revision of the family asteroseriidae (anthozoa, scleractinia, … · 2015. 5. 6. · by zlatarski...

21GEODIVERSITAS • 2013 • 35 (1) © Publications Scientifiques du Muséum national d’Histoire naturelle, Paris. www.geodiversitas.com

MOTS CLÉSScleractinia,

Asteroseriidae,coraux,

Crétacé,Thétys.

KEY WORDSScleractinia,

Asteroseriidae,corals,

Cretaceous,Tethys.

Löser H. 2013. — Revision of the family Asteroseriidae (Anthozoa, Scleractinia, Cretaceous). Geodiversitas 35 (1): 21-29. http://dx.doi.org/10.5252/g2013n1a2

ABSTRACTThe Cretaceous coral family Asteroseriidae (suborder Fungiina) is revised on the basis of its type genus and respective type species. The family encompasses small solitary patellate corals with compact septa in a regular hexameral symmetry. Septa are made of large trabeculae expressed in a regularly ornamented upper septal margin and septal lateral face. Synapticulae are present. In addition to the name-giving genus Asteroseris de Fromentel, 1867, the genera Actinoseris d’Orbigny, 1849 and Microseris de Fromentel, 1867, from the Cretaceous of France are assigned to the family. All genera are revised on the basis of the types of their respective type species, as far as available. Possible species of all three genera are presented. The family ranges from Hauterivian to Cenomanian of the Tethys.

RÉSUMÉRévision de la famille Asteroseriidae (Anthozoa, Scleractinia, Crétacé).Une révision de la famille Asteroseriidae (sous-ordre Fungiina) du Crétacé est proposée sur la base de son genre type et de son espèce type, respectivement. La famille est caractérisée par de petits polypiers solitaires et patelles avec éléments radiaires compacts avec une symétrie hexamérale régulière. Les trabécules son grands et les éléments radiaires montrent une forte ornementation sur leur face latérale et leur bord distal. La famille contient les genres Asteroseris de Fro-mentel, 1867, Actinoseris d’Orbigny, 1849 et Microseris de Fromentel, 1867 du Crétacé de la France. Tous les genres sont révisés sur la base des types de leurs espèces types quand elles sont disponibles. Les espèces de tous les genres sont présentées. La famille a une distribution stratigraphique allant du Hauterivien au Cénomanien de Téthys.

Hannes LÖSEREstación Regional del Noroeste, Instituto de Geología,

Universidad Nacional Autónoma de MéxicoBlvd. Luis Donaldo Colosio S/N y Madrid, Campus UNISON

83250 Hermosillo, Sonora (Mexico)[email protected]

Revision of the family Asteroseriidae (Anthozoa, Scleractinia, Cretaceous)

www.geodiversitas.comhttp://dx.doi.org/10.5252/g2013n1a2mailto:[email protected]

22 GEODIVERSITAS • 2013 • 35 (1)

Löser H.

INTRODUCTION

Small cupolate, discoid or patellate solitary corals are uncommon in siliclastic environments in the Cretaceous. They are difficult to separate from initial states of any other solitary or even colo-nial corals and therefore can only be recognised as distinct species when occurring in consider-able quantity. Their small size makes sometimes examination difficult. Sections and thin sections often result in nearly complete loss of the sample. All these factors combine to make this type of coral poorly known. The present study compiles data on the members of the family Asteroserii-dae, which includes species arrayed among three very similar genera occurring from Hauterivian to Cenomanian in the Tethys.

MATERIAL AND METHODS

The material comes from various localities. Most of them are listed, commented upon, and provided with additional references in Löser et al. (2005). Only details not reported in the 2005 publication are mentioned here. If no sample number is given, the material from the locality concerned was not available for study. Each sample number refers to only one specimen.

Material exaMinedFrance, Aube, Vallières. Calcaire à Spatangus, basal Hauterivian, Radiatus zone (F.2077).

France, Aude, Padern, SE Le Crès, 1.45 km WSW Padern. Early Albian, Mammillatum zone. This lo-cality very probably corresponds to that reported by Alloiteau (1948). The locality was sampled by the author in 1997: SMF 75576, 75617, 75632, 75639, 75640, 75651, 75658.

France, Sarthe, Le Mans. Middle to Late Cenomanian, Rhotomagense-Geslinianum zone (F.283): MNHN.F.M03501, M03523, A49177, A49178.

