JOINT HAECKEL and EHRENBERG PROJECT: Reexamination of the Haeckel and Ehrenberg Microfossil Collections
as a Historical and Scientific Legacy, edited by Y. Tanimura and Y. Aita, pp. 55–69. National Museum of Nature andScience Monographs, No. 40, Tokyo 2009

Radiolarian Studies by Christian Gottfried Ehrenberg (1795–1876) andValue of his Species as Type Species of Genera
Noritoshi Suzuki
Institute of Geology and Paleontology, Graduate School of Science, Tohoku University,
6–3 Aoba Aramaki, Aoba-ku, Sendai 980–8578, Japan(e-mail: suzuki.noritoshi@nifty.com)

Abstract Since there are a few reports on historical reviews of Ehrenberg’s studies for polycystineradiolarians, his studies are reviewed herein. Christian Gottfried Ehrenberg (1795–1876) published 41
papers concerning polycystine radiolarians. He described 72 genera and 501 species in his life. I list
the type species of 134 genera whose type species are selected from Ehrenberg’s species, and summarize taxonomic issues for these genera.
Key words:

Classification, Ehrenberg, Radiolaria, Type species.

Introduction
Since the first fossil polycystine radiolarians were described from Caltanisetta, Oran and
Zante by Ehrenberg (1839), more than 10,000 species have been described over the last 170 years(Suzuki and Aita, submitted). Christian Gottfried Ehrenberg (1795–1876) published 41 papers
regarding polycystine radiolarians in the modern sense from 1839 to 1876 (Lazarus and Suzuki,2009). A total number of 528 polycystine species were found from Recent sea-bottom sediments,
Neogene on-land samples and upper Paleogene Barbados samples, and 486 of them are still taxonomically available names (Ogane et al., 2009; Suzuki et al., 2009). Many of Ehrenberg’s species
have been used in the subsequent literature as age-diagnostic or paleoceanographic markers.
Many of these subsequent papers cited Ehrenberg’s papers with incorrect titles, conflicting pages
ranges, inaccurate journal names and even to nonexistent descriptions, presumably due to repeated indirect citation without directly checking the sources, which were until recently not readily
obtainable. As described in Lazarus and Suzuki (2009 in this volume) the correct titles, dates andcontent of Ehrenberg’s publications are now readily available, and are used in this paper throughout.
Ehrenberg’s radiolarian studies largely relied on samples which other scientists and explorers
sent to him. His primary research goals were to reveal the diversity of ‘Infusoria’, including polycystine radiolarians, and by comparing assemblages worldwide, to document the geographicaldistributions of Infusoria, and to attempt, albeit from bottom sediments, to infer their vertical distribution in the water column (e.g. Cole, 1926). His early work focused on description of new
species from many localities, which were summarized in his monograph Mikrogeologie Ehrenberg (1854c) and its supplement published in 1856. Radiolarian studies by Ehrenberg are thuspart of this broader research context.

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Historical review of the radiolarian studies
The first publication regarding polycystine radiolarians (Fig. 1) in the current sense is Ehrenberg (1839). This work was presented earlier orally in the Prussian Academy of Sciences(“Akademie der Wissenschaften”), Berlin, on 20th December, 1838, and was formally distributed
as publication in 1839 (Lazarus & Suzuki, 2009). He described four genera and 9 radiolarianspecies from Caltanisetta (Sicily), Oran (Africa) and Zante of John Island (pullout table in Ehrenberg, 1839). He proposed the “family Polycystina” as a member of the higher (also from Ehrenberg) taxon Polygastrica (p. 128 in Ehrenberg, 1839). Polycystina included Lithocampe, Cornutella, Haliomma. In this paper, a diatom, Coscinodiscus, and a silicoflagellate, Dictyocha, wereregarded as members of Polycystina. The remaining genus, Flustrella, was considered to belong
to Ehrenberg's higher taxon Polythalamia. Polygastrica was defined by presence of many stomachs in an individual whereas Polythamalia by presence of many chambers which can live individuals in each “chamber”. Although 9 radiolarian species were described, only one specimen,
named Lithocampe radiculata, was illustrated in this paper.
Ehrenberg (1840) described 274 Polygastrica species, and of these, 10 species belong to
Polycystina in the current scheme. In this paper, all the Polycystina which were described at thattime were noted. The four new species were newly described from Greece (‘E marga Graeciae’ inthe original spelling). In this paper, Flustrella was first moved to Polygastrica from Polythalamia
because these forms were noted to have a siliceous test (p. 210 in Ehrenberg, 1840). It is noteworthy that Ehrenberg (1840) described “FLUSTRELLA spiralisFlustrella concentrica Kr. B.
1838. p. 76. ex parte spirali.” This is interpreted in this paper that the specimen which was identified as F. concentrica on page 76 of Ehrenberg (1839) was now transferred to a new species, F.
spiralis. Ehrenberg (1840) also described Haliomma ovatum as a new species. The genus
Haliomma was first described by Ehrenberg (1839) and included two species Haliomma crenatum and Haliomma medusa, and consequently, Haliomma species could be presumed to belong
to Polycystina. However, Haliomma ovatum is now considered to be a siliceous sponge spicule.
Similar to H. ovatum, Ehrenberg (1841) described Haliomma (?) radians as a new species fromthe North Sea (Atlantic) “Nordsee bei Cuxhaven”, but this species belongs to siliceous sponge

Fig. 1.

The first illustrated fossil polycystine radiolarian specimen in the history, from Ehrenberg (1839).

