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Hayward, B.W.; Holzmann, M.; Pawlowski, J.; Parker, J.H.; Kaushik, T.; Toyofuku, M.S.; Tsuchiya, M. (2021). Molecular and morphological taxonomy of living Ammonia and related taxa (Foraminifera) and their biogeography. Micropaleontology. 67: 109-313.
405477
Hayward, B.W.; Holzmann, M.; Pawlowski, J.; Parker, J.H.; Kaushik, T.; Toyofuku, M.S.; Tsuchiya, M.
2021
Molecular and morphological taxonomy of living Ammonia and related taxa (Foraminifera) and their biogeography
Micropaleontology
67: 109-313.
Publication
Available for editors  PDF available [request]
Globally, one of the two most common shallow-marine and estuarine foraminiferal genera is Ammonia. Over the past 50 years, the majority of workers have identified specimens in this genus as belonging to just 1–3 cosmopolitan species – A. beccarii, A. tepida and A. parkinsoniana. This has been partly because of the problems of discriminating the Ammonia species based entirely on shell morphology and partly because of a 1974 laboratory study that claimed to have shown that all morphologies were merely ecophenotypic variants of one species – a conclusion that molecular studies have proven to be unequivocally wrong. In this study we recognize, describe and figure sixty-seven living species and infraspecies of Recent Ammonia and two closely-related genera (Acarotrochus, Pseudoeponides) from around the world and summarize their ecological and biogeographic distribution. Twenty-six species and two subspecies are recognized by DNA sequencing and shown to be morphologically distinguishable. A further 39 morphospecies and one subspecies, that have not yet been sequenced, have sufficiently distinct morphology to be recognized. Canonical variates analysis using 42 measured or assessed morphological characters shows that the majority of these species can be readily discriminated by their test morphologies, although a few of the less-ornamented molecular species are verging on being pseudocryptic. Molecular sequencing of the type species of Challengerella (C. bradyi) and Helenina (H. anderseni) places them within our Ammonia clade. Here we continue to recognize the morphologically highly distinct genera Pseudoeponides (subjective senior synonym of Helenina) and allied Acarotrochus. Twenty new species or subspecies are described (molecular T types in brackets): Ammonia abramovichae (T8), A. akitaae, A. aoteana australiensis (T5A), A. arabica (T26), A. ariakensis quiltyi, A. buzasi (T11), A. fajemilai, A. goldsteinae, A. goodayi, A. haigi (T25), A. hattai, A. jorisseni (T23), A. justinparkeri, A. kitazatoi (T10), A. morleyae (T12), A. shchedrinae, A. turgida almogilabinae (T22M), Acarotrochus lippsi, Pseudoeponides hottingeri and P. dubuissoni. Aneotype is designated for A. veneta (Schultze 1854) (T1).We recognize 67 Ammonia and related taxa in this study but speculate that there may be 30 or more additional living species that we are not yet confident to discriminate without molecular sequencing. Ammonia species live in most parts of the world between 62 °N (Faeroe Islands) and 55 °S (Strait of Magellan), where seasonal sea-surface temperatures are 4–10° C and above. One estuarine species (A. veneta, T1) is cosmopolitan, euryhaline and eurythermic. Several species are widespread in one or two ocean regions (e.g., Atlantic and Mediterranean; South Pacific), whereas the majority are endemic to smaller areas (e.g., eastern Mediterranean; Caribbean-Gulf of Mexico). Eleven biogeographic “provinces” are recognized by cluster analysis of presence/absence records with the highest diversities in the Australian and northwest Pacific provinces with 18 and 19 species each). Levels of endemism in our “provinces” range between 0 (temperate Atlantic) and 44% (Australian).
