PROCEEDINGS

OF THE

CALIFORNIA ACADEMY OF SCIENCES

Volume 51

Published 1999 by the

California Academy of Sciences
Golden Gate Park
San Francisco, California

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PROCEEDINGS

OF THE

CALIFORNIA ACADEMY OF SCIENCES

Volume 51

Published 1999 by the

California Academy of Sciences
Golden Gate Park
San Francisco, California

SCIENTIFIC PUBLICATIONS DEPARTMENT

Alan E. Leviton, Editor
Katie Martin, Managing editor
Juliet Knowles, Editorial assistant

ISSN 0068-547X

© California Academy of Sciences
Golden Gate Park
San Francisco, California 94118

PRINTED IN THE UNITED STATES OF AMERICA
BY ALLEN PRESS, INC., LAWRENCE, KANSAS 66044

TABLE OF CONTENTS
VOLUME 51

1. FERRARIS, JR., CARL and MARIO C.C. DE PINNA. Higher-level names for catfishes (Actinop-
fenyoi/Ostaniophysis siluntormes): (lus) March 295 1990F ae ee se sone 1-17
2. WILLIAMS, GARY C. Index Pennatulacea: annotated bibliography and indexes of the sea pens
(Coelenterata: Octocorallia) of the world 1469-1999. (Illus.)
cae) MOOS eh Aare xt ah Meee Rasen 1A os ms peeenye athe ole Sete Mere syne ieee CAA 19-103
3. IWAMOTO, TOMIO and ALAN WILLIAMS. Grenadiers (Pisces, Gadiformes) from the continental
slope of western and northwestern Australia. (Illus.)
JW) ZO SASS rece bce By eae RC iS oy ech SOP RAE aL te rere otra 105-243
4. GAROVOY, JOCELYN B., ANGEL VALDES and TERRENCE M. GOSLINER. Two new species of
Gargamella (Mollusca, Nudibranchia) from South Africa. (Illus. )
Vitel LO) aS IOS ee esata eres aca is sere ana ey eshte) Ae MS RS IRE PR nec 245-257
5. BRADBURY, MARGARET G. A review of the fish genus Dibranchus with descriptions of new
species and a new genus, Solocisquama (Lophiiformes, Ogcocephalidae). (Illus. )

PROV OUIS Mal ete, area a Sah ie Ie Sc date pista totes gt COO. WAL ST we Ghetto ata ene aa 259-310
6. BOUNDY, JEFF. Systematics of the garter snake Thamnophis atratus at the southern end of its
FAC ee ILS) PAU OUSE 2 Or MOOG ei 2 ope Rue ae ee a Reece SpA eh ts eae 311-336

7. WILLIAMS, GARY C. and PHILIP ALDERSLADE. Revisionary systematics of the western Pacific
soft coral genus Minabea (Octocorallia: Alcyoniidae), with descriptions of a related new genus
an@mpeciesmiom: the Indo-Paciic: (lllus:) October 341999. se. 2 ss sees ee 337-364

8. FAHEY, SHIREEN J. and TERRENCE M. GOSLINER. Descriptions of three new species of
Halgerda from the western Indian Ocean with a redescription of Halgerda formosa, Bergh
HS SOM Muss) KOCIObER.8.lOQO me oo ale downy Veda krdste een rotary ee oe sees 365-383

9. HEOK HEE, NG. A review of the southeastern asian catfish genus Ceratoglanis (Siluriformes:
Siluridae), with the description of a new species from Thailand. (Illus.)

Weiobe4ng WslOGO eran tent nor ACAD kesh wk es A Noes ON aac ren ones Oy atop eee eee 385-395
10. FERRARIS JR., CARL J. and KATHRYN E. RUNGE. Revision of the south Asian bagrid catfish
genus Sperata, with the description of a new species from Myanmar. (IIlus.)
CiSO) SIE A IIS a egrets ee te ae ei Pn ASAP Ene Renae a oe Ne Aen ER oss. 397-424
11. FAHEY, SHIREEN J. and TERRENCE M. GOSLINER. Preliminary phylogeny of Halgerda (Nudi-
branchia: Halgerdidae) from the tropical Indo-Pacific, with descriptions of three new species.
MULES ENOVEMIDCTEL Oe OOF ee re Se cot Ms Sie Be Be Nae Gee che ytd eas ee ae a 425-448
12. BROWN, WALTER C., ANGEL C. ALCALA, and ARVIN C. DIESMOS. Four new species of the
genus Platymantis (Amphibia: Ranidae) from Luzon Island, Philippines. (IIlus.)
INH LA OLS 2 ae LS 21) ees See an pean ithe ic son ae ey et ccna Ca etcain croacla. Fiano io -6' Gch 449-460
13. VALDES, ANGEL, ERNESTO MOLLO, and JESUS ORTEA. Two new species of Chromodoris
(Mollusca, Nudibranchia, Chromodorididae) from southern India, with a redescription of
Chromodoris trimarginata (Winckworth, 1946). (Illus.) November 23, 1999 ....... 461-472
14. RANDALL, JOHN E. and JOHN L. EARLE. Acanthurus reversus, a new species of surgeonfish
(Perciformes: Acanthuridae) from the Marquesas Islands. (Illus.) December 9, 1999.. 473-481
15. DANIEL, THOMAS F. Taxonomic and distributional notes on neotropical Justicia (Acan-
thaceac)(liltus) December 999 Was eae aan oes sie ie ee eee eo ae 483-492

Indexsto, Volume sil May tO! 20004 <5 sn oc bate Bo oa aig shale ey At agee ene eel 493-505

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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES

Volume 51, No. 1, pp. 1-17. March 29, 1999

Higher-level Names for Catfishes
(Actinopterygii: Ostariophysi: Siluriformes)

by AP;
Carl J. Ferraris, Jr.

Department of Ichthyology, California Academy of Sciences
Golden Gate Park, San Francisco, California 94118

and

Mario C. C. de Pinna
Department of Zoology, University of Sao Paulo, Caixa Postal 11461,
Sao Paulo, SP 05422-970, Brazil

A comprehensive list is provided of all family-group and other suprageneric names
proposed for the order Siluriformes. The list contains 169 names including both available
and unavailable names, whether in current usage or not. Dates and authorship for all
names are provided. In view of the present intense research activity on catfish phylogeny
and classification, this list will be a useful reference for future work, avoiding unnecessary
proposal of new names and facilitating decisions on priority.

During the past few decades, there has been a growing interest in the study of higher-level
relationships among catfishes (Ostariophysi: Siluriformes). This interest has resulted in several
doctoral dissertations and publications on phylogenetic relationships of one or more families of
catfishes. A number of these studies have included revisions in the classification of families, with the
recognition of additional monophyletic groups above the generic level. Many of these newly-recog-
nized suprageneric groups have previously been assigned formal family-level names. Others with
informal names will likely be given formal names in the future. The tracing of family-level groups
and their dates can be difficult. A number of obscure, but available, names have been proposed in rare
or inaccessible publications and others are buried in texts on unrelated groups making their discovery
unlikely for the specialist. Because of that, the assignment of family-group names to the classification
of various catfish groups has on occasion been mistaken in both the older and modern literature. We
believe that the piecemeal correction of these errors is detrimental to the stability of the classification
of catfishes.

In anticipation of future phylogenetic studies, the results of which may necessitate the recognition
of more family-level groups of catfishes, we have attempted to collect all previously proposed names
among catfishes. This provides a comprehensive list that will serve as a general reference for future
classificatory works on siluriforms. We made an effort to locate all relevant names, whether or not
in current use. As groupings of catfish are reexamined, it is possible that two or more currently used
names may be synonymous at some level. Following the rules of priority, it would then be necessary
to establish which name was proposed first. To that end, we have attempted to provide the earliest
use for each of the names.

Unlike generic and specific names, use of family-level names has historically been rather
informal in ichthyology and, we believe, in zoology as a whole. Names are often cited without

2 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 1

authorship or with no indication of the date of first usage. The initial proposal of family-level names
often lacks an indication that the author intended to coin a new name. The rules that govern the
formation of new family-level names are considerably less stringent than those for names at the
generic and specific level. While not advocating more strict requirements, we believe that everyone
benefits if new names are proposed with care, and that proper attribution should be made when
family-level names are used.

METHODS

Throughout the paper, we refer to the third edition of the International Code of Zoological
Nomenclature (ICZN 1985) as the Code.

The name list was built on the basis of statements in the literature, deliberate search for additional
names in references, and fortuitous encounters during our reading on other topics. We benefited
greatly from access to a copy of Henry W. Fowler’s draft list of fishes of the world that is on file at
the California Academy of Sciences. Fowler attempted to document the earliest usage of every
family-level name and the first proposed usage at each level within the range of family-level taxa.
We checked each citation by Fowler, and in many cases found earlier usage of a name. In some cases
we have not been able to confirm the names listed in the publications which he cited.

In the process of our search, we considered only works that are unquestionably published, thus
excluding dissertations and similar documents. This follows the criteria of publication of the Code
(Arts. 8, 9). Only names that were clearly proposed as suprageneric are included. Accordingly, all
names proposed in Cuvier (1816) and Cuvier and Valenciennes (1840), except Siluroides, are
unavailable because there is no evidence that they were applied at a suprageneric rank.

Names in the following list represent the earliest form we encountered of what we believe to be
a suprageneric scientific name. We have included names that have their basis in a generic name, as
well as names that have been proposed without an underlying generic name. All of the latter are
unavailable by our reading of the Code, and some of the former are likewise unusable for other reasons.
We chose to include all names in order to provide the most complete record of names proposed for
catfishes. The Code changes, and it is possible that future versions of the Code may change the status
of currently unavailable names.

In attempting to determine the availability of the names listed below, we encountered two
provisions of the Code that relate to several names, and we think it best to summarize our thoughts
here, rather than repeat our statements in each relevant account. Article 13 of the Code requires that
all new names published after 1930 be “accompanied by a description or definition that states in words
characters that are purported to differentiate the taxon.” This requirement was first stated in the second
edition of the International Code of Zoological Nomenclature (ICZN 1961), but backdated to 1930.
No equivalent provision was in force for family-group names previous to 1961, although a recom-
mendation to that effect was put forth in the Copenhagen Decisions on Zoological Nomenclature
(Hemming 1953). Therefore, a family-group name first published between 1930 and 1961 that was
based on an available generic name but without a diagnosis was validly proposed when published,
but is not now available because of the provisions of Art. 13. Within catfishes, family-group names
published in Fowler (1935), Hoedeman (1949), Fowler (1951) and Fernandez-Yépez and Martin
(1953) fall into this category. By our reading of the Code these names may be made available again
by use of the name in conjunction with a diagnosis, as was done in Lundberg et al. (1991), for the
name Pseudopimelodinae.

Article 29 of the Code requires that family-group names be formed by the addition of a prescribed
suffix to the stem derived from a generic name. Names first proposed prior to 1900 are exempt from
the requirement of a suffix [Art. 11(f)iii], provided the names have been subsequently adopted with
an appropriate suffix. The first proposal of several catfish family-group names (e.g., Loricaria

FERRARIS AND DE PINNA: CATFISH FAMILIES 3

Rafinesque, 1815, Siluroides Cuvier, 1816, and Hypostomiden Kner, 1853) fit into this category and
are, by our interpretation, available. An unusual case is provided in the work of Bleeker (1858).
Suprageneric names of the ranks of subfamily, stirps, and cohors were provided with standardized
suffixes added to generic stems. The type genera of the names were not designated but can be inferred
from the generic stem. However, in two cases, subfamily-level names are somewhat ambiguous with
regard to the type genus. The Silurichthyoidei included the genera Silurus and Silurichthys, and the
Bagrichthyoidei included Bagrus and Bagrichthys. Bleeker (1858) uniformly used the complex suffix
-ichthyoidei for subfamily names (e.g., Sisorichthyoidei for Sisor, and Plotosichthyoidei for Ploto-
sus). As such, we conclude that Si/urus and Bagrus were the intended types for the Silurichthyoidei
and Bagrichthyoidei, respectively.

FORM OF PRESENTATION

In the list below, we provide the family-level name as originally spelled, its author, date of
publication and, when specified, original rank. We list the names in alphabetical order, and at the end
of each account we provide the current familial allocation of its type genus. For a name with no type
genus, we inferred the current familial placement on the basis of included taxa, even though the name
is not available. For a name that is based on a generic name, we also include the type genus with
corresponding author and date of publication.

Information relating to the type genus was not always complete in the publication in which the
family-level name was first used. We copied all pertinent information provided in the publication,
whether it was associated directly with the proposed name or found elsewhere in the text. Additional
information about the type genus that was not found in the publication, but inferred by us, is enclosed
in brackets. In all cases, we agree with Eschmeyer and Bailey (in Eschmeyer 1998) on the source and
date of publication of the generic names. Therefore, literature citations that relate only to generic
names are not included here, but can be found in Eschmeyer (1998). Generic names preceded by a
dagger (f) indicate a name proposed for a fossil taxon.

Unless otherwise noted, we consider the names listed to be available. Appendix | shows the same
list arranged according to currently valid families. The spelling provided for currently-used family
names follows Steyskal (1980). We note that the spellings recommended in Steyskal (1980) are not
uniformly adopted (e.g., Nelson 1996; Eschmeyer 1998). However, as no alternative explanation of
the derivation of these names has been more recently published, we continue to use Steyskal’s
spellings.

FAMILY-LEVEL NAMES OF SILURIFORMES

Acanthicini Bleeker, 1862:2 (stirps). Type genus: Acanthicus Spix [Spix and Agassiz, 1829].
Loricariidae.

Acestridiinae Isbriicker and Nijssen, 1974:67, 68 (subfamily). Type genus: Acestridium Haseman,
1911. Loriicariidae.
Acestrini Bleeker, 1862:4 (stirps). Type genus: Acestra Kner, [1853]. Permanently invalid, following
Art. 39 of the Code. Acestra Kner is preoccupied by Dall, 1852, in Hemiptera. Loricariidae.
Acrochordonichthyini de Pinna, 1996:61 (tribe). Type genus: Acrochordonichthys Bleeker, 1858.
Akysidae.

Ageneiosi Bleeker, 1862:14 (phalanx). Type genus: Ageneiosus Lac [La Cépede, 1803]. Auchenip-
teridae.

Ailichthyoidei Bleeker, 1858:1x, 49, 248 (subfamily). Type genus: Ailia Gray, [1831]. Also proposed
in the same publication as phalanx Ailianini Bleeker, 1858:1x, 49. Schilbidae.

Akyses Gill, 1861b:52 (group). Type genus: Akysis Bleeker, [1860]. Akysidae.

4 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 1

Amblycepinae Day, 1873:cclxviii (subfamily). Type genus: Amblyceps Blyth, 1858. Amblycipitidae.

Amiurina Giinther, 1864:98. Type genus: Amiurus [Ameiurus Rafinesque, 1820]. The family-level name
must be corrected to Ameiurina, following Art. 35(d) of the Code. Ictaluridae.

Amphiliidae Regan, 1911:565 (family). Type genus: Amphilius [Ginther, 1864]. Also proposed in same
publication as subfamily Amphiliinae. Amphiliidae.

Anchariidae Glaw and Vences, 1994:380 (family). Type genus: Ancharius Steindachner, 1881. Glaw and
Vences cite de Pinna’s unpublished dissertation and a then-unpublished manuscript by Stiassny as
the source of this family name. Stiassny and Raminosoa (1994:140, footnote 4) also use the name,
but do not provide characters that purport to differentiate the family, as required in Art. 13 of the
Code. Thus, it appears that Glaw and Vences provide the first published use of the name Anchariidae
that satisfies the requirements of availability. Anchariidae.

Ancistri Kner, 1854:6 (Untergruppe). Type genus: Ancistrus Kner, 1854. Proposed as an alternate name
for Lictores. First used in Kner (1853:282) but not available from that publication because Ancistrus
was not yet published. Loricariidae.

Andinichthyidae Gayet, 1988:833 (family). Type genus: t Andinichthys Gayet, 1988. Andinichthyidae.

Anesipoma Latreille, 1825:125 (tribe). Not available. Not based on a generic name. Incertae sedis.

Anodontes Bleeker, 1858:49, 245 (phalanx). Listed with Hypophthalmus as sole included genus. Not
available, not based on a generic name. Pimelodidae.

Argeini Bleeker, 1862:15 (stirps). Type genus: Arges Val [ Valenciennes, in Cuvier and Valenciennes,
1840]. Astroblepidae.

Arii Bleeker, 1862:7, 25 (phalanx). Type genus: Arius Val [ Valenciennes, in Cuvier and Valenciennes,
1840]. Artidae.

Ariobagri Bleeker, 1862:10 (phalanx). Not available, not based on a generic name. Incertae sedis.

Ariodontes Bleeker, 1858:49, 57 (phalanx). Not available, not based on a generic name. Incertae sedis.

Aspidoradidi Hoedeman, 1952:4 (tribe). Type genus: Aspidoras Ihering, 1907. Callichthyidae.

Aspredinae Swainson, 1838:332, 354 (subfamily). Type genus: Aspredo Artedi [= Aspredo Swainson,
1838, not Aspredo Scopoli, 1777]. Also in Swainson, 1839:189, 308. Permanently invalid, as the
name is based on a junior homonym (ICZN 1985, art. 39). Aspredinidae.

Aspredinidae Adams et al., 1854:107 (family). Type genus: [Aspredo? Scopoli, 1777]. It is not clear
whether this name is based on Aspredo Scopoli or on Aspredo Swainson, 1838. The diagnosis of the
family indicates the presence of a “first pectoral ray generally excessively developed,” which is not
consistent with the description of Swainson. If this name is not considered available, the next name
that dates to Aspredo Scopoli is Aspredinoidei (Bleeker, 1858:327; also Aspredinini). Bleeker’s
Aspredo comes from usage of Valenciennes (in Cuvier and Valenciennes, 1840), which is based on
Aspredo of Gronovius (1763), on which Scopoli also based his Aspredo. Aspredinidae.

Astephinae Grande and Lundberg, 1988:146 (subfamily). Type genus: + Astephus Cope, 1873. Ictaluridae.

Astroblepiformes Bleeker, 1862:15 (subfamily). Type genus: Astroblepus Humb [Humboldt, 1805].
Astroblepidae.

Astrophysi Bleeker, 1862:7 (phalanx). Type genus: Asterophysus Kner, [1858]. The family-level name
must be corrected to Asterophysi, following Art. 35(d) of the Code. Auchenipteridae.

Auchenipterini Bleeker, 1862:14 (stirps). Type genus: Auchenipterus Val [Valenciennes, in Cuvier and
Valenciennes, 1840]. Eigenmann and Eigenmann (1890) synonymized Euanemus Miller and Tro-
schel, 1842, into Auchenipterus and adopted the name Auchenipterinae over the older Euanemini
Bleeker, 1858. Since then, Euanemini has not been used. Article 40(b) of the Code allows for the
continued usage of a generally-accepted, junior family-level synonym over an unused older name and
provides an example that closely resembles the situation here. By this provision of the Code, the name
Auchenipterini is to be considered a senior synonym of Euanemini. Auchenipteridae.

Auchenoglanidinae Jayaram, 1966:1119 (subfamily). Type genus: Auchenoglanis Giinther, 1864. Claro-
teidae.

FERRARIS AND DE PINNA: CATFISH FAMILIES 5

Austroglanididae Mo, 1991:160 (family). Type genus: Austroglanis Skelton, Risch and Vos, 1984.
Austroglanididae.

Bagarina Giinther, 1864:3, 9, 183 (group). Type genus: Bagarius Bleeker, [1853]. Sisoridae.

Bagreidae Schultz, 1944:182 (family). Type genus: Bagre Oken, 1817. Ariidae.

Bagrichthyes Bleeker, 1862:8, 48 (phalanx). Type genus: Bagrichthys Bleeker, 1858. Bagridae.

Bagrichthyoidei Bleeker, 1858:49, 54 (subfamily). Type genus: Bagrus Blkr. [= Bosc, 1816]. Also
proposed as cohors Bagrini in the same publication (p. 49, 64). Based on the style of Bleeker’s
subfamily name generation, we consider this name available with Bagrus, and not Bagrichthys
Bleeker, as type. Bagridae.

Bagroidinae Jayaram, 1968:371 (subfamily). Type genus: Bagroides Bleeker, 1851. Bagridae.

Batasinae Tilak, 1967:431 (subfamily). Type genus: Batasio Blyth, 1860. Bagridae.

Batrachocephalinae Gill, 1893:132 (subfamily). Type genus: [Batrachocephalus] Bleeker, [1846].
Ariidae.

Bunocephalini Bleeker, 1858:xi, 328 (phalanx). Type genus: Bunocephalus Kner, [1855]. Aspredini-
dae.

Callichthini Bonaparte, 1838:131 (subfamily?). Type genus: [Callichthys Scopoli, 1777]. Callichthy-
idae.

Callophysinae Eigenmann, 1890:12 (subfamily). Type genus: [Callophysus, = Calophysus Miller
and Troschel, 1842]. Generic and subfamily names spelled consistently throughout Eigenmann,
1890 (also in Eigenmann and Eigenmann 1890:6, 7, 94) following changed spelling in Miller
and Troschel (1849), which is a misspelling (and not an unjustified emendation). Because of this,
the spelling Callophysinae is a based on a misspelled generic name and must be corrected to
Calophysinae. Pimelodidae.

Cascaduridi Hoedeman, 1952:3 (tribe). Type genus: Cascadura Ellis, 1913. Callichthyidae.

Centrochirinae Fowler, 1951:3 (subfamily). Type genus: Centrochir Agassiz, 1829. This name was
available when published, but it does not now meet the criterion for availability specified in Art.
13 of the Code and is therefore unavailable. Doradidae.

Centromochli Bleeker, 1862:7 (phalanx). Type genus: Centromochlus Kner, [1858]. Auchenipteri-
dae.

Cetopsini Bleeker, 1858:49, 250, 257 (phalanx). Type genus: Cetopsis Ag [Agassiz, 1829]. Cetopsi-
dae.

Chacini Bleeker, 1858:49, 310, 322 (phalanx). Type genus: Chaca CV [= Gray, 1831]. Chacidae.

Chaetostomidi Fowler, 1958:14 (tribe). Type genus: Chaetostomus Heckel, [1846]. Loricariidae.

Chiloglanidinae Riehl and Baensch, 1990:396 (Unterfamilie). Type genus: [Chiloglanis Peters,
1868]. Mochokidae.

Chrysichthyinae Regan, 1911:561. Type genus: Chrysichthys [Bleeker, 1863]. Also mistakenly
proposed as new in Jayaram (1966). Claroteidae.

Clariini Bonaparte, 1846:5 (subfamily?). Type genus: [Clarias Scopoli, 1777]. Clariidae.

Claroteini Bleeker, 1862:4 (stirps). Type genus: Clarotes Kner, [1855]. Claroteidae.

Clypeati Kner, 1853:282. Not available, not based on a generic name. Proposed as a subgroup of
Loricartidae for Sisor. Sisoridae.

Continae de Pinna, 1996:64 (subfamily). Type genus: Conta Hora, 1950. Erethistidae.

Copionodontinae de Pinna, 1992:179 (subfamily). Type genus: Copionodon de Pinna, 1992.
Trichomycteridae.

Corydoradinae Hoedeman, 1952:4 (subfamily). Type genus: Corydoras La Cépede, 1803. Also
proposed as a new tribe Corydoradidi in the same publication. Callichthyidae.

Cranoglanidae Myers, 1931:355 (family). Type genus: Cranoglanis Peters, 1880. Cranoglanididae.

Cyclopidae Eigenmann, 1910:416 (family). Type genus: Cyclopium Swainson, 1838. Astroblepidae.

Dianemidi Hoedeman, 1952:4 (tribe). Type genus: Dianema Cope, 1872. Callichthyidae.

6 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. |

Diplomystidae Eigenmann, 1890:14 (family). Type genus: Diplomystes [Bleeker, 1858]. Diplomystidae.

Diptéronotes Duméril, 1856:479 (family). Not available. Proposed for a broad array of catfish genera, but
not on an available generic name. Incertae sedis.

Doiichthyidae Weber 1913:532 (family). Type genus: Doiichthys, Weber 1913. Ariidae.

Doradini Bleeker, 1858:48, 52 (phalanx). Type genus: Doras CV [= La Ceépede, 1803]. Doradidae.

Doumeinae Regan, 1911:554. Type genus: [Doumea Sauvage, 1879]. Amphiliidae.

Eremophilini Bonaparte, 1846:5 (subfamily?). Type genus: [Eremophilus Humboldt, 1805}.
Trichomycteridae.

Erethistides Bleeker, 1862:13 (phalanx). Type genus: Frethistes Mill. Trosch. [Miller and Troschel,
1849]. Erethistidae.

Euanemini Bleeker, 1858:49, 189 (cohors). Type genus: Euanemus Miller and Troschel, [1842].
Auchenipteridae.

Exostomatina Giinther, 1864:264. Type genus: Exostoma Blyth, [1860]. Sisoridae.

Farlowellidi Fowler, 1958:14 (tribe). Type genus: Farlowella Eigenmann and Eigenmann, [1889].
Loricariidae.

Gephyroglanidini Jayaram, 1966:1099 (tribe). Type genus: Gephyroglanis Boulenger, 1899. Claroteidae.

Glanapteryginae Myers, 1944:592 (subfamily). Type genus: Glanapteryx Myers, 1927. Trichomycteridae.

Glani Latreille, 1825:125 (tribe). Not available. Proposed for a broad array of catfish genera, but not on a
generic name recognized as valid or even available at that time. Incertae sedis.

Glyptosterni Gill, 1861b:53 (group). Type genus: G/yptosternum McClelland, [1842]. Sisoridae.

Glyptothoracini de Pinna, 1996:64 (tribe). Type genus: G/yptothorax Blyth, 1860. Sisoridae.

Goniodontes Kner, 1853:279 (family). Not available, not based on a generic name. Used as an alternative
name for Loricata Kner, 1853. Loricartidae.

Harttiinae Boeseman, 1971:4, 10 (subfamily). Type genus: Harttia Steindachner, 1876. Loricariidae.

Helogenidae Regan, 1911:573 (family). Type genus: Helogenes [Ginther, 1863]. Cetopsidae.

Hemidoradinae Fowler, 1951:3 (subfamily). Type genus: Hemidoras Bleeker, 1858. Available when
published, but does not meet availability criterion as now specified in Art. 13 of the Code. Doradidae.

Hemiodontichthyina Isbriicker, 1979a:88, 89 (subtribe). Type genus: Hemiodontichthys Bleeker, 1862.
Loricariidae.

Hemipimelodinae Gill, 1861b:46 (subfamily). Type genus: [Hemipimelodus Bleeker, 1858]. This subfa-
milial name is mentioned briefly in the text, noting that only one genus 1s included. Ariidae.

Heptapterinae Gill, 1861b:54 (subfamily). Type genus: Heptapterus Bleeker, [1858]. Pimelodidae.

Heterobranchia Latreille, 1825:125 (tribe). Not available. Proposed for a group consisting of the genera
Plotose [sic] and Macropterotere [sic] and therefore not based on a genus recognized as valid in that
publication.

Heterobranchoidei Bleeker, 1858:33, 37, 41, 333 (family). Type genus: Heterobranchus Geoffr [Geoffroy
St. Hilaire, 1808]. The name Heterobranchini is also proposed in the same publication. Clariidae.

Heteropneustidae Hora, 1936:209 (family). Type genus: Heteropneustes Miller, 1840. Heteropneustidae.

Hopliancistrini Isbriicker and Nijssen, 1989:543 (tribe). Type genus: Hopliancistrus Isbriicker and
Nijssen, 1989. Loricariidae.

Hoplomizoninae Fernandez-Yépez, 1950:113 (subfamily). Type genus: Hoplomyzon [Myers, 1942].
Spelling of name incorrectly derived from type genus. Correct spelling is Hoplomyzontinae, as
currently recognized. Aspredinidae.

Hoplosterninae Miranda Ribeiro, 1959:1 (Grupo). Type genus: Hoplosternum Gill, 1858. Callichthyidae.

Hypophthalmini Bleeker, 1862:15 (stirps). Type genus: Hypophthalmus Spix [Spix and Agassiz, 1829].
Pimelodidae.

Hypoptopominae Eigenmann and Eigenmann, 1890:8, 12, 353, 388 (subfamily). Type genus: Hypop-
topoma Giinther, 1868. Spelling of name incorrectly derived from type genus. Correct spelling is
Hypoptopomatinae, as currently recognized. Loricariidae.

FERRARIS AND DE PINNA: CATFISH FAMILIES 7

Hypostomiden Kner, 1853:279. Type genus: Hypostomus La Cépede, 1803. Loricariidae.

Hypsidoridae Grande, 1987:28 (family). Type genus: + Hypsidoris Lundberg and Case, 1970. Also
as superfamily Hypsidoroidea in the same publication. Hypsidoridae.

Ichthyaelurinae Meek, 1904:10 (subfamily). Type genus: /chthyaelurus [= Ichthaelurus| Cope, 1868.
The family-group name must be corrected to Ichthyaelurinae, following Art. 35(d) of the Code.
Ictaluridae.

Ictaluri Gill, 1861b:49 (group). Type genus: /cfa/urus Raf [Rafinesque, 1820]. Ictaluridae.

Inermes Kner, 1853:282. Not available, not based on a generic name. Proposed as an alternate name
for Clypeati, a subgroup of Loricata, for Sisor Hamilton, 1822. Sisoridae.

Kryptopterini Bleeker, 1862:18, 85 (stirps). Type genus: Kryptopterus Bleeker, [1858]. Siluridae.

Laguviini de Pinna, 1996:65 (tribe). Type genus: Laguvia Hora, 1921. Erethistidae.

Lictores Kner, 1853:282, 1854:255. Not available, not based on a generic name. Proposed as an
alternate name for Ancistri. Loricartidae.

Lithodoradinae Fowler, 1951:3 (subfamily). Type genus: Lithodoras |Bleeker, 1862]. Available when
published, but does not meet availability criterion as now specified in Art. 13 of the Code.
Doradidae.

Lithogeneinae Gosline, 1947:80 (subfamily). Type genus: Lithogenes Eigenmann, 1909. Correct
spelling should be Lithogeninae, as currently adopted. Loricariidae.

Lithoxina Isbriicker, 1980:77 (subtribe). Type genus: Lithoxus Eigenmann, 1910. Loricariidae.

Loricaria Rafinesque, 1815:89 (subfamily). Type genus: Loricaria Linnaeus, 1758. Loricariidae.

Loricariichthyina Isbriicker, 1979a:87, 89 (subtribe). Type genus: Loricariichthys Bleeker, 1862.
Loricariidae.

Luciopimelodinae Driver, 1919:451 (subfamily). Type genus: Luciopimelodus Eigenmann and
Eigenmann, 1888. Pimelodidae.

Malapterurini Bleeker, 1858:1x (subfamily). Type genus: Malapterurus Lac [La Cepede, 1803].
Malapteruridae.

Metaloricariina Isbriicker, 1979a:88, 90 (subtribe). Type genus: Metaloricaria, Isbriicker 1975.
Loricariidae.

Mochokidae Jordan, 1923:150 (family). Type genus: Mochokus Joannis, 1835. Mochokidae.

Mystidae Fowler, 1935:275. Type genus [Mystus Scopoli, 1777]. This name was available when
published, but it does not meet the criterion for availability as now specified in Art. 13 of the
Code and is therefore unavailable. Mystini Hoedeman, 1949, is similarly unavailable. Bagridae.

Nangrina de Pinna, 1996:62 (subtribe). Type genus: Nangra Day, 1876. Sisoridae.

Nematogenyini Bleeker, 1862:16 (stirps). Type genus: Nematogenys Girard, 1854. Nematogenyidae.

Nematognathi Gill, 1861a:11, 56 (order). Not available as a family-group name, not based on a
generic name. Proposed as an alternate name for Siluriformes.

Neoplecostominae Regan, 1904:306 (subfamily). Type genus: Neoplecostomus Eigenmann and
Eigenmann, 1888. Loricartidae.

Olyrinae Gill, 1893:132 (subfamily). Type genus: Olyra, McClelland 1842. Bagridae.

Oplophoria Rafinesque, 1815:88 (family). Not available as a family-group name, not based on a
generic name. Apparently the first collective name for fishes now called Siluriformes.

Osteogeniosinae Fowler, 1951:3 (subfamily). Type genus: Osteogeneiosus Bleeker, 1846. Available
when published, but does not meet availability criterion as now specified in Art. 13 of the Code.
Ariidae.

Otocinclini Isbriicker, 1979b:114 (tribe). Type genus: Otocinclus Cope, 1871. Loricariidae.

Otothyrini Schaefer, 1991:31 (tribe). Type genus: Otothyris Myers, 1927. Loricartidae.

Pangasianodonidi Fowler, 1958:14 (tribe). Type genus: Pangasianodon Chevey, [1930]. Pangasiidae.

Pangasini Bleeker, 1858:49, 63 (cohors). Type genus: Pangasius CV [Valenciennes, in Cuvier and
Valenciennes, 1840]. Pangasiidae.

8 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. |

Parakysidae Roberts, 1989:141 (family). Type genus: Parakysis Herre, 1840. Akysidae.

Pareiodontinae Eigenmann, 1918:261 (subfamily). Type genus: Pareiodon Kner, 1855. Trichomycteridae.

Pelteobagrini Jayaram, 1966:1071 (tribe). Type genus: Pe/teobagrus Bleeker, 1865. Bagridae.

Phalacronotini Bleeker, 1862:18, 90 (stirps). Type genus: Phalacronotus Bleeker, [1858]. Siluridae.

Phreatobinae Reichel, 1927:383 (subfamily). Type genus: Phreatobius Goeldi, 1905. Pimelodidae.

Pimelodinae Swainson, 1838:331, 338 (subfamily). Type genus: Pimelodus [La Cépede, 1803]. Also as
Pimelodi in the same publication. Priority over Pimelodini Bonaparte, 1838, not established. Pi-
melodidae.

Pimelodini Bonaparte, 1838:131 (subfamily?). Type genus: [Pimelodus La Cépede, 1803]. See Pimelodi-
nae Swainson. Pimelodidae.

Pinirampidae Fernandez-Y épez, 1965:12 (family). Type genus: [Pinirampus Bleeker, 1858]. Pimelodidae.

Planiloricariina Isbriicker, 1979a:87, 89 (subtribe). Type genus: Planiloricaria Isbriicker, 1971. Loricari-
idae. .

Platystacinae Eigenmann and Eigenmann, 1890:9,13,20 (subfamily). Type genus: Platvstacus Bloch,
1794. Aspredinidae.

Plecostomiformes Bleeker, 1862:2 (subfamily). Type genus: Plecostomus Gron [Gronovius, 1763]. Also
as stirps Plecostomini in the same publication. Not available, based on a name in a publication that
was placed on the Official Index of Rejected and Invalid Works in Zoology, Opinion 261 (in Melville
and Smith, 1987). Loricartidae.

Plotosichthyoidei Bleeker, 1858:49, 310 (subfamily). Type genus: Plotosus Lacép [La Ceépede, 1803].
Also phalanx Plotosini in the same publication. We consider this name to be available, with Plotosus
as the type, based on the style of Bleeker’s publication and the definite reference to a single genus.
Plotosidae.

