chaumán, Isla Chiloé, Chiloé Province, Los Lagos Region, 41°55’44”S, 74°01’39”W. 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 Puñihuil, Isla Chiloé, Chiloé Province, Los Lagos Region, 41°55’44”S, 74°01’39”W. 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 Caro Province, Libertador General Bernardo O’Higgins Region, 33°54’38”S, 71°50’41”W. 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 O’Higgins Region, 33°54’23”S, 71°50’18”W. Collected by S. N. Nielsen 2000–2002. 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 O’Higgins Region, 33°54’15”S, 71°50’13”W. Collected by S. N. Nielsen, 2000–2002 and K. L. Finger, 2003. UCMP MF9005.

PPS: Navidad Formation. Coarse grey sandstone lens, coastal bluff, Punta Perro, Cardinal Caro Province, Libertador General Bernardo O’Higgins Region, 33°54’39”S, 71°50’38”W. Collected

TEXT-FIGURE 13 Dendrogram from Q-mode cluster analysis using Ward’s 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.

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by S. N. Nielsen 2000 & 2001 and K. L. Finger 2003. UCMP MF9009.

PPT: Navidad Formation. Top of grey siltstone interval, approx-imately 15m stratigraphically above PPP, coastal bluff, Punta Perro, Cardinal Caro Province, Libertador General Bernardo O’Higgins Region, 33°54’16”S, 71°50’9”W. Collected by A. Encinas, 2006. UCMP MF9006.

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 O’Higgins Region,

33°56’23”S, 71°51’4”W. Collected by S. N. Nielsen 2002 and K. L. Finger 2003. UCMP MF9011.

RAN: Ranquil Formation. Brown massive sandstones with intermittent beds of glauconitic sandstone, overlying RQT and transected by RQS, coastal bluff of Punta Huenteguapi, Ranquil, Arauco Peninsula, Arauco Province, Biobío Region, 37°30’25”S, 73°35’28”W. Collected by S. N. Nielsen 2000–2002 and K. L. Finger 2003. UCMP MF9023.

RAP: Navidad Formation. Grey, reddish-brown and dark-brown sandstones in an undifferentiated blockfall from steep cliffs along the coast north of Río Rapel, San Antonio Province, Valparaíso Region, 33°53’20”S, 71°49’34”W. Collected by S. N. Nielsen, 2000 & 2002. UCMP MF9004.

TEXT-FIGURE 14 Principle coordinates plot from NMDS analysis showing the relationships between localities based on the 52 dominant (common and abundant) spe-cies occurrences.

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RQK: Ranquil Formation. Highly fossiliferous sandstone boulders restricted to the northernmost part of the beach, Punta Huenteguapi, Arauco Peninsula, Arauco Province, Biobío Region, 37°30’20”S, 73°35’26”W. 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, Arauco Peninsula, Arauco Province, Biobío Region, 37°30’18”S, 73°35’24”W. Collected by K. L. Finger 2003. UCMP9020.

RQT: Ranquil Formation. Grey mudstones to siltstones like FRM, intertidal platform of Punta Huenteguapi, Ranquil, Arauco Peninsula, Arauco Province, Biobío Region, 37°30’18”S, 73°35’24”W. 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, 39°55’43”S, 73°07’32”W. Collected by S. N. Nielsen, 2001. UCMP MF9026.

METHODS

Sample processingForaminifera 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-µm openings) sieve and (3) drying by funneling through fast-flow filter paper, followed by oven-drying at 30˚C. 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 procedureThe 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 Brönnimann 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

TEXT-FIGURE 15 Principle coordinates plot from NMDS analysis showing relationships between species based on same data matrix as text-figure 14.

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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

foraminifera of Chile was also perused for this purpose (see following subsection).

Many of the benthic species identified in this study were orig-inally described from the Pacific (Oligocene–Recent), Caribbean

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.

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(Oligocene and Miocene), Mediterranean (Miocene–Recent) Car Nicobar Island in the Andaman Sea (Pliocene), and the Vienna Basin of the Central Paratethys (Oligocene–Miocene). A few species were first described from the Atlantic and polar regions, or from pre-Oligocene strata.

Previous taxonomic studies on the modern Chilean faunaMarchant, 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; d’Orbigny 1839c). Of the 81 new species described in that study, 56 were from the Atlantic and 25 were from the Pacific. D’Orbigny’s only sample taken off Chile is from Bahía de Valparaiso (33˚S), 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]

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.

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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 d’Orbigny (1839c) from the southwest Atlantic — one from Patagonia and eight from the Falkland Islands.

Additional species were documented in H. B. Brady’s (1884) tome on the foraminifera collected by the Challenger Expedition (1873–1876). That global venture included sample stations west of Chiloé and throughout the archipelago of southern Chile. Egger (1893) subsequently worked on samples from the Gazelle Expedition (1874–1876) that were collected off northern Chile (north of Valparaiso). Bandy and Rodolfo (1964) studied 32 trawl and core samples from depths of 179–6250m off Peru and Chile, but only as far south as Valparaiso (32.3ºS). 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 44–260m off central Chile (29º57’–42º16’S). 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 82–2286m on the northern Nazca plate (0–27ºS) and on the continental margin from 2–20ºS. More pertinent to the present study is the analysis by Ingle, Keller and Kolpack (1980) of bottom samples collected from depths of 135–4500m along three transects off central Chile (31.5–39.2ºS).

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 (33º41’S, 78º50’W), 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 (18º28’–31º56’S), having documented 151 species

from depths of 1–170m. 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 125–3485m in the south-central Chilean province (i.e., from Valparaiso to the southern end of Chiloé Island).

Taxonomic problemsSubjective 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 d’Orbigny’s concept of foraminiferal taxonomy. Attitudes changed when H. B. Brady’s (1884) Challenger tome recognized many of d’Orbigny’s 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,

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).

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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 studyIn 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.

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 statisticsSpecies 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 α 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). Fisher’s α is a diversity index defined by S = αln(1 + n/α), 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 Ward’s method, in the cluster analyses, and the correlation coefficient (Pearson’s 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 (UCMP50438-50499), 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 (165–2133; mean 610, median 471) and benthic species richness (20–138; mean 60, median 63). Planktic:benthic (P:B) ratios range 0–0.68 (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

TABLE 1Comparison of bathymetric zonation schemes.

TABLE 2Early foraminiferal monographs and their latest revisions, and the num-ber of their benthic species recognized in this study.

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TABLE 3Relative abundance of benthic species vs. localities checklist. VR = 1 specimen; R ≥2 specimens if <1%, or 2 specimens if ≥1%; F = 1–5%; C = 5–25%; A = 25–50%; VA >33%.

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TABLE 3Continued.

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TABLE 3Continued.

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TABLE 3Continued.

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TABLE 3Continued.

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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).

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 (5–25%) 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).

Benthic species previously known from Miocene–Holocene 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

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.

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 α values are within the normal range of 5–16 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 α (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 α correlated much better with species richness than with species number, possibly because the richness value is a factor of the number, not vice versa.

TABLE 4Planktic species presence vs. localities checklist. Relative abundances: VR = 1 specimen; R ≥2 specimens if <1%, or 2 specimens if ≥1%; F = 1–5%; C = 5–25%; A = 25–50%; VA >33%.

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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 latter’s 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 unitsIn Peru, Navidad molluscs such as Miltha vidali, Acanthina katzi, Olivancillaria claneophila, and Testallium cepa occur in Late Oligocene–Middle Miocene sections (Nielsen et al. 2003), but not in younger successions (DeVries and Frassinetti 2003).

Other Navidad species such as Ficus distans are restricted in Peru to the Early–Middle Miocene, whereas Eucrassatella ponderosa, Glycymeris ibariformis, and G. colchaguensis only occur in the Late Oligocene–Early Miocene. Encinas (2006) obtained Early Miocene ages of 24.7±0.4 and 20.4±0.5 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 5Numerical and statistical data tally for the 28 foraminiferal assemblages.

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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 Early–Middle Miocene age. I subsequently followed his suggestion that I peruse George Scott’s 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

TABLE 6Dominant (common and abundant) species in each assemblage.

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TABLE 7Upper-depth limits of 383 foraminiferal taxa living off the central Chilean margin between 33–44ºS. 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).

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TABLE 7Continued.

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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 Scott’s 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 Oligocene–Early Miocene interval. Suárez, 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 (Suárez 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 (22.2–15.9 Ma); the exception yielded an age of 11.06±0.19 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 12.2–11.3 Ma (Serravallian) in the equatorial Pacific (Barron, 2003). That interval encompasses the Sr age of 11.5±1.0 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 14.2–11.6 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 Parada’s interpretation of the LPER locality as Pliocene, which they based on benthic foraminifera.

For the Navidad Formation in its type area along Punta Perro, Martínez-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 Ibaraki’s report, even though it was associated with a subtropical molluscan assemblage recorded by Covacevich and Frassinetti (1980), which suggests that it preceded the global Mid-Miocene cooling event. Tsuchi (2002) correlated the molluscs with one of the relative abundance spikes of warm-water planktic

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.

Gutiérrez 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). Gutiérrez 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 (N4A–N5 FAD), Globoquadrina dehiscens (N4 FAD), transitional forms between Paragloborotalia nana (N6 LAD), Neogloboquadrina continuosa, (N6 FAD), Pg. bella (N4–N8), and the Pg. zealandica group (N5–N7). For the El Peral beds they are Orbulina universa (N9 FAD), Globorotalia miotumida (N9 FAD), and

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TABLE 8Paleobathymetric 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).

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Glr. praemenardii (N10–N13). 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, morpho-statistical 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. nana–Ng 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 long-ranging 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 (text-fig. 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 Oligocene–Miocene interval. In summary, 16 localities yielded Sr-isotope ages that at least partly overlap the concurrent range

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 paleoenvironmentPreviously, 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 (33ºS), Cabo Carranza (36ºS), and Valdivia (39ºS), 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.

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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. 11–15), 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 d’Orbigny (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 179–6250m off Peru and Chile, but only as far south as Valparaiso (32.3ºS). Ingle, Keller and Kolpack (1980) analyzed the fauna in bottom samples collected from depths of 135–4500m along three transects off central Chile (31.5–39.2ºS). Resig (1981) analyzed 121 core-top samples taken from depths of 82–2286m on the northern part of the Nazca plate (0–27ºS) and on the continental margin off Guayaquil, Ecuador (2ºS) to Iquique (20ºS), 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 (33º41’S, 78º50’W), 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 (18º28’–31º56’S). They documented 151 species from depths of 1–170m, 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 Chiloé, Figueroa et al. (2005, 2006) recorded 117 species of calcareous benthic foraminifera from multicores taken at depths of 125–3485m. 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 15–25 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 4–33 bathyal species with an average of 16; the numbers of middle or lower bathyal indicators per assemblage range 1–21 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 1900–2200m for normal slopes and between 2800–3600m 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 (1500–2000m) or lower bathyal (2000–4000m) 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,

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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 faunasThere 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 Whittaker’s (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.

BiogeographyBiogeographic 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 Cushman’s 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 Oligocene–Holocene, and their regional distributions are widely scattered across the globe. Although many of the modern offshore Chilean species or their homeomorphs are

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, nutrient-enriched subpolar water is transported northward by the Peru-Chile Current (PCC). The Coastal Current (CC) also flows northward but is significantly affected by an admixture of low-salinity waters from the Chile fjord region. In between them, 100–300km offshore, the Peru-Chile Counter Current (PCCC) transports subtropical surface water to the south. The poleward-flowing Gunther Undercurrent underlies these surface-water masses at depths of 100–400m and transports relatively low-oxygen and high-salinity water masses southward along the shelf edge. At depths of 400–1000m 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 well–documented 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 deep–water 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

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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 Petróleo) 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 obli-quispina, 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 sectionFor 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).

kenfinger

Sticky Note

quispinata

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Remarks: Distinguishing features or comments about the Chilean specimens, but excluding comparisons with other species, which are the next category.

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.

Occurrence: Refers to the geologic units and their localities where the species is recorded in this study (see text-figures 1–5).

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 1925Kingdom RhizARiA Cavalier-Smith 2002Superphylum REtARiA Cavalier-Smith 2002Phylum FoRAMiNiFERA (d’orbigny 1826) Lee 1990Class PoLythALAMEA Ehrenberg 1838order AStRoRhiziDA Brady 1881Family BAthySiPhoNiDAE Avnimelech 1952BAthySiPhoN M. Sars, in G.o. Sars 1872Type species: Bathysiphon filiformis M. Sars, in G.o. Sars 1872.

Bathysiphon giganteus CushmanPlate 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.

Upper depth limit: Lower bathyal; based on Bathysiphon sp. recorded off Chile by Bandy and Rodolfo (1964) and ingle, keller and kolpack (1980).

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.

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 fine-grained 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 1869Plate 1, Figures 4, 5

PLATE 1Figures 4–6, 15, 17, and 22 are photomicrographs; all other images are SEMs. Scale bars in µm.

1 Bathysiphon giganteus Cushman, uCMP50000, RAP.

2, 3 Bathysiphon sp.: 2, uCMP50001, PCB; 3, uCMP50002, FRA

4, 5 Rhabdammina abyssorum Sars, PPP: 4, uCMP50003. 5, uCMP50004.

6 Ammodiscus incertus (d’orbigny), uCMP50005, FRM.

7 Reophax agglutinatus Cushman, uCMP50006, FRA.

8 Reophax sp., uCMP50007, VAL.

9 Haplophragmoides impressus Voloshinova, uCMP50008, MoS

10 Haplophragmoides mexicanus kornfeld, uCMP50009, PtA.

11 Haplophragmoides pulicosus Saidova, uCMP50010, FRA.

12 Alveophragmium orbiculatum Shchedrina, uCMP50011, PCB.

13, 14 Haplophragmoides sp.: 13, uCMP50012, PPP. 14, uCMP-50013, CuC.

15–18 Cyclammina cancellata (Brady): 15, uCMP50014, Cho. 16, uCMP50015, MAt. 17, uCMP50016, Cho. 18, typical large specimen covered with matrix, uCMP50017, MAt.

19 Ammobaculites agglutinans (d’orbigny), uCMP50018, PPP.

20 Ammobaculites exilis Cushman and Brönnimann, uCMP-50019, MS10.

21 Vulvulina pacifica Cushman, uCMP50020, RQt.

22 Tritaxis challengeri (hedley, hurdle and Burdett), uCMP-50021, PPP.

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Plate 1Kenneth L. Finger

Micropaleontology, vol. 59, nos. 4–5

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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 koLPACk 1980, pl. 9, fig. 8. — JoNES 1994, p. 32, pl. 21, figs. 1– 8, 10–13.

Occurrence: Navidad Fm. (PPP).


order LituoLiDA Lankester 1885Superfamily AMMoDiSCoiDEA Reuss 1862Family AMMoDiSCiDAE Reuss 1862Subfamily AMMoVoLuMMiNiNAE Chernykh 1967AMMoDiSCuS Reuss 1862; emend. Rhumbler 1895 and Loeb-lich and tappan 1954.Type species: Ammodiscus infimus L. G. Bornemann 1874.

Ammodiscus incertus (d’orbigny 1839a)Plate 1, Figure 6

Operculina incerta D’oRBiGNy 1839a, p. 49, pl. 8, figs. 16, 17.Ammodiscus incertus (d’orbigny). 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 d’orbigny’s species and others is blurred by growth rate, growth stage, micro spheric vs. megalospheric dimorphism, and test composition derived from the local substrate. Possible synonyms of d’orbigny’s 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).

Remarks: Large specimens tend to be less flat and less circular than smaller ones.

Occurrence: Navidad Fm. (CPuP, MPuP, PPP), Ranquil Fm. (FRA, FRM), Santo Domingo Fm. (VAL), Lacui Fm. (Cho).


Superfamily hoRMoSiNoiDEA haeckel 1894Family hoRMoSiNiDAE haeckel 1894Subfamily REoPhACiNAE Cushman 1910aREoPhAX de Montfort 1808Type species: Reophax scorpiurus de Montfort 1808.

Reophax agglutinatus Cushman 1913aPlate 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 occur rence.

Remarks: Represented by a single, small specimen.

Occurrence: Ranquil Fm. (FRA).

Maximum relative abundance: Very rare.

Reophax sp.Plate 1, Figure 8

Distinguishing features: 4–5 subspherical chambers, rapidly increasing in size.

Occurrence: Santo Domingo Fm. (VAL).


Superfamily LituoLoiDEA de Blainville 1827Family hAPLoPhRAGMoiDiDAE Maync 1952hAPLoPhRAGMoiDES Cushman 1910aType species: Nonionina canariensis d’orbigny 1839b.

Haplophragmoides impressus Voloshinova, in Voloshinova and Budasheva 1961Plate 1, Figure 9

Haplophragmoides impressus Voloshinova, in VoLoShiNoVA and BuDAShEVA 1961, p. 192, pl. 5, figs. 5–7.

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 1931Plate 1, Figure 10

Haplophragmoides canariensis var. mexicana koRNFELD 1931, p. 83, pl. 13, fig. 4.

Type age and locality: Recent, Louisiana; neritic.

Remarks: Specimens match the holotype by being very compressed, sixchambered 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 1970Plate 1, Figure 11

Haplophragmoides pulicosus SAiDoVA 1970, p. 150, pl. 4, fig. 4.

Upper depth limit: Lower bathyal, based on Saidova’s (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

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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 4–5 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.Plate 1, Figures 12, 13

Remarks: these are rather nondescript internal molds of somewhat inflated planispiral forms, many of which may be juvenile Cyclammina cancellata.

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).

Maximum relative abundance: Common (MPuP, CuC).

Family LituoLiDAE de Blainville 1827Subfamily AMMoMARGiNuLiNiNAE Podobina 1978AMMoBACuLitES Cushman 1910a; emend. höglund 1947Type species: Spirolina agglutinans d’orbigny 1846.

Ammobaculites agglutinans (d’orbigny 1846); emend. Bartenstein 1952Plate 1, Figure 19

Spirolina agglutinans D’oRBiGNy 1846, p. 137, pl. 7, figs. 10–12.Haplophragmium agglutinans (d’orbigny). BRADy 1884, pl. 32, figs.

19, 20, 24–26. Ammobaculites midwayensis PLuMMER 1933, p. 63, pl. 5, figs. 7–11. ? Ammobaculites subagglutinans BANDy 1949a, p. 27, pl. 3, figs. 5a, b. Ammobaculites agglutinans (d’orbigny). BARtENStEiN 1952, p.

318, pl. 1, fig. 1; pl. 2, figs. 10–16. — PAPP and SChMiD 1985, p. 54, pl. 45, figs. 6–9. — JoNES 1994, p. 39, pl. 32, figs. 19, 20, 24– 26. — LoEBLiCh and tAPPAN 1987, p. 74, pl. 58, figs. 3–4. — kAMiNSki and GRADStEiN 2005, p. 324, fig. 70, 1–3. — hoLBouRN, hENDERSoN and MACLEoD 2013, p. 26–27.

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 Brönnimann 1948Plate 1, Figure 20

Ammobaculites exilis CuShMAN and BRöNNiMANN 1948, p. 39, pl. 7, fig. 9.

Type age and locality: Recent, trinidad, 0–12m.

Upper depth limit: Inner neritic; based on type locality.

Comparative species: Ammobaculites salsus Cushman and Brönnimann 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 1884Family CyCLAMMiNiDAE Marie 1941Subfamily CyCLAMMiNiNAE Marie 1941ALVEoPhRAGMiuM Shchedrina 1936Type species: Alveolophragmium orbiculatum Shchedrina 1936.

Alveolophragmium orbiculatum Shchedrina 1936Plate 1, Figure 12

Alveolophragmium orbiculatum ShChEDRiNA 1936, p. 315, text-fig. 1.

Type age and locality: Recent, Japan Sea, 57–200m.

Stratigraphic range: Early Miocene to Recent.

Upper depth limit: outer neritic.

Occurrence: Navidad Fm. (LBz), Lacui Fm. (PCB).


Subfamily CyCLAMMiNiNAE Marie 1941CyCLAMMiNA Brady 1879aType species: Cyclammina cancellata Brady 1879a.

Cyclammina cancellata Brady 1879aPlate 1, Figures 15–18

Cyclammina cancellata BRADy 1879a, p. 62, pl. 37, figs. 8–16. — BRADy 1884, p. 351, pl. 37, figs. 8–16. — GRADStEiN and BERGGREN 1981, p. 254, pl. 7, figs. 1–3. — 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. 2–6. — JoNES, p. 43, pl. 37, figs. 8–16. — zAPAtA and CEAR 2004, p. 4, fig. 12. — hoLBouRN, hENDERSoN and MACLEoD 2013, p. 222–223.

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

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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 4–5mm, 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 Stache’s 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.

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.

Superfamily SPiRoPLECtAMMiNoiDEA Cushman 1927bFamily SPiRoPLECtAMMiNiDAE Cushman 1927bSubfamily VuLVuLiNiNAE Saidova 1981VuLVuLiNA d’orbigny 1826Type species: Vulvulina capreolus Defrance, in d’orbigny 1826.

Vulvulina pacifica Cushman 1932bPlate 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.

Occurrence: Ranquil Fm. (RAN, RQt).


order tRoChAMMiNiDA Schwager 1877Superfamily tRoChAMMiNoiDEA Schwager 1877Family tRoChAMMiNiDAE Schwager 1877Subfamily tRoChAMMiNiNAE Schwager 1877tRitAXiS Schubert 1921Type species: Rotalina fusca Williamson 1858.

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 Brönnimann 1976Type species: Trochammina quadriloba höglund 1948.

Trochamminopsis quadriloba höglund 1948Plate 2, Figure 1

Trochammina quadriloba höGLuND 1948, p. 46 (nom. nov. pro T. pusilla höglund 1947, p. 201, pl. 17, fig. 4).

Type age and locality: Recent, Skagerrak (S of Sweden), 366m.


Upper depth limit: Upper bathyal; based on its type locality.

Occurrence: Ranquil Fm. (FRA), Santo Domingo Fm. (VAL).


order tEXtuLARiiNA Ehrenberg 1839Superfamily VERNEuiLiNoiDEA Cushman 1911Family PRoLiXoPLECtiDAE Loeblich and tappan 1985kARRERuLiNA Finlay 1940Type 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. 7–9.

Type age and locality: Recent, type locality not designated; localities noted in North Pacific at 3840–7224m.


Upper depth limit: Lower bathyal, based on Loeblich and tappan (1987) for genus.

Occurrence: Navidad Fm. (MPuP, PPP), Ranquil Fm. (MS10), Santo Domingo Fm. (VAL).


Family VERNEuiLiNiDAE Cushman 1911Subfamily VERNEuiLiNiNAE Cushman 1911GAuDRyiNA d’orbigny 1840Type species: Gaudryina rugosa d’orbigny 1840.

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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 Grönhagen and Luterbacher 1966GuPPyELLA Brönnimann 1951Type 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).


Superfamily tEXtuLARioiDEA Ehrenberg 1838Family EGGERELLiDAE Cushman 1937Subfamily DoRothiiNAE Balakhmatova 1972DoRothiA Plummer 1931; emend. Desai and Banner 1987Type species: Gaudryina bulletta Carsey 1926.

Dorothia cylindrica (Nuttall 1932)Plate 2, Figure 5

Gaudryina cylindrica NuttALL 1932, p. 7, pl. 2, fig. 7.