Radiolarian Studies by Ehrenberg

57

spicules, too. Ehrenberg (1842) introduced 39 new organisms from Aegina but he did not publish
any definitions for these species. Two new species: Lithobotrys cribrosa and Cornutella Lithocampe would presumably be polycystine radiolarians in current taxonomy, but they are apparently nomen nudum. Although Ehrenberg (1842) did not describe new species, he first noted that hisorganisms were routinely illustrated by him, and at a magnitude of 300X (1842, p. 265).
In the 1844 issues of the Prussian Academy of Sciences Proceedings journal “Bericht überdie zur Bekanntmachung geeigneten Verhandlungen der Königlich Preußischen Akademie derWissenschaften zu Berlin”, three polycystine radiolarian papers were published, on pages 57–97(Ehrenberg, 1844a), 182–207 (Ehrenberg, 1844b) and 252–274 (Ehrenberg, 1844c). Ehrenberg(1844a) principally compared the assemblage similarity of micro-organisms (his Infusoria)
among North America, Europe and Africa. The faunal similarity of polycystines was comparedamong Oran (Africa), Caltanisetta and Aegina. Ehrenberg (1844a) also described one new genus(Lithobotrys) and 14 new polycystine species from Caltanisetta and Aegina. This paper also introduced the name Haliomma cornutum on page 64, but this species has never been formally described. Ehrenberg (1844a) described “FLUSTRELLA bilobataHALIOMMA Lagena 1840”
on page 81. The meaning of this assignment was not solved solely with his published references
but also with the aid of Ehrenberg’s original drawings housed in the Museum, which indicate thatthe same individuals have two species names. Since ‘Haliomma lagena’ is an available name by
the description of Ehrenberg (1840), the name ‘Flustrella bilobata’ is a primary objective juniorsynonym of Haliomma lagena. Ehrenberg (1844b) is the first report of polycystine radiolarians
from the Southern Ocean. The materials were collected between 1841–1843 by captain Ross, andpolycystine radiolarians were found from ‘Pancake Ice’ which was obtained at 78°10S and
162°W. Lithobotrys (?) denticulata and Lithocampe antarctica were newly described in Ehrenberg (1844b). The presence of Lithocampe australis was noted in page 187 of Ehrenberg(1844b), but was not formally described in this paper. Ehrenberg (1844c) reported Infusoria fromthe Bermuda Islands. Three new polycystine radiolarians (Haliomma (?) amphisiphon, H. (?) nobile and Lithocampe aculeata) were newly described in this paper.
Ehrenberg (1845) listed many species of Infusoria from 14 locations, including Charles Darwin’s samples (p. 63–66). This paper also described many new genera and new species of Infusoria, and a species from ‘Westmoreland Virginiae’ was named as Lithocampe (?) stiligera.
Ehrenberg (1846) was the first paper which focused only polycystine radiolarians. Robert H.
Schomburgk donated samples from Barbados to Ehrenberg (Schomburgk, 1847), and Ehrenberg(1846) noted that 140 species were found from Barbados samples. In Ehrenberg (1846), Polycystina was divided into two groups, “Polycystina Solitaria” and “Polycystina Composita”, andthey were further subdivided into Halicalyptrina, Lithochytrina and Eucyrtidina in Solitaria while
Haliommatina, Spyridina and Lithoccylidina were created within Composita. This paper newly
described 26 genera without species (p. 385). Since the definition of these genera was depicted in
Ehrenberg (1846), these genera are considered to be formally published in this paper.
His taxonomy was revised, and new taxa were described in Ehrenberg (1847). Ehrenberg(1847) first transferred genera in some species on pages 42–43. The list on these pages is comparable to synonym list in the modern articles, but includes nine nomen nudum species. Ehrenberg(1847) discussed the formation of siliceous tests, added a list of polycystine radiolarians for thepreviously described samples (e.g. South Pole “Südpols’), and noted 14 new species from Barbados. Of these, 11 species were illustrated. Although these species were not associated with descriptions, they are regarded as taxonomically available names only with illustration on the basis
of ICZN 2000. A total of 14 new genera were also newly defined in this paper.

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Ehrenberg (1854a) is the first paper to concern the vertical distribution of polycystine radiolarians. He compared the infusorian assemblages between sediments which were recovered fromdifferent water depths of 433, 840, 1050, 6480, 8160, 9480, 10800 and 12000 feet (Fuß), fromthe ocean bottom (‘Meersgrund’). In this paper the vertical distribution of 40 polycystine radiolarians were shown. Twenty-eight of 40 species names first appeared in Ehrenberg (1854a), butwere not formally described. All of these species were described in Ehrenberg (1854b). The typematerials of these new species are recognized as the samples from 6480, 10800, and 12000 feet.
‘Flustrella subtilis’ was stated as having been found from the sample 8460 feet on page 242, but
the actual specimen was detected from the sample from 6480 feet in our project.
The ‘Mikrogeologie’ is a milestone for Ehrenberg’s studies on Infusoria since all the Infusoria previously published were summarized in this book. Ehrenberg had attached only 12 species(Ehrenberg, 1839, 1847) but Ehrenberg (1854c) showed the illustrations of 70 polycystine radiolarian species, including 23 new species. These illustrations are duplicated with the previously illustrated 12 species, and subsequently the illustrations of 58 species were first published in
Ehrenberg (1854c).
After publishing the ‘Mikrogeologie’, Ehrenberg continued to examine the diversity of infusorians in many regions. Five polycystine radiolarian species were found in an ocean bottom
sample which was taken from a water depth of 12,900 feet (Ehrenberg, 1855a), eleven species
were found from “Simbirsk” (now Ulyanovsk, a city in Russia on the Volga River) (Ehrenberg,
1855b), one “polycystine” (but now identified as a sponge spicule) from Java (Ehrenberg, 1855c)
and seven genera from Chile (Ehrenberg, 1856). Ehrenberg (1855b) noted four new speciesnames, but only one species, Eucyrtidium simbirscianum, was formally described in Ehrenberg(1876).
The vertical distributions of polycystine radiolarians were revisited by Ehrenberg (1858).
This paper examined seafloor samples from ‘Meersgund bei Candia’ (possibly modern Crete)and compared these with Ehrenberg (1854c). Ehrenberg (1858) published 13 new species names
which were described in Ehrenberg (1859).
A total of 79 polycystine radiolarians were found from the surface sediments (sounding sampling) of the Philippine Sea (18°03N, 129°11E) (Ehrenberg, 1861a). Although Ehrenberg(1861a) included 48 nomen nudum species, this paper is of particular importance in providing
data with the number of individuals for each species. Vertical distributions of Infusoria were stillof interest for Ehrenberg. Ehrenberg (1861b) listed the vertical distributions of 69 polycystine radiolarians from California Ocean, while Ehrenberg (1862a) showed the distributions of fourpolycystine radiolarians from the Gulf of Mexico, near Florida. Ehrenberg (1861a, 1861b, 1862a)
listed many new species, but these papers included neither illustrations nor descriptions. The definition of 22 genera was only depicted in Ehrenberg (1861b).
The significant observation that polycystine radiolarians live in similar oceanographic conditions at the species level was first noted in Ehrenberg (1862b). Ehrenberg (1862b) examined 8
sea-bottom samples which were collected during a preliminary survey for the construction oftelegraph lines. He also summarized the vertical distributions of Infusoria based on the water
depths of the sea-floor, and he identified seven species which were found in Kamchatka by Bailey(1856). These seven species were Cadium marinum Bailey, Botryocampe inflata (Bailey), Carpocanium cornutum (Bailey), Cornutella annulata Bailey, Eucyrtidium cuspidatum Bailey,
Eucyrtidium hyperboreum Bailey and Eucyrtidium tumidulum Bailey, which were considered as
arctic species. Ehrenberg (1862b) formally described 27 new species and variations. The type localities of these new species are now specified on page 819–820 in Ehrenberg (1861b) for ‘Cali-

