Cushman Foundation Special Publication No. 46 p. 481-494, 2018

Chapter 18

TAXONOMY, BIOSTRATIGRAPHY, AND PHYLOGENY OF OLIGOCENE CASSIGERINELLA

Paul N. Pearson1, Silvia Spezzaferri2, Brian T. Huber3, and Michal Kučera4

1School of Earth and Ocean Sciences, Cardiff University, Main Building, Park Place, Cardiff, CF10 3AT, U.K. Email: [email protected]

2Department of Geosciences, University of Fribourg, Ch du Musée 6, 1700 Fribourg, Switzerland. Email: [email protected]

3Department of Paleobiology, MRC 121, Smithsonian Museum of Natural History, Washington, D.C. 20013-7012, U.S.A. Email: [email protected]

4MARUM - Center for Marine Environmental Sciences, University of Bremen, Bremen, Germany. Email: [email protected]

ABSTRACT The taxonomy, biostratigraphy, and phylogeny of test microstructure, with minute-scale internal Oligocene Cassigerinella is reviewed. Cassigerinella ornamentation (‘endospikes’). Only two species is in Superfamily Guembelitrioidea and Family occur in the Oligocene and are recognized as valid Cassigerinellidae. It is distinguished by its ‘twisted in this work: Cassigerinella chipolensis (Cushman enrolled-biserial’ coiling mode and distinctive and Ponton) and Cassigerinella eocaenica Cordey.

INTRODUCTION common species, Cassigerinella chipolensis, was a surface dwelling form (Boersma and Shackleton, 1978; Cassigerinella is a distinctive genus with an Pearson and Wade, 2009) and it is likely that the other unusual coiling mode and wall texture. It is frequently members of the genus had a similar habitat. encountered in sediments of late Eocene to middle There has been some debate in the literature on Miocene age, within which interval subtle changes the style of chamber addition through ontogeny, which in size and shape are of considerable potential in on first inspection can appear haphazard. Pokorný regional biostratigraphic correlation (Premoli Silva and (1955:138) noted that later-formed chambers ‘alternate’, Spezzaferri, 1990; Spezzaferri, 1994). Its planktonic that is the chambers have reflective symmetry and habit is indicated by its distribution in open ocean successive apertures / foramina face each other rather marine sediments (Hughes and Jenkins, 1981) and than all facing the same way (hence apertures do not general test morphology, including the rather globular define an anterior direction and there is no dorsal or chambers (Li, 1986; Huber and others, 2006). Its ventral side). Pokorný suggested that the earliest whorl occurrence can be quite patchy both stratigraphically could be planispiral or trochospiral although it was and geographically (e.g., Jenkins, 1971) and it is likely difficult for him to be sure. Saito and Biscaye (1977) that it was an opportunistic genus, blooming in the described this distinctive coiling type as ‘enrolled- surface mixed-layer when trophic conditions were biserial’, indicating that the test was fundamentally suitable. Stable isotope evidence indicates that the most biserial but coiled back on itself. Li (1986:52, his fig. Pearson, Spezzaferri, Huber, and Kučera

1) elaborated on this by pointing out that the plane channels (Pearson and Wade, 2009). A distinctive of enrollment itself twists through ontogeny, most feature is the presence of small micron-sized spikes markedly in forms with inflated globular chambers scattered on the interior surface, called ‘endospikes’ (e.g., typical Cassigerinella chipolensis). Presumably by Pearson and Wade (2009). This is apparently a this additional twisting allows successive apertures to unique feature in foraminifera. The presence of these be positioned clear of the earlier chambers and for the endospikes on the chamber interiors implies an unusual test to maintain a globular shape overall. We agree with calcification mechanism. D’ Haenens and others (2012) Li (1986) that this ‘twisted enrolled-biserial’ coiling observed similar endospikes in a dissected specimen seems to be the pattern of chamber addition from the of Chiloguembelina wilcoxensis. That observation earliest ontogenetic stages onward and we have not helps cement a link between Chiloguembelina and observed an initially planispiral or trochospiral phase Cassigerinella. Presumably the latter evolved by in any of the specimens examined, despite the earlier developing a secondarily enrolled coiling mode, which suggestions of various authors including Pokorný maintains the fundamental biserial arrangement. Other (1955), Cordey (1968), and Blow (1979:1362). species of Chiloguembelina do not show endospikes Hofker (1963) made detailed studies of the (see Chapter 15, this volume), so presumably the apertural system in Cassigerinella and, working near ancestor was C. wilcoxensis itself, although a large the limit of resolution of the light microscope, claimed stratigraphic gap exists between the highest known to observe an internal “toothplate”; that is, a flange occurrence of that species and the lowest Cassigerinella extending inward from the aperture and either joining (Huber and others, 2006). Stratigraphic ranges and with, or approaching the foramen of the preceding inferred phylogenetic relationships are illustrated in chamber (See Hofker, 1963, pl. 1, figs. 2b, 2e, 2g, and Figure 18.1. 3a). Steineck and Darrell (1971) also recorded this The systematic taxonomy of Oligocene feature in C. chipolensis. If so, this observation would Cassigerinella is described in the following section. provide good evidence of a phylogenetic linkage with We note for completeness that an alleged species the benthic genus Cassidulina which has this character of Cassigerinella from the Oligocene of Cuba as well as enrolled-biserial coiling (Banner, 1982; for (Cassigerinella regularis Iturralde-Vinent, 1966) is not discussion of the toothplate in benthic cassidulinids see referable to the genus and is very probably a benthic Nomura, 1983, and Li, 1986). However, SEM studies species (Li, 1986). by Fleisher (1974), Saito and Biscaye (1977), Blow (1979), Li (1986), Spezzaferri (1994), and Pearson SYSTEMATIC TAXONOMY and Wade (2009) among others have failed to reveal any evidence of toothplates. Instead, apertures and Superfamily GUEMBELITRIOIDEA, foramina commonly possess an irregular lip with a Montanaro Gallitelli, 1957 ragged outward-directed flange (see, for example, Plate 18.1, Figs. 4-6, 8, and 12). DISCUSSION.— The discovery by D’Haenens and The external wall texture is essentially smooth others (2012) of endospikes in Chiloguembelina, but may be pustulose to various degrees or bearing pore virtually identical to those described in Cassigerinella mounds. In general, the more pustulose forms tend to be by Pearson and Wade (2009), is good evidence that from the Oligocene, whereas Miocene forms are usually the two genera belong to the same phylogenetic group, smoother. There is, however, substantial variability and that the wall texture of Cassigerinella is a subtype even on individual tests and we do not use the degree of the ototara-type wall. The ototara-type wall helps of smoothness to separate taxa (see discussion under define a distinct higher taxonomic group. For that Cassigerinella chipolensis, below). The wall texture in reason we assign the Cassigerinellidae to Superfamily cross-section was described in detail by Pearson and Guembelitrioidea. Wade (2009), and is named the chipolensis-subtype wall (see Chapter 15, this volume). Although some Family CASSIGERINELLIDAE Bolli, Loeblich, earlier workers (e.g., Hofker, 1963) thought the wall had and Tappan, 1957, revised Li, 1986 a radial texture, it is in fact microgranular (as originally observed by Blow, 1979:1361) with expanding pore Type genus: Cassigerinella Pokorný, 1955.