France, Yonne, 4 km E Leugny. Calcaire à Spatangus, basal Hauterivian, Radiatus zone. The locality is close to the classic outcrop area of Leugny and was sampled by the author in 1997/1998: BSPG 2003 XX 6058.

France, Yonne, Fontenoy. Calcaire à Spa-tangus, basal Hauterivian, Radiatus zone (F.448): MNHN.F.B14285.

Greece, Viotía, Arachova. Evangelistria conglom-erates, Early Aptian (GR.976): MB K583-2.

Spain, Alicante, (Valencia, Mun. Morella, Morella. Early Aptian (E.717): BSPG 2003 XX 4748, 4749.

The present study is – as far as possible – based on type material. However, thin sections could not be obtained from the types because that would have resulted in the loss of the specimen. Therefore, thin sections of topotypical material are also included. Several of these thin sections are taken from the collection of the MNHN. Thin sections were compared to types, as far as available, and to the original description to en-sure that the characteristics of the examined thin sectioned specimens fit into the concept of the concerned species and genus. As many species and samples have been included as possible. Because of the small size of the individual coralla, they are widely ignored in the fossil record, often not recognised as corals and in most cases not ad-equately described and illustrated. All diagnoses are based on the studied material.

abbreviations InstitutionsBSPG Bayerische Staatssammlung für Paläontologie

und Geologie, München;MB Museum für Naturkunde, Berlin;MNHN.F Muséum national d’Histoire naturelle, Paris,

Collection de Paléontologie;SMF Senckenbergmuseum, Frankfurt/M.

Other abbreviations(v) the material has been studied;c calicular diameter;s number of septa.

SYSTEMATICS

Order SCLERACTINIA Bourne, 1900 Suborder FUNGIINA Verrill, 1870

Family asteroseriidae Alloiteau, 1952

range. — Hauterivian to Cenomanian.

http://coldb.mnhn.fr/CatalogNumber/MNHN/F/M03501http://coldb.mnhn.fr/CatalogNumber/MNHN/F/M03523http://coldb.mnhn.fr/CatalogNumber/MNHN/F/A49177http://coldb.mnhn.fr/CatalogNumber/MNHN/F/A49178http://coldb.mnhn.fr/CatalogNumber/MNHN/F/B14285

23

Revision of the family Asteroseriidae

GEODIVERSITAS • 2013 • 35 (1)

diagnosis. — Small solitary patellate corallum, gener-ally free-living, but often attached to a small piece of shell debris or a benthic foraminifer. Calicular outline circular. Septa compact. Symmetry of septa radial and regularly hexameral with regular cycles. Septal cycles differ in length, but almost not in thickness. Septa of younger cyles often and regularly connected to septa of preceeding cycles. Septal upper margin with regular large dentations, lateral face with granulae arranged in vertical rows. Pali or paliform lobes absent. Synapticulae present, in a regular pattern. Columella formed by septal fusion in the centre of the calice or parietal. Endotheca absent. Wall compact, formation unknown.

reMarksMicrostructures are hardly preserved in all stud-ied specimens, but the coarsly ornamented upper margins of the septal distal border suggests that the septa are composed of large trabeculae.

systeMatic positionThe comparable large trabeculae, ornamentation of the lateral face and upper margin of the septa, and the pattern of the synapticulae clearly indi-cates that the family should be placed into the suborder Fungiina. The existence of fulturae (Gill 1981) could only be proven in one species due to preservation problems and the scarcity of samples. The Asteroseriidae are probably the ancestors of the modern Fungiidae.

coMparisonThe Micrabaciidae Vaughan, 1905 (suborder Fungii-na) have perforated septa and a perforated wall. The Funginellidae Alloiteau, 1952 (suborder Fungiina, currently only encompassing the genera Funginella d’Orbigny, 1849 and Elasmogyra Beauvais, 1982) are characterised (based on the type material of Cyclolites haueri Michelin, 1847, type species of Funginella) by much larger coralla showing septa with larger tra-beculae. The fine structure of the septa and presence or absence of synapticulae in these taxa are unknown. The members of the Cyclolitidae Milne-Edwards & Haime, 1849 (suborder Microsolenina) are also larger and show pennulae and perforated septa.

genera of the faMilyThe family was established for two solitary coral genera of the Cenomanian: Asteroseris de Fromentel,

1867 and Microseris de Fromentel, 1867. Both gen-era were monospecific when established. Included here is Actinoseris d’Orbigny, 1849, which was considered by Alloiteau (1956, unpubl.) to belong to the Siderastraeidae Vaughan & Wells, 1943, but by Zlatarski (1968) to belong to the Funginellidae. Actinoseris was established with two species. More species were later assigned to all three genera. The genera differ by the formation of their columella and the distribution of synapticulae. In Asteroseris septa of the first cycle are fused together in the centre of the calice, whereas in both other genera a trabecular columella exist. The distinction between Actinoseris and Microseris is weak; both genera differ only in the distribution of synapticulae.

