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Species Acarinina bullbrooki Bolli 1957


Diagnosis / Definition:
Pearson et al. (2006):
DESCRTPTION. Type of wall: Strongly muricate, non-spinose, normal perforate. Test morphology: Quadrate, low-trochospiral; 4 inflated, moderately embracing, hemispherical chambers in last whorl; intercameral sutures radial, straight, depressed; umbilicus narrow to moderately wide in some (usually) younger forms, no circumumbilical rim of concentrated muricae; aperture a low, rimmed opening extending towards, but not reaching, the periphery; spiral side with lunate shaped chambers distributed in 2-2 1/2 whorls; small, discrete opening(s) are observed rarely on well-preserved individuals; sutures weakly curved; edge view rounded to subangular; moderate marginall peripheral concentration of muricae gives impression of pseudocarina in some instances. Size: Maximum diameter of holotype 0.43 mm, thickness 0.32 mm.
Discussion / Comments:
Toumarkine & Luterbacher (1985):
Acarinina bullbrooki differs from A.spinuloinflata by a flatter spiral side, a much more conical umbilical side and more subangular chambers which increase more rapidly in size. The general shape of the test is more quadrangular. Globorotalia (Acarinina) matthewsae Blow, 1979 (Figs. 29.11-13) is very close to A. bullbrooki but has a less flat spiral side, a less conical umbilical side and a more rounded periphery in side view, but nevertheless its periphery is not as rounded as in A. spinuloinflata.
Pearson et al. (2006):
DISTINGUlSHlNG FEATURES.- Acarinina bullbrooki is distinguished by its (sub)quadrate test, closely appressed and embracing chambers, rounded to subangular periphery, and weak to moderate murical concentration on the pcriphery. It differs from A. praetopilensis and related forms in having less incised sutures, generally smaller muricae, and by populations generally being approximately randomly coiled. DISCUSSION.- Subbotina (1947, p. 129, pls. 8, 9; 1953, p. 223, pl. 21) was one of the first to recognize and illustrate the plexus of late early to middle Eocene acarininids now associated with the bullbrooki and punctocarinata group. She referred a suite of specimens (pl. 2 1, f gs. l a-7c) to the Pliocene-Holocene taxon crassaformis. The illustrated forms would appear to be referable to A. wilcoxensis/quetra (pl. 21, figs. la-c) from the G. marginodentata Subzone (lower Eocene); A. decepta (pl. 21, figs. 2a-3c; see Blow, 1979, pl. 149, figs. 1-7; pl. 154 (from the lower Eocene Zone of conical globorotaliids); Acarinina bullbrooki (pl. 21, figs. 4ac) from the Zone of conical globorotaliids; A. quetra (pl. 21, figs. 5a-c) fiom the lower part of the Zone of acarininids, middle Eocene; Acarininaprivzitiva (pl. 2 1, figs. 7a-c) from the upper middle Eocene Lyrolepis caucasica Beds, Zone of thin-walled pelagic foraminifera; and an apparent morozovelloidid referable to Morozovelloides crassatus (pl. 21, figs. 6a-c (from the Zone of thin-walled pelagic foraminifera [~l= Zone P 141). The status of this taxon has long remained in limbo because of the uncertainty surrounding the generic identity of the holotype of Globorotalia bullbrooki Bolli and its uncertain relationship with Pulvinulina crassata var. densa Cushman (see discussion in Blow, 1979). Pessagno (1 961) suggested that Globorotalia bullbrooki Bolli should be considered a junior synonym of densa, based on examination of the primary types at the USNM, a viewpoint accepted by Berggren (1 966, l 968, 1977). Blow (1979, p. 9 15) noted that although he had made a cursory examination of the holotype of densa during a visit the U.S. National Museum in 1965 it was not found during a subsequent visit to the museum in May 1970 and presumed to be missing, a view seemingly confirmed in a subsequent letter to Blow from the then curator Richard Cifelli that the holotype of densa was indeed "not in our collections and I have no knowledge of it" (letter by Cifelli to Blow, 15th September 1970). Blow (1 979: 9 15) suggested that unless the holotype was found it would be better to consider densa a nomen non conservandum, because of its similar appearance to decepta Martin, praetopilensis Blow, and pseudotopilensis Subbotina and the fact that it might be referable to either bullbrooki or any one of these three forms. Enigmatically the holotype of densa (Cushman Collection 3027) was described and redrawn only two years later by Cifelli (1972). The specimen appears to exhibit a beaded peripheral keel, hallmark