Porcinae Fowler, 1915:219 (subfamily). Type genus: Porcus Geoffroy St. Hilaire, 1808. Not available,
based on a generic name that was placed on the Official Index of Rejected and Invalid Generic Names
In Zoology, Opinion 1402 (in Melville and Smith, 1987). Bagridae.

Pseudacanthicini Isbriicker, 1980:76 (tribe). Type genus: Pseudacanthicus Bleeker, 1862. Also as subtribe
Pseudacanthicina in the same publication. Loricartidae.

Pseudauchenipterini Bleeker, 1862:6 (stirps). Type genus: Pseudauchenipterus Bleeker, [1862]. Also as
phalanx Pseudauchenipteri in the same publication. Auchenipteridae.

Pseudecheneidina de Pinna, 1996:64 (subtribe). Type genus: Pseudecheneis Blyth, 1860. Sisoridae.

Pseudopimelodidae Fernandez-Yépez and Martin, 1953:234 (family). Type genus: Pseudopimelodus
Bleeker, 1858. Available when published, but does not meet availability criterion as now specified
in Art. 13 of the Code. Mistakenly considered as earliest available proposition of the name by
Silfvergrip (1992:305). This name was also used in subsequent papers by A. Fernandez-Yepez, but
without associated diagnostic characters. Pimelodidae.

Pseudopimelodinae Lundberg et al., 1991:204 (subfamily). Type genus: Pseudopimelodus Bleeker, 1858.
Pimelodidae.

Pteronotidae Adams et al., 1854:107 (family). Type genus: [Pteronotus Swainson, 1839]. Permanently
invalid, as Pteronotus Swainson is a junior homonym of Pteronotus Gray, 1837 (Mammalia)
(McKenna and Bell, 1997). Pimelodidae.

Pygidiidae Eigenmann and Eigenmann, 1888:649 (family). Type genus: Pygidium Meyen, 1835. There is
apparently a discrepancy between Meyen’s (1835) original and Eigenmann and Eigenmann’s sub-
sequent notions of Pygidium. Refer to Tchernavin (1944) for a detailed account of the problem. The
name Pygidiidae in virtually all the literature refers to the group known today as Trichomycteridae.
Trichomycteridae.

Reganellina Isbriicker, 1979a:87, 89 (subtribe). Type genus: Reganella Eigenmann, 1905. Loricariidae.

Rhamdiae Bleeker, 1862:11, 60 (phalanx). Type genus: Rhamdia Bleeker, 1858. Pimelodidae.

Rhinoglanina Giinther, 1864:4, 10, 216 (group). Type genus: Rhinoglanis Giinther, 1864. Mochokidae.

FERRARIS AND DE PINNA: CATFISH FAMILIES 9

Ricolina Isbriicker, 1981:54 (subtribe). Type genus: Ricola Isbriicker and Nijssen, 1978. Loricartidae.

Rineloricariina Isbriicker, 1979a:87, 89 (subtribe). Type genus: Rineloricaria Bleeker, 1862. Lori-
cariidae.

Ritae Bleeker, 1862:8 (phalanx). Type genus: Rita Blkr. [Bleeker, 1858]. Bagridae.

Saccobranchini Bleeker, 1858:xi, 336 (phalanx). Type genus: Saccobranchus [Valenciennes, in
Cuvier and Valenciennes, 1840]. Heteropneustidae.

Sarcoglanidinae Myers and Weitzman, 1966:278 (subfamily). Type genus: Sarcoglanis Myers and
Weitzman, 1966. Trichomycteridae.

Schilbeini Bleeker, 1858:49, 250, 256 (phalanx). Type genus: Schilbe Valenc. ex parte [= Oken,
1817]. Schilbidae.

Scleracanthi Miranda-Ribeiro, 1917:49 Not available as a family-group name, not based on a generic
name. Proposed as an alternate name for Siluriformes or Nematognathi.

Scoloplacinae Bailey and Baskin, 1976:5 (subfamily). Type genus: Scoloplax Bailey and Baskin,
1976. Scoloplacidae.

Siluranodontinae Regan, 1911:568. Type genus: Si/uranodon [Bleeker 1858]. Schilbidae.

Silurichthyoidei Bleeker, 1858:49, 249 (subfamily). Type genus: Silurus L CV [= Linnaeus, 1758].
Both Silurus and Silurichthys were included in Bleeker’s subfamily Silurichthyoidei, leaving
some doubt regarding the type genus. We believe the name to be based on Silurus, because of
Bleeker’s convention of generating subfamily names by adding -ichthyoidei to the generic root.
Therefore, we do not consider this name to be available as a family-group name based on
Silurichthys. Siluridae.

Siluridae Anomalopterae Giinther, 1864:3, 7, 66 (subfamily). Not available, not based on a generic
name.

Siluridae Branchicolae Giinther, 1864:4, 12, 276 (subfamily). Not available, not based on a generic
name.

Siluridae Heteropterae Giinther, 1864:2, 5,30 (subfamily). Not available, not based ona generic name.

Siluridae Homalopterae Giinther, 1864:2, 5, 13 (subfamily). Not available, not based on a generic
name.

Siluridae Opisthopterae Giinther, 1864:4, 12,271 (subfamily). Not available, not based on a generic
name.

Siluridae Proteropodes Giinther, 1864:4, 11, 222 (subfamily). Not available, not based on a generic
name.

Siluridae Proteropterae Ginther, 1864:3, 7, 69 (subfamily). Not available, not based on a generic
name.

Siluridae Stenobranchiae Giinther, 1864:3, 10, 191 (subfamily). Not available, not based on a generic
name.

Siluroides Cuvier, 1816:199. Type genus: [Si/urus Linnaeus, 1758]. Available as a family-level name
for Siluridae. Originally used for the group generally known as Siluriformes. Siluridae.

Simuldentinae Taverne and Aloulou-Triki, 1974:64 (subfamily). Not available, not based ona generic
name. Mochokidae.

Sisorichthyoidei Bleeker, 1858:48, 50 (subfamily). Type genus: Sisor Buch. [Hamilton, 1822].
Sisoridae.

Sorubinae Swainson, 1838:356 (also Swainson, 1839:190, 309) (subfamily). Type genus: Sorubium
(= Sorubim) Spix [Spix and Agassiz, 1829]. Sorubium Swainson, 1838, is an emendation of
Sorubim, which Swainson (1838:356) concluded was “no doubt a misprint.” As Swainson’s
Sorubinae was based on an available generic name which he treated as valid, it is an available
name, but with Sorubium as its type. Pimelodidae.

Spectracanthicina Isbriicker and Nijssen, 1989:544 (subtribe). Type genus: Spectracanthicus Nijssen
and Isbriicker, 1987. Loricariidae.

10 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. |

Stegophilina Giinther, 1864:5, 12, 276 (group). Type genus: Stegophilus Reinhardt, 1858. Trichomycteri-
dae.

Synodontini Bleeker, 1862:6 (stirps). Type genus: Synodontis Cuv [Cuvier, 1816]. Mochokidae.

Tachysurinae Eigenmann, 1890:12 (subfamily). Type genus: Tachysurus La Cépéde, 1803. Ariidae.

Torpedininae Fowler, 1915:225 (subfamily). Type genus: Torpedo [Forsskal, 1775]. Permanently invalid,
as the name is based on a junior primary homonym of Torpedo Houttuyn, 1764.

Trachelyopterini Bleeker, 1858:49, 250, 257 (phalanx). Type genus: Trachelyopterini [ 7rachelyopterus]
CV [Valenciennes, in Cuvier and Valenciennes, 1840]. See also entry above on Auchenipteridae.
Auchenipteridae.

Trichomycterini Bleeker, 1858:49, 250, 257 (phalanx). Type genus: Trichomycterus CV [Valenciennes,
1833]. Trichomycteridae.

Tridentinae Eigenmann, 1918:275 (subfamily). Type genus: Tridens Eigenmann and Eigenmann, 1889.
Trichomycteridae. .

Trichogeninae Isbriicker, 1986:276 (subfamily). Type genus: Trichogenes Britski and Ortega, 1983.
Trichomycteridae.

Vandelliini Bleeker, 1862:17 (stirps). Type genus: Vandellia Valenciennes, 1846. Trichomycteridae.

Vorhisiidae Frizzell, 1965:179 (family). Type genus: + Vorhisia Frizzell, 1965. Ariidae.

ACKNOWLEDGMENTS

This paper benefitted from reviews of one or more versions of the manuscript by: Reeve Bailey,
Bill Eschmeyer, Susana Ferraris, Tomio Iwamoto, Maurice Kottelat, Alan Leviton, John Lundberg,
Richard Vari, and two anonymous reviewers. Isaac Isbriicker provided an exceptionally thorough
review of the manuscript and pointed out a number of names about which we were previously
unaware. We greatly appreciate the time each of these people took on our behalf.

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Wissenschaften in Wien 7:25 1—286 [1—36 of separate], pls. 1—5.

LATREILLE, P. A. 1825. Famillies naturelles du régne animal exposées succinctement et dans un ordre analytique,
avec |’indication de leurs genres. 2nd. ed. Paris. 570 pp.

LUNDBERG, J. G., A. H. BORNBUSCH, AND F. MAGOo-LecciA. 1991. Gladioglanis conquistador n. sp. from
Ecuador with diagnoses of the subfamilies Rhamdiinae Bleeker and Pseudopimelodinae n. subf. (Silurifor-
mes: Pimelodidae). Copeia 1991(1):190—209.

MCKENNA, M.C. AND S.K. BELL. 1997. Classification of mammals above the species level. Columbia

University Press, New York. 631 pp.

MEEK, S. E. 1904. The fresh-water fishes of Mexico north of the isthmus of Tehuantepec. Field Columbian

Museum, Zoological Series 5:itxii + 1-252, pls. 1-17.

MELVILLE, R. V. AND J. D. D. SMITH, eds. 1987. Official lists and indexes of names and works in zoology. The

International Trust for Zoological Nomenclature, London. 366 pp.

MEYEN, F. J. F. 1834-1835. Reise um die erde ausgefihrt auf dem K6niglich preussischen seehandlungs-schiffe
Prinzess Louise, commandirt von captain W. Wendt, in den jahren 1830, 1831 und 1832; Historischer
bericht. Sander’sche buchhandlung, Berlin. 2 vols.

MIRANDA RIBEIRO, A. DE. 1917. De Scleracanthis. Fluvio “Solimées” anno MCMVIII a cl. F. Machado da Silva
duce brasiliense inventis et in Museo Urbis “Rio de Janeiro” servatis. Revista de Sciencias 1:49—52.

FERRARIS AND DE PINNA: CATFISH FAMILIES 13

MIRANDA RIBEIRO, P. DE. 1959. Consideragdes sdbre Callichthyidae Gill, 1872 (Nematognathi). Boletim do
Museu Nacional, n. s. (206): 1—9.

Mo, T. 1991. Anatomy, relationships and systematics of the Bagridae (Teleostei, Siluroidei) with a hypothesis
of siluroid phylogeny. Koeltz Scientific Books, Koenigstein, Germany. vii + 216 pp. + 63 unnumbered
figures.

MULLER J. AND F. H. TROSCHEL. 1849. Horae ichthyologicae: Beschreibung und Abbildung neuer Fische,
Drittes Heft. Verlag von Veit and Co., Berlin. 28 pp., 5 pls.

Myers, G. S. 1931. On the fishes described by Koller from Hainan in 1926 and 1927. Lingnan Science Journal
10(2—3):255—262.

. 1944. Two extraordinary new blind nematognath fishes from the Rio Negro, representing a new
subfamily of Pygidiidae, with a rearrangement of the genera of the family, and illustrations of some
previously described genera and species from Venezuela and Brazil. Proceedings of the California Academy
of Sciences (Series 4) 23(40):591—602, pls. 52—56.

Myers, G. S. AND S. H. WEITZMAN. 1966. Two remarkable new trichomycterid catfishes from the Amazon basin
in Brazil and Colombia. Journal of Zoology (London) 149:277-287.

NELSON, J. S. 1994. Fishes of the World. 3rd ed. John Wiley and Sons, Inc., New York. xvii + 600 pp.

PINNA, M. C. C. DE. 1992. A new subfamily of Trichomycteridae (Teleostei, Siluriformes), lower loricarioid
relationships and a discussion of the impact of additional taxa for phylogenetic analysis. Zoological Journal
of the Linnean Society 106:175—229.

. 1996. A phylogenetic analysis of the Asian catfish families Sisoridae, Akysidae, and Amblycipitidae,
with a hypothesis on the position of the neotropical Aspredinidae (Teleostei, Ostariophysi). Fieldiana:
Zoology (n. s.) (84):1—83.

PINNA, M.C. C. DE AND R. P. VARI. 1995. Monophyly and phylogenetic diagnosis of the family Cetopsidae,
with synonymization of the Helogenidae (Teleostei: Siluriformes). Smithsonian Contributions to Zoology
(571): 1-26.

RAFINESQUE, C. S. 1815. Analyse de la nature, ou tableau de I’univers et des corps organises. Privately published,
Palerme. 224 pp.

REGAN, C. T. 1904. A monograph of the fishes of the family Loricariidae. Transactions of the Zoological Society
of London 17(part 3, number 1):191—350, pls. 9-21.

. 1911. The classification of the teleostean fishes of the order Ostariophysi. 2. Siluroidea. Annals and
Magazine of Natural History (8)8:553—577.

REICHEL, M. 1927. Etude anatomique du Phreatobius cisternarum Goeldi, silure aveugle du Bresil. Revue Suisse
de Zoologie 34(16):285—403, pls. 2-6.

RIEHL, R. AND H. A. BAENSCH. 1990. Mergus Aquarien Atlas, Band 3. Mergus Verlag, Melle, Germany. 1104

RoBERTS, T. R. 1989. The freshwater fishes of western Borneo (Kalimantan Barat, Indonesia). Memoirs of the
California Academy of Sciences (14):i—x1i + 1-210.

SCHAEFER, S. A. 1991. Phylogenetic analysis of the loricariid subfamily Hypoptopomatinae (Pisces: Siluroidei:
Loricariidae), with comments on the generic diagnoses and geographic distribution. Zoological Journal of
the Linnean Society 102(1):1-41.

SCHULTZ, L. P. 1944. The catfishes of Venezuela, with descriptions of thirty-eight new forms. Proceedings of
the United States National Museum 94(3172):173-338, pls. 1-14.

SILFVERGRIP, A. M. C. 1992. Zungaro, a senior synonym of Paulicea (Teleostei: Pimelodidae). Ichthyological
Explorations of Freshwaters 3(4):305—3 10.

STEYSKAL, G. C. 1980. The grammar of family-group names as exemplified by those of fishes. Proceedings of
the Biological Society of Washington 93:168—177.

STIASSNY, M. L. J. AND N. RAMINOSOA. 1994. The fishes of the inland waters of Madagascar. Annales du Musee
Royal de l’Afrique Centrale, Série in 8°, Sciences Zoologiques (275):133—149.

SWAINSON, W. 1838. The natural history and classification of fishes, amphibians, and reptiles, or monocardian
animals, volume 1. Longman, Orme, Brown, Green and Longmans, and John Taylor, London. vi + 368
pp.

. 1839. The natural history and classification of fishes, amphibians, and reptiles, or monocardian animals,
volume 2. Longman, Orme, Brown, Green and Longmans, and John Taylor, London. vi + 448 pp.

TAVERNE, L. AND A. ALOULOU-TRIKI. 1974. Etude anatomique, myologique et ostéologique du genre Synodontis

14 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. |

Cuvier (Pisces: Siluriformes, Mochocidae). Annales du Musée Royal de |’Afrique Centrale, série in 8°,
Sciences Zoologiques (210):1—69, pls. 1—2.

TCHERNAVIN, V. V. 1944. A revision of some Trichomycterinae based on material preserved in the British
Museum (Natural History). Proceedings of the Zoological Society (London) 114(1—2):234-275.

TILAK, R. 1967. Studies of the osteocranium and the Weberian apparatus of the fishes of the genus Batasio Blyth,
1860 (Pisces: Siluroidei) with remarks on the systematic position of the genus. Anatomische Anzeiger
121:415-434.

WeBER, M. 1913. Siisswasserfische aus niederlandisch Stid- und Nord-Neu-Guinea. Nova Guinea (Zool.)
9(4):513-613, pls. 12-14.

FERRARIS AND DE PINNA: CATFISH FAMILIES l

WN

APPENDIX |

Catfish family-group names arranged alphabetically within current familes. Arrangement of
families follows Nelson (1994), with modifications by Mo (1991), de Pinna and Vari (1995) and de
Pinna (1996). Available family-group names, except those that are ruled permanently invalid, are in
bold face. Namés originally based on incorrect spelling of generic root are corrected; variant spellings

not included.

Akysidae

Acrochordonichthyini de Pinna, 1996

Akyses Gill, 1861b

Parakysidae Roberts, 1989
Amblycipitidae

Amblycepinae Day, 1873
Amphiltidae

Amphiliidae Regan, 1911

Doumeinae Regan, 1911
Anchariidae

Anchariidae Glaw and Vences, 1994
Andinichthyidae

Andinichthyidae Gayet, 1988
Ariidae

Arii Bleeker, 1862

Bagreidae Schultz, 1944

Batrachocephalinae Gill, 1893

Doiichthyidae Weber, 1913

Hemipimelodinae Gill, 1861b

Osteogeniosinae Fowler, 1951

Tachysurinae Eigenmann, 1890

Vorhisiidae Frizzell, 1965
Aspredinidae

Aspredinae Swainson, 1838

Aspredinidae Adams et al., 1854

Bunocephalini Bleeker, 1858

Hoplomyzontinae Fernandez-Y épez, 1950

Platystacinae Eigenmann and Eigenmann,

1890

Astroblepidae

Argeini Bleeker, 1862

Astroblepiformes Bleeker, 1862

Cyclopidae Eigenmann, 1910
Auchenipteridae

Ageneiosi Bleeker, 1862

Asterophysi Bleeker, 1862

Auchenipterini Bleeker, 1862

Centromochli Bleeker, 1862

Euanemini Bleeker, 1858

Pseudauchenipterini Bleeker, 1862

Trachelyopterini Bleeker, 1858
Austroglanididae

Austroglanididae Mo, 1991
Bagridae

Bagrichthyes Bleeker, 1862

Bagrichthyoidei Bleeker, 1858

Bagroides Bleeker, 1851

Batasinae Tilak, 1967

Mystidae Fowler, 1935

Mystini Hoedeman, 1949

Olyrinae Gill, 1893

Pelteobagrini Jayaram, 1966

Porcinae Fowler, 1915

Ritae Bleeker, 1862
Callichthyidae

Aspidoradidi Hoedeman, 1952

Callichthini Bonaparte, 1838

Cascaduridi Hoedeman, 1952

Corydoradinae Hoedeman, 1952

Dianemidi Hoedeman, 1952

Hoplosterninae Miranda Ribeiro, 1959
Cetopsidae

Cetopsini Bleeker, 1858

Helogenidae Regan, 1911
Chacidae

Chacini Bleeker, 1858
Clariidae

Clariini Bonaparte, 1846

Heterobranchoidei Bleeker, 1858

Saccobranchini Bleeker, 1858
Claroteidae

Auchenoglanidinae Jayaram, 1966

Chrysichthyinae Regan, 1911

Claroteini Bleeker, 1862

Gephyroglanidini Jayaram, 1966
Cranoglanididae

Cranoglanidae Myers, 1931
Diplomystidae

Diplomystidae Eigenmann, 1890
Doradidae

Centrochirinae Fowler, 1951

Doradini Bleeker, 1858

Hemidoradinae Fowler, 1951

Lithodoradinae Fowler, 1951
Erethistidae

Continae de Pinna, 1996

Erethistides Bleeker, 1862

Laguviini de Pinna, 1996
Heteropneustidae

Heteropneustidae Hora, 1936

Saccobranchini Bleeker, 1858
Hypsidoridae

16 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. |

Hypsidoridae Grande, 1987 Pimelodidae
Ictaluridae Anodontes Bleeker, 1858
Amieurina Giinther, 1864 Calophysinae Eigenmann, 1890
Astephinae Grande and Lundberg, 1988 Heptapterinae Gill, 1861b
Ichthaelurinae Meek, 1904 Hypophthalmini Bleeker, 1862
Ictaluri Gill, 1861b Luciopimelodinae Driver, 1919
Loricariidae Phreatobinae Reichel, 1927
Acanthicini Bleeker, 1862 Pimelodinae Swainson, 1838
Acestridiinae Isbriicker and Nijssen, 1974 Pimelodini Bonaparte, 1838
Acestrini Bleeker, 1862 Pinirampidae Fernandez-Yeépez, 1965
Ancistri Kner, 1853 Pseudopimelodidae Fernandez-Y épez and
Chaetostomidi Fowler, 1958 Martin, 1953
Farlowellidi Fowler, 1958 Pseudopimelodinae Lundberg et al., 1991
Goniodontes Kner, 1853 Pteronotidae Adams et al., 1854
Harttiinae Boeseman, 1971 Rhamdiae Bleeker, 1862
Hemiodontichthyina Isbriicker, 1979a Sorubinae Swainson, 1838
Hopliancistrini Isbriicker and Nijssen, 1989 —_ Plotosidae
Hypoptopomatinae Eigenmann and Plotosichthyoidei Bleeker, 1858
Eigenmann, 1890 Schilbidae
Hypostomiden Kner, 1853 Ailichthyoidei Bleeker, 1858
Lictores Kner, 1853 Schilbeini Bleeker, 1858
Lithogeninae Gosline, 1947 Siluranodontinae Regan, 1911
Lithoxina Isbriicker, 1980 Scoloplacidae
Loricaria Rafinesque, 1815 Scoloplacinae Bailey and Baskin, 1976
Loricariichthyina Isbriicker, 1979a Siluridae
Metaloricariina Isbriicker, 1979a Kryptopterini Bleeker, 1862
Neoplecostominae Regan, 1904 Phalacronotini Bleeker, 1862
Otocinclini Isbriicker, 1979b Silurichthyoidei Bleeker, 1858
Otothyrini Schaefer, 1991 Siluroides Cuvier, 1816
Planiloricariina Isbriicker, 1979a Sisoridae
Plecostomiformes Bleeker, 1862 Bagarina Gunther, 1864
Pseudacanthicini Isbriicker, 1980 Clypeati Kner, 1853
Pseudoloricariina Isbricker, 1981 Exostomatina Ginther, 1864
Reganellina Isbriicker, 1979a Glyptothoracini de Pinna, 1996
Ricolina Isbriicker, 1981 Glyptosterni Gill, 1861b
Rineloricariina [sbriicker, 1979a Inermes Kner, 1853
Spectracanthicina Isbriicker and Nijssen, Nangrina de Pinna, 1996
1989 Pseudecheneidina de Pinna, 1996
Malapteruridae Sisorichthyoidei Bleeker, 1858
Malapterurini Bleeker, 1858 Trichomycteridae
Mochokidae Copionodontinae de Pinna, 1992
Chiloglanidinae Riehl and Baensch, 1990 Eremophilini Bonaparte, 1846
Mochokidae Jordan, 1923 Glanapteryginae Myers, 1944
Rhinoglanina Ginther, 1864 Pareiodontinae Eigenmann, 1918
Simuldentinae Taverne and Aloulou-Triki, Pygidiidae Eigenmann and Eigenmann, 1888
1974 Sarcoglanidinae Myers and Weitzman, 1966
Synodontini Bleeker, 1862 Stegophilina Ginther, 1864
Nematogenyidae Trichogeninae Isbriicker, 1986
Nematogenyini Bleeker, 1862 Trichomycterini Bleeker, 1858
Pangasiidae Tridentinae Eigenmann, 1918
Pangasianodonidi Fowler, 1958 Vandelliini Bleeker, 1862

Pangasini Bleeker, 1858

FERRARIS AND DE PINNA: CATFISH FAMILIES

UNASSIGNED FAMILY-LEVEL NAMES (INCLUDING SUPRAFAMILIAL NAMES)

Anesipoma Latreille, 1825
Ariobagri Bleeker, 1862
Ariodontes Bleeker, 1858
Dipteronotes Dumeril, 1856

Glani Latreille, 1825
Heterobranchia Latreille, 1825
Nematognathi Gill, 186la
Oplophoria Rafinesque, 1815
Scleracanthi Miranda-Ribeiro, 1917

Siluridae Anomalopterae Ginther, 1864
Siluridae Branchicolae Giinther, 1864
Siluridae Heteropterae Giinther, 1864
Siluridae Homalopterae Giinther, 1864
Siluridae Opisthopterae Giinther, 1864
Siluridae Proteropodes Ginther, 1864
Siluridae Proteropterae Gunther, 1864
Siluridae Stenobranchiae Giinther, 1864

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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES

Volume 51, No. 2, pp. 19-103, | fig., 14 plates. July 20, 1999

Index Pennatulacea
Annotated Bibliography and Indexes of the Sea Pens
(Coelenterata: Octocorallia) of the World 1469-1999

by

Gary C. Williams
Department of Invertebrate Zoology, California Academy of Sciences
Golden Gate Park, San Francisco, California 94118

A reasonably comprehensive bibliography of the living and fossil pennatulacean Octo-
corallia is presented, with the goal of including all published accounts regarding the biology
of the sea pens. This compilation of approximately 1000 citations represents 530 years of
published research. Complete unabbreviated citations for periodicals are used throughout.
Many of the citations are annotated with descriptive notes. Taxonomic, geographic, and
field-of-study indexes to the literature are included, as well as a synopsis of historical
periods in the study of the Pennatulacea.

TABLE OF CONTENTS

IRE C PULOTV CII Ss tae Ne Re hike ae Bisco Mglet Rath ch Rom el 2 2 ale es cent Sea eT AV Ad Groh Sent sae 21
PRC KUROSNAC GOIN) ENES:<. i... eye iren ae Het tee ese Pc ee) sera nie Gua ea eee 2D
GTO Cdl NCCOUNL 2 22's: eremrenen aint tua ern ea ciate OMe = Sd ocean cates Gy oes See 3)
Bre -einieanmee nods LA GSN 77 Sih) weenie casyato cuss tomes face eed ic hos ten stra eceses Oreo e aE cher c aaa + pe)
Mine CiineanibenOGWl75S— S58) pico. 4. eas ee eu ee eo eile ehatecers cui = Gea esha eas 25
Mihe- Darwinian ReclodiG@ls 5 9—1 S899) Ps er oy. cas cs chats eel cs ecernie ave) os Sub) age Seteme eters 6). 0 25
the Bartyaliwentiethn@ century (900 -V9AO) on aia ea 5 ye om eraertes weet one er 6 ook Os Slecieaas ayer DS
fiheiate twentieth Centunys(T9SO= V9 9 ye ai we eats ich am, cop ceo7s Pe, Stereos eR Usee cesuaue eau 29
Syuonvins and misspelled genene names ofPennatulacea...-. 2... 42.24. sge ceo an es 29
BIRDIES HUT Xe aaah csc yeh ust Gh ayo ic Os deklre ace e aera o. Ch Sie wee ESN el ROR IG cheep 33
Goo reA FOUN oA ONG 255 gana CR Ae Cork aoe) = Pegs nie GAS Nes est eh RMP eee 47
ESTES (SRS BOTTING 01 WER ags PORE cles MN Saari Sanna rs Un RAR Renee rn bees SURI ey =e ane eee 47
PRAT aON Ie SUOMI A ts esti eatery a esr ete au ara a sas at oh Arca eye tA AES Men cu pc eter APR, 22 acai cay 47
JNIRGUG (QO SEIT Ay Renee REA ea Oe ny One Nice re ne eee SS eer ea 47
FENES TAME AAS COMMEND SIGIN 5 oss corey seme ye L OUR ISG) SSeS dele RES, 5 RAE ce ee eae da ae 47
/ANTISIT Nh = SSIS ee a et ere Peer AMPaMU PIS Tests. oc 47
nido-paciie (ineludingathe Red Sea) i o.0c > 2 ny-8 oicyene Bhasin ks Sarai tis Saye Seon 47
AUR ATM Mr N se rays, Sige cal a RUEIY lap bice sakes Ra) les eos Gale ato SkeeamesE Tok ahha a AD Re 48
ING Weed lan Cire eey en wees Rs ecto, Ceti fh wh aheloia), ates SOG oe ear cta oes se eee ee 48
Neorg Nam Ti aA ial coca dvarantnvationsh ao vo$, kana eee oles ao a me 48
North Atlantic and Mediterranean (Europe and eastern North America) ..............-.. 48
Facil Gx@ Ce ai mOntine nine gexec « (acct aca tis de sas on, ass) Sa aniay Moma ate ek a ER ee 48
SOUP AMICI Ca ICAU AIMEE iy sho acts sitet an arsuioesa dy aucdsois09: Sie cscaouaapeerre eos Nine ea RO wea When wees 48
SOUEMBANII CHIC AN AC UTC) MeOe ets et 5 ox iebets eRe cat Sn: 4G Lats <5 Gale Ro eas ee RC Rm ee ae 48

20

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 2

PLATE 1

Umbellula antarctica
After Kiikenthal & Broch, 1911

WILLIAMS: INDEX PENNATULACEA 2]

Souther!’ Oceans (Antarctica andthe subantarctic)™.. a2: er. STR: 49
Westen inopical-Atlantio-and Caribbean 28 2etes OP U) SPS hte Se ee 49
Non) divide arate ee ee Ae apenas 2 ees tts HOP MISHIP NY Died OI eet me aT RE 49
SU DCU iMG eNom rena fot es <i 2 shod carat yee ho chen we ld edagiens ivia Mellons ot arate A 49
IBS EISATOTS pte gsc bmn dc he Aa A at acest draco eae ad a ir ae ee RS SE 49
|B [SVINGYOR C2) OLIN, Sas cane a Ai ae oases CUPRA aie oe en 2a ae a Sa Are en eR Ear 49
Biography (Regarding Authors of Pennatulacean Research) ...........4..0052.-02205- 49
Bro lmiminlescencey me) sina A pu ae eee eer Pont Paha ek oh PI UAE NR 49
Goraisdbre-leimnean General Natural Elistory)) 0 0h i. Pele ee EO 50
Ecole rvsandebione: Interactions 2°. ee ae ee ee ae Oe a IE ah ee 50
GeneralaNaturalelistory and General Biology? J..ih.-4 ee ee 2 ce ea Pe at
Bisse oy eerie omens ye oes Vl tes 1 8 RO MA a ord ml alae 1d cee As ete 5]
History of science (Regarding Coelenterates and/or Sea Pens) ..0:...25. 02200. 22248.: 5S]
Molecuiarmiolocyrand Genetics 2 £25) he gee LP oe er ed a 5]
Mermnolopy-andiuitrastrac rune s!07% 6 U1! Lael, Teed a) 20 8 Sal, SL Se a
Natural!Products Chemistry, Biochemistry, and Toxicology 2s... -...2..2.02)55022.% 5S]
Paleontology (Mesozoic and Cenozoic Fossil Taxa) 25.595). ee ee ae 52
Ralconolosy(V endian Frondlike Fossil Taxa) 220. Se eee. oo eek eee se lesen ee: 52
Photesraphy«(Color Photographs or LivingSea Pens)! . 2.0 25 e eh ee ee ee eas. 52
Bhyloceny. Cladistics, Evolution; and Biogeography 2 202.0... 0020850 oe ee ae 52
Rorsiolopysancd Cell BiOlo sy: c2.0) ore eke eee ee Bee NON ee oe 52
Folymorphism, Phenotypic Variability; and Biodiversity’). .2.).. 0202.22 hen. Jose e. 53
Wietcdhbassapese tains: ase nk 2 CA. IC. it ed) dan ne Cee 53
Reproductive and Developmental Biology “2. oan. s a. fos ae oe ese see aeieee 58
Taxonomy, Distributional and Bathymetric Records os. .2.7 5 Os ee 53
Ea banirenral Mayer rien Hemet rere ey 2 ae ve nareee EO AIS A tet ke I 54
INTRODUCTION

The pennatulaceans, commonly known as sea pens and sea pansies, are a highly specialized and
distinct group of sessile benthic coelenterates. They are distributed throughout the world’s oceans
from the polar seas to the tropics, and at all depths from the intertidal (certain virgulariids) to over
6200 meters (Umbellulidae, Pl. 1). Morphologically, sea pens are highly diverse and exhibit morpho-
logical changes as evolutionary events within different lineages. Examples are bilateral symmetry,
concentration and localization of feeding polyps, the development of lateral processes as polyp leaves
or ridges, and the reduction in the number and size of sclerites. As octocorallian coelenterates,
pennatulaceans are characterized by having eight pinnate tentacles surrounding the mouth of each
polyp, and eight mesenteries. Unlike other octocorals, however, mature colonies develop from a single
large polyp (the oozooid) that produces secondary zooids by lateral budding of its body wall. Also
unique to the Pennatulacea, is the unbranched muscular peduncle, which anchors the animal by
peristaltic contractions into soft substrata such as sand, mud, or abyssal ooze. The secondary polyps
(larger autozooids for feeding and reproduction, and smaller siphonozooids for internal water
circulation) are restricted to the distal rachis, which is also unbranched but may have laterally-pro-
duced polyp leaves.

The ability to inhabit soft substrata has allowed several abyssal-dwelling sea pens to have nearly
cosmopolitan distributions. Despite these very widespread geographic ranges, pennatulacean species
diversity in the deep-sea is relatively low, and may be attributable to a combination of factors
including: relatively low energy input and depauperate productivity in the abyssal environment,
coupled with a relative lack of ecological diversity.

22 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 2

Thirty-two genera in fifteen families of living pennatulaceans are currently recognized. Of the
436 nominal species names described in the literature, approximately one half are currently considered
valid. Major monographic works on the Pennatulacea include Kiikenthal (1915), Hickson (1916), and
Williams (1990, 1995a).

The sparsely-represented pennatulacean fossil record extends back to the Cretaceous Period and
consists primarily of virgulariid fossil taxa in the genera Graphularia and Virgularia. Bayer
(1956:228) observed that fossils of Graphularia closely resemble the axial structure of the genus
Stylatula, and the two may be congeneric. Problematic and controversial frondlike taxa that resemble
pennatulid sea pens are described from the Vendian Period of the late Precambrian. In fact, the coming
of age of Precambrian paleobiology in the 1980s has produced a wealth of publications dealing with
the Ediacaran or Vendian biota. Many of the Vendian frondlike taxa have been allied with the
Pennatulacea by various authors, but these interpretations remain highly disputatious. Because of the
importance of recent fossil discoveries, which have helped to bridge the gap between fossil biotas of
the Precambrian and the Cambrian, many of the references on these Vendian pennatulacean-like
organisms are included here, but the coverage should in no way be considered comprehensive.

The goal of this work is to produce a bibliography of all published works pertaining in part or in
full to the pennatulacean octocoral coelenterates. Although no work of this kind can claim to be
absolutely comprehensive, this bibliography can at least be considered reasonably complete, certainly
more so than any previously-published, similar work. The fine bibliography of octocorals by Bayer
(1996) concentrates on taxonomic works for all of the Octocorallia, but does not claim to be inclusive
for Pennatulacea or fields of study outside of taxonomy.