Genus Actinoseris d’Orbigny, 1849

type species. — Actinoseris cenomaniensis d’Orbigny, 1849, by monotypy. The type species is valid as combined description of genus and species.

range. — Hauterivian to Cenomanian.

diagnosis. — Solitary patellate corallum. Calicular outline circular. Septa compact. Microstructure of septa unknown. Septa in cross section externally slightly thicker, and slightly thinner towards the centre of the calice. Septal maximum thickness 250 µm. Symmetry of septa radial and regularly hexameral. Cycles of septa regular. Septal cycles differ in length, almost not in thickness. First three septal cycles extended to the centre of the calice, later cycles are shorter. Septa of younger cycles often and regularly connected to septa of preceeding ones. Septal upper border smooth (probably due to preservation), lateral face with numerous fine thorns, inner margin unknown. Pali or paliform lobes absent. Costae present. Costae ornamented with fine granules on their surface. Synapticulae present, irregularly distributed. Columella composed of a group of isolated granules. Endotheca unknown. Wall present, compact, structure unknown.

reMarksFor the type species, only the complete type speci-men and thin sections of topotypical specimens were available. The structure of the synapticulae and the endotheca could not be observed because no longitudinal thin sections could be prepared and because the type specimen is eroded and barely shows ornamentation. Species other than the type species


Löser H.

show a regular dentation of the septal upper margin and vertical lines of granulae on the septal lateral face corresponding to the dentations. The existence of fulturae is possible, but could not be proven with the available material; they are reported by Alloiteau (1957: 339). The endotheca is absent in species other than the type species, although Alloiteau (1957: 339) claimed it existed. Alloiteau obviously had more material on hand, but it could not be found at the MNHN. The type locality has been inaccessible for a long time and does not allow re-collection.

Actinoseris cannot be assigned to the Funginellidae as formerly proposed. Funginella d’Orbigny, 1849 was long considered congeneric with the genera of the Asteroseriidae assuming that F. neocomiensis d’Orbigny, 1850 is the type species (for instance Beauvais 1982: II, 117). Hence, the type species is Cyclolites haueri Michelin, 1847, as designated by Wells (1936). The genus cannot be assigned to the Siderastraeidae because of the comparably lower density of ornamentation of the lateral faces of septa and lower number of synapticulae.

Actinoseris alloiteaui Beauvais & Zlatarski, 1966

Actinoseris? alloiteaui Beauvais & Zlatarski, 1966: 1171, text-fig. 1, pl. 1, figs 1-10.

diMensions. — c 9 mm, s 48.

occurrence. — Late Barremian to Early Aptian: Pu-shevo, Bela river (Veliko Tarnovska oblast, Bulgaria).

reMarksThe species is distinguished from the other two species by the lower number of septa.

Actinoseris cenomaniensis d’Orbigny, 1849 (Fig. 1A-C)

Actinoseris cenomanensis d’Orbigny, 1849: 12 (v); 1850: 180 (v). — Alloiteau 1957: 339, fig. 229, pl. 1, fig. 6 (v).

Cycloseris Cenomanensis – de Fromentel 1863: 372, pl. 49, fig. 2. — ? Söhle 1897: 42, pl. 5, fig. 3.

? Asteroseris coronula – Tchechmedjeva 1995: 63, pl. 13, fig. 7.

diMensions. — MNHN.F.M03501, lectotype, c = 14 mm, s = 96 ; MNHN.F.A49177, topotype, c = 12 mm, s = 96.

occurrence. — (?) Early Cenomanian: Germany (Bayern) Ettal, Lichtenstättgraben; Bulgaria (Pleven) Novatchené.Middle to Late Cenomanian: France (Sarthe) Le Mans (MNHN.F.M03501, MNHN.F.A49177).

reMarksThe lectotype is an unsectioned specimen that is not very well-preserved. It does not allow any destructive examination. Better diagnostic features were found in a thin section (MNHN.F.A49177) labelled as Actinoseris cenomaniensis, the specimen being very similar in size and septal pattern to the lectotype. The material described by Söhle (1897) could not be found in the collections of the BSPG.