482 Chapter 18 - Cassigerinella

Cassigerinella in detail and contrasted it with the very

Cassigerinella different wall seen in the Globigerinitidae. Accordingly ( S u b ) ( S u b ) t r o p i c a l t r o p i c a l A n t a r c i they resurrected Family Cassigerinellidae, which we follow here. Pearson (Chapter 15, this volume) n e s

A g e ( M a ) named the distinctive wall texture of this Family Z o n e s

E p o c h as the chipolensis-subtype wall, which has so far i c a G P T S ( 2 0 5 ) been observed in Cassigerinella and one species of (WPBP, 2011) (WPBP, ( B K S A , 1 9 5 ) & C a n d e & K t ( 1 9 5 ) F o r m e P E , O a n d M Z o Chiloguembelina (D’ Haenens and others, 2012). The H u b e r & Q i l é v N Z o n e s ( K & S , 1 9 8 3 ) c h i p o l e n s e o c a n i e o c a n c h i p o l e n s Family is regarded as either monogeneric or possibly 22 Y also including the Miocene genus Riveroinella (see b b discussion under C. eocaena) and, contra Stolk (1965) 23 M 1 E A R L N 4 M I O C E N a and BouDagher-Fadel (2012:166), it does not include a 24 the middle Eocene genus Cassigerinelloita which has a different coiling pattern and wall texture (Li and O7 25 Radford, 1992; Huber and others, 2006:470).

P22 A O 4 26 Genus Cassigerinella Pokorný, 1955, T E O6

L A emended Li, 1986 27

b O5 A O 3 TYPE SPECIES.— Cassigerinella boudecensis 28 1 2

P Pokorný, 1955 (= junior subjective synonym of 29 a O4 Cassigerinella chipolensis Cushman and Ponton, O L I G C E N A O 2 1932). 30 P20 O3

Y DISCUSSION.— Pokorný (1955) and Hofker 31 P19 O2 (1963) suggested that the genus was characterized by E A R L 32 apertural toothplates but this was disproved by the SEM investigations of Saito and Biscaye (1977) and P18 A O 1 33 O1 Li (1986) among others. The genus was also discussed P17 by Huber and others (2006) in the Atlas of Eocene AE10 34 E16 Planktonic Foraminifera. The superficially similar TE Eocene genus Cassigerinelloita is unrelated as it has

L A P16 35 E15 AE9 E O C N a wall texture more similar to Guembelitria (Li and Radford, 1991; Huber and others, 2006). FIGURE 18.1. Stratigraphic ranges and inferred phylogenet- ic relationships of Oligocene Cassigerinella. BKSA, 1995 Cassigerinella chipolensis (Cushman and Ponton, = Berggren and others, 1995; K&S, 1983 = Kennett and 1932) Srinivasan, 1983; WPBP, 2011 = Wade and others (2011). Plate 18.1, Figures 1-16. DISCUSSION.— In their original concept of Family (Note: in this synonymy list we focus on Oligocene Cassigerinellidae, Bolli and others (1957) included occurrences and taxonomically important works) several genera that are now regarded as essentially unrelated to one another. Li (1986) restricted the Family ?Cassidulina globulosa Egger, 1857:296, pl. 7, figs. 4-7 [lower Miocene, Habühl, Bavaria, Germany]. to include just the type genus. In the Atlas of Eocene Cassigerinella globulosa Egger, 1857.—Rögl, 1985:319, Planktonic Foraminifera, Huber and others (2006:449) figs. 5.18a-c (re-illustration of holotype), 5.19 [lower moved Cassigerinella to the Globigerinitidae, implying Miocene Ottnangian, Mairhof, Bavaria, Germany], 5.20 a close relationship with Tenuitella, and in so doing [lower Miocene Ottnangian, Blättermergel, Bavaria, suppressed the Cassigerinellidae. Pearson and Wade Germany]. (2009:198) described the distinctive wall texture of Cassidulina chipolensis Cushman and Ponton, 1932:98, 483 Pearson, Spezzaferri, Huber, and Kučera

