Developayella elegans, from Tong (1995).

Belongs within: Diaphoretickes.
Contains: Bigyra, Pseudofungi, Diatomista, Chrysista, Alveolata, Cercozoa, Endomyxa, Radiozoa, Foraminifera.

The Sar clade is a group of protists united by molecular data, named for the initials of the major subclades Stramenopiles, Alveolata and Rhizaria. It has been defined as the least inclusive clade containing Bigelowiella natans, Tetrahymena thermophila and Thalassiosira pseudonana but not Homo sapiens, Dictyostelium discoideum, Arabidopsis thaliana, Euglena gracilis or Emiliania huxleyi (Adl et al. 2019).

The Ochrophyta includes the photosynthetic heterokonts (golden and brown algae, etc.) and their secondarily non-photosynthetic deerivatives. They possess either a single ciliary transition region helix (Phaeista) or lack this helix entirely (Khakista) (Cavalier-Smith & Chao 2006).

The riddle of the sieve
Published 29 April 2024

In the cosmos of radiolarian systematics, few names loom larger than that of Ernst Heinrich Philipp August Haeckel. Haeckel named somewhere in the area of 5000 species of Radiolaria, many of them in his monumental review of specimens collected during the Challenger expedition (Haeckel 1887). He also established a classificatory system that would be emulated by workers for generations afterwards. And yet, these days Haeckel’s legacy has become much diminished. His broader system has been abandoned as hopelessly artificial, and only a few percent of the species names he established remain in common use (Lazarus 2014). As an example of the issues with Haeckel’s work that would lead to such a volte-face, we might consider Sethomelissa.

Haeckel’s (1887) figure of Lithomelissa (Sethomelissa) decacantha.

The name Sethomelissa was first employed by Haeckel in 1882, in a provisional layout of his classificatory system. At this time, it was a genus name only without any associated species names (this would not be acceptable according to modern nomenclatural practice, but was considered more forgivable at the time). The name of the genus is derived from the Greek words for ‘sieve’ and ‘bee’, and presumably refers to the radiolarian’s resemblance to a bee-hive. Actual species would not be published for Sethomelissa until the 1887 monograph but in a manner that can only be described as utterly baffling. In separate places within the monograph, Sethomelissa appears as a distinct genus, including a single species S. hymenoptera, and as a subgenus of the genus Lithomelissa, including species L. corythium and L. decacantha. In both places, Haeckel attributes the name to the 1882 publication (he also cites ‘Sethomelissa decacantha’ from that source, but that appears to have been an error). Both taxa have the thorax simply latticed, with three ribs in the thorax wall that are prolonged into wings, and three or more horns on the cephalis. However, Haeckel (1887) places the two taxa in separate subfamilies, supposedly distinguished by the thoracic opening: Lithomelissa sits in the subfamily Sethopilida, in which the terminal mouth of the thorax is a ‘simple wide opening’, whereas Sethomelissa the genus is placed in the Sethoperida, with the thorax ‘closed by a lattice-plate’. Just to confuse matters even further, the volume of plates for Haeckel (1887) provides figures for two ‘Sethomelissa’ species, ‘S. decacantha’ and ‘S. bütschlii’, though the latter of these is attributed in the text volume to a different subgenus of Lithomelissa.

Campbell (1951) would later attempt to correct Haeckel’s error by proposing the replacement name Corythomelissa for the subgenus of Lithomelissa. I am not certain that his method of doing so was entirely correct: he seems to have assumed that the separate genus Sethomelissa corresponded to the 1881 taxon, making the subgenus in Lithomelissa a later homonym, but without a formally designated type species I feel it could have gone either way. Nevertheless, there would be no reason other than shear perversity not to follow Campbell’s lead. Species attributed to Sethomelissa following Haeckel (1887) seem to have only been described in the context of the Lithomelissa subgenus, so recognition of Corythomelissa leaves Sethomelissa with the single species S. hymenoptera. Unfortunately, this species was not illustrated by Haeckel (1887) and seems to have never been identified since.

As noted above, modern authors now regard Haeckel’s (1887) classification as very artificial. Haeckel separated various high-level taxa on the basis of features that might be variable even within a single species (Lazarus 2014). The described similarity of Sethomelissa and Corythomelissa in features other than the thoracic aperture certainly raises questions about their distinction. After all, the presence or absence of a lattice over the aperture could easily relate to stages in the skeleton’s development, or indeed to the state of a specimen’s preservation. Haeckel’s reputation in the history of biological studies is a mixed one, and he has not been immune to questions about the validity of his observations. Unfortunately, most of the specimens that Haeckel used in his radiolarian descriptions have not been preserved to the present, whether because Haeckel himself never kept them or because they were subsequently lost. A recent Japanese research programme that examined Haeckel’s surviving specimens in the Museum für Naturkunde, as well as raw sediment samples from the Challenger expedition preserved in the British Museum of Natural History, was unable to identify material of any of the numerous species for which Haeckel did not supply illustrations. This lead Suzuki et al. (2021) to make the bold proposal that all such taxa should be regarded as nomina dubia and expunged from the record, never to be spoken of again. Among the ranks of these questionable casualties would be Sethomelissa hymenoptera, causing its genus to be forever more cast into the outer darkness, a fading echo in the void.

