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. 87–96. National Museum of Nature andScience Monographs, No. 40, Tokyo 2009

Reexamination of Ehrenberg’s Neogene Radiolarian Collections and
its Impact on Taxonomic Stability
Noritoshi Suzuki1, Kaoru Ogane1, Yoshiaki Aita2, Toyosaburo Sakai2 and David Lazarus3
1

Institute of Geology and Paleontology, Graduate School of Science, Tohoku University,
6–3 Aoba Aramaki, Aoba-ku, Sendai 980–8578, Japan2
Department of Geology, Faculty of Agriculture, Utsunomiya University, Utsunomiya 321–8505, Japan
3
Museum für Naturkunde, Humboldt University, Invalidenstrasse 43, 10115 Berlin, Germany
Corresponding Author: Noritoshi Suzuki (e-mail: suzuki.noritoshi@nifty.com)

Abstract All remaining Neogene polycystine radiolarian specimens which were examined by
Ehrenberg were searched for in the Ehrenberg Collection, Museum für Naturkunde, Humboldt University, Berlin, Germany. Almost all the illustrated specimens in his publications were located byusing published and unpublished source materials. These specimens are candidate lectotypes. However, rigid application of the International Code of Zoological Nomenclature (ICZN) would in somecase causes scientifically unfruitful confusion and dispute because of very different modern acceptedusage of several generic names (i.e. Acanthosphaera, Cenosphaera, Dictyophimus and Pterocanium).
For these alternate type species may be required. Some illustrated specimens in publications are also
broken by cracks in the embedding Canada balsam (i.e. Podocyrtis aegles) or are poorly preserved(i.e. Haliomma hexagonum) so that alternate type specimens may be required. For these reasons, we
do not formally designate types in our paper, only pointing out taxonomic problems where appropriate.
Key words:

Classification, Ehrenberg, Legacy Collection, Radiolaria, Taxonomy.

Introduction
Description of Neogene fossil polycystine radiolarians was initiated by Ehrenberg (1839),
and subsequent description of Neogene polycystines, including living ones, were actively carriedout by Ehrenberg until his death in 1876 (Suzuki, 2009). Prior to 1876, only a few taxonomists
described new polycystine radiolarian species (Müller, 1855a, 1855b, 1857, 1858a, 1858b; Bailey, 1856; Haeckel, 1860a, 1860a, 1862; Harting, 1863). Many species described by Ehrenberg
have still been used as valid species in current papers although Ehrenberg’s often simple drawings, ambiguous descriptions, and unspecified type materials are often noted as problematic bysubsequent workers. The purpose of our paper is to reillustrate all polycystine specimens which
Ehrenberg documented as having examined as the basis for his species descriptions, as these will
presumably be used in subsequent studies to set neotypes. Additionally we document some important species for age determination of the samples, and briefly clarify points for some species.
The detailed procedure how to find specimens examined by Ehrenberg himself is documented in
Lazarus (1998), Lazarus and Jahn (1998) and Suzuki et al. (2009).