The following annotated and alphabetical bibliography to the Pennatulacea of the world is derived
from a variety of sources—primarily the Zoological Record (1864-1998), as well as the author’s
research library. Other sources include Bayer (1981), Bayer (1996), and Hickson (1916). My
annotations are in square brackets at the end of many of the citations.

This work is intended for a variety of users including professional biologists, biology students,
fisheries biologists and marine resource managers. ;

The intent of the presentation is to make this work as user friendly as possible. In order to facilitate
this goal, full citations are given for the vast majority of the references listed, as abbreviations for
periodical publications have proved in the past to be confusing, inconsistent, or often unusable.

The Melvyl System, library data base of the University of California, [http://www.melvyl.
ucop.edu] was used extensively to acquire complete citations for periodicals. Other sources used for
abbreviations and complete citations of periodicals include the List of Serial Publications in the
BM(NH) Library, the World List of Sciéntific Periodicals 1900-1960, and Biosis Serial Sources.

For the most recently-recognized, valid species names of Pennatulacea, see Williams (1995a,
1997):

ACKNOWLEDGMENTS

I am grateful to Larry Currie (Librarian, California Academy of Sciences, San Francisco), who
was instrumental in providing expertise and in locating several references.

I thank Leen van Ofwegen (Nationaal Natuurhistorisch Museum, Leiden), Phil Alderslade
(Museum and Art Gallery of the Northern Territory, Darwin), Frederick Bayer (National Museum of
Natural History, Washington, D. C.), and Michael Ghiselin (California Academy of Sciences) for
their valuable contributions and comments.

I especially thank Alan Leviton (California Academy of Sciences) for his interest and patience,
regarding many hours spent at the computer preparing the figures.

WILLIAMS: INDEX PENNATULACEA 23

HISTORICAL ACCOUNT

Pre-Linnean Period (1469-1757)

In writing about the history of research regarding pennatulacean bioluminescence, E. Newton
Harvey (1952:170), related the following, “Although the Romans knew of sea pens, referring to them
as ‘Penna marina,” the sea feather, or as ‘Mentula alata,’ the winged penis, the ability to luminesce
does not appear to have been recorded. Probably the first mention of luminous sea pens comes to us
in the famed ‘De lunariis’ and the ‘Historia Animalium’ of Conrad Gesner, published in the middle
sixteenth century.”

The earliest published accounts describing aspects of the natural history of octocorals, including
the sea pens, are Pliny the Elder (1469), Guillaume Rondelet (1554, 1555), Conrad Gesner (1555,
1558) (PI. 2), Ferrante Imperato (1599), and Captain Lancaster (1601). Other noteworthy pre-Linnean
works dealing at least in part with the Octocorallia are those of Francisci Erasmi (1668), Paolo
Boccone (1674), Hans Sloane (1707), Jacques Barrelier (1714), Herman Boerhaave (1720, 1727),
Christlob Mylius (1753, 1754), and John Ellis (1753, 1755).

Pennatulacean bioluminescence is at least mentioned by Francois Boussuet (1558), Caspar
Bauhin (1620, 1671), Ulisse Aldrovandi (1642, 1648), Johann Bauhin (1650—51), and Thomas Shaw
(1738-46).

Louis Agassiz (1860) praised the perspicacity of early workers such as Rondelet and Gesner,
who first established the animal nature of the “acalephs” (many of the taxa that we now regard as
coelenterates), departing from the earlier views of Aristotle and Pliny, who compared them to plants
(the “zodphytes”).

Pertaining to Rondelet, Agassiz (1860:8) stated, ‘*... we are chiefly indebted to Rondelet for
contributions to the natural history of the Acalephs. He was, indeed, not only better acquainted with
the inhabitants of the Mediterranean than all his predecessors, but he knew them even more accurately
than any naturalist that lived before the present century.”

In his praise of sixteenth century naturalists, Agassiz (1860:10) continues, “It has been a source
of constant delight for me, while perusing the works of the earlier naturalists, to sympathize with the
genial spirit and the earnestness that pervade their writings, so free from egotism, and animosity
against their fellow-students. Their devotion to their studies is equal to the spirit of reverence with
which they look upon nature; and it is disgraceful to our age, that we must contrast with such
dispositions the ill-will, the jealousies, the quarrels for priority, and the profanation, which pervade
the discussions of certain modern authors. Moreover, in a systematic point of view, the great
naturalists of the sixteenth century deserve to be studied more fully than they have been thus far. It
is astonishing, for instance, to see how near Rondelet, in discussing the views of Aristotle upon the
affinities of animals, came to perceiving their true affinities, and their natural classification... .”

A good example of the evolution of thought in natural history is exemplified by various views
regarding the nature of corals in the seventeenth and eighteenth centuries. The concept of corals as
“zoophytes,” transitional between plants and animals, was established by the Roman scholars Sextus
Empiricus and Pliny the Elder (Caius Plinius Secundus), based on Aristotle’s notion of the gradation
from plants to higher animals. This undoubtedly served to propagate a state of confusion for nearly
two thousand years—the false view that corals as living beings share the natures of both animals and
plants (Agassiz, 1860:6—7). Paolo Boccone (1670) and John Woodward (1695) emphasized the
inanimate aspect of corals and regarded them as stones, while L. F. Marsigli (1725) considered them
to be plants, and the coral polyps as “flowers of the coral.” It was the perceptive observer, Jean-André
Peyssonnel (1753), who first recognized the animal nature of corals and their affinities to other
coelenterates, after observing the contraction, expansion and movement of the tentacles of coral
polyps (Hyman 1940:365).

24

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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 2

a ee o €
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téehothecs feats.
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eas tote u Soph . 2 ¥ : aes 2 2
Conrad Gesner (1516-1565). Courtesy Carl von Linné (Linnaeus) (1707-1778). Courtesy
Kraig Adler, Cornell University. Picture Collection, California Academy of Sciences.

Peter Simon Pallas (1741-1811). Courtesy Lazzaro Spallanzani (1729-1799). Courtesy
Kraig Adler, Cornell University. Michael Ghiselin, California Academy of Sciences.

WILLIAMS: INDEX PENNATULACEA 25

The Linnean Period (1758—1858)

The publication of the 10th edition of Systema Naturae by Linnaeus (Carl von Linné 1758) (PI.
2) is accepted as the starting point for our contemporary system of binominal nomenclature. Linnaeus
named the genus Pennatula, from which the ordinal name Pennatulacea is derived. Other contributors
during the Linnean Period include William Borlase (1758), Job Baster (1759-65); Joannes Baptist
Bohadsch (1761); John Ellis (1764); Peter Simon Pallas (1766) (PI. 2), first monographer of the
“zoophytes”; Pieter Boddaert (1771); I. Lepechin (1781); John Ellis and Daniel Solander (1786);
Lazzaro Spallanzani (1796) (PI. 2); Georges Cuvier (PI. 3), originally Jean Leopold Nicolas Frédéric
Cuvier (1797), who named the genera Umbellula and Veretillum; Jean Baptiste Pierre Antoine de
Monet de Lamarck (1816) (PI. 3), who named the genera Funiculina, Renilla, and Virgularia; Tilesius
von Tilenau (1819); S. della Chiaje (1827, 1841-44); M. W. Rapp (1827); H.-M. Ducrotay de
Blainville (1834); Christian Gottfried Ehrenberg (1834a) (Pl. 3), who introduced the family name
Pennatulina, later corrected by Dana, 1846 to Pennatulidae; Fredrich Sigismund Leuckart (1841);
James Dwight Dana (1846) (PI. 3), who corrected the familial name of Ehrenberg to Pennatulidae;
George Johnston (1847); John Edward Gray (1840-1873), who named the genus Sarcoptilus in 1848;
Henri Milne Edwards (PI. 4) and Jules Haime (1850), who named the fossil genus Graphularia;
Achille Valenciennes (1850), who named the veretillid genera Lituaria and Cavernularia; P. Chr.
Asbjornsen (1856), who named the genus Kophobelemnon; and Jan Adrianus Herklots (1858) (PI.
4), who named the genera Scytalium and Pteroeides.

The Darwinian Period (1859-1899)

The publication of the Origin of Species by Charles Darwin (1859) initiated what is here referred
to as the Darwinian Period.

The pre-eminent author of the Pennatulacea during this period was Rudolf Albert von KOlliker,
who produced major descriptive taxonomic works between 1865 and 1880. Kélliker described the
sea pens of the H. M. S. Challenger Expedition, in which thirty species were described. KOlliker also
named the genera Acanthoptilum, Anthoptilum, Halipteris, Protoptilum, Sclerobelemnon, Scleropti-
lum, and Stachyptilum. Other noteworthy contributions during this period include Pieter Bleeker
(1859) (Pl. 4), who described several species of Pteroeides; William More Gabb (1859-1864) (PI.
4); Jan Adrianus Herklots (1863); Josua Lindahl (1874); Johan Koren and Daniel C. Danielssen
(1847-1884); Addison Emery Verrill (1865—1 882), who named the genera Distichoptilum, Ptilosar-
cus, and Stylatula, and introduced the name Pennatulacea in 1865 as a subordinal name, later elevated
to ordinal status by Studer, 1887a; Sir Charles Wyville Thomson (1874), chief scientist of the
H. M. S. Challenger Expedition (PI. 5); Robert Edward Carter Stearns (1873-1883) (Pl. 5); Am-
brosius Arnold Willem Hubrecht (1885) (Pl. 5), who named the genus Echinoptilum; G. Herbert
Fowler (1888, 1894); A. Milnes Marshall (1883a, 1883b); A. Milnes Marshall and G. Herbert Fowler
(1887, 1889); and A. Milnes Marshall and William P. Marshall (1882); T. H. Tizard et al. (1885); and
Theéophile Studer (1891), who named the genus Gyrophyllum.

The Darwinian Period produced numerous studies of evolutionary relationships for many groups
of organisms. Important works pertaining to pennatulacean phylogeny during this time included
R. A. von Kolliker (1870, 1880), Gottlieb von Koch (1878), and A. Milnes Marshall (1887).

The Early Twentieth Century (1900-1949)

Significant contributions in octocorallian systematics (including pennatulaceans) during the early
part of the Twentieth Century include Hector F. E. Jungersen (1904, 1907, 1917); Charles Gravier
(1906), who named the genus Scytaliopsis; Charles Cleveland Nutting (1908), who named the genus
Calibelemnon; Heinrich Balss (1909, 1910); Hjalmar Broch (1910-1961); Willy Kitkenthal
(1902-1925) (Pl. 6), who named the genera Actinoptilum, Amphiacme, and Chunella; W. Kiikenthal
and H. Broch (1911), who named the genus Cavernulina; Sydney John Hickson (1883-1940);

26

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PLATE 3

Baron Georges Cuvier (1769-1832). Courtesy Jean Baptiste Pierre Antoine de Monet de Lamarck (1744-

GS Myers/AE Leviton Portrait File in Natural 1829). Courtesy GS Myers/AE Leviton Portrait File in
History, California Academy of Sciences. Natural History, California Academy of Sciences.

Christian Gottfried Ehrenberg (1795-1876). Courtesy James Dwight Dana (1813-1895). Courtesy
of the Linnean Society of London. Smithsonian Institution Archives.

WILLIAMS: INDEX PENNATULACEA

PLATE 4

Henri Milne Edwards (1800-1885).
Courtesy Nationaal Natuurhistorisch Museum, Leiden.

Pieter Bleeker (1819- 1878). Courtesy
Nationaal Natuurhistorisch Museum, Leiden.

~ William Gabb (1839-1878). Courtesy Department of

Jan Adrianus Herklots (1820-1872).
Courtesy Nationaal Natuurhistorisch Museum, Leiden.

Invertebrate Zoology and Geology, California
Academy of Sciences.

28 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 2

PLATE5

Se cha

HMS Challenger (1872-1876), Sir Charles Wyville Thomson, Chief scientist (inset).
After “Voyage of the Challenger,” Narrative, Vol. | (1885) and “The Atlantic,” Vol. 1 (1877) respectively.

Robert Edwards Carter Stearns (1827-1909). Courtesy Ambrosius Arnold Willem Hubrecht (1853-1915).
Bancroft Library, University of California, Berkeley. Courtesy Nationaal Natuurhistorisch Museum, Leiden.

WILLIAMS: INDEX PENNATULACEA 29

Elisabeth Deichmann (1936, 1941) (PI. 6); and J. Arthur Thomson (1905—1931). The latter published
in collaboration with several co-authors including George Crane, Doris L. Mackinnon, Nita I. Rennet,
W.D. Henderson, James Ritchie, and James J. Simpson. F. M. Bayer (pers. comm.) stated,
“Deichmann ... is the 20th Century link with the 19th Century in octocoral systematics. Her
Blake report (1936) is pivotal for the western Atlantic.”

It was during this period that the proposal was introduced by Libbie Henrietta Hyman (1940:365)
to abandon the name “Coelenterata” of Rudolph Leuckart in favor of “Cnidaria,” introduced by
Berthold Hatschek. This argument has apparently convinced many zoologists to substitute a name
long in use with a more recently created one. Hyman stated, “Leuckart (1847) clearly grasped the
fundamental differences between the two great radiate groups, the coelenterates and the echinoderms,
and separated them, creating the name Coelenterata. Leuckart’s Coelenterata, however, included the
sponges and the ctenophores ... . The proper splitting of Leuckart’s Coelenterata was achieved by
Hatscheck (1888), who recognized three phyla: Spongiaria, Cnidaria, and Ctenophora. We therefore
consider Cnidaria to be the most suitable name for the phylum... .”

The acceptance of this line of reasoning is both unnecessary and inconsistent. As an example,
Ernst Haeckel’s “Scyphozoa” originally included both the Scyphozoa and the Anthozoa, yet the name
Scyphozoa is still retained by most zoologists. This is just one example of a major animal group in
which the name has long usage and is still recognized today, even though it orginally encompassed
more taxa than are now accepted.

The Late Twentieth Century (1950-1999)

Previous to this period, publications on the Pennatulacea dealt predominantly with descriptive
taxonomy or general natural history (particularly bioluminescence). This most recent period has
produced a flourishing of works in fields of study other than taxonomy—including behavior, ecology,
morphology, physiology, cell biology, natural products biochemistry, and evolutionary biology.
Important contributions in systematics during this period include Frederick M. Bayer (1955—1996)
(Pl. 6); Huzio Utinomi, also known as Fujio Hiro (1956-1982) (Pl. 6); Manfred Grasshoff
(1972-1991); F. M. Bayer, M. Grasshoff, and Jakob Verseveldt (1983); F. M. Bayer and M. Grasshoff
(1997); Marie-José d’Hondt (1984); Andrée Tixier-Durivault (1954-1987), who named the genera
Crassophyllum and Malacobelemnon, and Gary C. Williams (1986-1999). The latter author intro-
duced cladistic methodology to the study of evolutionary relationships in the Pennatulacea.

SYNONYMS AND MISSPELLED GENERIC NAMES OF PENNATULACEA

Revised from Williams, 1995a:101; synonyms are listed on the left with the valid genera to which
they belong shown in parentheses; * = misspelled names or transcription errors; ** = see Bayer
and Grasshoff, 1997.

*Acanthoptilon Traquair in Zoological Record 7, 1870 (Acanthoptilum)
Actinoptilon Kikenthal, 1910 (Actinoptilum)

*Actinoptinum Day et al., 1970 (Actinoptilum)

Amphianthus Kiikenthal, 1902 (Amphiacme)

Argentella Gray, 1870 (Pteroeides)

Balticina Gray, 1870 (Halipteris)

Bathypenna Marion, 1906 (possibly synonymous with Gyrophyllum according to Kiikenthal, 1915)
Bathyptilum Kolliker, 1872 (Kophobelemnon)

Benthoptilum Verrill, 1885 (Anthoptilum)

*Benthoptillum Haddon in Zoological Record 22, 1885 (Anthoptilum)
Cladiscus Koren and Danielssen, 1877 (Virgularia)

30

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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PLATE 6

Willy Kikenthal (1861-1922). Courtesy Elizabeth Deichmann (1896-1975). Courtesy
Nationaal Natuurhistorisch Museum, Leiden. Frederick M. Bayer, Smithsonian Institution.

Frederick M. Bayer | Huzio Utinomi
Courtesy Frederick M. Bayer, Smithsonian Institution.

WILLIAMS: INDEX PENNATULACEA 3]

Clavella Gray, 1870 (Lituaria)

Crinillum Harting, Miquel and Hoeven, 1861 (Umbellula)

Crispella Gray, 1870 (Pteroeides)

Deutocaulon Marshall and Fowler, 1888 (Virgularia)

Diibenia Danielssen and Koren, 1884 (Sty/atula)

Encrinus Lamarck, 1801 (Umbellula)

Fusticularia Simpson, 1905 (probably synonymous with Cavernularia according to Kiikenthal,
1915)

*Godefrovia Leuckart, 1872 (Pteroeides)

Godeffroyia Kolliker, 1870 (Pteroeides)

Géndul Koren and Danielssen, 1883 (Halipteris)

Gunneria Danielssen and Koren, 1884 (Kophobelemnon)

Halisceptrum Herklots, 1863 (Virgularia)

Helicoptilum Nutting, 1912 (Distichoptilum)

Herklotsia Gray, 1860 (Renilla)

Juncoptilum Thomson and Henderson, 1905 (Distichoptilum)

*Leioptilum Verrill, 1865 (Ptilosarcus)

*Leioptillum Verrill, 1868 (Ptilosarcus)

Leioptilus Gray, 1860 (Pennatula)

Leioptilus of authors other than Gray, 1860 (Prilosarcus)

Leptoptilum KOlliker, 1880 (Funiculina)

*Lioptilum KOolliker, 1872 (Ptilosarcus)

Lygomorpha Koren and Danielssen, 1877 (Halipteris)

Lygus Herklots, 1858 (Virgularia)

Mesobelemnon Gravier, 1907 (considered synonymous with Sclerobelemnon by Hickson, 1916)

Microptilum K6lliker, 1880 (Halipteris)

Norticina Gray, 1870 (Halipteris)

Ombellulaires Cuvier, 1817 (Umbellula)

** Ombellula Cuvier, 1797 (Umbellula)

Ombellularia Lamarck, 1836 (Umbellula)

Osteocella Gray, 1870 (Halipteris)

Parabelemnon Thomson and Simpson, 1909 (possibly synonymous with Cavernularia according
to Kiikenthal, 1915) (PI. 7)

Pavonaires Cuvier, 1817 (Funiculina)

Pavonaria Schweigger, 1820 (Funiculina)

Pavonaria KOlliker, 1869 (Halipteris)

Penna Bohadsch, 1761 (Pennatula, in part) (name unavailable as the work was suppressed by
ICZN)

Phosphorella Gray, 1870 (Pennatula)

Policella Gray, 1870 (Veretillum)

Prochunella Balss, 1909 (Calibelemnon)

Protocaulon KOlliker, 1880 (Virgularia)

*Preroides Pfeffer, 1886 (Pteroeides)

Pteromorpha Herklots, 1858 (Pteroeides)

Ptilella Gray, 1870 (Pennatula)

*Renila Schweigger, 1820 (Renilla)

Renillina Gray, 1860 (possibly a young example of the alcyoniid soft coral Sarcophyton according
to Kiikenthal, 1915)

Sarcobelemnon Herklots, 1858 (Cavernularia)

32 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES

Volume 51, No. 2

PLATE 7

Veretillum

Lituaria hicksoni OO
After Thomson & Simpson, (ue) SE SL
1909 ’ & Broch, 1911

Parabelemnon indicum
After Thomson & Simpson,
1909

Actinoptilum molle
After Williams, 1995

LR op exerci

WILLIAMS: INDEX PENNATULACEA 33

Sarcophyllum K6lliker, 1870 (Sarcoptilus)

Sceptonidium Richiardi, 1869 (Virgularia)

Stephanoptilum Roule, 1905 (Anthoptilum)

Stichoptilum Grieg, 1887 (Halipteris)

Struthiopteron Broch, 1910 (Pteroeides)

Stylobelemnon Kolliker, 1872 (Cavernularia)

Stylobelemnoides J. A. Thomson and Henderson, 1905 (possibly synonymous with Cavernularia
according to Kiikenthal, 1915)

Svava Danielssen and Koren, 1884 (Virgularia)

Svavopsis Roule, 1908 (Virgularia)

Thesioides J. A. Thomson and Henderson, 1906 (Anthoptilum)

Trichoptilum K6lliker, 1880 (Funiculina)

Umbellaria Schweigger, 1820 (Umbellula)

Umbellularia Lamarck, 1801 (Umbellula)

*Verrilia Litken in Zoological Record 10, 1873 (Halipteris)

Verrillia Stearns, 1873 (Halipteris)

Vorticella Linnaeus, 1767 (Umbellula, in part)

TAXONOMIC INDEX

Original descriptions and other selected taxonomic references—testricted primarily to taxa
considered valid by Williams (1995a, 1997f, and the present work); + = fossil taxa; { = taxa with both
living and fossil representation.

Acanthopitilum Kolliker, 1870
[See also Balss (1910); Bayer (1957); Ktikenthal (1915); Williams (1995a)]
Acanthoptilum album Nutting, 1909
Acanthoptilum agassizi Kolliker, 1872
Acanthoptilum annulatum Nutting, 1909
Acanthoptilum gracile (Gabb, 1863)
Acanthoptilum oligacis Bayer, 1957
Acanthoptilum pourtalesii Kolliker, 1870
Acanthoptilum scalpelifolium Moroff, 1902
Actinoptilum Kikenthal in Kiikenthal and Broch, 1911
[See also Ktikenthal (1910, 1915); Williams (1990, 1995a)]
Actinoptilum molle (Kiikenthal, 1910) (PI. 7)
Amphiacme Kiikenthal, 1903
[See also Hickson (1916); Kiikenthal (1902a, 1915); Kiikenthal and Broch (1911); Williams
(1990, 1995a)]
Amphiacme abysorrum (Kiikenthal,1902)
Anthoptilum Kolliker, 1880 (PI. 9, 10)
[See also Grasshoff (1982a, 1982b); Hickson (1916); Kiikenthal (1915); Kiikenthal and Broch
(1911); Williams (1990, 199Sa)]
Anthoptilum grandiflorum (Verrill, 1879) (Pl. 9)
Anthoptilum murrayi Kolliker, 1880
+Bensonularia Hamilton, 1958
tBensonularia spatulata Hamilton, 1958 [Eocene of New Zealand]
Calibelemnon Nutting, 1908
[See also Balss (1910); Hickson (1916); Ktikenthal (1915); Williams (1990, 1995a)]

34 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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PLATE 8

Echinoptilum echinatum
After Williams, 1995a

Renilla amethystina
After Williams, 1995a

Umbellula thomsoni
After Tizard et al., 1885

Kophobelemnon stelliferum

After Marshall, 1883b | Chunella biflora
; After Hickson, 1916

WILLIAMS: INDEX PENNATULACEA 35

Calibelemnon hertwigi (Balss, 1909)

Calibelemnon indicum (Thomson and Henderson, 1906)

Calibelemnon symmetricum Nutting, 1908

Cavernularia Valenciennes in Milne Edwards and Haime, 1850
[See also Hickson (1916); Hondt (1984b); Kikenthal (1915); Lopez-Gonzalez, Gili, and
Williams (in press); Williams (1989b, 1990, 1995a)]

Cavernularia capitata Williams, 1989

Cavernularia clavata Kiikenthal and Broch, 1911

Cavernularia chuni Kikenthal and Broch, 1911

Cavernularia dayi Tixier-Durivault, 1954

Cavernularia dedeckeri Williams, 1989

Cavernularia elegans (Herklots, 1858)

Cavernularia glans Kolliker, 1872

Cavernularia habereri Kolliker, 1872

Cavernularia luetkeni Kolliker, 1872 (also spelled C. /iitkeni and C. lutkeni)

Cavernularia malabarica Fowler, 1894

Cavernularia mirifica Tixier-Durivault, 1963

Cavernularia obesa Valenciennes in Milne Edwards and Haime, 1850

Cavernularia pusilla (Philippi, 1835)

Cavernulina Kiikenthal and Broch, 1911
[See also Hickson (1916); Hondt (1984b); Kitikenthal (1915); Williams (1989b, 1990, 1995a)}

Cavernulina cylindrica Kikenthal and Broch, 1911

Cavernulina darwini (Hickson, 1921)

Cavernulina grandiflora Hondt, 1984b

Cavernulina orientalis Thomson and Henderson, 1909

Chunella Kikenthal, 1902 (PI. 8)
[See also Ktikenthal (1915); Ktikenthal and Broch (1911); Hickson (1916); Williams (1990,
1995a, 1997c)]

Chunella gracillima Kikenthal, 1902

Crassophyllum Tixier-Durivault, 1961
[See also Williams (1995a, 1995d)}

Crassophyllum cristatum Tixier-Durivault, 1961

Crassophyllum thessalonicae Vafidis and Koukouras, 1991

Distichoptilum Verrill, 1882
[See also Hickson (1916); Ktikenthal (1915); Kiikenthal and Broch (1911); Williams (1990,
1995a)]

Distichoptilum gracile Verrill, 1882

Echinoptilum Hubrecht, 1885
[See also Balss (1910); Hickson (1916); Kiikenthal (1915); Kiikenthal and Broch (1911); Sachs
(1913); Williams (1990, 1995a)]

Echinoptilum asperum Hickson, 1916

Echinoptilum echinatum (Kikenthal, 1910) (PI. 8)

Echinoptilum elongatum Hickson, 1916

Echinoptilum macintoshi Hubrecht, 1885

Echinoptilum minimum Hickson, 1916

Echinoptilum roseum Hickson, 1916

Funiculina Lamarck, 1816
[See also Balss (1910); Herklots (1858); Hickson (1916); Jungersen (1904); Kolliker (1872);
Kiikenthal (1915); Kiikenthal and Broch (1911); Williams (1990, 1995a)]

36 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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PLATE 9

Anthoptilum grandiflorum
After Kiikenthal & Broch, 1911

Protoptilum cyaneum
After Kiikenthal & Broch, 1911

WILLIAMS: INDEX PENNATULACEA Su)

PLATE 10

Anthoptilum thomsoni
After KGlliker, 1880

38 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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Funiculina armata Verrill, 1879

Funiculina parkeri Kikenthal, 1909

Funiculina quadrangularis (Pallas, 1766)

+Graphularia Milne Edwards and Haime, 1850

+ Graphularia ambigua (Morton, 1830) [Cretaceous/Tertiary of southeastern North America]
[See also Shapiro and Ramsdell (1965)]

+Graphularia badenia Strand, 1928

+Graphularia belgica Vincent (1893) [Eocene of Belgium]

*+Graphularia crecelii Andrée, 1912 [Oligocene of Germany]

+Graphularia desertorum Kuhn, 1949

+Graphularia groenwalli Nielsen , 1914 [Cretaceous of Denmark]

+Graphularia incerta Malaroda, 1951 [Oligocene of Italy]

+ Graphularia irregularis Nielsen, 1914 [Cretaceous of Denmark]

+Graphularia kalimnae Chapman and Crespin, 1928 [Tertiary of Australia]

+Graphularia longissima Squires, 1958 [Cretaceous/Tertiary of New Zealand]

+Graphularia meijeri Voight, 1958

+Graphularia nigra Malaroda, 1951 [Oligocene of Italy]

+ Graphularia quadrata Voigt, 1958

+Graphularia salisburgensis Traub, 1938 [Paleocene of Austria]

+Graphularia sp. [Eocene of Hungary]

tGraphularia sulcata Nielsen, 1914 [Cretaceous of Denmark]
[See also Kolosvary (1949)]

+ Graphularia wetherelli Milne-Edwards and Haime, 1850 [Eocene of England]
[See also Davis (1936)]

+ Graphularia? yamakawai Yabe and Sugiyama, 1937 [Pleistocene of Japan]

Gyrophyllum Studer, 1891
[See also Hickson (1916); Ktikenthal (1915); Roule (1905); Studer (1901); Williams (1995a,
1995d)]

Gyrophyllum hirondellei Studer, 1891

Gyrophyllum sibogae Hickson, 1916 (PI. 14)

Halipteris Kolliker, 1869
[See also Willams (1990, 1995a)]

Halipteris africana (Studer, 1879)

Halipteris californica (Moroff, 1902)

Halipteris christii (Koren and Danielssen, 1847)

Halipteris finmarchica (Sars, 1851)

Halipteris heptazooidea Acuna and Zamponi, 1992

Halipteris willemoesi KOlliker, 1870

Kophobelemnon Asbjornsen, 1856
[See also Balss (1910); Danielssen and Koren (1884); Herklots (1858); Hickson (1916); Jungersen
(1904); Kolliker (1872); Kiikenthal (1915); Kiikenthal and Broch (1911); Thomson and Simpson
(1909); Williams (1990, 1995a)]

Kophobelemnon affine Studer, 1894

Kophobelemnon heterospinosum Kiikenthal, 1910

Kophobelemnon hispidum Nutting, 1912

Kophobelemnon irregulatus Keller, Pasternak and Naumov, 1975

Kophobelemnon leucharti Cecchini, 1917

Kophobelemnon macrospinosum J. A. Thomson, 1927

Kophobelemnon molanderi Pasternak, 1975

WILLIAMS: INDEX PENNATULACEA 39

Kophobelemnon pauciflorum Hickson, 1916

Kophobelemnon stelliferum (Miller, 1776) (PI. 8)

Lituaria Valenciennes in Milne Edwards and Haime, 1850
[See also Balss (1910); Gray (1870); Hickson (1916); Hondt (1984b); Light (1921); Kolliker
(1872); Kiikenthal (1915); Thomson and Simpson (1909); Williams (1990, 1995a)]

Lituaria amoyensis Koo, 1935

Lituaria australasiae (Gray, 1860)

Lituaria breve Light, 1921

Lituaria habereri Balss, 1910

Lituaria hicksoni Thomson and Simpson, 1909 (PI. 7)

Lituaria kuekenthali Light, 1921

Lituaria molle Light, 1921

Lituaria phalloides (Pallas, 1766)

Lituaria philippinesis Light, 1921

Lituaria valenciennesi Hondt, 1984

Malacobelemnon Tixier-Durivault, 1965
[See also Williams (1995a)]

Malacobelemnon stephensoni Tixier-Durivault, 1965

+Octobasis Malecki, 1982 [Upper Cretaceous of Poland]

+Pavonaria portisi Angelis, 1895? [Tertiary of Italy]

+Pavonaria? singularis Malaroda, 1951 [Oligocene of Italy]

Pennatula Linnaeus, 1758 (PI. 12, 13)
[See also Balss (1910); Gray (1870); Herklots (1858); Kolliker (1869); Lamarck (1816); Leuckart
(1872); Hickson (1916); Kiikenthal (1915); Kiikkenthal and Broch (1911); Williams (1990, 1995a,
1995d)]

Pennatula aculeata Danielssen, 1860

Pennatula argentina Acuna and Zamponi, 1992

Pennatula delicata Tixier-Durivault, 1966

Pennatula fimbriata Herklots, 1858

Pennatula grandis Ehrenberg, 1834

Pennatula indica Thomson and Henderson, 1906

Pennatula inflata Kiikenthal, 1910 (PI. 13)

Pennatula mollis Alder, 1867 (considered a junior synonym of Pennatula phosphorea by

Cornelius and Garfath, 1980, after examination of the two well-preserved syntypes)

Pennatula phosphorea Linnaeus, 1758 (PI. 13)

Pennatula prolifera Jungersen, 1904

Pennatula rubra (Ellis, 1764)

+Pennatulites Nelli, 1903

+Pennatulites manzonii Nelli, 1903 [Miocene of Italy]

+ Prographularia Frech, 1890

+ Prographularia tradica Frech, 1890 [Triassic]

Protoptilum Kolliker, 1872
[See also Balss (1910); Hickson (1916); Jungersen (1904); K6lliker (1880); Kitikenthal (1915);
Kikenthal and Broch (1911); Williams (1995a)]

Protoptilum carpenteri Kélliker, 1872

Protoptilum celebense Hickson, 1916

Protoptilulm cyaneum Kikenthal, 1910 (PI. 9)

Protoptilum denticulatum Jungersen, 1904

Protoptilum smitti Kolliker, 1872

40 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 2

Protoptilum thomsoni KOlliker, 1872

Pteroeides Herklots, 1858 (Pl. 14)
[See also Gray (1870); Hickson (1916); Hondt (1984a); K6lliker (1872); Kiikenthal (1915);
Kiikenthal and Broch (1911); Leuckart (1872); Williams (1990, 1995a, 1995d)]

Pteroeides acutum Tixier-Durivault, 1966

t Pteroeides argenteum (Ellis and Solander, 1786) [Tertiary of Kei Islands]
[See also Bayer (1955a)]

Pteroeides bankanense Bleeker, 1859

Pteroeides bestae Hondt, 1984

Pteroeides breviradiatum KOlliker, 1869

Pteroeides caledonicum KOlliker, 1869

Pteroeides carnosum Tixier-Durivault, 1972

Pteroeides crossieri Tixier-Durivault, 1966

Pteroeides densum Tixier-Durivault, 1966

Pteroeides duebeni Kolliker, 1869

Pteroeides dofleini Balss, 1909

Pteroeides durum Kolliker, 1872

Pteroeides esperi Herklots, 1858

Pteroeides flexuosum Tixier-Durivault, 1966

Pteroeides humesi Tixier-Durivault, 1966

Pteroeides hymenocautlon Bleeker, 1859

Pteroeides isosceles J. S. Thomson, 1915

Pteroeides jungerseni Broch, 1910

Pteroeides laboutei Hondt, 1984

Pteroeides latissimum Kolliker, 1869

Pteroeides lusitanicum Broch, 1910

Pteroeides malayense Hickson, 1916

Pteroeides morbusus Tixier-Durivault, 1961

Pteroeides oblongum Gray, 1860

Pteroeides sagamiense Moroff, 1902

Pteroeides sparmanni Kolliker, 1869

Pteroeides spicatum Tixier-Durivault, 1972

Pteroeides spinosum (Ellis, 1764)

Pteroeides tenerum KOlliker, 1869

Pteroeides timorense Hickson, 1916

Pteroeides triangulum Tixier-Durivault, 1972

Ptilosarcus Verrill, 1865
[See also Leuckart (1872); Nutting (1909); Verrill (1865); Williams 1995a, 1995d)]

Ptilosarcus gurneyi (Gray, 1860)

Ptilosarcus undulatus (Verrill, 1865)

Renilla Lamarck, 1816 (PI. 8)
[See also Ehrenberg (1834); Eisen (1876); Gray (1860); Herklots (1858); Kélliker (1871b);
Kiikenthal (1915); Kiikenthal and Broch (1911); Miiller (1864); Williams (1995a); Zamponi and
Pérez (1996); Zamponi, Pérez, and Capitoli (1997)]

Renilla koellikeri (also spelled R. kollikeri or R. kollikeri) Pfeffer, 1886

Renilla muelleri (also spelled R. mulleri or R. miilleri) Kélliker, 1872

Renilla musaica Zamponi and Pérez, 1995

Renilla octodentata Zamponi and Pérez, 1995

Renilla reniformis (Pallas, 1766)

WILLIAMS: INDEX PENNATULACEA 4]