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59

fornico’ (California Ocean) and page 224–225 in Ehrenberg (1862a) for Mexicano’ (Gulf of
Mexico, near Florida), and page 276 in Ehrenberg (1862b) for “Groenland” (Davis strait), respectively.
After 1862, Ehrenberg did not publish papers on polycystine radiolarians until 1873. Ehrenberg (1873a) did not examine any new materials. Instead, he summarized his studies since 1836
and described 249 infusorians, including 109 new polycystine radiolarians from ‘Zanguebarica
13200’, ‘Insulam Milonem 1440’, ‘Philippine Ocean 19800’, ‘California Ocean 15600’, ‘Camorta Nicobarica’, ‘Florida 9066’, ‘Atlantic Ocean 9540’, and ‘Atlantic Ocean 9780’ He counted a
total of 287 polycystines that were described by his studies excepting Barbados. The illustrationsof some of these species were shown in Ehrenberg (1873b).
Ehrenberg (1873b) is a comprehensive monograph of his studies on marine micro-organisms.
The earlier part of Ehrenberg (1873b) summarized the assemblages examined after Ehrenberg(1854c). The examined samples were mainly collected from the sea-floor. The middle part of thepaper are summary cross tables (Fig. 5) between geographic zones and water depths of 1924 organisms (605 Polythalamia, 656 Polygastina, 279 Polycystina, 219 Phytolitharia, 56 Geolithia, 39
Zoolitharia, 51 Molluska, 1 Annulata, 2 Entomostraca, 6 Radiata, 1 Anthozoa, and 9 Bryozoa).
Ehrenberg (1873b) listed 279 polycystine radiolarian species, but not all of them are taxonomically available species because he marked 44 species without descriptions in these tables. Thesesummary tables are also very useful because the first description and the illustrated papers were
given in these tables. This list shows that the highest diversity was recorded in the tropical zone
and the diversity increases in deeper waters, although this latter estimation is not based on any direct water column data. The later part of Ehrenberg (1873b) discussed various aspects of his data.
Remarks on polycystine radiolarians are given from page 339 to 346. Based on his observations,
Ehrenberg (1873b) criticized the observations of living polycystine radiolarians by J. Müller andHaeckel in Messina (e.g. Müller, 1858; Haeckel, 1862). On page 344, Ehrenberg comments that
their conclusions were prematurely derived from local phenomena and their claim of rich radiolarians in the surface water is contradicted his compiled data that radiolarians increased in diversity toward deeper waters. Ehrenberg has not used the term “Radiolaria” instead of Polycystina inthat Polycystina has a priority (page 345 in Ehrenberg, 1873b). Ehrenberg (1873b) concluded
that Polycystina are closely related to Echinodermata based on a similar degree of complexity.
Another of Ehrenberg’s monographic works on polycystine radiolarians are his studies of radiolarians from Barbados. Brief notes and illustrations were already published in Ehrenberg(1846, 1847, 1854c), but only a few species were formally described and illustrated in these publications. Ehrenberg (1874) comprehensively described 265 species from Barbados samples. All
the genera used in Ehrenberg (1874) were previously published in Ehrenberg (1847, p. 53). According to the final paragraphs of this paper (p. 263), almost all the names in this paper and original drawings were already prepared before publication of his 1846 and 1847 papers. Ehrenberg(1874), however, has no illustrations.
Ehrenberg (1876) illustrated 85 specimens of 81 polycystine radiolarian species from Barbados, including 14 species (3 nomen nudum species). Ehrenberg (1876) gives an occurrence list offossil polycystine radiolarians from Barbados samples (coded as 1, 3, 14, 15, 16, 25, 26, 30, 43,
51, 59, 60, 74, 81, 94 and 95) and detailed information for these samples. Ehrenberg (1876) also
described occurrences from Nicobarica (p. 116–120), and discussed the systematic classificationof his Polycystine from page 151 to 157. Ehrenberg (1876) introduced the division of Spumellaria and Nassellaria on p. 156, and summarized the classification of all genera which were proposed by himself. This summary table excluded any genera which were proposed by other au-

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thors (Meyen, 1834; Huxley, 1851; Claparede, 1856; Müller, 1857, 1858; Haeckel, 1860a, 1860b,
1862). Ehrenberg (1876) principally is a comprehensive summary monograph of his fossil studies through his life. Ehrenberg (1876) compiled the worldwide occurrences of more than a thousand micro-organisms with geological ages and regions. A total of 326 polycystine radiolarians
were listed as fossils in this paper (p. 65–85) from Barbados, Bermuda, ‘Java, Nicobarien.
Scheduba’, the southern States of North America (‘Süd-Staaten North America’), Greece(Griechenland), Italey (Italien), Russia (Rußland), South America (Süd-America), and the southwestern coast of South Africa (‘Oran. Saldanha-Bay. Agulhas-Bank’). Ehrenberg (1876) was
probably the first paper to list the geological distributions of polycystine radiolarians, but his results are not usable any longer because the geological time scale used in Ehrenberg (1876) was
only divided it into Primary, Secondary, Tertiary and Quaternary, and thus all the fossil polycystine radiolarians were classified into the ‘Tertiary” period.
Type species using Ehrenberg’s species
Since the concept of type species for a genus did not become common in the 19th century,
Ehrenberg himself did not designate any type species for his genera. A total of 131 species, including 6 nomen nudum species, have been selected, mostly by subsequent designation by various later authors, as the type species of 134 genera. These are summarized in Table 1. The inconsistent numbers between species and genus are caused by duplicated designation. Based on thecondition of type designation, these species can be categorized into four groups, namely, typespecies (7 species), monotype (13 species), subsequent monotype (8 species) and subsequent
designation (118 species), following ICZN 2000. The definition of each term is cited below:
Type species (Glossary in ICZN 2000): The nominal species that is the name-bearing typeof a nominal genus or subgenus.
Monotypy (ICZN 2000 Art 68.3): When an author establishes a new nominal genus-grouptaxon for a single taxonomic species and denotes that species by an available name,the nominal species so named is the type species.
Subsequent monotypy (ICZN Art 69.3): If only one nominal species was first subsequently included in a nominal genus or subgenus established without included species, that
nominal species is automatically fixed as the type species, by subsequent monotypy.
Subsequent designation (ICZN 2000 Art 69.1): If an author established a nominal genus
or subgenus but did not fix its type species, the first author who subsequently designates one of the originally included nominal species validly designates the type speciesof that nominal genus or subgenus (type by subsequent designation), and no later designation is valid.
As shown Table 1, Ceratospyris Ehrenberg 1846, Cycladophora Ehrenberg 1846, Dictyospyris Ehrenberg 1846, Haliomma Ehrenberg 1839, Lithobotrys Ehrenberg 1844a, Lithomelissa Ehrenberg 1847, Lithostrobus Bütschli 1882, Lychnocanium Ehrenberg 1846 and Spongosphaera Ehrenberg 1847 have more than two type species. We don’t solve this issue in ourpaper because radiolarian communities have been familiar with the current usage, even this iswrong under ICZN 2000, rather than unfamiliar designations of type species. We only note thatthe following rules in ICZN 2000 are helpful to evaluate better usage of type designation.