Plate 18.1 Cassigerinella chipolensis (Cushman and Ponton, 1932) 484 Chapter 18 - Cassigerinella

pl. 15, figs. 2a-c [lower Miocene Chipola Fm., Alum 1996, pl. 1, fig. 9 [lower middle Miocene, Cadell Marl, Bluff Group, Calhoun County, Florida].—Cushman South Australia].—Huber and others, 2006:484-486, and Ellisor, 1945:570, pl. 78, fig. 1 [‘middle’ Oligocene, pl. 16.4, figs. 23, 24 (holotype re-illustrated by SEM), Anahuac Fm., Chambers County, Texas].—Cushman fig. 25 [upper Eocene Zone E15/16, ODP Hole 1053B, and Stainforth, 1945:64, pl. 12, fig. 5 [lower Oligocene, Blake Nose, North Atlantic Ocean], figs. 26, 27 [upper Cipero Fm., Trinidad]. Eocene Zone E14/15, Atlantic City borehole, New Cassigerinella chipolensis (Cushman and Ponton).— Jersey, ODP 150X, 1303.4-.5ft].—Miller and others, Bolli, 1957:108, pl. 22, figs. 3a-c [lower Oligocene 2008: fig. 6N [lower Oligocene Zone O1, St Stephens Globorotalia opima opima Zone, Cipero Fm., Quarry, Alabama].—Pearson and Wade, 2009:198-200, Trinidad].—Bermúdez, 1961:1222-1223, pl. 8, fig. pl. 1, figs. 1a-2f [upper Oligocene Zone O6 (=O7 in this 4a-c [‘middle’ Oligocene, Tinguaro Fm., Matanzas work), Cipero Fm., Trinidad], pl. 1, figs. 3a-3b [upper Province, Cuba].—Blow and Banner, 1962:81-82, pl. 15, Oligocene Zone O5, Juana Diaz Fm., Puerto Rico].— figs. M, N [lower OligoceneGlobigerina oligocaenica Pearson and Wade, 2015, fig. 30.7a-8 [lower Oligocene Zone, Lindi area, Tanzania].—Hofker, 1963, pl. 1, figs. Zone O1, TDP Site 12, Stakishari, Tanzania]. 1a-2f [lower Oligocene Globorotalia opima opima Cassigerinella boudecensis Pokorný, 1955:138, figs. zone, Cipero Fm., Trinidad].—Blow, 1969:377, pl. 51, 1-3 [‘middle’ Oligocene, Pausramer Marl, Czech fig. 5 [lower Oligocene Globigerina ampliapertura Republic].—Bolli and others, 1957:30-31, pl. 3, figs. Zone, Cipero Fm., Trinidad].—Jenkins, 1971:73, pl. 6a-c [‘middle’ Oligocene, Czech Republic].—Hofker, 1, fig. 30 [lower Miocene Awamoan series, Awamoa 1963, pl. 1, figs. 1a-2f [Oligocene ‘flysch’ east of Popice, Creek, New Zealand].Jenkins and Orr, 1972:1084, Czech Republic].—Li, 1986:61, pl. 1, figs. 1-18; pl. 2, pl. 1, figs. 5-8 [lower Oligocene Pseudohastigerina figs. 1-14; pl. 3, figs. 1-4 [lower Oligocene Zone P21, barbadoensis Zone, DSDP Site 77B, eastern equatorial Cipero Fm., Trinidad].—BouDagher-Fadel, 2012, pl. Pacific Ocean].Fleisher, 1974:1015, pl. 4, fig. 3 5.10, fig. 6 (reproduced without attribution from Li, [lower Oligocene Zone P18/19, DSDP Site 219, Arabian 1986, pl. 1, fig. 1, see above, wrongly recorded as Sea].Quilty, 1976:651, pl. 19, figs. 11, 12 [lower Oligocene Zone P19, Czech Republic), pl. 6.10, fig. 13 Oligocene Zone P19, DSDP Site 321, southeastern (reproduced without attribution from Li, 1986, pl. 1, fig. Pacific Ocean].—Saito and Biscaye, 1977, text-fig. 1, see above, wrongly recorded as Miocene Zone N4, 2a-c; pl. 2, figs. 3-7 [Oligocene, JOIDES Hole 3, Blake Czechoslovakia, despite being the same specimen as the Plateau, North Atlantic Ocean].—Blow, 1979:1362- misattributed occurrence listed above). 1363, pl. 51, fig. 5; pl. 247, figs. 5-8 [lower Oligocene, Cassigerinella globolocula Ivanova, 1958:57, pl. 11, figs. Zone P19-P20, Lindi, Tanzania].—Bolli and Saunders, 1a-3c [upper Oligocene, Kosmach series, Vorotyshche 1985:185, figs. 16.1-2 [re-illustration of holotype and River, Carpathian Mountains, USSR (modern Ukraine)]. paratype].—Li, 1986:6, pl. 3, figs. 7-11; pl. 4, figs. 1-17 Not Cassigerinella chipolensis (Cushman and Ponton).— [lower Oligocene Zone P21, Cipero Fm., Trinidad].— Huber and others, 2006:484-486, pl. 16.4, fig. 25 [upper Loeblich and Tappan, 1988:488, pl. 533, figs. 11-13 Eocene Zone E15/16, ODP Hole 1053B, Blake Nose, (re-illustration from Bolli and others, 1957), 14 (re- North Atlantic Ocean], figs. 26, 27 [upper Eocene Zone illustration from Saito and Biscaye, 1977).—Premoli E14/15, Atlantic City borehole, New Jersey, ODP 150X, Silva and Spezzaferri, 1990:444, pl. 1, figs. 3a-c [upper 1303.4-.5ft] (= C. eocaenica). Oligocene Zone P22, ODP Hole 709B, Madingley Rise, equatorial Indian Ocean].—Spezzaferri and Premoli DESCRIPTION. Silva, 1991:236, pl. 8, figs. 6a-c [upper Oligocene Type of wall: Microperforate to medio- Subzone P21b, DSDP Hole 538A, Gulf of Mexico].— perforate with pore diameters ranging from 0.5-2.0 Leckie and others, 1993:123, pl. 7, figs. 18, 19 [lower µm; surface may be smooth or covered in pore Oligocene, ODP Hole 628A, Little Bahama Bank, western North Atlantic Ocean].—Spezzaferri, 1994:64, mounds; microgranular internal structure with minute pl. 29, figs. 5a-c [upper Oligocene Zone P22, ODP Hole endospikes distributed on the interior chamber surfaces 709B, equatorial Indian Ocean].—Li and McGowran, (chipolensis-type wall; see Chapter 15, this volume).