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Acarotrochus Kawagata, 2005 (taxonomy source)
Acarotrochus lippsi Hayward & Holzmann, 2021 in Hayward et al., 2021 (original description)
Acarotrochus lobulatus Kawagata, 2005 (taxonomy source)
Ammonia abramovichae Hayward & Holzmann, 2021 in Hayward et al., 2021 (original description)
Ammonia agoiensis Yanko, 1990 accepted as Ammonia neobeccarii Shchedrina & Mayer, 1975 (source of synonymy)
Ammonia akitaae Hayward, Frenzel & Holzmann, 2021 in Hayward et al., 2021 (original description)
Ammonia angulata (Kuwano, 1950) accepted as Ammonia ketienziensis (Ishizaki, 1943) (source of synonymy)
Ammonia aoteana subsp. australiensis Hayward & Holzmann, 2021 (original description)
Ammonia arabica Kaushik, 2021 in Hayward et al., 2021 (original description)
Ammonia ariakensis subsp. quiltyi Hayward & Holzmann, 2021 in Hayward et al., 2021 (original description)
Ammonia asanoi (Uchio, 1951) (taxonomy source)
Ammonia astera Shchedrina, 1984 (taxonomy source)
Ammonia avalonensis (Natland, 1950) (taxonomy source)
Ammonia batava subsp. compacta (Hofker, 1964) accepted as Ammonia batava (Hofker, 1951) (source of synonymy)
Ammonia bulloides (d'Orbigny in Fornasini, 1906) (taxonomy source)
Ammonia buzasi Hayward & Holzmann, 2021 in Hayward et al., 2021 (original description)
Ammonia catesbyana (d'Orbigny, 1839) (taxonomy source)
Ammonia caucasica Yanko, 1990 accepted as Ammonia karadagica (Dolgopolskaya & Pauli, 1931) (source of synonymy)
Ammonia crebra Shchedrina, 1984 accepted as Ammonia confertitesta Zheng, 1978 (source of synonymy)
Ammonia cristobalensis McCulloch, 1981 (taxonomy source)
Ammonia faceta He, Hu & Wang, 1965 † accepted as Pseudoeponides japonicus Uchio in Kawai et al., 1950 † (additional source)
Ammonia fajemilai Hayward & Holzmann, 2021 in Hayward et al., 2021 (original description)
Ammonia flevensis (Hofker, 1930) (taxonomy source)
Ammonia formosa Shchedrina, 1984 accepted as Ammonia confertitesta Zheng, 1978 (source of synonymy)
Ammonia gaudichaudi (d'Orbigny in Fornasini, 1906) (taxonomy source)
Ammonia goldsteinae Hayward & Holzmann, 2021 in Hayward et al., 2021 (original description)
Ammonia goodayi Hayward & Holzmann, 2021 in Hayward et al., 2021 (original description)
Ammonia granuloumbilica Zheng, 1978 (taxonomy source)
Ammonia haigi Hayward & Holzmann, 2021 in Hayward et al., 2021 (original description)
Ammonia hattai Hayward & Holzmann, 2021 in Hayward et al., 2021 (original description)
Ammonia heterogeneus (Shchedrina, 1984) (additional source)
Ammonia jorisseni Hayward, Holzmann & Renjaan, 2021 in Hayward et al., 2021 (original description)
Ammonia juliae McCulloch, 1981 accepted as Ammonia turgida (Hofker, 1951) (source of synonymy)
Ammonia justinparkeri Hayward & Holzmann, 2021 in Hayward et al., 2021 (original description)
Ammonia kitazatoi Hayward & Holzmann, 2021 in Hayward et al., 2021 (original description)
Ammonia lalibertadensis McCulloch, 1977 accepted as Ammonia decorata Golik, 1977 (source of synonymy)
Ammonia langeri Hayward & Holzmann, 2019 accepted as Pseudoeponides langeri (Hayward & Holzmann, 2019) (additional source)
Ammonia limbatobeccarii (McLean, 1956) † accepted as Ammonia sobrina (Shupack, 1934) (additional source)
Ammonia limnetes (Todd & Brönnimann, 1957) accepted as Ammonia advena (Cushman, 1922) (source of synonymy)
Ammonia morleyae Hayward & Holzmann, 2021 in Hayward et al., 2021 (original description)
Ammonia multicella Zheng, 1978 (taxonomy source)
Ammonia nantongensis He, Hu & Wang, 1965 (taxonomy source)
Ammonia neobeccarii subsp. huanhajensis Shchedrina & Mayer, 1975 (source of synonymy)