GenusAcanthosphaera haliphormis Ehrenberg (1862b)
Lithomelissa microptera Ehrenberg (1854c)
Polysolenia setosa Ehrenberg (1873a)
Amphicentria salpa Ehrenberg (1862b)
Lithocampe clava Ehrenberg (1874)
Lithobotrys denticulata Ehrenberg (1844b)
Anthocyrtis mespilus Ehrenberg (1847)
Anthocyrtis mespilus Ehrenberg (1847)
Anthocyrtis ophirensis Ehrenberg (1873a)
Anthocyrtis serrulata Ehrenberg (1874)
Halicalyptra cancellata Ehrenberg (1854b)
Eucyrtidium profundissimum Ehrenberg (1873a)
Lithornithium loxia Ehrenberg (1854c)
Lithocampe aurita Ehrenberg (1844a)
Astromma aristotelis Ehrenberg (1847)
Astromma entomocora Ehrenberg (1854c)
Lithobotrys inflatum Bailey (1856)
Lithobotrys nucula Ehrenberg (1874)
Botryocyrtis caput-serpentis Ehrenberg (1873a)
Ceratospyris ocellata Ehrenberg (1874)
Calocyclas turris Ehrenberg (1874)
Eucyrtidium ampulla Ehrenberg (1862a)
Lithocampe solitaria Ehrenberg (1839)
Cenosphaera plutonis Ehrenberg (1854b)
Cornutella trochus Ehrenberg (1873a)
Cornutella cucullaris Ehrenberg (1874)
Ceratospyris pentagona Ehrenberg (1873a)
Haliomma radicatum Ehrenberg (1844a)
Chilomma saturnus Ehrenberg (1862b)
Ceratospyris ramosa Ehrenberg (1874)
Clathrocanium squarosum Ehrenberg (1873a)
Eucyrtidium trochus Ehrenberg (1873a)
Cornutella clathrata Ehrenberg (1839)
Cornutella trochus Ehrenberg (1873a)
Lithomelissa corythium Ehrenberg (1874)
Cryptoprora fundicola [nomen nudum] Ehrenberg (1873b)

Type species

The Ehrenberg’s species which are designated as the type species of polycystine radiolarians.

Acanthosphaera Ehrenberg (1862b)
Acromelissa Haeckel (1887)
Acrosphaera Haeckel (1882)
Amphicentria Ehrenberg (1862b)
Amphipternis Foreman (1973a)
Antarctissa Petrushevskaya (1967)
Anthocyrtella Haeckel (1887)
Anthocyrtis Ehrenberg (1846)
Anthocyrtissa Haeckel (1887)
Anthocyrtoma Haeckel (1887)
Archiphormis Haeckel (1882)
Artocyrtis Haeckel (1887)
Artoperina Campbell (1951)
Artostrobium Haeckel (1887)
Astractinium Haeckel (1887)
Astromma Ehrenberg (1846)
Botryocampe Ehrenberg (1861b)
Botryocella Haeckel (1887)
Botryocyrtis Ehrenberg (1861b)
Brachiospyris Haeckel (1882)
Calocyclas Ehrenberg (1847)
Calocycloma Haeckel (1887)
Carpocanium Ehrenberg (1846)
Cenosphaera Ehrenberg (1854b)
Ceratastrum Haeckel (1882)
Ceratocyrtis Bütschli (1882)
Ceratospyris Ehrenberg (1846)
Ceratospyris Ehrenberg (1846)
Chilomma Ehrenberg (1847)
Cladospyris Ehrenberg (1846)
Clathrocanium Ehrenberg (1873a)
Conarachnium Haeckel (1882)
Cornutella Ehrenberg (1839)
Cornutellium Haeckel (1882)
Corythomelissa Campbell (1951)
Cryptoprora Ehrenberg (1847)

Table 1.

Ehrenberg (1862b)
Foreman (1973a)
Petrushevskaya (1967)
Petrushevskaya and Kozlova (1972)
Campbell (1954)
Campbell (1954)
Frizzell and Middour (1951)
Campbell (1954)
Frizzell and Middour (1951)
Campbell (1954)
Campbell (1954)
Campbell (1954)
Campbell (1954)
Ehrenberg (1862b)
Campbell (1954)
Campbell (1954)
Frizzell and Middour (1951)
Campbell (1954)
Campbell (1954)
Campbell (1954)
Ehrenberg (1854b)
Petrushevskaya (1981)
Petrushevskaya (1971)
Campbell (1954)
Ehrenberg (1847)
Ehrenberg (1862b)
Campbell (1954)
Campbell (1954)
Campbell (1954)
Ehrenberg (1839)
Petrushevskaya and Kozlova (1972)
Campbell (1954)
Campbell (1954)

Ehrenberg (1862b)
Petrushevskaya (1975)

Assigned paper
Monotype
Subsequent designation
Subsequent designation
Monotype
Type species
Type species
Subsequent designation
Subsequent designation
Subsequent designation
Subsequent designation
Subsequent designation
Subsequent designation
Type species
Subsequent designation
Subsequent designation
Subsequent designation
Subsequent monotype
Subsequent designation
Subsequent designation
Subsequent designation
Subsequent designation
Subsequent designation
Subsequent designation
Monotype
Subsequent designation
Subsequent designation
Subsequent designation
Subsequent monotype
Subsequent monotype
Subsequent designation
Subsequent designation
Subsequent designation
Monotype
Subsequent designation
Type species
Subsequent designation

Type mode

Radiolarian Studies by Ehrenberg
61

GenusContinued.