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 1937EGGERELLA Cushman 1933bType species: Verneuilina bradyi Cushman 1911.

Eggerella bradyi (Cushman 1911)Plate 2, Figure 6

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. 4–7. — RoBERtSoN 1998, p. 24, pl. 3, fig. 4. — hAyWARD, GRENFELL, SABAA, NEiL and BuzAS 2010, p. 144, pl. 5, figs. 23, 24. — hoLBouRN, hENDERSoN and MACLEoD 2013, p. 232–233.

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. 5–7.Eggerelloides scaber (Williamson). JoNES 1994, p. 52, figs. 15–17.

Type age and locality: Recent, type locality not designated, but localities given in United Kingdom, depths not indicated.




kARRERiELLA Cushman 1933b Type species: Gaudryina siphonella Reuss 1851b.

Karreriella biglobata Finger, n. sp.Plate 2, Figure 2

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.

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.

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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. 1–4. — RoBERtSoN 1998, p. 26, pl. 4, fig. 1. — hAyWARD, GRENFELL, SABAA, NEiL and BuzAS 2010, p. 145, pl. 5, figs. 25–27. — hoLBouRN, hENDERSoN and MACLEoD 2013, p. 318–319.

Type age and locality: Recent; localities given have an average depth of 1829m off hawaiian islands, Galápagos 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)Plate 2, Figures 9, 10

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 1933bType species: Clavulina communis d’orbigny 1826.

Martinottiella communis (d’orbigny 1846)Plate 2, Figure 12

Clavulina communis D’oRBiGNy 1846, p. 196, pl. 12, figs. 1, 2.Martinottiella communis (d’orbigny). iNGLE, kELLER and koL

PACk 1980, pl. 4, fig. 14, 15. — PAPP and SChMiD 1985, p. 74, pl. 66, figs. 1–8. — kohL 1985, p. 33, pl. 4, fig. 2. — LoEBLiCh and tAPPAN 1987, pl. 190, figs. 3, 4. — JoNES 1994, p. 52, pl. 66, figs. 1–8. — hoLBouRN, hENDERSoN and MACLEoD 2013, p. 350–351.


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 2Figures 7 and 15 are photomicrographs; all other images are SEMs. Scale bars in µm.

1 Trochamminopsis quadriloba (höglund), uCMP50022, FRA.

2 Karreriella biglobata Finger, n. sp., holotype uCMP50023, FRM.

3 Karrerulina apicularis Cushman, uCMP50024, MS10.

4 Guppyella crassa (Cushman and Renz), uCMP50025, PPP.

5 Dorothia cylindrica (Nuttall), uCMP50026, RAP.

6 Eggerella bradyi (Cushman), uCMP50027, RAN.

7 Eggerelloides scabrus (Williamson), crushed specimen, uCMP50028, PPP.

8 Karreriella bradyi (Cushman), uCMP50029, FRA.

9, 10 Karreriella subcylindrica (Nuttall): 9, without biserial stage uCMP50030, PtA. 10, with biserial stage, uCMP50031, FRA.

11 Martinottiella juncea (Cushman), uCMP50032, FRA.

12 Martinottiella communis (d’orbigny), uCMP50033, PPP.

13 Gaudryina sp., uCMP50034, FRM.

14 Textularia lythostrota (Schwager), uCMP50035, FRA.

15 Martinottiella pallida (Cushman), uCMP50036, PPP.

16 Textularia miozea Finlay, uCMP50037, NAV5.

383

Kenneth L. Finger Plate 2


384


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 Bermúdez 1937; Eocene, Cuba).

Occurrence: Navidad Fm. (PPt), Ranquil Fm. (FRA).


Martinottiella pallida (Cushman 1927f)Plate 2, Figure 15

Clavulina communis var. pallida CuShMAN 1927f, p. 138, pl. 2, fig. 1.

Type age and locality: Recent, off California at 850m.

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. (PPP, RAP), Ranquil Fm. (RQt), Santo Domingo Fm. (VAL).


Family tEXtuLARiiDAE Ehrenberg 1838Subfamily tEXtuLARiiNAE Ehrenberg 1838tEXtuLARiA Defrance, in de Blainville 1824Type 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 d’orbigny 1839aPlate 3, Figures 1, 2

Textularia agglutinans D’oRBiGNy 1839a, p. 144, pl. 1, fig. 17, 18, 32, 34. — JoNES 1994, p. 48, pl. 43, figs. 1–3.

Type age and locality: Recent, Caribbean, depth not recorded.


Occurrence: El Peral beds (NLP), Navidad Fm. (CPuP, MPuP, PPP, PPt, PtA), Ranquil Fm. (RAN), Lacui Fm. (Cho).


Textularia lythostrota (Schwager 1866)Plate 2, Figure 14

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: Early to Middle Pliocene, Car Nicobar.

Occurrence: Navidad Fm. (NAV5, PPP, PPt, PtA), Ranquil Fm. (FRA), Lacui Fm. (PCB).


Textularia miozea Finlay 1939cPlate 2, Figure 16

Textularia cf. T. miozea FiNLAy 1939c, p. 326, pl. 129, figs. 159–161.

Type age and locality: Middle Miocene, New zealand.

Occurrence: Navidad Fm. (CPuP, NAV5, PPt), Ranquil Fm. (RQt).


Textularia schencki Cushman and Valentine 1930Plate 3, Figure 3

Textularia schencki CuShMAN and VALENtiNE 1930, p. 8, pl. 1, fig. 3.

Type age and locality: Recent, California, 5–37m.


1, 2 T extularia agglutinans d’orbigny: 1, uCMP50038, Cho. 2, uCMP50039, PtA.

3 Textularia schencki Cushman and Valentine, uCMP50040, PCB.

4 Textularia sp. A, uCMP50041, PNh.

5 Textularia sp. B., uCMP50042, NLP.

6 Textularia sp. C, uCMP50043, RQt.

7 Textularia? sp., uCMP50044, PtA.

8 Pseudoclavulina mexicana (Cushman), uCMP50045, FRA.

9 Cornuspira libella Finger, n. sp., holotype uCMP50046, FRA.

10 Cornuspira planorbis Schultze, uCMP50047, FRA.

11 Cornuspiroides foliaceus (Philippi), uCMP50048, RAP.

12 Cornuspira veleronis (McCulloch), uCMP50049, PPP.

13 Nummulopyrgo globulus (Bornemann), uCMP50050, NLP.

14 Spiroloculina robusta Brady, uCMP50051, NLP.

15 Spiroloculina incisa Cushman, uCMP50052, FRA.

16 Quinqueloculina akneriana d’orbigny, uCMP50053, MiB.

17 Quinqueloculina badenensis d’orbigny, uCMP50054, RAP.

18 Quinqueloculina cf. Q. benwestonensis McCulloch, uCMP- 50055, FRA.

385



386


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 d’orbigny 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).


Textularia sp. APlate 3, Figure 4

Occurrence: Ranquil Fm. (RAN), Lacui Fm. (PNh).


Textularia sp. BPlate 3, Figure 5

Occurrence: El Peral beds (NLP).


Textularia sp. CPlate 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 biserialtouniserial species in the fauna; hence, the generic placement of this specimen is uncertain.

Occurrence: Lacui Fm. (Cho).


Family PSEuDoGAuDRyiNiDAE Loeblich and tappan 1985Subfamily PSEuDoGAuDRyiNiNAE Loeblich and tappan 1985PSEuDoCLAVuLiNA Cushman 1936Type 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. 1–3 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.

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).


order MiLioLiDA Lankester 1885Superfamily CoRNuSPiRoiDEA von Schultze 1854Family CoRNuSPiRiDAE von Schultze 1854Subfamily CoRNuSPiRiNAE von Schultze 1854CoRNuSPiRA von Schultze 1854Type species: Orbis foliaceus Philippi 1844.

Cornuspira libella Finger, n. sp.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 (Galápagos, 24m), which has the spiral nodes but lacks the spiral ridge. Spirillina nodosa terquem 1880 (Recent, northern France) has 6–7 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 1854Plate 3, Figure 10

Cornuspira planorbis VoN SChuLtzE 1854, p. 40, pl. 2, fig. 21.

Type age and locality: Recent, off Mozambique, depth not indicated.

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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)Plate 3, Figure 12

Cyclogyra veleronis McCuLLoCh 1981, p. 31, pl. 270, fig. 31.

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 1928Type 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 1839Family SPiRoLoCuLiNiDAE Wiesner 1920NuMMuLoPyRGo hofker 1983Type species: Nummulopyrgo globulus (Bornemann 1855).

Nummulopyrgo globulus (Bornemann 1855)Plate 3, Figure 13

Biloculina globulus BoRNEMANN 1855, p. 349, pl. 19, fig. 3.

Type age and locality: oligocene, Germany.

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


SPiRoLoCuLiNA d’orbigny 1826Type species: Spiroloculina depressa d’orbigny 1826.

Spiroloculina incisa Cushman 1921Plate 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)


Upper depth limit: outer neritic; based on type locality (shelf edge for Spiroloculina sp. in ingle, keller and kolpack 1980).

Comparative species: Spiroloculina depressa d’orbigny 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 d’orbigny 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 1884Plate 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 3–4 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 ujiié 1990, but differs in coiling mode.

Occurrence: El Peral beds (NLP), Navidad (PPP, PPt)


Family hAuERiNiDAE Schwager 1876Subfamily hAuERiNiNAE Schwager 1876QuiNQuELoCuLiNA d’orbigny 1826Type species: Serpula seminulum Linnaeus 1758.

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).

Quinqueloculina akneriana d’orbigny 1846 Plate 3, Figure 16

Quinqueloculina akneriana d’orbigny 1846, p. 100, p. 290, pl. 18, figs. 16–21. — PAPP and SChMiD 1985, pl. 95, figs. 1–5.

388



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.

Occurrence: El Peral beds (LPER, NLP), Navidad Fm. (CPuP, 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).

Quinqueloculina badenensis d’orbigny 1846Plate 3, Figure 17

Quinqueloculina badenensis D’oRBiGNy 1846, p. 299, pl. 20, figs. 10–12. — PAPP and SChMiD 1985, p. 105, pl. 101, figs. 6–10.


Comparative species: Quinqueloculina bicostata d’orbigny 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 1977Plate 3, Figure 18

Quinqueloculina benwestonensis MCCuLLoCh 1977, p. 482, pl. 211, fig. 1.

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 d’orbigny 1846Plate 4, Figure 1

Quinqueloculina boueana D’oRBiGNy 1846, p. 293, pl. 19, figs. 7–9. — PAPP and SChMiD 1985, p. 101, pl. 96, figs. 8, 9. — JoNES 1994, p. 23, pl. 7, fig. 13.


Distinguishing features: Distinctly costate; subacute edges, slightly concave sides.

Occurrence: Navidad Fm. (PPN, RAP).

Maximum relative abundance: Common (RAP).

Quinqueloculina cf. Q. flexuosa d’orbigny 1839cPlate 4, Figure 2

Quinqueloculina flexuosa D’oRBiGNy 1839c, p. 73, pl. 4, figs. 4–6. (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).



1 Quinqueloculina boueana d’orbigny, uCMP50056, RAP.

2 Quinqueloculina cf. Q. flexuosa d’orbigny, uCMP50057, RAN.

3 Quinqueloculina magellanica d’orbigny, uCMP50058, CUC.

4 Quinqueloculina opulenta McCulloch, uCMP50059, NAV5.

5 Quinqueloculina suborbicularis d’orbigny, uCMP50060, MiB.

6 Quinqueloculina sp. A, uCMP50061, MiB.

7 Quinqueloculina sp. B, uCMP50062, RAN.

8 Quinqueloculina sp. C, uCMP50063, RAP.

9 Biloculinella labiata (Schlumberger), uCMP50064, PPP.

10 Cribromiliolinella subvalvularis (Parr), uCMP50065, MiB.

11, 12 Miliolinella suborbicularis (d’orbigny): 11, uCMP50066, RAP. 12, uCMP50067, CuC.

13 Pseudotriloculina cf. P. cyclostoma (Reuss), uCMP-50068, MiB.

14 Pyrgo depressa (d’orbigny), uCMP50069, RAN.

15 Pyrgo lunula (d’orbigny), uCMP50070, RQt.

389



390


Quinqueloculina magellanica d’orbigny 1839cPlate 4, Figure 3

Quinqueloculina magellanica D’oRBiGNy 1839c, p. 77, pl. 9, figs. 19–21.

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 1977Plate 4, Figure 4

Quinqueloculina opulenta MCCuLLoCh 1977, p. 500, pl. 216, figs. 2, 3, 6.

Type age and locality: Recent, Galápagos islands, 29m.

Distinguishing feature: Flat peripheral edge.

Occurrence: NAV5.


Quinqueloculina suborbicularis d’orbigny 1905Plate 4, Figure 5

Quinqueloculina suborbicularis D’oRBiGNy 1826, p. 302, modèle no. 29 (nomen nudem).

Quinqueloculina suborbicularis d’orbigny 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. APlate 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. BPlate 4, Figure 7

Comparative species: Quinqueloculina inaequalis d’orbigny 1839b (Recent, Canary islands) has a sinuous profile in apertural view.

Occurrence: Ranquil Fm. (FRA), Santo Domingo (VAL),


Quinqueloculina sp. CPlate 4, fig. 8

Occurrence: Navidad Fm. (RAP), Ranquil Fm. (RQt).


Subfamily MiLioLiNELLiNAE Vella 1957BiLoCuLiNELLA Wiesner 1931Type species: Biloculina labiata Schlumberger 1891.

Biloculinella labiata (Schlumberger 1891)Plate 4, Figure 9

Biloculina labiata SChLuMBERGER 1891, p. 556, pl. 9, figs. 60–62; p. 556 tf. 13; pl. 557 tf. 14.

Type age and locality: Recent, Mediterranean Sea, 555m.

Comparative species: Biloculina isabelleana d’orbigny 1839c (Recent, Falkland islands) has a rounded edge.

Occurrence: Navidad Fm. (MoS, PPP).


CRiBRoMiLioLiNELLA Saidova 1981Type species: Triloculina subvalvularis Parr 1950.

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. 16–18.

Type age and locality: Recent, off Antarctica, 193m.

Occurrences: Navidad Fm. (CPuP), Ranquil Fm. (MiB, RQt).


MiLioLiNELLA Wiesner 1931Type species: Vermiculum subrotundum Montagu 1803.

Distinguishing features: Early stage quinqueloculine, later planispiral with 4–7 chambers visible.

Miliolinella suborbicularis (d’orbigny 1839a)Plate 4, Figures 11, 12

Triloculina suborbicularis D’oRBiGNy 1839a, p. 177, type figure in v. 8, pl. 10, figs. 9–11.

Type age and locality: Recent, Caribbean.

Distinguishing features: Compressed, rounded edge, fine costae.

Occurrence: Navidad Fm. (MoS, PPP, RAP), Lacui Fm. (CuC).


PSEuDotRiLoCuLiNA Cherif 1970Type species: Triloculina lecalvezae kaaschieter 1961.

Pseudotriloculina cf. P. cyclostoma Reuss 1850Plate 4, Figure 13

391


Biloculina cyclostoma REUSS 1850, p. 382, pl. 49, fig. 6.Pseudotriloculina cyclostoma (Reuss). LoEBLiCh and tAPPAN

1987, p. 342, pl. 352, figs. 6–12.


Distinguishing features: only two chambers visible.

Remarks: Differs from P. cyclostoma by being less inflated.

Comparative species: Triloculina inornata d’orbigny 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 1824Type species: Pyrgo laevis Defrance, in de Blainville 1824.

Pyrgo clypeata (d’orbigny 1846)Plate 5, Figures 1, 2

Biloculina clypeata D’oRBiGNy 1846, p. 266, pl. 15, figs. 19–21.Pyrgo clypeata (d’orbigny). PAPP and SChMiD 1985, p. 89, pl. 82,

figs. 4–6. — hAyWARD, GRENFELL, SABAA, NEiL and BuzAS 2010, p. 150, pl. 7, figs. 7–10.


Comparative species: D’orbigny 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 chamber’s inward ledge and the edge-view 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).

Occurrence: El Peral beds (NLP), Navidad Fm. (CPuP, PPP, PtA, RAP), Ranquil Fm. (FRA, FRM, LEB, MiB, RAN, RQk, RQt), Santo Domingo Fm. (VAL), Lacui Fm. (PNh).


Pyrgo depressa (d’orbigny 1826)Plate 4, Figure 14

Biloculina depressa D’oRBiGNy 1826, p. 298, modèle no. 91.Pyrgo depressa (d’orbigny). 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.

Upper depth limit: Upper middle bathyal; based on occurrence recorded off Chile by Bandy and Rodolfo (1964).

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 (d’orbigny 1846)Plate 4, Figure 15

Biloculina lunula D’oRBiGNy 1846, p. 264, pl. 15, figs. 22–24.Biloculina scutella kARRER 1868, p. 134, pl. 1, fig. 7.Pyrgo lunula (d’orbigny). PAPP and SChMiD 1985, p. 89, pl. 82, fig.

7–9.

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 1866Plate 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. 20–123.

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.

Occurrence: El Peral beds (LPER), Navidad Fm. (MoS, RAP), Ranquil Fm. (FRA, MiB, RQk, RQt).


PyRGoELLA Cushman and White 1936Type species: Biloculina sphaera d’orbigny 1839c.

Pyrgoella sphaera (d’orbigny 1839c)Plate 5, Figure 3

Biloculina sphaera D’oRBiGNy 1839c, p. 66, pl. 8, figs. 13–16.Pyrgoella sphaera (d’orbigny). JoNES 1994, p. 18, pl. 2, fig. 4. —

hAyWARD, GRENFELL, SABAA, NEiL and BuzAS 2010, p.

392


152, pl. 8, figs. 9–11.Pyrgoella(?) generalis MCCuLLoCh 1977, p. 522, pl. 242, figs. 9–18.

(Recent, Pacific off Mexico)

Type age and locality: Recent, Falkland Islands, depth not recorded.


Upper depth limit: outer neritic; based on records off New zealand (hayward et al. 2010).

Comparative species: Pyrgoella dokici (= Biloculina dokici Pavlović 898; tertiary, yugoslavia) is not as spherical and its apertural end protrudes outward.

Occurrence: El Peral beds (NLP), Navidad Fm. (CPuP, MoS, PPP), Ranquil Fm. (FRA, RQt), Santo Domingo Fm. (VAL).


tRiLoCuLiNA d’orbigny 1826; emend. Łuczkowska 1972Type species: Miliola (Miliolites) trigonula Lamarck 1804.

Triloculina cf. T. brochita Carter 1964Plate 5, Figure 5

Triloculina brochita CARtER 1964, p. 59, pl. 1, figs. 3, 4.

Type age and locality: Lower Miocene, Victoria, Australia.

Remarks: In apertural view, Triloculina brochita has a subtriangular outline. the apertural tooth characteristic of the genus is not preserved.

Occurrence: Lacui Fm. (CuC).


Triloculina lucernuloides (ujiié 1990)Plate 5, Figure 7

Pyrgo lucernuloides uJiié 1990, p. 15, pl. 3 figs. 7, 8.

Type age and locality: Late Pleistocene, Ryuku islands.

Stratigraphic range: Early Miocene to late Pleistocene.

Upper depth limit: Bathyal; based on typedescription.

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: Its shape resembles that of Spiroloculina robusta Brady 1884, but T. lucernuloides has a sinuous outline and a different mode of coiling.

Occurrence: Navidad Fm. (CPuP, MAt, MoS, NAV5, PPP, PtA), Ranquil Fm. (MS10. RQt), Santo Domingo Fm. (VAL).


Triloculina oblonga (Montagu 1803)Plate 5, Figure 6

Vermiculum oblongum MoNtAGu 1803, p. 522, pl. 19, fig. 9.

Type age and locality: Recent, England, depth not recorded.


Comparative species: this form is similar to Triloculina oblonga d’orbigny 1839a (Recent, Cuba) and T. chemnitziana d’orbigny 1839b (Recent, Canary islands). Montagu’s type description and figure are inadequate for comparison and d’orbigny does not comment on Montagu’s species. Assuming they are synonymous, Montagu’s nomen has seniority.

Occurrence: Navidad Fm. (NAV5), Ranquil Fm. (FRA).


Triloculina striatotrigonula Parker and Jones 1941Plate 5, Figure 8


1, 2 Pyrgo clypeata (d’orbigny): 13, uCMP50071, MiB. 14, uCMP50072, FRA.

3 Pyrgoella sphaera (d’orbigny), 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 (ujiiè), uCMP50077, PPP.

8 Triloculina striatotrigonula Parker and Jones, orally distorted specimen, uCMP50078, FRA.

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.

393



394


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, 69–73m.

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 LAMARCk 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 subroundedsubtriangular, as all three sides are convex and the corners are well rounded.

Occurrence: (RAP)


tRiLoCuLiNELLA Riccio 1950Type species: Triloculinella obliquinodus Riccio 1950.

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.

Triloculinella bornemanni (Bosquet 1859)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).


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. APlate 5, Figure 13

Remarks: Juveniles display the broad apertural flap.

Comparative species: Similar in subglobose form to Triloculina flavescens d’orbigny (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. BPlate 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 Ushaped, 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),


Subfamily SiGMoiLiNitiNAE Łuczkowska 1974NuMMoLoCuLiNA Steinmann 1881Type species: Biloculina contraria d’orbigny 1846.

Nummoloculina contraria (d’orbigny 1846)Plate 6, Figure 3

Biloculina contraria d’orbigny 1846, p. 266, pl. 16, figs. 4–6.Nummuloloculina contraria (d’orbigny). PAPP and SChMiD 1985, p.

90, pl. 83, figs. 7–9. — LoEBLiCh and tAPPAN 1987, pl. 355, figs. 17–23. — hAyWARD, GRENFELL, SABAA, NEiL and BuzAS 2010, p. 150, pl. 7, figs. 4–6. — hoLBouRN, hENDERSoN and MACLEoD 2013, p. 376–377.



Distinguishing features: three to four chambers visible in the outer whorl.


395



SiGMoiLiNitA Seiglie 1965Type species: Quinqueloculina tenuis Cžjžek 1848.

Sigmoilinita tenuis (Cžjžek 1848)Plate 6, Figure 1

Quinqueloculina tenuis CžJžEk 1848, p. 149, pl. 13, figs. 31–34.Sigmoilina miocenica CuShMAN 1946 (Miocene, Florida).Spirosigmoilina tenuis (Cžjžek). 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. — hAyWARD, GRENFELL, SABAA, NEiL and BuzAS 2010, p. 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 Cžjžek’s 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 1983Type species: Biloculina vespertilio Schlumberger 1891.

Sigmopyrgo vespertilio (Schlumberger 1891)Plate 5, Figure 13

Biloculina vespertilio SChLuMBERGER 1891, p. 561, pl. 10, figs. 74–76; p. 561, tfs. 20–21; 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. 14–18. — JoNES 1994, p. 18, pl. 2, fig. 8.

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 1957SiGMoiLoPSiS Finlay 1947Type species: Sigmoilina schlumbergeri A. Silvestri 1904a.