of a morozovellid rather than an acarininid. Although poorly preserved, we have illustrated the holotype in SEM and have assigned it (questionably) to Morozovelloides crassatus (see Pearson and Berggren, Chapter 10, this volume). The holotype of bullbrooki (USNM P5742) has a subangular peripheral margin and lacks a peripheral muricocarina. The paratype of bullbrooki (USNM P5743) is somewhat smaller and has a subrounded periphery in edge view. Toumarkine and Bolli have deposited a suite of specimens of bullbrooki in the Cushman Collection at the USNM from Site 313 of DSDP Leg 32 which includes individuals with muricate (but unkeeled) peripheral margins varying fi-om angulate to subangular to subrounded, reinforcing the understanding that Bolli included a wide range of variation in his concept of hullbrooki, an interpretation which we follow here. Blow (1979, p. 926-928) treated the taxon Acarinina decepta (Martin) in some detail, the essential features of which are listed below: l ) Acarinina decepta was interpreted as ancestral to A. matthewsae by way of intermediate morphotypes of A. nitida; 2) A. decepta is distinguished from its ancestor A. pseudotopilensis by a slight increase in the number of chambers in the last whorl, reduction in the tangential elongation of the later chambers and in having a more quadrag (as opposed to lobulate) peripheral outline; 3) The morphotypes decepta and nitida lie at the early end of the morphogenesis that leads from pseudotopilensis to matthewsae; 4) A. decepta also forms a link between A. pseudotopilensis and A. cuneicamerata; 5) A. pseudotopilensis evolved into the angulate A. topilensis without passing through a "decepta" intermediate stage; in similar fashion A. pseudotopilensis evolved into A. bullbrooki at stratigraphic levels prior to the "decepta/nitida" morphotypes; 6) The name decepta serves as a useful means of recognizing the complex of forms from which the extreme end members matthewsae and cuneicamerata differentiate from the pseudotopilensis rootstock. The various illustrations provided by Blow (1979) of the taxa cited above do point to the intermediate nature of the morphology of A. decepta. Reference to the illustrations of Martin (1943, pl. 7) and Blow (1979, pls. 149, 154) suggest closest affinities of decepta with bullbrooki (compare Blow, 1979, pl. 149, fig. 4 with Blow, 1979, pl. 155, fig. 6, for example). In this work we have included A. matthewsae as a junior synonym of A. bullbrooki. In view of the intermediate morphologies expressed in A. decepta and its enigmatic affinities we include A. decepta here provisionally as a dubious prior synonym of A. bullbrooki. Globigerina spinuloinflata Bandy, 1949 has long been considered a junior synonym of either densa or a senior synonym of bullbrooki as Bandy (1964, p. 6) pointed out. (Re)illustration / drawing of the holotype of spinuloinflata Bandy by Cifelli (1972) shows marked similarities with bullbrooki, even though Cifelli called attention to the somewhat more rounded periphery of spinuloinflata in edge view as did Toumarkine and Bolli (1975, p. 130). We have been able to illustrate the holotype of Globigerina spinuloinflata Bandy from the Tallahatta Formation ofAlabama. While it is difficult to place this form, we believe it may be conspecific with bullbrooki as Bandy proposed, in which case it could be regarded as the senior synonym. We retain bullbrooki, however, in the interests of nomenclatural stability, because in contrast to spinuloinflata, bullbrooki is a wellknown form with historically consistent usage in the works of Bolli and coauthors and the holotype of spinuloinflata is not a common or central morphotype among the middle Eocene acarininids that have historically been referred to bullbrooki. Acarinina bullbrooki is a widely used taxon in middle Eocene biostratigraphy. However, its biostratigraphic utility has been somewhat compromised by uncertainty regarding its taxonomic affinities. With the recognition that densa, decepta, and spinuloinflata cannot be definitively placed, the biostratigraphic utility of bullbrooki can now be re-established. PHYLOGENETTC RELATIONSHIPS.- Probably evolved from A. boudreauxi (in Zone E7). STRATIGRAPHIC RANGE.- ZONE E7 to Zone E l l. GEOGRAPHIC DISTRIBUTION.- Cosmopolitan distribution (see synomy above). STABLE ISOTOPE PALEOBIOLOGY.- Boersma and others (1987) and Pearson and others (1993,2001) record this species as having isotope ratios indicative of a surface mixed layer habitat.
Systematics:

