Pillai: Serpulid polychaetes from the Australian Kimberleys 
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Systematic account 
The scheme for dividing serpulimorph polycheates into 
three subfamilies, namely, the Serpulinae, Filograninae, 
and Spirorbinae under the Serpulidae, and which had been 
used by some authors recently, is essentially Fauvelian. 
On the basis knowledge available at that time, Fauvel 
(1927: 346-403) includes widely different genera such as 
Serpula, Mercierella (presently Ficopomatus ), Vermiliopsis, 
Pomatostegus, Placostegus, Pomatoceros and Ditrupa 
under the Serpulinae (pp. 350-375); those included under 
the subfamily Filograninae (pp. 375-387) are Filograna, 
Salmacina, Josephella, Protula and Apomatus, while the 
Spirorbinae consists of the single genus Spirorbis divided 
into subgenera. 
However, research on serpulids carried out by numerous 
scientists during the period of over eighty years since then, 
had resulted in accrual of a vast amount of knowledge on 
them and alternative schemes for their classification, and 
there appear to be no sound reasons for reverting to Fauvel’s 
(1927) classification rather than moving on, on the basis of 
knowledge available since then. 
An important morphological character that needs to be 
taken into consideration with regard to classification of 
serpulids in general is that of opercular insertion. Although 
ten Hove (1984:184-189) did not categorically state that the 
operculum in all serpulimorphs is derived from the second 
dorsalmost branchial radiole, his arguments regarding it in 
certain taxa, supported by drawings, could, perhaps, lead one 
to such a conclusion. Indeed, Fitzhugh (1989:11) concludes 
that “ten Hove (1984) has shown that the serpulid operculum 
is actually a modified second, dorsalmost radiole. It is only 
in large-bodied serpulid species that the opercular stalk 
migrates during development to a position, which causes 
it to appear as though it originates from the first radiole.” 
Rouse (2000: 187) came to the same conclusion: “ten Hove 
(1984) argued that in all serpulids the operculum is derived 
from the second most dorsal radiole, though its position may 
alter subsequently.” However, this is, in reality, not the case 
with with all serpulids, as shown by Matjjasic & Sket (1966), 
Orrhage (1980) and Pillai (2008, 2009). 
In the latter context, Fauvel’s inclusion of Protula under 
the Filograninae causes some problems. According to 
Kupriyanova & Jirkov (1997), in Protula, an operculum, 
when present, is vesicular and occurs in the position of the 
second branchial radiole, whereas Filograna has “two spoon¬ 
shaped opercula on the first non-modified pinnulate branchial 
radioles, one on each branchial radiole”. Likewise, Fauvel’s 
subfamily Serpulinae includes genera, such as, Serpula 
and Spirobranchus in which the operculum is inserted in 
the position of the second radiole as well as those, such 
as, Ditrupa, Vermiliopsis, Pomatostegus and Mercierella 
(currently Ficopomatus ) in which it inserted in the position 
of the first radiole. 
Uchida (1978) erects several new serpulid subfamilies, 
the acceptance of which has been met with some reticence. 
However, it is to be expected that adequate scientific 
reasoning be provided in the event of acceptance or non- 
acceptance any of his specific proposals. In the meantime, 
classification of the Family Serpulidae into subfamilies 
in this paper is limited to a few that could adequately 
be explained and accepted in terms of the International 
Rules for Zoological Nomenclature. In the latter context, 
the subfamily Serpulinae, Macleay, 1840, sensu stricto is 
used for genera that constitute the Serpula-Spiraserpula- 
Crucigera-Hydroides clade. 
The descriptions of taxa in this paper broadly follow 
current phylogenetically based serpulimorph classifications. 
Their groupings range from genera that are currently 
believed by various authors to be more primitive, e.g., the 
Filograninae, to those less primitive and possess either simple 
to complex chitinous opercula, or calcareous opercula, 
besides other characters. 
They are dealt with alphabetically in two sections: the 
Kimberleys genera and species, followed by comparisons 
with species, especially of Spirobranchus, from other 
locations in Australia and the Indo-West Pacific. Within 
each genus, the species are dealt with in alphabetical order. 
Subfamily Filograninae Rioja, 1923 
Type genus. Filograna Oken, 1815. 
Type species. Filograna implexa Berkeley, 1828. See 
Hartman (1959: 576), Kupriyanova & Jirkov (1997: 209), 
and Nogueira & ten Hove (2000: 151). 
Diagnosis. Operculum absent, or present. When present, 
may be a terminal swelling at the end of each of several 
branchial radioles as in Salmacina, or modification of first 
branchial (dorsalmost) radiole, as in type genus, Filograna ; 
neither chitinous nor calcareous. Peduncle pinnulate; inter- 
radiolar membranes absent or present; Maximum number of 
thoracic chaetigers high compared with less plesiomorphic 
taxa, even up to 12 or more. Thoracic membranes: absent 
or present; apron absent or present. Special collar chaetae 
consist of either simple blades only (i.e., lacking a fin-shaped 
basal part) or a blade and basal fin-shaped process. Posterior 
thoracic chaetigers may or may not bear sickle-shaped 
chaetae. Thoracic uncini either rasp-shaped only; or both 
rasp-shaped and saw-shaped. Abdominal neurochaetae: 
geniculate. 
Remarks. As a group considered by most workers to be 
most plesiomorphic among serpulimorphs, the subfamily 
Filograninae currently includes taxa that are variable with 
regard to characters used to separate less primitive (more 
apomorphic) taxa. Notable among them are the presence or 
absence of an operculum, thoracic membranes, collar chaetae 
possessing a fin and blade, sickles in the posterior thoracic 
chaetigers, and an apron; also, whether thoracic membranes 
are absent, as originally described for Salmacinopsis or, 
present as in the new genus Kimberleya described in this 
paper. 
The genera Filograna Oken, 1815, and Salmacina 
Claparede, 1870, had in the past been included under the 
Filograninae. McIntosh (1923) believed Filograna implexa 
Berkeley and Salmacina dysteri (Huxley) to be “synonymous 
on the grounds of their similarities in most respects and their 
frequent occurrence in the same mass of tubes. Fauvel (1930 
& 1932) did not support McIntosh’s view but distinguished 
the two genera on the basis that Filograna possesses a kind 
of operculum that is lacking in Salmacina. 
Recently, Kupriyanova & Jirkov (1997: 209) distinguish 
the genera Filograna and Salmacina “by the presence of 
two (rarely one) membranous opercula in the former and 
the absence of the operculum in the latter.” Furthermore, 