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Results
Ehrenberg (1873b) summarized the occurrences of his previously described Neogene polycystine radiolarians, and we examined all the mica strips from the localities listed in this publication that could still be found during our study. We also included material from Bermuda, Caltanisetta and Camorta, which were listed as fossil localities in Ehrenberg (1876). Neogene polycystine radiolarian specimens were found and digitally captured from the locations: Südpol
(South Pole; mica tray’s ID: K01B01), Camorta (Nicobar Islands; K08B10), Mexikanischer Golfstrom (Mexican Gulfstream; K16B08 and K16B09), Phillipinischer Ozean (Philippine Sea;
K09B06), Oran (Algeria; K11B05), Bermuda I (K15B04), Bermuda II (K15B05), Californischer
Stiller Ozean (Pacific Ocean off California; K25B07 and K25B08), Meeresgrund II (“ocean bottom sediments in the Atlantic Ocean”; K27B02), Meeresgrund I (“ocean bottom sediments in the
Atlantic Ocean”; K27B03), Davisstraße I (Davis Straits; K27B17), Davisstraße II (K27B16),
Caltanisetta (Sicily; K28B06), Zante (Greece; K31B07), Aegina (Greece; K31B09), Meeresgrund Mittelmeer (Mediterranean ocean bottom sediments; K32B08) and Zanguebar 13200’(MB ES 912). This last is not a mica strip but a glass slide stored with the cited Ehrenberg collection sample. Although details of the geologic ages of the examined samples will be publishedin a separate paper, the sea-bottom sediment samples are probably Quaternary or Holocene.
These include the mica strips labeled Philippinischer Ozean, Californischer Ozean, Meersgrund I
and II, Davisstraße I and II, Meeresgrund Mittelmeer, and Zanguebarica. Miocene or younger
polycystines were observed samples from land section, i.e. in Caltanisetta, Zante and Aegina.
Caltanisetta (K28B06): Plates 1 to 21
As the first illustrated fossil polycystine ever published is Lithocampe radicula from Caltanisetta (Ehrenberg, 1839), Caltanisetta is a sample with particular historic value for radiolarianstudies. The new species found from Caltanissetta were described in Ehrenberg (1839, 1840,
1844a) and Ehrenberg (1854b). Ehrenberg (1873b) included the species names from Caltanisetta
in his subsequent geographical and vertical distribution tables of recent polycystine radiolarians,
although most of these species are not found in sea-bottom sediments. In his comprehensive
monograph of fossil polycystines (Ehrenberg, 1876) he noted that the Caltanisetta specimens
were Tertiary fossils.
Ehrenberg’s mica strips are now between 140–170 years old. As noted previously (Lazarus,
1998; Lazarus and Jahn, 1998) they are brittle and sometimes were damaged due to improper
storage and use in the 1970s and 1980s, as well as fire damage in the mid 1980s. The worst preserved strips encountered in this study were those from Caltanisetta. Several of the micas havebecome detached from their mica strips. Many of the micas are cracked, and Ehrenberg did notrecord the taxon name on the strip labels in the mica tray. The paper rings marking specimen locations have also partially detached from the micas. To safeguard these materials for the future,
we looked through all the micas and recorded all the polycystine specimens which were encountered. Our monograph illustrates all the specimens from Caltanisetta, and thus provides a virtual
copy of the radiolarian content which can largely replace further examination of this material forradiolarian studies.
Lithocampe radicula Ehrenberg 1839, the first illustrated fossil polycystine specimen in thehistory of science, is shown in this study on pl. 13, figs. 7a–7c. This form was first regarded as aforaminifera. The sketch was simple, and identifying the actual specimens required considerationof the probable inclination of the specimen used for Ehrenberg’s drawing (see Suzuki et al.,Ehrenberg’s Neogene Radiolarian Collections

892009). Ehrenberg’s sketch (pl. 13, fig. 7c in our paper) resembles the specimen illustrated in pl.
13, figs. 8a–8b, but the former differs from the latter by the former possessing three to four longitudinal pores in a segment, and lacking a tube in the final segment. Other sketches reflect the actual specimens reasonably well: Perichlamydium limbatum (Ehrenberg 1844a) (pl. 8, figs. 1a–1d)
and Perichlamydium praetexta (Ehrenberg 1844a) (pl. 8, figs. 3a–3d), for example.
Several micas could not be located in the Caltanisetta mica trays, and thus several important
species were not found in our examination. Eucyrtidium auritum (Ehrenberg 1844a) (pl. 19, fig. 9
in our paper), for example, is missing because the indicated mica strip is missing. We can only
find a single specimen (pl. 19, figs. 10a–10b) which is assignable to this species from all the remaining mica strips in Caltanisetta.
Zante (K31B07): Plates 22 and 23
A simple species list of polycystines from “Zante” was initially published in Ehrenberg(1839), while the new species from Zante were described in Ehrenberg (1844a) and illustrated in
Ehrenberg (1854b). Zante contains a few, poorly preserved polycystines, so that we looked at all
the mica strips to find and record all the polycystines as digital images; however we only found
18 specimens. Cyrtocapsella tetrapera Haeckel 1887 were found in the Zante material (pl. 23,
figs. 1a–2b), suggesting an Early to Middle Miocene age at least. A single specimen of Flustrella
concentrica Ehrenberg 1844a (published in pl. 20, fig. 42 of Ehrenberg (1854b) (pl. 22, figs.
3a–3c in our paper) was found.
Aegina (K31B09): Plates 24 to 26
New species from Aegina were described in Ehrenberg (1839, 1840, 1859), and illustrated in
Ehrenberg (1854b). As with Zante, all specimens encountered in the Aegina micas were digitally
recorded and are illustrated in our paper. The depositional age of Aegina is not determined yet
due to lack of age-diagnostic polycystines. The first spiral polycystine probably belonging to themodern genus concept Lithelius was described as Flustrella spiralis Ehrenberg 1840, but the illustrated specimen (pl. 24, figs. 4a–4c) is not well preserved. Unpublished Ehrenberg’s drawing
ID 1563 illustrated “Pterocanium graecum” but it is a nomen nudum (pl. 24, figs. 4a–4d). Eucyrtidium seriolatum was formally described in Ehrenberg (1859), but has never been illustrated. We
show for the first time in a publication the original drawing and photographs of the specimen inpl. 24, figs. 5a–5c.
The type species of the genus Lithornithium Ehrenberg 1847 was subsequently designated as
Lithocampe hirundo Ehrenberg 1840 by Campbell (1954). The specimen of this species illustrated in our paper is shown on pl. 26, figs. 1a–1f. This species probably belongs to the modern
genus concept Lychnocanoma, but the specimen is (originally) broken. Furthermore, Ehrenberg’s
concept of Lithocampe hirundo also included “Pterocanium praetextum” (pl. 26, figs. 3a–3c).
Südpol I; 78°10S.B. 162 W.L. (K01B01): Plates 27 and 28
New species from the sample Südpol I were described in Ehrenberg (1844b) and illustrated
in Ehrenberg (1854b). All polycystine specimens which were indicated by handwriting on themica tray or in the Ehrenberg’s drawing were searched for and illustrated in our paper. This sample is largely dominated by the genus Antarctissa, and other species are rare. Ehrenberg indicatedthe location of specimens of “Lithobotrys denticulata Ehrenberg 1844b” (Antarctissa denticulata (Ehrenberg)) where a few specimens are indeed present within the color paper ring of themica strip.