Actinoseris escosurae (Mallada, 1887) (Fig. 1D-H)

Cycloseris Escosurae Mallada, 1887: 167; 1890: pl. 61, figs 5-7. — Bataller 1947: 67, text-fig. — Russo et al. 1996: 200, figs 4-7, pls 1-4.

diMensions. — BSPG 2003 XX 4748, c = 11 mm, s = 96.

occurrence. — Early Aptian: Spain (Valencia, Ali-cante) Mun. Alicante, Loma de Guyón; Mun. Morella, Morella (BSPG 2003 XX 4748, 4749; Spain (Valencia, Castellón) Pont de Taules.(?) Early Albian: Spain (Cataluña, Tarragona) Com. Baix Penedés, Marmellà, Can Xuec.

reMarksThe material is well-preserved. The septal upper margin shows regular dentations that correspond to vertical lines of granulae on the septal lateral faces. The endotheca is absent. The wall is compact and with regular granulae at the surface (aboral face of the corallum).

The type material should be kept at the Insti-tuto Tecnológico GeoMinero de España (Madrid, Spain), but this could not be verified. Dimensions and illustrations are based on topotypical material from the type locality. The species is distinguished from A. cenomaniensis by a much smaller columella.

http://coldb.mnhn.fr/CatalogNumber/MNHN/F/M03501http://coldb.mnhn.fr/CatalogNumber/MNHN/F/A49177http://coldb.mnhn.fr/CatalogNumber/MNHN/F/M03501http://coldb.mnhn.fr/CatalogNumber/MNHN/F/A49177http://coldb.mnhn.fr/CatalogNumber/MNHN/F/A49177

25



Fig 1. — A-C, Actinoseris cenomaniensis d’Orbigny, 1849; A, B, MNHN.F.M03501, lectotype, in oral (A) and aboral (B) views; C, trans-verse thin section (MNHN.F.A49177); D-H, Actinoseris escosurae (Mallada, 1887), BSPG 2003 XX 4749 in oral (D) and aboral (E) views; F, BSPG 2003 XX 4748, transverse thin section; G, H, BSPG 2003 XX 4749, longitudinal thin sections; G, lateral, H, central; I, Actinoseris neocomiensis (d’Orbigny, 1850), BSPG 2003 XX 6058, oral view; J, K, Asteroseris coronula (de Fromentel, 1863); J, transverse thin section (MNHN.F.A49178); K, BSPG 2003 XX 4640, transverse thin section; L, Asteroseris sp., MB K583-2, transverse thin section; M, Microseris hemisphaerica de Fromentel, 1867, MNHN.F.M03523, syntype, transverse slab. Scale bars: 2 mm.

A B C

FED

G

H

K L M

JI

http://coldb.mnhn.fr/CatalogNumber/MNHN/F/M03501http://coldb.mnhn.fr/CatalogNumber/MNHN/F/A49177http://coldb.mnhn.fr/CatalogNumber/MNHN/F/A49178http://coldb.mnhn.fr/CatalogNumber/MNHN/F/M03523


Löser H.

Russo et al. (1996) reported the autecology of the coral and assigned it with question to Cycloseris, which is ruled out here. Russo et al. (1996) did not consider members of the Asteroseriidae family as possible genera for Cycloseris escosurae. Cycloseris is an extant coral genus, just recently (Gittenberger et al. 2006) taken out of synonymy with the very similar genus Fungia.

Actinoseris neocomiensis (d’Orbigny, 1850) (Fig. 1I)

Funginella neocomiensis d’Orbigny, 1850: 91 – Alloiteau 1957: 335, pl. 9, fig. 5 (v). — ? Fricot et al. 1995: 32, text-fig. 36.