Renilla tentaculata Zamponi, Pérez, and Capitoli, 1997
Sarcoptilus Gray, 1848
[See also Williams (1995a, 1995c, 1995d)]
Sarcoptilus bollonsi (Benham, 1906)
Sarcoptilus grandis Gray, 1848
Sarcoptilus nullispiculatus Williams, 1995
Sarcoptilus rigidus Williams, 1995
Sarcoptilus shaneparkerii Williams, 1995
Sclerobelemnon KOlliker, 1872
[See also Balss (1910); Hickson (1916); K6lliker (1872); Ktikenthal (1915); Kiikenthal and Broch
(1911); Thomson and Henderson (1906b); Williams (1995a, 1995d)]
Sclerobelemnon burgeri (Herklots, 1858)
Sclerobelemnon elongatum Hickson, 1916
Sclerobelemnon gracile (Gravier, 1908)
Sclerobelemnon gravieri Hickson, 1916
Sclerobelemnon koellikeri Thomson and Henderson, 1906
Sclerobelemnon magniflorum Hickson, 1916
Sclerobelemnon schmeltzi Kolliker, 1872
Sclerobelemnon theseus Bayer, 1959
Scleroptilum Kolliker, 1880
[See also Balss (1910); Kiikenthal (1915); Ktikenthal and Broch (1911); Williams (1990, 1995a)|
Scleroptilum grandiflorum Kolliker, 1880
Scytaliopsis Gravier, 1906
[See also Gravier (1908); Kiikenthal (1915); Williams (1990, 1995a)}
Scytaliopsis djiboutiensis Gravier, 1906
Scytaliopsis ghardagensis (?Gravenhorst, 1821) (also spelled S. ghardaqana; a probable nomen
nudem since it 1s unclear whether this citation represents an original description of a new taxon,
or merely the use of an unsubstantiated name)
[See also Atiya (1994); Schuhmacher and Hinterkircher (1996)]
Scytalium Herklots, 1858
[See also Balss (1910); Hickson (1916); K6lliker, 1870; Kikenthal (1915); Kiikenthal and Broch
(1911); Thomson and Simpson (1909); Williams (1995a)]
Scytalium balssi Hickson, 1916
Scytalium martensi K6lliker, 1870
Scytalium sarsii Herklots, 1858
Scytalium tentaculatum KOlliker, 1880 (Pl. 11)
Stachyptilum Kolliker, 1880
[See also Kiikenthal (1915); Kiikenthal and Broch (1911); Williams (1995a)}
Stachyptilum dofleini Balss, 1909
Stachyptilum macleari KOlliker, 1880
Stachyptilum superbum Studer, 1894
Stvlatula Verrill, 1864
[See also Balss (1910); Jungersen (1904); Kolliker (1870); Kiikenthal (1915); Kiikenthal and
Broch (1911); Williams (1995a)]
Stvlatula antillarum KOolliker, 1870
Stylatula brasiliensis (Gray, 1870)
Stylatula darwini Kolliker, 1870
Stylatula diadema Bayer, 1959
Stylatula elegans (Danielssen, 1860)

42 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 2

PLATE 11

Scytalium tentaculatum
After Kélliker, 1880

WILLIAMS: INDEX PENNATULACEA 43

PLATE 12

Pennatula naresi
After Kélliker, 1880

44 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 2

PLATE 13

(a3 Sia Bebe ie

Virgularia schultzei, upper left; Virgularia schultzei, lower left; Pennatula inflata, center; Pennatula phosphorea, right.
After Kiikenthal & Broch, 1911

WILLIAMS: INDEX PENNATULACEA 45

PLATE 14

Pteroeides punctatum
After Thomson & Simpson, 1909

Gyrophyllum sibogae
After Hickson, 1916

46 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 2

Stylatula elongata (Gabb, 1862)

Stvlatula gracilis (Gabb, 1864)

Stylatula kinbergi Kolliker, 1870

Stylatula lacazi K6lliker, 1870

Stylatula polyzoidea Zamponi and Pérez, 1996

Umbellula Cuvier, 1797 (see Bayer and Grasshoff, 1997) (PI. 1, 8)
[See also Broch (1957, 1958a); Cuvier (1800); Ellis (1753, 1755); Grasshoff (1971, 1973, 1982a,
1982b); Heezen and Hollister (1971); Hickson (1916); K6lliker (1875, 1880); Kiikenthal (1902b,
1915); Kiikenthal and Broch (1911); Lamarck (1801); Mylius (1755); Pasternak (1975); Williams
(1990, 1993b, 1995a, 1997a, 1997c); Williams and Rogers (1989)]

Umbellula durissima K6lliker, 1880

Umbellula encrinus Linnaeus, 1758

Umbellula hemigymna Pasternak, 1975

Umbellula huxleyi Kolliker, 1880

Umbellula lindahli Kolliker, 1874

Umbellula monocephalus Pasternak, 1975

Umbellula pellucida Kiikenthal, 1902

Umbellula spicata Kiikenthal, 1902

Umbellula thomsoni KOolliker, 1874 (PI. 8)

Veretillum Cuvier, 1797
[See also Balss (1910); Gray (1870); Herklots (1858); Hickson (1916); Kukenthal (1915);
Kiikenthal and Broch (1911); Williams (1990, 1995a)]

Veretillum australis (Gray, 1870)

Veretillum cynomorium (Pallas, 1766) (PI. 7)

Veretillum leloupi Tixier-Durivault, 1960

Veretillum malayense Hickson, 1916

Veretillum manillensis (K6lliker, 1872)

Veretillum tenuis (Marshall and Fowler, 1889)

Veretillum vanderbilti Boone, 1938

Virgularia Lamarck, 1816
[See also Herklots (1858); K6lliker (1880); Koren and Danielssen (1877); Kiikenthal (1915);
Kikenthal and Broch (1911); Richiardi (1869); Williams 1990, 1995a)]

Virgularia abies Kolliker, 1870

Virgularia brochi Kikenthal, 1915

Virgularia bromleyi Kolliker, 1880

Virgularia densa Tixier-Durivault, 1966

Virgularia galapagensis Hickson, 1930

Virgularia glacialis K6lliker, 1870

Virgularia gracillima Kolliker, 1880

Virgularia gustaviana (Herklots, 1863)

Virgularia halisceptrum Broch, 1910

Virgularia juncea (Pallas, 1766)

Virgularia kophameli May, 1899

Virgularia loveni Utinomi, 1971

Virgularia mirabilis (Miller, 1776)

Virgularia patagonica (a probable nomen nudem since an original description cannot be found and

a type specimen has not been located)

[See also Barrattini and Ureta, 1960; Darwin, 1860]

Virgularia patachonica (see Virgularia patagonica)

WILLIAMS: INDEX PENNATULACEA 47

t Virgularia presbytes Bayer, 1955a [Tertiary of Trinidad]
[See also Bayer (1957); Belem and Figueiredo Alvarenga (1973)]
Virgularia reinwardti Herklots, 1858
Virgularia rumphi Kolliker, 1870
Virgularia schultzei Kikenthal, 1910 (Pl. 13)
Virgularia tuberculata Marshall, 1883

GEOGRAPHIC INDEX

Africa — eastern (see Indo-Pacific)

Africa — northern (see North Atlantic and Mediterranean)

Africa — southern
Branch et al. (1994); Broch (1939, 1940); Day (1974b); Day et al. (1970); Molander (1929);
J.S. Thomson (1915, 1917, 1924); Tixier-Durivault (1954, 1960); Williams (1987, 1989a,
1989b, 1990, 1992, 1993a); Williams and Rogers (1989).

Africa — western
Broch (1910a; 1914b, 1958b); Buchanan (1955); Dollfus (1938); Lopez-Gonzalez et al. (in
press); Molander (1929); Pax and Miller (1954); Studer (1878); Tixier-Durivault (1961b, 1963);
Williams (1987, 1989b, 1990, 1992).

Africa — Atlantic (see Africa — western)

Antarctica (see Southern Oceans)

Arctic Ocean
Broch (1913c, 1955, 1956); Jungersen (1907, 1915, 1917); Lindahl (1874c); Madsen (1948);
May (1900); Mylius (1753, 1754); Pasternak (1980); Verrill (1922); Yashnov (1948).

Asia — northeastern (see Pacific Ocean — northern; or Japan)

Atlantic — eastern (see North Atlantic and Mediterranean or Africa — western)

Atlantic — northern (see North Atlantic and Mediterranean)

Atlantic — western temperate (see North Atlantic and Mediterranean or South America — Atlantic)

Atlantic — western tropical (see Western Tropical Atlantic and Caribbean)

Australia — eastern (see Indo-Pacific)

Australia — northern (see Indo-Pacific)

Australia — southern
Broch (1910c); Briggs (1915); Godfrey (1943); Gray (1848); McCoy (1892); Pasternak (1966);
Utinomi (1971); Utinomi and Shepherd (1982); Williams (1995c).

Australia — western
Broch (1910c); Williams (1995c).

Caribbean Sea (see Western Tropical Atlantic and Caribbean)

Cosmopolitan (see Worldwide)

East Africa (see Africa — eastern)

East Asia (see Indo-Pacific)

Eastern Australia (see Indo-Pacific)

Eastern Pacific — western North America (see North America — Pacific)

Eastern Pacific — western South America (see South America — Pacific)

Europe (see North Atlantic and Mediterranean)

Indo-Pacific (including Red Sea*)
Alcock (1902); Allen and Steene (1994); *Atiya (1994); Briggs (1915); Clastres et al. (1984);
Colin and Arneson (1995); Dinamani (1965); Fowler (1894); Gosliner et al. (1996); Gravely
(1941); *Gravenhorst (1821); *Gravier (1906a, 1906b, 1906c, 1907a, 1908, 1912a); Hickson
(1905); Hondt (1984a, 1984b); Hornell (1922); Humes (1978); Imahara (1991); Koo (1935,

48 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 2

1940); *Kiikenthal (1913b); Light (1921); *Magnus (1966); A. M. Marshall and Fowler (1889);
Mather and Bennet (1993); Nakasone and Yu (1987); Nutting (1908); Pasternak (1964); Ridley
(1883); Roule (1908); Sankolli and Neelakantan (1971); *Schuhmacher and Hinterkircher
(1996); Simpson (1905); J. A. Thomson (1905); J. A. Thomson and Crane (1909a, 1909b); J. A.
Thomson and Henderson (1905a, 1905b, 1906a, 1906b); Tixier-Durivault (1960, 1965, 1966,
1972): Tixier-Durivault and Hondt (1974a); Utinomi (1956b).

Japan
Balss (1910); Fisher (1874); Hubrecht (1885); Kumano (1937); Moroff(1902b); Nutting (1912);
Okutani (1969); Stearns (1883); Stimpson (1855); J. A. Thomson and Rennet (1927); Utinomi
(1956a, 1958, 1961, 1964).

Mediterranean Sea (see North Atlantic and Mediterranean)

New Zealand .
Benham (1906, 1907); Dendy (1897); Hamilton (1958); Hutton (1904); Powell (1947); Williams
(1995c).

North America — temperate Atlantic (see North Atlantic and Mediterranean)

North America — Pacific
Allen (1969); Anonymous (1898); Batie (1972); Belcik (1977); Blake (1872, 1873); Brusca
(1980); Dawson (1872); Fautin et al. (1987); Flora and Fairbanks (1966); Gabb (1862, 1864);
Gotshall (1987, 1994); Gotshall and Laurent (1979); Hartman (1960); Hickson (1930a);
Hochberg and Ljubenkov (1998); Human (1973); Johnson and Snook (1935); Kerstitch (1989);
Kozloff (1974, 1983, 1987); Kiikenthal (1913a); Lam et al. (1982); Nutting (1909); Ricketts et
al. (1985); Sclater (1872, 1873); Shimek (1998); Stearns (1873a); Strathmann (1988); Studer
(1894): Verrill (1865, 1868a, 1868b).

North Atlantic and Mediterranean (Europe and eastern North America)
Abel (1963); Alder (1862); Alder (1863); Altuna-Prados (1994); Andrée (1912); Angelis
(1895?); Arculeo et al. (1990); Atkinson (1989); Broch (1913a, 1914a, 1953); Baluk and Pisera
(1984); Bellini (1905); Berenguier (1954); Broch (1953); Demir (1952); Dyer et a | (1981):
Eckelbarger et al. (1998); Ellis (1753, 1755, 1764); Field (1949); Fischer (1889); Forbes and
Goodsir (1851); Gili and Pages (1987); Gili et al. (1987); Grasshoff (1981, 1982a, 1982b, 1989);
Grieg (1887, 1892, 1893, 1896); Jahn (1970); Keller et al. (1975); Koren and Danielssen (1847,
1856, 1874, 1877, 1883, 1884); Kramp (1932, 1933, 1950); Lacaze-Duthiers (1891); Langton et
al. (1990); Lindahl (1874a, 1874b, 1874c); Luther and Fiedler (1961); Madsen (1948); Manuel
(1981); Marion (1906); Mylius (1753, 1754); Nobre (1931); Patterson (1986); Pax (1936); Pax
and Miiller (1953, 1955a, 1955b, 1955c, 1959, 1962); Pérés and Picard (1958); Perrier (1936);
Rice et al. (1992); Riedl (1963, 1983); Rossi (1971); Roule (1905); Rowe(1971); Sars (1846,
1851); Stephens (1909); Tixier-Durivault and Lafargue (1968); Vafidis and Koukouras (1991);
Vafidis et al. (1994); Verrill (1878, 1879, 1882a, 1882b,1884a, 1884b, 1885a).

Northern Pacific (see Pacific Ocean — northern)

Pacific Ocean — eastern (see North America — Pacific)

Pacific Ocean — northern
Blake (1872, 1873); Broch (1935); Pasternak (1960, 1961a, 1970); Rho and Song (1976, 1977).

Pacific Ocean — western (see Indo-Pacific)

South America (Atlantic)
Barttini and Ureta (1960); Acufia and Zamponi (1992); ); Belem and Figueiredo Alvarenga
(1973); Darwin (1860); Pasternak (1975b, 1993); Sanchez (1994); Tommasi et al. (1972);
Zamponi and Perez (1996); Zamponi et al. (1997).

South America (Pacific)
Pasternak (1975b); Pérez (1996); Zuniga (1948).

Southeast Asia (see Indo-Pacific)

WILLIAMS: INDEX PENNATULACEA 49

Southern Asia (see Indo-Pacific)

Southern Africa (see Africa — southern)

Southern Australia (see Australia — southern)

Southern Oceans (Antarctica and the subantarctic)
Acuna and Zamponi (1992); Branch and Williams (1993); Ktikenthal (1912a); May (1900);
Pasternak (1961b, 1962); J. A. Thomson and Rennet (1931); Zamponi and Perez (1995a, 1995b).

Subantarctic (see Southern Oceans)

West Africa (see Africa — western)

Western Australia (see Australia — western)

Western Tropical Atlantic and Caribbean
Bayer (1957, 1959, 1961); Cairns, Hartog, and Arneson (1986); Deichmann (1936a, 1936b);
Fowler (1888); Kaplan (1982, 1988); Keller et al. (1975); Pasternak (1975a); Sanchez (1994);
Voss (1976).

Worldwide
Bayer (1956); Kiikenthal (1915); Williams (1995a, 1997e, 1997f).

SUBJECT INDEX

NOTE: The largest percentage, approximately 37%, of the citations listed pertain to
taxonomy and geographic records (see Fig. 1).

Anatomy (see Morphology and Ultrastructure)

Behavior

P. A. V. Anderson (1976); P. A. V. Anderson and Case (1975); Buisson (1974, 1976, 1980,
1988): Darwin (1860); Couet (1979); Dickinson (1978); Hoare and Wilson (1977); Kastendiek
(1975a, 1975b, 1976); Lancaster (1601); Langton et al. (1990); Mori (all citations); Mori and Ondo
(1957); Mori and Tanase (1973); Pantin (1950); Pavans de Ceccatty and Buisson (1965); Takada and
Mori (1956, 1957).

Bibliography
Bayer (1981la, 1996); Hickson (1916); Kiikenthal (1915); D. W. Thompson (1885); Williams
(1990).

Biography (Regarding Authors of Pennatulacean Research)
Boeseman (1973); Forest (1996); Fransen et al. (1997); Gardiner (1940); Holthuis (1993, 1995);
Smit (1979); Zimmer (1925).

Bioluminescence

Agassiz (1850); Aldrovandi (1642, 1648); Anctil et al. (1982); J. M. Anderson and Cormier
(1978); J. M. Anderson et al. (1974, 1978); P. A. V. Anderson and Case (1975); Awad and Anctil
(1993a, 1993b); Batie (1972); C. Bauhin (1620, 1671); J. Bauhin (1650-51); Bilhaut (1975a, 1975b);
Blainville (1834); Boussuet (1558); Buck (1973); Chiaje (1827); Charbonneau (1981); Cormier
(1978); Davenport and Nicol (1956); DeLuca et al. (1976); Dittrich (1888); Ehrenberg (1834c); Ellis
(1764); Forbes (1847); Forbes and Goodsir (1851); Germain and Anctil (1988); Gesner (1555, 1558):
Grant (1827, 1829); Grober (1990a, 1990b); Hart et al. (1979); Harvey (1917, 1920, 1940, 1952);
Hastings (1968, 1983); Hastings and Morin (1969); Herdman (1913a); Herring (1978, 1991); Hori
and Cormier (1973); Hori et al. (1972, 1973); Imperato (1599); Korotneff (1887); Kreiss and Cormier
(1967); Krukenberg (1887, 1888); Mangold (1910); N. B. Marshall (1979); Matern (1984); Matthews

50 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 2

OTHER FIELDS

p

BIOCHEMISTRY & NATURAL PRODUCTS TAXONOMY & GEOGRAPHIC RECORDS

MORPHOLOGY & ULTRASTRUCTURE —

ECOLOGY & BIOTIC INTERACTIONS

Ly

PHYSIOLOGY & CELL BIOLOGY

’ GENERAL NATURAL HISTORY & BIOLOGY

BIOLUMINESCENCE ~.>|
FIGURE |. Quantitative analysis of the literature by field of study.

et al. (1977); Morin (1974, 1976, 1998); Nealson et al. (1986); Nicol (1955a, 1955b, 1955c, 1958);
Panceri (1871b, 1871c, 1872a, 1872b, 1872c); Parker (1920c); Péron (1804); Pratje (1923); Rapp
(1827); Royal Society (1870); Satterlie and Case (1979); Shaw (1838—46); Shimomura and Johnson
(1975, 1979); Soares and Sawaia (1975); Spallanzani (1784, 1796); Spurlock and Cormier (1975);
Tilesius von Tilenau (1819); Titschack (1965, 1966); Tizard et al. (1885); Wampler et al. (1971,
1973); Ward (1979); Ward and Cormier (1978a, 1978b); Williams (1990); Wyville Thomson (1874).

Corals (Pre-Linnean General Natural History)
Boccone (1670); Marsigli (1725); Peyssonnel (1753); Rondelet (1554-1555); Woodward (1695).

Ecology and Biotic Interactions

P. K. Anderson (1989); Atkinson (1989); Bertsch (1968, 1982); Best (1988); Birkeland (1969,
1971, 1974); Brafield (1969); Creed and Coull (1984); Davis (1978); Davis et al. (1982); Dawydoff
(1930, 1938); Day (1974a); Dube and Ball (1971); Fager (1968); Fujita and Ohta (1988); ); Gili et al.
(1987); Gilluly (1970); Gosliner et al. (1996); Hornell (1922); Howson and Davies (1991); Humes
(1978); Jones (1960); Kastendiek (1975a, 1975b, 1976, 1982); Lam et al. (1982); Laubier (1972);
Magnus (1966); Mariscal and Bigger (1977); McDonald and Nybakken (1978); Miyajima (1897,
1900); Mori (all citations); Mori and Ondo (1957); Mori and Tanase (1973); Nakasone and Yu (1987);
Okutani (1969); Ramesh et al. (1985); Rittschof et al. (1986, 1988); Rowe (1971); Sankolli and
Neelakantan (1971); Shepherd (1983); Shimek (1998); Short and Trower (1986); Standing et al.
(1984); Thompson et al. (1988); Tyler et al. (1995).

WILLIAMS: INDEX PENNATULACEA 5]

General Natural History and General Biology

Alcock (1902); Allen (1969); Arculeo et al. (1990); Arndt (1912); R. D. Barnes (1987); R. S. K.
Barnes et al. (1988); Bayer (1973, 1981a); Bayer et al. (1983); Bayer and Owre (1968); Bellomy
(1974); Bourne (1900); Branch et al. (1994); Brusca and Brusca (1990); Colin and Arneson (1995);
Coleman and Teague (1973); Cooke (1889); Dakin (1953); Dalyell (1848); Darwin (1860); Delage
and Herouard (1901); Donovan (1995); Dunn (1982); Faulkner and Chesher (1979); Fossa and Nilsen
(1995); Frische (1991); Gage and Tyler (1991); Gesner (1565); Hadi (1994); Hardy (1967); Gosliner
et al. (1996): Gotshall (1987, 1994); Gotshall and Laurent (1979); Gravier (1912b); Haeckel (1904);
Hardy (1965, 1967); Heezen and Hollister (1971); Hickson (1906, 1909, 1930b); Huxlely (1907);
Hyman (1940); Johnson and Snook (1935); Kaplan (1982, 1988);Kerstitch (1989); Koren and
Danielssen (1877); Kozloff (1974, 1983, 1987, 1990); Kiikenthal (1923-1925); Lancaster (1601);
Luther and Fiedler (1961); Lutz (1986); Lydekker (no date); MacGinitie (1938); MacGinitie and
MacGinitie (1968); N.B. Marshall (1979); W.P. Marshall (1895); Mather and Bennet (1993);
Meglitsch (1972); Milne and Milne (no date); Milne Edwards and Haime (1857); Ming (1993);
Modeer (1786); H. B. Moore (1937); Moseley (1872); Moss (1878); Mylius (1753, 1754); Nordgaard
(1905); Omori (1991); Pearse et al. (1987); Philippi (1835); Pimentel (1967); Ricketts et al. (1985);
Ried! (1963); Ricketts et al. (1985); Rondelet (1554-1555); Rumphius (1705); Schechter (1959);
Sch6mann (1949): Schuhmacher and Hinterkircher (1996); Sclater (1872, 1873); Smith (1964): Smith
and Carlton (1975); Sphon (1964); Sprung and Delbeek (1997); Thurston (1890); Tilesius (1812);
Tizard et al. (1885); Tixier-Durivault (1987); A. Trembley (1744); Tyler and Zibrowius (1992);
Utinomi (1956a, 1964): Voss (1976); Wilkins and Birkholz (1986); Williams (1986, 1993a, 1995a,
1997c); Wyville Thomson (1874); Zim and Ingle (1955).

Histology
Hasama (1944); Kdlliker (1865); Korotneff (1887); Lyke (1965); Niedermeyer (1913, 1914);
Titschack (1970).

History of Science (Regarding Coelenterates and/or Sea Pens)
Agassiz (1860); Bayer (1981a); Hickson (1916); Hyman (1940); Shapiro and Ramsdell (1965);
Williams (1993b, 1995a, 1997c).

Molecular Biology and Genetics
Lorenz et al. (1991).

Morphology and Ultrastructure

Alonso (1979); Beklemishev (1969a, 1969b); Buisson (1970); Buisson and Franc (1969); Bujor
(1901); Chia and Crawford (1977); Crawford and Chia (1974); Bullough (1950); Dunkelberger and
Watabe (1972, 1974): Fautin and Mariscal (1991); S. France (1979); S. Franc et al. (1971, 1974, 1985);
Franzén (1967); Germain and Anctil (1988); Hickson (1883); Ivester and Dunkelberger (1971);
Jungersen (1888a, 1888b); Koch (1878, 1889, 1890); Kélliker (1865, 1871b, 1872, 1874); Korotneff
(1887); Ledger and Franc (1978); Lenhoff et al. (1971); Lightbown (1918); Lyke (1965); Mariscal
(1974, 1979); Mariscal and Bigger (1977); Marks et al. (1949); Niedermeyer (1911, 1912); Panceri
(1870); Parker (1919); Roule (1907); Satterlie et al. (1976); Shapeero (1969); Spurlock and Cormier
(1975); Titschack (1966, 1968, 1970); P. Trembley (1941, 1942); Watabe and Dunkelberger (1979);
Wilbur (1976); Wilson (1884).

Natural Products Chemistry, Biochemistry, and Toxicology
Anctil (1987, 1989a, 1989b); Anctil et al. (1982, 1984, 1991); J. M. Anderson and Cormier
(1978); J. M. Anderson et al. (1974, 1978); P. A. V. Anderson (1976); Awad and Anctil (1993a,

52 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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1993b); Bernheimer and Arigad (1981); Bullock (1970); Charbonneau (1981); Clastres et al. (1984);
Coan and Travis (1970); Datta et al. (1990); DeLuca et al. (1976); Fu et al. (in press); Goswami et al.
(1995); Grimmelikhuijzen and Groeger (1987); Grimmelikhuijzen et al. (1987); Harmon etal. (1984);
Hart et al. (1979); Hori and Cormier (1973); Hori et al. (1972, 1973); Huang and Mir (1972); Jones
et al. (1979): Karkhanis and Cormier (1971); Keifer et al. (1986); Kittredge et al. (1962); Kreiss and
Cormier (1967); Morin (1976); Kumar et al. (1990); Lorenz et al. (1991); Mackie (1987); Pani and
Anctil (1994); Shapeero (1969); Shimomura and Johnson (1975, 1979); Standing et al. (1984);
Thompson et al. (1988); Tillet-Barret et al. (1992); Vanderah and Djerassi (1977); Vidal et al. (1992);
Waele et al. (1987); Wampler et al. (1971); Wekell (1974, 1978); Wratten et al. (1977a, 1977b).

Paleontology (Mesozoic and Cenozoic Fossil Taxa)

Andrée (1912); Angelis (1895?); Baluk and Pisera (1984); Bayer (1955a, 1955b, 1956); Bradley
(1980, 1981); Branco (1885); Chapman and Crespin (1928); Davis (1936); Fowler (1911); Frech
(1890); Gabb (1859, 1861); Gregorio (1890); Hamilton (1958); Hantzschel (1958); Howell (1947);
Kolosvary (1949); Kuhn (1949); Malaroda (1951); Malecki (1982); Milne Edwards and Haime
(1850); Morton (1830, 1834); R.C. Moore (1956); Nelli (1903); Nielsen (1914); Shapiro and
Ramsdell (1965); Roemer (1880); Squires (1958); Strand (1928); Traub (1938); P. Trembley (1941,
1942): Valenciennes (1850); Vincent (1893); Voigt (1958); Yabe and Sugiyama (1937).

Paleontology (Vendian Frondlike Fossil Taxa)

Arthur (1997); Bergstrom (1989, 1991); Buss and Seilacher (1994); Conway Morris (1991,
1993); Deng and Chen (1981); Dzik (1991); Fedonkin (1992, 1996); Glaessner (1958a, 1958b, 1959,
1961, 1984); Glaessner and Daily (1959); Glaessner and Wade (1966); Jenkins (1985, 1992); Jenkins
and Gehling (1978); Lewin (1984); Liu (1981, 1983); Richter (1955); Runnegar (1992); Seilacher
(1989); Simonetta and Conway Morris (1991); Weiguo (1986); Williams (1995b, 1997c, 1997d);
Wright (1997).

Photography (Color Photographs of Living Sea Pens)

Allen and Steen (1994): Branch et al. (1994); Colin and Arneson (1995); Faulkner and Chesher
(1979): Gosliner et al. (1996); Fossa and Nilsen (1995); Gotshall (1987,1994); Gotshall and Laurent
(1979); Kerstitch (1989); Ming (1993); Nishimura (1992); Schumacher and Hinterkircher (1996);
Shimek (1998); Sprung and Delbeek (1997); Weinberg (1996); Williams (1990, 1996); Wilkins and
Birkholz (1986).

Phylogeny, *Cladistics, Evolution, and Biogeography

Altuna-Prados (1994); Bergstrém (1989, 1991); Bourne (1900); Broch (1913b); Conway Morris
(1991); Grasshoff (1973, 1991); Hickson (1916, 1930b); Kinoshita (1912); Koch (1878); Kolliker
(1869-72, 1872, 1880); Kiikenthal (1912b, 1914, 1915a, 1921); Kiikenthal and Broch (1910, 1911);
A. M. Marshall (1883b); Niedermeyer (1913); Patterson (1986); Williams (1992, *1993b, 1995a,
*1995b, *1995d, 1997a, 1997b, *1997c, 1997d, 1997e).

Physiology and Cell Biology

Anctil (1987, 1989a, 1989b); Anctil et al. (1982, 1984, 1991); J. M. Anderson and Cormier
(1978); J. M. Anderson et al. (1974); P. A. V. Anderson (1976); P. A. V. Anderson and Case (1975);
Awad and Anctil (1993a, 1993b); Bilhaut and Pavans de Ceccatty (1971a, 1971b); Brafield (1969);
Brafield and Chapman (1967); Case and Morin (1966); Chapman (1972); Child (1951); Buck and
Hanson (1967); Buisson (1964, 1969, 1971a, 1971b, 1973, 1976, 1979, 1988); Carlgren (1940);
Charbonneau (1981); Crawford and Chia (1974); Dickinson (1978); J. M. Franc (1979); S. Franc
(1970, 1973, 1979, 1980); Hagiwata et al. (1981); Honjo (1940); Imafuku (1973, 1975, 1976); Ivester

WILLIAMS: INDEX PENNATULACEA 53

(1977); Korotneff (1887); Krukenberg (1887); Ledger and Franc (1978); A. R. Moore (1926); Mori
(all citations); Mori and Ondo (1957); Mori and Tanase (1973); Musgrave (1909); Nicol (1955b,
1955c); Parker (1920a, 1920b); Pavans de Ceccatty and Buisson (1964a, 1964b, 1965); Pavans de
Ceccatty et al. (1963); Pratt (1909); Satterlie et al. (1976, 1980); Senut and Franc (1985); Takada and
Mori (1956, 1957); Titschack (1965, 1966, 1968, 1970); Umbriaco et al. (1990); Waele et al. (1987).

Polymorphism, Phenotypic Variability, and Biodiversity
Hickson (1903b, 1903c); Human (1973); Jaworski (1939); A. M. Marshall (1883); Pasternak
(1989); Williams (1992, 1993b, 1997b).

Quoted Passages

Agassiz (1860); Harvey (1952); Hyman (1940); Lankaster (1601); N. B. Marshall (1979); Peron
(1804); C. W. Thomson (1874); Shapiro and Ramsdell (1965) quoted under Gregorio (1890); Tizard
et al. (1885), Williams (1993b).

Reproductive and Developmental Biology (including Growth Stages and Regeneration)

Birkeland (1969, 1971); Chia and Crawford (1973); Dalyell (1839); Delage and Herouard (1901);
Eckelbarger et al. (1998); Franzen (1967); Jungersen (1888a, 1888b); Korschelt (1936); Lacaze-Duth-
iers (1865, 1887); Mori and Tanase (1973); Roule (1932); Satterlie and Case (1979); Strathmann
(1988): Tarent and Tarent (1980); Torrey (1901); Tyler et al. (1995); Willemoes-Suhm (1875); Wilson
(1880, 1881, 1882a, 1882b, 1883b, 1903).

Taxonomy, Distributional and Bathymetric Records

Abel (1963); Acufia and Zamponi (1992); Alder (1861, 1862, 1863, 1867); Allen (1969);
Anonymous (1898); Arculeo et al. (1990); Arndt (1912); Asbornsen (1856); Atiya (1994); Atkinson
(1989): Balss (1909, 1910, 1911); Barattini and Ureta (1960); Barreira y Castro (1990); Batie (1972);
Bayer (1955b, 1956, 1957, 1959, 1961); Bayer and Grasshoff (1997); Belcik (1977); Belem and
Alvarenga (1973); Belyaev (1972); Benham (1906, 1907); Boone (1933, 1938); Branch et al. (1994);
Branch and Williams (1993); Broch (all citations); Buchanan (1955); Cairns, Hartog, and Arneson
(1986); Carpine and Grasshoff (1985); Castro (1981); Colin and Arneson (1995); Costa Soares
(1979): Cuvier (1797, 1800); Dana (1846); Danielssen and Koren (1884); Dawson (1966); Deich-
mann (1936a, 1936b, 1941); Ehrenberg (1834a, 1834b); Ellis and Solander (1786); Erhardt and
Moosleitner (1995); Fautin et al. (1987); Fowler (1888, 1894); Fu et al. (in press); Gabb (1862, 1864);
Gili (1986); Gili and Pages (1987); Gosliner et al. (1996); Gotshall (1987, 1994); Gotshall and Laurent
(1979); Grasshoff (all citations); Gravenhorst (1821); Gravier (all citations); Gray (all citations);
Hartman (1960); Herklots (1858, 1863); Hickson (1890, 1894, 1900, 1903a, 1904, 1905, 1907, 1911,
1914,1916, 1921, 1922, 1930a, 1936, 1937, 1940); Hoare and Wilson (1977); Hochberg and
Ljubenkov (1998); Hondt (1984a, 1984b); Hubrecht (1885); Imahara (1991); Jahn (1970); Johnston
(1847); Jungersen (1904, 1905, 1907, 1915, 1917); K6lliker (all citations); Koo (1935, 1940); Koren
and Danielssen (1847, 1856, 1874, 1877, 1883, 1884); Kiikenthal (all citations); Kramp (1932, 1933,
1950); Kumano (1937); Lacaze-Duthiers (1891); Lamarck (1816, 1836); Langton et al. (1990);
Lepechin (1781); Leuckart (1841); Leunis (1886); Light (1921); Lindahl (1874a, 1874b, 1874c):
Linnaeus (1758, 1767); Lopez-Gonzalez et al. (in press); Madsen (1948); Manuel (1981); A. M.
Marshall (1883b); A. M. Marshall and Fowler (1888); A. M. Marshall and Marshall (1882); May
(1899, 1900); Milne Edwards and Haime (1850, 1857); Molander (1929); Moroff (1902a, 1902b);
O.F. Miiller (1776); F. Miller (1866); Naumov (1955); Nobre (1931); Norman (1867); Nutting (1908,
1909, 1912); Pallas (1766, 1787); Panceri (1871a); Pasternak (all citations); Pax (1936); Pax and
Miiller (1955b, 1955c, 1962); Pérez (1996); Pfeffer (1886); Poche (1914, 1915a, 1915b); Quoy and
Gaimard (1827); Rho and Song(1976, 1977); Richiardi (1869); Richmond (1997); Ridley (1883):

54 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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Ried! (1963, 1983); Riveros Zunica (1948); Robertson (1887); Rossi (1971); Roule (1905, 1906,
1908); Sachs (1913); Sars (1846,1851); M. Schultze (1871); F. E. Schulze (1875); Simpson (1905);
Stearns (1873a, 1873b, 1873c, 1874, 1882, 1883); Stephens (1909); Stiasny (1937, 1938); Stimpson
(1855); Studer (all citations); J. A. Thomson (1905, 1927); J. A. Thomson and Crane (1909a, 1909b);
J. A. Thomson and Henderson (1905a, 1905b, 1906a, 1906b); J. A. Thomson and Mackinnon (1911);
J. A. Thomson and Rennet (1927, 1931); J. A. Thomson and Ritchie (1906); J. A. Thomson and
Simpson (1909); J. S. Thomson (1915, 1917, 1924); Tilesius (1826); Tixier-Durivault (all citations);
Tixier-Durivault and Hondt (1974a, 1974b); Tixier-Durivault and Lafargue (1968); Utinomi (all
citations); Utinomi and Shepherd (1982); Vafidis and Koukouras (1991); Van Soest (1977); Verrill
(all citations); Waterman (1950); Williams (all citations); Wiktor (1974); Yashnov (1948); Zamponi
and Pérez (1995a, 1995b,1996); Zamponi et al. (1997); Zuniga (1948).

