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Georgescu:Maquetación 1 Revista Española de Micropaleontología, 43 (3), 2011, pp. 173-207 ©Instituto Geológico y Minero de España ISSN: 0556-655X Iterative evolution, taxonomic revision and evolutionary classification of the praeglobotruncanid planktic foraminifera, Cretaceous (late Albian-Santonian) Marius Dan Georgescu Department of Geosciences, University of Calgary, 2500 University Drive NW, Calgary, Alberta T2N 1N4, Canada. [email protected] RESUMEN La aparión de conchas de praeglobotruncánidos tuvo un desarrollo iterativo en tres géneros/linajes direccionales durante el intervalo Albiense tardío-Santoniense. Se describe un nuevo género/linaje direccional, Bermudeziana, y una nueva especie, Fingeria praeglo- botruncaniformis. Como herramienta para describir los cambios morfológicos dentro de una misma estirpe y comparar el desarrollo en distintos linajes en los cuales ciertas características se alcanzaron de forma independiente, se desarrolla un nuevo sistema de tipos de especies obteniéndose una clasificación evolutiva. Palabras clave: Foraminíferos planctónicos, Cretácico, clasificación evolutiva, nuevos linajes direccionales, nueva especie. ABSTRACT The praeglobotruncanid test appearance was iteratively developed in three genera/directional lineages during the late Albian-Santon- ian. A new genus/directional lineage, Bermudeziana, and a new species, Fingeria praeglobotruncaniformis are described. A new sys- tem of species types is developed for the evolutionary classification, as a tool to describe the morphological changes within a lineage and compare the developments in lineages in which certain features were achieved independently. Keywords: Planktic foraminifera, Cretaceous, evolutionary classification, new directional lineage, new species. 1. INTRODUCTION tulates taxa grouping at various levels based only on mor- phological resemblances; such a practice tends, for exam- Iterative evolution is a widespread pattern among the Cre- ple, to bring together into the same genus distantly related taceous planktic foraminifera, which results in the repeti- species that independently developed one or a few mor- tive development of certain morphological features in phological features considered of taxonomic significance distant lineages. Its existence is a major challenge to a tax- by a knowledgeable and experienced specialist. Therefore, onomist who seeks to define natural, monophyletic units a new approach is necessary to develop a classification above the species level. This challenge is de facto gener- framework able to accommodate the results of the iterative ated by the use of the Linnaean classification, which pos- evolution. 173 Revista Española de Micropaleontología / v. 43 / nº 3 / 2011 A crucial aspect in refining the classification framework prominent on the earlier chambers and a main aperture is the resolution at which the test morphological features considered by various authors in either umbilical-extraum- are observed. The classical taxonomic frameworks for the bilical or extraumbilical-umbilical position (Bermúdez, Cretaceous planktic foraminifers were developed entirely 1952; Banner & Blow, 1959; Loeblich & Tappan, 1964, at the resolution of the optical stereomicroscope (Cush- 1987; Robaszynski & Caron, 1979; Caron, 1985; Korcha- man, 1927, 1948; Sigal, 1952a; Brönnimann, 1952; Gan- gin, 2003). The genus concept remained relatively stable dolfi, 1955; Reiss, 1957; Banner & Blow, 1959; over nearly 60 years and different author opinions focused Brönnimann & Brown, 1956; Bolli et al., 1957; Loeblich rather on the species to be included within it. A wider in- & Tappan, 1964, Pessagno, 1967). The introduction of the terpretation of Praeglobotruncana was given by Banner & scanning electron microscope in the current practice of Blow, 1959, who regarded it as consisting of three subgen- this foraminiferal group resulted in a significant increase era: P. (Praeglobotruncana), P. (Hedbergella) Brönnimann of observation resolution; such technological break- & Brown, 1958 and P. (Clavihedbergella) Banner & Blow, through provided us with the first high-detailed observa- 1959. The origin of Praeglobotruncana was considered tions of the test wall ultrastructure, ornamentation, among the globular-chambered tests of pre-Albian age porosity as well as a reevaluation of the delicate test fea- (i.e., Globigerina s.l.) by Reiss (1957, p. 142). Banner & ture, such as peripheral and periapertural structures, etc Blow (1959, p. 10) detailed the evolutionary origin and (Eicher & Worstell, 1970; Eicher, 1973, Michael, 1973; role of P. (Praeglobotruncana) by considering it transitional Longoria, 1973, 1974, etc). Despite this advance, the sub- between P. (Hedbergella) and Rotalipora. Praeglobotrun- sequent taxonomic reviews by