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Bermuda I (K15B04), Bermuda II (K15B05) and Oran (K11B05): Plate 29
Bermuda I, II and Oran were regarded as Tertiary fossil material in the comprehensive monograph of Ehrenberg (1876). The new species from Bermuda were described in Ehrenberg (1844c,
1854b). The specimen from Oran was illustrated in Ehrenberg (1854b).
The mica trays in the Ehrenberg collection are separated into Bermuda I and II, but they arederived from a single sample, based on the handwriting on the mica tray. Astromma entomocora
Ehrenberg 1854b (pl. 29, figs. 1a–1c in our paper) and Pterocanium aculeatum (Ehrenberg
1844c) (pl. 29, figs. 3a–4d) were first described from the Bermuda sample, but they have neverbeen illustrated in the literature. Pterocanium aculeatum is the type species of the genus Pterocanium, and is apparently different from the current usage of the genus Pterocanium (i.e. Pterocanium praetextum (Ehrenberg)). A community consensus will be needed to accept, or to reject thisspecimen for stability of taxonomy. Astromma entomocora is also the type species of the genus
Astromma by the subsequent designation of Campbell (1954), and this species belongs to thesubfamily Artiscinae. Haliomma nobile Ehrenberg 1844c appeared to be from Barbados because
Ehrenberg (1876) illustrated the form with Barbados species, but we confirmed that the illustrated specimen is actually in the mica strip Bermuda I.
A single specimen was found from Oran representing Lithocampe lineata Ehrenberg 1839
(pl. 29, figs. 8a–8c in our paper). An illustration of this specimen was published on pl. 21, fig.
56b in Ehrenberg (1854b).
Meeresgrund II (K27B02): Plate 30
Mica tray K27B02 holds mica strips from several samples. Of these, the sample “Azores Islands 6480” was the only one used for radiolarian studies by Ehrenberg. The new species from
Azores Island 6480’ were described in Ehrenberg (1854a) and illustrated in Ehrenberg (1854b).
This sample contains the type species of Cenosphaera, namely Cenosphaera plutonis Ehrenberg
1854a (our paper: pl. 30, figs. 1a–1d). As shown in these photographs, this specimen has three
concentric shells, and is probably the same as the genus Thecosphaera. Flustrella subtilis Ehrenberg 1854a was formally described by Ehrenberg (1854a), but is first illustrated in our paper on
pl. 30, figs. 3a–3c.
Meeresgrund II (K27B03): Plates 31 to 36
This tray (K27B03) also contains micas derived from several samples. Polycystine radiolarians were examined by Ehrenberg from the samples “Atlantic Ocean 10800’, Bay of Biscay”(plates 31 to 34 in our paper) and “Nord Ocean 12000” (plates 35 and 36). Many species wererecorded on the labels of the mica tray, although only a few specimens were illustrated and published. All the specimens recorded on the mica trays for these two samples were searched for.
Ehrenberg (1854a) described 9 new species from the sample “Atlantic Ocean 10800’ ” while
Ehrenberg (1854a, 1854b) described six new species from “Nord Ocean 12000’.” The typespecies of the genus Dictyophimus, Dictyophimus crisiae, is first illustrated on pl. 35, figs. 1–1din our paper. Several species described in Ehrenberg (1854a) have not been illustrated in any
prior publications, and several of these species (Haliomma quadruplex on pl. 31, figs. 1a–1d;
Eucyrtidium heteroporum on pl. 32, figs. 2a–2d; Eucyrtidium galatheae on pl. 35, figs. 5a–5c)
appear to be different species from the modern usage of these names. Both the “Atlantic Ocean
10800’ ” and “Nord Ocean 12000’ ” samples generally yield broken polycystines (pl. 31, figs.
3a–3b), cracked specimens (pl. 31, figs. 8a–8c, pl. 32, figs. 9a–9c) and poorly preserved specimens (pl. 32, figs. 11a–11d, pl. 35, figs. 1a–1d, pl. 36, figs. 2a–2d): these specimens are probablyEhrenberg’s Neogene Radiolarian Collections