Sigmoilopsis schlumbergeri (A. Silvestri 1904a)Plate 6, Figure 2

Sigmoilina schlumbergeri A. SiLVEStRi 1904a, p. 267, 269; type-figures in SChLuMBERGER 1887, pl. 7, figs. 12–14; 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. 1–4. — 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, 600–1200m.


Upper depth limit: Neritic, but in greatest abundance at bathyal depths (van Morkhoven, Berggren and Edwards 1986; hayward et al. 2010).



order LAGENiDA Lankester 1885Superfamily NoDoSARioiDEA Ehrenberg 1838Family ChRySALoGoNiiDAE Mikhalevich 1993ANAStoMoSA hayward, in hayward et al. 2012Type 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. 20–25.

Type age and locality: Early Miocene, trinidad.

Stratigraphic range: Late Eocene to Early Miocene.

Upper depth limit: Unknown.

Occurrence: El Peral beds (NLP)


Anastomosa lamellata (Cushman and Stainforth 1945)Plate 6, Figures 5, 6

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, kAWAGAtA, SABAA, GRENFELL, VAN kERCkhoVEN, JohNSoN and thoMAS 2012, p. 116, pl. 3, figs. 10–14.

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 1908Type species: Nodosaria polystoma Schwager 1866.

396


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.

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. 9–16.


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.

Occurrence: Navidad (MPuP), Ranquil Fm. (FRM), Lacui Fm. (CuC).


Chrysalogonium equisetiformis (Schwager 1866)Plate 6, Figure 13

Nodosaria equisetiformis SChWAGER 1866, p. 231, pl. 6, fig. 66.Chrysalogonium equisetiformis (Schwager). hAyWARD, kAWAGA


Nodosaria spirostriolata CuShMAN 1917b, p. 656, pl. 38, fig. 4. (Recent, Philippines, 903m)


Stratigraphic range: Late Cretaceous to early Pleistocene.

PLATE 6Figures 2a, 3a,b, 4–6, 15, 18, 19, 24b, 34, and 37 are photomicrographs;

all other images are SEMs. Scale bars in µm.

1 Sigmoilinita tenuis (Cžjžek), uCMP50086, RQt.

2 Sigmoilopsis schlumbergeri (A. Silvestri), uCMP50087, NLP.

3 Nummoloculina contraria (d’orbigny), uCMP50088, PPP.

4 Anastomosa brevilocula (Cushman and Jarvis), uCMP-50089, NLP.

5, 6 Anastomosa lamellata (Cushman and Stainforth), PPP: 5. megalospheric, uCMP50090. 6, microspheric, uCMP-50091.

7, 8 Lotostomoides pyrulus (d’orbigny): 7, uCMP50092, PtA. 8, uCMP50093, PPP.

9 Neugeborina longiscata (d’orbigny), uCMP50094, FRA.

10, 11 Lotostomoides asperulum (Neugeboren): 10, uCMP-50095, PtA. 11, uCMP50096, NAV5.

12 Chrysalogonium deceptorium (Schwager), uCMP50097, FRM.

13 Chrysalogonium equisetiformis (Schwager), uCMP50098, RAN.

14 Chrysalogonium rudis (d’orbigny), uCMP50099, FRA.

15 Orthomorphina jedlitschkai (thalmann), uCMP50100, PPP.

16 Orthomorphina perversa (Schwager), uCMP50101, PtA.

17, 18 Dentalina aciculata d’orbigny, PPP: 17, uCMP50102. 18, uCMP50103.

19 Dentalina albatrossi (Cushman), uCMP50104, MPuP.

20 Dentalina mutsui hada, uCMP50105, FRA.

21 Dentalina obliquecostata (Stache), uCMP50106, RAP.

22, 23 Dentalina striatissima Stache, FRM: 22, uCMP50107. 23, uCMP50108.

24 Enantiodentalina muraii uchio, uCMP50109, PPP.

25 Fingerina weaveri (Finger and Lipps), uCMP50110, PPP.

26, 27 Pyramidulina acuminata (hantken): 26, microspheric form, uCMP50111, FRM. 27, megalospheric form, uCMP50112, MiB.

28 Dentalina flintii (Cushman), uCMP50113, PPP.

29 Laevidentalina cf. L. planata (Blake), uCMP50114, PPN.

30 Laevidentalina roemeri (Neugeboren), uCMP50115, MiB.

31 Laevidentalina sp. A, uCMP50116, PPP.

32 Laevidentalina sp. B, uCMP50117, FRM.

33 Laevidentalina advena (Cushman), uCMP50118, PPP.

34, 35 Laevidentalina communis (d’orbigny): 34, uCMP50119, PPP. 35, uCMP50120, PtA.

36, 37 Laevidentalina elegans (d’orbigny): 36, uCMP50121, FRA. 37, uCMP50122, PPP.

38 Laevidentalina inflexa (Reuss), uCMP50123, PPP.

39 Laevidentalina inornata (d’orbigny), uCMP50124, FRA.

397



398



Distinguishing features: Approximately 30 striae continue across sutures of barrelshaped 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 (d’orbigny 1846)Plate 6, Figure 14

Nodosaria rudis D’oRBiGNy 1846, p. 33, pl. 1, figs. 17–19. — PAPP and SChMiD 1985, p. 24, pl. 4, fig. 5.

Siphonodosaria setosa SChWAGER 1866, p. 219, pl. 5, fig. 40. (Pliocene, Car Nicobar)

Chrysalogonium rudis (d’orbigny). hAyWARD, kAWAGAtA, SABAA, GRENFELL, VAN kERCkhoVEN, JohNSoN and thoMAS 2012, p. 121, pl. 5, figs. 6–13.

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. 2012Type 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)Plate 6, Figures 10, 11

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. 191–200.

(Pliocene, italy)Lotostomoides asperulum (Neugeboren). hAyWARD, kAWAGAtA,

SABAA, GRENFELL, VAN kERCkhoVEN, JohNSoN and thoMAS 2012, p. 125, pl. 6, figs. 20–23.

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.

Comparative species: Nodosaria insecta Schwager 1866 (Pliocene, Car Nicobar) has a radiate aperture and more ovate chambers. Singlechamber segments could be mistakenly identified as N. fichteliana Neugeboren 1852 (tertiary, Romania). Chrysalogonium rudis (= Nodosaria rudis d’orbigny 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 (d’orbigny 1826)Plate 6, Figures 7, 8

Nodosaria pyrula D’oRBiGNy 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 d’orbigny. RoBERtSoN 1998, p. 46, pl. 15, fig. 8.Grigelis pyrulus (d’orbigny). 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 d’orbigny 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, Rögl, Rupp and Ctyroka 1998

Neugeborina longiscata (d’orbigny 1846)Plate 6, Figure 9

Nodosaria longiscata D’oRBiGNy 1846, p. 32, pl. 1, figs. 10–12. — kohL 1985, p. 43, pl. 6, fig. 3. — PAPP and SChMiD 1985, p. 23, pl. 3, figs. 1−5.

Ellipsonodosaria longiscata (d’orbigny). CuShMAN and CEDERStRoM 1945, p. 32, pl. 4, figs. 19−21.

Ellipsonodosaria cf. E. longiscata (d’orbigny). CuShMAN 1948, p. 239, pl. 19, figs. 12, 13.

Nodosaria? longiscata d’orbigny. RoBERtSoN 1998, p. 44, pl. 15, fig. 5.Neugeborina longiscata (d’orbigny). kAWAGAtA, hAyWARD and

GuPtA 2006, p. 237, pl. fig. X. — hAyWARD, kAWAGAtA, SABAA, GRENFELL, VAN kERCkhoVEN, JohNSoN and thoMAS 2012, p. 134, pl. 8, figs. 19–23.


Stratigraphic range: Late Cretaceous to middle Pleistocene.


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Remarks: Some of the Chilean specimens display very faint striae.

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 d’orbigny 1902 (in Fornasini 1902; nom. nov. pro N. ovicula d’orbigny 1826), may also be variants of Neugeborina longiscata.

Occurrence: El Peral beds (LPER, NLP), Navidad Fm. (CPuP, 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 1952Type species: Nodogenerina havanensis Cushman and Bermúdez 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. 24–27.

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).


Orthomorphina perversa (Schwager 1866)Plate 6, Figure 16

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. 35–38; pl. 9, figs. 1–4.



Upper depth limit: upper bathyal, 500m (hayward et al. 2012)

Occurrence: Navidad Fm. (PtA), Lacui Fm. (PCB).


Family NoDoSARiiDAE Ehrenberg 1838Subfamily NoDoSARiiNAE Ehrenberg 1838DENtALiNA Risso 1826

Type species: Nodosaria (les Dentalines) cuvieri d’orbigny 1826.

Distinguishing features: Elongate arcuate test, radiate aperture, longitudinal costae.

Dentalina aciculata (d’orbigny 1826)Plate 6, Figures 17, 18

Nodosaria (Dentaline) aciculata D’oRBiGNy 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. 11–14.



Occurrence: Navidad Fm. (MPuP).


Dentalina flintii (Cushman 1923)Plate 6, Figure 28

Nodosaria flintii CuShMAN 1923, p. 85, pl. 14, fig. 23.Dentalina flintii (Cushman). JoNES 1994, p. 76, pl. 64, figs. 20–22.

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.

400


Occurrence: Navidad Fm. (PPP), Ranquil Fm. (MiB, RAN, RQt).


Dentalina mutsui hada 1931Plate 6, Figure 20

Dentalina mutsui hADA 1931, p. 97, pl. 97, text-fig. 50. Type age and locality: Recent, Japan, 27–46m.


Upper depth limit: Inner neritic, based on type occurrence.

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.

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 1865Plate 6, Figures 22, 23

Dentalina striatissima StAChE 1865, p. 208, pl. 22, fig. 38.


Stratigraphic range: Miocene to Pliocene(?).

Comparative species: Dentalina sulcata Nilsson 1826 (Cretaceous, 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: 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 1941Type species: Enantiodentalina communis Marie 1941.

Distinguishing features: Early stage biserial; surface smooth.

Enantiodentalina muraii uchio 1953Plate 6, Figure 24

Enantiodentalina muraii uChio 1953, p. 152, pl. 14, fig. 2.

Distinguishing features: Early chambers of all specimens are biserial and slightly compressed, some tests are apiculate; sutures oblique, depressed.

Occurrence: Navidad Fm. (CPuP, PPP, PtA), Ranquil Fm. (RQt), Lacui Fm. (Cho).


FiNGERiNA hayward, in hayward et al. 2012Type 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 1990Plate 6, Figure 25

Nodosaria weaveri Finger and Lipps, in FiNGER, WEAVER, LiPPS and MiLLER 1990, p. 28, pl. 1, figs. 19, 20. — hAyWARD, kAWAGAtA, SABAA, GRENFELL, VAN kERCkhoVEN, JohNSoN and thoMAS 2012, p. 138, pl. 9, figs. 7–10.

Type age and locality: Early Miocene, California.

Stratigraphic range: Early Miocene to early Pleistocene.


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 d’orbigny 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 1986bType 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.

401


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 (d’orbigny 1826)Plate 6, Figures 34, 35

Nodosaria (Dentaline) communis D’oRBiGNy 1826, p. 254; no type-figure or description given, but earliest are in d’orbigny 1840, p. 13, pl. 1, fig. 4.

Dentalina communis d’orbigny. FiNGER 1992, p. 68, pl. 2, fig. 9.

Type age and locality: Recent, Adriatic Sea.


Comparative species: Laevidentalina filiformis (= Dentalina filiformis terquem 1878; Late Pliocene, Greece) has barrel-shaped 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 (d’orbigny 1846)Plate 6, Figures 36, 37

Dentalina elegans D’oRBiGNy 1846, p. 28, , p. 45, pl. 1, figs. 52–56. — PAPP and SChMiD 1985, p. 28, pl. 10, figs. 1–8.



Upper depth limit: outer neritic; based on 122–155m range reported by Parr (1950).

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).


Laevidentalina inflexa (Reuss 1866)Plate 6, Figure 38

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.

Occurrence: Navidad Fm. (PPP, PPt), Ranquil Fm. (FRM, MiB, MS10).


Laevidentalina inornata (d’orbigny 1846)Plate 6, Figure 39

Nodosaria inornata D’oRBiGNy 1846, p. 28, p. 2, pl. 1, fig. 5. — PAPP and SChMiD 1985, pl. 9, figs. 5–8. — hAyWARD, GRENFELL, SABAA, NEiL and BuzAS 2010, p. 171, pl. 12, figs. 48–520.



Upper depth limit: Upper middle bathyal; based on California occurrence (ingle 1980).

Comparative species: Similar forms have often been referred to Dentalina communis (= Nodosaria (Dentaline) communis d’orbigny 1826; Recent, Adriatic Sea) for which subsequent figures in d’orbigny (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.

Occurrence: El Peral beds (LPER, NLP), Navidad Fm. (CPuP, 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).


Laevidentalina roemeri (Neugeboren 1856)Plate 6, Figure 30

Dentalina roemeri NEuGEBoREN 1856, p. 82, pl. 2, figs. 13–17. — FiNGER 1992, p. 69, pl. 2, figs 19–22.

Type age and locality: Miocene to Pliocene, Romania.


Comparative species: Laevidentalina obliqua (= Nodosaria (Dentaline) obliqua d’orbigny 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

402


neugeboreni Schwager 1866; Pliocene, Car Nicobar) is another possible synonym.

Occurrence: Ranquil Fm. (MiB), Lacui Fm. (Cho).


Laevidentalina sp. APlate 6, Figure 31

Distinguishing features: Large, robust, nearly straight test with slightly oblique sutures and nearly equidimensional chambers.

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 d’orbigny 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. BPlate 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 1878Plate 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 1894Type species: Pyramidulina eptagona Fornasini 1894.

Distinguishing features: Same as Nodosaria but with distinct longitudinal costae.

Pyramidulina acuminata (von hantken 1876)Plate 6, Figures 26, 27

Nodosaria acuminata VoN hANtkEN 1876, p. 28, pl. 2, fig. 9; pl. 13, fig. 5.

Type age and locality: Early oligocene, hungary.

Stratigraphic range: Early oligocene to Recent.

Upper depth limit: Lower bathyal in the Southwest Pacific (hayward 2002).

Remarks: this species resembles Nodosaria affinis d’orbigny 1846 and N. bacillum d’orbigny 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. Vénec-Peyré (2005) could not locate any type specimens of N. affinis in d’orbigny’s collection at the Muséum National d’histoire 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, deeper-water species, Anastomosa lamellata (= Nodosaria lamellata Cushman and Stainforth 1945; new name for Dentalina carinata Neugeboren 1856).

Comparative species: Pyramidulina raphana (= Nautilus raphanus Linnaeus 1758; Recent, Mediterranean Sea) and P. latejugata (= Nodosaria latejugata von Gümbel 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 1955bType 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)Plate 7, Figures 2, 3

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 Hornibrook’s (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.

403


Occurrence: Navidad Fm. (CPuP, MoS, MPuP, NAV5, PPP, PPt, PtA), Ranquil Fm. (MS10, FRA, RAN, RQk, RQt), Santo Domingo Fm. (VAL).


PSEuDoNoDoSARiA Boomgaart 1949Type 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 (d’orbigny 1846)Plate 7, Figures 5, 6

Dentalina brevis D’oRBiGNy 1846, p. 48, pl. 2, figs. 9, 10. — PAPP and SChMiD 1985, p. 30, pl. 12, figs. 8–11.

Nodosaria radicula var. glanduliniformis DERViEuX 1894, p. 599, pl. 5, figs. 3–7. (Miocene, italy)

Glandulina aperta StAChE 1865, p. 188, pl. 22, fig. 11. (Late tertiary, New zealand)


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

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. 1–5.Pseudonodosaria comatula (Cushman). RoBERtSoN 1998, p. 50, pl.

17, fig. 1, 2.



Upper depth limit: Upper bathyal; based on type occurrence.

Occurrence: Navidad Fm. (CPuP, MoS, NAV5, PPP, PPt, PtA), Ranquil Fm. (FRA, RQt), Lacui Fm. (PCB).


Subfamily LiNGuLiNiNAE Loeblich and tappan 1961LiNGuLiNA d’orbigny 1826; emend. Sellier de Civrieux and Dessauvagie 1965; emend. Norling 1972.Type species: Lingulina carinata d’orbigny 1826.

Lingulina sirakawaensis Nakamura 1942Plate 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 d’orbigny 1839b (Recent, Canary islands) is similar but noncarinate.



PSEuDoLiNGuLiNA McCulloch,1977Type species: Pseudolingulina advena McCulloch 1977.

Pseudolingulina digitata (d’orbigny 1826)Plate 7, Figure 9

Frondicularia digitata D’oRBiGNy 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 concave-upward, 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)

Maximum relative abundance: Very rare (one specimen).


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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 1965Type species: Lingulina costata d’orbigny 1846.

Tollmannia costata (d’orbigny 1846)Plate 7, Figure 11

Lingulina costata D’oRBiGNy 1846, p. 62, pl. 3, figs. 1–5. — PAPP and SChMiD, pl. 20, figs. 1–5.

Tollmannia costata (d’orbigny). LoEBLiCh and tAPPAN 1987, pl. 442, figs. 16–24.



Occurrence: Navidad Fm. (PPP, PPt).


Family PLECtoFRoNDiCuLARiiDAE Montanaro-Gallitelli 1957MuCRoNiNA Ehrenberg 1839Type species: Nodosaria (les Mucronines) hasta d’orbigny 1826.

Mucronina acuta (Cushman and Stainforth 1945)Plate 7, Figure 12

Plectofrondicularia nuttalli var. acuta CuShMAN and StAiNFoRth 1945, p. 39, pl. 5. fig. 24.

Mucronina acuta (Cushman and Stainforth). hAyWARD, kAWAGA

tA, SABAA, GRENFELL, VAN kERCkhoVEN, JohNSoN and thoMAS 2012, p. 143, fig. 63, pl. 9, figs. 22–25.

Type age and locality: oligocene, trinidad.


Upper depth limit: Probably upper bathyal based on M. com-pressa.

Comparative species: Mucronina morreyae (= Plectofron-dicularia morreyae Cushman 1929; tertiary, Ecuador) lacks a medial ridge.

Occurrence: Navidad Fm. (PPP), Ranquil Fm. (MiB).


Mucronina compressa (Costa 1855) Plate 7, Figure 13

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. 30–32; pl. 10, figs. 1–6.




PLATE 7Figures 4, 10, 14, 17, 18, 20, 22a, and 23b are photomicrographs; all other images are SEMs. Scale bars in µm.

1 Nodosaria splendidula Schwager, uCMP50125, FRA.

2, 3 Pandaglandulina obliquesuturata (Stache): 2, uCMP50126, RQt. 3, uCMP50127, MS10.

4 Pseudonodosaria aequalis (Reuss), uCMP50128, RAN.

5, 6 Pseudonodosaria brevis (d’orbigny), PPP: 5, uCMP50129. 6, uCMP50130.

7 Pseudonodosaria comatula (Cushman), uCMP50131, FRA.

8 Lingulina sirakawaensis Nakamura, uCMP50132, PPP.

9 Pseudolingulina digitata (d’orbigny), uCMP50133, CPuP.

10 Pseudolingulina nielseni Finger, n. sp., holotype uCMP-50134, CPuP.

11 Tollmannia costata (d’orbigny), uCMP50135, PPP.

12 Mucronina acuta (Cushman and Stainforth), uCMP50136, MiB.

13 Mucronina compressa (Costa), uCMP50137, FRA.

14 Mucronina spatulata (Costa), uCMP50138, MoS.

15–17 Mucronina striata (d’orbigny): 15, uCMP50139, PPP. 16, uCMP50140, Cho. 17, aberrant form, uCMP50141, PPP.

18 Plectofrondicularia californica Cushman and Stewart, uCMP50142, CPuP.

19, 20 Plectofrondicularia digitalis (Neugeboren): 19, uCMP- 50143, FRA. 20, uCMP50144, PPP.

21 Cristellariopsis petersonae Finger, n. sp., holotype uCMP50145, MAt.

22 Lenticulina calcar (Linné), uCMP50146, PPP.

23 Lenticulina douglasi Finger, uCMP50147, CuC.

405



406


Comparative species: Plectofrondicularia vaughani Cushman 1927d (Eocene, Mexico) lacks the short costae at the apical end.

Occurrence: El Peral beds (LPER, NLP), Navidad Fm. (CPuP, MoS, PPP, PPt, ), Ranquil Fm. (FRA, FRM, MiB, MS10)


Mucronina spatulata (Costa 1855)Plate 7, Figure 14

Frondicularia spatulata CoStA 1855, p. 372, pl. 2, fig. 19.Mucronina spatulata (Costa). hAyWARD, kAWAGAtA, SABAA,

GRENFELL, VAN kERCkhoVEN, JohNSoN and thoMAS 2012, p.151, fig. 63; pl. 12, figs. 15–22.


Stratigraphic range: Late Eocene to middle Pleistocene.

Upper depth limit: upper bathyal (400m).



Mucronina striata (d’orbigny 1826)Plate 7, Figures 15–17

Frondicularia striata D’oRBiGNy 1826, p. 256, pl. 10, fig. 67.Mucronina striata (d’orbigny). hAyWARD, kAWAGAtA, SABAA,

GRENFELL, VAN kERCkhoVEN, JohNSoN and thoMAS 2012, p. 152, fig. 62; pl. 13, figs. 3–8.

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).

Maximum relative abundance: Common (MS10).

PLECtoFRoNDiCuLARiA Liebus 1902Type species: Plectofrondicularia concava Liebus 1902.

Plectofrondicularia californica Cushman and Stewart 1926Plate 7, Figure 18

Plectofrondicularia californica CuShMAN and StEWARt 1926, p. 39, pl. 6, figs. 1–8. — hAyWARD, kAWAGAtA, SABAA, GRENFELL, VAN kERCkhoVEN, JohNSoN and thoMAS 2012, p.

156, figs. 64, 65; pl. 14, figs. 1–8.Plectofrondicularia floridana CuShMAN 1930a, p. 41, pl. 8, fig. 1

(Miocene, Florida). — kohL 1985, p. 45, pl. 9, figs. 6, 7. — RoBERtSoN 1998, p. 56, pl. 18, fig. 7. — LoEBLiCh and tAPPAN 1987, pl. 443, figs. 1, 2.

Type age and locality: Pliocene, California.

Stratigraphic range: oligocene to earliest middle Pleistocene.


Occurrence: El Peral beds (LPER, NLP), Navidad Fm. (CPuP).


Plectofrondicularia digitalis (Neugeboren 1850)Plate 7, Figures 19, 20

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

tA, SABAA, GRENFELL, VAN kERCkhoVEN, JohNSoN and thoMAS 2012, p. 157, figs. 64, 65; pl. 14, figs. 14–20.

Type age and locality: Middle Miocene, Austria.

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

Compressed with prominent ornamentation ..........Percultazonaria

Compressed without ornamentation ........................Cristellariopsis

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

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Slightly compressed ± ornamentation ...................... Vaginulinopsis

Early stage not coiled .............................................. VAGiNuLiNiNAE

Compressed bladelike to palmate ................................. Vaginulina

Subfamily LENtiCuLiNiNAE Chapman, Parr and Collins 1934CRiStELLARioPSiS Rzehak 1895Type 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: Navidad Fm. (MAt).

Maximum relative abundance: Rare (3 specimens).

Type specimens: holotype uCMP50145; paratype uCMP50430 (from MAt).

