Description of A New Species of Thordisa (Nudibranchia: Discodorididae) from Panama

Abstract

A new species of Thordisa is described from the Caribbean coast of Panama. The phy-logeny of the genus Thordisa from the tropical Indo-Pacific and eastern Pacific is revised. Morphological and anatomical data from Thordisa species were used to con-struct a phylogeny with increased resolution. The phylogenetic analysis demonstrates the monophyly of Thordisa and its relationship to outgroups Asteronotus, Halgerda, and Hoplodoris. The dorid nudibranch genus Thordisa Bergh, 1877, consists of 31 species currently recognized as valid. The genus has been monographed recently (Chan and Gosliner, in press). Several species of Thordisa have been recently described by different authors (Chan and Gosliner, in press; Ortea and Valdés 1995; Cervera and García-Gómez 1989; Behrens and Hendersen 1981; Lance 1966). One new species of Thordisa is described from the Caribbean coast of Panama. A revised analysis of the phylogeny of the genus Thordisa is presented here, using three outgroup taxa: Asteronotus cespitosus van Hasselt, 1824, Halgerda dalanghita Fahey and Gosliner, 1999, and Hoplodoris estrelyado Gosliner and Behrens, 1998. Additional data were taken from the original publications on Thordisa (Marcus 1955; Pease 1860). The following species were examined directly: T. bimaculata Lance, 1966; T. filix Pruvot-Fol, 1951; T. rubescens Behrens and Henderson, 1981; T. sanguinea Baba, 1955; T. azmanii Cervera y García-Gómez, 1989; T. oliva Chan and Gosliner, 2006; T. luteola Chan and Gosliner, 2006; T. albomacula Chan and Gosliner, 2006; T. nieseni Chan and Gosliner, 2006; and T. tahala Chan and Gosliner, 2006. Thus, previous literature and direct observation and dissection of 14 species of Thordisa and members of the outgroup have provided the information on the characters for the present study. Thordisa harrisi Chan and Gosliner, sp. nov.

Full text

Description of A New Species of Thordisa (Nudibranchia:
Discodorididae) from Panama
Jamie M. Chan and Terrence M. Gosliner
Department of Invertebrate Zoology and Geology, California Academy of Sciences,
875 Howard St., San Francisco, California 94103; Email: tgosliner@calacademy.org
A new species of Thordisa is described from the Caribbean coast of Panama. The phy-
logeny of the genus Thordisa from the tropical Indo-Pacific and eastern Pacific is
revised. Morphological and anatomical data from Thordisa species were used to con-
struct a phylogeny with increased resolution. The phylogenetic analysis demonstrates
the monophyly of Thordisa and its relationship to outgroups Asteronotus, Halgerda,
and Hoplodoris.
The dorid nudibranch genus Thordisa Bergh, 1877, consists of 31 species currently recognized
as valid. The genus has been monographed recently (Chan and Gosliner, in press). Several species
of Thordisa have been recently described by different authors (Chan and Gosliner, in press; Ortea
and Valdés 1995; Cervera and García-Gómez 1989; Behrens and Hendersen 1981; Lance 1966).
One new species of Thordisa is described from the Caribbean coast of Panama. A revised analysis
of the phylogeny of the genus Thordisa is presented here, using three outgroup taxa: Asteronotus
cespitosus van Hasselt, 1824, Halgerda dalanghita Fahey and Gosliner, 1999, and Hoplodoris
estrelyado Gosliner and Behrens, 1998.
Additional data were taken from the original publications on Thordisa (Marcus 1955; Pease
1860). The following species were examined directly: T. bimaculata Lance, 1966; T. filix Pruvot-
Fol, 1951; T. rubescens Behrens and Henderson, 1981; T. sanguinea Baba, 1955; T. azmanii
Cervera y García-Gómez, 1989; T. oliva Chan and Gosliner, 2006; T. luteola Chan and Gosliner,
2006; T. albomacula Chan and Gosliner, 2006; T. nieseni Chan and Gosliner, 2006; and T. tahala
Chan and Gosliner, 2006. Thus, previous literature and direct observation and dissection of 14
species of Thordisa and members of the outgroup have provided the information on the characters
for the present study.
Thordisa harrisi Chan and Gosliner, sp. nov.
Figures 1–4.
MATERIAL EXAMINED.— HOLOTYPE: CASIZ 171542 one preserved specimen, Bocas del Toro,
Panama, 2 m depth, 19 June 2005, Larry Harris, length 35 mm (dissected).
DISTRIBUTION.— This species is only known from the Caribbean coast of Panama (present
study).
ETYMOLOGY.— This species was named for Larry Harris, Professor of Zoology, University of
New Hampshire, who has been a mentor and a colleague of the second author for many years. Larry
collected the holotype of this new species.
EXTERNAL MORPHOLOGY.— The body of the living animal is oval with a low profile (Fig. 1A).
PROCEEDINGS OF THE CALIFORNIAACADEMY OF SCIENCES
Fourth Series
Volume 57, No. 35, pp. 981–990, 4 figs., 2 tables December 28, 2006
981