Cycladophora Ehrenberg (1846)
Cycladophora Ehrenberg (1846)
Dendrospyris Haeckel (1882)
Dictyastrum Ehrenberg (1861b)
Dictyocephalus Ehrenberg (1861b)
Dictyocoryne Ehrenberg (1861b)
Dictyocryphalus Haeckel (1887)
Dictyomitrella Haeckel (1887)
Dictyophimus Ehrenberg (1846)
Dictyopodium Ehrenberg (1847)
Dictyospyrella Haeckel (1887)
Dictyospyris Ehrenberg (1846)
Dictyospyris Ehrenberg (1846)
Dictyospyrissa Haeckel (1887)
Dipospyris Haeckel (1887)
Disolenia Ehrenberg (1861b)
Druppatractona Haeckel (1887)
Euchitonia Ehrenberg (1861b)
Eucyrtidium Ehrenberg (1846)
Eucyrtis Haeckel (1882)
Flustrella Ehrenberg (1839)
Giraffospyris Haeckel (1882)
Glycobotrys Campbell (1951)
Halicalyptra Ehrenberg (1846)
Haliomma Ehrenberg (1839)
Haliomma Ehrenberg (1839)
Haliomma Ehrenberg (1839)
Haliphormis Ehrenberg (1846)
Hexaspyridium Haeckel (1887)
Hexastylus Haeckel (1882)
Histiastrum Ehrenberg (1846)
Hymenastromma Haeckel (1887)
Hymeniastrum Ehrenberg (1846)
Lamprodiscus Ehrenberg (1861b)
Litharachnidium Haeckel (1887)
Litharachnium Haeckel (1861b)
Lithobotrys Ehrenberg (1844a)

Table 1.

Cycladophora (?) davisiana Ehrenberg (1862b)
Cycladophora stiligera Ehrenberg (1874)
Ceratospyris stylophora Ehrenberg (1874)
Dictyastrum angulatum Ehrenberg (1873a)
Cornutella obtusa Ehrenberg (1844a)
Dictyocoryne profunda Ehrenberg (1873a)
Cornutella obtusa Ehrenberg (1844a)
Eucyrtidium articulatum Ehrenberg (1874)
Dictyophimus crisiae Ehrenberg (1854b)
Dictyopodium eurylophus Ehrenberg (1874)
Dictyospyris tristoma Ehrenberg (1874)
Dictyospyris ceratospyris [nomen nudum] Ehrenberg (1873b)
Dictyospyris trilobata Ehrenberg (1854c)
Dictyospyris fenestra Ehrenberg (1874)
Dictyospyris ceratospyris [nomen nudum] Ehrenberg (1873b)
Disolenia follis [nomen nudum] Ehrenberg (1873b)
Stylosphaera laevis Ehrenberg (1874)
Euchitonia furcata Ehrenberg (1873a)
Lithocampe acuminata Ehrenberg (1844a)
Lithocampe acuminata Ehrenberg (1844a)
Flustrella concentrica Ehrenberg (1839)
Ceratospyris heptaceros Ehrenberg (1874)
Lithobotrys geminata Ehrenberg (1874)
Halicalyptra virginica Ehrenberg (1854c)
Haliomma aequorea Ehrenberg (1844a)
Haliomma beroes Ehrenberg (1854b)
Haliomma medusa Ehrenberg (1839)
Haliphormis calva [nomen nudum] Ehrenberg (1854b)
Ceratospyris setigera Ehrenberg (1874)
Acanthosphaera setosa Ehrenberg (1873a)
Histiastrum quaternarium Ehrenberg (1874)
Histiastrum ternarium Ehrenberg (1874)
Hymeniastrum pythagorae Ehrenberg (1854c)
Lamprodiscus monoceros Ehrenberg (1873a)
Carpocanium arachnodiscus Ehrenberg (1862b)
Carpocanium arachnodiscus Ehrenberg (1862b)
Lithobotrys geminata Ehrenberg (1874)

Type species
Lombari and Lazarus (1988)
Campbell (1954)
Campbell (1954)
Strelkov and Reshetnyak (1959)
Ehrenberg (1861b)
Campbell (1954)
Loeblich and Tappan (1961)
Campbell (1954)
Riedel and Sanfilippo (1970)
Campbell (1954)
Campbell (1954)
Campbell (1954)
Sanfilippo et al. (1973)
Campbell (1954)
Campbell (1954)
Campbell (1954)
Campbell (1954)
Campbell (1954)
Frizzell and Middour (1951)
Frizzell and Middour (1951)
Ehrenberg (1839)
Campbell (1954)
Campbell (1954)
Campbell (1954)
Campbell (1954)
Strelkov and Reshetnyak (1959)
Ehrenberg (1839)
Campbell (1954)
Petrushevskaya (1981)
Kozur and Mostler (1979)
Campbell (1954)
Campbell (1954)
Strelkov and Reshetnyak (1959)
Campbell (1954)
Petrushevskaya (1981)
Campbell (1954)
Petrushevskaya (1981)

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Type species
Lithobotrys quadriloba Ehrenberg (1844a)
Lithocampe radicula Ehrenberg (1839)
Lithochytris vespertilio Ehrenberg (1874)
Lithobotrys galea Ehrenberg (1844a)
Lithocyclia ocellus Ehrenberg (1854c)
Lithomelissa (?) tartari Ehrenberg (1854b)
Lithomelissa microptera Ehrenberg (1854c)
Eucyrtidium pachyderma Ehrenberg (1874)
Eucyrtidium acephalum Ehrenberg (1874)
Lithopera bacca Ehrenberg (1873a)
Lithocampe hirundo Ehrenberg (1840)
Eucyrtidium argus Ehrenberg (1874)
Eucyrtidium picus Ehrenberg (1874)
Eucyrtidium antilope Ehrenberg (1872)
Eucyrtidium stephanophorum Ehrenberg (1874)
Lophophaena galea Ehrenberg (1854b)
Lophophaena radians Ehrenberg (1874)
Lychnocanium falciferum Ehrenberg (1854c)
Lychnocanium lucerna Ehrenberg (1847)
Mazosphaera laevis Ehrenberg (1873a)
Cornutella cassis Ehrenberg (1844a)
Podocyrtis aculeata Ehrenberg (1874)
Ommatocampe polyarthra Ehrenberg (1873a)
Ommatogramma navicularis Ehrenberg (1873a)
Ommatospyris apicata Ehrenberg (1873a)
Petalospyris confluens Ehrenberg (1874)
Astromma pentactis Ehrenberg (1874)
Podocyrtis pentacantha Ehrenberg (1874)
Flustrella praetexta Ehrenberg (1844a)
Periphaena decora Ehrenberg (1874)
Petalospyris platyacantha Ehrenberg (1874)
Petalospyris foveolata Ehrenberg (1854c)
Podocyrtis papalis Ehrenberg (1847)
Polysolenia setosa Ehrenberg (1873a)
Dictyocephalus galeatus Ehrenberg (1873a)
Pteractis elegans Ehrenberg (1873a)