Type age and locality: Early Miocene; Navidad Fm. Formation in coastal bluff at Los Goterones (Navidad Fm. locality MAt), about 1km north of Matanzas.

Etymology: Named for the late Dawn E. Peterson, who was a great assistance in processing the Chilean samples for microfaunal study.

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. 3g–i, 4l–n; 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. 9–12. — RoBERtSoN 1998, p. 62, pl. 20, fig. 4. — hAyWARD, GRENFELL, SABAA, NEiL and BuzAS 2010, p. 177, pl. 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 5–6 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 subcircularsubpolygonal outline.

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 1992Plate 7, Figure 23

Lenticulina douglasi FiNGER 1992, p. 70, pl. 4, figs. 32–39.

Type age and locality: Miocene, California.


Diagnostic features: Subcircularsubpolygonal periphery with narrow, rounded carina; usually 6 chambers, with slightly depressed sutures.

Comparative species: Differs from Lenticulina subcultrata (= Robulus subcultratus d’orbigny 1839c; Recent, Falkland islands and Canary islands) by its lateral outline.

Occurrence: Ranquil Fm. (MS10), Navidad Fm. (MoS, MPuP, PPP, PtA), Ranquil Fm. (FRM, MS10, RQt), Lacui Fm. (ChE, CuC).


Lenticulina foliata (Stache 1865)Plate 8, Figure 1

Cristellaria (Robulina) foliata StAChE 1865, p. 245, pl. 23, fig. 24.


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.

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 (d’orbigny 1839a)Plate 8, Figure 2

Cristellaria gibba D’oRBiGNy 1839a, p. 40; type-fig. in v. 8, pl. 7, figs. 20, 21.

Lenticulina gibba (d’orbigny). 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.

408


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; 9–10 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).


Lenticulina grandis (Bornemann 1860)Plate 8, Figure 4

Robulina grandis BoRNEMANN 1860, p. 156, pl. 6, fig. 1.

Type age and locality: Early oligocene, Germany.

Stratigraphic range: Early oligocene to Early Miocene.

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)


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,

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.

Occurrence: Ranquil Fm. (MiB, RQt).


Lenticulina neopolita Finger, new namePlate 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 8Figures 1b, 2b, 3b, 4a, 6b, 7b, 9b, 10a, 11a, 12b, 13a, and 14c are photomicrographs; all other images are SEMs. Scale bars in µm.

1 Lenticulina foliata (Stache), uCMP50148, FRA.

2 Lenticulina cf. L. gibba (d’orbigny), uCMP50149, LEB.

3 Lenticulina glaucina (Stache), uCMP50150, FRA.

4 Lenticulina grandis (Bornemann), uCMP50151, PPN.

5 Lenticulina halophora (Stache), uCMP50152, CPuP.

6 Lenticulina miyagiensis (Asano), uCMP50153, MiB.

7 Lenticulina neopolita Finger, new name, uCMP50154, MiB.

8 Lenticulina nuttalli (Cushman and Renz), uCMP50155, MS10.

9 Lenticulina stellata (Seguenza), uCMP50156, RQt.

10, 11 Lenticulina subcultrata (d’orbigny): 10, carinate, uCMP- 50157, FRA. 11, noncarinate, uCMP50158, RQt.

12 Lenticulina tangens (LeRoy), uCMP50159, MiB.

13 Lenticulina thalmanni (hessland), uCMP50160, NAV5.

14 Lenticulina variabilis (Reuss), uCMP50161, PPP.

409



410


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 Schwager’s 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 6–7 chambers with short inner margins that rise around a pluglike 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 9–11 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 (d’orbigny 1839c)Plate 8, Figures 10, 11

Robulina subcultrata D’oRBiGNy 1839c, p. 98, pl. 5, figs. 19, 20.Lenticulina clericii (Fornasini). WhittAkER 1988, pl. 5, figs. 5, 6

(Miocene–Pliocene, 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 8–10 chambers in final whorl, sutures slightly curved, ranging from oblique to nearly radial.

Remarks: it is difficult to consistently differentiate d’orbigny’s 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 d’orbigny 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 13–15 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, 5–7 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.


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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 6–7 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.

Occurrence: Navidad Fm. (LBz, MoS, NAV5, PPP), Ranquil Fm. (RQt).

Maximum relative abundance: Common (LBz).

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. 11–16.Lenticulina variabilis Romanova, in GLAzuNoVA et al. 1960, p. 73,

pl. 12, figs. 3–8.

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. APlate 9, Figure 1

Lenticulina convergens (Bornemann). hoLBouRN, hENDERSoN and MACLEoD 2013, p. 332–333.

Distinguishing features: outline rounded, but final two chambers have nearly straight outer edges that adjoin at 135°; carinate, fusiform in edge view; 5½–7½ translucent chambers in final whorl; sutures slightly curved, flush; umbo large, flush, opaque; surface smooth.



Lenticulina sp. BPlate 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.



Lenticulina sp. CPlate 9, Figure 3

Distinguishing features: Sublacrimate outline, slightly compressed; outer whorl comprised of four chambersm, slight keel; surface smooth, translucent.



Lenticulina sp. DPlate 9, Figure 4

Distinguishing features: outline slightly elongate, noncarinate. moderately compressed in edge view; 5 tightly coiled, translucent chambers; moderatley arched, slightly depressed.

Occurrence: El Peral beds (LPER), Navidad Fm. (NAV5), Ranquil Fm. (FRA, LEB).


NEoLENtiCuLiNA McCulloch 1977Type species: Neolenticulina chathamensis McCulloch 1977.

Neolenticulina peregrina (Schwager 1866)Plate 9, Figures 5, 6

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 EDWARDS 1986, p. 92, pl. 27, figs. 1, 2.

Neolenticulina peregrina (Schwager). — hoLBouRN, hENDERSoN and MACLEoD 2013, p. 368–369.

Type age and locality: Middle Pliocene, Car Nicobar.



Occurrence: El Peral beds (NLP), Navidad Fm. (CPuP, PPP, PtA), Ranquil Fm. (FRA, FRM, RQt).


PERCuLtAzoNARiA Loeblich and tappan 1986Type species: Cristellaria subaculeata Cushman 1923.

Remarks: Differs from Vaginulinopsis by its more prominent coil and elevated ornamentation.

Percultazonaria encinasi Finger, n. sp.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 hookshaped test, more inflated later chambers, and weak sutural nodules. Cristellaria spinulosa Costa (in (Fornasini) 1894 (tertiary, italy)

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has ornamentation similar to the Chilean species, but it does not appear to uncoil.

Occurrence: Navidad Fm. (CPuP, LBz, MAt, MoS, NAV5, PPN, PPP, PPt, PtA, RAP), Ranquil Fm. (FRA, RAN, RQt), Lacui Fm. (Cho, CuC).


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 Concepcíon).

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. hornibrook’s (1971) illustration of a topotype that is slightly uncoiled and noncarinate might not be the same species. Both the Chilean form and karrer’s species appear to be juveniles that have yet to uncoil;

however, neither resemble the planispiral stage of the uncoiling Percultazonaria species they are associated with.

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.



Percultazonaria obliquispinata Finger, n. sp.Plate 9, Figure 9

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 (= Cris-tellaria 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).


Type specimens: holotype uCMP50170; paratypes uCMP50436 (from FRM) and uCMP50437 (from CPuP).

PLATE 9Figures 1a, 3b, 4b, 5b, 11a, 15b, 16b, 17b, and 18a are photomicrographs; all other images are SEMs. Scale bars in µm.

1 Lenticulina sp. A, uCMP50162, MiB.

2 Lenticulina sp. B, uCMP50163, FRM.

3 Lenticulina sp. C, uCMP50164, MiB.

4 Lenticulina sp. D, uCMP50165, PNh.

5, 6 Neolenticulina peregrina (Schwager): 5, uCMP50166, FRA; 6, juvenile form, uCMP50167, PPP.

7 Percultazonaria encinasi Finger, n. sp., holotype uCMP- 50168, CuC.

8 Percultazonaria cf. P. mamilligera (karrer), uCMP50169, FRM.

9 Percultazonaria obliquispina Finger, n. sp., holotype uCMP50170, RQt.

10 Percultazonaria vaughani (Cushman), uCMP50171, PPP.

11, 12 Saracenaria schencki Cushman and hobson, PPP: 11, uCMP50172. 12, uCMP50173.

13 Amphicoryna badenensis (d’orbigny), uCMP50174, RQt.

14 Amphicoryna cf. A. scalaris (Batsch), uCMP50175, PPP.

15 Astacolus crepidulus (Montfort), uCMP50176, LEB.

16 Astacolus cymboides (d’orbigny), uCMP50177, FRA.

17 Astacolus jordai (Colom), uCMP50178, FRM.

18 Astacolus cf. A. mayi (Cushman and Parker), uCMP50179, RAN.

19 Astacolus mexicanus (Nuttall), uCMP50180, FRM.

kenfinger

Sticky Note

obliquispinata

413



414


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 populations.

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 1824Type species: Saracenaria italica Defrance, in de Blainville 1824.

Saracenaria schencki Cushman and hobson 1935Plate 9, Figures 11, 12

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).

Type age and locality: Eocene (oligocene?), California.

Stratigraphic range: Eocene (oligocene?)–Middle Miocene.

Upper depth limit: outer neritic (for Saracenaria spp. occurrences off California (ingle 1980) and New zealand (hayward et al. 2010).

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.

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).


Subfamily MARGiNuLiNiNAE Wedekind 1937AMPhiCoRyiNA Schlumberger, in Milne-Edwards 1881Type species: Marginulina falx Jones and Parker 1860.

Amphicoryna badenensis (d’orbigny 1846)Plate 9, Figure 13

Nodosaria badenensis D’oRBiGNy 1846, p. 38, pl. 1, figs. 34, 35. — PAPP and SChMiD 1985, p. 27, pl. 7, figs. 1–8.


Comparative species: Amphicoryna scalaris (= Nautilus scalaris Batsch 1791; probably Recent, italy) and Amphicoryna striaticollis (= Nodosaria striaticollis d’orbigny 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 d’orbigny 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

Nautilus (Orthoceras) scalaris BAtSCh 1791, p. 1, 4, pl. 2, fig. 4.

Type age and locality: oligocene, hungary.

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: Astacolus tunicata (= Cristellaria (Marginulina) tunicata von hantken 1868; oligocene, hungary) is narrower and devoid of costae.



AStACoLuS de Montfort 1808Type species: Astacolus crepidulatus de Montfort 1808.

Astacolus crepidulus (Fichtel and Moll 1798)Plate 9, Figure 15

Nautilus crepidula FiChtEL and MoLL 1798, p. 107, pl. 19, figs. g–i. Astacolus crepidulus (Fichtel and Moll). RöGL 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. — hAyWARD, GRENFELL, SABAA, NEiL and BuzAS 2010, p. 175, pl. 14, figs. 1, 2.

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Type age and locality: Recent, Italy.


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 (d’orbigny 1846)Plate 9, Figure 16

Cristellaria cymboides D’oRBiGNy 1846, p. 85, pl. 3, figs. 30, 31.Lenticulina cymboides (d’orbigny). PAPP and SChMiD 1985, p. 39,

pl. 24, figs. 1–3.Type age and locality: Middle Miocene (Badenian), Vienna 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 d’orbigny 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. 1–5, 14, 15, pl. 24, figs. 74–76.

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 d’orbigny 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. 3–5.

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).


Astacolus mexicanus (Nuttall 1932)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.

Occurrence: Navidad Fm. (CPuP), Ranquil Fm. (FRA, FRM).


Astacolus multicameratus (Cushman and Stainforth 1945)Plate 10, Figure 1

Marginulina sublituus var. multicamerata CuShMAN and StAiNFoRth 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 Mar-ginulina 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. APlate 10, Figure 3



Astacolus sp. BPlate 10, Figure 4


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hEMiRoBuLiNA d’orbigny 1826Type species: Cristellaria arcuatula Stache 1865.

Hemirobulina pedum (d’orbigny 1846)Plate 10, Figure 5

Marginulina pedum D’oRBiGNy 1846, p. 68, pl. 3, figs. 13, 14.Vaginulinopsis pedum (d’orbigny). 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, Galápagos)Type age and locality: Middle Miocene, Austria.


Occurrence: Ranquil Fm. (FRA, FRM, MiB, RAN), Santo Domingo Fm. (VAL)


Hemirobulina similis (d’orbigny 1846)Plate 10, Figures 6, 7

Marginulina similis D’oRBiGNy 1846, p. 69, pl. 3, figs. 15, 16. — PAPP and SChMiD 1985, p. 37, pl. 21. figs. 10–12.

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. 346–347.


Comparative species: Hemirobulina glabra (= Marginulina glabra d’orbigny 1826; Pliocene, italy) has more chambers and transverse sutures.

Occurrence: El Peral beds (NLP), Navidad Fm. (CPuP, NAV5, PtA).


Hemirobulina yabei (Asano 1936a)Plate 10, Figure 8

Dentalina yabei ASANo 1936a, p. 329, pl. 36, figs. 10–12.

Type age and locality: Pliocene, Japan.


Remarks: this form is nearly uniserial with slightly diagonal sutures.

Occurrence: Navidad Fm. (CPuP, MoS, RAP).


MARGiNuLiNA d’orbigny 1826 Type species: Marginulina raphanus d’orbigny 1826.

Marginulina cubana Palmer 1940Plate 10, Figures 9, 10

Marginulina cubana PALMER 1940, p. 278, pl. 52, fig. 18.

PLATE 10Figures 1b, 3b, 4b, 16, 18a, 19a, 20a, and 22b are photomicrographs; all other images are SEMs. Scale bars in µm

1 Astacolus multicameratus (Cushman and Stainforth), uCMP50181, LBz.

2 Astacolus novambiguus Finger, n. sp., holotype uCMP-50182, PtA.

3 Astacolus sp. A, uCMP50183, FRA.

4 Astacolus sp. B, uCMP50184, RQt.

5 Hemirobulina pedum (d’orbigny), uCMP50185, FRA.

6, 7 Hemirobulina similis (d’orbigny): 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 Vaginulinopsis lueneburgensis (Clodius), uCMP50193, PtA.

14 Vaginulinopsis subelegans? (Parr), uCMP50194, VAL.

15 Vaginulinopsis sp. A, uCMP50195, VAL.

16 Vaginulinopsis sp. B, uCMP50196, NLP.

17 Vaginulinopsis sp. C, uCMP50197, MS10.

18 Planularia cassis (Fichtel and Moll), uCMP50198, PPP.

19 Prismatomorphia tricarinata (d’orbigny), uCMP50199, NLP.

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.

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418


Type age and locality: Late oligocene, Cuba.

Stratigraphic range: Late oligocene to Early Miocene.

Remarks: this appears to be a dimorphic species, as some specimens have an incomplete early spiral stage.

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.

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 1986Type species: Vaginulina tricarinata d’orbigny 1826.

Prismatomorphia tricarinata (d’orbigny 1826)Plate 10, Figure 19

Vaginulina tricarinata D’oRBiGNy 1826, p. 258; type-figure in PARkER, JoNES, and BRADy 1865, pl. 1, fig. 34 (after d’orbigny’s modèle no. 4).

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 1904aType species: Vaginulina soluta var. carinata A. Silvestri 1898.

Vaginulinopsis cf. V. chetae (Basov 1964)Plate 10, Figure 11

Lenticulina (Marginulinopsis) chetae BASoV 1964, p. 80, 81, pl. 1, fig. 7.

Type age and locality: Late Jurassic, Siberia.

Remarks: this robust form is very similar to Basov’s Jurassic species.

Comparative species: Resembles Vaginulinopsis sp. A (Pl. 10. Fig. 15) in lateral view but has an acute, not rounded, periphery.

Occurrence: Lacui Fm. (PCB).


Vaginulinopsis costatus (Batsch 1791)Plate 10, Figure 12

Nautilus (Orthoceras) costatus BAtSCh 1791, p. 1, 4, pl. 1, fig. 1.

Type age and locality: Neither designated; probably Recent, Adriatic coast of italy.

Distinguishing featuress: Slightly compressed test with 16 strong, continuous costae, but missing its later chambers.

Occurrence: Lacui Fm. (PNh).


Vaginulinopsis lueneburgensis (Clodius 1922)Plate 10, Figure 13

Cristellaria lüneburgensis 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).


Vaginulinopsis subelegans? (Parr 1950)Plate 10, Figure 14

Vaginulina subelegans PARR 1950, p. 326, pl. 11, fig. 20.

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. APlate 10, Figure 15



Vaginulinopsis sp. BPlate 10, Figure 16



Vaginulinopsis sp. CPlate 10, Figure 17

Occurrence: Quiriquina island, off Ranquil Fm. (MS10).


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

419


Planularia cassis (Fichtel and Moll 1798)Plate 10, Figure 18

Nautilus cassis FiChtEL and MoLL 1798, p. 95, figured vars. α, β, γ, δ, ε


Remarks: Rögl and hansen (1984) provide a thorough review of Fichtel and Moll’s 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 1860VAGiNuLiNA d’orbigny 1826; emend. Reuss 1862, 1874 and Marie 1941Type species: Nautilus legumen Linnaeus 1758.

Vaginulina alazanensis Nuttall 1932Plate 10, Figure 20

Vaginulina alazanensis NuttALL 1932, p. 17, pl. 1, figs. 11, 15.

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. 21–23.


Occurrence: Navidad Fm. (LBz, MAt, PPP, PtA, RAP), Ranquil Fm. (FRA, RAN), Lacui Fm. (Cho, CuC).

Maximum relative abundance: Common (Cho).

FAMiLy LAGENiDAE Reuss 1862LAGENA Walker and Boys 1784; emend. A. Silvestri 1902Type species: Serpula (Lagena) sulcata Walker and Jacob, in kanmacher 1798.

Lagena cf. L. alcocki White 1956Plate 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 1875Plate 10, Figure 24

Lagena alternans tERQuEM 1875, p. 425, pl. 1, fig. 4.

Type age and locality: Recent, off northern France.

Distinguishing features: Subspherical with numerous striae that extend from base and alternate between those that reach neck and those that stop about ¾ of the way.

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 1974Plate 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 1863aPlate 11, Figure 2

Lagena filicosta REuSS 1863a, p. 328, pl. 4, figs, 50, 51.

420


Type age and locality: Not designated; levels given in Recent and Pliocene.

Occurrence: Navidad Fm. (PPP, PPt, PtA).


Lagena perlucida (Montagu 1803)Plate 11, Figures 3, 4

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 ¼ of the way from base to neck, with 8–14 bladed costae and a long neck with costae coiled at 45º (Pl. 11, Fig. 4).



Lagena semistriata Williamson 1848Plate 11, Figure 5

Lagena striata var. semistriata var. ß 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 ¾ up from the base.

Occurrence: Ranquil Fm. (FRA), Lacui Fm. (Cho, PCB).


Lagena striata (d’orbigny 1839c)Plate 11, Figures 6, 7

PLATE 11Figures 27 and 30–34 are photomicrographs; all other images are SEMs. Scale bars in µm.

1 Lagena bassensis Collins, uCMP50205, NAV5.

2 Lagena filicostata Reuss, uCMP50206, PPP.

3, 4 Lagena perlucida (Montagu), LEB: 3, uCMP50207. 4, uCMP50208.

5 Lagena semistriata Williamson, uCMP50209, Cho.

6, 7 Lagena striata (d’orbigny): 6, uCMP50210, RAN. 7, uCMP50211, FRA.

8 Lagena cf. L. striata (d’orbigny), 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 (d’orbigny), uCMP50216, PtA.

13 Lagena sp. A, uCMP50217, PtA.

14 Lagena sp. B, uCMP50218, PCB.

15 Lagena sp. C, uCMP50219, PtA.

16 Lagena sp. D, uCMP50220, RAN.

17 Procerolagena distomum (Parker and Jones), uCMP50221, Cho.

18 Procerolagena ingens (Buchner), uCMP50222, FRA.

19 Procerolagena multilatera (McCulloch), uCMP50223, RAN.

20 Procerolagena? sp., uCMP50224, PtA.

21 Pygmaeoseistron asperoides (Galloway and Morrey), uCMP50225, FRA.

22 Pygmaeoseistron gibberum (Buchner), uCMP50226, FRA.

23 Pygmaeoseistron globulohispidum (McLean), uCMP-50227, FRA.

24 Pygmaeoseistron hispidum (Reuss), uCMP50228, PPP.

25 Pygmaeoseistron 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.

31 Pseudopolymorphina atlantica Cushman and ozawa, uCMP50235, PPN.

32 Pseudopolymorphina sp. A, uCMP50236, NLP.

33 Pseudopolymorphina sp. B, uCMP50237, MAt.

34 Sigmomorphina trinitatensis Cushman and ozawa, uCMP50238, VAL.

421



422


Oolina striata D’oRBiGNy 1839c, p. 21, pl. 5, fig. 12.Lagena striata (d’orbigny). 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.

Type age and locality: Recent, Falkland Islands, depth not given.


Distinguishing features: Subsperical with 20–30 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 (d’orbigny 1839c)Plate 11, Figure 8

Oolina striata D’oRBiGNy 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 Cžjžek 1848; tertiary, Austria) has nearly twice as many striae. Lagena ampliformis McCulloch 1977 (Recent, Galápagos 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

Oolina striatula EGGER 1857, p. 269, pl. 5, figs. 3–8.

Type age and locality: Miocene, Germany.

Distinguishing features: Spherical, with about 14 striae extending onethird to more than twothirds up from the aboral end. Upper part of neck missing on all recovered specimens.

Remarks: the type figures of L. striatula show three distinct morphotypes. the Chilean form matches Egger’s first illustrated specimen (his figs. 3 and 4).



Lagena substriata Williamson 1848Plate 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.

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.

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.

Occurrence: Navidad Fm. (MAt, PPt), Ranquil Fm. (RAN, RQk, RQt).


Lagena sulcata Walker and Jacob 1798Plate 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, 25–27, 33, 34. — hoLBouRN, hENDERSoN and MACLEoD 2013, p. 324–325.


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 d’orbigny 1939 (Recent, Falkland islands), which has a slightly less-rounded aboral end. those species might be variants of L. sulcata.

Occurrence: Navidad Fm. (CPuP), Ranquil Fm. (FRA, RAN, RQt).


Lagena vilardeboana (d’orbigny 1839c)Plate 11, Figure 12

Oolina vilardeboana D’oRBiGNy 1839c, p. 19, pl. 5, figs. 4, 5.Lagena vilardeboana (d’orbigny). 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.

Comparative species: See Lagena sulcata (Pl. 11, Fig. 11).

Occurrence: Navidad Fm. (PtA), Ranquil Fm. (RAN).


Lagena sp. APlate 11, Figure 13

423


Diagnostic features: Globular; ~20 faint striae; short, smooth collar around base of neck.

Occurrence: Navidad Fm. (PPP, PtA), Lacui Fm. (PCB).


Lagena sp. BPlate 11, Figure 14

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.

Remarks: Neck and aperture of single specimen missing, base eroded.

Occurrence: Lacui Fm. (PCB)


Lagena sp. CPlate 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.

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.

Occurrence: Ranquil Fm. (PtA).


Lagena sp. DPlate 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.

Type age and locality: Not designated; lectotype from Recent, coast of Norway.