Cyclolites neocomiensis (v) – de Fromentel 1863: 355, pl. 57, fig. 2. — ? Karakash 1907: 255, pl. 23, fig. 14. — ? Kuzmicheva 1960a: 138, pl. 6, fig. 7; 1960b: 55, pl. 3, fig. 4.

diMensions. — MNHN.F.B14285, c = 15.5 mm, s = 96; BSPG 2003 XX 6058, c = 9.5 mm, s = 96.

occurrences. — Early Hauterivian: (?) France (Aube) Troyes, Vallières; France (Yonne) Fontenoy (MNHN.F.B14285) and Leugny (BSPG 2003 XX 6058);(?) Early Hauterivian: Ukraine (Krymskaya) Crimea Mts.

reMarksThe available material has not been sectioned. Several specimens were lost during preparation and did not deliver well-preserved thin sections. The sample figured by de Fromentel (1863: pl. 57: 2) is considered the lectotype, which is supported by the existence of an old label that gives a number of the d’Orbigny collection. A well-preserved sample (here Fig. 1I) shows about 20 dentations on the septal upper margin of a septum of the first cycle and a corresponding number of vertical lines of granulae.

Actinoseris provincialis d’Orbigny, 1850

Actinoseris provincialis d’Orbigny, 1850: 203.

Cycloseris provincialis (v) – de Fromentel 1863: 371, pl. 49, fig. 3, pl. 79, fig. 1.

reMarksThe species was established without giving a type locality. The type material could not be located and is very probably lost. The MNHN online catalogue lists various specimens (from different localities) depicted by de Fromentel (1863) as type specimens, but these are probably incorrect. Examination of these specimens has revealed that they differ from each other. To clarify this species, thin sections must be prepared and a lectotype needs to be selected.

Genus Asteroseris de Fromentel, 1867

type species. — Stephanoseris coronula de Fromentel, 1863 by monotypy.

range. — Aptian-Cenomanian.

diagnosis. — Solitary patellate corallum. Calicular outline circular. Septa compact. Microstructure of septa unknown. Septa in cross section externally slightly thicker, and slightly thinner towards the centre. Septal maximum thickness 250 µm. Symmetry of septa radial and regularly hexameral. Cycles of septa regular. Septal cycles differ in length, almost not in thickness. Septa of first cycle extended to the centre of the calice, later cycles are shorter. Septa of younger cycles often and regularly connected to septa of preceeding cycles. Septal upper margin unknown, lateral face occasionally with small thorns, inner margin unknown. Pali or paliform lobes absent. All septa of the first cycle are attached to the columella. Costae unknown. Synapticulae present. Columella formed by septal fusion in the centre of the calice. Endotheca unknown. Wall unknown.

Asteroseris coronula (de Fromentel, 1863) (Fig. 1J, K)

Stephanoseris coronula de Fromentel, 1863: pl. 48, fig. 3a-e.

Asteroseris coronula (v) – Alloiteau 1957: 339, text figs 230-232, pl. 3, figs 4, 5. — non Tchechmedjeva 1995: 63, pl. 13, fig. 7. — Löser 2013: 16, fig. 6a.

? Funginella? isfahanensis – Yazdi et al. 2011: 229, pl. 1, figs 1a-e, 2, 3, pl. 2, figs 1-3.

diMensions. — MNHN.F.A49178, topotype, c = 5 mm, s = 48; BSP 2003 XX 4640, c = 8.7 mm, s = 48.

http://coldb.mnhn.fr/CatalogNumber/MNHN/F/B14285http://coldb.mnhn.fr/CatalogNumber/MNHN/F/B14285http://coldb.mnhn.fr/CatalogNumber/MNHN/F/A49178

27



occurrences. — Early Albian (Mammillatum zone): France (Aude) Padern (SMF 75576, 75617, 75632, 75639, 75640, 75651, 75658);(?) Late Albian: Kolah-Qazi section (Esfahan, Iran);Middle to Late Cenomanian: France (Sarthe), Le Mans (MNHN.F.A49178).

reMarksThe type material is not available. The reference specimen is a thin section (MNHN.F.A49178) that was also used by Alloiteau (1957). The material de-scribed by Tchechmedjeva (1995) shows a parietal columella that contradicts the concept of the genus. This material may belong to Actino seris. The material described by Yazdi et al. (2011) is poorly preserved; and their discussion on the systematic position of the new species is not based on type material. It is here preliminarily assigned to Asteroseris coronula.

Asteroseris formosus Reig Oriol, 1995

Asteroseris formosus Reig Oriol, 1995: 37, pl. 6, figs 6, 7 (v).

reMarksThe holotype is the only available specimen. Fine or-namentation is not preserved and it is not possible to confirm its systematic position even within the family.

Asteroseris vidali Reig Oriol, 1989

Asteroseris vidali Reig Oriol, 1989: 30, pl. 1, figs 3, 4, pl. 7, figs 4, 5 (v).

reMarksThe type material consists of only one small un-sectionned cupolate corallum that does not allow any classification.