Plate 18.1 Cassigerinella chipolensis (Cushman and Ponton, 1932)

1 (re-illustration of type specimen of Cassidulina globulosa Egger, 1857 [nomen dubium non conservandum]; no scale available); 2, 3 (holotype of Cassidulina chipolensis Cushman and Ponton, 1932, reproduced from Huber and others, 2006), lower Miocene Chipola Fm., Florida; 4, Oligocene, Atlantic Slope Project, New Jersey, Sample 5B/20H, 18"; 5-8, Zone O1, Sample NKK1-33B, 45-105 cm, Kali Kunir, Nanggulan Fm., Java; 9-11, Zone O7, Sample PP07/T5, Cipero Fm., Trinidad (9, Pearson and Wade, 2009, pl. 1, fig. 1a);12 , Oligocene, Istra More-3 well, 968-974 m, Adriatic Sea; 13-16, Oligocene, Pausramer Marl, Czech Republic (topotypes of Cassigerinella boudecensis Pokorný). Scale bar: 2-16 = 50 µm. 485 Pearson, Spezzaferri, Huber, and Kučera

Test morphology: Test small, globular with on plate 7, not plate 11 of his work as has generally irregular outline. From the earliest stages chambers been reported. are in an enrolled-biserial arrangement with successive Pokorný (1955) distinguished his species chambers facing one another; the axis of enrollment boudecensis (the type species of Cassigerinella) from commonly changes through ontogeny; 6-8 chambers chipolensis on the basis of its supposed pustulose per enrollment, relatively compressed and compact wall texture. Various authors have maintained this initially but becoming strongly inflated or ovate in distinction (e.g., Rögl, 1985; Li, 1986) but others last whorl. Sutures straight and depressed to incised. have combined the two taxa. Pokorný himself, in Aperture interiomarginal, asymmetrical, highly arched a personal communication reported by Blow and to virguline in shape, surrounded by an irregular lip, Banner (1962:83), said that boudecensis was “probably commonly with laterally asymmetrical flanges. conspecific with chipolensis”; others who have Size: Small, mostly <200 µm. combined the taxa are Blow and Banner (1962), Cordey (1968), Jenkins (1971), Raju (1971), Saito and Biscaye DISTINGUISHING FEATURES.— This species is (1977), Blow (1979), Li and McGowran (1996), Huber distinguished from Cassigerinella eocaenica Cordey and others (2006), and Pearson and Wade (2009). by having a larger, more globular test, more inflated Note that while Li (1986) upheld the distinction, in chambers, faster chamber size increase and higher a later work (Li and McGowran, 1996) he subsumed arched aperture. the two species into one. Huber and others (2006:485) pointed out that the distinction is hard to maintain with DISCUSSION.— This species has been widely the light microscope and has never been proven to be recognized in the literature under the name of stratigraphic value. Pearson and Wade (2009:198) Cassigerinella chipolensis (Cushman and Ponton). argued that the degree of pustule development was a However it was probably first described as long ago variable feature on individual tests, among populations, as 1857 as Cassidulina globulosa Egger. Egger (1857) and between localities, making it difficult to sustain illustrated a specimen in three views and also provided the distinction. To further investigate this issue we an interpretative sketch of the irregular chamber have obtained a population of ~40 topotype specimens arrangement. Although he did not provide a scale, he from the type locality of boudecensis and examined a noted that test size ranges up to 250 µm. As pointed number of them in SEM. We find the degree of pustule out by Pokorný (1955:138) the specimen illustrated by development to be quite variable even in this material, Egger is undoubtedly referable to Cassigerinella (Plate with some nearly smooth forms and others distinctly 18.1, Fig. 1). Rögl (1985:319) suggested that Egger’s pustulose, hence we maintain the species in synonymy taxon is specifically distinct from chipolensis, being (compare Plate 18.1, Figs. 13-16; see also Chapter 15, distinguished by its more compressed, ovate chambers, this volume, Plate 15.3, Figs. 4a-5b). and in making this distinction he was followed Premoli Ivanova (1958:57) described a new species Silva and Spezzaferri (1990) and Spezzaferri (1994) Cassigerinella globolocula but did not provide criteria (see discussion below under Cassigerinella eocaenica). for distinguishing it from other species. Pokorný (in However, the evidence for this morphological a personal communication reported by Blow and distinction is unclear either in Egger’s type illustrations Banner, 1962) indicated that globolocula was “certainly or on the small additional illustrations of specimens conspecific” with his species boudecensis and hence, from Bavaria provided by Rögl (1985). In the tax- in this work, also with chipolensis. Blow (1969:212) onomy presented here, and following Huber and suggested that the paratypes of globolocula were others (2006), the more compressed morphotypes of referable to Cassigerinella eocaena (sic, = eocaenica) Cassigerinella are assigned instead to C. eocaenica. of Cordey but the type material of this species has We regard globulosa as a probable prior synonym of apparently been lost and we cannot confirm this chipolensis, but we note that Egger’s specimen is lost observation. Ivanova’s illustrations appear to us to and nomenclatorial stability is best served by regarding fall within the normal range of variation observed in it as nomen dubium non conservandum. We note also chipolensis. that Egger’s identification has often been erroneously As noted above, the morphospecies C. recorded as being from 1957, and the illustrations are chipolensis is distinguished from the other valid species

