Morfología, taxonomía y distribución del género de coral Holocoenia (Scleractinia) del Cretácico Temprano y su primer registro en el Caribe
Hannes Löser
Estación Regional del Noroeste, Instituto de Geología, Universidad Nacional Autónoma de México, Blvd. Luis Donaldo Colosio S/N y Madrid, 83250 Hermosillo, Sonora, Mexico. loeser@paleotax.de
Manuscript received: April 14, 2008 ]]>
Corrected manuscript received: September 17, 2008
Manuscript accepted: September 19, 2008
ABSTRACT
Although ten species are currently assigned to the Early Cretaceous coral genus Holocoenia, its characteristics are poorly known. Using material from the type locality of the type species Astrea micrantha along with described and undescribed material from France, Mexico, Poland and Spain, the genus is revised. It has a cerioid form with small calices, compact septa, a styliform columella, and an incomplete septothecal to synapticulothecal wall. Provisionally, it is assigned to the family Thamnasteriidae, being closely related to Mesomorpha and Thamnasteria. The genera Stereocaenia and Paretallonia are considered junior synonyms of Holocoenia. According to the present revision the genus contains only two species, which range from the Valanginian to the Aptian. Holocoenia micrantha is restricted to the central Tethys whereas Holocoenia jaccardi extends geographically from South America (Aptian of Argentina) and southern North America (Aptian of Puebla, Mexico) to the eastern Tethys (Hauterivian of Georgia). The indication of the genus in the San Juan Raya area in Puebla is the first indication in Central America. While the genus has been indicated in only eleven outcrop areas, making it rather rare, in many of these localities samples of Holocoenia are common.
Key words: corals, Scleractinia, Holocoenia, Early Cretaceous.
RESUMEN
Aunque actualmente existen diez especies asignadas al género de coral Holocoenia, poco sabemos sobre sus características. En este trabajo se revisa este género usando muestras de la localidad tipo de la especie Astrea micrantha, así como material nuevo de Francia, Polonia y España. Este género es cerioide con cálices pequeños, septos compactos, una columnila estiliforme y una pared incompleta de septotecal a sinapticulotecal. De manera provisional, el género está asignado a la familia Thamnasteriidae, muy cercano a Mesomorpha y Thamnasteria. Los géneros Stereocaenia y Paretallonia se consideran como sinónimos de Holocoenia. Según la revisión presente, el género tiene solamente dos especies, que tienen un rango desde el Valanginiano hasta el Aptiano. Holocoenia micrantha está restringido al mar de Tethys central, mientras que Holocoenia jaccardi muestra una distribución más amplia desde Sudamérica (Aptiano de Argentina) y América central (Aptiano de Puebla) hasta el mar de Tethys oriental (Hauteriviano de Georgia). Las evidencias del género en el área de San Juan Raya en Puebla, representan los primeros registros para América central. Aunque el género ha sido registrado solamente en once localidades y por tanto se considera raro, en muchas de estas localidades las muestras de Holocoenia son abundantes.
Palabras clave: corales, Scleractinia, Holocoenia, Cretácico Temprano.
]]>INTRODUCTION
The early Cretaceous coral genus Holocoenia is poorly known and interpreted by various authors in different ways. Even though the genus is rare, its species are distributed over a wide area and are punctually abundant. Because the genus was never properly revised, the species are distributed among three synonymous genera. The systematic study of multiple coral collections and the sampling of various outcrops have brought together a considerable amount of material allowing the present revision at the generic and species level.
The genus Holocoenia was first established by Milne–Edwards and Haime (1851: 99) on the basis of the species Astrea micrantha Roemer, 1841. The type species was designated by Milne–Edwards and Haime (1851) and the genus was monospecific when created. Astrea micrantha derives from the Hils conglomerates, a shallow marine, siliciclastic transgressional sediment which cropped out in Lower Saxon (Germany). These sediments were formed during the transgression of the very Late Valanginian to very Early Hauterivian (late Paucinodum / early Amblygonium zone; Michael 1974). The type locality of Astrea micrantha is Berklingen, a small village a few kilometers southeast of Braunschweig. The outcrop, probably a sand or clay pit, does not exist anymore. The description of the new genus by Milne–Edwards and Haime (1851) is brief and more a differentiating diagnosis: in comparison to Thamnasteria the genus Holocoenia is characterised by having a prominent styliform columella and septa with "entire edges" (a smooth distal margin).
