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A Taxonomic Revision of the Unilocular Foraminifera

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A Taxonomic Revision of the Unilocular Foraminifera Journal of Foraminiferal Research, v. 17, no. 3, p. 212-226, pl. 1-5, July 1987 A TAXONOMIC REVISION OF THE UNILOCULAR FORAMINIFERA R. TIMOTHY PATTERSON AND RANI H. RICHARDSON Department of Earth and Space Sciences, University of California, Los Angeles, CA 90024 ABSTRACT dance, high species diversity, and small size. As a re- Unilocular foraminifera are varied and diverse, yet sult, the group generally has been ignored by most among the least understood gronp of the foraminifera. foraminiferal workers. Part of the problem has been the inadequate number Prior to Cushman's reclassification of the forami- of taxonomic divisions within this varied group. We nifera in 1928, most authors followed H. B. Brady propose a new taxonomic framework to make the group (1884) in referring all single-chambered foraminifera more useful to taxonomists and stratigraphers alike, to Lagena Walker and Jacob. Most of these early work- and to eventually lead to a better understanding of the ers recognized relatively few species, but a large num- group as a whole. ber of varieties. Monographs on Lagena. such as that Rather than subdividing the unilocular foraminifera by Sidebottom (1912, 1913), clearly illustrate this ap- into more than one family, we reinstate the family La- proach. genidae Reuss, 1862, to include all unilocular forms, Cushman (1928) limited the use of Lagena to species and recognize three subfamilies. The first subfamily, with or without a neck, having a rounded, radiate, el- Lageninae Reuss, 1862, encompasses genera that lack liptical, or slitlike terminal aperture. He placed Oolina an entosolenian tube, including Lagena Walker and d'Orbigny, Fissurina Reuss, Trigonulina Seguenza, Jacob, and the reinstated Procerolagena Puri. The sec- Amphorina d'Orbigny, Vermiculum Montagu, and ond subfamily, Ellipsolageninae Silvestri, 1923, accom- other genera in synonomy with Lagena. To accom- modates genera with an entosolenian tube at some stage modate species with a rounded test and internal tube, of the life cycle, including Oolina d'Orbigny, Cush- Cushman reinstated Entosolenia Williamson and manina Jones, Fissurina Reuss, Galwayella Patterson transferred it from the family Lagenidae to the Buli- and Pettis, I renita Jones, Lagenosolenia McCulloch, minidae. Cushman also referred lagenid foraminifera Parafissurina Parr, Pseudofissurina Jones, Pseudooli- with an internal tube and projecting hood to Ellipso- na Jones, Pseudosolenina Jones, Solenina Jones, Ven- lagena Silvestri, which he placed in the family Ellip- trostoma Schnjtker, Walterparria Jones, and Wiesner- soidinidae. ina Jones. Four new genera are also included in the Parr (1947) recognized Lagena, Dolina, Fissurina, Ellipsolagenidae: Duplella (type species D. apexadina, and Parafissurina Parr and placed them all in the fam- n. sp.), Palliolatella (type species P. arita, n. sp.), Pris- ily Lagenidae. In the conclusion of his revision, Parr tinosceptrella (type species P. hispida, n. sp.), and Va- acknowledged the inadequacy of the classification of sicostella (type species V. helophoromarginata (Jones)). unilocular foraminifera and wrote". the writer re- We also describe two additional new species belonging alizes that the genera he has recognized could justifi- to this subfamily: Vasicostella singulara and Wiesner- ably be subdivided into a larger number of genera, ... ina carinata. (the) double wall seen in some species, ... the marginal We propose a new third subfamily, Sipholageninae, tubes in (some) species, . the development of hori- to include those unilocular genera with double walls zontal tubules (of others) . are all much more radical connected by a network of pillars. This subfamily con- differences. than the differences on which Dolina. tains Sipholagena Moncharmont-Zei and Sgarrella and Lagena. Fissurina, and Parafissurina are separated." Pytine Moncharmont-Zei and Sgarrella. Loeblich and Tappan (1964) recognized Dolina, Fis- surina, and Parafissurina and placed them in the subfamily Oolininae in the family Glandulinidae, based INTRODUCTION on the presence of entosolenian tubes. They placed Lagena in the subfamily Nodosariinae in the family Unilocular foraminifera are varied and diverse, yet Nodosariidae. Procerolagena Puri was placed in syn- one of the least understood groups of foraminifera. The onomy with Lagena. low number of taxonomic divisions within this varied In the most ambitious revision to date, Jones (1984) group suggested to us that a rigorous reexamination recognized 23 genera of unilocular foraminifera. In the was needed. We examined several thousand specimens subfamily Oolininae, Jones recognized Dolina. Fis- from Holocene localities around the world to gain