486 Chapter 18 - Cassigerinella in the genus, C. eocaenica, by being more inflated – Miocene boundary (Scott, 2001). According to and globular overall, and slightly larger. The two Cushman and Ponton (1932) their sample was from morphospecies intergrade throughout their combined the lower Miocene. range, and in the absence of biometric data, it is not at all obvious that these features are consistent through GEOGRAPHIC DISTRIBUTION.— This species is time and diagnose distinct biospecies. We maintain most abundant in the lower latitudes (Premoli Silva the taxonomic distinction because it seems to have and Boersma, 1988; Spezzaferri, 1995). It is absent biostratigraphic utility insofar as the C. chipolensis from high latitude sites (e.g. the Southern Ocean and morphospecies is seemingly absent in the Eocene (see ODP Site 647 in the Labrador Sea) although it occurs discussion below). sporadically as far south as the latitude of New Zealand (Jenkins, 1971; Jenkins and Srinivasan, 1986). PHYLOGENETIC RELATIONSHIPS.— Probably a direct descendant of Cassigerinella eocaenica and STABLE ISOTOPIC PALEOBIOLOGY.— part of the same evolving lineage (Cordey, 1968; Blow, Cassigerinella chipolensis has stable isotope ratios 1979). indicative of a warm surface water habitat (Boersma and Shackleton, 1978; Pearson and Wade, 2009). It STRATIGRAPHIC RANGE.— Most confirmed probably bloomed opportunistically in response to occurrences of chipolensis are in the Oligocene and seasonal productivity peaks (Pearson and Wade, 2009). Miocene: Eocene occurrences of Cassigerinella have generally been referred to C. eocaenica. This REPOSITORY.— Holotype (USNM 16326) deposited apparently neat distinction may be more an artifact of at the Smithsonian Museum of Natural History, fine taxonomic discrimination and the way in which Washington, D.C. the biozonation developed historically than it is for necessarily providing a clear and useful biohorizon. Cassigerinella eocaenica Cordey, 1968 The overlap in the ranges of Cassigerinella chipolensis and Pseudohastigerina spp. was for a time part of the Plate 18.2, Figures 1-12 standard zonation (Blow and Banner, 1962; Bolli, 1966; Bolli and Saunders, 1985; see discussion in Cassigerinella eocaenica Cordey, 1968:369, figs. 1a-e Chapter 2, this volume). Hence when Cassigerinella [Eocene, core hole drilled off northern Florida on specimens were discovered in the Eocene it was Blake Nose, western North Atlantic Ocean].—Bolli found convenient to refer them to a different species and Saunders, 1985:185, fig. 16.5-6 (re-illustration of based on relatively minor features. Sexton and others holotype).Huber and others, 2006:486-487, pl. 16.4, figs. 18, 19 [paratype, USNM 643514, upper Eocene, (2006) and Huber and others (2006) have recorded Blake Plateau, western North Atlantic Ocean], figs. 20, occasional “C. chipolensis” from the middle and upper 21 [reillustration of pl. 1, figs. 1, 2 of Li and McGowran, Eocene but in this work we assign those occurrences 1996, nannofossil Zone NP23, South Australia], fig. 22 to C. eocaenica. No detailed morphometric study of [upper Eocene Zone E14/E15, Atlantic City borehole, the early evolution of Cassigerinella has yet been New Jersey, ODP Hole150X]. conducted. The extinction of chipolensis has been Cassigerinella eocaena (sic) Cordey, 1968.—Blow, 1969, recorded diachronously at various levels in the middle pl. 51, fig. 6 [upper Eocene Zone P16, Core Lamont Miocene (e.g., Bolli, 1957; Martinotti, 1989; Chaisson A. 167-21, southeast of Blake Plateau, North Atlantic and Leckie, 1993; Spezzaferri, 1994; Turco and others, Ocean], figs. 7, 8 [upper Eocene Zone P16, Core 2002) but has not been investigated by us in detail. JOIDES Mo. 6, Blake Plateau, North Atlantic Ocean]. Cassigerinella sp.Quilty, 1976:651, pl. 19, figs. 13, Intervals of relatively large-sized individuals may be 14 [lower Oligocene Zone P19, DSDP Site 321, of biostratigraphical utility for fine-scale correlation southeastern Pacific Ocean].Leckie and others, (Premoli Silva and Spezzaferri, 1990; Spezzaferri, 1993:123, pl. 7, fig. 17 [lower Oligocene Zone P18, 1994; Pearson and Chaisson, 1997). ODP Hole 628A, Little Bahama Bank, Atlantic Ocean]. Cassigerinella winniana Howe.Blow, 1979:1363-1364, TYPE LEVEL.— The Chipola Formation of the Alum pl. 51, figs. 6-8, pl. 246, figs. 8-10 [upper Eocene Bluff Group is currently thought to span the Oligocene Zone P16, Core Lamont A. 167-21, southeast of Blake 487 Pearson, Spezzaferri, Huber, and Kučera