Systematics of Sar
<==Sar [Amphitrematides, Diplophryacea, Harosa]AB19
    |    |--Heterokonta [Actinodinea, Heterokontophyta, Labyrinthulidea, Stramenipila, Stramenopiles]C-SCL15
    |    |    |--BigyraC-SC06
    |    |    `--GyristaC-S18
    |    |         |--Bigyromonada [Bigyromonadea]C-S18
    |    |         |    |--Pirsonia [Pirsonea, Pirsoniaceae, Pirsoniales, Pirsoniida]C-S18
    |    |         |    |    `--P. diademataC-SC06
    |    |         |    `--Developayellidae [Developayellida, Developea]C-S18
    |    |         |         |--DevelorapaxC-S18
    |    |         |         `--Developayella Tong 1995C-S18, KC01 [Developayellaceae, Developayellales]
    |    |         |              `--D. elegansC-SC06
    |    |         |--PseudofungiC-SC06
    |    |         `--Ochrophyta (see below for synonymy)AB19
    |    |              |--DiatomistaC-S18
    |    |              `--ChrysistaC-S18
    |    `--AlveolataAS12
    `--Rhizaria (see below for synonymy)AS12
         |  i. s.: Amphitholus Haeckel 1887AA05
         |         Wetzelodinium Deflandre 1936FT93
         |         ActinolophusAS12
         |         Acaenispongus Kozur & Mostler 1981AA05
         |         Acanthocorallium Haeckel 1887 (n. o.)AA05
         |         Acanthocorythium Haeckel 1887 (n. o.)AA05
         |         Acanthopyle Vinassa de Regny 1898AA05
         |         Acanthosphaera Ehrenberg 1858AA05
         |         Acrocorona Haeckel 1881AA05
         |         Actinopyramis Haeckel 1887 (n. o.)AA05
         |         Actinosphaera Hollande & Enjument 1960AA05
         |         Actissa Haeckel 1887AA05
         |         Acutacapsula Tikhomirova 1983AA05
         |         Adelocyrtis Pantanelli 1880AA05
         |         Afens Riedel & Sanfilippo 1974AA05
         |         Alatipicapora Tekin 1999AA05
         |         Albatrossidium Sanfilippo & Riedel 1992AA05
         |         Amphiactura Haeckel 1881AA05
         |         Amphicarydiscus Lipman 1972AA05
         |         Amphicraspedum Haeckel 1881AA05
         |         Amphicyclia Haeckel 1881AA05
         |         Amphipyle Haeckel 1881AA05
         |         Amphipylonium Haeckel 1881AA05
         |         Amphipylura Haeckel 1887AA05
         |         Amphisphaera Haeckel 1881AA05
         |         Amphistylus Haeckel 1882AA05
         |         Amphitholonium Haeckel 1887AA05
         |         Amphymenium Haeckel 1881AA05
         |         ‘Andromeda’ Baumgartner 1980 non Gistel 1834AA05
         |         Angulocircus Lahm 1984AA05
         |         Anisicyrtis Kozur & Mostler 1981AA05
         |         Annikaella De Wever 1988AA05
         |         Spongocyclia Haeckel 1862 [incl. Annulatospira Clark & Campbell 1945, Spongospira Stohr 1880]AA05
         |         Annulopoulpus Kozur & Mostler 1981AA05
         |         Annulosaturnalis Dosztaly 1991AA05
         |         Anthocyrtidium Haeckel 1881 [incl. Anthocyrtissa Haeckel 1887]AA05
         |         Arachnopegma Haeckel 1881AA05
         |         Arachnopila Haeckel 1887AA05
         |         Arcanicapsa Takemura 1986AA05
         |         Archicapsa Haeckel 1881AA05
         |         Archicorys Haeckel 1881AA05
         |         Archipera Haeckel 1881 (n. o.)AA05
         |         Archiphaena Haeckel 1881AA05
         |         Archiphormis Haeckel 1881AA05
         |         Archistephus Haeckel 1887 (n. o.)AA05
         |         Ares De Wever 1982AA05
         |         Artophaena Haeckel 1881AA05
         |         Astractura Haeckel 1881AA05
         |         Astrocapsa Haeckel 1887AA05
         |         Astrocentrus Kozur & Mostler 1979AA05
         |         Astrocyclia Haeckel 1881AA05
         |         Astrosestrum Haeckel 1881AA05
         |         Atactodiscus Haeckel 1882AA05
         |         Atichocapsa Campbell & Clark 1944AA05
         |         Aulophacus Pessagno 1962AA05
         |         Axellipsis Haeckel 1887AA05
         |         Axodiscus Haeckel 1887AA05
         |         Bacus Wu 1986AA05
         |         Bagotella Tikhomirova 1987AA05
         |         Belonostaurus Haeckel 1887AA05
         |         Bipedis De Wever 1982AA05
         |         Borgertella Dumitrica 1973AA05
         |         Botryocephalina Petrushevskaya 1965AA05
         |         Braginastrum Tekin 1999AA05
         |         Braginella Sugiyama 1997AA05
         |         Bullacapsula Tikhomirova 1983AA05
         |         Byssulentactinia Nazarov 1988AA05
         |         Calocapsa Haeckel 1887 (n. o.)AA05
         |         Calompterium Clark & Campbell 1942 (n. o.)AA05
         |         Calpophaena Haeckel 1881AA05
         |         Caltrop Carter in Carter, Cameron & Smith 1988AA05
         |         Canarachnium Caulet 1971AA05
         |         Canesium Blome 1984AA05
         |         Cannocapsa Haeckel 1887AA05
         |         Cantharospyris Haeckel 1887AA05
         |         Carinaheliosoma Kozur & Mostler 1981AA05
         |         Caryomma Haeckel 1887AA05
         |         Caryosphaera Haeckel 1881AA05
         |         Caryostylus Haeckel 1881AA05
         |         Castrum Blome 1984AA05
         |         Catoma Blome 1983AA05
         |         Caucasicus Mamedov 1984AA05
         |         Cavabracchia Kito & De Wever 1992AA05
         |         Cenellipsis Haeckel 1882AA05
         |         Cenocapsa Haeckel 1887AA05