Lithobotrys Ehrenberg (1844a)
Lithocampe Ehrenberg (1839)
Lithochytris Ehrenberg (1846)
Lithocorythium Ehrenberg (1847)
Lithocyclia Ehrenberg (1846)
Lithomelissa Ehrenberg (1847)
Lithomelissa Ehrenberg (1847)
Lithomitra Bütschli (1882)
Lithomitrella Haeckel (1887)
Lithopera Ehrenberg (1846)
Lithornithium Ehrenberg (1847)
Lithostrobus Bütschli (1882)
Lithostrobus Bütschli (1882)
Lophoconus Haeckel (1887)
Lophocyrtis Haeckel (1887)
Lophophaena Ehrenberg (1847)
Lophophaenoma Haeckel (1887)
Lychnocanium Ehrenberg (1846)
Lychnocanium Ehrenberg (1846)
Mazosphaera Ehrenberg (1861b)
Microlecitos Issel (1890)
Octalacorys Haeckel (1887)
Ommatocampe Ehrenberg (1861b)
Ommatogramma Ehrenberg (1861b)
Ommatospyris Ehrenberg (1861b)
Patagospyris Haeckel (1882)
Pentactura Haeckel (1882)
Pentalacorys Haeckel (1882)
Perichlamydium Ehrenberg (1846)
Periphaena Ehrenberg (1874)
Petalospyrella Haeckel (1887)
Petalospyris Ehrenberg (1846)
Podocyrtis Ehrenberg (1846)
Polysolenia Ehrenberg (1861b)
Psilomelissa Haeckel (1882)
Pteractis Ehrenberg (1873a)

Continued.

Genus

Table 1.

Campbell (1954)
Campbell (1954)
Campbell (1954)
Petrushevskaya (1981)
Campbell (1954)
Campbell (1954)
Petrushevskaya and Kozlova (1979)
Campbell (1954)
Petrushevskaya and Kozlova (1979)
Campbell (1954)
Campbell (1954)
Campbell (1954)
Strelkov and Reshetnyak (1959)
Frizzell and Middour (1951)
Campbell (1954)
Campbell (1954)
Petrushevskaya and Kozlova (1979)
Campbell (1954)
Ehrenberg (1847)
Campbell (1954)
Issel (1890)
Campbell (1954)
Campbell (1954)
Campbell (1954)
Campbell (1954)
Campbell (1954)
Campbell (1954)
Campbell (1954)
Campbell (1954)
Ehrenberg (1874)
Campbell (1954)
Campbell (1954)
Campbell (1954)
Campbell (1954)
Campbell (1954)
Ehrenberg (1873a)

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63

GenusContinued.

Pterocanium Ehrenberg (1846)
Pterocodon Ehrenberg (1847)
Pterocyrtidium Bütschli (1882)
Pylosphaera Ehrenberg (1858)
Rhabdolithis Ehrenberg (1847)
Rhopalastrum Ehrenberg (1846)
Rhopalocanium Ehrenberg (1846)
Rhopalodictyum Ehrenberg (1861b)
Schizomma Ehrenberg (1861b)
Sestropyramis Haeckel (1887)
Sethoconus Haeckel (1887)
Sethodisculus Haeckel (1887)
Sethodiscus Haeckel (1882)
Sethopyramis Haeckel (1882)
Solenosphaera Haeckel (1887)
Spirillina Ehrenberg (1858)
Spongaster Ehrenberg (1862a)
Spongatractus Haeckel (1887)
Spongodiscus Ehrenberg (1854b)
Spongosphaera Ehrenberg (1847)
Spongosphaera Ehrenberg (1847)
Staurolonchidium Haeckel (1887)
Stephanastrum Ehrenberg (1846)
Stylactis Ehrenberg (1873a)
Stylocyclia Ehrenberg (1847)
Stylodictya Ehrenberg (1846)
Stylosphaera Ehrenberg (1846)
Theocoronium Haeckel (1887)
Theocorys Haeckel (1882)
Theocotylissa Foreman (1973a)
Theocyrtis Haeckel (1887)
Tholodiscus Petrushevskaya and Kozlova (1972)
Thyrsocyrtis Ehrenberg (1847)
Triospyrium Haeckel (1887)
Xiphatractylis Haeckel (1887).

Table 1.

Pterocanium proserpinae Ehrenberg (1858)
Pterocodon campana Ehrenberg (1847)
Pterocanium barbadense Ehrenberg (1874)
Pylosphaera mediterranea Ehrenberg (1858)
Rhabdolithis pipa Ehrenberg (1854c)
Haliomma lagena Ehrenberg (1840)
Rhopalocanium ornatum Ehrenberg (1847)
Rhopalodictyum abyssorum Ehrenberg (1873a)
Schizomma quadrilobum Ehrenberg (1862b)
Cornutella scalaris Ehrenberg (1874)
Eucyrtidium trochus Ehrenberg (1873a)
Haliomma radians Ehrenberg (1841)
Haliomma radiatum Ehrenberg (1854c)
Cornutella scalaris Ehrenberg (1874)
Disolenia follis [nomen nudum] Ehrenberg (1873b)
Spirillina imperforata Ehrenberg (1858)
Spongaster tetras Ehrenberg (1862b)
Spongosphaera pachystyla Ehrenberg (1874)
Spongodiscus resurgens Ehrenberg (1854b)
Spongosphaera pachystyla Ehrenberg (1874)
Spongosphaera polyacantha Müller (1858)
Haliomma perspicuum Ehrenberg (1874)
Stephanastrum rhombus Ehrenberg (1847)
Stylactis pacifica Ehrenberg (1873a)
Stylocyclia dimidiata Ehrenberg (1874)
Stylodictya gracilis Ehrenberg (1854c)
Stylosphaera hispida Ehrenberg (1854b)
Eucyrtidium turgidulum Ehrenberg (1873a)
Eucyrtidium turgidulum Ehrenberg (1873a)
Eucyrtidium ficus Ehrenberg (1874)
Eucyrtidium barbadense Ehrenberg (1874)
Stylodictya ocellata Ehrenberg (1874)
Thyrsocyrtis rhizodon Ehrenberg (1874)
Ceratospyris furcata Ehrenberg (1874)
Stylosphaera spinulosa Ehrenberg (1874)