D'Orbigny (1850a) established about 300 new species, all with very brief descriptions and lacking illustrations. Among many others, he created a species Centrastrea collinaria with the type locality Fontenoy in the Yonne department (France). The small village Fontenoy is practically surrounded by fields where the Calcaire à Spatangus, brownish–yellow oolithic marls, crops out. These sediments are extremely rich in fossils and contain a very diverse coral fauna (Löser, 2001). They belong to the basal Hauterivian (zone of Acanthodiscus radiatus) and represent remains of the same transgressions that formed the Hils conglomerates. The holotype of Centrastrea collinaria (MNHN, Coll. d'Orbigny B14279, old number 5292) could not be found (but recorded in the online catalogue of the MNHN).
The first illustration of Centrastrea collinaria was provided by de Fromentel (1857). He assigned the species to Holocoenia and discussed also the synonymy of the three species (C. collinaria, C. excavata, C. microphyllia) established by d'Orbigny (1850a). De Fromentel (1857) also mentioned Holocoenia micrantha providing a quoted description. Both species, H. collinaria and H. micrantha have the same calicular dimensions and the same number of septa, but de Fromentel (1857) did not discuss the synonymy of these species, probably because of the absence of correct data on H. micrantha.
Bölsche (1866) revised the corals from the Hils conglomerates and gave a detailed description, but no illustration of Holocoenia micrantha. He did, however, mention that Centrastrea collinaria and Holocoenia micrantha are the same species.
De Fromentel (1883) described and illustrated both species again, but he gave no calicular diameter for Centrastrea collinaria (but did for H. micrantha). In the following years, Centrastrea collinaria was quoted with Centrastrea or Holocoenia as the genus name, whereas Holocoenia micrantha always retained the same genus name. Both species were mentioned occasionally in the literature, but they were almost always from the Late Valanginian / Early Hauterivian of Lower Saxon or from the Early Hauterivian of the Paris basin.
Through the years, various species were attributed to Holocoenia, such as Holocoenia indica Stoliczka, 1873, Holocoenia ramosa Stoliczka, 1873, Holocoenia jaccardi Koby, 1897, Holocoenia chelussii Prever, 1909, Holocoenia formai Prever, 1909, and Holocoenia polymorpha Prever, 1909.
Alloiteau (1952: 629) mentioned that Holocaenia (sic) is a genus with a poorly known structure, differing from Thamnasteria by the compactness of septa, which are always in a low number. Practically en passant Alloiteau (1952) established the genus Stereocaenia. As the type species, Alloiteau designated Holocaenia collinaria de Fromentel. However, de Fromentel never established such a species and therefore Alloiteau will say, that de Fromentel had misidentified his material: it did not belong to Centrastrea collinaria d'Orbigny, 1850 sensu stricto but to another species for which Alloiteau (1952) created a new genus. According to the ICZN (1999, article 11.10) Stereocaenia collinaria therefore becomes available as the type species with Alloiteau (1952) as its author. The type is represented by the sample depicted by de Fromentel (1857), kept at the MNHN under the number M03564. The description of the new genus is short: "differs from the previous [= Holocoenia] by the lateral fusion of rudimentary septa with the numerous septa of the first and second cycle and the presence of numerous scattered calcification centres". The illustration (Alloiteau 1952, fig. 112) is poor. The "fusion of rudimentary septa" are interpreted herein as the septa of the second generation which are connected by synapticulae to the septa of the first generation. The microstructural interpretation is suspect because in material from the Hauterivian of the Paris basin, microstructures are rarely preserved. As can easily be counted in this illustration, the total number of septa is eighteen, with little difference from Holocoenia micrantha, and there is no trace of septa connected to each other.