         |         Cenolarcopyle Tan Sin Hok 1931AA05
         |         Cenolarcus Haeckel 1887AA05
         |         Centracontarium Popofsky 1912AA05
         |         Centracontium Popofsky 1912AA05
         |         Cephalospyris Haeckel 1881AA05
         |         Ceratobotrys Nishimura 1990AA05
         |         Chaenicosphaera Haeckel 1887AA05
         |         Choenicosphaera Haeckel 1887AA05
         |         Cladarachnium Haeckel 1881AA05
         |         Clathrocyrcus Haeckel 1881AA05
         |         Clathrosphaera Haeckel 1881AA05
         |         Clistophaena Haeckel 1881AA05
         |         Coccolarcus Haeckel 1887AA05
         |         Conoactinomma Gorbunov 1979AA05
         |         Conocaryomma Haeckel 1887AA05
         |         Cornucapsula Tikhomirova 1983AA05
         |         Cornutellium Haeckel 1881AA05
         |         Coronidium Haeckel 1881AA05
         |         Cortiniscus Haeckel 1887 (n. o.)AA05
         |         Coscinomma Haeckel 1887AA05
         |         Cristallosphaera Popofsky 1912AA05
         |         Cromyatractus Haeckel 1887AA05
         |         Cromydruppocarpus Campbell & Clark 1944AA05
         |         Cromyechinus Haeckel 1881AA05
         |         Cromyocarpus Haeckel 1887AA05
         |         Cromyodrimus Haeckel 1881AA05
         |         Cromyostaurolonche Clark & Campbell 1944AA05
         |         Cromyostaurus Haeckel 1881AA05
         |         Cromyostylus Haeckel 1881AA05
         |         Crubus Yeh 1987AA05
         |         Crucidiscus Haeckel 1887AA05
         |         Cryptocarpium Sanfilippo & Riedel 1992AA05
         |         Cryptoprora Ehrenberg 1860AA05
         |         Cubaxonium Haeckel 1887AA05
         |         Cubotholonium Haeckel 1887AA05
         |         Cubotholus Haeckel 1887AA05
         |         Cyphinidium Haeckel 1887AA05
         |         Cyphocolpus Haeckel 1887AA05
         |         Cyphonium Haeckel 1887AA05
         |         Cyrcotympanum Haeckel 1887AA05
         |         Cyrthentactinia Foreman 1963AA05
         |         Cyrtopera Haeckel 1881AA05
         |         Deflandrastrum De Wever 1983AA05
         |         Dendrocyrcus Haeckel 1881AA05
         |         Desmartus Haeckel 1887AA05
         |         Desmocampe Haeckel 1887AA05
         |         Dicapnuchosphaera Tekin 1999AA05
         |         Distympanium Haeckel 1887AA05
         |         Dictyastrum Ehrenberg 1860 [incl. Dictyastrella Haeckel 1887, Dictyastromma Haeckel 1887]AA05
         |         Dictyodedalus O’Dogherty 1994AA05
         |         Dictyophimium Haeckel 1887AA05
         |         Dictyospyrella Haeckel 1887AA05
         |         Didactylum Baumgartner 1980AA05
         |         Diplactura Haeckel 1881AA05
         |         Diplellipsis Popofsky 1908AA05
         |         Diploplegma Hinde 1890AA05
         |         Diplostrobus Squinabol 1903AA05
         |         Dipospyris Haeckel 1881AA05
         |         Discopyle Haeckel 1887AA05
         |         Discozonium Haeckel 1887AA05
         |         Disolenia Ehrenberg 1860AA05
         |         Dispongia Popofsky 1912AA05
         |         Distriactis Haeckel 1887AA05
         |         Distylocapsa Squinabol 1904AA05
         |         Dizonites Haeckel 1887AA05
         |         Dizonium Haeckel 1881AA05
         |         Doracantha Haeckel 1881AA05
         |         Doryconchidium Vinassa de Regny 1898AA05
         |         Dorydictium Hinde 1890AA05
         |         Dorydiscus Carnavale 1908AA05
         |         Dorydruppa Vinassa de Regny 1900AA05
         |         Dorylonchidium Vinassa de Regny 1898AA05
         |         Doryphacus Carnavale 1908AA05
         |         Doryplegma Hinde 1890AA05
         |         Dorysphaera Hinde 1890 [incl. Monostylus Cayeaux 1897]AA05
         |         Dreyerella Kozur & Mostler 1979AA05
         |         Dreyeropyle Kozur & Mostler 1979AA05
         |         Drulanta Yeh 1987AA05
         |         Druppalonche Hinde 1899AA05
         |         Druppastylus Cayeux 1897AA05
         |         Druppatractylis Haeckel 1887AA05
         |         Druppocarpus Haeckel 1887AA05
         |         Druppula Haeckel 1887AA05
         |         Dryomyomma Jörgensen 1905AA05
         |         Dumitricaella De Wever 1982AA05
         |         Dumitricasphaera Kozur & Mostler 1979AA05
         |         Dyostephus Haeckel 1881 (n. o.)AA05
         |         Dystympanum Haeckel 1887AA05
         |         Eastonerius Empson-Morin 1981AA05
         |         Echinactura Haeckel 1887AA05
         |         Echinomma Haeckel 1881AA05
         |         Elatomma Haeckel 1887AA05
         |         Ellipsostygma Hinde 1899AA05
         |         Ellipsostylus Haeckel 1887AA05
         |         Ellipsoxiphus Dunikowski 1882AA05
         |         Elodium Carter in Carter et al. 1988AA05
         |         Enneaplagia Haeckel 1881 (n. o.)AA05
         |         Enoplocampe Sugiyama 1997AA05
         |         Eostichomitra Empson-Morin 1981AA05
         |         Eucecryphalium Haeckel 1887AA05
         |         Eutympanium Haeckel 1881AA05
         |         Excingula Kozlova 1994AA05
         |         Falsocromyodrymus O’Dogherty 1994AA05
         |         Fontinella Carter 1993AA05
         |         Foremaniella Deflandre 1972AA05
         |         Foremina Empson-Morin 1981AA05
         |         Gigi De Wever 1982AA05