Type species
Campbell (1954)
Ehrenberg (1847)
Petrushevskaya and Kozlova (1972)
Ehrenberg (1858)
Sanfilippo and Riedel (1973)
Strelkov and Reshetnyak (1959)
Campbell (1954)
Campbell (1954)
Ehrenberg (1862b)
Campbell (1954)
Petrushevskaya and Kozlova (1972)
Campbell (1954)
Kozur and Mostler (1979)
Petrushevskaya and Kozlova (1972)
Riedel (1971)
Ehrenberg (1858)
Ehrenberg (1862b)
Sanfilippo and Riedel (1973)
Frizzell and Middour (1951)
Campbell (1954)
Müller (1858)
Campbell (1954)
Campbell (1954)
Ehrenberg (1873a)
Campbell (1954)
Campbell (1954)
Campbell (1954)
Frizzell and Middour (1951)
Frizzell and Middour (1951)
Foreman (1973a)
Campbell (1954)
Petrushevskaya and Kozlova (1972)
Campbell (1954)
Campbell (1954)
Campbell (1954)

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Doubtfully included species (ICZN 2000 Art 67.2.5): A nominal species is deemed not tobe originally included if it was doubtfully or conditionally included.
Type fixation overlooked (ICZN 2000 Art 70.2): If it is found that an earlier type species
fixation has been overlooked, the overlooked fixation is to be accepted and any later
fixations are invalid.
Under ICZN Art 67.2.5 and Art 70.2, Cycladophora (?) davisiana Ehrenberg 1862b (typespecies of the genus Cycladophora. Subsequent designation by Lombari and Lazarus, 1988) andLithomelissa (?) tartari Ehrenberg 1854b (the type species of the genus Lithomelissa, subsequent
designation by Campbell, 1954) need consideration. Nomen nudum species are also designatedas type species, e.g. of Cryptoprora Ehrenberg 1847, Dictyospyris Ehrenberg 1846, Dipospyris
Haeckel 1887, Disolenia Ehrenberg 1861b, Haliphormis Ehrenberg 1846 and Solenosphaera
Haeckel 1887. Except for Dictyospyris, no genus has replaced [none of these genera has been
provided with a valid] type species. This issue also needs discussion by the radiolarian community.
A total 12 species are designated as the type species of more than two genera, but it is acceptable by the following rule in ICZN 2000.
Nominal species that are already type species (ICZN 2000, Art 67.11): The fact that anominal species is the type species of a nominal genus or subgenus does not prevent itfrom being the type species of another. In such a case, the genus-group names are objective synonyms of one another.
Based on the ICZN 2000 Art 67.11, the following combinations are objective synonyms. The
underlined genus is the junior objective synonym when we rigidly follow ICZN 2000.
Anthocyrtis Ehrenberg 1846 and Anthocyrtella Haeckel 1887 (type species: Anthocyrtis
mespilus Ehrenberg 1847),
Conarachnium Haeckel 1882 and Sethoconus Haeckel 1887 (type species: Eucyrtidium
trochus Ehrenberg 1873a),
Dictyocephalus Ehrenberg 1861b and Dictyocryphalus Haeckel 1887 (type species: Cornutella obtusa Ehrenberg 1844a),
Eucyrtidium Ehrenberg 1846 and Eucyrtis Haeckel 1882 (type species: Lithocampe acuminata Ehrenberg 1844a),
Litharachnium Haeckel 1861b and Litharachnidium Haeckel 1887 (type species: Carpocanium arachnodiscus Ehrenberg 1862b),
Lithomelissa Ehrenberg 1847 and Acromelissa Haeckel 1887 (type species: Lithomelissa
microptera Ehrenberg 1854b),
Polysolenia Haeckel 1882 and Acrosphaera Haeckel 1882 (type species: Polysolenia setosa
Ehrenberg 1873a),
Sethopyramis Haeckel 1882 and Sestropyramis Haeckel 1887 (type species: Cornutella
scalaris Ehrenberg 1874),
Spongosphaera Ehrenberg 1847 and Spongatractus Haeckel 1887 (type species: Spongosphaera pachystyla Ehrenberg 1874), andTheocorys Haeckel 1882 and Theocoronium Haeckel 1887 (type species: Eucyrtidium
turgidulum Ehrenberg 1873a),

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Furthermore, the genera Conarachnium, Eucyrtis, Spongosphaera and Theocorys have othertype species (Sethoconus rayianus Haeckel 1887 for Conarachnium by subsequent designation ofCaulet, 1971; Eucyrtis conoidea Rüst 1885 for Eucyrtis by subsequent designation of Foreman,
1973b; Spongosphaera polyacantha Müller 1858 by subsequent monotype; and Theocorys
morchellula Rüst 1885 by subsequent designation of Campbell 1954 and Theocorys veneris
Haeckel 1887 by subsequent designation of Strelkov and Reshetnyak 1959 for Theocorys). Typespecies issues have been resolved for only a few of these genera (e.g. Spongosphaera and Eucyrtidium). Although we don’t fix these taxonomic issues in our paper, proper type species should beselected in consideration of not only the precise application of ICZN 2000 but also consensus in
the radiolarian community.
Concluding remarks
Ehrenberg described a total of 72 genera and 501 species, excluding 31 nomen nudum
species in his life. He described 544 species in Ehrenberg (1873a, 1874), but it includes species
that were described by other authors. The precise number of the species described by Ehrenberg
is variably estimated in papers (530 new species in Tanimura et al, 2006; about 500 species at p.
clxxiv in Haeckel, 1887), which is probably due to presence of nomen nudum. Ehrenberg’s contribution to the study of polycystine radiolarians also include proposals for a higher classification(e.g. Ehrenberg, 1847, 1876) and the first reports to stress their geographical and geological distributions (e.g. Ehrenberg, 1874, 1876). Ehrenberg also commented the taxonomic position of
Polycystina and probable physiological functions, but these comments were strongly criticized inother studies (e.g. Haeckel, 1862, 1887). Haeckel (1887, page clxxiii to clxxv) summarized hisview of problems in Ehrenberg’s studies:
1. Ehrenberg ignored any contrary observations and remarks, specifically those regarding
living radiolarians.
2. Ehrenberg often changed the taxonomic of position of Polycystina between Echinoderma, Bryozoa and Oscillaria.
3. Ehrenberg believed that Polycystina were multi-cellular animals and that the pores of thesiliceous test were comparable to cells, disregarding the extensive detailed observationsof living radiolarians by Haeckel, Huxley and Müller.
4. Ehrenberg’s diagnoses for genera were inadequate.
Although Haeckel (1887) criticized Ehrenberg using rather emotional words (e.g. strange, remained entirely ignorant, the invincible obstinacy, his preconceived opinion, etc.), Haeckel recognized the accuracy of Ehrenberg’s sketches (p. clxxv in Haeckel, 1887). Since the species published by Ehrenberg have been extensively used up to the modern day, they are still importanttaxonomic concepts. His simple sketches are often insufficient for modern taxonomic work, andsubsequently re-examination of the original specimens of Ehrenberg’s species is necessary.