]]> Alloiteau (1957) provided a detailed description of Holocoenia based on material from the Hauterivian of the Paris basin which Alloiteau (1957: 205) believed to belong to Holocoenia micrantha ["spécimens … que nous avons déterminés d'après les descriptions et figurations originales (celle–ci très mauvaises)"; "we have determined them after the original descriptions and illustrations (which are very poor)"]. Thus it is questionable how Alloiteau managed to determine material using poor descriptions and illustrations. The original description and depiction are indeed poor, and the differentiating diagnosis given by Milne–Edwards and Haime (1851) does not provide much additional information. Only the description provided by Bölsche (1866) gives precise information. He was the only researcher who studied material from the type locality of the type species of Holocoenia despite of Roemer. However, Alloiteau (1957) provided a diagnosis different from that of Bölsche (1866), suggesting that Alloiteau most likely had material different from Holocoenia micrantha in hand, and with this material it was easy to separate Holocoenia from Stereocaenia. Bölsche (1866) had already mentioned that one could easily become confused because Holocoenia micrantha looks like Actinastrea (e.g., Actinastrea cornueliana d'Orbigny, 1850, has the same appearance and dimensions as Holocoenia micrantha), and it is likely that Alloiteau (1957) examined a species of this genus. Since he did not give an illustration of his material, it remains uncertain what Alloiteau considered Holocoenia.Alloiteau (1957) gave also a detailed description of Centrastrea collinaria d'Orbigny, 1850, and assigned it as type species to the genus Stereocaenia. Since he had previously assigned (Alloiteau, 1952) Centrastrea collinaria d'Orbigny, 1850 sensu de Fromentel, 1857 as type species, this second designation is invalid. It is confusing that the illustrations of Stereocaenia collinaria in Alloiteau (1957) are all labeled as "Stereocaenia (Holocoenia) collinaria de From. sp.", indicating a possible synonymy of both genera. The distinction between Stereocaenia and Holocoenia as proposed by Alloiteau (1957: 207) is not based on any type material but only on the imagination which had Alloiteau about both genera.
After Alloiteau (1957), Holocoenia was not generally accepted by coral taxonomists. Instead of using Holocoenia, the genus name Stereocaenia was applied. Holocoenia species were attributed to Stereocaenia, such as Centrastrea collinaria d'Orbigny 1850 (by Fricot et al., 1995) or Astrocoenia triboleti Koby, 1897 (by Morycowa, 1964). To increase the confusion, Sikharulidze (1972) established the genus Paretallonia with the type species P. bendukidzeae from the Hauterivian of Georgia, which fits perfectly into the concept of Holocoenia. After Sikharulidze (1972), true Holocoenia material was assigned to Paretallonia (e.g., by Baron–Szabo 1997) making the confusion complete.
The diagnosis given by Baron–Szabo (2002) for Holocoenia is based on Holocoenia polymorpha Prever, 1909. She assigned the genus to the Actinastraeidae family. It is correct that all syntypes of Holocoenia polymorpha belong to the genus Actinastrea, but H. polymorpha is not the type species of Holocoenia and can therefore not be used to provide an updated diagnosis or to redefine the systematic position of Holocoenia.
MATERIAL
Unpublished material mentioned under occurrences in the systematic part comes from the following localities:
1. Berklingen (Lower Saxon, Germany). Hils conglomerates. Late Valanginian to earliest Hauterivian (late Paucinodum / early Amblygonium zone. For details see Michael (1974).
2. Fields east of the Fontenoy village (Yonne, France). Calcaire à Spatangus. Basal Hauterivian (Radiatus zone). For details see Corroy (1925).
3. San Juan Area (Puebla, Mexico). San Juan Raya Formation. Aptian. For details see Calderón–García (1956) and Reyeros–Navarro (1963). The stratigraphy is only an approximation; a (Late) Barremian to Early Aptian instead of Aptian age is also possible. Detailed investigation on the stratigraphy of the San Juan Raya Formation is under progress.
]]>SYSTEMATIC PALAEONTOLOGY
The following institutional abbreviations are used: FGUB: Facultad de Geologia de la Universitad de Barcelona, Spain; BSP: Bayerische Staatssammlung für Paläontologie und Geologie München, Germany; GIN: Geologicheskij Institut, Tbilisi, Georgia; IGM: Colección Nacional de Paleontología, Instituto de Geología, UNAM, Ciudad de México, Mexico; MB: Naturkundemuseum der Humboldt–Universität Berlin, Germany; MGSB: Museo Geológico del Seminario de Barcelona, Spain; MHNG: Muséum d'histoire naturelle de la Ville de Genève, Switzerland; MNHN: Muséum National d'Histoire Naturelle, Paris, France. The following abbreviations are applied in dimensions of the corals: c: calicular diameter; ccd: distance of calicular centres; cm: calices per 25 mm2; s: number of septa; sd: density of septa. The abbreviations used in the synonymy lists follow Matthews (1973): *: earliest valid publication of the species name; ?: the assignation of this description to the species is doubtful (so marked quotations are not reflected in the stratigraphic and palaeobiogeographic distribution); non: the described material does not belong to the species concerned; p: the described material belongs only in part to the species concerned; v: the specimen was observed by the author. A year in italics indicates that the quotation is provided with neither a description nor an illustration.