1
 Superregnum Eukaryota
  Regnum Protoctista
   Phylum Ciliophora
    Subphylum Postciliodesmatophora
     Ordo Globigerinida
      Superfamilia Globorotaliaceae
       Superfamilia Nonionacea
        Familia Truncorotaloididae
         Genus Acarinina
          Species Acarinina bullbrooki

15
  Classis Foraminifera
   Genus Acarinina
    Species Acarinina bullbrooki

35
  Ordo Foraminiferida
   Superfamilia Globigerinaceae
    Familia Truncorotaloididae
     Genus Acarinina
      Species Acarinina bullbrooki
Synonym list:
Toumarkine & Luterbacher (1985):
1957 Globorotalia bullbrooki Bolli. - Bolli : p.167 pl 38 figs 5a-b (type reference)
1985 Acarinina bullbrooki Bolli. - Toumarkine & Luterbacher : p.129 figs 29.4-10
Pearson et al. (2006):
1930 Globorotalia crassata var. densa Cushman. - Cushman & Barksdale : p.68 pl. 12; fig. 8a-b [Martinez Fm., Martinez, California]
1939 Globorotalia crassata var. densa Cushman. - Cushman : p. 74, 75 pl. 12; fig. 20a-b [northwest Atlantic Ocean, off eastern North America]
? 1943 Globigerina decepta Martin. - Martin : p.114 pl. 7; fig. 2a-c [lower Eocene, Lodo Fm, Fresno Co., California]
1948 Globorotalia crassata var. densa Cushman. - Cushman & Renz : p.40 pl. X; fig. 7, 8 [Navet Fm., Trinidad]
1949 Globorotalia crassata var. densa Cushman. - Bandy : p.80 pl. 12; fig. 4a-c [Tallahatta Fm., Clarke County, Alabama]
1949 Globorotalia (Truncorotalia) crassata densa Cushman. - Cushman & Bermudez : p.38 pl. 7; fig. 10-12 [Matanzas Province, Cuba]; [Not Cushman, 1925]
? 1949 Globigerina spinuloinflata Bandy. - Bandy : p.122 pl. 23; fig. 1a-c [Tallahatta Fm., Claiborne Group, Little Stave Creek, Clarke County, Alabama]
p 1953 Globorotalia crassaformis Galloway & Wissler. - Subbotina : p.223 pl. 21; fig. 4a-c [Zone of conical globorotaliids, Green Group, Kuban River, North Caucasus]; [Not Galloway and Wissler, 1927]
1957 Globorotalia bullbrooki Bolli. - Bolli : p.167 pl. 38; fig. 4a-c and 5a-b (holotype) [Zone P10, Navet Fm., Trinidad]
1959 Globorotalia crassata var. densa Cushman. - Hamilton & Rex : p.793 pl. 252; fig. X9 [Sylvania Guyot, Bikini Atoll, Marshall Islands]
1960 Globorotalia crassata var. densa Cushman. - Todd & Low : p.850 pl. 259; fig. 13a-c [Eniwetok Atoll, Marshall Islands]
1960 Globorotalia densa Cushman. - Pessagno : p.99 pl. 5, fig. 3 [south-west Puerto Rico]; pl. 1, fig. 1-3 [Jacaguas Group, south central Pucrto Rico] [Not Cushman, 1925]
1962 Globorotalia crassata var. densa Cushman. - Saito : p. 214-215 pl. 33; fig. 5a-11c [Haha Jima, Bonin Islands, western Pacific Ocean]
1962 Globorotalia crassata var. densa Cushman. - Aubert : p.58 pl. 2; fig. 4a-c [middle Eocene, Morocco]; [Not Cushman, 1925]
? 1964 Globorotalia spinuloinflata Bandy. - Bandy : p.6 pl. 4; fig. 19 (reillustration of holotype)
? 1971 Globorotalia (Acarinina) spinuloinflata Bandy. - Jenkins : p. 83, 84 pl. 4, fig. 99-101; pl. 5, fig. 119-122 [G. (C.) index Zone, Hampden Beach, lower part of Bortonian Stage, New Zealand; = tightly coiled, 4 1/2 chambered form]