Ammonia novoeuxinica Yanko in Yanko & Troitskaya, 1987 (taxonomy source)
Ammonia pauciloculata subsp. major He, Hu & Wang, 1965 † (taxonomy source)
Ammonia pauperata (Parr, 1941) (taxonomy source)
Ammonia penangensis McCulloch, 1981 (additional source)
Ammonia pseudotepida (Ujiie, 1992) accepted as Ammonia paucipora Zheng, 1979 (source of synonymy)
Ammonia rupta Lei & Li, 2016 accepted as Ammonia ketienziensis (Ishizaki, 1943) (source of synonymy)
Ammonia shchedrinae Hayward & Holzmann, 2021 in Hayward et al., 2021 (original description)
Ammonia takanabensis (Ishizaki, 1948) accepted as Ammonia ketienziensis (Ishizaki, 1943) (source of synonymy)
Ammonia tepida subsp. bombayensis McCulloch, 1981 (taxonomy source)
Ammonia tepida subsp. cochensis McCulloch, 1981 (taxonomy source)
Ammonia tepida subsp. nitae McCulloch, 1981 accepted as Ammonia aberdoveyensis Haynes, 1973 (source of synonymy)
Ammonia turgida subsp. almogilabinae Hayward & Holzmann, 2021 in Hayward et al., 2021 (original description)
Ammonia umbilicata (d'Orbigny in Fornasini, 1902) (taxonomy source)
Challengerella Billman, Hottinger & Oesterle, 1980 accepted as Ammonia Brünnich, 1771 (source of synonymy)
Cyclohelenina formosaensis Chien, Chang & Yang, 2020 accepted as Pseudoeponides liujunensis (Sun & Zhang in Gou & Tang, 2000) (source of synonymy)
Helenina anderseni (Warren, 1957) accepted as Pseudoeponides anderseni Warren, 1957 (source of synonymy)
Helenina davescottensis van Hengstum & Bernhard, 2016 accepted as Pseudoeponides davescottensis (van Hengstum & Bernhard, 2016) (additional source)
Helenina liujunensis Sun & Zhang in Gou & Tang, 2000 accepted as Pseudoeponides liujunensis (Sun & Zhang in Gou & Tang, 2000) (additional source)
Heleninidae Loeblich & Tappan, 1987 accepted as Ammoniidae Saidova, 1981 (source of synonymy)
Pararotalia capetownensis (McCulloch, 1977) accepted as Pararotalia armata (d'Orbigny in Parker, Jones & Brady, 1865) (source of synonymy)
Pseudoeponides Uchio in Kawai et al., 1950 (additional source)
Pseudoeponides anderseni Warren, 1957 (additional source)
Pseudoeponides angulatus Shchedrina, 1984 accepted as Ammonia veneta (Schultze, 1854) (source of synonymy)
Pseudoeponides compressum Zheng, 1978 accepted as Pseudoeponides anderseni Warren, 1957 (source of synonymy)
Pseudoeponides dubuissoni Baccaert, 2021 in Hayward et al., 2021 (original description)
Pseudoeponides falsobeccarii Rouvillois, 1974 accepted as Ammonia falsobeccarii (Rouvillois, 1974) (source of synonymy)
Pseudoeponides heterogeneus Shchedrina, 1984 accepted as Ammonia heterogeneus (Shchedrina, 1984) (additional source)
Pseudoeponides hottingeri Hayward & Holzmann, 2021 in Hayward et al., 2021 (original description)
Pseudoeponides japonicus Uchio in Kawai et al., 1950 † (additional source)
Pseudoeponides ketienziensis (Ishizaki, 1943) accepted as Ammonia ketienziensis (Ishizaki, 1943) (additional source)
Pseudoeponides langeri (Hayward & Holzmann, 2019) (new combination reference)
Pseudoeponides liujunensis (Sun & Zhang in Gou & Tang, 2000) (additional source)
Pseudoeponides whittakeri Bhatia, 1985 accepted as Pseudoeponides anderseni Warren, 1957 (additional source)
Pseudohelenina Collins, 1974 accepted as Ammonia Brünnich, 1771 (additional source)
Pseudohelenina collinsi (Parr, 1932) accepted as Ammonia collinsi (Parr, 1932) (additional source)
 Neotype

Neotype proposed from Lagoon of Venice by Hayward et al. (2012) = T1 phylotype.  [details]

 Taxonomic remark

Junior synonym in opinion of Hofker (1958) and Schweizer et al. (2011) but discounted by Hayward et al. (2021) [details]

 Taxonomic remark

When Pseudoeponides was considered to be a junior synonym of Ammonia, then this species became a secondary homonym ... [details]