         |         Grandicapsula Kazintsova 1985AA05
         |         Guangxitrisphaera Wang 1993AA05
         |         Guexella Baumgartner 1984AA05
         |         GuttacapsaAA05
         |         Haeckelella Chabakov 1937AA05
         |         Hagiastromma Haeckel 1887 [incl. Hagiastrella Haeckel 1887]AA05
         |         Halidictyum Ichikawa 1950AA05
         |         Haliodictya Hojnos 1916AA05
         |         Haliodictyum Ichikawa 1950AA05
         |         Haliphormis Ehrenberg 1847AA05
         |         Helenofore Nazarov & Ormiston 1983AA05
         |         ‘Heliaster’ Hollande & Enjument 1960 non Gray 1840AA05
         |         Heliosestarium Campbell & Clark 1944AA05
         |         Heliosoma Haeckel 1881AA05
         |         Helminentactinia Li 1994AA05
         |         Hergelia Nazarov & Ormiston 1985AA05
         |         Heterosestrum Clark & Campbell 1945AA05
         |         Hexacaryum Haeckel 1881AA05
         |         Hexacontarium Haeckel 1887AA05
         |         Hexacorys Haeckel 1887 (n. o.)AA05
         |         Hexacromyum Haeckel 1881AA05
         |         Hexactura Haeckel 1882 [incl. Hexalastromma Haeckel 1887, Hexalastrum Haeckel 1881 (nom. inv.)]AA05
         |         Hexacyclia Totchilina 1970AA05
         |         Hexadendron Haeckel 1881AA05
         |         Hexadoras Haeckel 1881AA05
         |         Hexadoridium Haeckel 1881AA05
         |         Hexadorium Campbell & Clark 1944AA05
         |         Hexalatractus Haeckel 1887AA05
         |         Hexaloncharium Haeckel 1887AA05
         |         Hexalonchidium Haeckel 1881AA05
         |         Hexaplagia Haeckel 1881 (n. o.)AA05
         |         Hexaplecta Haeckel 1887 (n. o.)AA05
         |         Hexaplegma Haeckel 1881 (n. o.)AA05
         |         Hexaporobranchia Kozur & Mostler 1979AA05
         |         Hexapylomella Kozur & Mostler 1979AA05
         |         Hexaspongonium Haeckel 1881AA05
         |         Hexaspyris Haeckel 1887AA05
         |         Hexastylarium Haeckel 1887AA05
         |         Hexastylidium Haeckel 1881AA05
         |         Hexastylurus Haeckel 1881AA05
         |         Hinedorcus Dumitrica, Kozur & Mostler 1980AA05
         |         Holdsworthium Furutani 1990AA05
         |         Homeoarchicorys Empson-Morin 1981AA05
         |         Homeoparonaella Baumgartner 1980AA05
         |         Hymenactura Haeckel 1881AA05
         |         Hymeniastrum Ehrenberg 1847AA05
         |         Intocapsa Tikhomirova 1987AA05
         |         Japonisaturnalis Kozur & Mostler 1972AA05
         |         Japonocampe Kozur 1984AA05
         |         Kafanella Tikhomirova 1987AA05
         |         Kahlerosphaera Kozur & Mostler 1979AA05
         |         Kashiwara Blome 1992AA05
         |         Katroma Pessagno & Poisson 1979AA05
         |         Kozurastrum De Wever 1984AA05
         |         Kozuricyrtium Tekin 1999AA05
         |         Krempelinella Empson-Morin 1981AA05
         |         Kuppelella Empson-Morin 1981AA05
         |         Lachnosphaera Haeckel 1881AA05
         |         Lampoxanthium Haeckel 1887AA05
         |         Lamprocyrtis Kling 1973AA05
         |         Laracospongus Haeckel 1887AA05
         |         Larcarium Haeckel 1887AA05
         |         Larcidium Haeckel 1887AA05
         |         Larnacalpis Haeckel 1887AA05
         |         Larnacantha Haeckel 1887AA05
         |         Larnacidium Haeckel 1887AA05
         |         Larnacoma Haeckel 1887AA05
         |         Larnacospongus Haeckel 1887AA05
         |         Larnacostupa Haeckel 1887AA05
         |         Liassosaturnalis Kozur & Mostler 1990AA05
         |         Lichnosphaera Haeckel 1881AA05
         |         Lithapium Haeckel 1887AA05
         |         Lithatractus Haeckel 1887AA05
         |         Lithocarpium Stohr 1880AA05
         |         Lithocorcys Ichikawa 1950 (n. o.)AA05
         |         Lithophyllium Müller 1858AA05
         |         Lupherium Pessagno & Whalen 1987AA05
         |         Lychnodictyum Haeckel 1881AA05
         |         Mesosaturnalis Kozur & Mostler 1981AA05
         |         Middourium Kozlova 1999AA05
         |         ‘Milax’ Bloome 1984 non Gray 1855AA05
         |         Minocapsa Matsuoka 1991AA05
         |         Mitrocalpis Haeckel 1881AA05
         |         Monaxonium Popofsky 1912AA05
         |         ‘Monobrachium’ Kozlova 1999 non Mereschkowsky 1877AA05
         |         Monocapnuchosphaera Tekin 1999AA05
         |         Monosera Takemura & Nakaseko 1986AA05
         |         Monostephus Haeckel 1881AA05
         |         Monotrabs Baumgartner 1984AA05
         |         Monozonites Haeckel 1887AA05
         |         Monozonium Haeckel 1887AA05
         |         Mostlericyrtium Tekin 1999AA05
         |         Muellericyrtium Kozur & Mostler 1981AA05
         |         Multastrum Vishnevskaya 1990AA05
         |         Multimonilis Yeh 1989AA05
         |         Nakasekoellus Kozur 1984 [incl. Xipha Blome 1984]AA05
         |         NatobaAA05
         |         Natraglia Pessagno in Pessagno, Finch & Abbott 1979AA05
         |         Nodocapnuchosphaera Tekin 1999AA05
         |         Noviforemanella Pessagno, Blome & Hull 1993AA05
         |         Novodiacanthocapsa Empson-Morin 1981AA05
         |         Octopylura Haeckel 1887AA05