The notum bears long, villous papillae and
rounded tubercles. The longest papillae are con-
centrated in the middle of the dorsum (Fig. 1B).
Some of the longest papillae are bifurcated at
the base. The rhinophores are perfoliate with
20-25 lamellae each. The rhinophoral sheath is
highly papillate, with 15 papillae along the
inner margin. The gill is completely retractile
and surrounded by a low even sheath. The six
gill leaves are tripinnate and do not extend
beyond the edge of the notum. The anterior
margin of the foot is bilabiate (Fig. 1C). The
oral tentacles are digitiform and do not extend
beyond the margin of the foot. The notum of the
living specimens has a bright orange ground
color with translucent papillae and tubercles.
There are small rust-colored spots dispersed
over the notum and two brown oval spots in the
center that extend from the anterior of the gill
pouch toward the rhinophores. The rhinophores
and gill are opaque brown. The color of the foot
of the live specimens is solid orange. On the
ventral side of the mantle is a small red ring of
dots surrounding the foot.
ANATOMY.— The labial cuticle is smooth
and devoid of rodlets. The radula formula is
34x 2-3.1.33-35.0.33-35.1.2-3 at the 15th row
from the anterior of the radula. The innermost
lateral teeth are smaller, hamate and have a
large, wide base (Fig. 2A). The middle lateral
teeth are hamate with slender bases and long
finger-like tips (Fig 2B). They slightly increase
in size toward the margin. The third tooth from
the margin is bifurcated at the tip (Fig. 2C). The
two to three outer teeth are thinner, having fine
pectination at the tips (Fig. 2D). The central
nervous system consists of partially fused cere-
bral and pleural ganglia. The pedal ganglia are
situated ventrally and extend outside the junc-
tion of the cerebropleural ganglia. They are
connected by a circum-esophageal nerve ring (Fig. 3B). The stomach is partly free and medial, and
rests on the digestive gland. The digestive gland is approximately twice the length of the stomach.
The intestine is straight and dorsally situated.
The ampulla is straight and then curves once before ending in the female gland mass (Fig. 3A)
via a short oviduct. The ampullary hermaphroditic duct is three times the length of the duct con-
necting the prostate. The base of the ampulla meets the uterine duct where the prostate gland begins.
The prostate is approximately the same length as the ampulla. The more proximal white portion of
982 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Fourth Series, Volume 57, No. 35
FIGURE 1. Photograph of living animal Thordisa harrisi
sp. nov. (CASIZ 171542) A. dorsal view. B. Dorsal/side view.
C. ventral view. Photos by Sigmer Quiroga.

CHAN & GOSLINER: NEW SPECIES OF THORDISA FROM PANAMA 983
FIGURE 2. Thordisa harrisi sp. nov. (CASIZ 171542) Scanning Electron Micrographs A. inner lateral teeth B. middle
lateral teeth C. outer lateral teeth. D. close up of outer lateral teeth.
FIGURE 3. Thordisa harrisi sp. nov. (CASIZ 171542). A. Reproductive System. Scale bar = 1.0 mm. Abbreviations: am,
ampulla; bc, bursa copulatrix; ej, ejaculatory duct; fg, female gland; pr, prostate; rs, receptaculum seminis; v, vagina; vg,
vestibular gland. B. Central nervous system. Scale bar = 1.0 mm. Abbreviations: cnr, circum-oesophageal nerve ring; e, eye;
lcpg, left cerebral-pleural ganglia; rpg, right pedal ganglia; lrn, left rhinophoral nerve.