Distinguishing features: Cylindrical with tapered ends; finely striate.

Comparative species: Hyalinonetrion ingens (= Lagena distoma var. ingens Buchner 1940; “fossil”, italy) is more inflated.

Occurrence: El Peral beds (NLP), Navidad Fm. (PPP, PtA), Ranquil Fm. (FRA, RQk), Lacui Fm. (Cho, PCB).

Maximum relative abundance: Rare

Procerolagena ingens (Buchner 1940)Plate 11, Figure 18

Lagena distoma var. ingens BuChNER 1940, p. 425, pl. 2, fig. 22.

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, 480–641m.

Upper depth limit: Lower bathyal.



Procerolagena? sp.Plate 11, Figure 20

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 1988Type 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.

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.

424


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. 48–50. — 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)Plate 11, Figure 23

Lagena globulohispida MCLEAN 1956, p. 331, pl. 39, fig. 7.

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. par-vulipora (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. 77–79.Lagena aspera (Reuss). BoLtoVSkoy and thEyER 1970, p. 338,

pl. 3, fig. 15.


Distinguishing features: Spherical with long, narrow neck and moderatesized 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).



Pygmaeoseistron parvuliporum (Bandy 1949b)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.

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 1956Type 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.

Type age and locality: Upper Cretaceous, Poland.

Remarks: this form has the deep radial grooves around the aperture, but lacks the apical protrusion of R. apiculata.



Family PoLyMoRPhiNiDAE d’orbigny 1839aSubfamily PoLyMoRPhiNiNAE d’orbigny 1839aALLANhANCoCkiA McCulloch 1977Type 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 d’orbigny 1839aType species: Polymorphina (les Globulines) gibba d’orbigny 1826.

Globulina pirula Egger 1857Plate 11, Figure 28

Polymorphina (Globulina) gibba var. pirula EGGER 1857, p. 290, pl. 13, figs. 11, 12.

Type age and locality: Miocene, Germany.

Occurrence: Lacui Fm. (PNh).

425



PoLyMoRPhiNA d’orbigny 1826Type species: Polymorphina burdigalensis d’orbigny 1826.

Remarks: Numerous genera have been differentiated among forms traditionally assigned to Polymorphina, but they are based on internal chamber arrangements determined by dissection or thinsectioning, tasks that are impractical in most studies, so I have retained the traditonally broad concept of Polymorphina here.

Polymorphina fistulosa Williamson 1858Plate 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 1928Type species: Pseudopolymorphina hanzawai Cushman and ozawa 1928.

Pseudopolymorphina atlantica Cushman and ozawa 1930Plate 11, Figure 31

Pseudopolymorphina atlantica CuShMAN and ozAWA 1930, p. 94, pl. 24, fig. 2.

Type age and locality: Recent, off eastern u.S. (161–8050m).

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.

Occurrence: Navidad Fm. (LBz, MAt, MPuP, PPN, PPP, RAP), Ranquil Fm. (LEB, RAN, RQt), Lacui Fm. (PCB, PNh).

Maximum relative abundance: Common (PPN).

Pseudopolymorphina sp. APlate 11, Figure 32

Occurrence: El Peral beds (NLP), Navidad Fm. (PPP), Ranquil Fm. (MiB).


Pseudopolymorphina sp. BPlate 11, Figure 33

Occurrence: Navidad (MAt), Lacui Fm.(PNh).

Maximum relative abundance: Common (PNh).

SiGMoMoRPhiNA Cushman and ozawa 1928Type species: Sigmomorphina (Sigmomorphina) yokoyamai Cushman and ozawa 1928.

Sigmomorphina trinitatensis Cushman and ozawa 1930Plate 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 (= Poly-morphina 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 d’orbigny 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 1884RAMuLiNA t.R. Jones, in J. Wright 1875; emend. Dakin 1906Type species: Ramulina laevis t.R. Jones, in J. Wright 1875.

Ramulina globulifera BradyPlate 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. 22–28. — RoBERtSoN 1998, p. 98, pl. 36, figs. 3, 4. — hAyWARD, GRENFELL, SABAA, NEiL and BuzAS 2010, p. 173, pl. 13, fig. 9.

Type age and locality: Recent, type locality not given; localities in North Atlantic or South Pacific, 265–1097m.

Upper depth limit: Neritic; based on occurrence off New zealand (hayward et al. 2010).

Distinguishing features: Spherical chamber with many tubular branches.

Occurrence: Navidad Fm. (PPP, PtA).


Ramulina pulchra Bermúdez 1949Plate 12, Figure 2

Ramulina globulifera var. pulchra BERMúDEz 1949, p. 164, pl. 11, fig. 12.

Type age and locality: Middle Miocene, Dominican Republic.

426


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 1923Subfamily ooLiNiNAE Loeblich and tappan 1961FAVuLiNA Patterson and Richardson 1988Type species: Entosolenia squamosa var. hexagona Williamson 1848.

Favulina favosopunctata (Brady 1881)Plate 12, Figure 3

Lagena favosopunctata 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 Brady’s specimen from the torres Strait, differing only in not having a multi-ringed 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. — RoBERtSoN 1998, p. 100, pl. 37, fig. 2. — zAPAtA and CEAR 2004, p. 31, pl. 11, fig. 12. — hoLBouRN, hENDERSoN and MACLEoD 2013, p. 382–383.

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 (d’orbigny 1839c)Plate 12, Figure 5

Oolina melo D’oRBiGNy 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 12Figures 1, 10, and 13 are photomicrographs; all other images are SEMs. Scale bars in µm.

1 Ramulina globulifera Brady, uCMP50239, PPP.

2 Ramulina pulchra Bermúdez, uCMP50240, FRA.

3 Favulina favosopunctata (Brady), uCMP50241, Cho.

4 Favulina hexagona (Williamson), uCMP50242, PPP.

5 Favulina melo (d’orbigny), uCMP50243, LEB.

6 Favulina squamosa (Montagu), uCMP50244, FRA.

7 Homalohedra sp., uCMP50245, PCB.

8 Lagnea cf. L. enderbiensis (Chapman), uCMP50246, PCB.

9 Lagnea sp., uCMP50247, RAN.

10 Oolina laevigata d’orbigny, uCMP50248, RQk.

11 Fissurina ambicarinata Finger, n. sp., holotype uCMP50249, RQt.

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.

18 Pseudoolina cf. P. fissurinea R. W. Jones, uCMP50255, VAL.

19 Pseudoolina sp., uCMP50256, PPP.

20 Parafissurina inermis (Buchner), uCMP50257, FRA.

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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. 28–32.

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 d’orbigny 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 1988Type 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 ¾ of the way to the aperture.



LAGNEA Popescu 1983Type species: Fissurina radiata Seguenza 1862.

Lagnea cf. L. enderbiensis (Chapman 1909)Plate 12, Figure 8

Lagena enderbiensis ChAPMAN 1909, p. 339, pl. 16, fig. 1.

Type age and locality: Recent, southern New zealand (155m).

Distinguishing features; Flat with fusiform outline and ridged perimeter.

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.



Lagnea sp.Plate 12, Figure 9

Distinguishing features; Lacrimate; no peripheral ridge.

Remarks: Represented by a single poorly preserved specimen.



ooLiNA d’orbigny 1839c Type species: Oolina laevigata d’orbigny 1839c.

Oolina laevigata d’orbigny 1839cPlate 12, Figure 10

Oolina laevigata D’oRBiGNy 1839c, p. 19, pl. 5, fig. 3. — LoEBLiCh 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 1923FiSSuRiNA Reuss 1850Type 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 ¾ the length of the chamber.

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 speciesgroup, as Fissurina carinata Reuss 1863a has priority.


Maximum relative abundance: Rare (4 specimens).

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).

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Etymology: Morphodescriptor derived from the Latin ambi (circumference) + carinata (carinate).

Fissurina cuculatta A. Silvestri 1902Plate 12, Figure 12

Fissurina cuculatta A. SiLVEStRi 1902, p. 146, figs. 23–25.

Type age and locality: Recent, tyrrenian Sea.

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



Fissurina marginata Seguenza 1862Plate 12, Figures 13, 14

Fissurina marginata SEGuENzA 1862, p. 66, pl. 2, figs. 27, 28. — LoEBLiCh and tAPPAN 1987, p. 465, figs. 5–7.

Type age and locality: Late Miocene, Sicily.


Remarks: the type figure is ovate and widest at midline, whereas the Chilean form is widest below midline.



Fissurina cf. F. marginata Seguenza 1862Plate 12, Figures 15

Fissurina cf. F. marginata SEGuENzA 1862, p. 66, pl. 2, figs. 27, 28.


Remarks: Differs from Fissurina marginata by being more lacrimate and compressed, and the worn carina appears to be ridged.



Fissurina cf. F. obvia Seguenza 1862Plate 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.



PSEuDooLiNA R. W. Jones 1984Type 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.



Pseudoolina sp.Plate 12, Figure 19

Distinguishing features: ovate outline, circular in apertural view; aperture a thickly lipped slit.

Occurrence: Navidad Fm. (PPP), Ranquil Fm. (RQk).


Subfamily PARAFiSSuRiNiNAE R.W. Jones 1984PARAFiSSuRiNA Parr 1947Type species: Lagena ventricosa A. Silvestri 1904b.

Parafissurina inermis (Buchner 1940)Plate 12, Figure 20

Lagena staphyllearia forma inermis BuChNER 1940, p. 523, pl. 25, fig. 522.

Type age and locality: Recent, Mediterranean.

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.



Family GLANDuLiNiDAE Reuss 1860Subfamily GLANDuLiNiNAE Reuss 1860GLANDuLiNA d’orbigny 1839aType species: Nodosaria (les Glandulines) laevigata d’orbigny 1826, subsequent designation by Cushman 1927f, p. 189.

Glandulina dentalinoides A. Silvestri 1903Plate 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: Spindleshaped with blunt aboral end.

Occurrence: Navidad Fm. (MoS). Lacui Fm. (Cho).

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Glandulina laevigata d’orbigny 1826Plate 13, Figures 2–5

Nodosaria (Glanduline) laevigata D’oRBiGNy 1826, p. 252, pl. 10, figs. 1–3.

Glandulina ovula D’oRBiGNy 1846, p. 29, pl. 1, figs. 4, 5.Glandulina laevigata d’orbigny. LoEBLiCh and tAPPAN 1987, pl.

468, figs. 1–4. — RoBERtSoN 1998, p. 112, pl. 43, fig. 4.

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 d’orbigny (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.

Occurrence: El Peral beds (LPER, NLP), Navidad Fm. (CPuP, MoS, MPuP, NAV5, PPP, PPt, PtA, RAP), Ranquil Fm. (all localities), Santo Domingo Fm. (VAL), Lacui Fm. (Cho, PCB, PNh).

Maximum relative abundance: Common (RQt).

Glandulina simplex hussey 1949Plate 13, Figure 6

Glandulina simplex huSSEy 1949, p. 130, pl. 26, fig. 22.

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 1980Superfamily CERAtoBuLiMiNoiDEA Cushman 1927cFamily CERAtoBuLiMiNiDAE Cushman 1927cSubfamily CERAtoBuLiMiNiNAE Cushman 1927cCERAtoBuLiMiNA toula 1915Type 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. 14–16. — RoBERtSoN 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 13Figures 1, 3–5, 9c, and 16 are photomicrographs; all other images are SEMs. Scale bars in µm.

1 Glandulina dentalinoides Silvestri, uCMP50258, MoS.

2–5 Glandulina laevigata d’orbigny: 2, uCMP50259, FRA. 3, 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 (d’orbigny), uCMP50266, FRA.

10, 11 Globigerina 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 Globorotalia praemenardii Cushman and Stainforth, uCMP50273, NLP.

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Type age and locality: Recent, indonesia (1061m).


Comparative species: Ceratobulimina haueri (= Rotalina hauerii d’orbigny 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 1937Subfamily EPiStoMiNiNAE Wedekind 1937hoEGLuNDiNA Brotzen 1948Type species: Rotalia elegans d’orbigny, 1826.

Hoeglundina elegans (d’orbigny 1826)Plate 13, Figure 9

Rotalia (Turbuline) elegans D’oRBiGNy 1826, p. 276, type-figure not given.

Hoeglundina elegans (d’orbigny). 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. 1–6. — VAN MoRkhoVEN, BERGGREN and EDWARDS 1986, p. 97, pl. 29, figs. 1a, b, 2a, b. — WhittAkER 1988, p. 110, pl. 14, figs. 17–19. — JoNES 1994, p. 104, pl. 105, figs. 3–6. —RoBERtSoN 1998, p. 114, pl. 44, fig. 2.— zAPAtA and CEAR 2004, p. 28, pl. 9, fig. 5–7. — hAyWARD, GRENFELL, SABAA, NEiL and BuzAS 2010, p. 183, pl. 16, figs. 16–21. — hoLBouRN, hENDERSoN and MACLEoD 2013, p. 298–299.

Type age and locality: Not given.


Diagnostic feature: Number of chambers in final whorl is 6–10.

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 1850Family RoBERtiNiDAE Reuss 1850Subfamily RoBERtiNiNAE Reuss 1850RoBERtiNA d’orbigny 1846; emend. höglund 1947Type species: Robertina arctica d’orbigny 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 d’orbigny 1846 (Middle Pliocene, Austria) emend. höglund 1947 has broader and more numerous chambers.



order GLoBiGERiNiDA Carpenter, Parker and Jones 1862Superfamily GLoBoRotALioiDEA Cushman 1927cFamily GLoBoRotALiiDAE Cushman 1927cGLoBoRotALiA Cushman 1927cType species: Pulvinulina menardii var. tumida Brady 1877.

Globorotalia miotumida Jenkins 1960Plate 13, Figure 14

Globorotalia menardii miotumida JENkiNS 1960, p. 362, pl. 4, figs. 9a–c.

Globorotalia miotumida Jenkins. — JENkiNS 1985, p. 278, figs. 7.12, 7.13 — SCott, BiShoP and BuRt 1990, p. 39, figs. 26–28.

Stratigraphic range: Middle Miocene to Early Pliocene (Jenkins 1985); base of N9–N18/19.

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 1939Plate 13, Figure 15

Globorotalia miozea Finlay 1939, p. 326, pl. 29, figs. 159–161.

Stratigraphic range: Globorotalia miozea ranges from late Early through Middle Pliocene (N6–N13).

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 1945Plate 13, Figure 16

Globorotalia praemenardii CuShMAN and StAiNFoRth 1945, p. 70, pl. 13, fig. 14. — kENNEtt and SRiNiVASAN 1983, p. 122, pl. 28, figs. 6–8. —BoLLi and SAuNDERS 1985, fig. 32.7.

Stratigraphic range: Late Middle Miocene zones N10–N13.



Globorotalia praescitula Blow 1959Plate 14, Figure

Globorotalia praescitula BLoW 1959, p. 221, pl. 19. figs. 128a–c. — kENNEtt and SRiNiVASAN 1983, p. 108, pl. 24, fig. 1; pl. 25, figs. 4–6. — SCott, BiShoP and BuRt 1990, p. 27, figs. 19, 30. — ChAiSSoN and LECkiE 1993, p. 162, pl. 4, figs. 7–11.

Globorotalia scitula praescitula Blow. BoLLi and SAuNDERS 1985, fig. 31.6.

Globorotalia praescitula Blow plexus. MAJEWSki 2010, p. 4, figs. 3, 5.

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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 1982Type 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) plexusPlate 14, Figure 2

Globorotalia bella JENkiNS 1967: 1069, fig. 3, nos. 1–6. — kENNEtt and SRiNiVASAN 1983, p. 174, pl. 43, figs. 1–3.

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 less-developed 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 fourchambered variant of Glr. mayeri, but in their 1985 publication they differentiated them and gave the Glr. opima nana-continuosa transitional forms a P22–N6 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 4–5 chambers in the outer whorl.

Occurrence: Navidad Fm. (NAV5), Ranquil Fm. (FRA, RQt), Lacui Fm. (Cho, PCB).


Paragloborotalia mayeri (Cushman and Ellisor 1939) plexusPlate 14, Figure 3

Globorotalia mayeri CuShMAN and ELLiSoR 1939, p. 11, pl. 2, fig. 4. — kENNEtt and SRiNiVASAN 1983, p. 174, pl. 43, figs. 4–6. — BoLLi and SAuNDERS 1985, p. 201, figs. 26.31–26.43.

Paragloborotalia mayeri (Cushman and Ellisor). SCott, BiShoP and BuRt 1990, p. 127, figs. 83, 84. — ChAiSSoN and LECkiE 1993, p. 164, pl. 8, figs. 16–20.

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 4–5.

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) plexusPlate 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. 3–5.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

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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) plexusPlate 14, Figures 5–8

Globorotalia zealandica hoRNiBRook 1958, p. 667, figs. 18 19, 30. — kENNEtt and SRiNiVASAN 1983, p. 108, pl. 25, figs. 1–3. —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 elongatearcuate 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 (N9–N15), 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 1957CAtAPSyDRAX Bolli, Loeblich and tappan 1957Type species: Globigerina dissimilis Cushman and Bermúdez 1937.

Catapsydrax dissimilis (Cushman and Bermúdez 1937)Plate 15, Figures 1−3

Globigerina dissimilis CuShMAN AND BERMúDEz 1937, p. 25, pl. 3, figs. 4–6.

Catapsydrax dissimilis (Cushman and Bermúdez). kENNEtt and SRiNiVASAN 1983, p. 22, pl. 2, figs. 3–8. — BoLLi and SAuNDERS 1985, p. 186, figs. 17.1–17.4. — 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 Bermúdez). BRöNNiMANN 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 1947Type 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. 7–9. — BoLLi and SAuNDERS 1985, figs. 15.4–15.7 — ChAiSSoN and LECkiE 1993, p. 159, pl. 9, figs. 14–16. — 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 14Figures 6a–c are photomicrographs; all other images are SEMs. Scale bars in µm.

1 Globorotalia praescitula Blow, uCMP50274, PPP.

2 Paragloborotalia bella (Jenkins), uCMP50275, FRA.

3 Paragloborotalia mayeri (Cushman and Ellisor) s.l., uCMP50276, PPt.

4 Paragloborotalia nana (Bolli), uCMP50277, NAV5.

5–8 Paragloborotalia zealandica (hornibrook) plexus: 5, 6, uCMP50278, PPP. 7, juvenile, uCMP50279, PPP. 8, Para-

globorotalia zealandica-Globorotalia miozea transitional, uCMP50280, PtA.

9 Globoquadrina praedehiscens Blow and Banner, uCMP- 50281, RAN.

10 Globoquadrina dehiscens (Chapman, Parr and Collins), uCMP50282, FRA.

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Globoquadrina praedehiscens Blow and Banner 1962Plate 14, Figure 9

Globoquadrina dehiscens praedehiscens BLoW and BANNER 1962, p. 116, pl. 15, figs. Q–S.

Globoquadrina praedehiscens Blow and Banner. kENNEtt and SRiNiVASAN 1983, p. 182, pl. 45, figs. 4–6. — BoLLi and SAuNDERS 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 1862Family GLoBiGERiNiDAE Carpenter, Parker and Jones 1862Subfamily GLoBiGERiNiNAE Carpenter, Parker and Jones 1862GLoBiGERiNA Delage and hérouard 1896Type species: Globigerina bulloides d’orbigny 1826.

Globigerina concinna Reuss 1850Plate 13, Figures 10, 11

Globigerina concinna REuSS 1850, p. 373, pl. 47, fig. 8.

Stratigraphic range: upper N6–N12, 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 (P19–N4B, 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 (N7–N12) 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 1962Plate 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. — kENNEtt and SRiNiVASAN 1983, p. 38, pl. 6, figs. 1–3. — 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: N7–N12, Late Early to late Middle Pliocene.

Comparative species: Globoturborotalita woodi (P21–N21) differs by its apertural lip and strongly pitted, cancellate texture (see Pl. 15, Fig. 8)

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 1937Plate 13, Figure 13

Globigerina venezuelana hEDBERG 1937, p. 681, pl. 92, fig. 72b. — BRöNNiMANN 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 SRiNiVASAN 1983, p. 180, pl. 44, figs. 5–7.

“Globigerina” venezuelana Hedberg. SPEzzAFERRi and PREMoLi-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 1927cType species: Globigerina aequilateralis Brady 1879b.

Globigerinella obesa (Bolli 1957)Plate 15, Figures 4, 5

Globorotalia obesa BoLLi 1957, p. 119, pl. 29, figs. 2, 3. — BRöNNiMANN 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. 3–5.

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-geri nella 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, umbilicalextraumbilical 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 1979Type species: Globigerina rubra d’orbigny 1839a.

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Globigerinoides primordius Blow and Banner 1962Plate 15, Figure 6

Globigerinoides quadrilobatus primordius BLoW and BANNER 1962, p. 115, pl. 9, figs. Dd–Ff ; p. 138, text-fig. 14: figs. Dd–Ff.

Globigerinoides primordius Blow and Banner. — kENNEtt and SRiNiVASAN 1983, p. 54, pl. 11, figs. 1–3. — BoLLi and SAuNDERS 1985, p. 195, fig. 20.16.

Stratigraphic range: Late oligocene to Early Miocene, zones P22 (N4A) and N5.

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, LPER), Navidad Fm. (CPuP, MAt, LBz, MS10, PPN, PPP, PPt, PtA), Ranquil Fm. (MiB, MS10, RQk).


Globigerinoides trilobus (Reuss 1850)Plate 15, Figure 7

Globigerina triloba REuSS 1850, p. 374, pl. 447, fig. 11.Globigerinoides trilobus (Reuss). — kENNEtt and SRiNiVASAN

1983, p. 62, pl. 13, figs. 1–3.

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).


GLoBotuRBoRotALitA hofker 1976Type species: Globigerina rubescens hofker 1956.

Globoturborotalita brazieri (Jenkins 1966)Plate 15, Figure 8

Globigerina brazieri JENkiNS 1966, p. 1098, fig. 6, nos. 43–51.

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)Plate 15, Figures 9, 10

Globigerina woodi JENkiNS 1960, p. 352, pl. 2, fig. 2. — kENNEtt and SRiNiVASAN 1983, p. 43, pl. 7, figs. 4–6. — 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

Jenkins 1964; Early Miocene, N4–N7). i have reidentified very largeapertured 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 largeapertured 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.

Occurrence: El Peral beds (NLP), Navidad Fm. (all localities), Ranquil Fm. (all localities except LBz), Lacui Fm. (ChE, Cho, CuC, PCB).


Subfamily oRBuLiNiNAE von Schultze 1854oRBuLiNA d’orbigny 1839aType species: Orbulina universa d’orbigny 1839a.

Orbulina universa d’orbigny 1839aPlate 15, Figure 12

Orbulina universa D’oRBiGNy 1839a, p. 106; v. 8, pl. 1, fig. 1. — kENNEtt and SRiNiVASAN 1983, p. 86, pl. 20, figs. 4–6. — JoNES 1994, p. 90, pl. 82, figs. 1–3. — BoLLi and SAuNDERS 1985, p. 201, figs. 23.1, 24.2.

Stratigraphic range: Early Middle Miocene zone N9 to Recent (kennett and Srinivasan 1983).

Occurrence: El Peral beds (LPER)


PRAEoRBuLiNA olsson 1964Type species: Globigerinoides glomerosa subsp. glomerosa Blow 1956.