         |         Odontosphaera Haeckel 1887AA05
         |         Oertlisphaera Kozur & Mostler 1979AA05
         |         Ommatocampe Ehrenberg 1860AA05
         |         Ommatogramma Ehrenberg 1860AA05
         |         Ormistonia Li 1994AA05
         |         Oroscena Haeckel 1887AA05
         |         Otosphaera Haeckel 1887AA05
         |         Ovulopyle Kozur & Mostler 1979AA05
         |         Palaeacanthus Davis 1950AA05
         |         Palaeoeuchitonia Kozur & Mostler 1978AA05
         |         Palaeofripus Goodbody 1986AA05
         |         Palaeopyramidatum Goodbody 1986AA05
         |         Palaeoxyphostylus Won 1983AA05
         |         Paleocenosphaera Nazarov 1973AA05
         |         Paleotripus Goodbody 1986AA05
         |         Paleoxiphosphaera Nazarov 1973AA05
         |         Panarium Haeckel 1881AA05
         |         Papiliocampe Tekin 1999AA05
         |         Parabipedis Yeh & Cheng 1996AA05
         |         Paracenodiscus Krascheninnikov 1960AA05
         |         Paraplafkerium Kozur & Mostler 1981AA05
         |         Pararchocyrtium Deflandre 1972AA05
         |         Parares Takemura 1986AA05
         |         Pararuesticyrtium Kozur & Mock in Kozur & Mostler 1981AA05
         |         Parastephanus Haeckel 1881AA05
         |         Parastomatosphaera Kozur & Mostler 1979AA05
         |         Parentactinia Dumitrica 1978AA05
         |         Parentactinosphaera Kozur & Mostler 1979AA05
         |         Paricrioma Tekin 1999AA05
         |         Parvifusus Takemura 1986AA05
         |         Patyomma Haeckel 1881AA05
         |         Pentactinosphaera Nagata & Nishimura 1983AA05
         |         Pentactura Haeckel 1881AA05
         |         Pentalastrum Haeckel 1881 [incl. Pentalastrella Haeckel 1887, Pentalastromma Haeckel 1887]AA05
         |         Pentalpagia Cachon & Cachon 1969AA05
         |         Pentasphaera Squinabol 1904AA05
         |         Pentatortis Kozur 1988AA05
         |         Pentophiastrum Haeckel 1887AA05
         |         Peridarium Haeckel 1887 (n. o.)AA05
         |         Peripanarium Haeckel 1887AA05
         |         Perispongidium Haeckel 1882AA05
         |         Perizona Haeckel 1881AA05
         |         Perseus Takemura & Nakaseko 1983AA05
         |         Pessagnalus Dumitrica 1982AA05
         |         Phacodiscinus Heackel 1887AA05
         |         Phacopyle Dreyer 1889AA05
         |         Phacotriactis Sutton 1896AA05
         |         Phalangites O’Dogherty 1994AA05
         |         Pharyngosphaera Heackel 1887AA05
         |         Phermostichoartus Campbell 1951AA05
         |         Phizosphaera Haeckel 1881AA05
         |         Phormospyris Haeckel 1881AA05
         |         Phormostichoartus Campbell 1951AA05
         |         Phractacantha Haeckel 1881AA05
         |         Phractaspis Haeckel 1881AA05
         |         Phyletripes Campbell 1951AA05
         |         Physematium Meyen 1834AA05
         |         Picapora Kozur & Mostler 1981AA05
         |         Pipetta Haeckel 1887AA05
         |         Pipettaria Haeckel 1887AA05
         |         Pipettella Haeckel 1887AA05
         |         Pityomma Haeckel 1881AA05
         |         Plafkerium Pessagno 1979AA05
         |         Plagonidium Haeckel 1881 (n. o.)AA05
         |         Plectocoronis Haeckel 1881AA05
         |         Plectoscenium Nishimura 1990AA05
         |         Plegmatium Campbell 1954 (n. o.)AA05
         |         Pobum De Wever 1984AA05
         |         Podocapsa Rüst 1885AA05
         |         Podocoronis Haeckel 1881 [incl. Tetracoronis Haeckel 1881]AA05
         |         ‘Pogonias’ O’Dogherty 1994 nec Lacépède 1802 nec Illiger 1811AA05
         |         Poloctostylus Dumitrica 1994AA05
         |         Polyalacorys Haeckel 1887AA05
         |         Polynura Haeckel 1887AA05
         |         Polyplagia Haeckel 1887 (n. o.)AA05
         |         ‘Polyzonium’ Haeckel 1887 non Brandt 1837AA05
         |         Porocapsa Haeckel 1887AA05
         |         Praeamphibrachidium Kozur & Mostler 1978AA05
         |         Praedruppatractylis Kozur & Mostler 1979AA05
         |         Praeflustrella Kozur & Mostler 1978AA05
         |         Praemesosaturnalis Kozur & Mostler 1981AA05
         |         Praeprotunuma Tekin 1999AA05
         |         Praerhopalastrum Kozur & Mostler 1978AA05
         |         Praesaturnalis Li 1994AA05
         |         Praexiphospira Kozur & Mostler 1978AA05
         |         Prismatium Haeckel 1862AA05
         |         Protosphaera Cayeux 1897AA05
         |         Prunocarpus Haeckel 1887AA05
         |         Prunulum Haeckel 1887AA05
         |         Pseudoacanthocircus Kozur & Mostler 1990AA05
         |         Pseudoacanthosphaera O’Dogherty 1994AA05
         |         Pseudoeucyrtis Pessagno 1977AA05
         |         Pseudogodia Tekin 1999AA05
         |         Pseudospongoprunum Wakamatsu, Sugiyama & Furutani 1990AA05
         |         Pseudostaurolonche Mamedov 1969AA05
         |         Pseudostaurosphaera Krascheninnikov 1960AA05
         |         Pseudostigmosphaera Kozur & Mostler 1979AA05
         |         Pseudotheocampe Empson-Morin 1981AA05
         |         Pterocampe Haeckel 1887AA05
         |         Pteroscenium Haeckel 1887AA05
         |         Pylocapsa Haeckel 1881AA05