Asteroseris sp. (Fig. 1L)

diMensions. — MB K583-2, c = 14 mm, s = 96.

occurrence. — Early Aptian: Greece (Viotía) Arachova (MB K583-2).

reMarksOnly one thin section is available; any other parts of the specimen no longer exist.

Genus Microseris de Fromentel, 1867

type species. — Microseris hemisphaerica de Fromentel, 1867 by monotypy.

diagnosis. — Solitary patellate corallum. Calicular outline circular. Septa compact. Microstructure of septa unknown. Septa in cross section externally slightly thicker, and slightly thinner towards the centre. Septal maximum thickness 100 µm. Symmetry of septa radial and regularly hexameral. Cycles of septa regular. Septal cycles differ in length, almost not in thickness. Septa of the first two cycles extended to the centre of the calice, later cycles are shorter. Septa of younger cycles often and regularly connected to septa of preceeding ones. Septal upper margin unknown, lateral face with fine thorns. Pali or paliform lobes absent. Synapticulae present, forming a more or less regular ring. Columella poorly defined; it is probably composed of some granulae. Costae un-known. Endotheca unknown. Wall present, compact, formation unknown.

range. — Middle to Late Cenomanian.

Microseris godoganiensis Bendukidze, 1956

reMarksThe type material of this species could not be lo-cated. The description and illustration do not allow assignment of this species to a genus.

Microseris hemisphaerica de Fromentel, 1867 (Fig. 11)

Microseris hemisphaerica de Fromentel, 1867: 368, pl. 80, fig. 1 (v).

Cycloseris hemisphaerica Fromentel – Termier & Termier 1960: 310, text-fig. 2201.

diMensions. — MNHN.F.M03523, syntype, c = 8-9 mm, s = 96.

occurrence. — Middle to Late Cenomanian: France (Sarthe), Le Mans (MNHN.F.M03523).

http://coldb.mnhn.fr/CatalogNumber/MNHN/F/A49178http://coldb.mnhn.fr/CatalogNumber/MNHN/F/A49178http://coldb.mnhn.fr/CatalogNumber/MNHN/F/M03523http://coldb.mnhn.fr/CatalogNumber/MNHN/F/M03523


Löser H.

reMarksAccording to Alloiteau (1941: 16) Microseris hemi-sphaerica and M. semiglobosa are synonymous. The inner structure of Microseris semiglobosa is unknown; therefore it cannot be determined if it is a senior synonym of Microseris hemisphaerica.

CONCLUSIONS

The rarity of specimen-rich populations and the small size of the specimens make the definition of genera and species in the Asteroseriidae fam-ily difficult. Moreover, the material bears the uncertainty that very young stages of any solitary or colonial coral may appear to be a coral of the Asteroseriida.

The autecology of one member of the fam-ily – Actinoseris escosurae – was studied in great detail by Russo et al. (1996). It is unknown whether the same pattern can also be assumed for the other members of the family. Most specimens are derived from a siliclastic environment where they occur with other solitary and colonial scleractinian cor-als (e.g., Hauterivian of the Paris Basin, Aptian of Greece, Albian of Aude, Cenomanian of Sarthe). The corals from the Albian of Aude are always attached to a shell fragment or a small gastropod, whereas the samples from the Hauterivian of the Paris Basin and the Cenomanian of Sarthe do not show these attachment marks.

AcknowledgementsI am grateful to various individuals and institu-tions. Didier Merle, Jean-Michel Pacaud and col-leagues gave access to the type collection of the Muséum national d’Histoire naturelle in Paris. The Universidad Nacional Autónoma de México (DGAPA) financially supported my work in Paris. Jacob Leloux (Leiden) gave information on new Actinoseris material from the Iran and on the status of Cycloseris. Fieldwork in Spain to obtain topotypes of Cycloseris escosurae was logistically supported by Elmar Schöllhorn (Bad Waldsee). Fieldwork in France was financed by DFG pro-ject FL 42/73. Grammatical correction was car-

ried out by Brian Hallmark (Tucson, Arizona). The paper was enhanced by reviews by Bogdan Kolodziej (Kraków), Annemarie Ohler (Paris) and an anonymous reviewer.

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Submitted on 3rd June 2011; accepted on 21st February 2012;

published on 29 March 2013.

revision of the family asteroseriidae (anthozoa, scleractinia, … · 2015. 5. 6. · by zlatarski...

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