Order Scleractinia Bourne, 1900
Herein the classification system proposed by Alloiteau (1952) is applied, and not that of Vaughan and Wells (1943) because the latter is much more dedicated to Tertiary corals than to those of the Cretaceous. The family Thamnasteriidae belongs to the suborder Fungiina because of the presence of synapticulae and pores in the septal blades.
Suborder Fungiina Verrill, 1868–70
Family Thamnasteriidae Reuss, 1864
Holocoenia Milne–Edwards and Haime, 1851 emend. de Fromentel, 1857
Type species. Astrea micrantha Roemer, 1841, original designation by Milne–Edwards and Haime (1851).
Synonyms. Stereocaenia Alloiteau, 1952; Paretallonia Sihkarulidze, 1972.
]]> Diagnosis. Cerioid colony with small calices. Septa compact, in radial symmetry and always two generations. Septal face with few granules. Wall incomplete, made of synapticulae and septa. Columella styliform. Endotheca present, made of thin dissepiments. Budding extracalicinal. Septal microstructure unknown.Remarks. Holocoenia forms small to medium large colonies, circular or with an irregular shape. In siliciclastic environments flat and laterally extended, often with pillar like elevations indicating a certain sediment stress. In carbonatic environments the colonies have a massive growth form. Colonies often with a wrinkled holotheca. The outer appearance of the colony might be cerioid or thamnasterioid. The colony is cerioid with well separated polygonal to circular calices. The calices are slightly depressed, the walls slightly elevated with costae running over the wall. The septa are compact and thick. They are almost always free and rarely connected to each other as in Thamnasteria or Mesomorpha. The Septa always appear in two size orders, one long which reach the columella and one shorter which reach about half or less of those of the first order. There exists no clear symmetry. Though it is radial, calices with eight, nine, ten or even eleven septa of the first order can be found in the same colony. Septal number in all Holocoenia species is always between 16 and 22, but 20 septa are most common. Septa and costae cannot be clearly distinguished; obviously septa are biseptal blades where the external part of one septum forms a septum of another calice. The columella is styliform, clearly visible, slightly elevated and large compared to the small diameter of the calices. Septa of the first order can be attached to it. The wall is septothecal with many synapticulae. It is not compact as it can be seen in longitudinal sections. Synapticulae are only found in the wall. The endotheca consists of few thin dissepiments. – The genus was originally described as Holocaenia, but the name was later changed by de Fromentel (1857) to Holocoenia, a change of spelling which was later generally accepted and which is therefore valid.
Comparison. Mesomorpha and Thamnasteria are closely related genera, and both form astreoid to thamnasterioid colonies without a well–defined wall. Moreover, septa are often (Mesomorpha) or sometimes (Thamnasteria) attached to other septa. Both genera have no clear septal size orders as in Holocoenia. Actinastrea is very similar but differs by the mostly compact wall, the absence of synapticulae, the presence of fine thorns at the lateral septal faces, and septal swellings.
Systematic position. The systematic position is preliminary. The genus also shares many characteristics with the suborder Archeocaeniina (Astrocoeniina p.p.). It is closely related to Etallonasteria Roniewicz, 1987, a Jurassic coral genus very similar to Holocoenia but without columella.
Species. Only two species are accepted here: Holocoenia micrantha (Roemer, 1841) and Holocoenia jaccardi, Koby, 1897. Both species are morphologically identical, and differ only in their calicular diameter. Though both species co–occur at some localities or in some regions, species can be clearly distinguished by their calicular dimensions. Other species assigned to Holocoenia or its synonymous genera belong to one of these two species or other genera. Species originally assigned to other genera but belonging to Holocoenia, are synonymous with one of the two species. In Table 1 the dimensions and some statistical values of the examined material are provided.
Distribution range. Valanginian to Early Aptian. Late Aptian is questionable.