Praeorbulina glomerosa Blow 1956, s.l.Plate 15, Figure 13

Globigerinoides glomerosa glomerosa BLoW 1956, p. 64. tf. 1, nos. 15–19; tf. 2, nos. 1, 2. — kENNEtt and SRiNiVASAN 1983, p. 82, pl. 18, figs. 5–8. — BoLLi and SAuNDERS 1985, p. 199, figs. 23.4, 24.5.

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 1958Superfamily BoLiViNoiDEA Glaessner 1937Family BoLiViNiDAE Glaessner 1937BoLiViNA d’orbigny 1839cType species: Bolivina plicata d’orbigny 1839c.

Remarks: Most species of Bolivina have their upper depth limit in the outer neritic or upper bathyal zone. Specimens of this

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genus are surprisingly rare in the Early Miocene assemblages from Chile.

Bolivina advena Cushman 1925Plate 16, Figure 1

Bolivina advena CuShMAN 1925, p. 29, pl. 5, fig. 1.


Occurrence: Navidad Fm. (NAV5), Lacui Fm. (PCB).


Bolivina aenariensis Costa 1856; emend. Sgarrella 1992Plate 16, Figures 2, 3

Bolivina aenariensis CoStA 1856, p. 297, pl. 15, fig. 1; emend. SGARRELLA 1992, p. 318, 320, 322, pl. 1, figs. 1–13, pl. 2, figs. 1–11.

Bolivina subaenariensis CuShMAN 1922a , p. 46, pl. 7, fig. 6. (Recent, Atlantic); see Sgarrella (1992).

Type age and locality: Age not indicated, italy; Sgarrella’s 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.

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


Bolivina alazanensis Cushman 1926bPlate 16, Figure 4

Bolivina alazanensis CuShMAN 1926b, p. 82, pl. 12, fig. 1.

Type age and locality: oligocene, Mexico.

Remarks: the illustrated specimen closely resembles that of Bolli, Beckmann and Saunders (1994, fig. 53.3).

Comparative species: the form recognized as Bolivina serrata (= B. subadvena var. serrata Natland 1938; Recent, California) by Whittaker (1988: pl. 13, figs. 1–3) 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 1930Plate 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 15Figures 13c, d are photomicrographs; all other images are SEMs. Scale bars in µm.

1–3 Catapsydrax dissimilis (Cushman and Bermúdez): 1, uCMP50283, RQt. 2, uCMP50284, RQt. 3, uCMP50285, MiB.

4, 5 Globigerinella obesa (Bolli): 4, uCMP50286, PPP. 5, uCMP50287, FRA.

6 Globigerinoides primordius Blow and Banner, uCMP50288, PPP.

7 Globigerinoides trilobus (Reuss), uCMP50289, PtA.

8 Globoturborotalita brazieri (Jenkins), uCMP50290, PPP.

9, 10 Globoturborotalita woodi (Jenkins): 9, uCMP50291, FRA. 10, uCMP50292, PPP.

11 Radiolarian, uCMP50293, RQk.

12 Orbulina universa d’orbigny, uCMP50294, LPER.

13 Praeorbulina glomerosa (Blow), uCMP50295, NLP.

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Bolivina pukeuriensis hornibrook 1961Plate 16, Figure 6

Bolivina pukeuriensis hoRNiBRook 1961, p. 76, pl. 9, figs. 172, 173.

Type age and locality: Early Miocene, New zealand.

Occurrence: Concepción (MS10), Navidad (PtA).

Maximum Relative Abundance: Few.

Bolivina tumida Cushman 1925Plate 16, Figure 7

Bolivina tumida CuShMAN 1925, p. 32, pl. 5, fig. 9.




Superfamily CASSiDuLiNoiDEA d’orbigny 1839aFamily CASSiDuLiNiDAE d’orbigny 1839aSubfamily CASSiDuLiNiNAE d’orbigny 1839aCASSiDuLiNoiDES Cushman 1927cType species: Cassidulina parkeriana Brady 1881.

Cassidulinoides californiensis Bramlette, in Woodring and Bramlette 1951Plate 16, Figure 8

Cassidulinoides californiensis Bramlette, in WooDRiNG and

BRAMLEttE 1951, p. 61, pl. 22, fig. 7.

Type age and locality: Late Miocene, California.

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).

Occurrence: Navidad Fm. (PtA).


Cassidulinoides porrectus (heron-Allen and Earland 1932)Plate 16, Figures 9, 10

Cassidulina crassa var. porrecta hERoN-ALLEN and EARLAND 1932, p. 358, pl. 9, figs. 34–37.

Cassidulinoides porrectus (heron-Allen and Earland). NoMuRA, 1999,

Type age and locality: Recent, between Punta Arenas and East Falkland Islands.



Globocassidulina Voloshinova 1960Type species: Cassidulina globosa von hantken 1876.

Remarks: Most Globocassidulina have their upper depth limits in the upper middle bathyal zone (ingle 1980).

PLATE 16Figures 5b, 9, 10, 11b, 12b, 13c, 14a, 15b, and 25 are photomicrographs; all other images are SEMs. Scale bars in µm.

1 Bolivina advena Cushman, uCMP50296, PCB.

2, 3 Bolivina aenariensis Costa: 2, uCMP50297, NLP. 3, uCMP50298, PPt.

4 Bolivina alazanensis Cushman, uCMP50299, PPP.

5 Bolivina arta MacFadyen, opposite side views, uCMP-50300, PPP.

6 Bolivina pukeuriensis hornibrook, uCMP50301, MS10.

7 Bolivina tumida Cushman, uCMP50302, MiB.

8 Cassidulinoides californiensis Bramlette, uCMP50303, PtA.

9, 10 Cassidulinoides porrectus (heron-Allen and Earland), PCB: 9, uCMP50304. 10, uCMP50305.

11 Globocassidulina chileensis Finger, n. sp., holotype uCMP- 50306, PNh.

12 Paracassidulina lobatula (Cushman), uCMP50307, MAt.

13 Globocassidulina quadrata (Cushman and hughes), uCMP50308, FRA.

14 Globocassidulina subglobosa (Brady), uCMP50309, PCB.

15 Planocassidulina curvicamerata Voloshinova, uCMP- 50310, PCB.

16 Ehrenbergina fyfei Finlay, uCMP50311, PPP.

17 Stainforthia cf. S. complanata (Egger), uCMP50312, MS10.

18 Rectuvigerina transversa (Cushman), uCMP50313, LEB.

19 Bulimina alazanensis Cushman, uCMP50314, CPuP.

20 Bulimina spicata Phleger and Parker, uCMP50315, FRA.

21, 22 Globobulimina pacifica Cushman, PPP: 21, uCMP50316; 22, uCMP50317.

23 Praeglobobulimina ovata (d’orbigny), uCMP50318, RQk.

24 Praeglobobulimina ovula (d’orbigny), uCMP- 50319, MiB.

25 Praeglobobulimina socialis (Bornemann): uCMP50320, RQk.

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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).

Type specimens: holotype uCMP50305; paratypes uCMP50432 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).

Etymology. Named for the country from which it is described.

Type age and locality: Early Miocene, Chile.

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 d’orbigny 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).

Maximum relative abundance: Very abundant.

Globocassidulina quadrata (Cushman and hughes 1925)Plate 16, Figure 13

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’). WhittAkER 1988, p. 107, pl. 14, figs. 8, 9.

Type age and locality: Pleistocene, California.

Occurrence: Ranquil Fm. (FRA, FRM, RQt).


Globocassidulina subglobosa (Brady 1881)Plate 16, Figure 14

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. 18–23. — JoNES 1994, p. 60, pl. 54, fig. 17. — WhittAkER 1988, p. 107, pl. 14, figs. 10, 11. — hoLBouRN, hENDERSoN and MACLEoD 2013, p. 264–265.

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).

PARACASSiDuLiNA Nomura 1983aType species: Cassidulina delicata Cushman 1927a.

Paracassidulina lobatula (kaiho 1984)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).


PLANoCASSiDuLiNA Gudina 1966; emend. Nomura 1999.Type species: Cassidulina norcrossi Cushman 1933c.

Planocassidulina curvicamerata (Voloshinova 1952)Plate 16, Figure 15

Cassidulina curvicamerata Voloshinova 1952, p. 92, pl. 2, fig. 9.

Type locality and age: Miocene, Sakhalin island, Russia.

Occurrence: Chiloe (PCB).


Subfamily EhRENBERGiNiNAE Cushman 1927cEhRENBERGiNA Reuss 1850Type species: Ehrenbergina serrata Reuss 1850.

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Ehrenbergina fyfei Finlay 1939cPlate 16, Figure 16

Ehrenbergina fyfei FiNLAy 1939c, p. 323, pl. 28, figs. 119, 121, 122.

Comparative species: Ehrenbergina compressa Cushman (1927a; Recent, Pacific off Panama, 783m) is of similar dimensions but the edges of its chamber are bluntly angled.

Type age and locality: Late Miocene, New zealand.

Occurrence: Navidad Fm. (CPuP, MoS, NAV5, PPP, PPt, PtA), Ranquil Fm. (MS10, RQt), Lacui Fm. (ChE, PCB).


Superfamily tuRRiLiNoiDEA Cushman 1927cFamily StAiNFoRthiiDAE Reiss 1963StAiNFoRthiA hofker 1956Type species: Virgulina concava höglund 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 1958Superfamily BuLiMiNoiDEA t. R. Jones, in J. Wright 1875Family SiPhoGENERiNoiDiDAE Saidova 1981.Subfamily SiPhoGENERiNiNAE Saidova 1981RECtuViGERiNA Mathews 1945Type 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. 8–10.

Rectuvigerina transversa (Cushman). FiNGER 1992, p. 82, pl. 21 , figs 1–6.

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).

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).

Occurrence: Navidad Fm. (MAt), Ranquil Fm. (FRA, FRM, LEB, RQk, RQt), Santo Domingo Fm. (VAL), Lacui Fm. (PCB).

Maximum relative abundance: Common (MAt).

Family BuLiMiNiDAE t. R. Jones, in J. Wright 1875BuLiMiNA d’orbigny 1826Type species: Bulimina marginata d’orbigny 1826.\

Bulimina alazanensis CushmanPlate 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). — MARtíNEz 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. 14–15. — hoLBouRN, hENDERSoN and MACLEoD 2013, p. 90–91.

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: Brady’s type figure of Bulimina rostrata shows a fusiform test that tapers at both ends and is finely pointed at its base. Bulimina truncana von Gümbel 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 1951Plate 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, downwardprojecting 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

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species are differentiated from B. spicata by the nature of their costae: B. inflata Seguenza 1862 (Pleistocene, italy) has morespinose 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 less-spinose 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. höglund 1947Type species: Globobulimina pacifica Cushman 1927c.

Remarks: Most Globobulimina have their upper depth limits in the upper middle bathyal zone (ingle 1980).

Globobulimina pacifica Cushman 1927gPlate 16, Figures 21, 22

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 1951Type species: Bulimina pyrula d’orbigny var. spinescens Brady 1884.

Praeglobobulimina ovata (d’orbigny 1846)Plate 16, Figure 23

Bulimina ovata D’oRBiGNy 1846, p. 85, pl. 11, figs. 13, 14. Praeglobobulimina ovata (d’orbigny), emend. hAyNES 1954, p. 150,

text-figs. 9–12; pl. 35, figs. 2, 3. (Paleocene, England)]Bulimina pyrula (d’orbigny). PAPP and SChMiD 1985, pl. 62, figs.

2–4.Praeglobobulimina ovata (d’orbigny). kohL 1985, p. 67, pl. 21, fig. 3.


Occurrence: Ranquil Fm. (MiB), Lacui Fm. (PCB).


Stratigraphic range: Paleocene to Middle Miocene.

Praeglobobulimina ovula (d’orbigny)Plate 16, Figure 24

Bulimina ovula D’oRBiGNy 1839c, p. 51, pl. 1, figs. 10, 11. — PAPP and SChMiD 1985, RESiG 1981, pl. 1, fig. 10.

Globobulimina ovula (d’orbigny). 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 d’orbigny 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 (d’orbigny). 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 d’orbigny; 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.

Occurrence: El Peral beds (LPER, NLP), Navidad Fm. (CPuP, 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. 13–16. — 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).

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PRotoGLoBoBuLiMiNA hofker 1951Type species: Bulimina pupoides d’orbigny 1846.

Protoglobobulimina pupoides (d’orbigny 1846)Plate 17, Figure 2

Bulimina pupoides D’oRBiGNy 1846, p. 185, pl. 11, figs. 11, 12.Bulimina pyrula (d’orbigny). PAPP and SChMiD 19785, p. 69, pl. 62,

figs. 2–4.Praeglobobulimina pupoides (d’orbigny). LoEBLiCh and tAPPAN

1987, pl. 572, figs. 1–6. — JoNES 1994, p. 55, pl. 50, figs. 14, 15. — hAyWARD, GRENFELL, SABAA, NEiL and BuzAS 2010, p. 192, pl. 17, figs. 35, 36.



Remarks: D’orbigny 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 d’orbigny’s 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).

Occurrence: Navidad Fm. (PPP, PtA), Ranquil Fm. (FRA, MiB, MS10, RQt) Lacui Fm. (PNh).


Family BuLiMiNELLiDAE hofker 1951EuBuLiMiNELLA Revets 1993Type 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 5–6 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 3–4 chambers per whorl.

Occurrence: Ranquil Fm. (RQk).


Family uViGERiNiDAE haeckel 1894Subfamily uViGERiNiNAE haeckel 1894CiPERozEA Vella 1961Type species: Siphogenerina ongleyi Finlay 1939c.

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).

Maximum relative abundance: Common (RAN).

Ciperozea multicostata (Cushman and Jarvis 1929)Plate 17, Figure 5

Siphogenerina multicostata CuShMAN and JARViS 1929, p. 14, pl. 3, fig. 6.

Rectuvigerina multicostata (Cushman and Jarvis). VAN MoRkhoVEN, BERGGREN and EDWARDS 1986, p. 115, pl. 36, figs. 1–4. — WhittAkER 1988, p. 72, pl. 9, figs. 12, 13. — RoBERtSoN 1998, p. 144, pl. 55, figs. 6, 7.

Ciperozoa multicostata (Cushman and Jarvis). LoEBLiCh and tAPPAN 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

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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 oligocene–Early 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 oligocene–Early Miocene, Puerto Rico) has a much more extensive uniserial stage and its costae are continuous across the sutures. Ciperozoa stonei (= Siphogenerina stonei Bermúdez 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 1951Type species: Uvigerina asperula var. ampullacea Brady 1884 (designated by thalmann 1952).

PLATE 17Figures 2, 3, 25, and 32a are photomicrographs; all other images are SEMs. Scale bars in µm.

1 Praeglobobulimina spinescens (Brady), uCMP50321, NAV5.

2 Protoglobobulimina pupoides (d’orbigny), uCMP50322, RQt.

3 Eubuliminella bassendorfensis (Cushman and Parker), uCMP50323, RQk.

4 Ciperozea basispinata (Cushman and Jarvis), uCMP-50324, FRA.

5 Ciperozea multicostata (Cushman and Jarvis), uCMP- 50325, CPuP.

6 Ciperozea ongleyi (Finlay), uCMP50326, PPP.

7 Neouvigerina auberiana (d’orbigny), uCMP50327, MiB

8, 9 Neouvigerina hispida (Schwager): 8, uCMP50328, MiB. 9, uCMP50329, PPP.

10 Neouvigerina gallowayi (Cushman), uCMP50330, PPP.

11 Neouvigerina schwageri (Brady), uCMP50331, NAV5.

12 Uvigerina hispidocostata Cushman and todd, uCMP-50332, LEB.

13, 14 Uvigerina kernensis Barbat and Estorff, PtA: 13, uCMP50333; 14, uCMP50334.

15 Uvigerina peregrina Cushman, uCMP50335, RQk.

16, 17 Trifarina angulosa (Williamson): 16, uCMP50336, NAV5. 17, uCMP50337, LEB.

18 Fursenkoina vicksburgensis (Cushman), uCMP50338, RQt.

19 Ellipsopolymorphina zuberi (Grzybowski), uCMP50339, FRA.

20 Virgulinella pertusa (Reuss), uCMP50340, Cho.

21 Obesopleurostomella 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 Siphonodosaria sentifera (Cushman and Parker), uCMP50347, LEB.

28, 29 Siphonodosaria lepidula (Schwager): 28, uCMP50348, PPP. 29, uCMP50349, FRA.

30, 31 Toddostomella hochstetteri (Schwager): 30, microspheric, uCMP50350, FRA. 31, megalospheric, uCMP50351, NLP.

32, 33 Cancris auriculus (Fichtel and Moll): 32, uCMP50352, NLP. 33, uCMP50353, PCB.

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Neouvigerina auberiana (d’orbigny 1839a)Plate 17, Figure 7

Uvigerina auberiana D’oRBiGNy 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. 6–9.

Siphouvigerina auberiana (d’orbigny). kohL 1985, p. 70, pl. 22, figs. 7, 8; pl. 23, fig. 1.

Type age and locality: Recent, type locality not designated; 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.

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.

Occurrence: Navidad Fm. (MoS, MPuP, PPP, PtA), Ranquil Fm. (MiB, RQk).


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.

Distinguishing features: Costae widely spaced, nearly linear, continuous.

Remarks: See remarks for Ciperozoa basispinata.


Maximum abundance: Few.

Neouvigerina hispida (Schwager 1866)Plate 17, Figures 8, 9

Uvigerina hispida SChWAGER 1866, p. 249, pl. 7, fig. 95. — MARtíNEz 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. 1–4. — RoBERtSoN 1998, p. 154, pl. 58, fig. 3. — hAyWARD, GRENFELL, SABAA, NEiL and BuzAS 2010, p. 203, pl. 21, figs. 4, 5.

Uvigerina rugosa d’orbigny. — WhittAkER 1988, p. 70, pl. 8, figs. 19, 20.


Stratigraphic range: Late oligocene zone P21 through Pleistocene zone N23.

Upper depth limit: Upper middle bathyal.

Comparative species: Neouvigerina rugosa (= Uvigerina rugosa d’orbigny; 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.

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

Uvigerina schwageri BRADy 1884, p. 575, pl. 74, figs. 8–10. — WhittAkER 1988, p. 71, pl. 8, fig. 11.

Type age and locality: Recent, Fiji islands, 384m.


Distinguishing features: Costae poorly defined, partial, few in number.

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 d’orbigny 1826Type species: Uvigerina pygmaea d’orbigny 1826.

Uvigerina hispidocostata Cushman and todd 1945Plate 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 MCCuLLoCh 1948, p. 267, pl. 34, fig. 3. (Recent, off southern California, 896m)

Uvigerina peregrina Cushman. WhittAkER 1988, pl. 8, fig. 1. (Recent, West Atlantic)

Type age and locality: Miocene, Jamaica.

Stratigraphic range: Miocene

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 1933Plate 17, Figures 13, 14

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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 1923Plate 17, Figure 15

Uvigerina peregrina CuShMAN 1923, p. 166, pl. 42, figs. 7–10. — MARtíNEz and PARADA 1968, pl. 1, figs. 10, 11. — WhittAkER 1988, pl. 8, fig. 3. — LoEBLiCh and tAPPAN 1987, pl. 573, figs. 24–28.

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 d’orbigny 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).


tRiFARiNA Cushman 1923Type species: Trifarina bradyi Cushman 1923.

Trifarina angulosa (Williamson 1858)Plate 17, Figures 16, 17

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. 5–9. — hAyWARD, GRENFELL, SABAA, NEiL and BuzAS 2010, p. 204, pl. 21, figs. 12–15.



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

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 1961Family FuRSENkoiNiDAE Loeblich and tappan 1961FuRSENkoiNA Loeblich and tappan 1961Type species: Virgulina squammosa d’orbigny 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.



Family ViRGuLiNELLiDAE Loeblich and tappan 1984ViRGuLiNELLA Cushman 1932aType 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. 20–22.

Virgulina miocenica CuShMAN and PoNtoN 1931, p. 32, pl. 4, figs. 14–16. (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?

Comparative species: Although the type description of Vir-gulinella 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.

Occurrence: Ranquil Fm. (RQk), Lacui Fm. (Cho).


Superfamily PLEuRoStoMELLoiDEA Reuss 1860Family ELLiPSoiDiNiDAE A. Silvestri 1923ELLiPSoPoLyMoRPhiNA A. Silvestri 1901Type 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)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

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Type age and locality: Early oligocene, Poland.

Stratigraphic range: Early oligocene to Early Miocene.


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. hayward’s 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.

Occurrence: Ranquil Fm. (FRA, FRM, RQt).


Family PLEuRoStoMELLiDAE Reuss 1860oBESoPLEuRoStoMELLA Hayward, in Hayward et al. 2012Type species: Pleurostomella bierigi Palmer and Bermúdez 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, SABAA, GRENFELL, VAN kERCkhoVEN, JohNSoN and thoMAS 2012, p. 221, pl. 33, figs. 3–8.




Comparative species: Obesopleurostomella recens (= Pleurosto-mella 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 1860Type species: Pleurostomella subnodosa (Reuss 1860).

Pleurostomella alternans Schwager 1866Plate 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. 10–15.


Stratigraphic range: Late Cretaceous–middle 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).


Family StiLoStoMELLiDAE Finlay 1947SiPhoNoDoSARiA A. Silvestri 1924Type species: Nodosaria abyssorum Brady 1881.

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)Plate 17, Figures 23, 24

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,



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)

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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 middlebathyal depth.




Siphonodosaria lepidula (Schwager 1866)Plate 17, Figures 28, 29

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.

1–7. — FiNGER, WEAVER, LiPPS and MiLLER 1990, pl. 1, figs. 43–45. — FiNGER 1992, p. 84, pl. 24, figs. 10–16. — FiNGER, NiELSEN, DEVRiES, ENCiNAS and PEtERSoN 2007, fig, 13F.

Siphonodosaria matanzana (Palmer and Bermúdez). RoBERtSoN 1998, p. 180, pl. 67, fig. 2.

Siphonodosaria subspinosa (Cushman). RoBERtSoN 1998, p. 180, pl. 67, fig. 3.

Siphonodosaria lepidula (Schwager). hAyWARD, kAWAGAtA, SABAA, GRENFELL, VAN kERCkhoVEN, JohNSoN and thoMAS 2012, p. 174, pl. 18, figs. 6–20.




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 (Miocene–Pliocene, 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 LeRoy’s 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,


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, kAWAGAtA, SABAA, GRENFELL, VAN kERCkhoVEN, JohNSoN and thoMAS 2012, p. 180, pl. 19, figs. 18–20.


Stratigraphic range: Late Cretaceous to Miocene.


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 d’orbigny 1846 (Middle Pliocene, Austria) is considerably stouter with less constricted sutures.



StRiCtoCoStELLA Patterson 1987Type species: Ellipsonodosaria modesta var. prolata Cushman and Bermúdez 1937.

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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. 7–10.

(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 karrer’s species anywhere in their book, but they reassigned Cushman and Laiming’s 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 d’orbigny 1826 (fossil, italy), which was based on synonymy with figures in Soldani (1789–1799), but a primary type was never described or figured; Parker, Jones and Brady (1865) subsequently examined Soldani’s 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. 2012Type species: Siphonodosaria chileana todd and kniker 1952.