? 1972 Globigerina spinuloinflata Bandy. - Cifelli : p.159 fig. 2a-c (reillustration of holotype); [lower Eocene, Tallahatta Fm., Claiborne Group, Little Stave Creek, Alabama]
1977 Acarinina densa Cushman. - Berggren : p.259 chart-no. 12 (reillustration of figured specimens in literature); [Not Cushman, 1925]
? 1979 Globorotalia (Acarinina) decepta Martin. - Blow : p.926 pl. 149, fig. 2-6 [lower Eocene, Zone P9, KANE 9-C piston core, Endeavour Seamount, equatorial Atlantic Ocean]; pl. 154, fig. 5 [middle Eocene, Zone P1 0, KANE 9-C piston core, Endeavour Seamount, equatorial Atlantic Ocean]
p 1979 Globorotalia (Acarinina) bullbrooki Bolli. - Blow : p.915 pl. 149, fig. 8 and 9; pl. 155, fig. 1-2, 6-8[Zone P9-P10, KANE 9, piston core 42, Endeavour Seamount, equatorial Atlantic Ocean]; pl. 17 1, fig. 1 -3 and 7-9 [Zone P 1 1, Kilwa area, Tanzania] (not figs. 3-5 =A. mcgowrani)
1979 Globorotalia (Acarinina) cf. bullbrooki Bolli. - Blow : p.915 pl. 187, fig. 6, 7 [DSDP Site 19, South Atlantic Ocean]
1979 Globorotalia (Acarinina) matthewsae Blow. - Blow : p.935 pl. 170, fig. 1 (holotype)-9; pl. 203, fig. 3 = detail of pl. 170, fig. 2 [Zone P11, sample RS. 24, Kilwa area, Tanzania]; pl. 179, fig. 1 and 2 [DSDP Hole 21A, South Atlantic Occan]; pl. 187, fig. 5 [Zone P 12, DSDP Site 19, South Atlantic Ocean], pl. 204, fig. 1-5; pl. 205, figs. 1-6 [Sample RS. 24, Kilwa area, Tanzania]
1979 Globorotalia (Acarinina) sp. . - Blow : p.915 pl. 143, fig. 1-5 ex interc. G (A.) pseudotopilensis (Subbotina) and G. (A.) bullbrooki (Bolli); [Zone P8b, DSDP Hole 20C, South Atlantic Ocean]
1991 Acarinina bullbrooki Bolli. - Huber : p.439 pl. 3; fig. 6, 7 [Acarinina collactea (AP10) Zone, ODP Site 738, Kerguelen Plateau, southern Indian Ocean]
1991 Acarinina matthewsae Blow. - Huber : p.439 pl. 3; fig. 3, 4 [middle Eocene Zone AP8, ODP Hole 738B, Kerguelen Plateau, southern Indian Ocean]
1992 Acarinina bullbrooki Bolli. - Berggren : p.563 pl. 2; fig. 9, 10 [Acarinina bullbrooki /A. matthewsae Zone; ODP Hole 748B, Kerguelen Plateau, South Indian Ocean]
1992 Acarinina matthewsae Blow. - Berggren : p.563 pl. 2; fig. 7, 8 [Acarinina bullbrooki Zone, middle Eocene, ODP Hole 749B, Kerguelen Plateau, southern Indian Ocean]
2004 Acarinina matthewsae Blow. - Pearson et al. : p.37 pl. 2; fig. 5, 6 [middle Eocene, Zone P11, Tanzania Drilling Project Site 2, Kilwa Masoko, Tanzania]
2006 Acarinina bullbrooki Bolli. - Pearson et al. : p.274 pl. 9.6; fig. 1-16 (Pl. 9.6, Figs. 1-3, 5-7: new SEMs of holotype and paratype of Globorotalia bullbrooki Bolli) (Pl. 9.6, Fig. 13: reillustration of holotype of Globorotalia (Acarinina) matthewsae Blow) (PI. 9.6, Fig. 16: new SEM of holotype of Globigerina spinu
Specimen:
Smithsonian Museum of Natural History, Washington, D.C., Inventory number: USNM P5742
References:

