The outgroup state is isolated (state 0) or biserial (state 2), implying that the more complex states of paired (state 3), whorled (state 4), and in leaves (state 5) are derived. (but see Results). The outgroup genera all have large crescent-shaped tentacles (state 0) that provide strength and protection to the tentacles (Alderslade, 1998), while large sclerites do not occur in the tentacles of primnoids (state 1). .
Primnoeides Studer and Wright in Studer, 1887
Ophidiogorgia Bayer, 1980
Aglaoprimnoa Bayer, 1996
Weakly developed, almost fl in operculum present; inner face of opercular scale rolled longitudinally. Marginal scales undifferentiated from other body wall scales, folded over bases of opercular scales (Figure 3r).
Armadillogorgia Bayer, 1980
Small Calytes covered with eight longitudinal rows of scales, the adaxial rows somewhat shorter; however, larger calyces insert additional body wall scales, rendering the longitudinal row structure indistinct and even appearing to spiral (Figure 3s). The outer surface of the body wall scales is smooth, but with radiating ridges near the distal margin, which engage similarly sized ridges on the inner distal surface of the more proximal scale, preventing lateral displacement of the scales as the calyx opens and closes ( Bayer, 1996a).
Ainigmaptilon Dean, 1926
She finally tentatively placed the new genus in the pennatulacean, allied to the Virgulariidae. Perhaps unaware of Dean's work, Molander (1929) described two new species with similar morphology, but after undergoing much the same soul-searching as Dean, he concluded that his two species belonged to a new primnoid genus (Lycurus) and new subfamily (Lycurinae ), related to the primnoid Callozostron.
Primnoella Gray, 1858
Ainigmaptilon has such an unusual mixture of characters that Dean noted that it "differs so remarkably from any other alkyonarian hitherto described that it is difficult to assign it to even any of the five great orders." She referred not only to the leaf-like arrangement of the polyps and axial canal structure, which linked it to the order Pennatulacea, but also to the scale-like sclerites and lack of siphonozoids, which linked it more closely to the family Primnoidae of the order Gorgonacea. Carlgren (1943) described still a fourth species, synonymized Ainigmaptilon and Lycurus, and elevated Molander's subfamily to family rank: Ainigmaptilonidae, with suggested affinities to Primnoidae.
Convexella Bayer, 1996
Kükenthal (1919) placed it in the Convexae section of the genus Primnoella (later named Convexella by Bayer, 1996b) because of its round-sectioned calyxes, but Bayer removed it from that genus in 1996 because it has a bare adaxial body wall. Apart from describing a separate genus for this species, we propose a tentative placement in Primnoella, but it differs from that genus in having a calyx that is round in cross-section and thus has four rows of body wall scales visible in abaxial view.
Dicholaphis Kinoshita, 1907
Characters used to distinguish species include number of polyps per whorl, number of abaxial body wall scales, number of adaxial body wall scales, size and shape of marginal scales and size of polyps (Cairns, 2006). Polyps completely covered with eight longitudinal rows of body wall scales, the rows becoming disorganized near the base of each calyx.
Callozostron Wright, 1885
Bayer (1996b) noted the similarity between Callozostron and Ainigmaptilon, particularly with regard to the basal fusion of their calyces; whereas some polyps of Callozostron are basally united, as are a few other primnoid genera (e.g. Arntzia, Onogorgia, Ophidiogorgia), the calyces of Ainigmaptilon are supported on distinct, fleshy, leaf-like structures. Furthermore, the marginal scales of Callozostron are much larger than those of Ainigmaptilon and have prominent apical spines.
Arntzia López-González, Gili and Orejas, 2002
Finally, Callozostron has a typical folded operculum, whereas the opercular scales of Ainigmaptilon bear such long apical spines that they cannot fold over the tentacles as most opercula do. Bayer (1996b) also compared Callozostron with Convexella (as Primnoella magelhaenica), and Callozostron differs from that genus in having fused basal sepals and spinose marginal scales that do not fold over the operculum.
Thouarella (Thouarella) Gray, 1870
Four species are known in this genus (Table 4), all of which are restricted to Antarctic and subantarctic regions. Parathouarella: Primnoa antarctica Valenciennes, 1846, based on the inclusion of the type species of the genus in this subgenus.
Thouarella (Euthouarella) Kükenthal, 1915
Polyps are protected by six longitudinal rows of body wall scales, with the adaxial face covered by enlarged inner lateral scales, resulting in complete coverage of the polyp. Coenenzyme scales in two layers: the outer layer consists of irregularly shaped scales with a granular outer surface; inner layer composed of small spheroids.
Thouarella (Diplocalyptra) Kinoshita, 1908
Thouarella (Epithouarella) Kükenthal, 1915
Metafannyella, new genus
Fannyella (Fannyella) Gray, 1872
Body wall scales are characteristically shaped, the outer surface with a dentate or smooth transverse crest that divides the scales into a distal, more or less concave or oblique cup-shaped part, and a proximal, tubercular base, these cup-like sclerites called ascus scales (Figure 8c– d). Distal surface of ascus scales smooth, proximal surface tuberculate, and rim of goblet cavity granular to finely toothed.
