Dai and Horng, 2009, p. 59


Colonial in nearly all species. Budding intracalicular, and may also be extracalicular. Corallites monomorphic or polymorphic; discrete, uniserial or organically united. Monticules mainly absent. Walls may be fused, separated to various degrees, or colonies may be phaceloid or flabello-meandroid. Coenosteum spinose if present. Calice width medium to large (≥ 4 mm), with varying relief. Costosepta may be confluent. Septa in varying cycles and abundances. Free septa irregular. Septa spaced ≤ 11 septa per 5 mm. Costosepta unequal in relative thickness. Columellae mainly trabecular and spongy (> three threads), of varying sizes, and discontinuous among adjacent corallites with lamellar linkage. Paliform (uniaxial) or septal (multiaxial) lobes may be weakly or moderately developed. Epitheca varies in development. Endotheca low-moderate (tabular) or abundant (vesicular). Tooth base at midcalice elliptical-parallel. Tooth tip at midcalice irregular; tip orientation parallel or forming multiaxial bulb. Tooth height medium to high (≥ 0.3 mm). Tooth spacing medium to wide (≥ 0.3 mm), with varying numbers of teeth per septum. Tooth shape may vary between first and third order septa. Tooth size may vary between wall and septum. Granules mainly scattered on septal face; weak (rounded), strong (pointed) or irregular. Interarea smooth or palisade. Walls formed by dominant paratheca and partial septotheca. Thickening deposits in concentric rings with extensive stereome. Costa centre clusters weak or strong; ≥ 0.3 mm between clusters; medial lines weak or strong. Septum centre clusters weak or strong; ≥ 0.3 mm between clusters; medial lines weak. Perpendicular crosses absent. Columella centres clustered.


There are five synapomorphies defining Lobophylliidae (bootstrap support of 95 and decay index of 5): (1) coenosteum spinose (likelihood of 1.00 based on the Mk1 model); (2) columellae discontinuous among adjacent corallites with lamellar linkage (likelihood 1.00); (3) tooth base at midcalice elliptical-parallel (likelihood 1.00); (4) tooth tip orientation parallel or forming multiaxial bulb (likelihood 1.00); and (5) thickening deposits in concentric rings with extensive stereome (likelihood 1.00). These comprise two macromorphological, two micromorphological and one microstructural features. All of these characters strongly support the monophyly of Lobophylliidae and are monomorphic within the clade. Furthermore, the subcorallite characters unequivocally distinguish Lobophylliidae from Merulinidae, which has circular tooth base at midcalice, tooth tip orientated perpendicular to the septum or as multiaxial threads, and thickening deposits that are thick fibrous. Mussidae (clade XXI) is an exclusively Atlantic clade, and in contrast to Lobophylliidae, has costate coenosteum, regular (pointed) midcalice tooth tip, transverse septal crosses (as clusters or carinae), and no extensive stereome thickening (Budd et al., 2012).


