A Generic Revision and Phylogenetic Analysis of the Primnoidae

40 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
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 fl at operculum over
the polyp. The calyces are quite small (only 0.4 mm long
and 0.25 mm in diameter) and are closely appressed to
the branch, almost adnate, having only very short (two or
three scales) adaxial rows of body wall scales. Coenenchymal
scales are elongate and have coarse granules on their
edges (Figure 11n). Finally, none of the sclerites appear to
have tubercles on their inner surface, a character that blurs
the distinction between Plumarella and Acanthoprimnoa.
Distribution. Western Pacifi c, Patagonia,
northwest Atlantic, 10–1,914 m.
Type Species. Gorgonia penna Lamarck,
1815, by subsequent designation (Wright and Studer,
1889:73). Type deposited at MNHNP (fi de Versluys,
1906:18) and a fragment of the type at the ZMA (Coel.
3085) (van Soest, 1979). The type species is well described
and fi gured by Versluys (1906:18–20) and Fabricius and
Alderslade (2001), the latter in color.
25. Callogorgia Gray, 1858
FIGURE 12A–D
Callogorgia Gray, 1858:286.—Bayer, 1956:F220; 1961:294 [illustrated key
to genus]; 1981b:938 [key to genus]; 1982:119–123 [in part: key to Indo-
Pacifi c species, genus synonymy].—Bayer and Stefani, 1989:455 [key to
genus].—Bayer, 1998:162–163.—Cairns and Bayer, 2002:841–845 [key to
western Atlantic species].
Calligorgia Gray, 1870:35 [unjustifi ed emendation].—Studer, 1887:51.
Callicella Gray, 1870:37.
Xiphocella Gray, 1870:56.
Caligorgia Wright and Studer, 1889:75–77 [in part; unjustifi ed emendation].—Versluys
1906:55–58 [in part].—Kinoshita, 1908a:34.—Kükenthal,
1915:146 [key to genus and species]; 1919:362–366 [in part: key to species];
1924:267–269 [in part: key to species].
Diagnosis. Colonies usually pinnately branched
in an alternating manner (e.g., type species), dichotomously
branched in one plane, or, rarely, opposite pinnately
branched. Calyces arranged in whorls of up to 12, the appressed
calyces facing upward (Figure 12a). Well-developed
operculum present, the operculars ridged on their inner surface.
Eight marginal scales present, which do not fold over
operculars. Polyps protected by four to eight longitudinal
rows of up to 12 body wall scales, the abaxials occurring
in fully developed rows but lateral and adaxial scale rows
progressively shorter toward adaxial side; in some species
adaxial and even inner-lateral body wall scales absent, resulting
in the lower adaxial body wall being naked. Outer
surface of body wall scales smooth, granular, or covered
with prominent radiating or reticulate ridges (Figure 12b);
distal edge of inner surface of body wall scales radially
ridged, these ridges sometimes continuous with the ridges
of the outer surface. Coenenchymal scales arranged in one
layer, outer surface usually granular.
Discussion. Many of the listings in the genus
synonymy must be listed as “in part” because species now
attributed to Fanellia were customarily placed in Callogorgia.
The 26 valid species known in the genus are listed in
Table 4, most of these keyed by either Bayer (1982) or
Cairns and Bayer (2002), both of whom extensively discuss
the characters used to identify species. Bayer (1982) distinguished
two groups of species within the genus based on the
sculpturing of the body wall scales (granular versus crested)
but did not suggest taxonomic distinction at any level.
Distribution. Indo-Pacifi c, North Atlantic,
37–2,472 m.
Type Species. Callogorgia: Gorgonia verticillata
Pallas, 1766, by monotypy. Type not traced. The type
species was well described and illustrated by Carpine and
Grasshoff (1975).
Callicella: C. elegans Gray, 1870, by monotypy. The
dry types are deposited at the BM (1965.12.15.6, 8).
Xiphocella: Gorgonia verticillata sensu Esper, 1797,
by monotypy. Type not traced.
26. Fanellia Gray, 1870
FIGURE 12E–M
Fanellia Gray, 1870:46.—Bayer, 1982:134–135 [key to species].—Bayer and
Stefani, 1989:470–471 [key to genus and all species].
Diagnosis. Colonies usually dichotomously
branched in one plane (e.g., type species), although some
species branch in a pinnate or bushy dichotomous manner.
Calyces arranged in whorls (type species) of up to 16
and, in some species, in pairs, the appressed calyces facing
upward. Well-developed operculum present; operculars
ridged on their inner surface. The eight marginal scales do
not fold over operculars. Polyps protected by four to eight
longitudinal rows of up to 11 thick body wall scales, the
abaxials constituting fully developed rows but lateral and
adaxial scale rows progressively shorter toward adaxial