         |         Pylosphaera Ehrenberg 1858AA05
         |         Pylospira Haeckel 1887AA05
         |         Pylozonium Haeckel 1887AA05
         |         Quadricaulis Caridroit & De Wever 1986AA05
         |         Quadrigastrum O’Dogherty 1994AA05
         |         Quadrilonche Haeckel 1887AA05
         |         Quarkus Pessagno, Blome & Hull 1993AA05
         |         Quasipetatus Blome 1984AA05
         |         Raoultius De Wever 1982AA05
         |         Rhabdolithis Ehrenberg 1847AA05
         |         Rhaphidococcus Haeckel 1862AA05
         |         Rhodosphaera Haeckel 1881AA05
         |         Rhopalodictyum Ehrenberg 1860 [incl. Triactinosphaera Dunikowski 1882]AA05
         |         Riedelipyle Kozur & Mostler 1979AA05
         |         Stauracontium Haeckel 1882 [incl. Ruestia Vinassa de Regny 1898, Xiphostaurus Vinassa de Regny 1898]AA05
         |         Sarla Pessagno 1979AA05
         |         Saturninus Haeckel 1887AA05
         |         Saturnosphaera Tichomirova 1975AA05
         |         Saturnulus Haeckel 1881AA05
         |         Schaafella Vishnevskaya 1990AA05
         |         Schadelfusslerus Empson-Morin 1981AA05
         |         Schaumellus Empson-Morin 1981AA05
         |         Senelella Tekin 1999AA05
         |         Sethocanium Nischimura 1990AA05
         |         Sethocephalus Haeckel 1887 (n. o.)AA05
         |         Sethoconus Haeckel 1881AA05
         |         Sethodiscus Haeckel 1881AA05
         |         Sethomelisa Haeckel 1881AA05
         |         Sethornithium Haeckel 1881 (n. o.)AA05
         |         Sethosphaera Haeckel 1881AA05
         |         Sethostaurus Haeckel 1881AA05
         |         Soreuma Haeckel 1881 (n. o.)AA05
         |         Sorolarcus Haeckel 1887AA05
         |         Sphaerocapsa Haeckel 1881AA05
         |         Sphaerostylus Haeckel 1881AA05
         |         Spicularina Donofrio 1991AA05
         |         Spinopoulpus Kozur & Mock in Kozur & Mostler 1981AA05
         |         Spinosiocapsa Ozvoldova 1975 (n. o.)AA05
         |         Spiremaria Kozlova 1960AA05
         |         Spiromultitunica Tochilina & Popova 1985AA05
         |         Spironium Haeckel 1887AA05
         |         Spiropyle Tochilina 1985AA05
         |         Spirotunica Tochilina & Popova 1985AA05
         |         Spongasteriscus Haeckel 1882 [incl. Spongastericulus Haeckel 1887]AA05
         |         Spongechinus Haeckel 1881AA05
         |         Spongellipsis Haeckel 1887AA05
         |         Spongiocanium Nishimura 1990AA05
         |         Spongioconcha Mast 1910AA05
         |         Spongiomma Haeckel 1887AA05
         |         Spongiopodium Nishimura 1990AA05
         |         Spongiostoma Carter, Cameron & Smith 1988AA05
         |         Spongoacanthus Squinabol 1903AA05
         |         Spongobrachium Haeckel 1881AA05
         |         Spongocyrtis Dunikowski 1862AA05
         |         Spongodendron Hollande & Enjument 1960AA05
         |         Spongodruppa Haeckel 1887AA05
         |         Spongodrymus Haeckel 1881AA05
         |         Spongolarcus Haeckel 1887AA05
         |         Spongolonchidium Chedia 1959AA05
         |         Spongophacus Haeckel 1881AA05
         |         Spongophortis Haeckel 1881AA05
         |         Spongopila Haeckel 1881AA05
         |         Spongoplegma Haeckel 1881AA05
         |         Spongosaturnalis Campbel & Clark 1944AA05
         |         Spongostaurus Haeckel 1881AA05
         |         Spongostichomitra O’Dogherty 1994AA05
         |         Spongostylidium Haeckel 1881AA05
         |         Spongostylus Haeckel 1881AA05
         |         Spongothamnus Haeckel 1887AA05
         |         Spongotractus Haeckel 1887AA05
         |         Spongoxiphus Haeckel 1887AA05
         |         Stauractura Haeckel 1881AA05
         |         Staurancistra Haeckel 1881AA05
         |         Stauroacanthocircus Kozur & Mostler 1983AA05
         |         Staurocaryum Haeckel 1881AA05
         |         Staurocromyum Haeckel 1881AA05
         |         Staurocyclia Haeckel 1881AA05
         |         Staurodictya Haeckel 1882AA05
         |         Staurodiscus Krascheninnikov 1960AA05
         |         Staurodoras Haeckel 1881AA05
         |         Staurolonche Haeckel 1881AA05
         |         Staurolonchidium Haeckel 1887AA05
         |         Stauromesosaturnalis Kozur & Mostler 1990AA05
         |         Stauroplegma Hinde 1890AA05
         |         Staurosphaera Haeckel 1881AA05
         |         Staurostylus Haeckel 1881AA05
         |         Staurotholonium Haeckel 1887AA05
         |         Staurotholus Haeckel 1887AA05
         |         Stauroxiphos Haeckel 1887AA05
         |         Stephaniscus Haeckel 1887 (n. o.)AA05
         |         Stichocampe Haeckel 1881AA05
         |         Stichoperina Haeckel 1887 (n. o.)AA05
         |         Stichophaenoma Haeckel 1887AA05
         |         Stomatosphaera Dreyer 1889AA05
         |         Streblonia Haeckel 1887AA05
         |         Streblopyle Haeckel 1887AA05
         |         StychopyliumAA05
         |         Stylartus Haeckel 1881AA05
         |         Stylocapsa Principi 1909AA05
         |         Stylocromyum Haeckel 1881AA05
         |         Stylocryptocapsa Tan Sin Hok 1926AA05
         |         Stylocyclia Ehrenberg 1847AA05
         |         Stylodiscus Haeckel 1887AA05