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inadequate for the designation of lectotypes or neotypes. New radiolarians slides should be madefrom the type materials or topotypic materials.
Meeresgrund Mittelmeer (K32B08): Plate 37
Ehrenberg (1854b, 1859) described seven new species from this sample. The precise skeletal
morphology of Pylosphaera mediterranea Ehrenberg 1859 could not be determined due to ahighly obliquely oriented specimen (pl. 37, figs. 2a–2d). Eucyrtidium creticum Ehrenberg 1859
looks very different appearance between the illustration (pl. 37, fig. 7a) and the actual specimen(pl. 37, figs. 7b–7d), but we are confident that this specimen is correctly identified as it occurs exactly where indicated by Ehrenberg- mica ring 320801.d.v.
Camorta (K08B10): Plate 38
Based on Ehrenberg’s handwritten mica tray labels, he examined polycystine radiolarians andspherical siliceous sponge remains from the samples “Camorta XIII” (pl. 38, figs. 1a–1b, 5a–5b,
9a–11b), “Camorta XIX” (pl. 38, figs. 4a–4b, 6a–6b, 8a–8b, 12a–13b, 20a–20b), and “Camorta
Nicobarien 18” (pl. 38, figs. 2a–3b, 7a–7b, 15a–16b, 19a–19b). The species names from Camorta
were first published in Ehrenberg (1876), the year of his death, and four species names (Spongodiscus argulus, Haliomma argulus, Eucyrtidium arugulus and Diauletes nicobarium) arenomen nudum.
Mexikanischer Golfstrom (K16B08 and K16B09): Plate 39
Ehrenberg examined polycystine radiolarians and spherical siliceous sponge remains fromthe samples “Golfstrom 9066” (pl. 39, figs. 1a–1b, 4a–4b, 6a–9d), “Golfstrom 2556” (pl. 39, figs.
2a–2b), “Golfstrom 1158” (pl. 39, figs. 3a–3b, 5a–5b) and “Golfstrom 840” (pl. 39, figs.
10a–11d). Three new species (Dictyocephalus pyrum on pl. 39, figs. 9a–9d; Rhopalodictyum
truncatum on pl. 39, figs. 10a–10b; Dictyocephalus hispidus on pl. 39, figs. 11a–11d) were described from Golfstrom 9066 and 840 by Ehrenberg (1862).
Davisstraße I (K27B17): Plates 40–43 and Davisstraße II (K27B16): Plates 44–45
New species from Davisstraße were described in Ehrenberg (1862) and illustrated in Ehrenberg (1873b). Polycystine radiolarians described from “Davisstraße ” include several oceanographic marker species (e.g. Cycladophora davisiana, pl. 42, figs. 9a–9d; Eucyrtidium lineatum
arachneum, pl. 44, figs. 3a–3d). These two trays hold 36 mica strips from many samples, andsample labeling is very complex. Worse, the location information of the specimens shown in the
Ehrenberg’s drawings is wrong for many of the mica strips in these trays in comparison with thespecimens morphologically matched to them from examination of the mica contents.
Polycystine radiolarians were examined from samples recovered from water depths of 600
feet, 9240 feet (1540 fathoms), 10998 feet (1833 fathoms) and 11040 feet (1840 fathoms).
The sample from 9240 feet was obtained at 62°06N, 52°21W on 18th October 1859, and thatfrom 10998 feet at 59°12N, 52°38W on 20th September 1859. The 9204 feet sample is further
subdivided into three parts, noted by codes, namely “A.b”, “B.a”, “B.b”, while the 10998 feetsample is subdivided into A and B. The 6000 feet sample is also subdivided into several parts, including b .II. Although we have not yet resolved the interpretation of most of these subdivided
sample codes, we note that the original location ID in Ehrenberg’s drawings is separately numbered for each subdivision. Flustrella (?) haliomma was described by Ehrenberg (1839), but onlya single drawing of a Davisstraße specimen is available for this species. This specimen is not the92