the prostate is longer and thinner than the distal yellow portion. The muscular portion of the vas
deferens is long and slightly curved. The penial sheath is unarmed. A bi-lobed, unarmed vestibular
gland connects to a separate vestibule adjacent to the genital atrium. The lobes of the gland are long,
slender and finger-like. The unarmed vaginal duct is straight and thinner than the vas deferens. It
leads to the bursa copulatrix, which is approximately twice as large as the receptaculum seminis.
The duct of the receptaculum seminis is short and straight.
DISCUSSION.— This is the second Thordisa species known from Panama. The first species
described from the Pacific coast is T. nieseni (Chan and Gosliner 2006), while the present species
is found on the Caribbean side of the isthmus. Thordisa harrisi, can be distinguished from T. nieseni
by its bright yellow mantle color, long slender vestibular glands, and bifurcated outer tooth.
Thordisa nieseni is a bright red mantle color and has pectinate outer teeth.
Thordisa harrisi is similar to a recently described discodorid species “aliciae“ (Dayrat 2005).
Discodorididae “aliciae” is found on the Pacific coast in Panama and has a similar external mor-
phology, yet lacks both pectination of the marginal teeth and a vestibular gland. We re-examined
the type material of “aliciae” housed in the collection at the California Academy of Sciences and
confirmed the absence of a vestibular gland. Discodorididae “aliciae” also has a series of black
spots on the undersurface of the mantle that are absent in Thordisa harrisi.
PHYLOGENETIC ANALYSIS OF THORDISA.— The character states recorded for each species of
Thordisa, were placed into a data matrix from MacClade version 4.0 (Maddison and Maddison
2000) (Table 2). All characters used have equal weight and are unordered (Table 1). Six characters
were deleted from the analysis due to being phylogenetically uninformative. The characters delet-
ed are the morphology of the rhinophore sheath (2), the rachidian tooth (4), the relative size of the
bursa copulatrix compared to the receptaculum seminis (16), the color of the mantle (19), the man-
tle pattern (20) and the rhinophore color (23). Characters 19 and 20 were omitted because their
states had many intermediate conditions. The data were analyzed by Phylogenetic Analysis Using
Parsimony (PAUP*) version 4.0 (Swofford 2003). A heuristic search was performed with the opti-
mality criterion of maximum parsimony. The stepwise addition option of random trees was used,
with 100 repetitions, starting from random start trees. Bremer support values (Bremer 1988) were
calculated to estimate branch support using PAUP*. Character tracing was performed to understand
the characters that united resulting clades.
To establish the polarity of the morphological characters used in this study, three outgroup taxa
(Asteronotus cespitosus, Hoplodoris estrelyado, and Halgerda dalanghita) were used based a
review of anatomical characters as described in Valdés and Gosliner, 2001; Gosliner and Behrens,
1988 and Fahey and Gosliner, 1999. We used these outgroups based on the phylogenetic study by
Valdés (2002). In Valdés’ study, these taxa were more closely related to Thordisa than other taxa.
The following characters were considered in this analysis:
[The characters preceded by an asterisk were deleted from the final analysis because they were phylogeneti-
cally uninformative) (0) = presumed pleisiomorphic (1) = apomorphic, ? = missing data or not applicable.]
1. Labial pits.— The derived character state is only found in T. oliva and T. diuda (1). In all
other species the pits are absent (0).
2. *Rhinophoral sheath.— There is observed intraspecific variation within the rhinophore
sheaths of Thordisa. Some species are observed to have scalloped edges (1) and others are straight.
The outgroups Asteronotus cespitosus, Hoplodoris estrelyado, and Halgerda dalanghita are all
known to have straight sheaths (0).
3. Gill.— All of the outgroup taxa, Asteronotus cespitosus, Hoplodoris estrelyado, and
Halgerda dalanghita have tripinnate gills (0). Four of the examined species, T. nieseni, T. azmanii,
984 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Fourth Series, Volume 57, No. 35

