Polyphyly of the traditional family Flabellinidae affects a major group of Nudibranchia: aeolidacean taxonomic reassessment with descriptions of several new families, genera, and species (Mollusca, Gastropoda)
Author
Korshunova, Tatiana
Author
Martynov, Alexander
Author
Bakken, Torkild
Author
Evertsen, Jussi
Author
Fletcher, Karin
Author
Mudianta, I Wayan
Author
Saito, Hiroshi
Author
Lundin, Kennet
Author
Michael Schroedl,
Author
Picton, Bernard
text
ZooKeys
2017
717
1
139
http://dx.doi.org/10.3897/zookeys.717.21885
journal article
http://dx.doi.org/10.3897/zookeys.717.21885
1313-2970-717-1
C19B43B1B3214CB1B1B2A246CEAC56BC
C19B43B1B3214CB1B1B2A246CEAC56BC
Family
Samlidae
fam. n.
Diagnosis.
Body narrow. Notum discontinuous. Cerata in separate clusters, on elevations. Rhinophores perfoliated. Anus pleuroproctic under the reduced notal edge or mixed (pleuroproctic in higher acleioproctic position). Distinct oral glands present, commonly penetrate below anterior cerata. Radula formula 1.1.1. Rachidian teeth usually with strong cusp, only rarely compressed by adjacent lateral denticles. Lateral teeth denticulated narrow or with attenuated process basally. Distal and proximal receptaculum seminis or only proximal receptaculum. Vas deferens usually short, with indistinct prostate. External permanent penial collar absent. Penis in many cases conical, narrow, always internal and unarmed.
Genera included.
Luisella
gen. n.,
Samla
Bergh, 1900.
Remarks.
The genus
Samla
and related taxa were invariably considered as synonyms of the traditional genus
Flabellina
throughout all the recent history of nudibranch taxonomy (e.g.,
Gosliner and Griffiths 1981
;
McDonald 2009
). Previously it was listed sometimes as valid (e.g.,
Burn 1964
) but without detailed discussion. However, the Indo-Pacific tropical genus
Samla
differs morphologically from the
Flabellinidae
s. str.
in
the absence of compound stalks, often non-compressed cusp of the rachidian teeth, and also by having a short vas deferens. The present molecular phylogenetic analysis not only confirmed such disparate morphology from true
Flabellinidae
, but also revealed a very profound problem: species of the genus
Samla
and some related taxa are basal to the
Aeolidacea
as a whole and the major complex of aeolidacean families, including
Aeolidiidae
,
Facelinidae
,
Tergipedidae
, and
Eubranchidae
, rendering the
Flabellinidae
polyphyletic! Such topology with minor variations has appeared invariably on all trees obtained in the present analysis and was hinted at as soon as
Flabellina babai
was sequenced (
Carmona et al. 2011
). This result is of great importance because it clearly implies that 1) Understanding of the real phylogenetic and taxonomic patterns of the traditional "
Flabellinidae
" is possible only with inclusion of all major families of the nudibranch suborder
Aeolidacea
in the analysis, as is performed in the present study, and 2) The traditional "
Flabellinidae
" cannot be by any means maintained as single family, but must be separated into several smaller morphologically and molecularly consistent families (Fig. 2). Therefore, the genus
Samla
Bergh, 1900 (
Bergh 1900b
,
1908
) is resurrected and the new family
Samlidae
is established here.
Apart from the taxa that belong to the genus
Samla
, which are clustered in a compact clade (Figs 1, 2, 45) the species
Flabellina babai
Schmekel, 1972 appears as a closely related clade.
Flabellina babai
differs considerably from
Samla
including the character of a distinct granulated prostate (
Schmekel 1972
;
Schmekel and Portmann 1982
), unique for the majority of the
Paracoryphellidae
,
Coryphellidae
and
Flabellinidae
. This feature is so far known only for the paracoryphellid genus
Chlamylla
, which is dramatically different from
Flabellina babai
externally. Therefore, a new genus for
F. babai
,
Luisella
gen. n., is proposed here (see below).