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original specimen for the description of Fl. (?) haliomma because the Davisstraße samples weretaken only in 1859, 20 years after the original description.
Philippinischer Ozean 19800 (K09B06): Plates 46–57
A total of 57 Neogene new species described by Ehrenberg were illustrated from the Philippinischer Ozean 19800’ sample. The descriptions of the new species were published in Ehrenberg (1873a) and the illustrations of them were included in Ehrenberg (1873b). The published illustrations of some species (Ehrenberg, 1873b) show some ambiguous morphological characters.
The drawing of Haliomma octacanthum Ehrenberg 1873a (pl. 48, fig. 3a in our paper) does notclearly indicate if the species is flat or spherical in shape: the photograph of this specimen clearlyshows a discoidal polycystine radiolarian, probably belonging to Heliodiscus. It has previously
been difficult to decide whether Acanthosphaera elliptica Ehrenberg 1873a (pl. 51, figs.
10a–10c) is a spherical siliceous sponge spicule or a polycystine radiolarian. Based on our examination, this species is a polycystine, probably related to the genus Lithelius or Larcopyle. The
drawing of Stylosphaera testudo Ehrenberg 1873b (pl. 51, figs. 12a–12c) showed a vertical structure inside the cortical shell, but it seems to be lacking in the actual specimen. The drawing of
Stylosphaera holosphaera Ehrenberg 1873a (pl. 53, figs. 2a–2c) was previously not regarded as
belonging to this genus. The actual specimen in fact has a saturnalid ring, and is probably conspecific with Saturnalis circularis Haeckel 1887 under modern usage.
Owing to a thin layer of embedding Canada balsam, the morphological characters of Stylosphaera megadictya Ehrenberg 1873a (pl. 53, figs. 3a–3d), Ceratospyris aculeata Ehrenberg
1873a (pl. 53, figs. 4a–4b) and Eucyrtidium cassis Ehrenberg 1873a (pl. 53, figs. 6a–6c) cannot
be recognized from the actual specimens. Ehrenberg created colorful drawings for several ofthese species as they were living specimens. Ommatospyris penicillata Ehrenberg 1873a (pl. 52,
figs. 6a–6c), Oms. profunda (pl. 52, figs. 7a–7c), Stylosphaera setosa Ehrenberg 1873a (pl. 52,
figs. 12a–12c), and Eucyrtidium pleuracanthus Ehrenberg 1873a (pl. 54, figs. 7a–7c) have yellowish matter within them that seems to be like protoplasmic intracapsulum, but this material appears to be aggregated clay minerals.
Californischer Stiller Ozean (K25B07 and K25B08): Plate 58–67
All the polycystine radiolarians which were recorded by Ehrenberg were digitally captured
and are shown on Plates 58 to 67 in our paper. Ogane et al. (2009) pointed out that Ehrenberg erroneously mixed the location of his drawing specimens between Philippinischer Ozean 19800
and Californischer Stiller Ozean. These two samples yield so many common species that the confusion might easily have happened. Our careful comparisons between drawings and actual specimens lead us to conclude that only eight species were illustrated from Californischer Stiller
Ozean. The details of Euchitonia furcata Ehrenberg 1873a (pl. 60, figs. 4a–4b) were alreadycommented in Ogane et al. (2009). Ommatocampe polyarthra Ehrenberg 1873a (pl. 63, figs.
2a–2d) seems to have a straight cylindrical form, but we cannot discriminate this species fromthe lateral view of three-armed discoidal species (e.g. pl. 62, figs. 4a–4c).
Zanguebar 13200 (MB ES 942): Plates 68–77
Zanguebar 13200 (database ID: MB ES 912) is also an important sample in that a total of 34
species were described and illustrated from it (Ehrenberg 1873a, 1873b). Differing from his usual
mica strips, “MB ES 912” is uniquely for his radiolarian specimens, a glass slide. Ehrenberg’s
drawings indicate the location of the specimens and the color of the paper rings that are attachedEhrenberg’s Neogene Radiolarian Collections