T. villosa and T. filix have bipinnate gills (1). T. sabulosa Burn, 1957 is the only other Thordisa that
has been described with bipinnate gills.
4. *Rachidian tooth.— The rachidian tooth is absent in all specimens (0).
5. Inner lateral teeth compared to outer lateral teeth.— The inner lateral teeth of most
species were considerably smaller than middle lateral teeth (1). Four Thordisa (T. albomacula, T.
azmanii, T. villosa, and T. tahala) have inner teeth that are equal to the size of the middle laterals
(0) as do the outgroup taxa.
6. Shape of inner lateral teeth.— The shape of the inner lateral teeth can be simply hamate,
denticulate or bifurcate. Only Thordisa oliva was observed to have consistently bifurcated inner
teeth (2). Thordisa nieseni can have bifurcated tips or simply hamate inner teeth (2). Thordisa albo-
macula, T. setosa, and T. tahala have denticulate inner teeth (1) whereas the remaining nine species
have simply hamate inner teeth (0).
7. Shape of middle lateral teeth.— The middle lateral teeth are either denticulate or simply
hamate. Asteronotus cespitosus, Hoplodoris estrelyado and Halgerda dalanghita have smooth mid-
dle teeth (0). Thordisa oliva, T. nieseni, T. villosa and T. luteola have smooth hamate middle teeth.
Thordisa tahala, T. albomacula, and T. setosa have denticulate middle lateral teeth (1).
8. Shape of outer lateral teeth.— The outer lateral teeth of all Thordisa are pectinate (1).
Asteronotus cespitosus, Hoplodoris estrelyado and Halgerda dalanghita have simple teeth (0).
CHAN & GOSLINER: NEW SPECIES OF THORDISA FROM PANAMA 985
TABLE 1. Characters and states considered for the phylogeny of Thordisa.

9. Vestibular gland.— The vestibular gland is present (1) in all Thordisa except T. hilaris (Kay
and Young 1969). A gland is also present in Asteronotus cespitosus and Hoplodoris estrelyado (1).
A vestibular gland is absent in Halgerda dalanghita (0).
10. Number of vestibular glands.— Thordisa has either one (0) or two vestibular glands (1).
Two vestibular glands occur in six of the recently described species. This character is not applica-
ble to Halgerda dalanghita (?).
11. Vestibular gland shape.— Vestibular glands were categorized into three distinct shapes:
pouch, coiled and lobate. T. filix is the only Thordisa observed to have a long and coiled vestibular
gland (1). Three species (T. albomacula,T. tahala and T. setosa) have lobate vestibular glands (2).
The remaining taxa have vestibular glands that are pouch shaped (0). This character is not applica-
ble to Halgerda dalanghita (?).
12. Vestibular gland spines.— The presence of vestibular glands spines occurs in seven
Thordisa species and Asteronotus cespitosus and Hoplodoris estrelyado (1). This character is not
applicable to Halgerda dalanghita and is unknown for Thordisa diuda and T. setosa (?). Absence
of spines is coded (0).
13. Number of vestibular gland spines.— The number of vestibular gland spines increases
with the number of vestibular glands. Thordisa azmanii, T. bimaculata, T. villosa, T. oliva, and T.
tahala all have one vestibular gland spine (0). T. nieseni, T. rubescens, and T. luteola all have two
vestibular gland spines (1). Halgerda dalanghita, Thordisa albomacula, T. diuda, T. filix, T. san-
guinea, T. harrisi and T. setosa do not possess vestibular gland spines. In several species, it is either
not applicable or unknown (?).
14. Vaginal spines.— Vaginal spines are present in Thordisa luteola, and T. rubescens and (1).
This character is unknown for Thordisa bimaculata, T. diuda, T. filix, and T. setosa (?). Absence of
spines is coded (0).
15. Penial spines.— Penial spines usually occur as a single large spine or a series of small
spines surrounding the penis. Penial spines occur in five of the Thordisa species (1). This character
is unknown for T. rubescens and T. setosa (?). Absence of spines is coded (0).
16. *Bursa copulatrix vs. receptaculum seminis.— The receptaculum seminis (rs) is small-
er than the bursa copulatrix (bc) in all the taxa (0) examined except for Thordisa sanguinea.
17. Papillae.— Villous papillae occur in thirteen of the Thordisa species (1). Thordisa bimac-
ulata and all the outgroup taxa have shorter papillae throughout their mantle (0).
18. Compound papillae.— Compound papillae are only found in Thordisa luteola and
Thordisa harrisi (1). Absence of papillae is coded (0).
19. *Mantle color.— Mantle color was categorized into four color groups, a brown/black/olive
group (0), a tan/translucent color (1), yellow colored (2) and an orange/red color (3).
20. *Mantle pattern.— There are distinctive mantle patterns (0) found in species studied with
the exception of Thordisa azmanii, T. filix, T. luteola, T. nieseni, and T. oliva, which all possess a
solid colored mantle (1).
21. Tubercle or ridge color vs. mantle color.— Tubercle or ridge color is a contrasting color
to mantle color (0) in all species with the exception of Thordisa filix, T. nieseni and T. sanguinea
(1). Halgerda dalanghita does not possess tubercles on its mantle (?).
22. Gill color.— The gill of all species was light in color (0) with the exception of Thordisa
luteola, T. rubescens, T. harrisi, and Hoplodoris estrelyado (1).
23. *Rhinophore color.— The rhinophore color is dark for Thordisa villosa, T. rubescens, T.
sanguinea, T. setosa, T. harrisi, Halgerda dalanghita, Asteronotus cespitosus and Hoplodoris
estrelyado (0). No variation among species was observed and this character was excluded.
986 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Fourth Series, Volume 57, No. 35