Holocoenia micrantha (Roemer, 1841) Figures 1, 2a–c
* 1841 Astraea micrantha Roemer, p. 113, pl. 16: 27
]]> 1849 Thamnastrea micrantha – Milne–Edwards and Haime, p. 1581850a Centrastraea collinaria d'Orbigny, (2), p. 93
1850a Synastrea micrantha – d'Orbigny, (2), p. 94
1850b Centrastrea collinaria – d'Orbigny, p. 177
1850b Synastrea micrantha – d'Orbigny, p. 178
1851 Astrea micrantha Roemer – Milne–Edwards and Haime, p. 99
1851 Thamnastrea ? collinaria – Milne–Edwards and Haime, p. 112
1851 Thamnastrea ? micrantha – Milne–Edwards and Haime, p. 111
1857 Holocoenia micrantha – de Fromentel, p. 53
v 1857 Holocoenia collinaria – de Fromentel, p. 54, pl. 7: 9, 10
]]> 1857 Holocoenia micrantha – Milne–Edwards, (2), p. 2501857 Thamnastraea ? collinaria – Milne–Edwards, (2), p. 583
1861 Holocoenia collinaria – de Fromentel, p. 200
1861 Holocoenia micrantha – de Fromentel, p. 200
1862 Holocoenia collinaria de Fromentel – de Fromentel, p. 410
1866 Holocoenia micrantha – Bölsche, p. 476
1867 Holocoenia micrantha Roem. sp. – Bölsche, p. 40
1868 Holocaenia collinaria (D'Orb.) E. de Fromentel – de Fromentel, p. 86, pl. 8: 3
1881 Holocoenia micrantha – Quenstedt, p. 996, pl. 182: 3
1883 Holocoenia collinaria – de Fromentel, p. 516, pl. 138: 1, pl. 139: 2
]]> 1883 Holocoenia micrantha – de Fromentel, p. 517? 1887 Centrastraea collinaria – Mallada, p. 170, pl. 62: 4
non v 1909 Holocoenia micrantha – Prever, p. 128, pl. 14: 8 [= Stylina sp.]
? 1937 Centrastraea collinaria d'Orbigny, 1849 – Bataller, p. 201
1952 Holocaenia collinaria de Fromentel – Alloiteau, p. 659, fig. 112
1956 Holocoenia collinaria Fromentel, 1867 – Wells, p. 372, fig. 263.2
1957 Centrastrea collinaria d'Orb. 1850 – Alloiteau, p. 206
p 1957 Stereocaenia collinaria de From. sp. – Alloiteau, fig. 272–274, pl. 19: 4, non pl. 1: 10 [= Mesomorpha or Thamnasteria sp.]
? 1978 cf. Holocoenia micrantha (Roemer) – Wingfield, Evans and Deegan, p. 30
non 1980 Stereocoenia collinaria (Fromentel, 1857) – Kuzmicheva, p. 100, pl. 37: 2 [= Mesomorpha sp.]
]]> non 1983 Stereocaenia collinaria (Fromentel, 1857) – Kuzmicheva, p. 34, pl. 3: 4 [not pl. 2: 4 as indicated] [= Mesomorpha or Thamnasteria sp.]non 1987 Stereocoenia collinaria (d'Orbigny, 1850) – Kuzmicheva, p. 246, pl. 5: 3 [= Mesomorpha sp.]
non 1988 Stereocoenia collinaria (Fromentel, 1857) – Kuzmicheva and Aliev, p. 168, pl. 5: 3 [= Mesomorpha sp.]
v 1995 Stereocaenia (= Holocoenia) collinaria d'Orbigny – Fricot, Colleté and Brossard, p. 32, text–fig. 35
v 1997 Paretallonia bendukidzeae Sikharulidze, 1972 – Baron–Szabo, p. 76, pl. 11: 1, 3, 4
v 2001 Holocoenia jaccardi Koby, 1897 – Löser, p. 45, pl. 2: 7
2002 Holocoenia micrantha (Roemer, 1841) – Löser et al. p. 347 [detailed synonymy here]
2002 Stereocaenia collinaria (d'Orbigny, 1850) – Löser et al., p. 616 [detailed synonymy here]
2002 Stereocaenia collinaria Alloiteau 1952 – Löser et al., p. 616 [detailed synonymy here]
v p 2006 Holocoenia jaccardi Koby 1897 – Löser and Ferry, p. 483, fig. 5.4 [not fig. 5 = Holocoenia jaccardi]
]]> Types. The type of Astrea micrantha is presumably lost. It was sought but not found in the following institutions: the Hildesheim Museum dedicated to F.A. Roemer, in the collections of the Naturkundemuseum Berlin, the collections of the Geologische Bundesanstalt (Berlin), the Geowissenschaftliche Sammlung of the University Bremen, the University collection of Göttingen, and the Staatlichen Naturhistorischen Museum Braunschweig, all in Germany. Topotypical material was sought in the same collections and only one small sample was found in the Naturkundemuseum Berlin, which is used here as a reference sample (MB K2466.1). The holotype of Centrastrea collinaria d'Orbigny, 1850 is probably not lost (according to written communication by E. Morycowa, Kraków), but it is not available. The holotype of Stereocaenia collinaria Alloiteau, 1952 (MNHN M03564) was available for study but there was no opportunity to make a polished section to clearly prove its systematic position. Thin sections made by Alloiteau were not available for study. The holotype of Paretallonia bendukidzeae Sikharulidze, 1972 is deposited in the Tbilisi Geological Institute (GIN) under the number 36/77, but was not available for study.Synonyms. Centrastraea collinaria d'Orbigny, 1850; Stereocaenia collinaria Alloiteau, 1952.