BIBLIOGRAPHY

Note: Author’s annotations are in square brackets following selected citations.

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and Berlin.

ACUNA, F. H. AND M. O. ZAMPONI. 1992. Pennatulacaea (Cnidaria: Octocorallia) de la region subantarctica:
nuevos taxa y registros. Iheringia (Série Zoologia) (73):47—S3. [Distributional data: Pennatula rubra,
Anthoptilum grandiflorum; original descriptions: Pennatula argentina, Halipteris heptazooidea. |

AGASSIZ, L. 1850. On the principles of classification in the animal kindgom on the structure of the halcyonoid
polypi; on the morphology of the Medusae. Press of Walker and James, Charleston, South Carolina. 19 pp.
[Sea pansy bioluminescence. ]

. 1860. Contributions to the natural history of the United States of America, Vol. 3. Little, Brown and
Company, Boston. 301 pp. [Pp. 1-35: a detailed discourse on the history of scientific investigation regarding
the coelenterates, including the major contributions of Rondelet, Gesner, Boccone, and others; see quoted
passages in Historical Account—-Pre-Linnean Period, above. ]

ALcock, A. 1902. A naturalist in Indian seas, or, four years with the Royal Indian marine survey ship
“Investigator.” John Murray, London. 328 pp. [Cited in Hickson, 1916:256.]

ALDER, J. 1861. Occurrence of a sea pen, new to Britain. Transactions of the Tyneside Naturalists’ Field Club
5:60—-61. [Halipteris christii, referred to by Alder as Virgularia christii.]

. 1862. Supplement to a catalogue of the zoophytes of Northumberland and Durham. Transactions of the

Tyneside Naturalists’ Field Club 5: 225-247.

. 1863. Report on the zoophytes. /n Report of the dredging expedition to Dogger Bank and the coasts of

Northumberland, H. T. Mennell, ed. Transactions of the Tyneside Naturalists’ Field Club 5:288—290.

. 1867. Notice of some Invertebrata, in connexion with the report of Mr Gwyn Jeffreys on dredging
among the Hebrides. Report of the British Association for the Advancement of Science, part 1:206—208.
[Original description of Pennatula mollis, considered a junior synonym of Pennatula phosphorea by
Cornelius and Garfath, 1908:274; see also Norman, 1867:196, 206. ]

ALDROVANDI, U. 1642. De reliquis animalibus exanguibus libri quatuor, post mortem ejus editi: nempe de
mollibus, crustaceis, testaceis et zoophytis ... Bononiae, Typis Jo. Baptistae Ferronii, sumptibus Marci
Antonii Berniae. 593 pp. [Pennatulacean bioluminescence. ]

. 1648. Musaeum metallicum in libros III distributum. Bartholomaeus Ambrosinus . .. composuit. . .
Marcus Antonius Bernia . . . in lucem edidit . .. Typis Jo. Baptistae Ferronii, Bononiae. 979 pp. [Pennatu-
lacean bioluminescence. ]

ALLEN, G. R. AND R. STEENE. 1994. Indo-Pacific coral reef field guide. Tropical Reef Research, Singapore. 378
pp. [Includes color photographs of Preroeides spp. and Virgularia spp.]

ALLEN, R. K. 1969. Common intertidal invertebrates of southern California. Peek publications, Palo Alto.
(Anthozoa, pp. 30-32). [Renilla kollikeri.]

ALONSO, C. 1979. Estudio morfologico y biométrico de las espiculas de Kollikeri K6lliker, 1872 (Anthozoa,
Pennatulacea). Revista brasileira de Biologia. 39(4):827—834. [Skeleton; sclerite morphology. ]

WILLIAMS: INDEX PENNATULACEA

nN
Nn

ALTUNA-PRADOS, A. 1994. Observaciones biogeograficas sobre los cnidarios bentonicos de la costa Vasca.
Kobie Ciencias Naturales 22:41—S7.

ANCTIL, M. 1987. Bioactivity of FMRFamide and related peptides on a contractile system of the coelenterate
Renilla kollikeri. Journal of Comparative Physiology, Part B: Biochemical, Systemic and Environmental
Physiology 157(1):31-38.

. 1989a. Modulation of a rhythmic activity by serotonin via cyclic AMP in the coelenterate Renilla

kollikeri. Journal of Comparative Physiology, Part B: Biochemical, Systemic and Environmental Physiol-

ogy 159(4):491—S00. [Renilla kollikeri, muscles, rachidial rhythmic contractions, modulatory mechanisms. |

. 1989b. The antiquity of monoaminergic neurotransmitters: evidence from Cnidaria. NATO ASI
(Advanced Science Institutes) Series A Life Sciences 188:141—155.

ANCTIL, M., D. BOULAY, AND L. LARIVIERE. 1982. Monoaminergic mechanisms associated with control of
luminescence and contractile activities in the coelenterate, Renilla kollikeri. Journal of Experimental
Zoology 223(1):11—24. [Nervous transmitter substances; monoaminergic drugs, effects on luminescence
and rachidial contraction. ]

ANCTIL, M., G. GERMAIN, AND L. LARIVIERE. 1984. Catecholamines in the coelenterate Renilla koellikeri. Uptake
and radioautographic localization. Cell and Tissue Research 238(1):69—80. [Nervous transmitter sub-
stances. |

ANCTIL, M., A. K. PANI, AND M. A. ALI. 1991. Modulation of rhythmic contractions by meltonin via cyclic GMP
in the coelenterate Renilla koellikeri. Journal of Comparative Physiology, Part B: Biochemical, Systemic
and Environmental Physiology 161(6):569—S75.

ANDERSON, J. M., H. CHARBONNEAU, AND M. J. CORMIER. 1974. Mechanisms of calcium induction of Renilla
bioluminescence. Involvemnent of a calcium-triggered luciferin binding protein. Biochemistry (American
Chemical Society) 13(6):1195—1200.

ANDERSON, J. M. AND M. J. CORMIER. 1978. Sodium gradient dependent calcium transport in Reni//a lumisomes.
Biochemical and Biophysical Research Communications 81(1):114—121. [Renilla reniformis; luminous
organs.

ANDERSON, J. M., K. HORI, AND M. J. CORMIER. 1978. A bioluminescence assay for PAP (3,5 -diphosphoade-
nosine) and PAPS (3 ’-phosphoadeny! sulfate). Methods in Enzymology 57:244—257. [Renilla reniformis;
luminescence assay. |

ANDERSON, P. A. V. 1976. An electrophysiological analysis of behavioural integration in colonial anthozoans.
Pp. 609-618 in Coelenterate Ecological Behavior, G.O. Mackie, ed. Plenum, New York. 744 pp. [Renilla
spp., Renilla kéllikeri.|

ANDERSON, P. A. V. AND J. F. CASE. 1975. Electrical activity associated with luminescence and other colonial
behavior in the pennatulid Renilla kollikeri. Biological Bulletin (Woods Hole, Mass. Marine Biological
Laboratory) 149(1):80—95.

ANDERSON, P. K. 1989 (1988). Deliberate foraging on macroinvertebrates by dugongs. National Geographic
Research 5(1):4—6. [Virgularia, predators, Dugong dugon, probable record, W. Australia. ]

ANDREE, K. 1912. Eine zweite Graphularia - Art (Gr. Crecelii n. sp.) aus dem mitteloligocanen Meeressand im
Mainzer Becken. Centralblatt fiir Mineralogie, Geologie und Paldontologie, Stuttgart 1912:202—207.
[Extinct sea pen Graphularia crecelii from the Oligocene of Germany. ]

ANGELIS, G. DE. 1895? I Corallari dei terreniterziari dell’Italia settentrionale. Collezione Michelotti. Museo
Geologico della R. Universita di Roma. Memorie Reale Academia dei Lincei, Roma S(1):164—280. [Extinct
sea pen Pavonaria portisi from the Tertiary of Italy, Pavonaria as used here is a synonym of Halipteris,
see Williams, 1995a:122.]

ANONYMOUS. 1898. A preliminary catalogue of the collections of natural history and ethnology in the Provincial
Museum, Victoria, British Columbia. The Government of the Province of British Columbia, Victoria,
British Columbia. Pp. 107-109. [Listed by family; Prilosarcus gurneyi, Verrillia blakei.]

ARCULEO, M., M. P. SPARIA, G. D’ANNA, C. PIPITONE, AND S. RIGGIO. 1990. Dati sulla pesca a strascico nel
Gofo di Gela (Sicilia sud orentale). Naturalista siciliano 14 (3—4):39—5S. [Pennatula phosphorea. |

ARNDT, W. 1912. Notiz tiber Virgularia mirabilis. Zoologischer Anzeiger 40(2/3):93—94. [Virgularia mirabilis. ]

ARTHUR, W. 1997. The origin of animal body plans—a study in evolutionary developmental biology. Cambridge
University Press, Cambridge. 338 pp. [Pp. 66-69, Arthur mentions the resemblance of the Vendian fossil
taxon Charniodiscus to modern sea pens. |

56 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 2

ASBJORNSEN, P. C. 1856. Beskrivelse over Kophobelemnon miilleri, en ny Sofjaerslaegt. /n Fauna Littoralis, M.
Sars, J. Koren, and D. C. Danielssen, eds. Norvegiae 2:81—85.

ATIYA, F. S. 1994. The Red Sea in Egypt. Part 2, Invertebrates. Farid F. S. Atiya, Cairo. 297 pp. [Color
photograph of specimen identified as Scytaliopsis ghardagensis. |

ATKINSON, R. J. A. 1989. Baseline survey of the burrowing megafauna of Loch Sween PMNR and an
investigation of the effects of trawling on the benthic megafauna. Nature Conservancy Council, Report No.
909:1—S9. [Virgularia mirabilis: distribution and trawling effect on density in the northeastern Atlantic. ]

AWAD, E. W. AND M. ANCTIL. 1993a. Identification of (beta)-like adrenoceptors associated with biolumines-
cence in the sea pansy Renilla koellikeri. Journal of Experimental Biology 177:181—200.

. 1993b. Positive coupling of (beta)-like adrenergic receptors with adenylate cyclase in the cnidarian
Renilla koellikeri. Journal of Experimental Biology 182:131—146.

BALss, H. 1909. Uber Pennatuliden des Miinchener Museums. Zoologischer Anzeiger 34(113/14):423—431.
[Lituaria habereri, Prochunella hertwigi, Pteroeides dofleini, Stachyptilum dofleini.]

. 1910. Japanische Pennatuliden. /n Beitrage zur Naturgeschichte Ostasiens. Abhandlungen der Mathe-

matisch-Physischen Classe der Kéniglich Sachsischen Gesellschaft der Wissenschaften, F. Doflein, ed.

1(10, suppl.): 1-106. [Taxonomic descriptions: Balticina willemoesi, Calibelemnon indicum, Calibelemnon

hertwigi, Cavernularia elegans, Cavernularia habereri, Cavernularia marquesarum, Cavernularia obesa,

Echinoptilum macintoshi, Funiculina quadrangularis, Kophobelemnon stelliferum, Lituaria habereri,

Lituaria phalloides, Pennatula fimbriata, Pennatula murrayi, Pennatula naresi, Pennatula phosphorea,

Pteroeides chinense, Pteroeides dofleini, Pteroeides lacazii, Pteroeides sagamiense, Sclerobelemnon

burgeri, Sclerobelemnon schmeltzii, Scytalium splendens, Stachyptilum superbum, Virgularia abies, Vir-

gularia gustaviana, Virgularia reinwardtii. |

. 1911. Notiz tiber einige Pennatuliden des zoologischen Museums der Kaiserlichen Annuaire du Musee
zoologique de I’ Akadémie impériale des sciences de St. Pétersbourg 16:158—159. [Virgularia mirabilis,
Virgularia gustaviana var. magnifolia.|

BALUK, W. AND A. PISERA. 1984. A new species of sea pen, Graphularia transaedina sp. n. from the Korytnica
Clays (Middle Miocene; Holy Cross Mountains, central Poland). Acta Geologica Polonica 34
(3—4):203-211.

BARATTINI, L. P. AND Y. E. H. URETA. 1960. La fauna de las costas Uruguayas del este (Invertebrados). Museo
Damaso Antonio Larrafaga, Publicaciones de Divulgacion Cientifica, Montevideo, 195 pp. [Pennatu-
laceans, pp. 30-31; Renilla danae, Virgularia patachonica.]

BARNES, R. D. 1987. Invertebrate Zoology, Sth ed. Saunders College Publishing, Philadelphia. 893 pp. [Octo-
corals, pp. 134-139; Renilla, Umbellula.]

BARNES, R. S. K., P. CALOW, AND P. J. W. OLIVE. 1988. The invertebrates: a new synthesis. Blackwell Scientific
Publications, Palo Alto. 582 pp. [Classification of anthozoans, p. 59.]

BARREIRA Y CASTRO, C. 1990. Revisao taxonomica dos Octocorallia (Cnidaria, Anthozoa) do litoral Sul-Ameri-
cano: da foz do Rio Amanona a foz do Rio da Prata. Ph. D. dissertation, Sao Paulo. 343 pp. [Brazilian sea
pens of the family Virgulariidae. ]

BARRELIER, J. 1714. Plantae per Galliam. Hispaniam et Italiam observatae, iconibus aeneus exhibitae a R. P.
Jacobo Barreliero Parisino ... opus posthumum. Accurante Antonio de Jussieu in lucem editum, & ad
recentiorum normen digestum. Cui accessit ejusdem auctoris specimen de insectis quibusdam marinis,
mollibus, crustaceis is & testaceis. Apud S. Ganeau, Parisiis [Paris]. 140 pp.

BASTER, J. 1759-1765. Natuurkundige Uitspanningen, behelzende eenige waarneemingen, over sommige
Zee-Planten en Zee-Insecten, benevens derzelver Zaadhuisjes en Eijernesten. J. Bosch, Te Haarlem, 2 vols.

BATIE, R. R. 1972. Investigations concerning the taxonomic status of the sea pen Prilosarcus gurneyi (Cnidaria:
Pennatulacea). Northwest Science 46(4):290—300.

BAUHIN, C. 1620. Prodromos theatri botanici, Caspari Bauhini Basileens; in quo plantae supra sexcentae ab ipso
primum descriptae cum plurimis figuris proponuntur. Francofurti ad Monum[Frankfurt am Main]: Typis
Pauli Iacobi, impensis Ioannis Treudelii. 160 pp. [Pennatulacean bioluminescence. ]

————. 1671. Pinax theatri botanici; sive, Index in Theophrasti Dioscoridis et botanicorum qui a seculo
scripserunt opera, plantarum circiter sex millium ab ipsis exhibitarum nomina cum earundem synoymiis &
differentiis methodice secundum genera & species proponens. Opus XL. annorum summopere expetitum
ad autoris autographum recensitum. Basileae [Basel], Impensis Johannis Regis. 518 pp. [Pennatulacean
bioluminescence. ]

WILLIAMS: INDEX PENNATULACEA 57

. 1650-51. Historia plantarum universalis, nova et absoutissima, cum consensu et dissensu circa eas,
auctoribus Ioh. Bauhino ... et loh. Henr. Cherlero ... quam recensuit et auxit Dominicus Charaeus .. .
Juris vero publici fecti Franciscus Lud. Graffenried ... Ebroduni, 3 vols. [Pennatulacaean biolumines-
cence. |

BAYER, F. M. 1955a. Remarkably preserved fossil sea-pens and their Recent counterparts. Journal of the
Washington Academy of Sciences 45(9):294—300. [Virgularia presbytes, Pteroeides argenteum. |

. 1955b. Contributions to the nomenclature, systematics, and morphology of the Octocorallia. Proceed-

ings of the United States National Museum 105 (3357):207—220. [Affinity between the extinct genus

Graphularia and the recent genus Sty/atula. |

. 1956. Octocorallia. Pp. 166-231 in Treatise on invertebrate paleontology, Part F: Coelenterata, R. C.

Moore, ed. Geological Society of America and the University of Kansas Press, Lawrence.

. 1957. Additional records of Western Atlantic octocorals. Journal of the Washington Academy of

Sciences 47(11):379-390. [Protoptilum thomsonii, Virgularia presbytes, Stvlatula antillarum, Stvlatula

elegans, Acanthoptilum agassizii, Acanthoptilum oligacis.]

. 1959. Octocorals from Surinam and the adjacent coasts of South America. Natuurwetenschappelijke

Studiekring voor Suriname en de Nederlandse Antillen, No. 20:1—43. [Renilla reniformis forma americana,

Kollikeri, Sclerobelemnon theseus, Stylatula cf. brasiliensis, Stvlatula diadema, Virgularia presbyvtes,

Virgularia kophameli.|

. 1961. The shallow-water Octocorallia of the West Indian region. A manual for marine biologists. Studies

on the fauna of Curacao and other Caribbean Islands 12:1—373. [Renilla reniformis, Renilla miilleri,

Sclerobelemnon theseus, Virgularia presbytes, Stvlatula diadema. |

. 1973. Colonial organization in octocorals. Pp. 69-93 in Animal colonies—development and function

through time, Boardman, Cheetham, and Oliver, eds. Sinauer, Sunderland, MA. 603 pp.

. 1981a. Status of knowledge of octocorals of world seas. Seminarios de Biologia Marinha, Academia

Brasileira de Ciencias Rio de Janeiro 1981:3—102.

. 1981b. Key to the genera of Octocorallia exclusive of Pennatulacea (Coelenterata: Anthozoa), with

diagnoses of new taxa. Proceedings of the Biological Society of Washington 94(3):902—947.

. 1996. Bibliography of Octocorallia from Pliny to the present. Smithsonian Institution. [WEB SITE;
http://nmnhgoph.si.edu/gopher-menus/Octocorals.html. ]

BAYER, F. M. AND M. GRASSHOFF. 1997. Umbellula Cuvier, [1797] (Cnidaria, Anthozoa): proposed conservation
as the correct original spelling, and corrections to the entries relating to Umbellularia Lamarck, 1801 on
the Official Lists and Indexes of Names in Zoology. Bulletin of Zoological Nomenclature 54(1):14—18.
[Use of Umbellula supercedes that of Ombellula.]

BAYER, F. M., M. GRASSHOFF, AND J. VERSEVELDT. 1983. Illustrated trilingual glossary of morphological and
anatomical terms applied to Octocorallia. E. J. Brill/Dr. Backhuys, Leiden. 75 pp.

BAYER, F. M. AND H. B. OwRE. 1968. The free-living lower invertebrates. The Macmillan Company, London.
[Pennatulaceans, pp. 81—83, 93; Pennatula.|

BEKLEMISHEV, W. N. 1969a. Principles of comparative anatomy of invertebrates, Vol. 1: Promorphology. The
University of Chicago Press. 490 pp. [Pennatulaceans, pp. 35-36, 100-102, 486490; Funiculina quad-
rangularis, Pennatula phosphorea, Pteroeides griseus. |

. 1969b. Principles of comparative anatomy of invertebrates, Vol. 2: Organology. The University of
Chicago Press. 529 pp. [Pennatulaceans, pp. 42, 197, 253, 264; Funiculina, Veretillum.]

BELCIK, F. P. 1977. A distribution study of the Octocorallia of Oregon. Publications of the Seto Marine Biological
Laboratory 24(1/3):49-52. [Kophobelemnon hispidum, Kophobelemnon sp., Anthoptilum grandiflorum,
Anthoptilum sp., Funiculina armata, Helicoptilum rigidum, Scleroptilum sp., Umbellula spp., Virgularia
sp., Stvlatula elongata, Balticina pacifica, Pennatula sp., Pennatula phosphorea, Ptilosarcus gurneyi.|

BELEM, M. J. DA COSTA AND L. C. DE FIGUEIREDO ALVARENGA. 1973. Contribuigao ao conhecimento da fauna
de Cnidarios dos Estados da Guanabara e do Rio de Janeiro, Brasil. I. Virgularia presbytes Bayer, 1955
(Anthozoa, Pennatulacea, Virgulariidae). Actas da Sociedade de Biologia do Rio de Janeiro 17(1):41—S1.
[Brazilian Virgulariidae; Virgularia presbytes; redescription; southwest Atlantic Ocean; Brazil, Guanabara
Bay, first record. |

BELLINI, R. 1905. Le varie Facies del Miocene medio nelle Colline de Torino. Bolletino della Societa geologica
italiana 24:607—653. [Pavonaria portisi from the Middle Miocene of Italy. ]

BELLOMY, M. 1974. The curious sea pansy. Marine Aquarist 5(3):14—20.

58 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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BELYAEV, G. M. 1972. Hadal bottom fauna of the world ocean. Akademiya Nauk SSSR Institut Okeanologil.
Published for the Smithsonian Institution and the National Science Foundation, Washington, D. C. by the
Israel Program for Scientific Translations, Jerusalem. [Translated into English from the Russian; pp. 48-49,
Umbellula sp. juv., Umbellula sp., bathymetric records, 6010-6730 m, Peru-Chile Trench and Kermadec
Trench. |

BENHAM, W.B. 1906. On a new species of Sarcophyilum from New Zealand. Zoologisher Anzeiger
31(2/3):66—-67. [Sarcophyllum bollonsi.|

. 1907. On a new species of pennatulid (Sarcophyllum bollonsi). Transactions and Proceedings of the
New Zealand Institute 39:193—195. [Sarcophyllum bollonsi.|

BERENGUIER, A. 1954. Contribution a l’étude des octocoralliaires de Méditerranée Occidentale. Recueil des
travaux de la Station Marine d’Endoume 12:53—96. [Pennatula rubra, Pennatula phosphorea, Pteroeides
griseum. |

BerG, S. E. 1941. Die Entwicklung und Koloniebildung bei Funiculina quadrangularis (Pallas). Zoologiska
bidrag fran Uppsala 20:1—100. [Funiculina quadrangularis, Kophobelemnon stelliferum, Pennatula phos-
phorea, Virgularia mirabilis, Virgularia cladiscus.]

BERGER, J. 1915. Zur Kenntnis des feineren Baues von Stv/atula. Inaugural-Dissertation zur Erlangung der
Doktorwiirde der Hohen Philosophischen Fakultét der Schlesischen Friedrich-Wilhems-Universitat zu
Breslau. Druck von Wilh. Gottl. Korn, Breslau. 72 pp.

BERGSTROM, J. 1989. The origin of animal phyla and the new phylum Procoelomata. Lethaia 22(3):259-269.

. 1991. Metazoan evolution around the Precambrian-Cambrian transition. Pp. 25-34 in The early
evolution of Metazoa and the significance of problematic taxa, A. M. Simonetta and S. Conway Morris,
eds. Proceedings of an International Symposium held at the University of Camerino 27-31 March 1989.
Cambridge University Press, Cambridge. 296 pp. [Charniodiscus, supposed pennatulaceans of the Precam-
brian Vendian Period).

BERNHEIMER, A. W. AND L. S. AVIGAD. 1981. New cytolysins in sea anemones from the west coast of the United
States. Toxicon 19(4):529-534. [Renilla koellikeri, toxins and venoms; cytolysin, first recorded analysis;
California. }

BERTSCH, H. 1968. Effect of feeding by Armina californica on the bioluminescence of Renilla koellikeri. Veliger
10:440-441. [Predator/prey interaction: nudibranch mollusk and sea pansy. ]

. 1982. Estudios de ecosistemas bentonicos a lo largo de la costa noroccidental de Baja California,
Mexico: distribution y presa de varios invertebrados marinos. Ciencias marinas 8(2):91—123. [In Spanish
with English summary; Stylatula elongata; northern Pacific; Baja California; distribution and range
extensions. |

Best, B. A. 1988. Passive suspension feeding in a sea pen: effects of ambient flow on volume flow rate and
filtering efficiency. Biological Bulletin (Woods Hole, Mass. Marine Biological Laboratory) 75(3):332-342.
[Ptilosarcus gurneyi, filter feeding, filtering speed and efficiency, effect of water flow rate and size, north
Pacific. }

BILHAUT, A. 1975a. Etude de la bioluminescence chez l’octocoralliaire Veretillum cynomorium Pall. |. - Les
réponses lumineuses des autozoides isolés de la colonie. Archives de zoologie expérimentale et generale.
116(1):27-42.

————. 1975b. Etude de la bioluminescence chez |’ octocoralliaire Veretillum cynomorium Pall. 2. Les réponses
lumineuses de la colonie. Archives de zoologie expérimentale et générale 1 16(3):321—341.

BILHAUT, A. AND M. PAVANS DE CECCATTY. 1971a. Les récepteurs sensoriels de |’octocoralliaire Veretillum
cynomorium Pall. Comptes rendus hebdomadaires des séances de |’Academie des sciences, Paris
272D:3150-3153.

———. 1971b. Les différenciations cellulairs de type choanocytaire chez l’octocoralliaire Veretillum cyno-
morium Pall. Comptes rendus hebdomadaires des séances de |’Academie des sciences, Paris
272D:3053—3056.

BIRKELAND, C. 1974. Interactions between a sea pen and seven of its predators. Ecological Monographs
44(2):211—232. [Predators on the sea pen Ptilosarcus gurneyi, including four asteroid species: Hippasteria
spinosa, Mediaster aequalis, Crossaster papposus, and Dermasterias imbricata; and three nudibranch
gastropods: Hermissenda crassicornis, Armina californica, and Tritonia festiva; also includes an underwa-
ter photograph of a dense bed of Ptilosarcus gurneyi on p. 213.]

WILLIAMS: INDEX PENNATULACEA 59

BIRKELAND, C. E. 1969. Consequences of differing reproductive and feeding strategies for the dynamics and
structure of an association based on the single prey species, Ptilosarcus gurneyi (Gray). Ph.D. dissertation,
University of Washington, Seattle. 99 pp.

. 1971. Consequences of differing reproductive and feeding strategies for the dynamics and structure of
an association based on the single prey species, Ptilosarcus gurneyi (Gray). Dissertation Abstracts
International 31B:5925—5926.

BLAINVILLE, H. M. D. DE. 1834. Manuel d’actinologie, ou de zoophytologie. F. G. Levrault, Strasbourg. 644 pp.
[Bioluminescence in sea pens. ]

BLAKE, J. 1872. On the supposed new marine animal from Barraud’s Inlet. Nature 7:67—68. [Axis of Halipteris
willemoesi.]

. 1873. On the structure of Verrillia blakei. Proceedings of the California Academy of Sciences
5:149-150.

BLEEKER, P. 1859. Over eenige nieuwe soorten van Zeeveders of Pennatulina (polypi natantes) van den Indische
Archipel. Natuurkundig tijdschrift voor Nederlandsch-Indie 20:399-404. [Summary in Latin under the title
“Sur des especes nouvelles de Pennatulina (polypi natantes) de l’Archipe indien,” in Revue et magasin de
zoologie pure et appliquée (2)14(1):38—42; 1862; original descriptions of several species of Preroeides.|

BLUMENBACH, J. F. 1779. Handbuch der Naturgeschichte. H. Dieterich, Gottingen. 743 pp. [Umbellula as
Encrinus.]|

Boccone, P. 1670. Recherches et observations d’histoire naturelle touchant le Corail. Paris.

————. 1674a. Recherches et observations curieuses sur la nature du corail blanc & rouge vray de Dioscoride;
et sur la sangsue qui se trouve attachée au posson xiphias, avec son anatomie & autres choses fort rare,
proposées & examinées a diverses fois dans 1’ Academie de M. |’abbe Bourdelot. Claudi Barbin, Paris. 69
pp.

. 1674b. Recherches et observations naturelles de Monsieur Boccone Bentilhomme Sicilien; touchant le
Corail, la Pierre Etoilée, les Pierres de figure de Coquilles, la Corne d’ Ammon, |’ Astroite Undulatus, les
Dents de Poissons petrifiees, les Herissons alterez, |’Embrasement du Mont Etna, la Sangsue du Xiphias.
l’Alcyonium stupposum, le Bezoar mineral, & les Plantes qu’on trouve dans la Sicili, avec quelques
Reflexions sur la Vegetation des Plantes. Examinées a diverses fois dans |’Assemblée de Messieurs de
Society Royale de Londres, & Conferences dans les de Monsieur |’ Abbe Bourdelot a paris. Jean Jansson
a Waesberge, Amsterdam. 328 pp.

BODDAERT, P. 1771. Brief aan den Schryver der Bedenkingen, betreffende den dierlyken Oorsprong der
Koraalgewassen. A. van Paddenburgh, Utrecht. 57 pp.

BOERHAAVE, H. 1720. Index alter plantarum quae in Horto Academico Lugduno-Batavo aluntur conscriptus ab
Hermanno Boerhaave . . . 2 vols. Lugduni Batavorum: Sumptibus auctoris.

. 1727. Historia plantarum quae in Hort Academico Lugduni-Batavorum crescunt cum earum charac-
teribus, & medicinalibus virtutibus. 2 vols.( 408 pp.; 409-698). Romae: apud Franciscum Gonzagam.
BOESEMAN, M. 1973. The autobiography of Pieter Bleeker [With additional notes and remarks]. Collected fish

papers of Pieter Bleeker |:1—71. [Biographical material on Pieter Bleeker. ]

BOHADSCH, J. B. 1761. De quibusdam animalibus marinis, eorumque proprietabibus, orbi litterario vel nondum
vel minus notis, liber cum nonnullis tabulis aeri incisis, ab auctore super vivis animalibus delineatis.
Dresdae: apud G.C. Walther. [This post-Linnean work was suppressed by the ICZN for nomenclatural
purposes. |

BOONE, L. 1933. Scientific results of the cruise of the yachts “Eagle” and “Ara,” 1921—1928. Coelenterata,
Echinodermata and Mollusca. Bulletin of the Vanderbilt Marine Museum, Huntington 4:1—217. [Prilosar-
cus gurneyi, Pavonaria californica. |

. 1938. The marine algae, Coelenterata .. . of the world cruises of the yachts “Ara,” 1928-1929, and
“Alva,” 1931-1932, “Alva” Mediterranean Cruise, 1933, and “Alva” South American Cruise, 1935,
William K. Vanderbilt, Commanding. Bulletin of the Vanderbilt Marine Museum, Huntington 7:27—76.
[Veretillum vanderbilt of the Philippine Islands. ]

BorLASE, W. 1758. The natural history of Cornwall. Printed for the author by W. Jackson, Oxford. 326 pp.

BourNE, G. C. 1900. The Anthozoa. /n A treatise on zoology, Part II: The Porifera and Coelenterata, E. R.
Lankester, ed. Adam and Charles Black, London. [Pennatulaceans, pp. 30—34.]

60 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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BoussueT, F. 1558. De natura aquatilium carmen, in universam Gulielmi Rondeletii ...quam de piscibus
marinis scripsit historiam: cum vivis eorum imaginibus, opusculum nunc primum in luce emissum. Apud
Mattiam Bonhomme, Lugduni, 2 vols. [Pennatulacean bioluminescence. ]

BRADLEY, J. 1980. Scolicia and Phycodes trace fossils of Renilla (Pennatulacea). Pacific Geology 14:73—86.

. 1981. Radionereites, Chondrites and Phycodes; trace fossils of anthoptiloid sea pens. Pacific Geology
15:1—16.

BRAFIELD, A. E. 1969. Water movements in the pennatulid coelenterate Preroides griseum. Journal of Zoology
158:317-325.

BRAFIELD, A. E. AND G. CHAPMAN. 1967. The respiration of Prteroides griseum (Bohadsch) a pennatulid
coelenterate. Journal of Experimental Biology 46:97—104.

BRANCH, G. M., C. L. GRIFFITHS, M. L. BRANCH, AND L. E. BECKLEY. 1994. Two oceans—a guide to the marine
life of southern Africa. David Philip, Cape Town. 360 pp. [P. 23, color photographs of Virgularia schultzei,
as well as the soft coral Eleutherobia aurea mislabeled as Actinoptilulm molle.]

BRANCH, M. L. AND G. C. WILLIAMS. 1993. The Hydrozoa, Octocorallia and Scleractinia of subantarctic Marion
and Prince Edward Island: illustrated keys to the species and results of the 1982—1989 University of Cape
Town surveys. South African Journal of Antarctic Research 23(1, 2):3-24. [Illustrated key, taxonomic
description, depth distribution: Halipteris sp., Pennatula inflata, Pennatula phosphorea. |

BRANCO, W. 1885. Ueber einige neue Arten von Graphularia und tiber Tertiare Belemniten. Zeitschrift der
Deutchen Geologischen Gessellschaft 37:422—432. [The extinct sea pen Graphularia ambigua.]

BRIEN, P. 1947. Guide des Travaux Pratiques de Zoologie. Paris. 218 pp. [Coelenterates, pp. 18—28; Veretillum
cynomorium. |

BricGs, E. A. 1915. Report on the Alcyonarians obtained by F. I. S. “Endeavour” on the Eastern and Southern
Coasts of Australia. Part I. Trade and Customs (Fisheries) Australia. Biological Results of the Fishing
Experiments carried on by the F. 1. S. “Endeavour” 1909-14, 3(2):59-94. [Kophobelemnon schmeltzii,
Godeffroyia elegans, Sarcophyllum grande. |

BrocH, H. 1910a. Anthozoa (Nachtrag): Pennatuliden. /n Schultze, L. Zoologische und anthropologische
Ergebnisse einer Forschungsreise im westlichen und zentralen Siidafrika, 1903-1905. 4. Systematik und
Tiergeographie. XIII. Denkschriften der Medicinisch-naturwissenschaftlichen Gesellschaft zu Jena
16:229-235.

. 1910b. Diagnosen von neuen oder weniger bekannten Pennatulidae. Zoologischer Anzeiger 36:60—65.

[Leioptilum solidum - locality unspecified, Pteroeides lusitanicum - Portugal, Pteroeides heteroradiatum -

locality unspecified, Pteroeides jungerseni - Matupi, Struthiopteron, Virgularia halisceptrum. |

. 1910c. Pennatulida. Jn Die Fauna Sudwest-Australiens, W. Michaelsen and R. Hartmeyer, eds. Er-

gebnisse der Hamburger sudwest-australischen Forschungsreise 1905, 3(2):109-122. Verlag von Gustav

Fischer, Jena. [Sarcophyllum roseum - Australia. ]

. 1913a. Pennatulacea. Report on the Scientific Results of the ‘Michael Sars’ North Atlantic Deep Sea

Expedition 1910 3(1):3—-8.

. 1913b. Die Alcyonarien des Trondhjemsfjordes, III: Pennatulacea; IV. Biogeographische Ubersicht.

Norske Videnskabers Selskab, Trondheim. Skrifter 1912(10):1-S9. [Funiculina quadrangularis, Ko-

phobelemnon stelliferum, Pavonaria christii, Pavonaria finmarchica, Pennatula grandis, Pennatula phos-

phorea, Pennatula aculeata, Stylatula elegans, Virgularia mirabilis.)

. 1913c. Arktiske Alcyonarier 1 Tromso museum. Tromso museums aarshefter 34:179—186.