Plate 18.2 Cassigerinella eocaenica Cordey, 1968 and Riveroinella martinezpicoi Bermúdez and Seiglie, 1967 488 Chapter 18 - Cassigerinella

Plateau, North Atlantic Ocean; metatypes from upper from the earliest stages formed in an attenuated Eocene, JOIDES hole Mo. 6, Blake Plateau, North enrolled-biserial arrangement with successive Atlantic Ocean; lower Oligocene Zone P18, DSDP Site chambers facing one another; axis of enrollment 14, central South Atlantic Ocean].Li and McGowran, fairly constant through ontogeny; chambers 6-8 per 1996:97-103, pl. 1, figs. 1-12 [lower Oligocene, Port enrollment, compressed and sometimes reniform. Willunga Fm., South Australia].Li and others, 2003:16, pl. 2, fig. 14 [upper Eocene Zone P15/P16, Sutures straight to moderately curved, depressed. ODP Site 1134A, Great Australian Bight].Li and Aperture interiomarginal, asymmetrical, highly arched others, 2005:16, pl. 1, fig. 3 [lower Oligocene Subzone to virguline in shape, surrounded by an irregular lip P21a, ODP Hole 1148A, South China Sea; designation with laterally asymmetrical flanges. according to the figure caption; note the citation on p. 18 Size: Small, mostly <150 µm. assigns this specimen to C. chipolensis].—BouDagher- Fadel, 2012, pl. 5.10, fig. 5 (reproduced without DISTINGUISHING FEATURES.— This species attribution from Blow, 1979, pl. 246: fig. 9) [see above]; is distinguished from Cassigerinella chipolensis [wrongly recorded as Oligocene Zone P19, Louisiana]. (Cushman and Ponton) by its more equatorially [Not Howe, 1939.] compressed test. This is partly because the chambers are Cassigerinella globulosa (Egger).Premoli Silva and Spezzaferri, 1990:300, pl. 1, figs. 1a-c [upper less inflated and partly because the axis of enrollment Oligocene Zone P22, ODP Hole 707A, equatorial Indian does not tilt markedly during ontogeny as it does in Ocean].Spezzaferri, 1994:64, pl. 29, figs. 3a-c [lower chipolensis (see discussion above) so the enrollment Miocene Zone N6, ODP Hole 709C, equatorial Indian tends to follow approximately the same plane and Ocean]. [Not Egger, 1857.] can appear pseudo-planispiral. The periphery tends to Cassigerinella martinezpicoi (Bermúdez and Seiglie). be subrounded to subacute, compared with the more Premoli Silva and Spezzaferri, 1990:444, pl. 1, figs. rounded profile in C. chipolensis. It is distinguished 2a-c [lower Oligocene Zone P19 (=O1 in this study), from Riveroinella martinezpicoi by lacking the ODP Hole 706A, Mascarene Plateau, equatorial Indian very flattened test, pinched and acute periphery, and  Ocean]. Spezzaferri, 1994:64, pl. 29, figs. 4a-c (re- protruding, continuous flange-like lip around the illustrated from Premoli Silva and Spezzaferri, 1990). [Not Bermúdez and Seiglie, 1967.] aperture. Cassigerinella chipolensis (Cushman and Ponton).—Sexton and others, 2006, pl. 1, figs. 16, 17 [upper Eocene DISCUSSION.— The genus Cassigerinella was Zone E14, ODP Sample 1052B 10H-5, 43-46 cm. (P. originally thought to be restricted to the Oligocene Sexton, personal communication, 2012), Blake Nose, and early Miocene and the first appearance of C. North Atlantic Ocean].—Huber and others, 2006 chipolensis was proposed as a marker for the basal (partim):484-486, pl. 16.4, fig. 25 [upper Eocene Zone Oligocene (Blow and Banner, 1962). However Saito E15/16, ODP Hole 1053B, Blake Nose, North Atlantic and Bé (1963) observed specimens of Cassigerinella Ocean], figs. 26, 27 [upper Eocene Zone E14/15, in the Eocene. This was confirmed by Cordey (1968), Atlantic City borehole, New Jersey, ODP 150X, 1303.4- but he distinguished them at the species level as C. .5ft]. [Not Cushman and Ponton, 1932.] eocaenica based on the slightly smaller size and DESCRIPTION. less inflated chambers, flatter enrollment (which he Type of wall: Microperforate with pore described, erroneously in our view, as planispiral in the diameters ranging from 0.5-1.0 µm; surface smooth early ontogenetic stages) and less rounded periphery. (chipolensis-subtype wall; see Chapter 15, this volume; Subsequently it became typical for workers to refer note that the internal structure has yet to be confirmed Oligocene specimens to chipolensis and Eocene for this species). specimens to eocaenica (see discussion above under Test morphology: Test small, flattened, petaloid chipolensis). However, as discussed by Huber and others in outline, periphery rounded to subacute. Chambers (2006:486-487), compressed forms of Cassigerinella