Dimensions. c: (1.0) 1.4–1.5 (1.7) mm, ccd: (1.2) 1.5–1.6 (2.0) mm, cm: 10–11, s: 18–20, sd: 4–5 / 1 mm.
Remarks. Holocoenia collinaria d'Orbigny, 1850 was unified by de Fromentel (1857) with Centrastrea microphyllia d'Orbigny, 1850 and Centrastrea excavata d'Orbigny, 1850. Centrastrea microphyllia remains an unknown species. Its type is not available. The illustration provided by de Fromentel (1887) is poor and the description is incomplete since it starts on the very last page of the 16th (and last) instalment of the revision of the Cretaceous corals within the Paléontologie française, which is also the last page of the whole publication that was not completed. Centrastrea excavata d'Orbigny, 1850 is considered a species different from Holocoenia and was in recent literature assigned to Mesomorpha. The species is not revised: its type (MNHN 5291) is a small unsectioned sample that may belong to Mesomorpha or Thamnasteria. The material mentioned by Bataller (1937; probably only in reference to Mallada 1887) was sought in the MGSB (Barcelona), but not found. It is not registered in the files and according to Sebastian Calzada (Barcelona, pers. comm. June 2007) this is a sure indication that the material is not held in the collection. The age of the La Avellà outcrop is also questionable. It was originally published as Hauterivian (Götz et al., 2005), but new material kindly provided by S. Tomás (Barcelona) included a Felixigyra species, a genus which is absent in the Hauterivian elsewhere.
Occurrence. Astrea micrantha Roemer, 1841: Early Valanginian of Arzier, La Violette quarries (Vaud, Switzerland); Latest Valanginian to Earliest Hauterivian (late Paucinodum / early Amblygonium zone) of Apelnstedt and Berklingen near Braunschweig (Lower Saxon, Germany); Early Hauterivian (Radiatus zone) of the various outcrops near the villages Fontenoy, Gy–l'Evêque, and Les Saints (Yonne, France) and near Vallières and Marolles (Aube, France); Hauterivian (?Barremian) of the La Avellà section near Catí (Valencia, Castellón, Spain; FGUB AV–C–6); Late Barremian of Pont de Laval near St.Remèze (Ardèche, France); Latest Barremian to Early Aptian (Weissi zone) of the outcrops Mahdtal, Mitteleck, Obere Gottesackerwände, Seealpe in the Allgäu Mountain range (Bavaria, Germany). – Additional occurrences indicated for Centrastrea collinaria d'Orbigny, 1850: Early Hauterivian (Radiatus zone) of Morancourt and Saint Dizier (Haute–Marne, France) and outcrops near the villages Chenay, Leugny, Saint–Sauveur and Venoy (Yonne, France).
Range. Valanginian to Early Aptian (Weissi zone).
Holocoenia jaccardi Koby, 1897 Figures 2d–f
v p 1850a Astrocoenia Cornueliana d'Orbigny, p. 92
*1897 Holocoenia Jaccardi Koby, p. 35, pl. 4: 4, 5
v 1897 Astrocoenia Triboleti Koby, p. 62, pl. 14: 6–8
]]> 1928 Astrocoenia cf. Triboleti Koby – Gerth, p. 7v p 1935 Astrocoenia Cornueliana d'Orb. – Cottreau, pl. 74: 7 [non fig. 6]
v non 1936 Holocoenia ex. aff. jaccardi Koby, 1898 – Hackemesser, p. 21
? 1959 Holoccenia [sic] Jaccardi Koby – Bonev, p. 478, pl. 2: 1
v 1964 Stereocaenia triboleti (Koby, 1896) – Morycowa, p. 77, pl. 23: 1–3
v 1970 Stereocaenia triboleti (Koby) – Czerminski, p. 129
1972 Paretallonia bendukidzeae Sikharulidze, p. 643, text–fig. 1, 2
1985 Paretallonia Bendukidzeae Sikh., 1972 – Sikharulidze, p. 62, pl. 28: 2
v 1989 Stereocaenia triboleti (Koby, 1896) – Morycowa, p. 65, pl. 27: 1, 2
non 1996 Paretallonia bendukidzeae Sikharulidze, 1972 – Császár and Turnsek, p. 434, fig. 4 [= Cryptocoenia sp.]