Toddostomella hochstetteri (Schwager 1866)Plate 17, Figures 30, 31

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,



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).

Comparative species: Toddostomella spinosa (= Dentalina spinosa d’orbigny 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 1838Superfamily DiSCoRBoiDEA Ehrenberg 1838Family BAGGiNiDAE Cushman 1927cSubfamily BAGGiNiNAE Cushman 1927cCANCRiS de Montfort 1808Type species: Cancris auriculatus de Montfort 1808 = Nautilus auriculus Fichtel and Moll 1798.

Cancris auriculus (Fichtel and Moll 1798)Plate 17, Figures 32, 33

Nautilus auricula FiChtEL and MoLL 1798, p. 108, pl. 20, figs. a–f.Rotalia brongniartii D’oRBiGNy 1826, p. 273; type fig. not given.

(fossil, italy)Rotalia sagra D’oRBiGNy 1839a, p. 77, pl. 5, figs. 13–15. (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. 8–10; 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. 1–3. — JoNES 1994, p. 105, pl. 106, fig. 4. — zAPAtA and CEAR 2004, p. 19, pl. 3, fig. 1. — hoLBouRN, hENDERSoN and MACLEoD 2013, p. 134–135.

Cancris sagra (d’orbigny). WhittAkER 1988, p. 115, pl. 15, figs. 13–15 (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 Moll’s species illustrated in Le Calvez (1977) and Rögl and hansen (1984).

Occurrence: El Peral beds (LPER, NLP), Navidad Fm. (PPP, PtA), Lacui (PCB).


VALVuLiNERiA Cushman 1926dType species: Valvulineria californica Cushman 1926d.

Valvulineria ecuadorana Cushman and Stevenson 1948Plate 18, Figure 1

Valvulineria ecuadorana CuShMAN and StEVENSoN 1948, p. 64, pl. 10, fig. 12.

Type age and locality: Early Miocene, Ecuador.

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Stratigraphic age: Early Miocene.

Upper depth limit: Neritic (for V. araucana and V. inflata in zapata and Cear 2004).

Occurrence: Navidad Fm. (CPuP, MoS, PPP), Ranquil Fm. (MS10).


Family DiSCoRBiDAE Ehrenberg 1838EPoNiDES Montfort 1808Type species: Eponides repandus (Fichtel and Moll 1798)

Eponides orientalis Asano 1937Plate 18, Figure 2

Eponides orientalis ASANo 1937, p. 117, pl. 16, fig. 8.

Type age and locality: Recent, Japan



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 1932Plate 18, Figure 3

Eponides ouachitaensis hoWE and WALLACE 1932, p. 69, pl. 13, fig. 8.

Stratigraphic range: Eocene to Early Miocene

Upper depth limit: Neritic

Occurrence: Ranquil Fm. (RQk).


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 antil-larum d’orbigny 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).


oRBitiNA Sellier de Civrieux 1977Type species: Orbitina carinata Sellier de Civrieux 1977.

Orbitina parri (Collins 1974)Plate 18, Figure 5

Rosalina parri CoLLiNS 1974, p. 46, 47, pl. 3, fig. 36 (plesiotype); holotype figured in Parr 1945, pl. 10, fig. 3.

Type age and locality: holocene, off southeastern Australia (depth not given).


Remarks: Differs from holotype in having much finer radial grooves on the umbilical side.

Occurrence: Ranquil Fm. (LEB), Lacui Fm. (PCB).


Family RoSALiNiDAE Reiss 1963GAVELiNoPSiS hofker 1951Type species: Discorbina praegeri heron-Allen and Earland 1913.

Gavelinopsis alhamensis González-Donoso 1968Plate 18, figs. 6

Gavelinopsis alhamensis GoNzáLEz-DoNoSo 1968, p. 74, 76, pl. 1, fig. 7.

Type age and locality: Late Miocene, Spain.

Stratigraphic range: Early to Late Miocene.

Occurrence: Navidad Fm. (NAV5), Ranquil Fm. (LEB), Santo Domingo Fm. (VAL).


RoSALiNA d’orbigny 1826Type species: Rosalina globularis d’orbigny 1826.

Rosalina rugosa d’orbigny 1839cPlate 18, Figure 7

Rosalina rugosa D’oRBiGNy 1839c, p. 42, pl. 2, figs. 12–14.

Type age and locality: Recent, coast of Patagonia, Argentina (depth not given).


Comparative species: Rosalina candeina d’orbigny 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).

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Family SPhAERoiDiNiDAE Cushman 1927cSPhAERoiDiNA d’orbigny 1826Type species: Sphaeroidina bulloides d’orbigny 1826.

Sphaeroidina bulloides d’orbigny 1826Plate 19, Figure 8

Sphaeroidina bulloides D’oRBiGNy 1826, p. 257, modèle 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. 7–12. — VAN MoRkhoVEN, BERGGREN and EDWARDS 1986, p. 80, pl. 24, figs. 1, 2. — LoEBLiCh and tAPPAN 1987, pl. 617, figs. 1–6. — WhittAkER 1988, p. 107, pl. 14, figs. 12, 13. — JoNES 1994, p. 91, pl. 84, figs. 1–5. — 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 1952Family PARRELLoiDiDAE hofker 1956CiBiCiDoiDES thalmann 1939Type species: Truncatulina mundula Brady, Parker and Jones 1888.

Cibicidoides bradyi (trauth 1918)Plate 18, Figures 8–10

Truncatulina bradyi tRAuth 1918, p. 235; type-figure not designated.

Cibicidoides bradyi (trauth). iNGLE, kELLER and koLPACk 1980, pl. 6, figs. 11, 12. — WhittAkER 1988, p. 141, pl. 22, figs. 13–15. — 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. 1–3.

Cibicidoides robertsonianus (Brady). kohL 1985, p. 98, pl. 35, fig. 4.Cibicidoides ferasini (Conato). RoBERtSoN 1998, p. 202, pl. 78, fig.

4.

Type age and locality: Recent, type locality not designated; noted localities include the Atlantic, SW Pacific and Coral Sea.

Stratigraphic range: Late Early Eocene zone P9 through Pleis-tocene 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 6–10 chambers in the final whorl and a broadly rounded axial periphery and it grades into C. robertsonianus (Brady 1881), which has 13–14 chambers in the final whorl and an angular to carinate periphery. the Chilean Neogene specimens show the full range of edge variations but only 5–6 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 kolpack’s (1980) data on the Peru-Chile trench indicate.

PLATE 18Figures 2, 3, 4c, 5c, and 6c are photomicrographs; all other images are SEMs. Scale bars in µm.

1 Valvulineria ecuadorana Cushman and Stevenson, uCMP- 50354, MS10.

2 Eponides orientalis Asano, uCMP50355, Cho.

3 Eponides ouachitaensis (howe and Wallace), uCMP-50356, RQk.

4 Eponides parantillarum Galloway and heminway, uCMP- 50357, RAP.

5 Orbitina parri (Collins), uCMP50358, LEB.

6 Gavelinopsis alhamensis González-Donoso, uCMP- 50359, VAL.

7 Rosalina rugosa d’orbigny, uCMP50360, PtA.

8–10 Cibicidoides bradyi (trauth): 8, uCMP50361, PtA. 9, uCMP50362, NLP. 10, uCMP50363, FRA.

11, 12 Cibicidoides havanensis (Cushman and Bermúdez), NLP: 11, uCMP50364. 12, uCMP50365.

455



456


Comparative species: Cibicidoides robertsonianus (= Planorbulina (Truncatulina) robertsonianus Brady 1881; Caribbean, 2340m) has an acute, noncarinate edge and 13–14 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)Plate 19, Figures 1–4

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. 166–167.

Cibicidoides floridanus (Cushman). WhittAkER 1988, p. 143, pl. 20, figs. 4–6. — 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 10–16 chambers that are typically 2–4 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. 4–12) 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; oligocene–Miocene, 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 oligocene–Early Miocene, Venezuela, Mexico, and trinidad)

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).

Occurrence: Navidad Fm. (CPuP, MAt, MoS, MPuP, NAV5, PPP, PPt, PtA), Ranquil Fm. (all except MS10), Lacui Fm. (ChE, Cho, PCB, PNh).

Maximum relative abundance: Abundant (LEB).

Cibicidoides havanensis (Cushman and Bermúdez 1937)Plate 19, Figures 11, 12

Cibicides havanensis CuShMAN and BERMúDEz 1937, p. 28, pl. 3, figs. 1–3.

Cibicidoides havanensis (Cushman and Bermúdez). VAN MoRkhoVEN, BERGGREN and EDWARDS 1986, p. 189, pl. 64A, figs. 1–4; pl. 64B, figs. 1, 2. — RoBERtSoN 1998, p. 204, pl. 80, fig. 1. — hoLBouRN, hENDERSoN and MACLEoD 2013, p. 180–181.

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 deepwater 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)Plate 19, Figures 5, 6

Planulina renzi CuShMAN and StAiNFoRth 1945, p. 72, pl. 15, fig. 1. — VAN MoRkhoVEN, BERRGREN and EDWARDS 1986, p. 133, pl. 43A, figs. 1–5; pl. 43B, figs. 1, 2. — RoBERtSoN 1998, p. 214, pl. 85, fig. 2. — hoLBouRN, hENDERSoN and MACLEoD 2013, p. 410–411.

Cibicidoides renzi (Cushman and Stainforth). WhittAkER 1988, p. 147, pl. 21, figs. 1–6.

Type age and locality: Middle Miocene, trinidad.

457


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, 9–10 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 1952Subfamily PSEuDoPARRELLiNAE Voloshinova, in Voloshinova and Dain 1952PSEuDoPARRELLA Cushman and ten Dam 1948Type species: Pulvinulinella subperuviana Cushman 1926d.

Pseudoparrella naraensis kuwano 1950Plate 19, Figure 10

Pseudoparrella naraensis kuWANo 1950, p. 317, text-fig. 6 on p. 313.



Distinguishing features: 5–6 chambers in outer whorl, nearly smooth periphery, rounded edges.

Comparative species: Pseudoparrella exigua (= Pulvinulina exigua Brady 1884; Recent, 12 localities given in Atlantic, Pacific, and Southern oceans, 117–5011m) has a lobulate, subacute edge. Pulvinulinella subperuviana Cushman 1926d (Miocene, California) has 10–11 chambers in its outer whorl and a subacute edge.

Occurrence: Navidad Fm. (MAt, MoS, NAV5).


Family DiSCoRBiNELLiDAE Sigal 1952Subfamily DiSCoRBiNELLiNAE Sigal 1952LAtiCARiNiNA 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, BERGGREN and EDWARDS 1986, p. 89, pl. 26, fig. 1. — LoEBLiCh and tAPPAN 1987, pl. 631, figs. 1–4. — WhittAkER 1988, p. 128, pl. 17, figs. 19–21. — RoBERtSoN 1998, p. 210, pl. 84, figs. 1, 2. — hAyWARD, GRENFELL, SABAA, NEiL and BuzAS 2010, p. 213, pl. 24, figs. 19–21. — hoLBouRN, hENDERSoN and MACLEoD 2013, p. 328–329.

Type age and locality: Recent, type locality not designated; noted localities are in the Atlantic and indian oceans.

Stratigraphic range: Early oligocene zone P19 through Pleis-tocene 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 bathyalabyssal 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 1877Family PLANuLiNiDAE Bermúdez 1952PLANuLiNA d’orbigny 1826Type species: Planulina ariminensis d’orbigny 1826.

Planulina sp.Plate 19, Figure 11

Planulina sp. 3 of VAN MoRkhoVEN, EDWARDS and BERGGREN 1986, p. 55, pl. 17, figs. 1–3.

Stratigraphic range: Early Miocene through Pleistocene.

Upper depth limit: Bathyal.

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 1927cSubfamily CiBiCiDiNAE Cushman 1927cCIBICIDES de Montfort 1808Type species: Cibicides refulgens de Montfort 1808.

Cibicides cicatricosus (Schwager 1866)Plate 20, Figures 1–4

Anomalina cicatricosa SChWAGER 1866, p. 260, pl. 7, figs. 4, 108.Truncatulina akneriana (d’orbigny). 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. 7–9.

458


Cibicidoides cicatricosus (Schwager). RoBERtSoN 1998, p. 200, pl. 78, fig. 1. — hoLBouRN, hENDERSoN and MACLEoD 2013, p. 164–165.


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.

Occurrence: El Peral beds (LPER, NLP), Navidad Fm. (CPuP, LBz, PtA, RAP), Ranquil Fm. (all except MS10), Lacui Fm. (PCB, PNh, ChE).

Maximum relative abundance: Common (LBz, PCB)

Cibicides lobatulus? (Walker and Jacob 1798)Plate 20, Figure 5

Nautilus spiralis lobatulus Walker and Jacob, in kANMAChER 1798, p. 642, pl. 14, fig. 36.

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 1940Plate 20, Figures 6–8

Cibicides mediocris FiNLAy 1940, p. 464, pl. 67, figs. 198, 199.

Type age and locality: Middle Miocene, New zealand.

Stratigraphic range: Early to Middle Pliocene.

Remarks: Populations of this species include forms ranging from slightly concavo-convex to biconvex, with 6–9 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)Plate 20, Figures 9, 10

Operculina (?) umbonifera hoWChiN and PARR 1938, p. 309, pl. 18, figs. 3, 4, 6, 13, 14.

Type age and locality: Miocene, Australia.

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.


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

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.

Occurrence: Ranquil Fm. (LEB, RAN), Lacui Fm. (ChE, PCB)

PLATE 19Figs. 3a, 5a, 9b, and 11a are photomicrographs; all other images are SEMs.

1–4 Cibicidoides compressus (Cushman and Renz): 1, uCMP-50366, 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.

8 Sphaeroidina bulloides d’orbigny, uCMP50373, FRA.

9 Laticarinina pauperata (Parker and Jones), uCMP50374, FRA.

10 Pseudoparrella naraensis kuwano, uCMP50375, NAV5.

11 Planulina sp., uCMP50376, NLP.

459



460



FALSoCiBiCiDES Poignant 1958Type species: Falsocibicides aquitanicus Poignant 1958.

Falsocibicides sp.Plate 21, Figure 2

Upper depth limit: Cibicidids that conform to a hard substrate are generally restricted to innerneritic depths.

Occurrence: El Peral beds (NLP), Navidad Fm. (LBz, MoS), Ranquil Fm. (LEB, RAN), Lacui Fm. (CuC).


FoNtBotiA González-Donoso and Linares 1970Type 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. 10–12, 16–21.

Planulina wuellerstorfi (Schwager). VAN MoRkhoVEN, BERGGREN and EDWARDS 1986, p. 48, pl. 14, figs. 1, 2. — RoBERtSoN 1998, p. 216, pl. 86, fig. 2. — hoLBouRN, hENDERSoN and MACLEoD 2013, p. 416–417.

Fontbotia wuellerstorfi (Schwager). LoEBLiCh and tAPPAN 1987, pl. 634, figs. 10–12.

Cibicides wuellerstorfi (Schwager). hAyWARD, GRENFELL, SABAA, NEiL and BuzAS 2010, p. 210, pl. 23, figs. 17–22.


Stratigraphic range: Early Miocene through Pleistocene.

Upper depth limit: usually lower bathyal; based on van Mork-hoven, 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).


Superfamily NoNioNoiDEA von Schultze 1854Family NoNioNiDAE von Schultze 1854Subfamily NoNioNiNAE von Schultze 1854NoNioNELLA Cushman 1926cType species: Nonionella miocenica Cushman 1926c.

Nonionella miocenica Cushman 1926cPlate 21, Figures 4, 5

Nonionella miocenica CuShMAN 1926c, p. 64; type-figure not given. — iNGLE, kELLER and koLPACk 1981, pl. 2, figs. 15–18. — SAuNDERS and MüLLER-MERz 1982, p. 270, pl. 2, figs. 9–13. — LoEBLiCh and tAPPAN 1987, pl. 689, figs. 18–21. — WhittAkER 1988, p. 166, pl. 24, figs. 16–18.

Nonionella pulchella hADA 1931, p. 120, text-fig. 79. (Recent, Japan, 7–60m).

Nonionella auris (d’orbigny 1839c). BoLtoVSkoy and thEyER 1970, p. 323, pl. 4, fig. 8. (Recent, Chile, 88 & 120m).



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 1930Plate 21, Figure 6

Nonionella miocenica var. stella CuShMAN and MoyER 1930, p. 56, pl. 7, fig. 17.

Type age and locality: Recent, California, 64–166m.

Upper depth limit: Inner neritic.


Remarks: Cushman and Moyer (1930) believed the finger-like 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 20All images are SEMs. Scale bars in µm.

1–4 Cibicides cicatricosus (Schwager): 1, uCMP50377, RAN. 2, uCMP50378, FRA. 3, uCMP50379, FRA. 4, uCMP50380, FRM.

5 Cibicides lobatulus? (Walker and Jacob), uCMP50381, PCB.

6–8 Cibicides mediocris Finlay: 6, uCMP50382, PtA; 7, uCMP50383, Cho; 8, uCMP50384, PtA.

9, 10 Cibicides umboniferus (howchin and Parr): 8, uCMP-50385, PCB. 9, uCMP50386, RAN.

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462



PSEuDoNoNioN Asano 1936bType species: Pseudononion japonicum Asano 1936b.

Pseudononion communis (d’orbigny 1846)Plate 21, Figure 7

Nonionina communis D’oRBiGNy 1846, p. 106, pl. 5, figs. 7, 8.Nonion pizarrensis BERRy 1928, p. 269, text-figs. 1–3. (Recent, Peru,

48m) Nonionina pizarrense var. basispinata CuShMAN and MoyER

1930, p. 54, pl. 7, fig. 18. (Recent, California, 64–91m).Nonion commune (d’orbigny). PAPP and SChMiD 1985, p. 45, pl. 34,

figs. 1–5.Pseudononion pizarrensis (Berry). WhittAkER 1988, p. 171, pl. 24,

figs. 1–3, 9–11.


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. 2–5) illustrate specimens from d’orbigny’s 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, 210–300m); 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 Müller-Merz 1982Plate 21, Figure 8

Pseudononion cuevasensis SAuNDERS and MüLLER-MERz 1982, p. 267, pl. 4, figs. 34–38.

Pseudononion atlanticum (Cushman 1947). SAuNDERS and MüLLER-MERz 1982, p. 264, 268, pl. 3, figs. 19–25.

Florilus grateloupi (d’orbigny). kohL 1985, p. 92, pl. 32, fig. 3.Pseudononion grateloupi (d’orbigny). 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 d’orbigny 1839a (Recent, Caribbean) do not indicate the presence of extraumbilical pustules characteristic of Pseudononion, nor do the photomicrograph and description of its lectotype (Le Calvez 1977). Saunders and Müller-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 d’orbigny 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 Pliocene–Recent, 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)Plate 21, Figures 9, 10

Nonionella novo-zealandica CuShMAN 1936a, p. 88, pl. 13, fig. 16.

Type age and locality: Miocene, New zealand.


PLATE 21All images are SEMs. Scale bars in µm.

1 Cibicides sp., uCMP50387, RAN.

2 Falsocibicides sp., uCMP50388, LEB.

3 Fontbotia wuellerstorfi (Schwager), uCMP50389, FRA.

4, 5 Nonionella miocenica Cushman, NAV5: 4, uCMP50390. 5, uCMP50391.

6 Nonionella stella Cushman and Moyer, uCMP50392, Cho.

7 Pseudononion communis (d’orbigny), uCMP50393, NLP.

8 Pseudononion cuevasensis Saunders and Müller-Merz, uCMP50394, RAP.

9, 10 Pseudononion novozealandicum (Cushman): 9, uCMP- 50395, PPP. 10, uCMP50396, Cho.

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Comparative species: Pseudononion papillatum Galloway and heminway 1941 (Late oligocene, Puerto Rico) displays the earlier whorl on the nonpapillate spiral side.

Occurrence: Navidad Fm. (CPuP, MAt, MoS, MPuP, NAV5, 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.Plate 22, Figure 1

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.

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 1962Type species: Nonionella parri Cushman 1936a.

Zeaflorilus chiliensis (Cushman and kellett 1929)Plate 22, Figures 2–4

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 GutiéRREz 1995, p. 27, pl. 6, figs. 1–3.

Nonionella parri CuShMAN 1936a, p. 89, pl. 13, fig. 17. (Recent shore sand, New zealand)

Type age and locality: Recent, off Chile.


Upper depth limit: Inner neritic (based on Zapata, Zapata and Gutiérrez 1995).

Remarks: Most of the Chilean Miocene specimens have 14–18 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 d’orbigny 1846; Middle Miocene, Vienna Basin) has an

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).

Maximum relative abundance: Very abundant (ChE, PPN).

Subfamily AStRoNoNioNiNAE Saidova 1981FiJiNoNioN hornibrook 1964Type species: Astrononion fijiensis Cushman and Edwards 1937.

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.

Occurrence: El Peral beds (NLP), Lacui Fm. (PCB).


Subfamily PuLLENiNAE Schwager 1877MELoNiS de Montfort 1808Type species: Melonis etruscus de Montfort 1808 = Nautilus pompiliodes Fichtel and Moll 1798.

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,

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figs. 1, 2. — RESiG 1981, pl. 4, figs. 8, 9. — WhittAkER 1988, pl. 24, figs. 25–27. — kohL 1985, p. 100, pl. 36, fig. 4. — RoBERtSoN 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. 354–355.

Type age and locality: Recent, British Isles.Stratigraphic range: oligocene to Recent.

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 1958Plate 22, Figure 6

Nautilus pompilioides FiChtEL and MoLL 1798, p. 31, pl. 2, figs. a–c.

Nonionina soldanii D’oRBiGNy 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). MARtíNEz and PARADA 1968, pl. 1, figs. 31, 32. — iNGLE, kELLER and koLPACk 1980, pl. 9, fig. 14. — RöGL 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, GRENFELL, 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. 358–359.

Type age and locality: Pliocene, italy and Recent, Mediterranean Sea.

Stratigraphic range: Late oligocene through Pleistocene; Early Miocene to Pliocene for f. sphaeroides.

Upper depth limit: Globally neritic for Melonis pompilioides s.s., middle bathyal for f. sphaeroides (see remarks below), but the

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 Moll’s 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 Brady’s 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 1862Type species: Nonionina bulloides d’orbigny 1846.

Pullenia bulloides (d’orbigny 1846)Plate. 21, Figure 9

Nonionina bulloides D’oRBiGNy 1846, p. 107, pl. 5, figs. 9, 10.Pullenia bulloides (d’orbigny). iNGLE, kELLER and koLPACk

1980, pl. 5, fig. 7. — RESiG 1981, pl. 7, fig. 13. — PAPP and SChMiD 1985, p. 45, pl. 34, figs. 6–9. — WhittAkER 1988, p. 173, pl. 24, figs. 20–32. — JoNES 1994, p. 92, pl. 84, figs. 12, 13. — hAyWARD, GRENFELL, SABAA, NEiL and BuzAS 2010, p. 225, pl. 29, figs. 16, 17.



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).


Pullenia subcarinata (d’orbigny 1839c)Plate 22, Figures 10, 11

Nonionina subcarinata D’oRBiGNy 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.