         |         Stylodruppa Kasinzova 1979AA05
         |         Stylosphaerella Haeckel 1887AA05
         |         Stylospongidium Haeckel 1882AA05
         |         Stylostaurus Haeckel 1881AA05
         |         Stylotrochus Haeckel 1862AA05
         |         Stypolarcus Haeckel 1887AA05
         |         Tauridastrum Tekin 1999AA05
         |         Teichertus Hull 1997AA05
         |         Tessarospyris Haeckel 1881AA05
         |         Tetraedrina Haeckel 1881 (n. o.)AA05
         |         Tetrahedrina Haeckel 1881 (n. o.)AA05
         |         Tetrapteroma Haeckel 1881 (n. o.)AA05
         |         Tetrapylissa Haeckel 1887AA05
         |         Tetrapylonium Haeckel 1881AA05
         |         Tetraregnon Ormistone & Lane 1976AA05
         |         Tetrarhabda Haeckel 1881 (n. o.)AA05
         |         Tetrasphaera Popofsky 1912AA05
         |         Tetraspongonium Haeckel 1882AA05
         |         Tetraspyris Haeckel 1881AA05
         |         Thaisphaera Sashida & Igo 1992AA05
         |         Thalassolampe Haeckel 1862AA05
         |         Thalassopila Haeckel 1881AA05
         |         Thalassoplancta Haeckel 1862AA05
         |         Thecacapsula Tikhomirova 1983AA05
         |         Thecoentactinia Nazarov 1975AA05
         |         Thecosphaerella Haeckel 1887AA05
         |         Theocapsa Haeckel 1882AA05
         |         Theocotylissa Foreman 1973AA05
         |         Theodiscus Haeckel 1887AA05
         |         Theosyringium Haeckel 1881AA05
         |         Tholartus Haeckel 1887AA05
         |         Tholodes Haeckel 1887AA05
         |         Tholospira Haeckel 1887AA05
         |         Tholostaurus Haeckel 1887AA05
         |         Tiarospyris Haeckel 1881AA05
         |         Tirodella Kozur & Mostler 1981AA05
         |         Tlecerina Furutani 1983AA05
         |         Triacanthocircus Kozur & Mostler 1990AA05
         |         Triacornicyrtium Tekin 1999AA05
         |         Triadiscus Haeckel 1887AA05
         |         Triaenosphaera Deflandre 1973AA05
         |         Trialatus Yeh 1990AA05
         |         Triarcella Kozur & Mock in Kozur & Mostler 1981AA05
         |         Triassocrucella Kozur 1984AA05
         |         Triborella Dumitrica, Kozur & Mostler 1980AA05
         |         Tricapsula Wu 1986AA05
         |         Tricirconium Haeckel 1887 (n. o.)AA05
         |         Tricolocamptra Haeckel 1887AA05
         |         Tricyclonium Haeckel 1887 (n. o.)AA05
         |         Trigonactura Haeckel 1881AA05
         |         Trigonocyclia Haeckel 1881AA05
         |         Trigonodiscus Neviani 1900AA05
         |         Trilobatum Popofsky 1912AA05
         |         Trilonche Hinde 1899AA05
         |         Trimiduca Kozur & Mostler 1979AA05
         |         Trimulus O’Dogherty 1994AA05
         |         Triodiscus Haeckel 1887AA05
         |         Triolena Haeckel 1887AA05
         |         Triopyle Haeckel 1881AA05
         |         Triostephus Haeckel 1881 (n. o.)AA05
         |         Triplanospongos Sashida & Tonishi 1988AA05
         |         Triplecta Haeckel 1881 (n. o.) [Triplectinae]AA05
         |         Tripocoronis Haeckel 1881 (n. o.)AA05
         |         Tripodictya Haeckel 1881AA05
         |         Tripodiscium Haeckel 1881AA05
         |         Tripodoconus Nishimura 1990AA05
         |         Tripodonium Haeckel 1881 (n. o.)AA05
         |         Tripospyrantha Haeckel 1887 (n. o.)AA05
         |         Tripospyrissa Haeckel 1887 (n. o.)AA05
         |         Tripospyromma Haeckel 1887 (n. o.)AA05
         |         Tripterocalpis Haeckel 1881AA05
         |         Trissopilium Haeckel 1881 (n. o.)AA05
         |         Tristephaniscus Haeckel 1881 (n. o.)AA05
         |         Tristylospyrium Haeckel 1887 (n. o.)AA05
         |         Trizonaris Haeckel 1887AA05
         |         Turbocapsa O’Dogherty 1994AA05
         |         Udan Renz 1976AA05
         |         Veghia Kozur & Mostler 1981AA05
         |         Verticillata Popofsky 1913AA05
         |         Vinassapongus Kozur & Mostler 1979AA05
         |         Vistularia Gorka 1989AA05
         |         Wildeus Empson-Morin 1982AA05
         |         Wuranella Kozur & Mostler 1981AA05
         |         Xastrum Neviani 1900AA05
         |         Xanthiosphaera Haeckel 1881AA05
         |         Xiphodictya Haeckel 1881AA05
         |         Xiphoptera Haeckel 1881AA05
         |         Xiphosphaera Haeckel 1881AA05
         |         Xiphospira Haeckel 1887AA05
         |         Yaosaturnalis Kozur & Mostler 1983AA05
         |         Yeharaia Nakaseko & Nishimura 1979AA05
         |         Zhamojdasphaera Kozur & Mostler 1979AA05
         |         Zigocircus Butschli 1882AA05
         |         Zigostephaniscus Haeckel 1881 (n. o.)AA05
         |         Zonarium Haeckel 1887AA05
         |         Zonidium Haeckel 1887AA05
         |         Zoniscus Haeckel 1887AA05
         |         Zonodiscus Haeckel 1887AA05
         |         Zygacantha Müller 1858AA05
         |         Zygartus Haeckel 1881AA05
         |         Zygocampe Haeckel 1887AA05
         |         Zygospyris Haeckel 1881AA05
         |         Zygostaurus Haeckel 1887AA05
         |         Zygostephanium Haeckel 1881AA05
         |         Aquavolon Tikhonenkov, Mylnikov & Bass 2018 [Aquavolonida]AB19