]]> v non 1996 Paretallonia bendukidzeae Sikharulidze – Wilmsen, p. 360, pl. 3: 3 a, b, 4 [= Actinastrea sp.]v non 1997 Paretallonia cf. bendukidzeae Sikharulidze – Wilmsen, pl. 32: 3, 4 [= Actinastrea sp.]
2002 Holocoenia micrantha (Roemer, 1841) – Löser et al. p. 347 [detailed synonymy here]
2002 Paretallonia bendukidzeae Sikharulidze, 1972 – Löser et al., p. 498 [detailed synonymy here]
2002 Stereocaenia triboleti (Koby, 1896) – Löser et al., p. 617 [detailed synonymy here]
v 2005 Holocoenia jaccardi Koby, 1897 – Götz, Löser and Schmid, p. 876, fig. 8C
v p 2006 Holocoenia jaccardi Koby, 1897 – Löser and Ferry, p. 483, fig. 5.5 [not fig. 4 = Holocoenia micrantha]
Holotype. The holotype of Holocoenia jaccardi is officially not available. The specimen of the collection of the Muséum d'Histoire naturelle de Neuchâtel (No. 26735) is suspected to be the holotype, even though it is not marked as such and its identity with the sample figured by Koby cannot be assured. Syntypes of Astrocoenia Triboleti are available at the Muséum d'Histoire naturelle de la Ville de Genève (MHNG 4852), but they are poorly preserved.
Synonyms. Astrocoenia triboleti Koby, 1897; Paretallonia bendukidzeae Sikharulidze, 1972.
Dimensions. c: (0.8) 0.9–1.2 (1.3) mm, ccd: (0.9) 1.0–1.4 (2.0) mm, cm: 12–14, s: (16) 18–20, sd: 5–6 / 1 mm.
]]> Remarks. Holocoenia jaccardi and Holocoenia triboleti were described by the same author in the same year. Priority is given to the first species not only because of page precedence but because it has been assigned by Koby to Holocoenia, whereas the other species to Astrocoenia. Moreover, the (? type) material available of Holocoenia jaccardi is much better preserved than those of Holocoenia triboleti. The material described by d'Orbigny as Astrocoenia Cornueliana consists of various syntypes of which one (MNHN R09357 = Coll. d'Orbigny 5284A) belongs to Holocoenia. Specimen MNHN A24821 was selected as lectotype of Astrocoenia Cornueliana according to the collection database of the MNHN without giving a reference, which makes this designation according to the ICZN (1999; Art. 72.4.7) invalid. The material described by Gerth is kept at the Nationaal Natuurhistorisch Museum (Leiden, The Netherlands; RGM.143057). With the kind help of Jacob Leloux (Leiden) photographs of the sample and thin sections were made available to the author.Occurrence. Early Hauterivian (Radiatus zone) of the outcrops near the village Gy–l'Evêque (Yonne, France); Hauterivian of Shkmeri (Racha, Georgia), Kvemo–Chalovani and Sachkhere (Imereti, Georgia), Wieliczka, Buków (Malopolskie, Wieliczka, Poland). Hauterivian (?Barremian) of the La Avellà section near Catí (Valencia, Castellón, Spain); Barremian of Morteau (Doubs, France); Early Barremian of the section NW Archiane in the Vercors Mountain range (Drôme, France); Late Barremian of outcrops at Pont de Laval, Belvédère du Serre–de–Tourre and Belvédère du Gaud near St.Remèze (Ardèche, France), Trzemesna near Tarnów (Malopolskie, Tarnów, Poland); Early Aptian of Jastrzebia near Lanckorona (Malopolskie, Wadowice, Poland); Aptian (? Late Barremian) of the San Juan Raya area (Puebla, Mexio; IGM 9253); Aptian of Sierra Vaca Muerta (Neuquén, Argentina).
Range. Hauterivian to Aptian (range into the Late Aptian is questionable).
OTHER SPECIES
Species originally or currently assigned to Holocoenia, but not being synonymous with one of the above described species and which do not belong to Holocoenia (all Cretaceous material was available to the author):
Holocoenia chelussii Prever, 1909 – both syntypes belong to Columnocoenia.
Holocoenia indica Stoliczka, 1873 – the species belongs very probably to Astreopora.
Holocoenia polymorpha Prever, 1909 – all five syntypes belong to Actinastrea.