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on the cover slip of the slide. However, it was impossible to locate all of the illustrated specimens
only based on these indications. We therefore examined all the polycystine specimens embeddedon the slide MB ES 912. It is fortunate that only one specimen was found for many species,
while several broken specimens provided unique characteristics which made it possible to determine unambiguously which specimen had been illustrated by Ehrenberg. The quality of our photographs in plates 68–77 is better than any of our other plates made from mica strips because theflat glass cover slip of slide MB ES 912, mounted on the glass slide provides, like modern microscopic slides, a superior optical mount. The mounting medium is also more transparent than theuncovered Balsam present on the micas.
The drawing of Haliphormis hexacanthus Ehrenberg 1873a (pl. 69, figs. 1a–1d) was not useful to examine this species’ internal structure. The specimen of this species illustrated by Ehrenberg does not have any internal structure, but other, unfigured specimens that we think are conspecific possess three concentric shells (pl. 69, figs. 2a–2c), suggesting assignment to the genus
Hexacontium. Ceratospyris pentagona Ehrenberg 1873a (pl. 71, figs. 5a–5d) is often found in living and Quaternary samples, but the illustrated specimen is poorly preserved. We found Acanthosphaera (?) fenestrata Ehrenberg 1873a (pl. 71, figs. 8a–8c), but the classification of species
to modern taxonomy was not possible because the major part of this specimen was broken off.
Concluding Remarks
Many Neogene species that were described by Ehrenberg have been designated as the typespecies of genera (e.g. Campbell, 1954). The designation of Ehrenberg species as generitypes has
been frequently criticized or even rejected since the quality of Ehrenberg’s drawings are not always adequate and some of the species subsequently designated by Campbell (1954) were not illustrated. Our reexamination of the Neogene polycystine radiolarians in the Ehrenberg Collection
has the potential to clarify the precise meaning of these generic names. On the other hand, we
emphasize the hazards that a rigid application of the International Code of Zoological Nomenclature would bring, potentially adding confusion and damaging modern taxonomic usage. The solution of this problem requires, in addition to formal rulings of the commission, a consensus
among radiolarian specialists, and for this reason we do not in this paper propose any formal revisions to radiolarian taxonomy: Instead, we only comment on differences between modern usage
and the actual specimens.
One surprising result is to have found internal structure in the type species of the genera
Cenosphaera Ehrenberg 1854a and Acanthosphaera Ehrenberg 1862. These two genera havelong been considered to have nothing inside the cortical shell, but Cenosphaera plutonis (pl. 30,
figs. 1a–1d), the type species of Cenosphaera, apparently consists of three concentric shells, andconsequently is consistent with the current definition of the genus Thecosphaera Haeckel 1882.
The type species of Acanthosphaera is Acanthosphaera haliphormis Ehrenberg 1862 (pl. 42, figs.
1a–1c). The actual specimen of this species is so poorly preserved that the detailed structure ofthis specimen is unclear, but two concentric structures are likely to be present inside the test. This
species was recovered from the Davisstraße, and it is most likely a specimen of the modern taxon
Actinomma boreale Cleve or a related species. These morphological characters of Acs.
haliphormis are thus the same as the genus Actinomma Haeckel 1860a. If the Code’s rules wererigidly applied, all current Thecosphaera species would have to be moved to the ‘true’
Cenosphaera and the current Cenosphaera would be moved to the other genus. In this case, morethan 58 Thecosphaera species would move to Cenosphaera and more than 170 Cenosphaera