——. 1914a. Scyphomedusae, Pennatulacea and Hydroida from the “Michael Sars” North Atlantic Deep Sea
Expedition 1910. Report on the Scientific Results of the “Michael Sars” North Atlantic Deep Sea
Expedition, 1910 3(1) papers 4-6: 1—24 + 1-9 + 1-18. [Umbellula giintheri.]

. 1914b. Pennatulacea. Pp. 51—56 in Beitrage zur Kenntnis der Meerefauna Westafrikas, W. Michaelson,
ed. L. Friederichsen and Co., Hamburg.

———.1927. The Folden Fiord. Coelenterata, II: Corals. Troms@ museums skrifter 1(12):7—8. [Kophobelemnon
stelliferum, Virgularia cladiscus.|

= . 1935. Oktokorallen des nérdlichsten pazifischen Ozeans und ihre Beziehungen zur atlantischen Fauna.
Avhandlinger utgitt av det Norske videnskaps-akademi I Oslo 1935(1):1—S3. [Pavonaria finmarachica,
Pavonaria willemoesi? sp. juv.]

——. 1939. Some South African shallow water octactinians. Kungliga fysiografiska sallskapets | Lund
forhandlingar 9(6):1—32.

WILLIAMS: INDEX PENNATULACEA 61

——. 1940. Some South African shallow water Octactinians. Kungliga fysiografiska sallskapets | Lund
forhandlingar 9:48—79. [Actinoptilum molle, Virgularia schultzei. |

. 1953. Octocorals and stony corals of the high Adriatic trawling grounds. Split, Yugoslavia. Institut za

Oceanogrfiju i Ribarstvo. Ribarstveno-Bioloska Ekspedicija M/B “hvar,” 1948-1949. Izvjesca-Reports

6(2):1—22. [Pennatula phosphorea, Pteroeides griseum, Veretillum cynomorium.|

. 1955. Octocorallia from the Northern Ocean collected by the Soviet expeditions in the years 1929-1935.

Travaux de |’ Institut zoologique de |’ Academie des sciences de l' URSS = Trudy Zoologicheskogo instituta

18:26-35. [In Russian - Umbellula encrinus, Virgularia glacialis, Virgularia tuberculata.|

. 1956. Oktokorallen russischer Expeditionen em Polarmeer wahrend der Jahre 1929-1935. Avhandlin-

gar utgitt av det Norske videnskaps-akademi i Oslo...1. Matematisk-natur videnskapelig klasse

1956(4):1—18. [Umbellula encrinus, Virgularia glacialis, Virgularia tuberculata.]|

. 1957. Pennatularians (Umbellula). Reports of the Swedish Deep Sea Expedition, 1947-48

2(21):348-364. [Umbellula guntheri, U. carpenteri, U. durissima.]

. 1958a. The pennatularian genus Umbellula. Nature, London 182(4630):267-268. [Umbellula duris-

sima, U. thomsoni, U. huxleyi, U. spicata, U. pellucida, U. lindahli.|

. 1958b. Octocorals, Part I: Pennatularians. Discovery Reports 29:245—280. [Actinoptilum molle, Cav-

ernularia elegans, Pennatula rubra, Pteroeides griseum, Renilla miilleri, Veretillum cynomorium. |

. 1961. Umbellula encrinus pavist i heyantarktiske farvann. Fauna (Norsk Zoologisk Forenings
Tidsskrift, Oslo) 14(2):33-37.

Brusca, R. C. 1980. Common intertidal invertebrates of the Gulf of California, 2nd ed. University of Arizona
Press, Tucson. [Cnidaria, pp. 48-66; Ptilosarcus sp., Stylatula sp.|

Brusca, R. C. AND G. J. BRUSCA. 1990. Invertebrates. Sinauer Associates Publishers, Sunderland, Mass. 922
pp. [Anthozoa, pp. 214-261; Renilla, Ptilosarcus, Pennatula.]

BUCHANAN, J. B. 1955. The Pennatulacea of the Gold Coast. Revue de zoologie et de botanique africaines
51(1—2):388-390. [Cavernularia elegans, Veretillum cynomorium, Virgularia rumphii.|

Buck, J. 1973. Bioluminescent behavior in Renilla, 1: Colonial responses. Biological Bulletin (Woods Hole,
Mass. Marine Biological Laboratory), Boston 144(1):19-42.

Buck, J. AND F. E. HANSON. 1967. Zooid response in Renilla. Biological Bulletin (Woods Hole, Mass. Marine
Biological Laboratory), Boston 133:459.

BUISSON, B. 1964. Données physilogiques sur l’intégration et la polarité dans les colonies de Veretillum
cynomorium Pall. (Cnidaire, Pennatulidae). Comptes rendus hebdomadaires des seances de |’ Academie des
sciences, Paris (Série D) Sciences Naturelles 259:3361—3362.

. 1969. Aspects de la régénération chez Veretillum cynomorium (Cnidaria, Pennatulacea). Le Naturaliste

Canadien, Québec 96:785—797.

. 1970. Les supportes morphologiques de I’intégration dans la colonie de Veretillum cynomorium Pall.

(Cnidaria, Pennatularia). Zeitschrift fiir Morphologie der Tiere 68:1—36.

. 1971a. Les activités rhythmiques comportementales de la colonie de Veretillum cynomorium (Cnidaire,

Pennatulidae). Cahiers de Biologie Marine 12:1 1-48.

. 1971b. Activités électriques spontanées et réactions motrices au cours de rythmes comportmentaux de

la colonie Veretillum cynomorium Pall. (Cnidaire Pennatulidae). Vie et Milieu (Serie A) Biologie marine

22(2):327-372. [In French with English and German summaries; circadian functions; contraction expansion
rhythm, control and stimulation effects. |

. 1973. Données sur le comportement rythmique du polype isolé de la colonie de Veretillum cynomorium

Pall. (Cnidaria Pennatularia). Comptes rendus hebdomadaires des séances de |’ Academie des sciences, Paris

(Série D) Sciences Naturelles 277(15):541—1544.

. 1974. The behaviour and the nervous system of Veretillum cynomorium Pall. (Cnidaria Pennatularia).

Zoologischer Anzeiger 192(3—4):165—174.

. 1976. Note on the behavoural physiology of Pteroides griseum (Bohadsch) (Cnidaria Pennatularia).

Zoologische Jahrbiicher, Abteilung fiir Allgemeine Zoologie und fysiologie der Tiere 80(3):223-230.

. 1979. Sur l’origine des pulsations du polype isolé de la colonie de Veretillum cynomorium (Pall.)

(cnidaire anthozoaire). Comptes rendus hebdomadaires Seances Academie des Sciences, Paris (Serie D)

Sciences Naturelles 288(10):891—894. [Polyp; isolated, contraction generation, influencing factors. ]

. 1980. Data on the rhythmic behaviour of a young sea-pen (Veretillum cynomorium). Zoologischer

Anzeiger 205(1—2):20-26.

62 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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. 1988. Sur le rhythme d’expansion-contraction circadien du cnidaire octocoralliaire Veretillum cyno-
morium Pall: la recherche de photorecepteurs. Bulletin de la Société Zoologique de France 113(3):279-283.

BUISSON, B. AND S. FRANC. 1969. La structure et lultrastucture des cellules mesenchymateuses et nerveuses
intramesogléennes de Veretillum cynomorium Pall. (Cnidaire, Pennatulidae). Vie et Milieu 20(2-A):
279-291.

Busor, P. 1901. Sur Vorganisation de la Véretille (Veretillum cynomorium (Pall.) Cuv. var. stvlifera Kollik.).
Archives de zoologie expérimentale et générale, Notes (3)9:xlix—lx. [Morphology; Veretillum cyno-
morium.|

BULLOCK, E. 1970. Occurrence of free porphyrins in certain coelenterates. Comparative Biochemistry and
Physiology 33:711—712. [Biochemistry; Balticina finmarchia.]

BULLOUGH, W. S. 1950. Practical invertebrate anatomy. MacMillan and Company, London. 463 pp. [Pennatu-
laceans, pp. 74-76; Pennatula.|

Buss, L. W. AND A. SEILACHER. 1994. The phylum Vendobionta: a sister group of the Eumetazoa? Paleobiology
20(1): 1-4. [Classification of the Vendian biota. ]

CAIRNS, S., J. C. DEN HARTOG, AND C. ARNESON. 1986. Class Anthozoa (Corals, anemones). /n Marine Fauna
and Flora of Bermuda, W. Sterrer, ed. John Wiley and Sons, Inc., New York. 742 pp. [Anthozoans, pp.
159-194; Sclerobelemnon cf. theseus.]

CARLGREN, O. 1940. A contribution to the knowledge of the structure and distribution of the cnidae in the
Anthozoa. Acta Universitatis Lundensis 36(3):1—62. [Kophobelemnon stelliferum - cnidae.]

. 1945. Polypdyr (Coelenterata), III: Koraldyr. Danmarks Fauna Kobenhavn 51:1—167. [Funiculina
quadrangularis, Kophobelemnon stelliferum, Pavonaria finmarchica, Pavonaria christii, Pennatula phos-
phorea, Pennatula aculeata, Pennatula grandis, Protoptilum thomsoni, Stylatula elegans, Virgularia
mirabilis, Virgularia tuberculata. |

CARPINE, C. AND M. GRASSHOFF. 1985. Catalogue critique des Octocoralliaires des collections du Musée
océanographique de Monaco. |. Gorgonaires et pennatulaires. Bulletin de l'Institut oceéanographique,
Monaco 73(1435):1—71.

Case, J. AND J. MORIN. 1966. Glutamate suppression and neuroeffector processes in a coelenterate. American
Zoologist 6:525 (abstract). [Renilla koellikeri. |

Castro, C. B. 1981. Notas sobre Stylatula brasiliensis (Gray, 1870) (Coelenterata, Octocorallia, Pennatulacea).
In Resumos du VI Congreso Brasileiro de Zoologia, Sociedade Brasileira de Zoologia, Brasilia. 80 pp.

CECCHINI, C. 1917. Gli Alcionari e i Madreporari raccolti nel Mediterraneo dalla R. N. “Washington”
(1881-1883). Archivio zoologico italiano, Napoli 9:123—157. [Funiculina quadrangularis, Kophobelem-
non leuckarti.]

CHAPMAN, F. AND I. CRESPIN. 1928. The Sorrento Bore, Mornington Peninsula, with a description of new or
little-known fossils. Victoria Geological Survey, Records 5(1):89—-90, 14, 175. [Graphularia kallimnae
from the Tertiary of Australia. ]

CHAPMAN, G. 1972. A note on the oxygen consumption of Renilla kollikeri Pfeffer. Comparative Biochemistry
and Physiology (A)42(4):863—866.

CHARBONNEAU, H. 1981. The calcium receptors in colelenterate bioluminescence and plants: a study of plant
calmodulin and the luciferin binding protein from Renilla. Dissertation Abstracts International (B) Sciences
and Engineering 42(2):466. [Renilla reniformis; calcium triggered luciferin binding protein, purification
and characterization. |

CHIA, F.-S. AND B. J. CRAWFORD. 1973. Some observations on gametogenesis, larval development and substra-
tum selection of the sea pen Ptilosarcus guerneyi. Marine Biology 23(1):73—-82.

———. 1977. Comparative fine structural studies of planulae and primary polyps of identical age of the sea pen,
Ptilosarcus gurneyi. Journal of Morphology 151(1):131—157.

CHIAJE, S. DELLA. 1827. Memorie sulla storia e notomia degli animali senza vertebri del regno di Napoli. Napoli.

. 1841-44. Descrizione e notomia degli animali invertebrati delle Sicilia citeriore osservati vivi negli anni
1822-1830, 2nd ed. 8 vols. Napoli.

CHILD, C. M. 1951. Oxidation-reduction indicator patterns in three coelenterates. Physiological Zoology
24:97—115. [Stvlatula elongata. |

CHUN, C. 1903. Aus den Tiefen des Weltmeeres. Schilderungen von der Deutschen Tiefsee-Expedition, 2nd ed.
Verlag von Gustav Fischer. 592 pp.

WILLIAMS: INDEX PENNATULACEA 63

CLASTRES, A., A. AHOND, C. POUPAT, P. POTIER, AND S. K. KAN. 1984. Invertébrés marins du lagon Néo-
Calédonien, 2. Etude structurale de trois nouveaux diterpenes isoles du pennatulaire Preroeides laboutei.
Journal of Natural Products 47(1):155-161. [Pteroeides laboutei, lipid and fatty acid content, New
Caledonia. ]

CLASTRES, A., P. LABOUTE, A. AHOND, C. POUPAT, AND P. POTIER. 1984. Invertébrés marins du lagon
Néo-Calédonien, 3. Etude structurale de trois nouveaux diterpenes isolés du pennatulaire, Cavernulina
grandiflora. Journal of Natural Products 47(1):162—166. [Cavernulina grandiflora.|

CoAN, M. H. AND J. TRAVIS. 1970. Comparative biochemistry of proteases from a coelenterate. Comparative
Biochemistry and Physiology 32:127—139.

COLEMAN, D. E. ANDC. TEAGUE. 1973. Sea pansies. Pacific Discovery 26(5):28—29. [The genus Renilla, includes
black and white photographs of living animals. ]

COLIN, P. L. AND C. ARNESON. 1995. Tropical Pacific invertebrates—a field guide to the marine invertebrates
occurring on tropical Pacific coral reefs, seagrass beds and mangroves. Coral Reef Press, Beverly Hills.
296 pp. [Color photographs of living animals: Cavernularia, Cavernulina, Pteroeides, Virgularia.]

CONWAY Morris, S. 1991. Problematic taxa: a problem for biology or biologists? Pp. 19-24 in The early
evolution of Metazoa and the significance of problematic taxa, A. M. Simonetta and S. Conway Morris,
eds. Proceedings of an International Symposium held at the University of Camerino 27-31 March 1989.
Cambridge University Press, Cambridge. 296 pp. [Fossil taxa resembling pennatulaceans: Charniodiscus,
Charnia, Pteridinium. |

. 1993. Ediacaran-like fossils in Cambrian Burgess Shale-type faunas of North America. Palaeontology
36(3):593—635. [Fossil taxon Thaumaptilon walcotti, resembling a sea pen.]

CooKE, M. C. 1889. Toilers in the sea. Society for promoting Christian knowledge, London. 373 pp. [Pennatu-
laceans, pp. 207-214; Pennatula phosphorea, Pavonaria quadrangularis. |

CorMIER, M. J. 1978. Applications of Renilla bioluminescence: an introduction. Methods in Enzymology
57:237-244. [Luminescence; proteins and substrates characteristics and energy transfer. ]

CORNELIUS, P. F.S. AND J. B. Garfath. 1980. The coelenterate taxa of Joshua Alder. Bulletin of the British
Museum (Natural History), Zoology series 39(5):273-291. [P. 274, status of Pennatula mollis Alder,
1867:207 (also of Norman, 1867:196, 206), considered a junior synonym of Pennatula phosphorea
Linnaeus, 1758:818.]

CORNELIUS, P. F. S. AND J. W. WELLS. 1988. Ellis and Solander’s *Zoophytes,’ 1786: six unpublished plates and
other aspects. Bulletin of the British Museum (Natural History), History series 16(1):17—87.

CosTA, O.G. 1841a. Storia e notomia delle Pennatule. /n Frammenti di anatomia comparata. Napoli, 3 fascicles.

. 1841b. Sur les Pennatules. I’ Institut 9(407):345.

CosTA Soares, V. M. F. 1979. Stylatula antillarum KOlliker, 1870 (Octocorallia, Pennatulacea, Virgulariidae).
in Resumos du VI Congreso Brasileiro de Zoologia, Brasilia. 80 pp.

CoTTe, J. 1909. Sur le mot Veretillum. Bulletin de la Société linnéenne de Provence, Marseille 1:31—34.
[ Veretillum.]

COuET, H. G. DE. 1979. Biologie. Der wandernde Korallenstock. Tauchen 2(2):46—47. [Pennatulaceans: bur-
rowing; colony formation; photoperiodic behavior. ]

CoUuLON, L. 1928. Sur quelques animaux curieux du Musée d’ Elbeuf. Société d’Etude des Sciences Naturelles
d’Elbeuf, Bulletin 46:92—93. | Pennatula rubra.]}

COWARD, W. E. 1909. “On Ptilocodium repens.” Verslangen en mededeelingen der Koninklije akademie van
wetenschappen, Afdeeling natuurkunde 17:635—641. [Cited by Hickson (1916:256) with unspecified
pagination. ]

CRAWFORD, B. J. AND F.-S. CHIA. 1974. Fine structure of the mucous cell in the sea pen, Ptilosarcus guerneyi,
with special emphasis on the possible role of microfilaments in the control of mucus release. Canadian
Journal of Zoology 52(12):1427—1432.

CREED, E. L. AND B. C. COULL. 1984. Sand dollar, Melitta quinquiesperforata (Leske), and sea pansy, Renil-
lareniformis (Cuvier) effects on meiofaunal abundance. Journal of Experimental Marine Biology and
Ecology 84(3):225—234. [Behavioral interactions, disturbance effects on meiofaunal abundance in South
Carolina. |

Cuvier, G. L. C. F. D. 1797. Tableau elémentaire de l’histoire naturelle des animaux. Bandouin, Paris. 710 pp.

. 1800. Lecons d’anatomie comparée de G. Cuvier. Recueilles et publiées sous ses yeux, par C. Dumeéril.

Vol. 1. Bandouin, Paris. 521 pp.

64 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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CZECZUGA, B. 1973. Comparative studies of carotenoids in the fauna of the Gullman Fjord (Bohus, Sweden). 1.
Alcyonium digitatum and Pennatula phosphorea (Anthozoa). Marine Biology 19(3):206—209.

DAKIN, W. J. AND OTHERS. 1953. Australian seashores. A guide for the beach lover, the naturalist, the shore
fisherman, and the student. Angus and Robertson, London. 372 pp. [Coelenterata pp. 120-139; Cavernu-
laria obesa.|

DALYELL, J. G. 1839. On the reproduction of the Virgularia or Pennatula mirabilis. Edinburgh New Philosophi-
cal Journal 27:379-381. [Also published in 1840 in Froriep’s Notizen 14:65—68. ]

. 1848. Rare and remarkable animals of Scotland . .. with . . . observations on their nature | and 2. J. Van
Voorst, London.

Dana, J.D. 1846. Zoophytes. United States Exploring Expedition during the years 1838-1842, under the
command of Charles Wilkes, U. S. N., Vol. 7. C. Sherman, Philadelphia. 740 pp. [On p. 586, Dana corrects
the familial name of Ehrenberg (1834) from Pennatulina to Pennatulidae. ]

DANIELSSEN, D.C. 1860. Beskrivelse over en ny Art Virgularia. Forhandlinger 1 Videnskabs-selskabet 1
Christiania 1859:251.

. 1864. Fra den Norske Nordhavs-expedition, Kristiania: 1884:1—46. [Umbellula encrinus: 2306 mm in
length. ]

DANIELSSEN, D. C. AND J. KOREN. 1884. Pennatulida. Den Norske Nordhavs-Expedition, 1876-1878, Zoologi
12:1—-84. Christiania. [Umbellula.]

DARWIN, C. R. 1860. The voyage of the Beagle, annotated and with an introduction by Leonard Engel; 1962
republication. Anchor, Doubleday and Company, New York. 524 pp. [Behavior and description of a sea
pen (“Virgularia patagonica’’) at Bahia Blanca, Argentina, pp. 100, 101 and 202 in the 1889 edition of
“The Voyage” as cited by Hickson 1916:28.]

DatTTA, P. K., A. K. RAY, A. K. BARUA, S. K. CHOWDHURI, AND A. PATRA. 1990. Isolation of a bioactive sterol
from a sea pen Preroeides esperi. Journal of Natural Products (Lloydia) 53(5):1347-1348.

DAVENPORT, D. AND J. A. C. NICOL. 1956. Observations on luminescence in sea pens (Pennatulacea). Proceed-
ings of the Royal Society (B) 144:480-496.

Davis, A. G. 1936. The London clay of Sheppey and the location of its fossils. Proceedings of the Geological
Association of London 47(4):340-345. [Graphularia wetherelli.]

Davis, N. 1978. Studies of the southern California nearshore sand bottom community. Dissertation Abstracts
International (B) 39(3):1080. [Stvlatula elongata; northern Pacific Ocean of California; artificial reef,
effects on population. |

Davis, N., G. R. VAN BLARICOM, AND P. K. DAYTON. 1982. Man-made structures on marine sediments: effects
on adjacent benthic communities. Marine Biology 70(3):295-303. [Stv/atula elongata; population changes,
California. ]

Dawson, C. E. 1966. Additions to the known marine fauna of Grand Isle, Louisiana. The Proceeding of the
Louisiana Academy of Sciences 29:175-180. [Virgularia mirabilis, new record to Louisiana. |

Dawson, J. W. 1872. The pennatulid from Washington Territory. Nature 6:516. [Axis of Halipteris willemoesi.|

DaAwypboFF, C. 1930. Coeloplana duboscqui nov. sp., Coeloplanide Provenant du Golfe de Siam, Commensale
des Pennatules. Archives de Zoologie Expérimentale et Générale 70:87—90.

. 1938. Les Coeloplanides Indochinoises. Archives de Zoologie Expérimentale et Generale. Paris
80:149-161. [Preroeides sp.]

Day, J. H. 1974a. The ecology of Morrumbene Estuary, Mozambique. Transactions of the Royal Society of
South Africa 41(1):3—97.

. 1974b. A guide to marine life on South African shores, 2nd ed. A. A. Balkema, Cape Town. [Octocorals,
pp. 33-35; Pennatula spp., Virgularia schultzei, Virgularia gustaviana. |

Day, J. H., J.G. FIELD, AND M. J. PENRITH. 1970. The benthic fauna and fishes of False Bay, South Africa.
Transactions of the Royal Society of South Africa 39(1):1—108.

DEICHMANN, E. 1936a. Notes on Pennatulacea and Holothuroidea collected by the first and second Bingham
Oceanographic Expeditions 1925—1926. Bulletin of the Bingham Oceanographic Collection, New Haven
5(3):1-11. [Stylatula antillarum, Stylatula elongata, Stylatula brasiliensis, Stvlatula gracilis, Stvlatula
darwini, Umbellula guntheri.|

——. 1936b. The Alcyonaria of the western part of the Atlantic Ocean. Memoirs of the Museum of
Comparative Zoology, Harvard 53:1—317. [Acanthoptilum pourtalesii, Acanthoptilum agassizii, Anthopti-
lum grandiflorum, Anthoptilum murrayi, Anthoptilum sertum, Balticina finmarchica, Balticina christii,

WILLIAMS: INDEX PENNATULACEA 65

Distichoptilum gracile, Funiculina quadrangularis, Gyrophyllum hirondellei, Kophobelemnon stelliferum,

Ptilosarcus (Leioptilus), Pennatula grandis, Pennatula aculeata var. laxa, Pennatula bellissima, Pennatula

americana, Pennatula phosphorea, Pennatula rubra, Pennatula prolifera, Protoptilum carpenterti, Pro-

toptilum aberrans, Protoptilum thomsonii, Protoptilum denticulatum, Protoptilum smitti, Pteroeides
griseum, Pteroeides lusitanicum, Ptilosarcus undulatus, Ptilosarcus guerneyi, Ptilosarcus brevicaulis,

Ptilosarcus sinuosus, Ptilosarcus grayi, Renilla miilleri, R. reniformis forma typica, Renilla reniformis

forma americana, Renilla kéllikeri, Scleroptilum grandiflorum, Scleroptilum gracile, Stvlatula elegans,

Stylatula brasiliensis, Stvlatula darwini, Stylatula antillarum, Umbellula giintheri, Umbellula lindahlii,

Umbellula gracilis, Umbellula crassiflora, Umbellula thomsoni, Umbellula encrinus var. ambigua, Veretil-

lum binghami sp. nov. from California, Virgularia mirabilis, Virgularia kophameli, Virgularia sp..

Virgularia tuberculata.]}

. 1941. Coelenterates collected on the Presidential cruise of 1938. Smithsonian miscellaneous collections
99(10):1—17. [Leioptilus undulatus, Renilla kéllikeri var. tigrina.]

DEKAY, J. E. 1822. Observations on the Pennatule fleche (P. sagitta of LaMarck), in the cabinet of Dr. Mitchell.
American Journal of Science and Arts 4:87—89.

DELAGE, Y. AND E. HEROUARD. 1901. Traité de zoologie concrete, Les Coelentérés 2(2). Librairie C. Reinwald,
Paris. 848 pp. [Pp. 429-458 dedicated to morphology and anatomy in Pennatulacea, with detailed illustra-
tions of development in Renilla; other taxa illustrated include Veretillum, Umbellula, Kophobelemnon,
Funiculina, Pennatula, Pteroeides, Virgularia, and Stylatula; plate 54 - morphology of Kophobelemnon. |

DELUCA, M., M. E. DEMPSEY, K. Hort, AND M. J. CORMIER. 1976. Source of oxygen in the CO) produced during
chemiluminescence of firefly luciferyl-adenylate and Renilla luciferin. Biochemical and biophysical
research communications 69(1):262—267.

Demir, M. 1952. The invertebrate benthos of the Bosporus and of the litoral of the Sea of Marmara closer to the
Bosporus. Istanbul Universitesi fen Fakiiltesi Hidrobiologie Arastirma Enstitiisii yayiinlariidan. 3:1-615.
[In Turkish. ]

DENDY, A. 1897. On Virgularia gracillima in Lyttelton Harbour. Transactions and Proceedings of the New
Zealand Institute 29 (new series 12):256—257. [Abstract in Zoologisches Centralblatt 5:251; Virgularia
gracillima.|

DE WAELE, J.-P. et al. (See WAELE, J.-P. DE et al.)

D’HONDT, M.-J., or DDHONDT, M.-J. (See HONDT, M.-J. D’.)

DICKINSON, P. 1978. Conduction systems controlling expansion-contraction behavior in the sea pen Prilosarcus
gurneyi. Marine Behaviour and Physiology 5(2):163—183. [Nervous electrophysiology; nerve net control
of expansion contraction behaviour. }

DINAMANI, P. 1965. On a collection of Umbellula from the Arabian Sea. Nytt Magasin for Zoologi 12
(1964/65):30-34.

DING, Q. AND Y. CHEN. 1981. Discovery of soft metazoan from the Sinian system along eastern Yangtze Gorge,
Hubei. Earth Science, Journal of Wuhan College of Geology 1981 (2)15:53—S7. [Charnia dengyingensis
sp. n.; China; Proterozoic; Precambrian fossil resembling a sea pen.]

DITTRICH, R. 1888. Uber das Leuchten der Tiere. Wissenschaftliche Beilage zum Program d. Realgymm. am
Zwinger zu Breslau. 70 pp. [Bioluminescence. ]

DoLLrus, R. 1938 (or 1939). Sur un octocoralliaire du genre Cavernularia, commun sur les fonds cotiers de
1’ Atlantique Marocain. Travaux de la Station zoologique de Wimereux 13:243-265. [Cavernularia pusilla. ]

DODERLEIN, L. 1902. Ueber die Beziehungen nahe verwandter Tierformen zu einander. Zeitschrift fiir Morpholo-
gie und Anthropologie Bd. 4:394.

DONOVAN, P. 1995. Sea pens. Tropical Fish Hobbyist 43(7):52, 54-57, 60, 62.

Dube, M. A. AND E. BALL. 1971. Desmarrestia sp. associated with the sea pen Ptilosarcus gurneyi (Gray).
Journal of Phycology 7:218—220. [Ecological interaction: rhodophyte alga with sea pen.]

DUNKELBERGER, D. AND N. WATABE. 1972. Electron microscope study of the coenenchyme in the pennatulid
colony Renilla reniformis, with special emphasis on spicule formation. American Zoologist 12:716—717.

. 1974. An ultrastructural study on spicule formation in the pennatulid colony Renilla reniformis. Tissue
and Cell 6(4):573—5S86.

DUNN, D. F. 1982. Cnidaria. /n Synopsis and Classification of Living Organisms, S. A. Parker, ed. McGraw-Hill
Book Co., New York. 1:669—706.

66 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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Dyer, M. F., W. G. FRY, AND P. D. FRY. 1981. Results from North Sea benthos surveys. Porcupine Newsletter
2(1):12-14. [Pennatula phosphorea; distribution and population density in the North Sea. ]

Dzik, J. 1991. Is fossil evidence consistent with traditional views of the early metazoan phylogeny? Pp. 47-S6
in Proceedings of an International Symposium held at the University of Camerino 27-31 March 1989,
A. M. Simonetta and S. Conway Morris, eds. Cambridge University Press, Cambridge. 296 pp. [Discusses
the enigma of the paucity of Cambrian/Precambrian coelenterates. ]

ECKELBARGER, K. J., P. A. TYLER, AND R. W. LANGTON. 1998. Gonadal morphology and gametogenesis in the
sea pen Pennatula aculeata (Anthoza: Pennatulacea) from the Gulf of Maine. Marine Biology
132(4):677-690.

EHRENBERG, C. G. 1834a. Beitrage zur physiologishcen Kenntniss der Corallenthiere im allgemeinen, und
besonders des rothen Meeres, nebst einem Versuche zur physiologischen Systematik derselben. Abhan-
dlungen der K6niglichen Akademie der Wissenschaften zu Berlin. Aus dem Jahre 1832. Erster
Theil:225—380. [Introduction of the familial name Pennatulina on p. 287; later corrected to Pennatulidae
by Dana, 1846:586.]

_1834b. Uber die Natur und Bildung der Corallenbanke des rothen Meeres. Abhandlungen der

K6niglichen Akademie der Wissenschaften zu Berlin. Aus dem Jahre 1832. Erster Theil:381—432.

. 1834c. Das Leuchten des Meeres. Abhandlungen der Kéniglichen Akademie der Wissenschaften zu
Berlin 1834:411—572. [Bioluminescence. |

EISEN, G. 1876. Bidrag till Kannedomen om Pennatulid-slagtet Renilla. Bihang till Kongl. Svenska vetenskaps-
akademiens handlingar (n.f.)13(1):1—15. [Renilla from California. ]

ELLIS, J. 1753. A letter from Mr. John Ellis to Mr. Peter Collinson F. R. S., concerning a Cluster-Polype found
in the sea near the coast of Greenland. Philosophical Transactions of the Royal Society 48(1):305—308. [See
also the abridged 1809 account of the Transactions 10:409; this is a description of the original find of
Umbellula; see also Mylius 1753 and 1754.]

. 1755. An account of clustered polype, found in the North Seas, near the Pole, Pp. 96-99 in An essay

towards a natural history of the corallines, and other marine productions of the like kind, commonly found

on the coasts of Great Britain and Ireland . . . to which is added the description of a large Polype taken near
the North Pole by the Whalefishers. For the author, London. 103 pp.

. 1764. An account of the sea pen, or Pennatula phosphorea of Linnaeus; likewise a description of a new
species of sea pen, found on the coast of South-Carolina, with observations on sea-pens in general. In a
letter to the honourable Coote Molesworth, Esq; M. D. and F.R. S. from John Ellis, esq; F. R. S. and
member of the Royal Academy of Upsal. Philosophical Transactions of the Royal Society of London
53:419-435. [Pennatulacean bioluminescence. |

ELLIS, J. AND D. SOLANDER. 1786. The natural history of many curious and uncommon zoophytes, collected
from various parts of the globe by the late John Ellis . . . systematically arranged and described by the lated
Daniel Solander . . . London, printed for Benjamin White and Son, at Horace’s Head, Fleet-Street; and Peter
Elmsly, in the Strand. 206 pp. [Pennatula phosphorea, Pennatula rubra, Pennatula spinosa, Pennatula
mirabilis, Pennatula antennina, Pennatula cynomorium, Pennatula reniformis, Pennatula argentea, Pen-
natula encrinus. See Cornelius and Wells (1988).]

ERASMI, F. 1668. Ost- und West-Indischer wie auch Sinesischer Lust- und Stats-Garten, mit einem Vorgesprach
von mancherlei lustigen Discursen; In drei Haupt = Theile unterschieden. Der erste Theil Begrifft in sich
die edelsten Blumen, Krauter, Baume . . . in Ost-Indien, Sina und America: .. . aus den furnemsten, alten
und neuen, indianischen Geschicht- Land- und Reisbeschreibungen, mit Fleiss zusammengezogen, und aus
annehmliche Unterredungs-Art eingerichtet. Johann Andreae Endters, und Wolfgang dess Jugern Sel.
Erben, Nurnberg. 1762 pp.

ERHARDT, H. S. AND H. MOOSLEITNER. 1995. Meerwasser Atlas Band 2. Wirbellose Tiere. Mergus Verlag,
Melle, Germany. 736 pp. [Sarcoptilus sp., Pennatula sp., Pteroeides sp. Cavernularia obesa, Veretillum
cynomorium, Virgularia sp.; color photographs of living animals. ]

Esper, E. J. C. 1791. Die Pflanzenthiere in Abbildungen nach der Natur mit Farben erleuchtet nebst Beschreibun-
gen, Theil 2:1-96.

FAGER, E. W. 1968. A sand-bottom epifaunal community of invertebrates in shallow water. Limnology and
Oceanography 13:448-64. [Stvlatula elongata. |

FAULKNER, D. AND R. CHESHER. 1979. Living corals. Clarkson N. Potter, Inc., New York. 310 pp. [Pennatu-
laceans: plate 62 is a large-format color photograph of Ptilosarcus gurneyi, text on p. 276.]

WILLIAMS: INDEX PENNATULACEA 67

FAUTIN, D. G. AND R. N. MARISCAL. 1991. Cnidaria: Anthozoa. Pp. 267-358 in Microscopic Anatomy of
Inveretebrates, Vol. 2: Placozoa, Porifera, Cnidaria and Ctenophora, W. Harrison and J. A. Westfall, eds.
Wiley-Liss, New York.

FAUTIN, D. G., A. E. SIEBERT, AND E. N. KOZLOFF. 1987. Class Anthozoa. /n Marine Invertebrates of the Pacific
northwest, E. N. Kozloff, ed. University of Washington Press, Seattle and London. [Taxonomic listing, pp.
68-71 on Anthozoa: Kophobelemnon affine, Kophobelemnon biflorum, Kophobelemnon hispidum, Anthop-
tilum grandiflorum, Funiculina parkeri, Helicoptilum rigidum, Scleroptilum sp., Umbellula lindahli,
Umbellula loma, Umbellula encrinus, Umbellula carpenteri, Umbellula magniflora, Balticina californica,
Balticina septentrionalis, Verrillia blakei, Stvlatula elongata, Stylatula columbiana, Virgularis spp..,
Pennatula phosphorea, Ptilosarcus gurneyi.]|

FEDONKIN, M. A. 1992. Vendian faunas and the early evolution of Metazoa. /n Origin and early evolution of the
Metazoa. Topics in Geobiology, vol. 10, J. H. Lipps and P. W. Signor, eds. Plenum Press, New York. 570
pp. [Interpretation of Vendian fossils. ]

. 1996. The oldest fossil animals in ecological perspective. /n Biology as history No. 2 - new perspectives
on the history of life: essays on systematic biology as historical narrative, M. T. Ghiselin, and G. Pinna,
eds. Memoirs of the California Academy of Sciences 20:1—107. [Interpretation of Vendian fossils. |

FIELD, L. R. 1949. Sea anemones and corals of Beaufort, North Carolina. Bulletin of the Duke University Marine
Station, Durham, North Carolina, No. 5:1—39. [Renilla reniformis.]