Plate 18.2 Cassigerinella eocaenica Cordey, 1968 and Riveroinella martinezpicoi Bermúdez and Seiglie, 1967

Cassigerinella eocaenica 1-3, holotype (1,2) and paratype (3) from Cordey, 1968, text-fig. 1a-b, 1e;4-12 , lower Oligocene Zone O1, ODP Hole 706A/3H/4, 93-95 cm, Mascarene Plateau, equatorial Indian Ocean; 13-15, new SEMs of the holotype of Riveroinella martinezpicoi Bermúdez and Seiglie, 1967. Scale bars: 1-15 = 50 µm. 489 Pearson, Spezzaferri, Huber, and Kučera that are similar to eocaenica also occur in the illustrated by Saito and Biscaye (1977) from a similar Oligocene. In some cases these have been documented Miocene level in JOIDES Hole 3, Blake Plateau, in the under the name Cassigerinella winniana but Huber and western North Atlantic Ocean. From this we conclude others (2006) showed that the holotype of winniana is that martinezpicoi is a distinct form that is so far known a benthic form referable to Cassidulina. Hence Huber only from Subzone M5b of the subtropical North and others (2006) extended the range of eocaenica up Atlantic Ocean and Caribbean. We have searched for into the Oligocene. According to our observations, specimens of this species in several samples lent to us compressed morphotypes of Cassigerinella referable by W.A. Berggren from very close to the level of Saito to this species are in fact quite common throughout the and Biscaye’s specimens in JOIDES Hole 3. Although Oligocene and on into the Miocene. Cassigerinella is quite common in this material (both Premoli Silva and Spezzaferri (1990) and C. chipolensis and C. eocaenica being present), no Spezzaferri (1994) recognized Cassigerinella globulosa specimens comparable to martinezpicoi were found. Egger as valid and distinct from chipolensis. According Hence it seems that the latter must have a patchy to Spezzaferri (1994) the distinguishing features of occurrence in that core. globulosa are its more flattened test, reniform chambers Bermúdez and Seiglie (1967) used martinezpicoi and rounded to slightly subacute peripheral margin. to typify their genus Riveroinella. Saito and Biscaye These are all characteristic of eocaenica according to (1977) regarded it as a compressed Cassigerinella, and Huber and others (2006) and the taxonomy presented so subsumed Riveroinella in Cassigerinella. Although here. As discussed above under chipolensis, Egger’s the forms appear to be related, Loeblich and Tappan type material is lost and his illustrated specimen appears (1988) upheld the generic distinction on the basis of more similar to chipolensis than eocaenica. the peculiar morphological features, including apertural Premoli Silva and Spezzaferri (1990) and features, in martinezpicoi. We follow Loeblich and Spezzaferri (1994) also recognized the species Tappan (1988) and suggest that Riveroinella was Cassigerinella martinezpicoi (Bermúdez and Seiglie). descended from Cassigerinella some time in the lower According to Spezzaferri (1994:64) this differs from Miocene. The peculiar compressed morphology, patchy other Cassigerinella by its “strongly reniform to occurrence and restricted geographic distribution (as subtriangular chambers, more curved sutures, an almost so far determined; see Li, 1986:62) could indicate that subacute peripheral margin and a very flattened test Riveroinella had a secondarily benthic habit, although (more flattened than C. globulosa)”. The specimens that has yet to be tested using stable isotopes. illustrated by Premoli Silva and Spezzaferri (1990) and Spezzaferri (1994) from tropical Indian Ocean PHYLOGENETIC RELATIONSHIPS.— The origin sites show these features very clearly and here we of this species, the first of its Superfamily, is unknown. illustrate three more specimens from ODP Hole 706A The most likely possibility is that it evolved from a (Mascarene Plateau, Indian Ocean) (Plate 18.2, Figs. benthic foraminifer in the genus Cassidulina, with 4-12). Nevertheless these specimens do not approach which it shares the enrolled-biserial coiling mode. The the extremely flat and peripherally acute morphology species gave rise to Cassigerinella chipolensis in the of the type series of Riveroinella martinezpicoi. The earliest Oligocene and may also have been ancestral to holotype of the latter (which is from a relatively Riveroinella martinezpicoi in the early Miocene. high stratigraphic level in what is now termed the Praeorbulina glomerosa Subzone, M5b, in upper part STRATIGRAPHIC RANGE.— The lowest reported of the lower Miocene) is illustrated here for the first occurrences are in upper Eocene Zone E14 (Sexton time in SEM for comparison (Plate 18.2, Figs. 13-15). and others, 2006). It ranges into the Miocene. We have We note that this specimen and the five paratypes observed it as high as Subzone M5b in the Atlantic originally illustrated by Bermúdez and Seiglie (1967) Ocean but have not investigated its extinction level in all possess a peculiar slit-like aperture surrounded detail as part of this project. by a protruding, flange-like lip and strongly acute profile which are features not seen in the other species TYPE LEVEL.— Upper Eocene, together with under consideration. Specimens of nearly identical Hantkenina alabamensis, Hantkenina primitiva, and morphology to the type series were described and Turborotalia cerroazulensis (probably Zone E16).