Fannyella (Scyphogorgia), new subgenus
Eight marginal scales fold over the base of the opercular scales, forming a circumoperculum; distal edges of marginals pointed or rounded, but never spinose. Polyps protected by six longitudinal rows of body wall scales, the two abaxial rows consisting of up to 16 scales, the outer and inner-lateral rows having fewer scales; adaxial row usually absent, resulting in a bare adaxial face (Figure 8b).
Fannyella (Cyathogorgia), new subgenus
Cyathogorgia is monotypic (Table 4), differing from the named subgenus in having spinose marginal (and submarginal) scales that have a keeled inner surface and in eight rows of body wall scales that completely cover the polyp. Because the differences are so small, a middle ground is taken in this, that is, the establishment of a subgenus for this species.
Onogorgia, new genus
Pyrogorgia, new genus
Amphilaphis Studer and Wright in Studer, 1887
Mirostenella Bayer, 1988
Although similar to the family Isididae in the nature of its joint axis, the scales of Mirostenella have the typical primnoid cruciform extinction pattern. Ignoring the unique character of organic nodes, Mirostenella is remarkably similar to Plumarella, as noted by Bayer (1988), which differs mainly in having sepals arranged in whorls and pairs, not in an alternate biserial fashion.
Acanthoprimnoa Cairns and Bayer, 2004
Plumarella Gray, 1870
Its opercular scales are uniquely shaped (Figure 11o–p) and so slender that they do not form a complete covering of the polyp; they articulate to form a flat operculum over the polyp. Finally, none of the sclerites appear to have tubercles on their inner surface, a character that blurs the distinction between Plumarella and Acanthoprimnoa.
Callogorgia Gray, 1858
Fanellia Gray, 1870
When the inner lateral scale is absent, this area is usually covered by an adaxial winged projection of the outer lateral scale. As discussed by Bayer (1982), Fanellia is morphologically very similar to Callogorgia, but differs mainly in the nature of the sculpture of the outer sclerite surface, which is tuberous to nodular in Fanellia instead of granular to smooth in Callogorgia.
Paranarella Cairns, 2007
Outer surface of body wall scales, opercular and coenenchymal scales all covered with prominent tubercles, often arranged in ridges (Figure 12h, j–k). Coenenchymal scales in two layers, including an inner layer of tubercular spheroids (Figure 12m) separating the stem canals;.
Primnoa Lamouroux, 1812
Also, species of Fanellia are mostly dichotomous in branching, while species of Callogorgia are mostly legume, although both genera contain species with alternate mode of branching. Eight Fanellia species are listed in Table 4, all of which are listed by Bayer and Stefani (1989).
Australogorgia, new genus
Narella Gray, 1870
Arthrogorgia Kükenthal, 1908
There are two or more transverse rows of unpaired, crescent-shaped infrabasal scales, which often form a shallow conical tube on which the calyx sits (Figure 14h). Arthrogorgia differs from Paracalyptrophora primarily by having four or more unpaired infrabasal scales that form a hollow basal tube and by having a pinnate branching pattern.
Paracalyptrophora Kinoshita, 1908
Calyptrophora Gray, 1866
Polyps protected by two pairs of crescent-shaped sclerites (each sclerite with its own separate center of calcification), basal (Fig. 15m) and buccal, each pair formed by inseparable fusion of two sclerites ab- and adaxially, result- in solid rings; no additional adaxial sclerites are present and the adaxial symphysis is narrow, leaving the adaxial face relatively bare. Bayer (2001) divided the species in this genus into two species complexes: the “japonica complex,” which includes the type species and consists of species with upturned calyxes, and smaller ones.
Tokoprymno Bayer, 1996
Species within this genus can usually be distinguished on the basis of branching pattern, calyx orientation, and whorl of body scale (number of spines, length, and whether smooth or covered with toothed ridges; see Kükenthal, 1924; Cairns, 2007a, 2007b).
Parastenella Versluys, 1906
The type species was beautifully described and illustrated by Utinomi b), who considered the alternation between marginal and opercular scales to be of generic importance. Characters used to distinguish species include colony shape, arrangement of calyces, size and presence of marginal grooves, and the number of body wall scale rows.
Candidella Bayer, 1954
1981b), who considered the alternation between marginal and opercular scales to be of generic importance.
Microprimnoa Bayer and Stefani, 1989
The original description reports that the polyps are arranged in whorls of two, but this is a different way of pairing than is coded in the phylogenetic analysis.
Pterostenella Versluys, 1906
Perissogorgia Bayer and Stefani, 1989
Dasystenella Versluys, 1906
The polyps are completely covered by five longitudinal rows of body wall scales (one abaxial row, two lateral rows and two adaxial rows, the adaxial scales being much smaller than the others). Dasystenella, originally considered by Versluys (1906) as one of the four subgenera of Stenella sensu Gray, 1870, was elevated to a generic category by Bayer (1981b), distinguished by having only five marginal scales and as brush-branched colonies .
Pseudoplumarella Kükenthal, 1915
New species of Calyptrophora (Coelenterata: . Octocorallia: Primnoidae) from the western Atlantic. A review of the genus Primnoa (Octocorallia: . Gorgonacea: Primnoidae), with description of two new species.