Ochrophyta [Chrysomonada, Chrysomonadida, Chrysophycophyta, Chrysophyta, Dictyochiae, Marista, Ochrista, Ochrophytina]AB19

Rhizaria [Actinopoda, Granuloreticulosa, Granuloreticulosia, Hydromyxales, Leucodictyida, Radiolaria, Reticularea, Reticularia]AS12

*Type species of generic name indicated


[AB19] Adl, S. M., D. Bass, C. E. Lane, J. Lukeš, C. L. Schoch, A. Smirnov, S. Agatha, C. Berney, M. W. Brown, F. Burki, P. Cárdenas, I. Čepička, L. Chistyakova, J. del Campo, M. Dunthorn, B. Edvardsen, Y. Eglit, L. Guillou, V. Hampl, A. A. Heiss, M. Hoppenrath, T. Y. James, A. Karnkowska, S. Karpov, E. Kim, M. Kolisko, A. Kudryavtsev, D. J. G. Lahr, E. Lara, L. Le Gall, D. H. Lynn, D. G. Mann, R. Massana, E. A. D. Mitchell, C. Morrow, J. S. Park, J. W. Pawlowski, M. J. Powell, D. J. Richter, S. Rueckert, L. Shadwick, S. Shimano, F. W. Spiegel, G. Torruella, N. Youssef, V. Zlatogursky & Q. Zhang. 2019. Revisions to the classification, nomenclature, and diversity of eukaryotes. Journal of Eukaryotic Microbiology 66: 4–119.

[AS12] Adl, S. M., A. G. B. Simpson, C. E. Lane, J. Lukeš, D. Bass, S. S. Bowser, M. W. Brown, F. Burki, M. Dunthorn, V. Hampl, A. Heiss, M. Hoppenrath, E. Lara, E. Le Gall, D. H. Lynn, H. McManus, E. A. D. Mitchell, S. E. Mozley-Stanridge, L. W. Parfrey, J. Pawlowski, S. Rueckert, L. Shadwick, C. L. Schoch, A. Smirnov & F. W. Spiegel. 2012. The revised classification of eukaryotes. Journal of Eukaryotic Microbiology 59 (5): 429–493.

[AA05] Afanasieva, M. S., E. O. Amon, Y. V. Agarkov & D. S. Boltovskoy. 2005. Radiolarians in the geological record. Paleontological Journal 39 (Suppl. 3): S135–S392.

Campbell, A. S. 1951. New genera and subgenera of Radiolarida. Journal of Paleontology 25 (4): 527–530.

[C-S10] Cavalier-Smith, T. 2010. Kingdoms Protozoa and Chromista and the eozoan root of the eukaryotic tree. Biology Letters 6 (3): 342–345.

[C-S18] Cavalier-Smith, T. 2018. Kingdom Chromista and its eight phyla: a new synthesis emphasising periplastid protein targeting, cytoskeletal and periplastid evolution, and ancient divergences. Protoplasma 255: 297–357.

[C-SC06] Cavalier-Smith, T., & E. E.-Y. Chao. 2006. Phylogeny and megasystematics of phagotrophic heterokonts (kingdom Chromista). Journal of Molecular Evolution 62: 388–420.

[C-SCL15] Cavalier-Smith, T., E. E. Chao & R. Lewis. 2015. Multiple origins of Heliozoa from flagellate ancestors: new cryptist subphylum Corbihelia, superclass Corbistoma, and monophyly of Haptista, Cryptista, Hacrobia and Chromista. Molecular Phylogenetics and Evolution 93: 331–362.

[FT93] Fensome, R. A., F. J. R. Taylor, G. Norris, W. A. S. Sarjeant, D. I. Wharton & G. L. Williams. 1993. A classification of living and fossil dinoflagellates. Micropaleontology Special Publication 7: i–viii, 1–351.

Haeckel, E. 1882. Entwurf eines Radiolarien-Systems auf Grund von Studien der Challenger-Radiolarien. Jenaische Zeitschrift für Naturwissenschaft 15: 418–472.

Haeckel, E. 1887. Report on the Radiolaria collected by H.M.S. Challenger during the years 1873–1876. Second part.—Osculosa. (Nassellaria and Phaeodaria). Report on the Scientific Results of the Voyage of H.M.S. Challenger during the years 1873–76. Zoology 18 (2): 889–1803.

[KC01] Kirk, P. M., P. F. Cannon, J. C. David & J. A. Stalpers. 2001. Ainsworth & Bisby’s Dictionary of the Fungi 9th ed. CAB International: Wallingford (UK).

Lazarus, D. 2014. The legacy of early radiolarian taxonomists, with a focus on the species published by early German workers. Journal of Micropalaeontology 33: 3–19.

Suzuki, N., L. O’Dogherty, J.-P. Caulet & P. Dumitrica. 2021. A new integrated morpho- and molecular systematic classification of Cenozoic radiolarians (class Polycystinea)—suprageneric taxonomy and logical nomenclatorial acts. Geodiversitas 43 (15): 405–573.

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