Morphological distinction of molecular types in Ammonia – towards a taxonomic revision of the world’s most commonly misidentified foraminifera

  • publication
  • 01-02-2004

Bruce W. Hayward, Maria Holzmann, Hugh R. Grenfell, Jan Pawlowski, Chris M. Triggs. Marine Micropaleontology, Volume 50, Issues 3–4, 2004, Pages 237-271, ISSN 0377-8398,

In this study, morphometric analysis has been performed on 178 Ammonia specimens belonging to 12 different molecular types, plus non-sequenced type specimens of Ammonia beccarii and A. tepida. Molecular type distinction is based on phylogenetic analysis of 267 partial LSU rDNA sequences, obtained from 202 living Ammonia specimens, sampled in 30 localities from 17 countries bordering the Pacific Ocean, Atlantic Ocean, Mediterranean Sea, Caribbean Sea and North Sea. Restriction fragment length polymorphism (analysis was carried out for another seven specimens. Morphometric analysis was based on measurements or assessments of 37 external test characters in spiral, umbilical, profile and close-up Scanning Electron Microscopic views. Cluster analysis, canonical variates analysis, and detrended correspondence analysis, performed on the morphological data set, suggest that each molecular type can be distinguished morphologically and can be regarded as a separate species. Primary types of A. tepida and topotypes of A. beccarii are shown to be morphologically separate from any of the molecular types so far recognised. We are aware of at least 9 more distinctive morphotypes that have not yet been sequenced, and thus we infer that the total number of genetically distinct and morphologically separable living species of Ammonia worldwide is likely to exceed 25–30. At this stage not all molecular types can be unequivocally assigned to formally described species. Several genetically-based species can be distinguished by the presence of one distinct character, but most are discriminated on the basis of a combination of many different characters. Morphological characters (e.g. test shape, chamber shape, porosity, prolocular diameter, folium shape, radial furrow length, umbilical diameter) are shown to be slightly more valuable in separating the molecular types than surficial ornament (beads, pustules, bosses, secondary calcite). One highly distinctive group (2–3 species – beccarii, batava, ?inflata) is readily discriminated on the basis of its large test size, strongly beaded and grooved ornament, and the presence of fissures along the sutures on the spiral side. The results of this study imply that the widespread practice of recognising only one, two or three species of Recent Ammonia worldwide should be abandoned. The most commonly used name, Ammonia beccarii, should be restricted to a large, compressed, highly ornamented species, so far not recognised beyond its type locality in the Adriatic Sea. Other commonly used names, such as A. parkinsoniana and A. tepida, apply to species with far more restricted distributions than the literature would suggest.

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