References
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Sci. Arts Ser. 2, 22 (64): 1–6.

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Bütschli, O., 1882. Beiträge zur Kenntnis des Radiolarienskelette, insbesondere der der Cyrtida. Zeit. Wiss. Zool., 36:
485–540.
Campbell, A. S., 1951. New genera and subgenera of Radiolaria. Jour. Paleont., 25: 527–530.
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Ehrenberg, C. G., 1840. Hierauf 274 Blätter von ihm selbst ausgeführter Zeichnungen von eben so vielen Arten in dem
1838 erschienenen grösseren Infusorienwerke noch nicht abgebildeter Infusorien. Ber. Königl. Preuß. Akad. Wiss.
Berlin, 1840: 197–219.
Ehrenberg, C. G., 1841. Über noch jetzt zahlreich lebende Thierarten der Kreidebildung und den Organismus der Polythalamien. Abh. Königl. Akad. Wiss. Berlin, 1839: 81–174.
Ehrenberg, C. G., 1842. Über einen plastischen Kreidemergel von Ägina aus mikroskopischen Organismen und über die
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Kunstdenkmäler aus gebrannter Erde (Terracotten) mit bisher unbekannter Sicherheit zu bestimmen. Ber. Königl.
Preuß. Akad. Wiss. Berlin, 1842: 263–268.
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Akad. Wiss. Berlin, 1844: 57–97.
Ehrenberg, C. G., 1844b. Einige vorläufige Resultate seiner Untersuchungen der ihm von der Südpolreise des CaptainRoss, so wie von den Herren Schayer und Darwin zugekommenen Materialien über das Verhalten des kleinsten
Lebens in den Oceanen und den grössten bisher zugänglichen Tiefen des Weltmeeres. Ber. Königl. Preuß. Akad.
Wiss. Berlin, 1844: 182–207.
Ehrenberg, C. G., 1844c. Untersuchungen über die kleinsten Lebensformen im Quellenlande des Euphrats und Araxes, sowie über eine an neuen Formen sehr reiche marine Tripelbildung von den Bermuda-Inseln. Ber. Königl. Preuß.
Akad. Wiss. Berlin, 1844: 253–275.
Ehrenberg, C. G., 1845. Neue Untersuchungen über das kleinste Leben als geologisches Moment. Ber. Königl. Preuß.
Akad. Wiss. Berlin, 1845: 53–87.
Ehrenberg, C. G., 1846. Über eine halibiolithische, von Herrn R. Schomburgk entdeckte, vorherrschend aus mikroskopischen Polycystinen gebildete, Gebirgsmasse von Barbados. Ber. Königl. Preuß. Akad. Wiss. Berlin, 1846: 382–385.
Ehrenberg, C. G., 1847. Über die mikroskopischen kieselschaligen Polycystinen als mächtige Gebirgsmasse von Barbados und über das Verhältniss der aus mehr als 300 neuen Arten bestehenden ganz eigenthümlichen Formengruppe
jener Felsmasse zu den jetzt lebenden Thieren und zur Kreidebildung. Eine neue Anregung zur Erforschung desErdlebens. Ber. Königl. Preuß. Akad. Wiss. Berlin, 1847: 40–60.
Ehrenberg, C. G., 1854a. Über das organische Leben des Meeresgrundes in bis 10800 und 12000 Fuss Tiefe. Ber. Königl.
Preuß. Akad. Wiss. Berlin, 1854: 54–75.
Ehrenberg, C. G., 1854b. Systematische Charakteristik der neuen mikroskopischen Organismen des tiefen atlantischen
Oceans. Ber. Königl. Preuß. Akad. Wiss. Berlin, 1854: 236–250.
Ehrenberg, C. G., 1854c. Mikrogeologie. Voss, Leipzig, 374 pp.
Ehrenberg, C. G., 1855a. Über die weitere Entwicklung der Kenntniss des Grünsandes als grüner PolythalamienSteinkerne, über braunrothe und corallrothe Steinkerne der Polythalamien-Kreide in Nord-Amerika, und über den
Meeresgrund aus 12,900 Fuss Tiefe. Ber. Königl. Preuß. Akad. Wiss. Berlin, 1855: 172–178.
Ehrenberg, C. G., 1855b. Über ein europäisches marines Polygastern-Lager und über verlarvte Polythalamien in den
marinen Polygastern Tripeln von Virginien und Simbirsk. Ber. Königl. Preuß. Akad. Wiss. Berlin, 1855: 292–305.
Ehrenberg, C. G., 1855c. III. Nähere Bestimmung der Mischung des frischen Auswurfs des Schlamm-Vulkans von Poorwadadi auf Java. Ber. Königl. Preuß. Akad. Wiss. Berlin, 1855: 570–576.
Ehrenberg, C. G., 1856. Über 2 neue südamerikanische Gebirgsmassen aus mikroskopischen Organismen, eine aus
Meeresorganismen in Chile und eine als mit gefritteten Süsswasserorganismen gemischten vulkanischen essbaren

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Tuff aus Honduras in Centro-Amerika. Monat. Königl. Preuß. Akad. Wiss. Berlin, 1856: 425–431.
Ehrenberg, C. G., 1858. Über die organischen Lebensformen in unerwartet grossen Tiefen des Mittelmeeres. Monat.
Königl. Preuß. Akad. Wiss. Berlin, 1857: 538–570.
Ehrenberg, C. G., 1859. Kurze Characteristik der 9 neuen Genera und der 105 neuen Species des ägäischen Meeres und
des Tiefgrundes des Mittel-Meeres. Monat. Königl. Preuß. Akad. Wiss. Berlin, 1858: 10–40.
Ehrenberg, C. G., 1861a. Über die organischen und unorganischen Mischungsverhältnisse des Meeresgrundes in 19800
Fuss Tiefe nach Lieut. Brookes Messung. Monat. Königl. Preuß. Akad. Wiss. Berlin, 1860: 765–774.
Ehrenberg, C. G., 1861b. Über den Tiefgrund des stillen Oceans zwischen Californien und den Sandwich-Inseln aus bis
15600' Tiefe nach Lieut. Brooke. Monat. Königl. Preuß. Akad. Wiss. Berlin, 1860: 819–833.
Ehrenberg, C. G., 1862a. Beitrag zur Übersicht der Elemente des tiefen Meeresgrundes im Mexikanischen Golfstrome
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