© The Norwegian Academy of Science and Letters • Zoologica Scripta, 31, 3, July 2002, pp271–319 271
Dorgan, K. M., Valdés, Á. & Gosliner, T. M. (2002). Phylogenetic systematics of the genus
Platydoris (Mollusca, Nudibranchia, Doridoidea) with descriptions of six new species. —
Zoologica Scripta, 31, 271– 319.
The genus Platydoris Bergh, 1877 is characterized by a flat body, wide mantle margin, and ele-
vated branchial and rhinophoral sheaths. Examination of a number of specimens and a review
of the literature have shown that there are at least 20 valid species of the genus Platydoris
world-wide. Some of them have previously been described by other authors and are re-
described or discussed here: P. argo, P. scabra, P. cruenta, P. angustipes, P. ellioti, P. formosa,
P. inframaculata, P. pulchra, P. sanguinea, P. townsendi, P. capricornensis, P. esakii, P. macfarlandi,
P. galbanus, P. carolynae. Six new species are described here: P. inornata, P. ocellata, P. annulata,
P. sabulosa, P. cinereobranchiata, P. rolani. Platydoris tabulata is a nomen dubium. Species removed
from Platydoris include: Doris hepatica, D. murrea, P. variegata, P. punctatella, P. papillata,
P. spinulosa, P. carinata, P. immonda, P. spongilla, P. maculata, P. stomascuta. A phylogenetic analysis
was conducted, showing the genus to be monophyletic. There was strong support for a clade
of all species of Platydoris included in the analysis except for the nodes containing P. annulata
and P. macfarlandi, which are found in deeper water than the rest of the genus. This major clade
was further divided into a clade of Atlantic and Eastern Pacific species and a clade of Indo-
Pacific species, showing a strong biogeographical trend. Important characters separating the
clades include ventral pigment around the foot, large ventral spots, the presence of vaginal
hooks, and whether the accessory gland enters the atrium on the side of the vagina, the side
of the deferent duct, or between the two.
Kelly M. Dorgan & Terrence M. Gosliner, Department of Invertebrate Zoology and Geology, Cali-
fornia Academy of Sciences, Golden Gate Park, San Francisco, CA 94118, USA. E-mail:
tgosline@calacademy.org
Ángel Valdés, Natural History Museum of Los Angeles County, 900 Exposition Boulevard, Los
Angeles, CA 90007, USA. E-mail: avaldes@nhm.org
Blackwell Science, Ltd
Phylogenetic systematics of the genus Platydoris (Mollusca,
Nudibranchia, Doridoidea) with descriptions of six new species
KELLY M. DORGAN, ÁNGEL VALDÉS & TERRENCE M. GOSLINER
Accepted: 2 January 2002
Introduction
The genus Platydoris Bergh, 1877 contains several generally
large species of dorid nudibranchs, distributed throughout
tropical and temperate areas in the Atlantic, Eastern Pacific
and Indo-Pacific. Species of Platydoris are easily recognized
by their flattened, normally rigid (or coriaceous) body. This
genus is also characterized by the dorsum covered with very
small caryophyllidia (Foale & Willan 1987), a generally
lobate branchial sheath, a non-tubular prostate with two
different portions, a penis (and in some cases a vagina) armed
with hooks, a lobate accessory gland, a smooth labial cuticle,
and hamate radular teeth (Bergh 1877a; Valdés & Gosliner
2001). Species of Platydoris are mostly found in shallow
waters, but some species have been described from deep
waters (e.g. P. macfarlandi, P. annulata).
A review of the literature reveals that most species of
Platydoris are poorly described and are either unidentifiable
or synonyms of other species of this or other genera. This paper
attempts to review the taxonomic status of the described
species of Platydoris, based on a re-examination of available
type material and the original description of each species.
In addition, six new species from the tropical Atlantic and
Indo-Pacific are described, and 11 species are re-described.
Platydoris was recently shown to be monophyletic, and
closely related to Baptodoris Bergh, 1884 and Gargamella
Bergh, 1894 (Garovoy et al. 1999; Valdés & Gosliner 2001).
However, the phylogenetic relationships within the members
of Platydoris were unknown. This paper examines the phylog-
eny of the species of Platydoris.
Materials and methods
The studied specimens are deposited at the Department of
Invertebrate Zoology of the California Academy of Sciences
(CASIZ). Type specimens of several species were obtained