FISCHER. P. 1889. Note sur le Pavonaria quadranularis et sur les Pennatulides des cotes de France. Bulletin de
la Societe Zoologique de France 14:34—38. [Presumably refers to Funiculina quadrangularis from France. |

FISHER, W. J. 1874. On a new species of alcyoniid polyp. Proceedings of the California Academy of Sciences
5:418. [Virgularia ornata from Japan. ]

FLORA, C. J. AND E. FAIRBANKS. 1966. The sound and the sea. Pioneer, Bellingham, Washington. 455 pp.
[Ptilosarcus gurneyi listed as Leioptilus guerneyi.]

FORBES, E. 1847. [Jn Johnson (1847); a published letter from Edward Forbes regarding observations on
bioluminescence in Pennatula phosphorea.|

FORBES, E. AND J. GOoDsIR. 1851. On some remarkable marine Invertebrata new to the British Seas. Transactions
of the Royal Society of Edinburgh 20(2):307-315. [Pennatulacean bioluminescence. }

Forest, J. 1996. Henri Milne Edwards (23 October 1800—29 July 1885). Journal of Crustacean Biology
16(1):208—213. [Biographical sketch of Henri Milne Edwards. |

Fossa, S. A. AND A. J. NILSEN. 1995. Korallenriff-Aquarium - Band 4 - Nesseltiere im Korallenriff und fiir das
Korallenriff-Aquarium. Birgit Schmettkamp, Bornheim. 448 pp. [Pp. 228-229, color photographs of
Cavernularia sp., possibly Veretillum sp., and Pteroeides sp.]

FOWLER, G. H. 1888. Ona new Pennatula from the Bahamas. Proceedings of the Zoological Society of London
1888:135—140. [Pennatula bellissima. ]

. 1894. On two sea-pens of the family Veretillidae from the Madras Museum. Proceedings of the
Zoological Society of London 1894:376-379. [Abstract in: Journal of the Royal Microscopical Society
1894:576; Cavernularia malabarica, Lituaria phalloides.|

FOWLER, H. W. 1911. A description of the fossil fish remains of the Cretaceous, Eocene, and Miocene formations
of New Jersey. New Jersey Geological Survey Bulletin 4:1-192. [Specimens of the extinct sea pen
Graphularia ambigua, thought to be spines of a batoid fish by Fowler; see Shapiro and Ramsdell (1965). }

FRANC, J.-M. 1979. Collagene intracellulaire chez un invertébré marin (cnidaire). Biologie cellulaire 35(2):6a.
[Abstract in French; Veretillum cynomorium; protein metabolism; collagen fibrils synthesis and intracellular
localization. |

FRANC, S. 1970. Les evolutions cellularies au cours de la regénération du pédoncle de Veretillum cynomorium
Pall. Vie et Milieu 21(I-A):49—93.

. 1973. Essai de culture in vitro de cellules de Veretillum cynomorium Pall. (Cnidaire anthozoaire).

Comptes rendus hebdomadaires des séances de |’ Academie des sciences, Paris (Série D) Sciences Naturelles

276(4):559-562.

. 1979. Genése de la matrice conjonctive de Veretillum cynomorium Pall. (cnidaires-anthozoaires). Etude
ultrastructuale et autoradiographique. Archives D’ Anatomie Microscopique et de Morphologie Experimen-
tale 68(4):237—256. [Tissues; ultrastructural and autoradiographic study; genesis of connective matrix. ]

FRANC, S., R. GARRONE, AND A. Huc. 1971. Observations ultrastructurales sur la mésoglée de Veretillum
cynomorium Pall. (Cnidaire, anthozoaire). Journal de Microscopie, Paris 11:57. [Abstract only. ]

68 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 2

FRANC, S., A. HUC, AND G. CHASSAGNE. 1974. Etude ultrastructurale et physico-chemique de |’axe squelettique
de Veretillum cynomorium Pall. (cnidaire, anthozoaire): cellules, calcite, collagene. Journal de Microscopie,
Paris 21(1):93—110.

FRANC, S., P. W. LEDGER, AND R. GARRONE. 1985. Structural variability of collagen fibres in the calcareous
axial rod ofa sea pen. Journal of Morphology 184(1):75—84. [Skeletal rod collagen compared with mesoglea
fibrils. |

FRANSEN, C. H. J. M., L. B. HOLTHUIS, AND J. P. H. M. ADEMA. 1997. Type-catalogue of the Decapod Crustacea
in the collections of the Nationaal Natuurhistorisch Museum, with appendices of pre-1900 collectors and
material. Zoologische Verhandelingen 311:1—344. [Appendix 2 contains biographical sketches of several
authors of pennatulacean research—Pieter Bleeker, Jan Adrianus Herklots, Sydney John Hickson, Am-
brosius Arnold Willem Hubrecht, Willy Kiikenthal, and Henri Milne Edwards, including portraits of all
except Hickson. ]

FRANZEN, A. 1967. Remarks on spermiogenesis and morphology of the spermatozoon among the lower Metazoa.
Arkiv for zoologi 19:335—342. [Funiculina quadrangularis.|

FRECH. F. 1890. Die Korallenfauna der Trias. Palaeontographica 37(1—4):1—116. [Prographularia triadica from
the Triassic. ]

FRISCHE, J. 1991. Seefedern. Eine Randgruppe in der Meerwasser-Aquaristik. Aquarium (Minden) No. 267
1991:49-S0.

Fu, X., F. J. SCHMITZ, AND G. C. WILLIAMS. In press. Malayenolides A-D, novel diterpenes from the Indonesian
sea pen Veretillum malayense. Journal of Natural Products. [New diterpenes found to be toxic to brine
shrimp; includes a brief assessment of systematics in the genus Veretillum.|

FusITA, T. AND S. OHTA. 1988. Photographic observations of the life style of a deep-sea ophiuroid Asteronyx
loveni (Echinodermata). Deep-Sea Research, Part A, Oceanographic Research Papers 35(12):2029-2043.
[Anthoptilum, Funiculina quadrangularis, ecological associations with Asteronyx loveni in the western
Pacific. ]

GABB, W. M. 1859. Catalogue of the invertebrate fossils of the Cretaceous formation of the United States, with
references. Issued with: Proceedings of the Academy of Natural Sciences of Philadelphia 11:1—20. [The
extinct sea pen Graphularia ambigua listed by Gabb as Belemnites ambiguus; see Shapiro and Ramsdell
(1965:2).]

. 1861. Synopsis of the Mollusca of the Cretaceous formations, including the geographical and strati-

graphical range and synonymy. Proceedings of the American Philosophical Society 8:57—257. [The extinct

sea pen Graphularia ambigua listed by Gabb as Belemnites ambiguus; see Shapiro and Ramsdell (1965:2).]

. 1862. Description of two new species of Pennatulidae from the Pacific coast of the United States.

Proceedings of the California Academy of Natural Sciences 2:166—167. [Pennatula tenua, Virgularia

elongata. |

. 1864. Description of new a species of Virgularia from the coast of California. Proceedings of the
California Academy of Natural Sciences 3(2):120. [Virgularia gracilis.]

GAGE, J.D. AND P. A. TYLER. 1991. Deep-sea biology, a natural history of organisms at the deep-sea floor.
Cambridge University Press, Cambridge. 504 pp. [Kophobelemnon stelliferum, Pennatula aculeata, Um-
bellula.|

GARDINER, J. S. 1940. Prof. Sydney J. Hickson, F. R. S. Nature, London 145(3670):338—339. [Biographical
sketch of Sydney Hickson. ]

GELDIAY, R. AND A. KOCATAS. 1972. Note préliminaire sue les peuplements benthiques du golfe d’Ismir.
Scientific Monographs of the Faculty of Science, Ege University 12:1—34.

GERMAIN, G. AND M. ANCTIL. 1988. Luminescent activity and ultrastructural characterization of photocytes
dissociated from the coelenterate Renilla kollikeri. Tissue and Cell 20(5):701—720. [Renilla kollikeri,
luminous organs, photocyte luminescent activity and ultrastructural characterization. |

GESNER, C. 1555. De raris and admirandis herbis: quae, sive quod noctu luceant, sive alias ob causas, lunariae
nominantur and obiter de aliis etiam rebus quae in tenebris lucent, commentariolus. Cum iconibus
quibusdam Herbarum novis. Editione hac secunda emendatior. Typis Matthiae Godicchenii, Impensis Petri
Hauboldi, Hafniae. 82 pp. [Subsequent printing dated 1669; Penna marina bioluminescence. ]

—. 1558. Historiae animalium liber III. quae est de piscium and aquatilium animantium natura. Cum
iconibus singulorum ad vivum expressis. Continentur in hoc volumine, Gulielmi Rondeletii and Petri

WILLIAMS: INDEX PENNATULACEA 69

Bellonii Cenomani de aquatilium singulis scripta. Apud C. Froschoverum, Tiguri. 20 lvs. + 1297 pp.

[Pennatulacean bioluminescence. ]

. 1565. De rerum fossilium, lapidum et gemmarum maxime, figuris and similitudinibus Liber: non solum
Medicis, sed omnibus rerum Naturae ac Philologiae studiosis, vtilis and iucundus futurus. Jac. Gesner,
Tiguri. 7 + 169 lvs.

GILI, J. M. 1986. Estudio sistematico y faunistico de los cnidarios de la costa catalana. Ph. D. dissertation,
Universidad Autonoma de Barcelona, Espafia. 565 pp.

GILI, J. M. AND F. PAGES. 1987. Pennatulaceos (Cnidaria, Anthozoa) recolectados en la plataforma continental
Catalana (Mediterraneo occidental). Miscel-lania Zoologica (Barcelona) | 1:25—39. [Taxonomic descrip-
tions: Cavernularia pusilla, Funiculina quadrangularis, Kophobelemnon stelliferum, Pennatula phos-
phorea, Pennatula rubra, Pteroeides spinosum, Veretillum cynomorium, and Virgularia mirabilis.]

GILI, J. M., J. D. Ros, AND F. PAGES. 1987. Types of bottoms and benthic Cnidaria from the trawling grounds
(littoral and bathyal) off Catalonia (NE Spain). Vie et Milieu 37(2):85—98. [Ecology: Cavernularia pusilla,
Funiculina quadrangularis, Kophobelemnon stelliferum, Pennatula phosphorea, Pteroeides griseum,
Veretillum cynomorium, Virgularia mirabilis. |

GILLULY, R. H. 1970. Umbellula in its deep-sea habitat. Science News 97(24):586-587. [Deep-sea ecology of
Umbellula.]

GLAESSNER, M. F. 1958a. The oldest fossil faunas of South Australia. Geologische Rundschau 47(2):522-531
[Precambrian fossils Preridinium and Rangea, resembling sea pens. ]

. 1958b. Precambrian Coelenterata from Australia. Transactions of the Royal Society of Australia

81:185—188. [Precambrian fossils resembling sea pens. ]

. 1959. Precambrian Coelenterata from Australia, Africa and England. Nature, London

183(4673):1472—1473. [Precambrian fossils Charnia masoni, Charniodiscus, Rangea, and Pteridinium,

resembling sea pens. |

. 1961. Pre-Cambrian animals, Pp. 63-69 in The fossil record and evolution—teadings from Scientific

American. W. H. Freeman and Company, San Francisco. 225 pp. [Sea pens, Pennatula rubra, Pennatula

aculeata; and fossil organisms resembling sea pens, Rangea arborea and Charnia.]

. 1984. The dawn of animal life—a biohistorical study. Cambridge University Press, London. 244 pp.
[Precambrian colonial cnidarians, some of which resemble pennatulaceans, pp. 52, 56—57.]

GLAESSNER, M. F. AND B. DAILY. 1959. The geology and late Precambrian fauna of the Ediacara Fossil Reserve.
Records of the South Australian Museum 13(3):369-401. [Precambrian fossils that resemble pennatu-
laceans. }

GLAESSNER, M. F. AND M. WADE. 1966. The Late Precambrian fossils from Ediacara, South Australia.
Palaeontology 9:599-628. [Vendian fossils resembling sea pens: Rangea longa, Rangea grandis, Arborea
arborea, Pteridium cf. simplex.]

GopFREY, F. K. 1943. Sea-Pens. The South Australian Naturalist, Adelaide 21(2):13—15. [Sarcophyllum
grande. |

GOLL, R. M. 1980. Pliocene-Pleistocene radiolarians from the East Pacific Rise and the Galapagos Spreading
Center, Deep Sea Drilling Project Leg 54. Initial Reports of the Deep Sea Drilling Project 54:425—453.

GOSLINER, T. M., D. W. BEHRENS, AND G. C. WILLIAMS. 1996. Coral reef animals of the Indo-Pacific, animal
life from Africa to Hawai’i exlusive of the vertebrates. Sea Challengers, Monterey. 314 pp. [Underwater
color photographs of living animals; sea pens, pp. 56-60: Cavernularia cf. obesa, Cavernularia sp.,
Pteroeides spp., Sclerobelemnon burgeri, Sclerobelemnon sp., Scytalium cf. sarsii, Veretillum spp.,
Virgularia gustaviana, Virgularia spp.; p. 93 (no. 307) shows a color photograph of an ecological interaction
between an attached jellyfish (cf. Lipkea sp.) and the sea pen Preroeides sp., taken underwater in Indonesia;
and on page 227 (no. 826) shows commensal procelain crabs Porcellanella picta on the sea pens, Veretillum
sp. and Pteroeides sp., underwater photographs taken in the Philippines and Indonesia. ]

GoswaMlI, A., S. SAHA, S. MUKHOPADHYAY, AND S. C. PAKRASHI. 1995. Marine products from Bay of Bengal,
3: Sterol composition of four molluscs and a sea pen. Indian Journal of Marine Sciences 24(4):238-239.

GOTSHALL, D. W. 1987. Marine animals of Baja California—a guide to the common fishes and invertebrates,
2nd ed. Sea Challengers, Monterey. 113 pp. [Octocorals, pp. 86-91; color photograph of Ptilosarcus
undulatus.]}

70 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 2

GOTSHALL, D. W. 1994. Guide to marine invertebrates Alaska to Baja California. Sea Challengers, Monterey.
105 pp. [Octocorals, pp. 31-36; color photographs of Renillia koellikeri, Ptilosarcus gurneyi, Stylatula
elongata. |

GOTSHALL, D. W. AND L. L. LAURENT. 1979. Pacific coast subtidal marine invertebrates—a fishwatcher’s guide.
Sea Challengers, Los Osos. 112 pp. [Octocorals, pp. 35-39; color photographs of Renillia koellikeri,
Ptilosarcus gurneyi, and Stvlatula elongata.|

GRANT, R. E. 1827. Notice regarding the structure and mode of generation of the Virgu/aria and the Pennatula
phosphorea. Edinburgh Journal of Science 7:330-335. [Also published in Frorieps Notizen 19:337—341,
pennatulacean bioluminescence. ]

. 1929. Further observations on the generation of the Virgularia mirabilis. Edinburgh Journal of Science
10:350-351. [Also published in Frorieps Notizen 24:247.]

GRASSHOFF, M. 1972. Eine Seefeder mit einem einzigen Polypen: Umbellula thieli n. sp. Die von F. S. ‘Meteor’
1967-1970 im ostlichen Nordatlantik gedredschten Pennatularia (Cnidaria: Anthozoa). Meteor Forschung-
sergebnisse (D)12:1—11.

. 1973. Schopf-Seefedern - Erforschungsgeschichte und Evolution einer Tiefseetiergruppe. Natur und

Museum 103(2):58-64. [Umbellula; evolutionary trends; deep-sea benthos. ]

. 1981. Gorgonaria und Pennatularia (Cnidaria: Anthozoa) vom Mittelatlantischen Riicken SW der

Azoren. Steenstrupia 7(9):213-230. [Funiculina quadrangularis, distribution patterns in the Atlantic

Ocean. |

. 1982a. Die Gorgonaria, Pennatularia und Antipatharia des Tiefwassers der Biskaya (Cnidaria, Antho-

zoa). Ergebnisse der franzésischen Expeditionen Biogas, Polygas, Geomanche, Incal, Noratlante und

Fahrten der ‘Thalassa.’ I. Allgemeiner Teil. Bulletin du Museum national d’histoire naturelle, Section A:

Zoologie, Biologie et Ecologie Animales 3(3):731—766. [Pennatulacea, bathyal zone, geographic and

bathymetric distribution, world marine zones: Kophobelemnon macrospinosum, Anthoptilum grandiflorum,

Anthoptilum murrayi, Funiculina quadrangularis, Protoptilum carpenteri, Distichoptilum gracile, Sclerop-

tilum grandiflorum, Umbellula monocephalus, Umbellula durissima, Umbellula thomsoni, Umbellula

lindahli, Pennatula grandis, Pennatula aculeata.|

. 1982b. Die Gorgonaria, Pennatularia und Antipatharia des Tiefwassers der Biskaya (Cnidaria, Antho-
zoa). Ergebnisse der franzésischen Expeditionen Biogas, Polygas, Geomanche, Incal, Nortatlante und
Fahrten der ‘Thalassa.’ I]. Taxonomischer Teil. Bulletin du Muséum national d’histoire naturelle, Section
A: Zoologie, Biologie et Ecologie Animales (4)3(A)4:94 1-978. [Taxonomic descriptions: Kophobelemnon
macrospinosum, Anthoptilum grandiflorum, Anthoptilum murrayi, Protoptilum carpenteri, Distichoptilum
gracile, Scleroptilum grandiflorum, Umbellula monocephalus, Umbellula durissima, Umbellula thomsoni,
Umbellula lindahli, Pennatula grandis.|

————. 1989. Die Meerenge von Gibraltar als Faunen-Barriere: Die Gorgonaria, Pennatularia und Antipatharia
der BALGIM-Expedition (Cnidaria: Anthozoa). Senckenbergiana Maritima 20(5—6):201—223. [Listing of
sea pens with distributional and bathymetric data: Kophobelemnon stelliferum, Funiculina quadrangularis,
Protoptilum thomsoni, Stylatula elegans, Umbellula thomsoni, Umbellula pallida, Virgularia mirabilis,
Virgularia gracilis, Pennatula aculeata, Pennatula phosphorea, Gyrophyllum hirondellei. |

.1991a. Die Evolution der Cnidaria. I. Die Entwicklung zur Anthozoen-Konstruktion. Natur und

Museum 121(8): 225-236. [Anthozoan evolution. ]

.1991b. Die von E. J.C. Esper 1788-1809 beschriebenen Anthozoa (Cnidaria). Senckenbergiana
Biologica 71:325—368.

GRAVELY, F. H. 1941. Shells and other animal remains found on the Madras Beach, I: Groups other than snails,
etc. (Mollusca Gastropoda). Madras Government Museum Bulletin, N.S. Natural History Section
5(1):1-112. [Coelenterates, pp. 6-18; Cavernularia sp.]

GRAVENHORST, J. L. C. 1821. De natura vegetabili-Gorgoniarum. Memorie della Accademia della Scienze di
Torino 26:41 1—432. [Scytaliopsis ghardagensis. |

GRAVIER, C. 1906a. Sur un type nouveau de Virgulaire. Bulletin du Muséum national d’histoire naturelle
12(5):291-293. [Scytaliopsis djiboutiensis. |

. 1906b. Sur la biologie des virgulaires. Bulletin du Muséum national d’histoire naturelle 12:391—395.

———. 1906c. Sur un type nouveau d’Alcyonaire de la famille des Virgularidae. Comptes rendus hebdo-
madaires des Seances de |’ Academie des sciences, Paris (Série D) Sciences Naturelles 142:1290—1291.
[Scytaliopsis gen. nov.]

WILLIAMS: INDEX PENNATULACEA 71

. 1907a. Sur un genre nouveau de Pennatulidés (Mesobelemnon nov. g. gracile nov. sp.). Bulletin du

Muséum national d’histoire naturelle 13(2):159—161 [Also published as: Sur un genre nouveau de

Pennatulidés. Comptes rendus hebdomadaires des Seances de |’Academie des sciences, Paris (Serie D)

Sciences Naturelles 144:439-440; Mesobelemnon gracile. |

. 1907b. Sur les pennatulidés de la famille des Kophobelemnonidae KOlliker. Bulletin du Muséum

national d’histoire naturelle, Paris 13:161—164. [Kophobelemnidae: the genera composing the family. ]

. 1908. Recherches sur quelques Alcyonaires du Golfe de Tadjourah. Archives de zoologie expérimentale

et générale (4)8(2): 179-266. [Mesobelemnon gracile, Scytaliopsis djiboutiensis.|

. 1912a. Sur une nouvelle famille des Pennatulidés (Scytaliopsidae). Proceedings of the Seventh Inter-

national Zoological Congress. Cambridge, Mass. 1912:819—821.

. 1912b. Sur la biologie des Pennatulidés. Proceedings of the Seventh International Zoological Congress.
Cambridge, Mass. 1912:822-825.

GRAY, J. E. 1840. The third room (radiated animals). Pp. 66—77 in Synopsis of the contents of the British Museum
(Ed. 42), London. 273 pp.

. 1848. Description of Sarcoptilus, a new genus of Pennatulidae. Proceedings of the Zoological Society

of London 1848(16):45. [Also published in 1849 in Annals and Magazine of Natural History (2)3:76—77. |

. 1859. On the arrangement of zoophytes with pinnated tentacles. Annals and Magazine of Natural

History, ser. 3, 4:439-444.

. 1860. Revision of the family Pennatulidae, with some descriptions of some new species in the British

Museum. Annals and Magazine of Natural History, ser. 3, 5:20—25.

. 1862. Notes on some specimens of claviform Pennatulidae (Veretillidae) in the collection of the British

Museum. Annals and Magazine of Natural History, ser. 3, 10:73—76.

. 1870. Catalogue of sea-pens or Pennatulariidae in the collection of the British Museum. British Museum,

London. 40 pp. [Funiculina quadrangularis, Funiculina forbesii, Funiculina philippinensis, Balticina

finmarchica, Norticina christii, Scytalium sarsii, Virgularia juncea, Virgularia reinwardtii, Virgularia

elegans, Virgularia philippinensis, Virgularia pusilla, Virgularia gracilis, Virgularia elongata, Virgularia
patachonica, Lygus mirabilis, Lygus vanbenedenii, Lygus ellisii, Lygus brasiliensis, Stvlatula gracilis,

Stylatula elongata, Pennatula rubra, Phosphorella phosphorea, Ptilella borealis, Leioptilus fimbriatus,

Leioptilus undulatus, Argentella elegans, Argentella grandis, Argentella jukesii, Pteromorpha dringii,

Pteromorpha grisea, Pteromorpha expansa, Pteroeides latipinnarum, Pteroeides spinosum, Pteroeides

esperi, Pteroeides japonicum, Crispella sieboldii, Sarcoptilus grandis, Ptilosarcus gurnevi, Ptilosarcus

sinuosus, Kophobelemnon stelliferum, Kophobelemnon burgeri, Kophobelemnon clavatum, Veretillum
cynomorium, Veretillum obesa, Veretillum australasiae, Veretillum cantoriae, Veretillum stimpsonii,
Veretillum baculatus, Veretillum valenciennesii, Veretillum luteum, Veretillum pusillum, Lituaria phal-
loides, Policella australis, Clavella australasia, Renilla reniformis, Renilla amethystina, Renilla danae,

Renilla peltata, Renilla patula, Renilla australasica, Herklotsia edwardsii, Renillina sinuata, Umbellula

groenlandica, Crinillum siedenburgii, Osteocella cliftoni.]

. 1872a. On the genus Osteocella. Annals and Magazine of Natural History, ser. 4, 9:405—406. [Refers

to the genus Halipteris. |

. 1872b. Additional note on Osteocella. Annals and Magazine of Natural History, ser. 4, 10:76, 406.

[Refers to the genus Halipteris.]

. 1872c. The clustered sea-polype (Umbellula groenlandica). Annals and Magazine of Natural History,

ser. 4, 10:151, 469.

. 1873. On the stick-fish (Osteocella septentrionalis) and on the habits of sea-pens. Nature 9:13—14.
[Probably correctly identified as Halipteris willemoesi.]

GREGORIO, A. DE. 1890. Monographie de la faune Eocenique de |’Alabama. Annales de geologie et de
paleontologie 7/8:1—346. [Shapiro and Ramsdell (1965:2) stated, “In 1890, de Gregorio described a
pennatulid from the Eocene of Alabama, which he named Corallium perplexum. To judge from his figures,
de Gregorio’s species appears to be the same as Graphularia ambigua (Morton) and probably is a synonym
of the latter. De Gregorio gave no formation other than ‘Eocene’ or locality other than ‘Alabama’ for his
specimens. Unfortunately, de Gregorio’s specimens were unavailable for study at the time of this writing
... Yabe and Sugiyama, in 1937, in a listing of the known species of Graphularia, placed Corallium
perplexum de Gregorio in that genus.” ]

72 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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GRIEG, J. A. 1887. Bidrag til de norske Alcyonarier. Bergens Museums Aarsberetning for 1886:1—26. [Protop-
tilum tortum, Stichoptilum arcticum.]

. 1892. Ovsersigt over Norges pennatulider. Bergens Museums Aarsberetning for 1891(1):1—22.

. 1893. Bidrag til kjendskaben om de nordishke alcyonarier. Bergens Museums Aarbog 1893(2):1—21.

. 1896. On Funiculina and Kophobelemnon. Bergens Museum Aarbog 1896(3):1—11. [Abstracts in
Journal of the Royal Microscopical Society 1897:132, and Zoologisches Centralblatt 4:448-449; Funicu-
lina quadrangularis, Kophobelemnon stelliferum.|

GRIMMELIKHULZEN, C. J. P., D. GRAFF, A. GROEGER, ANDI. D. MCFARLANE. 1987. Structure, action and location
of coelenterate arg-phe-amide peptides. Bulletin de |’ Académie serbe des sciences et des arts, Classe des
sciences naturelles et mathématiques 96(29):13—24. [Renilla kollikeri, protein content, Arg-phe-amide
characteristics and possible neurological function. |

GRIMMELIKHUIJZEN, C. J. P. AND A. GROEGER. 1987. Isolation of the neuropeptide pGlu-Gly-Arg-Phe-amide
from the pennatulid Renilla kollikeri. Federation of European Biochemical Societies, Letters
211(1):105—108.

GRoBER, M. S. 1990a. Luminescent flash avoidance in the nocturnal crab Portunus xanthusii. 1. The effects of
luminescence and mechanical stimulation on heart rate. Journal of Experimental Biology 148:415—426.
[Renilla kollikeri, luminescence, flashes effect on crustacean heart rate. ]

. 1990b. Luminescent flash avoidance in the nocturnal crab Portunus xantusii. 2. Cardiac and visual
responses to variations in simulated luminescent flashes. The Journal of Experimental Biology
148:427-428. [Renilla kollikeri.]

HapI, N. 1994. Sekilas Tentang Penalaut (Pennatulacea) (A Glimpse of the Pennatulacea). Oceana 9(2):17—22
[In Indonesian with an English abstract. ]

HAECKEL, E. 1904. Kunstformen der Nature. Verlag des Bibliographischen Instituts, Leipzig and Vienna. [Also
published by Dover Publications, New York, 1974 as “Art Forms In Nature,” 100 plates; pennatulaceans,
plate 19: Stvlatula, Virgularia, Umbellula, Renilla, Pteroeides, and Scleroptilum.]

HAGIWATA, S., S. YOSHIDA, AND M. YOSHII. 1981. Transient and delayed potassium currents in the egg cell
membrane of the coelenterate, Renilla koellikeri. Journal of Physiology (Cambridge) 318:123—141.
[Physical properties of tissues and oocyte cell membrane. ]

HAMID, A. 1931. On Virgularia gracillima (K6lliker) from the Chilka Lake. Records Indian Museum Calcutta
33:483-487. [Virgularia gracillima.]

HAMILTON, D. 1958. An Eocene Sea-pen from Dunedin, New Zealand. Palaeontology 1(3):226—230. [An extinct
sea pen: Bensonularia spatulata from the Eocene of New Zealand. |

HANTZSCHEL, W. 1958. Oktokoralle oder Lebensspur? Mitteilungen aus dem Geologisch-Paladontogischen
Institut der Universitat Hamburg 27:77—87. [Extinct sea pens: Pennatulites longespicata?, Protovirgularia
dichotoma?, Virgularia presbytes?|

Harpy, A. 1965. The open sea: Its natural history, Part 2: Fish and fisheries. Houghton Mifflin Company, Boston.
322 pp. [Pennatula phosphorea.|

. 1967. Great waters. Collins. [Antarctic octocorals, pp. 372-373; Umbellula.]

HARMON, A. C., H. W. JARRETT, AND M. J. CORMIER. 1984. An enzymatic assay for calmodulin based on plant
NAD kinase activity. Analytical Biochemistry 141(1):168—178. [Renilla reniformis, biochemical tech-
niques. |

Hart, R.C., J.C. MATTHEWS, K. Hori, AND M. J. CORMIER. 1979. Renilla reniformis bioluminescence:
luciferase-catalysed production of nonradiating excited states from luciferin analogues and elucidation of
the excited state species involved in energy transfer to Renilla green fluorescent protein. Biochemistry
18(11):2204—2210.

HARTING, P. M. AND J. VAN DER HOEVEN. 1861. Organisch Voorwerp (Crinillum). Verslagen van de gewone
vergaderingen der Afdeeling Natuurkunde, Amsterdam 11:286.

HARTMAN, O. 1960. Systematic account of some marine invertebrate animals from the deep basins off Southern
California. Allan Hancock Pacific Expedition 22(2):69—2 15. [Coelenterates, beginning on page 76: includes
Distichoptilum verrillii. |

HARVEY, E. N. 1917. Studies on bioluminescence. 6. Light production by a Japanese pennatulid Cavernularia
habereri. American Journal of Physiology 42:349-358.

——. 1920. The nature of animal light. Monographs of Experimental Biology. J. B. Lippincott Company,
Philadelphia and London. 182 pp. [Bioluminescence in Cavernularia and Pennatula, pp. 74 and 103.]

WILLIAMS: INDEX PENNATULACEA 15

. 1940. Living light. Princeton University Press. 328 pp. [Pennatulaceans, pp. 48-53; Renilla sp. and
Pennatula sp., pp. 274-275; Pennatula, Veretillum, Cavernularia, Funiculina, Umbellula, Leioptilus (=
Ptylosarcus), and Pteroeides contain luminescent species; p. 50.]

. 1952. Bioluminescence. Academic Press, New York. 649 pp. [Pp. 168—180; a detailed history of the
literature pertaining to bioluminescence research in the Octocorallia including the following pennatulacean
taxa: Renilla amethystina—considered a synonym of R. muelleri by Zamponi and Pérez (1995:23);
Cavernularia, Veretillum, Funiculina, Umbellula, Virgularia, Stylatula, Pennatula, Ptilosarcus, and
Pteroeides; a quoted passage from this book appears above in Historical Account — Pre-Linnean Period. |

HASAMA, B. 1943. Uber die Biolumineszenz des Plocamophorus tilesii Bergh sowie der Cavernularia habereri
Moroff im Aktionsstrombild sowie 1m histologischen Bild. Cytologia 13:146—154.

HASTINGS, J. W. 1968. Bioluminescence. Annual Review of Biochemistry 37:597-630.

. 1983. Chemistry and control of luminescence in marine organisms. Bulletin of Marine Science
33(4):818-828. [Renilla; protein metabolism; luciferin oxidation to produce luminescence, ionic control. }

HASTINGS, J. W. AND J. G. MorRIN. 1969. Calcium-triggered light emission in Reni//a. A unitary biochemical
scheme for coelenterate bioluminescence. Biochemical and Biophysical Research Communications
37:493-498. [Renilla koellikeri.]

HEEZEN, B. C. AND C. D. HOLLISTER. 1971. The face of the deep. Oxford University Press, London, New York,
Toronto. 659 pp. [Umbellula.]

HERDMAN, W. A. 1913. Spolia Runiana, I: (Funiculina). Journal of the Linnean Society of London, Zoology
323103:

. 1913a. “Phosphorescence” of Pennatulida. Nature London 91:582. [Pennatula phosphorea and Funi-

culina quadrangularis.]

. 1913b. Spolia Runiana, I: Funiculina quadrangularis (Pallas) and the Hebridean Diazona violacea,

Savigny. Journal of the Linnean Society of London, Zoology 32:163—172. [Funiculina quadrangularis.]

. 1914. Spolia Runiana, II: Funiculina quadrangularis (Pallas); variation in Ascidia; and records of
various rare invertebrata. Journal of the Linnean Society of London, Zoology 32:269-285. [Funiculina
quadrangularis, Virgularia mirabilis, Pennatula phosphorea.|

HERKLOTS, J. A. 1858. Notices pour servir a |’etude des polypiers nageurs ou pennatulides. Bijdragen tot de
Dierkunde 7:1—31. [Original descriptions of the genera Scytalium and Pteroeides.|

. 1863. Descriptions de deux expeéces nouvelles de pennatulides des Mers de la Chine. Nederlandsch
Tijdschrift voor de Dierkunde 1:31—34.

HERRING, P. J., ed. 1978. Bioluminescence in action. Academic Press, London. 570 pp. [Stv/atula, Pennatula,
Pteroeides, Renilla, Acanthoptilum, Ptilosarcus, Cavernularia, Virgularia, and Veretillum.]

HERRING, P. J. 1991. Observations on bioluminescence in some deep-water anthozoans. Hydrobiologia
216-217:573—579. [Distichoptilum gracile, Funiculina quadrangularis, Umbellula huxleyi.]

HICKSON, S. J. 1883. On the ciliated groove (siphonoglyphe) in the stomodaeum of the Alcyonarians. Philo-
sophical Transactions of the Royal Society of London 174(3):693—705.

. 1890. Preliminary report on a collection of Alcyonaria from Port Philip. Proceedings of the Royal

Society of Victoria, Melbourne (new series) 2(2):136—140.

. 1894. A revision of the genera of the Alcyonaria Stolonifera, with a description of one new genus and

several new species. Transactions of the Zoological Society of London 13(9):325—347.

. 1900. The Alcyonaria and Hydrocorallinae of the Cape of Good Hope. Marine Investigations in South

Africa 1:67—96. [Virgularia schultzei as Virgularia reinwardti, Actinoptilum molle as Cavernularia elegans

and Cavernularia obesa.|

. 1903a. Presidential Address to Zoological Section of the British Association. Report of the seventy-third

meeting of the British Association for the Advancement of Science, Southport 1903:672.

. 1903b. Polymorphism in the Pennatulida. Report of the seventy-third meeting of the British Association

for the Advancement of Sciences, Southport 1903:688.

. 1903c. Report of the Southport meeting of the British Association. (Polymorphism in Pennatula

murrayi). Science 68:614. [Pennatula murrayi.]

. 1904. The Alcyonaria of the Cape of Good Hope. Part II. Marine Investigations in South Africa

3:211—239. [Anthoptilum grandiflorum.}

74 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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. 1905. The Alcyonaria of the Maldives. Part III. The families Muriceidae, Gorgonellidae, Melitodidae,
and the genera Pennatula, Eunephthya. In The Fauna and Geography of the Maldive and Laccadive
Archipelagoes, J. Stanley Gardiner, ed. 2(4):807—826. [Pennatula murrayi, Suvadiva, p. 823.]