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species have to be transferred to the other genus. This change also would make many synonyms
between Cenosphaera and Thecosphaera, and subsequently a rigid application of ICZN 2000 is
not practical procedure: it would create, rather than help settle taxonomic confusion. Similar
problems will arise in Acanthosphaera, Astromma (type species: Astromma entomocora Ehrenberg 1854b. See pl. 29, figs. 1a–1c in our paper), Dictyophimus (type species: Dictyophimus
crisiae Ehrenberg 1854a. See pl. 35, figs. 1a–1d in our paper), Lithocampe (type species: Lithocampe radiculata. See pl. 12, figs. 7a–7c), and Pterocanium (type species: Pterocanium aculeatum Ehrenberg 1844c. See pl. 29, figs. 3a–4d in our paper).
A rigid application of ICZN 2000 also would produce unfruitful confusion for many otherwell known species names. Ehrenberg (1873a, 1873b) described and illustrated species which
now belong to the subfamily Artiscinae. Ommatospyris laevis Ehrenberg 1873a (pl. 52, figs.
4a–4c), Ommatospyris setosa Ehrenberg 1873a (pl. 52, figs. 5a–5c), Ommatospyris penicillata
Ehrenberg 1873a (pl. 52, figs. 6a–6c) and Ommatospyris profunda Ehrenberg 1873a (pl. 52, figs.
7a–7c) are comparable to ontogenetic stages of a well-known species, Didymocyrtis tetrathalamus tetrathalamus (Haeckel 1887). Similar problems will happen with Lit. radicula and Lithocampe siculum Ehrenberg 1859 (pl. 13, figs. 1a–1c). L. siculum is morphologically similar to Stichocorys peregrina (Riedel 1953). Sti. peregrina has long been cited in numerous papers in the
Deep Sea Drilling Project, Ocean Drilling Program and Integrated Ocean Drilling Program, andthe replacement of the species name is not scientifically useful.
Our reexamination has largely succeeded in identifying the published specimens from the
Ehrenberg Collection, which have potential to be used as lectotypes for Ehrenberg’s species. For
taxonomic stability, not all the discovered specimens are always appropriate to use as lectotypes.
Podocyrtis aegles Ehrenberg 1854a (pl. 32, figs. 9a–9c) and Rhopalastrum lagenosum (Ehrenberg 1840) (pl. 24, figs. 10a–10b) for example have been damaged—they are cracked along thecenter of the test. Poorly preserved specimens are also inappropriate as lectotypes in the following species: Dictyophimus crisiae (pl. 35, figs. 1a–1d), Haliomma hexagonum Ehrenberg 1854a(pl. 36, figs. 2a–2d), Acanthosphaera haliphormis Ehrenberg 1862 (pl. 42, figs. 1a–1c), Mazosphaera laevis Ehrenberg 1873a (pl. 51, figs. 2a–2b), Stylosphaera setosa Ehrenberg 1873a (pl.
52, figs. 12a–12c), Ceratospyris setosa Ehrenberg 1873a (pl. 52, figs. 1a–1c), Stylosphaera
megadictya Ehrenberg 1873a (pl. 53, figs. 3a–3d), Ceratospyris aculeata Ehrenberg 1873a (pl.
53, figs. 4a–4b) and Eucyrtidium cassis Ehrenberg 1873a (pl. 53, figs. 6a–6c). Several specimens
are fixed at a difficult orientation to use as a standard specimen for classification: Carpocanium
arachnodiscus Ehrenberg 1862 (pl. 41, figs. 6a–6e) and Halicalyptra hexathyris Ehrenberg 1862
(pl. 43, figs. 6a–6g).
We tentatively conclude that formal taxonomic fixation of Ehrenberg’s polycystine radiolarian names must be done carefully, and in a way so as not to bring scientifically unfruitful confusion. This will only be possible if for some taxa, taxonomic stability of common use be given priority over rigid application of ICZN 2000 rules.
References
Bailey, J. W., 1856. Notice of microscopic forms found in the soundings of the Sea of Kamchatka-with a plate. Amer. J.
Sci. Arts Ser. 2, 22: 1–6.
Campbell, A. S., 1954. Radiolaria. In R. C. Moore (ed.) Treatise on Invertebrate Paleontology, Part D Protista 3. Protozoa(Chiefly Radiolaria and Tintinnina), Geological Society of America and University of Kansas Press, Kansas, p.
D1–D163.
Ehrenberg, C. G., 1839. Über die Bildung der Kreidefelsen und des Kreidemergels durch unsichtbare Organismen. Abh.