490 Chapter 18 - Cassigerinella

GEOGRAPHIC DISTRIBUTION.— Global, except Tappan, H., Beckmann, J.P., Bolli, H.M., Montanaro Gallitelli, in high latitudes. E., and Troelsen, J.C. (eds.): Studies in Foraminifera, U.S. National Museum Bulletin, U.S. Government Printing Office, Washington, D.C., v. 215, p. 3-50. STABLE ISOTOPE PALEOBIOLOGY.— No data , and Saunders, J.B., 1985, Oligocene to Holocene low available. latitude planktonic foraminifera, in Bolli, H.M., Saunders, J.B., and Perch-Nielsen, K., (eds.): Plankton Stratigraphy: REPOSITORY.— Holotype (BMNH P46838) and Cambridge University Press, Cambridge, p. 155-262. BouDagher-Fadel, M.K., 2012, Biostratigraphic and Geological paratype (BMNH P46839) deposited at the Natural Significance of Planktonic Foraminifera: Developments in History Museum, London. Palaeontology and Stratigraphy, v. 22, 400 p. 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(eds.): Proceedings of the Ocean Rögl, F., 1985, Late Oligocene and Miocene planktonic foraminifera Drilling Program, Scientific Results: Ocean Drilling Program, of the Central Paratethys, in Bolli, H.M. and Saunders, J.B. College Station, TX, v. 182, p. 1-28. (eds.): Plankton Stratigraphy: Cambridge University Press, , Jian, Z., and Su, X., 2005, Late Oligocene rapid Cambridge, p. 315-328. transformations in the South China Sea: Marine Saito, T., and Bé, A.W.H., 1963, Planktonic foraminifera from the Micropaleontology, v. 54, p. 5-25. American Oligocene: Science, v.145, p. 703-704. Loeblich, A.R., Jr., and Tappan, H., 1988, Foraminiferal Genera , and Biscaye, P.E., 1977, Emendation of Riveroinella and their Classification: Van Nostrand Reinhold Co., New martinezpicoi Bermúdez and Seiglie, 1967, and synonymy York, p. 1059. of Riveroinella with Cassigerinella Pokorný 1955: Martinotti, G.M., 1989, The last occurrence of Cassigerinella Micropaleontology, v. 23, p. 319-329. chipolensis in the Mediterranean region: Journal of Scott, T.M., 2001, Text to accompany the geologic map of Florida: Foraminiferal Research, v. 19, p. 180-184. Florida Geological Survey, open-File Report No. 80. Florida

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Geological Survey, Tallahassee, ISSN 1058-1391. Sexton, P.F., Wilson, P.A., and Pearson, P.N., 2006, Palaeoecology of late middle Eocene planktic foraminifera and evolutionary implications: Marine Micropaleontology, v. 60, p. 1-16. Spezzaferri, S., 1994, Planktonic foraminiferal biostratigraphy and taxonomy of the Oligocene and Lower Miocene in the oceanic record. An Overview: Palaeontographia Italica, v. 81, 187 p. , 1995, Planktonic foraminiferal paleoclimatic implications across the Oligocene-Miocene transition in the oceanic record (Atlantic, Indian, and South Pacific): Palaeogeography, Palaeoclimatology, Palaeoecology, v. 114, p. 43-74. , and Premoli Silva, I., 1991, Oligocene planktonic foraminiferal biostratigraphy and paleoclimatic interpretation from Hole 538A, DSDP Leg 77, Gulf of Mexico: Palaeogeography, Palaeoclimatology, Palaeoecology, v. 83, p. 217-263. Steineck, P.L., and Darrell, J.H., 1971, Cassigerinella winniana (Howe) from the Cook Mountain Eocene, Louisiana: Micropaleontology, v. 17, p. 357-360. Stolk, J., 1965, Contribution à l’étude des corrélations microfaunique du Tertiare inférieur de la Nigérie méridionale: Bureau Recherches Géologiques Minèrales Mémoire, v. 32, p. 247- 267. Turco, E., Bambini, A.M., Foresi, L.M., Iaccarino, S., Lirer, F., Mazzei, R., and Salvatorini, G., 2002, Middle Miocene high resolution calcareous plankton biostratigraphy at Site 926 (Leg 154, equatorial Atlantic Ocean): palaeoecological and paleobiogeographical implications: Géobios, v. 35, p. 257-276. Wade, B.S., Pearson, P.N., Berggren, W.A., and Pälike, H., 2011, Review and revision of Cenozoic tropical planktonic foraminiferal biostratigraphy and calibration to the geomagnetic polarity and astronomical time scale: Earth- Science Reviews, v. 104, p. 111-142.

Citation

Pearson, P.N., Spezzaferri, S., Huber, B.T., and Kučera, M., 2018, Taxonomy, biostratigraphy, and phylogeny of Oligocene Cassigerinella, in Wade, B.S., Olsson, R.K., Pearson, P.N., Huber, B.T. and Berggren, W.A. (eds.), Atlas of Oligocene Planktonic Foraminifera, Cushman Foundation of Foraminifer- al Research, Special Publication, No. 46, p. 481-494.

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