Research paperEvolution of the silicoflagellate naviculopsid skeletal morphology in the Cenozoic
Introduction
Silicoflagellates are a group of marine single-celled algae with a geologic record that extends to at least the late Early Cretaceous, about 115 million years ago (Ma) (McCartney et al., 1990, McCartney et al., 2014a). Of the diverse Cenozoic silicoflagellate skeletal morphologies, the simplest is the naviculopsid design characterized by a 2-sided basal ring with major-axis spines and simple apical structure, best known from Naviculopsis (Fig. 1), which ranges from late middle Paleocene (~59.9 million years ago, McCartney et al., 2018) to early Miocene (~17.9 Ma, below). The apical structure of this morphology consists of an arch composed of either a single element or plate, or two elements that meet at a triple junction and apex spine (see McCartney and Witkowski, 2016). Other skeletal morphologies occur within Naviculopsis to include species characterized by three or four struts that meet at an apex, and 4-strutted morphologies with minor-axis bridge. Since a triple junction is established by two basal elements and an arch, there is no basal spine on the minor-axis. Skeletons of other genera can also, as part of species variability or as aberrant specimens, display the naviculopsid morphology.
General silicoflagellate phylogenies for the Cenozoic (Lipps, 1970; McCartney et al., 1990, McCartney et al., 2015a; Prema, 1996) begin with the 3-sided Corbisema that survived the K/Pg boundary event (Harwood, 1988; McCartney et al., 2010, McCartney et al., 2015a). McCartney et al. (2018) provided more constrained ages for the sequence of evolutionary events during the Paleocene. The first divergence of a new silicoflagellate genus – Pseudonaviculopsis – occurs in the early Paleocene before ~63.9 Ma. Dictyocha appears in the late early Paleocene (~61.4 Ma), prior to the first appearance of Naviculopsis in the late middle Paleocene (~59.9 Ma). Gleser, 1962, Gleser, 1966 is the only researcher to previously illustrate phylogenetic interpretations of various Paleogene silicoflagellates, including Naviculopsis, at the species level. The present paper provides more detailed phylogenetic interpretation and follows a study of silicoflagellate evolution in the Cretaceous (McCartney et al., 2010). A more general paper on silicoflagellate Cenozoic evolution is in preparation.
Section snippets
Methods
This paper provides evolutionary interpretations for 36 taxa that commonly exhibit the naviculopsid skeletal morphology, with authorities for original descriptions, transfers and emendations listed in Table 1. Deep sea sediment samples from the Deep Sea Drilling Project (DSDP) and Ocean Drilling Program (ODP) were provided by the International Ocean Discovery Program (IODP), which administrates cores drilled at various Sites that can include multiple Holes. Specific samples examined for this
Naviculopsis (or naviculopsid) skeletal morphology
The naviculopsid morphology, best known within the genus Naviculopsis, has only two basal sides connected by an arch often composed of a single element (Fig. 1A). More complicated apical structures occur within Naviculopsis, and can include a curved band (Fig. 1B) that attaches to both sides or an arch composed of two elements that meet at an apical triple junction that includes an apex spine. The term 'naviculopsid‘ may be applied to sometimes aberrant skeletal morphologies included within
New perspectives on evolution of the naviculopsid skeletal morphology
Presented below are the geologic histories of three general silicoflagellate groups that commonly display the naviculopsid skeletal morphology. The first two of these, Pseudonaviculopsis and Naviculopsis, each descend from separate groups of Corbisema, which may have independent origins from the Late Cretaceous genus Cornua (McCartney et al., 2011b, fig. 5). Pseudonaviculopsis developed from the C. archangelskiana (Schulz) Frenguelli group of species with large skeletons and the star-of-David
Constraints on silicoflagellate skeletal design
Naviculopsis has, after Corbisema, the longest geological history of an extinct silicoflagellate genus. Both Naviculopsis and Corbisema became extinct during the Miocene, replaced by silicoflagellate genera that have more complicated skeletal structures. Essential questions concern the advantage offered by the Naviculopsis construction early in history of the group, and the advantage of replacement genera. Naviculopsis has an unusual basal design with an elongate axis while all other
New combinations
Corbisema cunicula subsp. alta (Ciesielski) McCartney comb. nov.
Basionym: Corbisema hastata subsp. alta Ciesielski, 1991, p. 75–76, pl. 4, figs. 1, 2; pl. 9, fig. 10.
Remarks: The late Paleocene to early Eocene includes diverse Corbisema with relatively long spines and small (~20 μm) basal ring. This diversity, placed in C. hastata subsp. cunicula by Bukry (1976a) and C. triacantha subsp. lepidospinosa by Ciesielski (1991), was observed by McCartney et al. (2018) to be difficult to consistently
Declaration of Competing Interest
The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper.
Acknowledgements
We thank the Deep Sea Drilling Project, Ocean Drilling Program and Integrated Ocean Drilling Program and in particular Phil Rumford for providing samples for comparative study. Marlena Świło and Krzysztof Owocki provided assistance in the SEM photography of specimens at the Institute of Paleobiology, Polish Academy of Science, Warsaw. Manfred Ruppel assisted with the SEM photography at J. W. Göthe-Universität, Frankfurt am Main, Germany. Małgorzata Bąk assisted with figures and other technical
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