Balanophyllia Searles Wood, 1844

Baron-Szabo, Rosemarie C., 2008, Dendrophylliina, Caryophylliina, Fungiina, Microsolenina, and Stylinina, Zootaxa 1952, pp. 1-244 : 40-42

Genus Balanophyllia Searles Wood, 1844

(= Eupsammia Milne Edwards & Haime, 1848d , Type species. Madrepora trochiformis Pallas, 1766 , Eocene of France); (= Ceratopsammia Alloiteau, 1958 , Type species. Ceratopsammia besairiei Alloiteau, 1958 , Campanian-Maastrichtian of

Madagascar)

Type species. Balanophyllia calyculus Searles Wood, 1844 , Lower Pliocene of England (Norfolk).

PLATE 3:

Fig. 1 Wadeopsammia nodosa (Wade, 1926) , holotype, NMNH, I32703, Maastrichtian of the USA, scale bar: 1.5 mm. 1a, longitudinal view; 1b, cross view (photographs courtesy S. Cairns). Fig. 2 Palaeopsammia zitteli ( Vaughan, 1900) , BMNH, K˥hn coll., R.30280 (holotype of Palaeopsammia fastigiata Kühn, 1933 ), Maastrichtian of Iran, scale bar: 3 mm. 2a, cross view, polished surface; 2b, longitudinal view. Fig. 3 Balanophyllia ponderosa Vaughan, 1900 , syntype, NMNH, Vaughan coll., I158364, Paleocene of the USA. 3a, cross view, scale bar: 3 mm; 3b, longitudinal view, scale bar: 4 mm. Fig. 4 Balanophyllia caulifera ( Conrad, 1847) (paratype of Balanophyllia schlosseri Traub, 1938 ), polished surface, cross view, BSP, Kch6, Middle Paleocene of Austria, scale bar: 5 mm. Fig. 5 Balanophyllia caulifera ( Conrad, 1847) (holotype of Eupsammia narindensis Alloiteau, 1936 ), MNHN, Mo5211, Upper Maastrichtian of Madagascar. 5a, longitudinal view, scale bar: 5 mm; 5b, cross view, scale bar: 4.5 mm. Fig. 6 Balanophyllia besairiei (Alloiteau, 1958) ,

syntype, MNHN, Mo 5028, Campanian-Maastrichtian of Madagascar, scale bar: 5 mm. 6a, longitudinal view; 6b, cross view .

Diagnosis. Solitary, variable conical, often trochoid, ceratoid, or turbinate. Base broad or narrow, attached or unattached. Asexual budding may occur from edge zone. Costae short, costosepta subcompact, Pourtalès plan present. Columella elongate, spongy. Wall synapticulothecal. Epitheca present or absent.

Remarks. The validity of the taxon Eupsammia and its possible synonymy with Balanophyllia have been discussed since the early 20 th century. In the generic revision and phylogenetic analysis of dendrophylliid genera, Cairns (2001) gave a detailed discussion of the taxonomic history of these two forms and provided descriptions for Balanophyllia and Eupsammia . While Cairns acknowledged the arguments that support the grouping of Balanophyllia with Eupsammia as its junior synonym (e.g., structurally they are identical, cooccurrence of the two forms in many regions since at least the Eocene), he separated the two forms on the basis of their attached or unattached living mode and differences in their early ontogenetical development, the latter idea of which had been proposed earlier by Durham (1949). According to Durham (1949, p. 139ff.) the taxon Balanophyllia shows a polycentric development (=”...formation of the adult corallum through two or more distinct stages....”) in its very first ontogenetical stages, whereas Eupsammia is considered to show a monocentric development in its earliest ontogenetical stages (=”....formation of the adult corallum by direct conical enlargement of the prototheca...”). However, recent investigations regarding ontogenetical stages of solitary and colonial scleractinians from Austria (Gosau Group, Upper Cretaceous) by Baron-Szabo (2003 and unpublished data) indicate that different septal and wall developments in the earliest ontogenetical stages are the result of whether the polyp was attached or was free-living (see remarks regarding the presumed characteristic of Cunnolites to be “free in ephebic stage” in the chapter of the Family Cunnolitidae Alloiteau ). The sediments of the Upper Cretaceous part of the Gosau Group are characterized by muddy, soft-bottom paleoenvironments that often contain bioclastics. As a result, unattached and attached corals occur side by side. In numerous outcrops, the author of this paper has observed specimens of the same taxa of, e.g., Cunnolites , that always showed the development of a single septal cycle in the earliest ontogenetical stage with no or only one poorly developed thecal ring, whereas in attached forms at least 2 or 3 cycles and generally well-developed thecal structures were observed in the earliest stages that could be studied. This observation corresponds to the statement by Durham (1949, p. 143) regarding the attached form of Balanophyllia elegans : “No specimens representing the earliest stages of the corallum of Balanophyllia elegans were found....” Therefore, it is assumed that Balanophyllia and Eupsammia represent only one taxon that has the ability to settle on soft and hard grounds, living attached and unattached, and that, resulting from this, the terms monocentric and polycentric have no taxonomic value for this genus.

With the exception of terms monocentric and polycentric, which are considered to have no taxonomic value for this genus, the diagnosis given for Balanophyllia represents a combination of the two descriptions provided by Cairns (2001, pp. 14 and 17) for Balanophyllia and Eupsammia .