RESULTS
One parsimonious tree was produced (Fig. 4). The tree required 37 steps and had a consisten-
cy index of 0.541 and retention index of 0.653. Bremer support was (1) for all branches except for
clades A and E, which have a Bremer support of (2), as does the clade containing Halgerda
dalanghita and Asteronotus cespitosus.
Clade A corresponds to the ingroup, Thordisa. Clade A is united by the presence of inner lat-
eral teeth that are smaller than the middle lateral teeth (5), pectinate outer laterals (8), one vestibu-
lar gland (10), and the presence of penial spines (15). This clade includes 14 taxa classified in the
genus Thordisa. Clade B is united by the presence of villous papillae (17). Clade C is united by the
absence of vestibular gland spines (12). Clade D is united by the presence of labial pits (1). Clade
E is united by the presence of denticulate inner lateral teeth (6), denticulate middle lateral teeth (7),
and a lobate vestibular gland (11). Clade F is united by a reversal from the possession of two
vestibular glands to a single one (13). Clade G is united by a reversal of the presence of inner lat-
eral teeth equal in size to middle lateral teeth (5). Clade H is united by the absence of penial spines
(15). Clade I is united by the presence of two vestibular glands (10). Clade J is united by the pres-
ence of vestibular gland spines (12), which are two in number (13). Clade K is united by the pres-
ence of bipinnate gills (3). Clade L is united by the presence of vaginal spines (14) and a dark gill
color (22). Clade M is united by the reversal of the presence of inner lateral teeth equal in size to
the middle lateral teeth (5).
CHAN & GOSLINER: NEW SPECIES OF THORDISA FROM PANAMA 987
TABLE 2. Data Matrix. Character states in species of Thordisa and the outgroup taxa
Asteronotus, Halgerda, and Hoplodoris.
? = missing or not applicable, 0 = presumed pleisiomorphic,
1 = apomorphic, 2 = apomorphic 3 = apomorphic

988 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Fourth Series, Volume 57, No. 35
FIGURE 4. Preliminary phylogeny of Thordisa. Strict consensus tree of one most parsimonious trees. Asteronotus,
Halgerda, and Hoplodoris were chosen as outgroup taxa to polarize the characters. T=Thordisa. Numbers above lines are
character numbers: underlined numbers indicate reversals and numbers in parentheses are character states (Table 1). Letters
A to J indicate the 10 clades that constitute the hierarchical pattern resulting from the analysis. Numbers below are Bremer
support values.