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2. (Pliocene, California).Pullenia quinqueloba JoNES 1994, p. 92, pl. 84, figs. 14, 15.Pullenia subcarinata (d’orbigny) zAPAtA 1999, fig. 27. —

BoLtoVSkoy and thEyER 1970, p. 352, pl. 5, fig. 18. — WhittAkER 1988, p. 173, pl. 24, figs. 33–38.



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 4–6 chambers, with 5 chambers most common.

Comparative species: Pullenia inglei Finger and Lipps (in Finger et al. 1990; Miocene, California) has 5–6 chambers in the outer whorl and a more rounded peripheral edge. Pullenia malkinae Coryell and Mossman 1942 (Pliocene, Pacific coast of Panama) has 7–8 chambers in the outer whorl.

Occurrence: El Peral beds (NLP), Navidad Fm. (PPP), Ranquil Fm. (FRA, FRM, LEB, MiB, RAN, RQk, RQt), Santo Domingo Fm. (VAL), Lacui Fm. (PCB).


Superfamily ChiLoStoMELLoiDEA Brady 1881Family ChiLoStoMELLiDAE Brady 1881Subfamily ChiLoStoMELLiNAE Brady 1881ALLoMoRPhiNA Reuss, in Cžjžek 1849Type species: Allomorphina trigona Reuss 1850 (first species named).

Allomorphina pacifica Cushman and todd 1949Plate 22, Figure 12

Allomorphina pacifica CuShMAN and toDD 1949, p. 68, pl. 12, figs. 6–9. — JoNES 1994, p. 61, pl. 55, figs. 24–26.

Type age and locality: Pliocene, Fiji Islands.


Remarks: the recovered Chile specimens are relatively large.

Occurrence: Navidad Fm. (PtA), Ranquil Fm. (FRA, FRM).


ChiLoStoMELLA Reuss, in Cžjžek 1849Type species: Chilostomella ovoidea Reuss 1850.

Chilostomella ovoidea Reuss 1850Plate 23, Figure 1

Chilostomella ovoidea REUSS 1850, p. 380, pl. 48, fig. 12.— LoEBLiCh and tAPPAN 1987, pl. 701, figs. 6, 7. — RoBERtSoN 1998, p. 234, pl. 93, fig. 3. — FiNGER 1992, p. 88, pl. 34, figs, 19–28. — JoNES 1994, p. 61, pl. 55, figs. 15–16, 19–23. — hAyWARD, GRENFELL, SABAA, NEiL and BuzAS 2010, p. 207, pl. 21, figs. 36–40.

Chilostomella oolina Schwager 1878. iNGLE, kELLER and koLPACk 1981, pl. 5, figs. 9, 10. — zAPAtA and CEAR 2004, p. 20, fig. 1. — hoLBouRN, hENDERSoN and MACLEoD 2013, p. 148–149.

Type age and locality: tertiary, Poland.

Stratigraphic range: Miocene or earlier to Recent.


Occurrence: Navidad Fm. (CPuP, MoS, PPP, PPt, PtA), Ranquil Fm. (MiB, RQk, RQt).


hiDiNA Gheorghian, iva and Gheorghian 1968Type species: Hidina variabilis Gheorghian, iva and Gheorghian 1968.

Hidina variabilis Gheorghian, iva and Gheorghian 1968Plate 23, Figure 2

Hidina variabilae GhEoRGhiAN, iVA and GhEoRGhiAN 1968, p. 195, pl. figs. 1–4.

Hidina variabilis Gheorghian, iva and Gheorghian. LoEBLiCh and tAPPAN 1987, pl. 701, figs. 11–14.

PLATE 22Figures 3a and 5 are photomicrographs; all other images are SEMs. Scale bars in µm.

1 Pseudononion ranquilensis Finger, n. sp., holotype uCMP- 50397, MiB.

2–4 Zeaflorilus chiliensis (Cushman and kellett): 2, uCMP-50398, PPN. 3, uCMP50399, LEB; 4, uCMP50400, RQS.

5 Fijinonion obesum (Carter), uCMP50401, NLP.

6 Melonis pompilioides (Fichtel and Moll), uCMP50402, NAV5.

7 Melonis affinis (Reuss), uCMP50403, PCB.

8 Melonis barleeanus (Williamson), uCMP50404, NAV5.

9 Pullenia bulloides (d’orbigny), uCMP50405, FRA.

10, 11 Pullenia subcarinata (d’orbigny): 10, 4-chambered variant (f. quinqueloba), uCMP50406, FRM. 11, uCMP50407, FRA.

12 Allomorphina pacifica Cushman and todd, uCMP50408, FRM.

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Type age and locality: Miocene, Romania.

Distinguishing features: Robust, subspherical test showing 3–4 chambers and pronounced flange-like apertural lip.

Comparative species: Hidina hidensis Popescu 1975 (Early Miocene, Romania) and Hidina cubensis (= Allomorphina cubensis Palmer and Bermúdez 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 1981QuADRiMoRPhiNA Finlay 1939bType 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 1964oSANGuLARiA Brotzen 1940Type 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. 1–6. — 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. 7–9. — hoLBouRN, hENDERSoN and MACLEoD 2013, p. 386–387.

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.

Occurrence: El Peral beds (NLP), Ranquil Fm. (FRA, MiB, RAN, RQk, RQt), Lacui (PNh).


Family oRiDoRSALiDAE Loeblich and tappan 1984oRiDoRSALiS Andersen 1961Type 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. 1–3.— zAPAtA and CEAR 2004, p. 32, pl. 12, fig. 5. — hAyWARD, GRENFELL, SABAA, NEiL and BuzAS 2010, p. 227, pl. 30, figs. 3–6. — hoLBouRN, hENDERSoN and MACLEoD 2013, p. 384–385.

Oridorsalis tener (Brady) iNGLE, kELLER and koLPACk 1980, pl. 5, figs. 5, 6.

Oridorsalis tenera (Brady) LoEBLiCh and tAPPAN 1987, pl. 708, figs. 9–11.

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 Gonzáles-Donoso 1969ANoMALiNoiDES Brotzen 1942Type 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). MARtíNEz and PARADA 1968, pl. 1,

figs. 15–17.Anomalinoides salinasensis (kleinpell). WhittAkER 1988, p. 140,

pl. 23, figs. 10–12. — FiNGER 1992, p. 88, pl. 35, figs. 30–35.

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 kleinpell’s type description, it is somewhat variable. the outer whorl of this form has 10–11 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

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figure, however, shows only the involute umbilical side and 9 chambers.

Occurrence: El Peral beds (LPER, NLP), Navidad Fm. (CPuP, PPP, NAV5, MPuP, CPuP), Ranquil Fm. (LEB), Santo Domingo Fm. (VAL), Lacui Fm. (PCB, PNh).


Family GAVELiNELLiDAE hofker 1956Subfamily GAVELiNELLiNAE hofker 1956GyRoiDiNA d’orbigny 1826Type species: Gyroidina orbicularis d’orbigny 1826.

Gyroidina laevigata d’orbigny 1826Plate 23, Figure 7

Gyroidina laevigata D’oRBiGNy 1826, p. 278; figure in PARkER, JoNES and BRADy 1871, pl. 12, fig. 130; also see text-fig. of FoRNASiNi 1898, p. 52.



Remarks: the Chilean forms are small with a circular outline and about 10 slightly inflated, nearly equidimensional chambers separated by slightly limbate, curved sutures.

Comparative species: Gyroidina laevis (= Rotalia laevis d’orbigny 1852; tertiary, localities given in italy) has curved sutures with chambers that are about twice as high as broad. Gyroidina orbicularis d’orbigny 1826 (Recent, Adriatic Sea) has 6 outer chambers that are 2–3 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.

Occurrence: El Peral beds (NLP), Ranquil Fm. (FRA, MiB, RAN), Lacui Fm. (PCB, PNh).


Gyroidina sp. Plate 22, Figure 8

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.



GyRoiDiNoiDES Brotzen 1942Type species: Rotalina nitida Reuss 1844

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.

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 1987Type species: Gyroidina soldanii d’orbigny 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. 16–18.

Gyroidinoides altiformis (R.E. and k.C. Stewart). kohL 1985, p. 95, pl. 34, fig. 3. — RoBERtSoN 1998, p. 242, pl. 98, fig. 1.

Type age and locality: Early Pliocene, California.

Stratigraphic range: Late oligocene to Recent.

Upper depth limit: Generally outer neritic.

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 (d’orbigny 1826)Plate 23, Figure 11

Gyroidina soldanii D’oRBiGNy 1826, p. 278; type-figure not designated. — iNGLE, kELLER and koLPACk 1980, pl. 7, figs. 12, 13.

Gyroidinoides soldanii (d’orbigny). MARtíNEz and PARADA 1968, pl. 1, figs. 1, 2. — JoNES 1994, p. 106, pl. 107, figs. 6, 7. — hoLBouRN, hENDERSoN and MACLEoD 2013, p. 278–279.

Hansenisca soldanii (d’orbigny). LoEBLiCh and tAPPAN 1987, p. 106, pl. 719, figs. 5–9.




Remarks: D’orbigny’s earliest representation of this species is modèle 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. D’orbigny’s (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

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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. 16–18) identified a biconvex form, lenticular in edge view, with nearly radial sutures in the Late oligocene–Pliocene of Ecuador as Gyroidina soldani (d’orbigny). 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 1944Type species: Hanzawaia nipponica Asano 1944.

Hanzawaia cf. H. nipponica Asano 1944Plate 24, Figures 1, 2

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: Miocene–Recent.

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 1981BuCCELLA Andersen 1961Type species: Eponides hannai Phleger and Parker 1951.

Buccella peruviana (d’orbigny 1839c)Plate 24, Figures 4, 5.

Rotalina peruviana D’oRBiGNy 1839c, p. p. 41, pl. 1, figs. 12–14.Buccella peruviana (d’orbigny). 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. 2A–2o.

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 (Calvco-Marcilese 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 23Figs. 4a, 6c, 7c, 10c, and 11c are photomicrographs; all other images are SEMs. Scale bars in µm.

1 Chilostomella ovoidea Reuss, uCMP50409, RQt.

2 Hidina variabilis Gheorghian, iva and Gheorghian, uCMP50410, MiB.

3 Quadrimorphina glabra (Cushman), uCMP50411, RQt.

4 Osangularia culter (Parker and Jones), uCMP50412, FRA.

5 Oridorsalis umbonatus (Reuss), uCMP50413, FRA.

6 Anomalinoides salinasensis (kleinpell), uCMP50414, CPuP.

7 Gyroidina laevigata d’orbigny, uCMP50415, FRA.

8 Gyroidina sp., uCMP50416, MiB.

9 Gyroidinoides umbonatus (A. Silvestri), uCMP50417, NAV5.

10 Hansenisca altiformis (R. E. and k. C. Stewart), uCMP50418, PPP.

11 Hansenisca soldanii (d’orbigny), uCMP50419, FRA.

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472


Pleistocene, oregon) has 8–10 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 1839Family ELPhiDiiDAE Galloway 1933Subfamily ELPhiDiiNAE Galloway 1933CRiBRoELPhiDiuM Cushman and Brönnimann 1948Type species: Cribroelphidium vadescens Cushman and Brönnimann 1948.

Cribroelphidium hauerinum (d’orbigny 1846)Plate 24, Figure 7

Polystomella hauerina D’oRBiGNy 1846, p. 122, pl. 6, figs. 1, 2.Elphidium hauerinum (d’orbigny). PAPP and SChMiD 1985, p. 49,

pl. 38, figs. 5–10.




Comparative species: Cribroelphidium trinitatensis Cushman and Brönnimann 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 1808Type species: Nautilus macellus var. β Fichtel and Moll 1798.

Elphidium macellum (Fichtel and Moll 1798)Plate 24, Figure 9

Nautilus macellus var. β FiChtEL AND MoLL 1798, p. 68, pl. 10, figs. h, i, k; type-figures also in RöGL 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 1839RotALiNoiDES Saidova 1975Type species: Rotalia papillosa Brady 1884.

Rotalinoides margaritifera (Brady 1881)Plate 24, Figures 6, 8, 10

Planorbulina (Truncatulina) margaritifera BRADy 1881, p. 66; type-fig. 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 91–1448m)

Rotalinoides compressiusculus (Brady). JoNES 1994, p. 106, pl. 107, figs. 1, 3.

Type age and locality: Recent, off the Philippines, 174–183m.



Remarks: Brady’s 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).



1, 2 Hanzawaia cf. H. nipponica Asano, uCMP50420, PPP.

3 Hanzawaia strattoni (Applin), uCMP50421, RAP.

4, 5 Buccella peruviana (d’orbigny): 4, uCMP50422, PPN. 5, uCMP50423, PtA.

6, 8, 10 Rotalinoides margaritifera (Brady): 6, uCMP- 50424, PPN. 8, uCMP50425, VAL. 10, uCMP50426, LEB.

7 Cribroelphidium hauerinum (d’orbigny), uCMP50427, CHE.

9 Elphidium macellum (Fichtel and Moll), uCMP50428, CHE.

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——— , 1911. A monograph of the foraminifera of the North Pacific Ocean. Part II – Textulariidae. Washington DC: united States National Museum. Bulletin 71(2), 108 pp.

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——— , 1918a. The smaller fossil foraminifera of the Panama Canal Zone. Washington DC: united States Geological Survey. Bulletin 103, 89–102.

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——— , 1921. Foraminifera of the Philippine and adjacent seas. Washington, DC: united States National Museum. Bulletin 100(4), 608 pp.

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——— , 1924. Samoan foraminifera. Washington DC: Carnegie institution. Publication 342 (Department of Marine Biology, Paper 21), 75 pp.

——— , 1925. Some textulariidae from the Miocene of California. Con-tributions from the Cushman Laboratory for Foraminiferal Re-search, 1: 29–35.

——— , 1926a. the Foraminifera of the Velasco Shale of the tampico embayment. American Association of Petroleum Geologists Bulle-tin, 10: 581–612.

——— , 1926b. Some fossil bolivinas from Mexico. Contributions from the Cushman Laboratory for Foraminiferal Research, 1: 81–85.

——— , 1926c. Miocene species of Nonionina from California. Contri-butions from the Cushman Laboratory for Foraminiferal Research, 1: 89–92.

——— , 1926d. Foraminifera of the typical Monterey of California. Contributions from the Cushman Laboratory for Foraminiferal Re-search, 2: 53–69.

——— , 1927a. Recent foraminifera from off the West Coast of America. La Jolla: Scripps institution of oceanography. technical Series Bulletin 1, 119–188.

——— , 1927b. Some new genera of the Foraminifera. Contributions from the Cushman Laboratory for Foraminiferal Research, 2: 77–85.

——— , 1927c. An outline of a re-classification of the foraminifera. Con-tributions from the Cushman Laboratory for Foraminiferal Re-search, 3: 1–105.

——— , 1927d. New and interesting foraminifera from Mexico and texas. Contributions from the Cushman Laboratory for Foraminiferal Research, 3: 111–119.

——— , 1927e. Some characteristic Mexican fossil foraminifera. Journal of Paleontology, 1: 147–172.

——— , 1927f. the designation of some genotypes in the foraminifera. Contributions from the Cushman Laboratory for Foraminiferal Re-search, 3: 189–190.

——— , 1928. Additional genera of the foraminifera. Contributions from the Cushman Laboratory for Foraminiferal Research, 4: 1–8.

——— , 1929. A late tertiary fauna of Venezuela and other related regions. Contributions from the Cushman Laboratory for Foramin-iferal Research, 5: 77–101.

——— , 1930a. The foraminifera of the Choctawhatchee Formation of Florida. tallahassee: Florida State Geological Survey. Bulletin 4, 89 pp.

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——— , 1932a. Notes on the genus Virgulina. Contributions from the Cushman Laboratory for Foraminiferal Research, 8: 7–23.

——— , 1932b. the genus Vulvulina and its species. Contributions from the Cushman Laboratory for Foraminiferal Research, 8: 75–85.

——— , 1933a. New Arctic foraminifera collected by Captain R. A. Bart-lett from Fox Basin and off the northeast coast of Greenland. Wash

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——— , 1933c. Some new Recent foraminifera from the tropical Pacific. Contributions from the Cushman Laboratory for Foraminiferal Re-search, 9: 77–95.

——— , 1936a. Some new species of Elphidium and related genera. Con-tributions from the Cushman Laboratory for Foraminiferal Re-search, 12: 78–89.

——— , 1936b. New genera and species of the families Verneuilinidae and Valvulinidae and of the subfamily Virgulininae. Sharon, Massachusetts: Cushman Laboratory for Foraminiferal Research. Special Publication 6, 71 pp.

——— , 1937. A monograph of the foraminiferal families Verneuilinidae and Valvulinidae and of the subfamily Virgulininae. Sharon, Massachusetts: Cushman Laboratory for Foraminiferal Research. Special Publication 7, 157 pp.

——— , 1946. the genus Sigmoilina and its species. Contributions from the Cushman Laboratory for Foraminiferal Research, 22: 29–45.

CuShMAN, J. A. and BERMúDEz, P. J., 1937. Further new species of foraminifera from the Eocene of Cuba. Contributions from the Cushman Laboratory of Foraminiferal Research, 13: 1–29.

CuShMAN, J. A. and BRöNNiMANN, P., 1948. Some new genera and species of foraminifera from brackish water of trinidad. Contri-butions from the Cushman Laboratory for Foraminiferal Research, 24: 68–75.

CuShMAN, J. A. and DAM, A. TEN, 1948. Pseudoparrella, a new generic name and a new species of Parrella. Contributions from the Cushman Laboratory for Foraminiferal Research, 24: 49–50.

CuShMAN, J. A. and EDWARDS, P. G., 1937. Astrononion, a new genus of the Foraminifera and its species. Contributions from the Cushman Laboratory for Foraminiferal Research, 13: 29–36.

——— , 1938. Notes on the oligocene species of Uvigerina and Angulo-generina. Contributions from the Cushman Laboratory for Fora-miniferal Research, 14: 74–89.

CuShMAN, J. A. and ELLiSoR, A. C., 1939. New species of foraminifera from the oligocene and Miocene. Contributions from the Cushman Laboratory for Foraminiferal Research, 15: 1–14.

CuShMAN, J. A. and hARRiS, R. W., 1927. Some notes on the genus Ceratobulimina. Contributions from the Cushman Laboratory for Foraminiferal Research, 3(4): 171–179.

CuShMAN, J. A. and hoBSoN, h. D. 1935, A foraminiferal faunule from the type San Lorenzo Formation, Santa Cruz County, California. Contributions from the Cushman Laboratory for Foraminiferal Research, 11: 53–64.

CuShMAN, J. A. and huGhES, D. D., 1925. Some later tertiary Cassidulinas of California. Contributions from the Cushman Laborato-ry for Foraminiferal Research, 1(5): 11–17.

CuShMAN, J. A. and JARViS, P. W., 1929. New foraminifera from trinidad. Contributions from the Cushman Laboratory for Fora-miniferal Research, 5: 6–17.

——— , 1934. Some interesting new uniserial foraminifera from trinidad. Contributions from the Cushman Laboratory for Foraminifer-al Research, 10: 71–75.

CuShMAN, J. A. and kELLEtt, B., 1929. Recent foraminifera from the west coast of South America. Proceedings of the United States National Museum, 75: 1–16.

CuShMAn, J. A. and KlEinpEll, R. M., 1934. new and unrecorded foraminifera from the California Miocene. Contributions from the Cushman Laboratory for Foraminiferal Research, 10: 1–23.

CuShMAn, J. A. and lAiMinG, B., 1931. Miocene foraminifera from los Sauces Creek, Ventura County, California. Journal of Pa-leontology, 5: 79-120.

CuShMAn, J. A. and McCullOCh, i., 1939. A report on some aren-aceous foraminifera. los Angeles: university of Southern Califor-nia Press. Allan Hancock Pacific Expeditions, 6(1): 1–113 pp.

——— , 1948. The species of Bulimina and related genera in the collec-tions of the Allan Hancock Foundation. los Angeles: university of Southern California Press. Allan Hancock Pacific Expeditions, 6(5), 231–294.

CuShMAn, J. A. and McGlAMERy, W., 1939. new species of fora-minifera from the lower Oligocene of Alabama. Contributions from the Cushman Laboratory for Foraminiferal Research, 15: 45–49.

CuShMAn, J. A. and MOyER, D. A., 1930. Some Recent foramin-ifera from off San pedro, California. Contributions from the Cush-man Laboratory for Foraminiferal Research, 6: 49–62.

CuShMAn, J. A. and OzAWA, y., 1928. An outline of a revision of the polymorphinidae. Contributions from the Cushman Laboratory for Foraminiferal Research, 4: 13–21.

——— , 1929. Some species of fossil and Recent polymorphinidae found in Japan. Transactions of the Japanese Journal of Geology and Ge-ography, 6: 79–83.

——— , 1930. A monograph of the foraminiferal family polymorphin-idae, Recent and fossil. Proceedings of the United States National Museum, 77: 1–195.

CuShMAn, J. A. and pARKER, F. l., 1931. Miocene foraminifera from the Temblor of the east side of the San Joaquin Valley, Califor-nia. Contributions from the Cushman Laboratory for Foraminiferal Research, 7: 1–16.

——— , 1936. Some species of Robertina. Contributions from the Cush-man Laboratory for Foraminiferal Research, 12: 92–100.

——— , 1937. notes on some Oligocene species of Bulimina and Bulimi-inella. Contributions from the Cushman Laboratory for Foramin-iferal Research, 12: 36–40.

——— , 1938, notes on some pliocene and pleistocene species of Bulim-ina and Bulimiinella. Contributions from the Cushman Laboratory for Foraminiferal Research, 14: 53–71.

CuShMAn, J. A. and pOnTOn, G. M., 1931. A new Virgulina from the Miocene of Florida. Contributions from the Cushman Labora-tory for Foraminiferal Research, 17: 1–27.

CuShMAN, J. A. and RENz, h. h., 1941. New oligocene-Miocene foraminifera from Venezuela. Contributions from the Cushman Laboratory for Foraminiferal Research, 7: 32–33.

CuShMAN, J. A. and StAiNFoRth, R. M., 1945. the Foraminifera of the Cipero Marl Formation of trinidad, British West indies. Con-tributions from the Cushman Laboratory for Foraminiferal Re-search, 14: 3–75.

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——— , 1930. Post-Miocene foraminifera from the Ventura Quadrangle, Ventura County, California. Journal of Paleontology, 4: 60–72.

CuShMAN, J. A. and toDD, R., 1942. the Recent and fossil species of Laticarinina: Contributions from the Cushman Laboratory for Foraminiferal Research, 18: 14–20.

——— , 1944. The genus Spiroloculina and its species. Sharon, Massachusetts: Cushman Laboratory for Foraminiferal Research. Special Publication 11, 82 pp.

——— , 1945. Miocene foraminifera from Buff Bay, Jamaica. Sharon, Massachusetts: Cushman Laboratory for Foraminiferal Research. Special Publication 15, 73 pp.

——— , 1949. the genus Sphaeroidina and its species. Contributions from the Cushman Laboratory for Foraminiferal Research, 25: 11–21.

CuShMAN, J. A. and VALENtiNE, W. W., 1930. Shallow-water fora-minifera from the Channel Islands of southern California. Stanford: Stanford University Press. Contributions from the Stanford University Department of Geology, 1: 5–51.

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