Cushman,J.A. and Barksdale,J.D.. (1930):
Eocene foraminifera from Martinez, California . Stanford University, Department of Geology, Contributions Vol. 1 p. 55-73

Cushman,J.A. (1939):
Eocene Foraminifera. from submarine cores off the eastern coast of North America . Contr. Cushman Lab. Foram. Res. Vol. 15 p. 49-76

Martin,L.T. (1943):
Eocene formainifera from the type Lodo Formation, Fresno County, California . Stanford University Publications Geological Sciences Vol. 3 p. 93-125

Cushman,J.A. and Renz,H.H. (1948):
Eocene Foraminifera of the Navet and Hospital Hill formations of Trinidad, B.W.I. . Cushman Lab. Foram. Res., Spec. Publ. Vol. 24 p. 1-42

Cushman,J.A. and Bermudez,P.J. (1949):
Some Cuban species of Globorotalia . Contributions from the Cushman Foundation for Foraminiferal Research Vol. 25 p. 26-44

Bandy,O.L. (1949):
Eocene and Oligocene foraminifera from Little Stave Creek, Clarke County, Alabama . Bulletins of American paleontology Vol. 32 p. 1-211

Subbotina,N.N. (1953):
Iskopaemye foraminifery SSSR (Globigerinidy, Khantkenininidy i Globorotaliidy) . Trudy Vsesoyznogo Nauchno-Issledovatel'skogo Geologo-razvedochnogo Instituta (VNIGRI) Vol. 76 p. 296

Bolli,H.M. (1957):
Planktonic Foraminifera from the Eocene Navet and San Fernando formations of Trinidad, B.W.I. . Bull. U.S. natl. Mus. Vol. 215 p. 155-172

Hamilton,E.L. and Rex,R.W. (1959):
Lower Eocene phosphatized Globigerina ooze from Sylvania buyot. . U. S. Geological Survey Professional Paper p. 785-798

Pessagno,E.A. (1960):
Stratigraphy and micropaleontology of the Cretaceous and lower Tertiary of Puerto Rico . Micropaleontology Vol. 6(1) p. 87-110

Todd,R. and Low,D. (1960):
Smaller foraminifera from Eniwetok drill hole . U.S. Geological Survey, Professional paper 260-X p. 799-861

Aubert,J.. (1962):
Les Globorotalia de la region prerifaine (Maroc septentrional) . Notes et Memoires du Service Geologique Maroc Vol. 22 p. 1-156

Saito,T. (1962):
Eocene planktonic foraminifera from Hahajima (Hillsborough Island) . Trans. Proc. Paleontol. Soc. Japan, news series Vol. 45 p. 209-225

Bandy,W.A. (1964):
The type of Globorotalia crassata (Cushman) . Contributions from the Cushman Foundation for Foraminiferal Research Vol. 15 p. 34-35

Jenkins,D.G. (1971):
New Zealand Cenozoic Planktonic Foraminifera . New Zealand Geological Survey Paleontological Bulletin Vol. 42

Cifelli,R. (1972):
The holotypes of Pulvinulina crassata var. densa Cushman and Globigerina spinuloinflata Bandy . Journal of Foraminiferal Research Vol. 2 p. 157-159

Berggren,W.A. (1977):
Atlas of Paleogene planktonic foraminifera. Some species of the genera Subbotina, Planorotalites, Morozovella, Acarinina and Truncorotaloides.
In: Oceanic Micropaleontology Eds: Ramsay, A.T.S. p. 205-299

Blow,W.H. (1979):
The Cainozoic Globigerinida. 3 Vols p. 1413 pp

Toumarkine,M. and Luterbacher,H.P. (1985):
Paleocene and Eocene Planktic Foraminifera.
In: Plankton Stratigraphy p. 87-154

Huber,B.T. (1991):
Paleogene and early Neogene planktonic foraminifer biostratigraphy of Sites 738 and 744, Kerguelen Plateau (southern Indian Ocean).
In: Proceedings of the Ocean Drilling Program, Scientific Results Vol. 119 Eds: Leckie, R.M.Sigurdsson, H.Acton, G.D.Draper, G. p. 427-449

Berggren,W.A. (1992):
Paleogene Planktonic Foraminifera Magnetobiostratigraphy of the Southern Kerguelen Plateau (Sites 747-749).
In: Proceedings of the Ocean Drilling Program, Scientific Results Vol. 120

Pearson,P.N.; Nicholas,C.J..; Singano,J.M..; Bown,P.R..; Coxali,H.K..; van Dongen,B.E..; Huber,B.T.; Karega,A..; Lees,J.A..; Misaky,E..; Pancost,R.D..; Pearson,M.. and Roberts,A.P.. (2004):
Paleogene and Cretaceous sediment cores from the Kilwa and Lindi areas of coastal Tanzania: Tanzania Drilling Project Sites 1-5 . Journal of African Earth Sciences Vol. 39 p. 25-62

Pearson,P.N.; Olsson,R.K.; Hemleben,C.; Huber,B.T. and Berggren,W.A. (2006):
Atlas of Eocene Planktonic Foraminifera. p. 1-513
Anonymous: Unedited TaxonConcept data
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