Masters (1977), Robaszyn- cana’s origin from species of Hedbergella is generally ac- ski & Caron (1979), Robaszynski et al. (1984) and cepted today (Caron, 1983; Robaszynski et al., 1990; Hart, Loeblich & Tappan (1987) remained mostly based on the 1999; Hart et al., 2002). gross test architectural features. Testing the occurrence of iterative evolution in the case of High-resolution, SEM-observable test features (i.e., ultra- Praeglobotruncana is challenging mainly due to the stable structure and porosity) were used for the first time in the nature of the genus over several decades. The SEM was Cretaceous planktic foraminiferal taxonomy on a large extensively used to make high resolution observations on scale by Banner and Desai (1988). This study opened a the test ultrastructure, ornamentation, porosity and other new way towards understanding the iterative evolution delicate and therefore, rarely preserved morphological among several genera of the upper Cretaceous structures. Such observations were made in stratigraphic foraminifera, such as Pseudoguembelina Brönnimann & context, in order to observe the discrete morphological Brown, 1953 (Georgescu, 2007), Laeviheterohelix Neder- changes through time. Ultimately, the identification of lin- bragt, 1991 (Georgescu, 2009a), Clavihedbergella Banner eages as natural units with significance in evolutionary & Blow, 1959 (Georgescu, 2009b) and Ventilabrella Cush- classification that independently led to the development man, 1928 (Georgescu, 2010a). These developments also of praeglobotruncanid tests demonstrates the occurrence led to the definition of two kinds of lineages as taxonomic of iterative evolution within this group. units with significance in evolutionary classification: di- rectional and branched (Georgescu, 2010a). These devel- opments demonstrated also the necessity of high resolution taxonomic studies in order to identify the oc- 2. ITERATIVE EVOLUTION AND currences of iterative evolution and therefore refine the CHALLENGES TO CLASSIFICATION Cretaceous planktic foraminiferal classification. APPROACH Praeglobotruncana Bermúdez, 1952 is a frequent genus in The in-use Cretaceous planktic foraminiferal classification the upper Albian-middle Turonian sediments and accord- is mostly Linnaean. In this approach the iterative evolu- ing to the present-day acceptance represents a distinct tion pattern was rarely recognized and taken into consid- plexus characterized by a trochospiral test, peripheral eration in the taxa definitions (Robaszynski & Caron, structures consisting of closely-spaced pustules more 1979; Robaszynski et al., 1984; Loeblich & Tappan, 1987; 174 Marius Dan Georgescu, Iterative evolution, taxonomic revision and evolutionary classification of the praeglobotruncanid planktic foraminifera, Cretaceous (late Albian-Santonian) Nederbragt, 1991, etc). Linnaean classification practice at A distinct evolution pattern in the Cretaceous planktic the genus and species level is based on the premise that foraminifera is the high rate of morphological changes species showing a certain degree of resemblance can be pertaining to gross test architecture in certain lineages, re- grouped into genera (Fig. 1-A). Occurrence of a taxonom- sulting in frequent crossings over the boundaries between ically significant morphological feature or set of features Linnaean genera or higher taxonomic categories (Randri- can provide the reason for grouping the species, irrespec- ansolo & Anglada, 1989; Georgescu & Huber, 2008; tive of their ancestry. Practically, this approach in species Georgescu, 2009b, 2010b). This pattern, together with the grouping takes into consideration only the effects of iter- iterative evolution, induces a major challenge to the Lin- ative evolution. One of the cases of taxonomic solution naean classification, because different test morphologies, that accommodates the iterative evolution is represented which potentially require formalization at the genus level, by the separation between two Late Cretaceous homeo- can be successively developed in the same lineage. Ac- morphous genera, Helvetoglobotruncana Reiss, 1957 and cordingly, maintaining unchanged the classical taxonom- Gansserina Caron, González Donoso, Robaszynski & ically significant features, invariably results in a significant Wonders, 1984 of the early Turonian and late Campan- proliferation of monospecific genera and/or genus-line- ian-Maastrichtian respectively (Robaszynski et al., 1984). ages (Fig. 1-B). The latter kind of unit, although a step for- Figure 1. Schematic presentation of the classification approaches and how they can accommodate the iterative evolution pattern. A: a certain “mor- phology” developed in independent lineages is used to define a genus. B: although two lineages are recognized,
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