Ehrenberg’s Neogene Radiolarian Collections

95

Königl. Akad. Wiss. Berlin, 1838: 59–147
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., 1844a. Über 2 neue Lager von Gebirgsmassen aus Infusorien als Meeres-Absatz in Nord-Amerika undeine Vergleichung derselben mit den organischen Kreide-Gebilden in Europa und Afrika. Ber. Königl. Preuß.
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., 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: 40–61.
Ehrenberg, C. G., 1854a. Systematische Charakteristik der neuen mikroskopischen Organismen des tiefen atlantischen
Oceans. Ber. Königl. Preuß. Akad. Wiss. Berlin, 1854: 236–250.
Ehrenberg, C. G., 1854b. Mikrogeologie. Voss, Leipzig, 374 pp.
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., 1862. Über die Tiefgrund-Verhältnisse des Oceans am Eingange der Davisstrasse und bei Island.
Monat. Königl. Preuß. Akad. Wiss. Berlin, 1861: 275–315.
Ehrenberg, C. G., 1873a. Mikrogeologische Studien als Zusammenfassung seiner Beobachtungen des kleinsten Lebens
der Meeres-Tiefgründe aller Zonen und dessen geologischen Einfluss. Monat. Königl. Preuß. Akad. Wiss. Berlin,
1872: 265–322.
Ehrenberg, C. G., 1873b. Mikrogeologische Studien über das kleinste Leben der Meeres-Tiefgründe aller Zonen unddessen geologischen Einfluss. Abh. Königl. Akad. Wiss. Berlin, 1872: 131–399.
Ehrenberg, C. G., 1876. Fortsetzung der mikrogeologischen Studien als Gesammt-Übersicht der mikroskopischen
Paläontologie gleichartig analysirter Gebirgsarten der Erde, mit specieller Rücksicht auf den Polycystinen-Mergel
von Barbados. Abh. Königl. Akad. Wiss. Berlin, 1875: 1–225.
Haeckel, E., 1860a. Über neue, lebende Radiolarien des Mittelmeeres und legte die dazu gehörigen Abbildungen. Monat.
Königl. Preuß. Akad. Wiss. Berlin, 1860: 794–817.
Haeckel, E., 1860b. Fernere Abbildungen und Diagnosen neuer Gattungen und Arten von lebenden Radiolarien des Mittelmeeres. Monat. Königl. Preuß. Akad. Wiss. Berlin, 1860: 835–845.
Haeckel, E., 1862. Die Radiolarien (Rhizopoda Radiolaria). Eine Monographie. Tafel 1, Reimer, Berlin, 572 pp.
Haeckel, E., 1882. Entwurf eines Radiolarien-Systems auf Grund von Studien der Challenger-Radiolarien. Jena. Zeit.
Naturwiss. Med. -Naturwiss. Gesell. Jena, 15: 418–472.
Haeckel, E., 1887. Report on the Radiolaria collected by H.M.S. Challenger during the years 1873–1876. Rep. Sci. Res.
Voyage H.M.S. Challenger 1873–1876, Zool., 18: 1–1803.
Harting, P., 1863. Bijdrage tot de kennis der mikroskopische fauna en flore van de Banda-Zee. Verh. Konin. Akad. Weten.,
10: 1–34.
ICZN, 2000. International Code of Zoological Nomenclature. Fourth Edition. The International Trust for Zoological
Nomenclature. London, 306 pp.
Lazarus, D. B., 1998. The Ehrenberg Collection and its curation. In D. M. Williams & R. Huxley (ed.) Christian Gottfried
Ehrenberg (1795–1876), The Man and His Legacy, Special Publication of the Linnean Society, Academic Press,
pp. 14–29.
Lazarus, D. B. & R. Jahn, 1998, Using the Ehrenberg Collection. Diatom Res., 13: 273–291.
Müller, J., 1855a. Über Sphaerozoum und Thalassicolla. Ber. Königl. Preuß. Akad. Wiss. Berlin, 1855: 229–254.
Müller, J., 1855b. Über die im Hafen von Messina beobachtenen Polycystinen. Ber. Königl. Preuß. Akad. Wiss. Berlin,
1855: 671–677.
Müller, J., 1857. Über die Thalassicollen, Polycystinen und Acanthometren des Mittelmeeres. Monat. Königl. Preuß.

96

N. Suzuki et al.

Akad. Wiss. Berlin, 1857: 474–503.
Müller, J., 1858a. Einige neue bei St. Tropez am Mittelmeer beobachtete Polycystinen und Acanthometren aus den Abbildungen. Monat. Königl. Preuß. Akad. Wiss. Berlin, 1858: 154–155.
Müller, J., 1858b. Über die Thalassicollen, Polycystinen und Acanthometren des Mittelmeeres. Abh. Königl. Akad. Wiss.
Berlin, 1858: 1–62.
Ogane, K., N. Suzuki, Y. Aita, D. Lazarus & T. Sakai, 2009, The Ehrenberg type species of flat-shaped radiolarian genera(Spongodiscidae and Stylodictyidae, Spumellaria, Polycystina). Jour. Syst. Palaeont., 7: 81–94.
Riedel, W. R., 1953. Mesozoic and Late Tertiary Radiolaria of Rotti. Jour. Paleont., 27: 805–813.
Suzuki, N., 2009. Radiolarian studies by Christian Gottfried Ehrenberg (1795–1876) and value of his species as typespecies of genera. In Y. Tanimura & Y. Aita (eds.), Joint Haeckel and Ehrenberg Project, Reexamination of the
Haeckel and Ehrenberg Microfossil Collections as a Historical and Scientific Legacy, National Museum of Natureand Science Monographs, 40: 55–69.
Suzuki, N., D. Lazarus, K. Ogane, Y. Aita & T. Sakai, 2009, General results of reexamination of Ehrenberg’s radiolariancollections: with instructions on efficient methods to find microfossils from the Collection. In Y. Tanimura & Y.
Aita (eds.), Joint Haeckel and Ehrenberg Project, Reexamination of the Haeckel and Ehrenberg Microfossil Collections as a Historical and Scientific Legacy, National Museum of Nature and Science Monographs, 40: 71–86.
(Note: All the plates and other tables are prepared as the attachments in the CD at the back of the monograph.)