DISCUSSION
Our analysis demonstrates the monophyly of Thordisa and its relationship to the outgroups
Asteronotus, Halgerda and Hoplodoris. Pectinate outer lateral teeth are a strong synapomorphy of
the genus. Characters of the reproductive system such as vestibular gland morphology and genital
armature can prove to be important for distinguishing species of Thordisa. The unique pits on the
mouth of Thordisa diuda and Thordisa oliva are significant for uniting the two sister species.
However, re-examination of other type species should be done to confirm absence or presence of
this feature. It is clear that there are several homoplastic characters in our analysis such as teeth size
and the number of vestibular glands. Much of the topology of our current tree is similar to the first
preliminary phylogeny of Thordisa (Chan and Gosliner, in press). The current tree has increased
resolution. In particular, the previous trichotomy containing Thordisa villosa, T. nieseni and T.
azmanii is now fully resolved to show T. azmanii and T. villosa as more closely related to each other
than to T. nieseni. The newly described species Thordisa harrisi is the most basal member of Clade
H. Many species of Thordisa are incompletely described and will need re-evaluation as more mate-
rial becomes available. Further character analysis and testing of the data are needed to strengthen
the phylogenetic hypothesis of the genus Thordisa.
ACKNOWLEDGEMENTS
The authors would like to thank Larry Harris and Shireen Fahey for collection and preserva-
tion of the specimen. This research was supported by the California Academy of Sciences and the
NSF PEET Grant # 0329054 to T. Gosliner and A. Valdés.
LITERATURE CITED
BABA, K. 1955. Opistobranchia of Sagami Bay. Supplement. Iwanami Shoten, Tokyo, Japan. 39 pp.
BEHRENS, D., AND R. HENDERSEN. 1981. Two new cryptobranch dorid nudibranchs form California. The Veliger
24(2):120–128.
BREMER, K. 1988. The limits of amino acid sequence data in angiosperm phylogenetic reconstruction.
Evolution 42:785–803.
CERVERA, J., AND J. GARCÍA-GÓMEZ. 1989. A new species of the genus Thordisa (Mollusca: Nudibranchia)
from the southwestern Iberian peninsula. The Veliger 34(4):382–386.
CHAN, J., AND T. GOSLINER. (In press.) Preliminary Phylogeny of Thordisa (Nudibranchia: Discodorididae)
with Descriptions of Five New Species. The Veliger 48(3).
DAYRAT, B. 2005. Advantages of naming species under the Phylocode: An example of a new species of
Discodorididae (Mollusca, Gastropoda, Euthyneura, Nudibranchia, Doridina) may be named. Marine
Biology Research 1:216–232.
FAHEY, S., AND T. GOSLINER. 1999. Description of three new species of Halgerda from the Western Indian
Ocean with a rediscription of Halgerda formosa, Bergh 1880. Proceedings of the California Academy of
Sciences, ser. 4, 51(8):365–383.
GOSLINER, T.M., AND D. BEHRENS. 1998. Two new discodorid nudibranchs from the western Pacific with a
redescription of Doris luteola Kelaart, 1858. Proceedings of the California Academy of Sciences, ser. 4,
50(11):279–293.
GOSLINER, T.M. 1987. Nudibranchs of Southern Africa. A Guide to the Opisthobranch Molluscs of Southern
Africa. Sea Challengers, Monterey, California, USA. 136 pp.
LANCE, J.R. 1966. New distributional records of some northeastern Pacific opisthobranchiata (Mollusca:
Gastropoda) with descriptions of two new species. The Veliger 9(1):69–81.
MADDISON, W., AND D. MADDISON. 2000. MacClade, version 4. Cambridge, Massachusetts, USA. [Distributed
by authors.]
CHAN & GOSLINER: NEW SPECIES OF THORDISA FROM PANAMA 989

MARCUS, E. 1955. Opistobranchia from Brazil. Zoologia 20:140–142 pl. 15.
ORTEA, J.A., AND A. VALDES. 1995. Una nueva especie de Thordisa Bergh, 1877 (Mollusca: Nudibranchia:
Discodorididae) de las costas Angola. Avicennia 3:35–41.
PEASE, W. 1860. Descriptions of new species of mollusca from the Sandwich Islands. Proceedings of the
Zoological Society of London 28:18–36.
SWOFFORD, D. 2003. PAUP* Phylogenetic Analysis Using Parsimony (*and other methods), version 4.0.
Sinauer Associates, Sunderland, Massachusetts, USA.
VALDÉS, Á. 2002. A phylogenetic analysis and systematic revision of the cryptobranch dorids (Mollusca,
Nudibranchia, Anthobranchia). Zoological Journal of the Linnean Society 136:535–636.
VALDÉS, Á., AND T.M. GOSLINER. 2001. Systematics and phylogeny of the caryophyllidia-bearing dorids
(Mollusca, Nudibranchia), with the description of a new genus and four new species from the Indo-Pacific
deep waters. Zoological Journal of the Linnean Society 133:103–198.
990 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Fourth Series, Volume 57, No. 35
Copyright © 2006 by the California Academy of Sciences
San Francisco, California, U.S.A.