Lobophylliidae was established by Dai and Horng (2009: 59) for six of the 13 genera in Mussidae sensu Veron (2000) and two of the five genera in Pectiniidae sensu Veron (2000). Licuanan (2009: 135) followed this scheme for the corals of northwestern Philippines. These taxa constitute the molecular clades XVIII, XIX and XX designated by Fukami et al. (2008) (for a list of all available lobophylliid nomina, valid and synonymised). For Mussidae sensu Veron (2000; see also Vaughan and Wells, 1943; Wells, 1956), Dai and Horng (2009) dealt only with the fauna in Taiwan (i.e. Lobophyllia de Blainville, 1830: 321, Acanthastrea Milne Edwards and Haime, 1848a, vol. 27: 495, Australomussa Veron, 1985: 171, Cynarina Brüggemann, 1877: 305, Scolymia Haime, 1852: 279, and Symphyllia Milne Edwards and Haime, 1848a, vol. 27: 491), so the remaining seven genera were not included in the new family. The Atlantic taxa, represented by four of these seven genera, Mussa Oken, 1815: 73, Isophyllia Milne Edwards and Haime, 1851a, vol. 5: 87, Mussismilia Ortmann, 1890: 292, and Mycetophyllia Milne Edwards and Haime, 1848a, vol. 27: 491, were placed in Mussidae by Budd et al. (2012) owing to the deep divergence between the Atlantic (clade XXI sensu Fukami et al., 2008) and Indo-Pacific fauna (Fukami et al., 2004b, 2008), and the status of Mussa as type genus of Mussidae Ortmann, 1890: 315. Blastomussa Wells, 1968: 276, was placed in family incertae sedis (Budd et al., 2012) because it is genetically distinct from lobophylliids and mussids, and most closely related to Physogyra, Plerogyra and Nemenzophyllia (clade XIV; Fukami et al., 2008; Benzoni et al., 2014). Also in family incertae sedis is Indophyllia Gerth, 1921: 405, now considered an extinct genus after I. macassarensis Best and Hoeksema, 1987: 394, was transferred into Cynarina by Budd et al. (2012). Micromussa Veron, 2000, vol. 3: 8, the final Mussidae genus (sensu Veron, 2000), was placed in Lobophylliidae by Budd et al. (2012). Further actions influenced the final generic composition of Lobophylliidae prior to the present study. Scolymia, one of the six genera that initially defined the family (Dai and Horng, 2009), was moved into Mussidae because its type, Madrepora lacera Pallas, 1766: 298 (see Vaughan, 1901: 6), is an Atlantic species (Budd et al., 2012). Its two Indo-Pacific members were redistributed into Homophyllia Brüggemann, 1877: 310, and Parascolymia Wells, 1964: 379. The two Pectiniidae genera (sensu Veron, 2000) initially assigned to Lobophylliidae by Dai and Horng (2009), Echinophyllia Klunzinger, 1879: 69, and Oxypora Saville Kent, 1871: 283, were joined by Echinomorpha Veron, 2000, vol. 2: 333 (Budd et al., 2012). Moseleya Quelch, 1884: 292, formerly in Faviidae sensu Veron (2000) was also placed in Lobophylliidae (Huang et al., 2011; Budd et al., 2012). Sclerophyllia Klunzinger, 1879: 4, was resurrected based on new molecular and morphological data collected for Sclerophyllia margariticola Klunzinger, 1879: 4, whose sister congener is Acanthastrea maxima Sheppard and Salm, 1988: 276 (Arrigoni et al., 2015). Arrigoni et al. (2014b) found Australomussa and Parascolymia to be genetically indistinguishable, and therefore considered the former to be a junior synonym of the latter. Finally, based on a morpho-molecular approach Arrigoni et al. (2016a) formally revised Homophyllia and Micromussa with the inclusion of Homophyllia bowerbanki (Milne Edwards and Haime, 1857), Micromussa lordhowensis (Veron and Pichon, 1982), M. multipunctata (Hodgson, 1985), as well as the new species M. indiana Benzoni and Arrigoni, and M. pacifica Benzoni and Arrigoni. The authors also established Australophyllia Benzoni and Arrigoni, to accommodate the highly divergent H. wilsoni. Drawing upon morphological and molecular phylogenies inferred in this study, as well as prior work carried out by Budd et al. (2012) and Arrigoni et al. (2012, 2014b, c, 2015, 2016a), we classify Lobophylliidae species into 11 genera. The major change over the most recent proposals by Arrigoni et al. (2014b, 2015) is the placement of all members of subclade I (sensu Arrigoni et al., 2014c) in Lobophyllia; our results show neither genetic nor morphological separation among Lobophyllia, Parascolymia and Symphyllia. Furthermore, they support the transfers of Acanthastrea ishigakiensis Veron, 1990: 132, into Lobophyllia, and Lobophyllia pachysepta Chevalier, 1975: 269, into Acanthastrea, which we carry out here. Lobophyllia thus becomes the most species-rich genus in Lobophylliidae but with relatively limited genetic differentiation between species (see Arrigoni et al., 2014b: fig. 9, 2014c: fig. 1). Lobophylliidae is widely distributed on reefs of the Indo-Pacific, and absent in the eastern Pacific.


This page has been in preparation since 02-Oct-2009 17:15

This version was contributed by Danwei Huang on 29-Jan-2016 13:42.

Page authors are: Ann Budd Danwei Huang. Please contact the editor if you would like to contribute to the diagnosis of this taxon.

The editor is: Ann Budd

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