Holocoenia ramosa Stoliczka, 1873 – both syntypes belong the same Columnocoenia species.
]]> It is very probably that the genus does not occur in the Jurassic. The three species established by de Fromentel (1856) without illustration do not fit well into the concept of the genus. Holocoenia cesaredensis Koby, 1904 was assigned by Geyer (1955) to the genus Convexastrea. Thamnastrea scita Milne–Edwards and Haime, 1851 was assigned by Beauvais (1966) to Stereocaenia. Thamnastrea scita belongs to Thamnasteria (for instance see Pandey and Fürsich 2003). The material described by Beauvais (1966) does not belong to Thamnasteria scita, but to Thamnasteria mettensis (according to Lathuilière 2000).Stratigraphic and palaeobiogeographic distribution
Though both species are restricted to a few outcrops, they are not rare in those outcrops. Holocoenia micrantha is common in the Early Hauterivian of the Paris basin. It is abundant in the recently investigated faunas of Ardèche and Drôme (Barremian, France) and Catí (Hauterivian, probably Barremian; Spain). Holocoenia jaccardi is abundant in the San Juan Raya area.
The genus was up to now only indicated in the Boreal (France and Germany) and the central (southern France, southern Germany, Poland, Spain, Switzerland) to eastern Tethys (Georgia); the indication of Holocoenia jaccardi in the Aptian (probably Late Barremian to Early Aptian) of Puebla (Mexico) is the first indication of the genus Holocoenia in the Caribbean realm at all (Figure 3). Both species differ in their distribution. Whereas Holocoenia micrantha shows a very limited geographical distribution occurring only in the Boreal and central Tethys, Holocoenia jaccardi shows a wider distribution and was found between South and southern North America and the eastern Tethys.
The genus occurs in outcrops with a siliciclastic facies (such as in the Boreal in northern France and northern Germany, in the northern Tethys in France and Poland, in the Caribbean in Mexico) and in carbonate ramps or platforms (such as in northern margin of the central Tethys in southern France, southern Germany, and Switzerland). It has not been indicated in the coral rich sediments of the centrals Tethys (such as the Early Aptian of Greece, the Hauterivian to Aptian of Spain).
Both species almost co–occur in time (Figure 4) with the exception that H. micrantha was already indicated in the Valanginian, whereas H. jaccardi might reach into the Late Aptian. But this is not certain as the whole outcrop area around San Juan Raya (Puebla, Mexico) needs revision of its stratigraphy and the outcrops in Argentina are imprecisely dated. Both species are most widely distributed in the Hauterivian, but also appear with a considerable distribution during the Late Barremian to Early Aptian.
CONCLUSIONS
The long story of this genus clearly demonstrates the importance of proper description and illustration to prevent confusion and the assignment of the same species to multiple genera. Above all, clear illustration is essential, since although the concepts of interpreting certain morphological elements may change through time, the fossil itself, and hence illustration, remains the same. After Holocoenia was established, it was twice created under different names (Stereocaenia, Paretallonia) due to insufficient knowledge of type material or misunderstandings of the literature. Unfortunately, this type of situation is not an exception and is very common in the study of Cretaceous corals.
]]> ACKNOWLDGEMENTS
It is a pleasure to thank Jacques Ayer and Jean–Paul Haenni (Neuchâtel), Sebastian Calzada (Barcelona), Danielle Decrouez (Genève), G. Haldar (Calcutta), Peter Jung and René Panchaud (Bâle), Dieter Korn (Berlin), Jacob Leloux (Leiden), Christine Perrin (Paris), Elzbieta Morycowa (Kraków), Daniele Ormezzano (Torino), Carmen Perrilliat (Mexico City), Sara Tomás (Barcelona), and Winfried Werner (München) who allowed me to study material of their respective museum or institute collections, which made this study possible. Juliane Fenner (Hannover), Fritz Krüger (Braunschweig), Mike Reich (Göttingen), and Jürgen Vespermann (Hildesheim) searched their collections for type or topotypical material of Astrea micrantha for which I am grateful. The quality of the paper was enhanced by reviews by Jacob Leloux (Leiden) and an anonymous reviewer. For grammatical correction I would like to thank Kimberly Franklin (Tucson, Arizona).
Field work in France and preparation of thin sections was covered by DFG project FL 42/73. Examination of the Mexican material was realised within the framework of CONACyT project 52442–Q. The ability to use the Jurassic Coral Database compiled by Bernard Lathuilière (Nancy) is greatly appreciated.
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