as surina, and Lagenosolenia along with nine new genera; wide an exposure as possible to the myriad morpho- Arthurina Jones (a junior synonym of Ventrostoma logic variations. In addition, some fossil unilocular Schnikter), Buchnerina Jones, Cushmanina Jones, Ire- foraminifera, as old as Late Cretaceous, were studied nita Jones, Pseudofissurina Jones, Pseudoolina Jones, to broaden the taxonomic interpretations. Pseudosolenina Jones, Solenina Jones, and Wiesnerina Jones. In the Lingulinidae Loeblich and Tappan, Jones TAXONOMIC HISTORY erected the subfamily Rimulininae to include Rimu- Taxonomic difficulties associated with unilocular fo- lina d'Orbigny and Rimulinoides Saidova. Rimulina raminifera arise in part from their relatively low abun- is based on a single specimen, R. glabra d'Orbigny, 212 TAXONOMIC REVISION OF UNILOCULAR FORAMINIFERA 213 and although originally described as being multilocu- TABLE 1. Percentage of Lagenosolenia incomposita Patterson and lar, Loeblich and Tappan (1955) found this not to be Pettis with entosolenian tube. evident. Due to the fact that no additional specimens have been found, and that the type level is uncertain, we do not recognize Rimulina within the Lagenidae. Rimulinoides is described as being cylindrical, with a few transverse growth lines, and with a sieve-like ap- erture having three radiating grooves. Although Sai- dova's (1975) original type illustration is poor, her description suggests a close affinity with Chrysalogo- nium (Nodosariinae). Jones (1984) described the biloc- ular genus Heteromorphina (Glandulininae), suggest- ing that it may be the microspheric generation of (1984) recognized that evolutionary pathways are by Oolina. We do not include Heteromorphina in our clas- no means proven, and that further work is required to sification due to the insufficient evidence in support of discern phylogenetic trends. Rather than separating the this hypothesis. Jones (1984) established the genus Phi- unilocular foraminifera into more than one family based alinea Jones and the subfamily Phialineinae in the on conjecture, we reinstate the family Lagenidae Reuss family (?)Eouvigerinidae Cushman. This genus, re- to include all unilocular taxa, and recognize within it, corded by Jones (1984) as ranging from the Miocene three subfamilies. The first subfamily, Lageninae Reuss, to Holocene, was based on Miliola elongata Ehrenberg includes Lagena and Procerolagena Puri, which lack (1844; figured in Ehrenberg, 1854), an unrecognizable an entosolenian tube. The second subfamily, Ellipso- fragment from Cretaceous strata of western Syria. Eh- lageninae Silvestri, is composed of genera that have an renberg (1844) gave the type locality as Kurdistan, entosolenian tube at some stage of the life cycle. We whereas Ehrenberg (1854, pl. 25, la, fig. 1) listed the propose a new third subfamily, Sipholageninae, to in- type locality of the species as Al Jabal Ash Sharqi (Anti- clude unilocular genera having double walls connected Lebanon) in western Syria. Jones placed Parafissurina by a network of pillars. and Walterparria Jones in the subfamily Parafissuri- ninae Jones, in the Pleurostomellidae. He placed La- CRITERIA FOR DISTINGUISHING gena and Pseudarcella Spandel in the Lagenidae, in the GENERA order Incertae Sedis. Pseudarcella is, in fact, a tintinnid (Lindenberg, 1965; Tappan and Loeblich, 1968). Jones Despite the great species diversity seen in the studied (1984) also assigned Cribrolagena Jones to the family samples and in the literature, some phenotypic group- Incertae Sedis, in the superfamily Nodosariacea. ings are recognizable. These groups are based on sur- face sculpture, test shape, wall structure, wall perfo- rations, apertural configuration, and carinal EVOLUTIONARY HISTORY development. We recognize 22 genera according to these criteria, including four new genera, four genera The earliest record of lagenid foraminifers is from recognized by Loeblich and Tappan (1964), 13 de- Lias of Germany (and elsewhere). Franke (1936) re- scribed since 1964, and one reinstated from synonomy. corded Liassic species having generally circular axial sections, some possessing entosolenian tubes. He also SIGNIFICANCE OF ENTOSOLENIAN TUBE reported a single species externally similar to Fissurina. By Early Cretaceous time, unilocular foraminifera were Live culture studies of Glabratella (Myers, 1940), more diverse, although they still closely resembled Ju- Tretomphalus (Myers, 1943), Rubratella (Grell, 1958), rassic species (Parr, 1947). Haeusler (1887) figured and Orbulina (Le Calvez, 1936, 1947) have shown that compressed, carinate forms from the Neocomian of some species (including unilocular forms), resorb in- Switzerland that are similar to those found in modern ternal calcareous structures during reproduction. In ad- oceans. dition, Taylor and others (1985) found that
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