. 1906. Coelenterata: Anthozoa and Actinozoa-general characters-Alcyonaria. /n The Cambridge Natural

History , S. F. Harmer and A. E. Shipley, eds. Macmillan and Co., London. 1:326—364.

. 1907. Coelenterata I.-Alcyonaria. National Antarctic (Discovery) Expedition, Natural History 3:1—15.

British Museum.

. 1909. Coelenterata: Anthozoa. /n The Cambridge Natural History , S. F. Harmer and A. E. Shipley, eds.

Macmillan and Co., London. 1:326411.

. 1911. Ona specimen of Osteocella septentrionalis (Gray). Memoirs and Proceedings of the Manchester

Literary and Philosophical Society 55(3)(No. 23):1—15. [Possibly Halipteris willemoesi.]

. 1914. On the Sagitta marina of Rumphius. Zoologischer Anzeiger 44(10):471.

. 1916. The Pennatulacea of the Siboga Expedition, with a general survey of the order. Siboga Expeditie

Monographs 14, Livr. 77:1—265. [Anthoptilum malayense, Chunella biflora, Echinoptilum roseum, Echi-

noptilum elongatum, Echinoptilum asperum, Echinoptilum minimum, Gyrophyllum sibogae, Kophobelem-

non pauciflorum, Protoptilum celebense, Pteroeides timorense, Pteroeides malayense, Sclerobelemnon
gravieri, Scleroptilum elongatum, Sclerobelemnon magniflorum, Scytalium balssii, Umbellula weberi,

Veretillum malayense, Virgularia rubra, Virgularia roulei nom. nov. for V. elegans.]

. 1921. On some Alcyonaria in the Cambridge Museum. Proceedings of the Cambridge Philosophical

Society 20(3):366—373. [Cavernularia darwinii.]

. 1922. On two sea-pens from West Australia. The Percy Sladen Trust Expedition to the Abrolhos Islands

(Indian Ocean). Journal of the Linnean Society of London, Zoology 35:21—23. [Veretillum malayense,

Pteroeides sp.? juv.]

. 1930a. Some alcyonarians from the Eastern Pacific Ocean. Proceedings of the Zoological Society of

London 1930(14):209—227. [Virgularia galapagensis. |

. 1930b. On the classification of the Alcyonaria. Proceedings of the Zoological Society of London

1930(1):229-252.

. 1936. Darwin’s Cavernularia. Nature London 137(3474):909. [Veretillum binghami synonynous with

Cavernularia darwinii var. binghami.]|

. 1937. The Pennatulacea. Scientific Reports. Scientific Reports of the John Murray Expedition,

1933-1934 4(5):109—130. [Cavernularia orientalis, Cavernularia darwinii, Cavernularia chuni, Cavernu-

laria malabarica, Chunella gracillima, Funiculina quadrangularis, Pennatula inflata, Protoptilum cy-

aneum, Pteroeides sp.? juv., Scytalium spendens, Scytalium martensii var. magnifolia, Scytalium sarsii,

Umbellula huxleyi, Umbellula spicata, Umbellula validiviae, Umbellula kollikeri, Umbellula pellucida,

Umbellula rigida, Umbellula purpurea, Umbellula elongata, Umbellula radiata, Umbellulid pendula,

Umbellula indica, Umbellula jordani, unidentified pennatulid. }

. 1940. The Gorgonacea with notes on two species of Pennatulacea. Scientific Reports of the John Murray
Expedition, 1933-1934 6:267-317. [Anthoptilum inermis, Kophobelemnon pauciflorum.]

HOARE, R. AND E. H. WILSON. 1977. Observations on the behaviour and distribution of Virgularia mirabilis O.F.
Muller (Coelenterata: Pennatulacea) in Holyhead Harbour, Anglesey. European Marine Biology Sympo-
sium (No. 11):329-337.

HOCHBERG, F. G. AND J. C. LIUBENKOV. 1998. Class Anthozoa: Subclass Octocorallia—Orders Stolonifera and
Pennatulacea. /n Taxonomic atlas of the benthic fauna of the Santa Maria Basin and the western Santa
Barbara Channel, Vol. 3: The Cnidaria, P. V. Scott and J. A. Blake, eds. Santa Barbara Museum of Natural
History, Santa Barbara, California. 150 pp. [Taxonomy and distribution of sea pens of southern California,
pp. 67-112: Kophobelemnon affine, Distichoptilum gracile, Funiculina parkeri, Stachyptilum superbum,
Umbellula magniflora, Anthoptilum grandiflorum, Halipterus californica, Pennatula californica, Ptilosar-
cus gurneyi, Virgularia agassizii, Virgularia californica, Acanthoptilum album, Acanthoptilum gracile,
Stylatula elongata, Stylatula gracilis. |

HOLTHUIS, L. B. 1993. History of the carcinological collections of the Rijksmuseum van Natuurlijke Historie,
Leiden, Netherlands (1820-1950). Crustacean Issues, Rotterdam 8:225—242. [Biographical notes on Jan
Adrianus Herklots. ]

———, 1995. 1820-1958 Rijksmuseum van Natuurlijke Historie. Nationaal Natuurhistorisch Museum, Leiden.
172 pp.[Biographical notes on Jan Adrianus Herklots. ]

WILLIAMS: INDEX PENNATULACEA ie)

HonptT, M.-J. D’. 1984a. Pteroeides (Octocorallia, Pennatulacea) de Nouvelle-Caledonie. Bulletin du Museum
National D’ Histoire Naturelle, Section A: Zoologie Biologie et Ecologie Animales 6(1):3—29.

. 1984b Contribution a la connaissance de certains genres de la famille Veretillidae. (Pennatulacea).
Description de Cavernulina grandiflora n. sp. et de Lituaria valenciennesi nom. nov. Bulletin du Museum
National D’ Histoire Naturelle, Section A: Zoologie Biologie et Ecologie Animales 6(3):625-640.

Honjo, I. 1940. Beitrage zur nervenmuskelphysiologie der kolonienbildenden tiere. I. Die Peristaltik von
Cavernularia. Annotationes zoologicae Japonenses 19(4):301—308.

Hori, K. AND M. J. CORMIER. 1973. Structure and chemical synthesis of a biologically active form of Renilla
(sea pansy) luciferin. Proceedings of the National Academy of Sciences of the United States of America
70(1):120-123.

Hori, K., Y. NAKANO, AND M. J. CORMIER. 1972. Studies on the bioluminescence of Renilla reniformis. Il.
Location of the sulfate group in luciferyl sulfate. Biochemica et Biophysica Acta 256(3):638-644.

Hori, K., J. E. WAMPLER, J.C. MATTHEWS, AND M. J. CORMIER. 1973. Identification of the product excited
states during the chemiluminescent and bioluminescent oxidation of Renilla (Sea pansy) luciferin and
certain of its analogs. Biochemistry 12(22):44634468.

HORNELL, J. 1922. Some commensals of Indian Alcyonarians and crabs. Journal of the Bombay Natural History
Society 28(4):926—936. [The pennatulacean genera: Cavernularia, Pennatula, Virgularia.]

HOWELL, B. J. 1947. Eocene Alcyonaria in New Jersey. Abstract in: Bulletin of the Geological Society of
America 58:1195. [Two unnamed species of Graphularia, both considered by Shapiro and Ramsdell
(1965:3) as synonyms of Graphularia ambigua.|

Howson, C. M. AND L. M. DAvies. 1991. Marine nature conservation review surveys of Scottish sea lochs. A
towed video survey of Loch Fyne, Vol. 1: Report. Nature Conservancy Council CSD Report 1189.
[Virgularia mirabilis. ]

HUANG, C. L. AND MIR, G. N. 1972. Toxicological and pharmacological properties of sea pansy Renilla mulleri.
Journal of Pharmacological Sciences 60:1620—22.

HUBRECHT, A. A. W. 1885. On a new pennatulid from the Japanese Sea. Proceedings of the Zoological Society
of London 1885:512—518. [Abstract in J.R. Microscopical Society [2]6:81; new genus and species
Echinoptilum macintoshii.]

HUMAN, V. L. 1973. Albinism in three species of marine invertebrates from southern California. California Fish
and Game 59:89—92. [Renilla koellikeri.]

HuMEs, A. G. 1978. Lichomolgid copepods (Cyclopoida), with two new species of Doridicola, from sea pens
(Pennatulacea) in Madagascar. Transactions of the American Microscopical Society 97(4):524-539.
[ Virgularia juncea and Pteroeides oblongum; associations with crustaceans, Malagasy Republic. ]

HUTTON, F. W., ed. 1904. Index faunae Novae Zealandiae. Dulau and Co., London. 372 pp. [Virgularia
gracillima, Protocaulon molle, Leptoptilum gracile.|

HUXLEY, T. H. 1907. A manual of the anatomy of invertebrated animals. D. Appleton and Company, New York.
596 pp. [Octocorals, pp. 143-145; Veretillum, Pennatula, Virgularia.|

HYMAN, L. H. 1940. The Invertebrates, Vol. 1: Protozoa through Ctenophora. McGraw-Hill, New York. 726 pp.
[Pennatulacean biology, pp. 557-565; quoted passage in Historical Account—The Early to Mid-Twentieth
Century, above. ]

IMAFUKU, M. 1973. On some physiological aspects in the daily rhythmic activity of the sea-pen, Cavernularia
obesa Valenciennes. Publications of the Seto Marine Biological Laboratory 20:43 1-454.

. 1975. Peristalsis in the monopolypid stage of the sea-pen Cavernularia obesa Valenciennes. Publica-

tions of the Seto Marine Biological Laboratory 22(1-4):195-216.

. 1976. On the mechnism of the activity rhythm of the sea-pen, Cavernularia obesa Valenciennes.

Publications of the Seto Marine Biological Laboratory 23(1—2):1—17.

. 1980. Activity rhythm of the sea-pen, Cavernularia obesa Valenciennes, under temperature and light
cycles. Publications of the Seto Marine Biological Laboratory 25(1—4):119—130.

IMAHARA, Y. 1991. Report on the Octocorallia from the Ryukyu Islands of Japan. Bull. Inst. Oceanic Research
and Development, Tokai Univ. (1991)11/12:59-94. [Cavernulina orientalis, Sclerobelemnon burgeri,
Virgularia rumphii, Struthiopteron caledonican. |

IMPERATO, F. 1599. Dellhistoria natvrale, di Ferrante Imperato . . . libri XXVIII, nella qvale ordinatamente se
tratta della diuersa condition di miniero, e pietre. Con alcune historie di piante, & animali; sin “hora non

76 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 2

date in luce... .C. Vitale, Napoli. 791 pp. [Also published in 1672, Presso Combi and La Nou, Venetia
696 pp.; pennatulacean bioluminescence. ]

IVESTER, M. S. 1977. Nematocyst differentiation in the anthozoan Renilla reniformis (Pallas). Transactions of
the American Microscopical Society 96(2):238-247.

IVESTER, S. AND D. DUNKELBERGER. 1971. Ultrastructure of the autozooid of the sea pansy, Renilla reniformia
(Pallas). American Zoologist 1 1:695—696.

JAHN, W. 1970. Umbellulidae distribution in the Atlantic. Nature, London 225:1068—1069. [Umbellula.]

JAwWorSKI, E. 1939. Geographische Rassen und Standortsmodifikationen bei Seefedern. Thalassia 3(7):1—24.
[Pennatula phosphorea var. rubella, Pennatula phosphorea var. variegata forma typica, Pennatula
phosphorea var. candida, Pennatula phosphorea var. californica, Pennatula phosphorea var. longispi-
nosa.|

JENKINS, R. J. F. 1985. The enigmatic Ediacaran (Precambrian) genus Rangea and related forms. Paleobiology
11(3):336-355. [Taxonomic position of Precambrian fossils that resemble sea pens, placed in order
Rangeomorpha. |

. 1992. Functional and ecological aspects of Ediacaran assemblages. /n Lipps, Origin and early evolution
of the Metazoa, Vol. 10: Topics in Geobiology, J. H. and P. W. Signor, eds. Plenum Press, New York. 570
pp. [Precambrian Vendian fossils resembling pennatulaceans. ]

JENKINS, R. J. F. AND J. G. GEHLING. 1978. A review of the frond-like fossils of the Ediacara assemblage. Records
of the South Australian Museum 17:347-359. [Precambrian Vendian fossils resembling pennatulaceans. ]

JOHNSON, M. E. AND H. J. SNOOK. 1935. Seashore animals of the Pacific Coast. MacMillan Co., New York, 659
pp. [Pennatulacea, pp. 89-94; Ptilosarcus quadrangularis, Renilla amethystina, Stylatula elongata, Sty-
latula gracile.|

JOHNSTON, G. 1847. A history of the British zoophytes, 2nd ed. 2 vols. John van Voorst, Paternoster Row,
London. 488 pp. [Virgularia mirabilis, Pavonaria quadrangularis, Pennatula phosphorea, Virgularia
mirabilis, Pavonaria quadrangularis.]

JONES, S. 1960. Note on animal associations. I. A porcellanid crab on the sea pen, Pteroeides esperi Herklots.
Journal of the Marine Biological Association of India 1(2):(for 1959):178—179. [Ecological interaction:
decapod crustacean and sea pen. }

JONES, H. P., J. C. MATTHEWS, AND M. J. CoRMIER. 1979. Isolation and characterization of Ca** -dependent
modulator protein from the marine invertebrate Renilla reniformis. Biochemistry 18(1):55—60. [Renilla;
enzymes; activator; modulator protein. ]

JUNGERSEN, H. F. E. 1888a. Om Bygningen og Udviklingen af Kolonien hos Pennatula phosphorea. Videnska-
belige Meddelelser fra den Naturhistoriske forening 1 Kjobenhavn for Aaret 1888, 40:154—181.

_1888b. Uber Bau und Entwicklung der Kolonie von Pennatula phosphorea L. Zeitschrift fiir wissen-

schaftliche Zoologie 47:626-649.

. 1904. Pennatulida. Danish Ingolf-Expedition 5(1):1—95. [Taxonomic descriptions: Halipteris christii

p. 45 Faroe Islands, Pavonaria finmarchica p. 39 Iceland, Protoptilum denticulatum p. 59, Protoptilum

carpenteri p. 51, Protoptilum thomsoni p. 55, Pennatula aculeata p. 11, Pennatula prolifera p. 18,

Virgularia cladiscus p. 79, Virgularia tuberculata p. 33, Umbellula lindahlii p. 75, Umbellula encrinus

p. 79.]

. 1905. Some results from the Ingolf Expeditions. Oversigt over det Kongeligh Danske videnskabernes

Selskab forhandlinger 2:127—135. [Pennatula aculeata.]

. 1907. Pennatulida. Expedition Antarctique Belge, Rapports Scientifiques: Zoologie. 12 pp. [Umbellula

carpenteri. |

. 1915. Alcyonaria, Antipatharia og Madreporaria. Conspectus Faunae Groenlandicae. Meddelelser om

Groénland 23:1156—1212.

. 1917. The Alcyonaria of East Greenland. Medd. Grfnl. 43(18):485—S03. (also published separately as
Danmark Exp. til Gronlands Nordéstkyst 1906-1908 3(18):487—-S03). [Umbellula encrinus, Virgularia
affinis.|

KAPLAN, E. N. 1982. A field guide to coral reefs, Caribbean and Florida—Peterson field guide series. Houghton
Mifflin Company, Boston. 289 pp. [Octocorals, pp. 86—95 including Renilla. ]

—_——. 1988. A field guide to southeastern and Caribbean seashores—Cape Hatteras to the Gulf Coast, Florida,
and the Caribbean—Peterson field guide series. Houghton Mifflin Company, Boston. 425 pp. [Octocorals,
pp. 210-211 including Renilla reniformis and Renilla muelleri.]

WILLIAMS: INDEX PENNATULACEA 77

KARKHANIS, Y. D. AND M. J. CORMIER. 1971. Isolation and properties of Renilla reniformis luciferase, a low
molecular weight energy conversion enzyme. Biochemistry 10:317—326.

KASTENDIEK, J. 1975a. The behavior, distribution, and predatory-prey interactions of Renilla kéllikeri. Ph. D.
dissertation, Zoology, University of California, Los Angeles, 235 pp.

. 1975b. The role of behaviour and interspecific interactons in determining the distribution and abundance

of Renilla kollikeri Pfeffer, a member of a subtidal sand bottom community. Dissertation Abstracts

International (B)36(6):2604—2605 [Abstract only. ]

. 1976. Behavior of the sea pansy Renilla kollikeri Pfeffer (Coelenterata: Pennatulacea) and its influence

on the distribution and biological interactions of the species. Biological Bulletin of the Marine Biological

Laboratory Woods Hole 151(3):518—537.

. 1982. Factors determining the distribution of the sea pansy Renilla kollikeri, in a subtidal sand-bottom
habitat. Oecologia 52(3):340—347. [Population density and vertical distribution; influencing biological and
physical factors, northern Pacific]

KEIFER, P. A., K. L. RINEHART JR., AND I. R. HOOPER. 1986. Renillafoulins, antifouling diterpenes from the sea
pansy Renilla reniformis (Octocorallia). Journal of Organic Chemistry 51(23):4450-4454.

KELLER, N. B., F. A. PASTERNAK, AND D. V. NAUMOV. 1975. Bottom deep-sea Coelenterata from the Caribbean
Sea and Gulf of Mexico (from material from the 14th expedition of the ‘Akademik Kurchatov’ ). /n Scientific
studies Caribbean Sea, Gulf of Mexico and adjacent waters. Transactions of the P. P. Shirshov Institute of
Oceanology (Trudy Instituta okeanologii im. P. P. Shirshova) 100:147—159. [In Russian with English
summary. |

KERSTITCH, A. 1989. Sea of Cortez marine invertebrates—a guide for the Pacific Coast, Mexico to Ecuador. Sea
Challengers, Monterey. 114 pp. [Octocorals, pp. 29-30; Prilosarcus undulatus, Stvlatula elongata. |

KILIAS, R. 1993. Stamm Cnidaria—Nesseltiere. Pp. 115—219 in Urania Tierreich. Wirbellose | (Protozoa bis
Echiurida). Urania, Leipzig. 666 pp. [Umbellula.]

KINOSHITA, K. 1912. Hasshasango-rui no keitohassei oyobi sono bunrui (Phylogeny and classification of
Alcyonaria). Dobutsugaku zasshi, Tokyo 24:433-441.

KITTREDGE, J. S., D. G. SIMONSON, E. ROBERTS, AND B. JELINEK. 1962. Free amino acids of marine invertebrates.
Pp. 176-186 in Amino acid pools: distribution, formation and function of free amino acids, J. T. Holden,
ed. Elsevier, Amsterdam. 815 pp. [Renilla.]

Kner, R. 1858. Uber Virgularia multiflora, n. sp. aus der Familie der Seefedern, Pennatulina. Verhandlungen
der Kaiserlich-K6niglichen Zoologisch-Botanischen Gesellschaft 1858:295—298.

Koch, G. VON. 1878. Notiz tiber die Zooide von Pennatula. Zoologischer Anzeiger 1:103—104.

. 1889. Kleinere Mittheilungen tiber Anthozoen. |. Zwei Entwicklungsstadien von Preroides spinulosus.

Morphologisches Jahrbuch 15: 646-649.

. 1890. Kleinere Mittheilungen tiber Anthozoen. 2. Terminalpolyp und zooid bei Pennatula und
Pteroides. -3. Einstiilpung der Tentakel bei Rhizoxenia rosea und Asteroides_calycularis. Morphologisches
Jahrbuch 16: 396-400.

KOLLIKER, R. A. VON. 1865. Icones histiologicae oder Atlas der vergleichenden Gewebelehre. Zweite Abthei-
lung. Der feinere Bau der hoheren Thiere. Erstes Heft. Die Bindesubstanz der Coelenteraten. Leipzig, Verlag
von Wilhelm Engelmann: 87-181. [Reviewed by Verrill, 1866.]

. 1869-72. Anatomisch-Systematische Beschreibung der Alcyonararien. Erste Abtheilung. Die Pen-
natuliden. Abhandlungen von der Senckenbergischen naturforschenden Gesellschaft 7:1 1 1-255; 487-602;
8:85-275 [Also issued in 1872 with consecutive pagination, 1-485; Acanthoptilum pourtalesi, Acanthop-
tilum agassizi, Argentella, Bathyptilum carpenteri, Cavernularia obesa, Cavernularia defilippii, Cavernu-
laria elegans, Cavernularia glans, Cavernularia haimii, Cavernularia luetkeni, Cavernularia
valenciennesi, Clavella australasiae, Crinillum seidenburgi, Crispella, Godeffroyia gen. nov. elegans sp.
nov., p. 116 Siam; Kophobelemnon burgeri, Kophobelemnon clavatum, Kophobelemnon leuckarti, Ko-
phobelemnon stelliferum, Leioptilum Gray, char. emend., p. 139; Leioptilum undulatum = Ptilosarcus
sinuosus, Lituaria phalloides, Pennatula phosphorea var. lancifolia, Pennatula mollis, Pennatula targionii,
Policella manillensis, Policella australis, Protoptilum carpenteri, Protoptilum smithi, Protoptilum thom-
soni, Leioptilum grayi, Pteroeides speciosum, p. 54 locality unknown; Preroeides nigrum,p. 56 locality
unknown; Prteroeides hartingii, p. 58 locality unknown; Pteroeides lacazii spinosum; p. 60 Australia;
Pteroeides lacazii molle, p. 60 Sumatra; Pteroeides multiradiatum, p. 63 Penang; Pteroeides schlegelii, p.
65 Japan; Pteroeides hystrix, p. 72 locality unknown; Preroeides longepinnatum, p. 74 locality unknown;

78

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 2

Pteroeides caledonican, p. 75 New Caledonia; Pteroeides diibenii, p. 77 Mozambique; Pteroeides gracile,
p. 77 Philippines; Preroeides brachycaulon, p.78 Philippines; Pteroeides breve, p. 78 Philippines;
Pteroeides pellucidum, p. 80 Philippines; Pteroeides manillense, p. 81 Philippines; Pteroeides brevira-
diatum, p. 82 Indian Ocean; Pteroeides tenerum, p. 84 locality unknown; Preroeides flavidum, p. 85 Java;
Pteroeides ferrugineum, p. 86 Java; Pteroeides herklotsii, p. 87 locality unknown; Pteroeides sparmannii,
p. 89 locality unknown; Pteroeides imbricatum, p. 89 Singapore; Pteroeides acuminatum, p. 91 Turan?,
Pteroeides lugubre, p. 94 New Holland; Pteroeides crassum, p. 95 Singapore; Pteroeides westermannii, p.
96 locality unknown; Preroeides bleekerii, p. 96 locality unknown; Pteroeides fusco-notatum, p. 99 Chinese
Sea; Pteroeides latissimum, p. 102 China; Pteroeides macandrewi, Pteroeides durum, Pteroeides grayi,
Pteroeides vogti, Pteroeides cornaliae, Pteroeides clausi, Pteroeides griseum, Pteroeides steenstrupi,
Pteroeides jukesi, Pteroeides oblongum, Pteroeides lacazii, Pteroeides pancerii, Pteroeides schlegeli,
Pteroeides crispum, Pteroeides spinosum, Pteromorpha, Ptilosarcus Gray, char. emend. p. 144; Renilla
edwardsi, Renilla deshayesi, Renilla muelleri, Renilla amethystina, Renilla patula, Renilla peltata, Renilla
sinuata, Renilla australasiae, Renilla africana, Renilla reniformis, Renilla mollis, Sarcophyllum gen. nov.,
p. 116 australe sp. nov., p. 121 Australia; Sarcoptilus grandis, Sclerobelemnon schmeltzi, Scytalium
mertensi, Stylatula lecazii, Stylatula kinbergi, Stylatula darwini = Virgularia patachonica, Stylatula
antillarum, Stylobellemnon pusillum, Veretillum cynomorium vat. astyla, Veretillum stimpsoni, Veretillum
baculatum, Virgularia lvungmanii, Virgularia strupi, Virgularia loveni, Virgularia rumphii.]

_1871b. Ueber den Bau der Renillen. Verhandlungen der Physikalisch-medicinischen Gesellschaft in
Wiirzburg 2:108—111. [Also printed in Annals and Magazine of Natural History (4)7:307—309; Renilla. ]

. 1872. Morphologie und Entwickelungsgeschichte des Pennatulidenstammes nebst allgemeinen Be-
trachtungen zu Descendenzlehre. Christian Winter, Frankfurt am Main. 87 pp. [A separate printing from
Kolliker, 1869-72. ]}

. 1874. Uber den Bau und die systematische Stellung der Gattung Umbellularia; eine vorlaufige
Mittheilung. Verhandlungen der Physikalisch-medizinischen Gesellschaft zu Wirzburg 8:13—18 (or
92-957).

. 1875. Die Pennatulide Umbellula und zwei neue Typen der Alcyonarien. /n Festschrift zur Feier des
fiinfundzwanzigjahrigen Bestehens der Physikalisch-medizinischen Gesellschaft in Wurzburg. 23 pp.
[Umbellula thomsoni, Umbellula encrinus and Umbellula_lindahli.|

. 1880. Report on the Pennatulida dredged by H. M. S. Challenger during the years 1873-1876. Report
of the Scientific Results of the Voyage of H. M. S. Challenger during the years 1873—76. Zoology 1(2):1-41.
[Taxonomic descriptions: Pteroeides esperi, Pteroeides breviradiatum, Pennatula naresi, Pennatula pear-
cevi, Pennatula murrayi, Virgularia_bromleyi, Scytalium sarsi, Stachyptilum gen. nov. macleari sp. nov.,
Anthoptilum gen. nov. thomsoni sp. nov., Anthoptilum murrayi, Kophobelemnon ferrugineum, Umbellula
durissima, Umbellula giintheri, Umbellula thomsoni, Umbellula leptocaulis, Umbellula simplex, Umbellula
huxleyi, Umbellula carpenteri, Umbellula magniflora, Protocaulon gen. nov. molle sp. nov., Microptilum
gen nov. willemdesi sp. nov., Leptoptilum gen. nov. gracile, Protoptilum aberrans, Protoptilum sp.,
Trichoptilum gen. nov. brunneum sp. nov., Scleroptilum gen. nov. grandiflorum sp. nov., Scleroptilum
durissimum, Renilla miilleri, Cavernularia obesa, Lituaria pohalloides, Clavella australasiae.|

KoLosvary, G. 1949. The Eocene corals of the Hungarian transdanubian province. Bulletin of the Hungarian

Geological Society 79(5—8):141—242. (In Hungarian with English summary) [Extinct sea pen Graphularia
sp. from the Eocene of Hungary. |

Koo, S. Y. 1935. On a new pennatulid (Lituaria) from Amoy. Natural Science Bulletin of the University of

Amoy 1(2):157—164. [Original description of Lituaria amoyensis.]

———. 1940. Some sea pens (Pennatulacea) from Amoy Island. China Journal 32(3):113—118. [Taxonomic

descriptions: Cavernularia habereri, Lituaria amoyensis, Pteroeides chinense, Pteroeides dofleini, Virgu-
laria gustaviana, Virgularia reinwardti.|

KOREN, J. AND D.C. DANIELSSEN. 1847. Virgularia christii, n. sp. Nyt Magazin for naturvidenskabernes

5(3):269-271 [Also published in Oken’s /sis 1848:208—209. ]

———. 1856. Virgularia Christii K. and D. Jn Fauna Littoralis Norvegiae, M. Sars, J. Koren, and D.C.

Danielssen, eds. 2:91—93.

——. 1874. Bidrag til de ved den norske Kyst levende Pennatuliders Naturhistorie. Nyt Magazin for

Naturvidenskabernes 12:422—427. [Taxonomic descriptions: Cladiscus gracilis, Lygomorpha sarsi, Batea
abyssicola var. smaragdina, Batea (Stylatula) elegans, Virgularia affinis = Virgularia glacialis and Virgu-

WILLIAMS: INDEX PENNATULACEA 79

laria steenstrupi KOll., Pennatula aculeata var. rosea, Pennatuladistorta, Ptilella grandis (Ehrbg) =

Pennatula borealis Sars.

. 1877. Contribution to the natural history of the Pennatulidae living on the Norwegian coast. /n Fauna

Littoralis Norvegiae, M. Sars, J. Koren, and D. C. Danielssen, eds. 3:82—102. [Ptilella grandis, Pennatula

aculeata var. rosea, Pennatuladistorta, Pennatula phosphorea var. variegata, Virgularia affinis, Dubenia

(=Batea olim) abyssicola var. smaragdina, Dubenia elegans, Lygomorpha sarsi, Cladiscus gracilis,

Umbellularia groenlandica|

. 1883. Nye Alcyonider, Gorgonider og Pennatulider tilhorende Norges Fauna. Bergens Museum.

Bergen, John Griegs Bogtrykkeri. 38 pp. (New Alcyonarians, Gorgonids and Pennatulids of the Norwegian

Seas. Bergen.) [Abstract in: Journal of the Royal Microscopical Society (2)4:239; Goendul new genus,

Goenduleae new family. ]

. 1884. Pennatulida. The Norwegian North-Atlantic Expedition, 1876-1878 4:1,83.

KOROTNEFF, A. 1887. Zur Anatomie und Histologie des Veretillum. Zoologischer Anzeiger 10:387-390.
[Anatomy, physiology, cell biology: Veretillum—differentiated nervous system, special cell elements
causing phosphorescence, sexual and sexless polyps. |

KORSCHELT, E. 1936. Vergleichende Entwicklungs-geschichte der Tiere, volume I:158—220. Jena. [Renilla sp.]

KOZLOFF, E. N. 1974. Keys to the marine invertebrates of Puget Sound, the San Juan Archipelago, and adjacent
regions. University of Washington Press, Seattle and London: 1—226.

———. 1983. Seashore life of the northern Pacific Coast—An illustrated guide to northern California, Oregon,
Washington, and British Columbia. University of Washington Press, Seattle and London: 1-370. [Prilosar-
cus gurneyi.|

. 1987. Marine invertebrates of the Pacific northwest. University of Washington Press, Seattle and

London. 511 pp.[Pennatulacea, key to the species from B. C., Wash., and Ore., pp. 70—71.]

. 1990. Invertebrates. Saunders College Publishing, Philadelphia. 866 pp. [Pennatulaceans, pp. 145—149;
Veretillum cynomorium, Renilla, Ptilosarcus gurneyi, Virgularia.]

KRAMP, P. L. 1932. The Godthaab Expedition, 1928. Alcyonaria, Antipatharia, and Madreporaria. Meddelelser
om Gronland 79(1):1—20. [Anthoptilum grandiflorum, Pennatula_grandis, Umbellula encrinus, Virgularia
tuberculata. |

. 1933. The Scoresby Sound Committee’s 2nd East Greenland Expedition in 1932 to King Chrstian IX’s

Land. Coelenterata, Ctenophora and Chaetognatha. Meddelelser om Groénland 104(11):1—20. [Virgularia

tuberculata. |]

. 1950. Polydyr. Pp. 500-517 in Braestrup, F. W. Vort Lands Dyre, Liv 2. Gyldendalske Boghandel,
Kobenhavn. [Pennatula phosphorea.]|

KREISS, P. AND M. J. CORMIER. 1967. Inhibition of Renilla reniformis bioluminescence by light: Effect on
luciferase and its substrates. Biochimica et Biophysica Acta 141:181—183.

KRUKENBERG, C. F. W. 1887. Die physiologischen Eigenthumlichkeiten des Leuchtvermogens bei Preroides
griseum L. In Vergleichend-Physiologische Studien. Carl Winter’s Universitats-Buchhandlung, Heidelberg
(2 Reihe) 4:83—105. [Bioluminescence in Preroeides.]

. 1888. Das Leuchten des Rotes Meeres. /n Vergleichend-Physiologische Studien. Carl Winter’s Univer-
sitats-Buchhandlung, Heidelberg (2 Reihe) 4:117—142, frontispiece.

KUHN, O. 1949. Lehrbuch der Palaozoologie. Stuttgart: 1-326. [Coelenterata, pp. 25-50; extinct sea pen
Graphularia desertorum.}

KUKENTHAL, W. 1902a. Diagnosen neuer Alcyonarien aus der Ausbeute der Deutschen Tiefsee-Expedition. 2.
Eine neue Familie der Penatuliden. Zoologischer Anzeiger 25(668):299-303. [Amphianthus abyssorum,
Chunella gracillima, Chunellidae, p. 302.]

. 1902b. Diagnosen neuer Umbelluliden aus der Ausbeuter der deutschen Tiefsee-Expedition. Zoologis-

cher Anzeiger 25(679):593—597. [Umbellula encrinus var. antarctica (Bovet Island), Umbellula pellucida,

Umbellula spicata, Umbellula valdiviae, Umbellula rigida, Umbellula kéllikeri (East Africa). ]

. 1903. Uber eine neue Nephthyidengattung aus dem siidatlantischen Ocean. Zoologischer Anzeiger

26:272-275. [Amphiacme.]

. 1910. Pennatuliden der Deutschen Tiefsee-Expedition. Zoologischer Anzeiger 36(2/3):51—S8. [Acti-

noptilon, Kophobelemnon heterospinosum, Pennatula inflata, Pennatula phosphorea var. antarcticum,

Protoptilum cyaneum, Virgularia schultzei. |

80 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 2

. 1911. Aleyonarien von der Aru- und Kei-Inseln nach den Sammlungen von Dr. H. Merton. Abhandlun-

gen der Senckenbergischen Naturforschenden Gesellschaft 33; 307-346.

. 1912a. Die Alcyonaria der deutschen Siidpolar-Expedition 1901—1903. Jn Deutsche Siidpolar Expedi-

tion 1901—1903, Erich von Drygalski, ed. 13 Zoologie 5 (3):289-349.

. 1912b. Der Stammbaum der Seefedern. Verhandlungen der international Zoologischen Kongress Jena

8:563—570.

. 1913a. Uber die Alcyonarienfauna Californiens und ihre tiergeographischen Beziehungen. Zoologische

Jahrbucher (Systematik) 35(2):219-270. [Funiculina parkeri, Pennatula phosphorea var. californica,

Leioptilum quadrangulare, Virgularia bromelyi, Stvlatula elongata, ?Acanthoptilum gracile, ?Acanthopti-

lum pourtalesii, Acanthoptilum album, Acanthoptilum scalpellifolium, Acanthoptilum annulatum, Pavon-

aria californica, Pavonaria willemoesi, Pavonaria sp. juv., Pavonaria sp. juv., Stachyptilum superbum,

Stachyptilum dofleini, Anthoptilum grandiflorum, Umbellula magniflora, Umbellula huxleyi, Umbellula

loma, Renilla amethystina. |

. 1913b. Alcyonaria des Roten Meeres. /n Expeditionen S. M. Schiff “Pola” in das Rote Meer. Zoolo-

gische Ergebnisse 29. Denkschriften der Kaiserlichen Akademie der Wissenschaften Mathematisch-Natur-

wissenschaftliche Klasse 89:1—31.

. 1914. Zur systematik der Umbelluliden. Zoologischer Anzeiger 43(13):630—632. [Umbellula spp.

tabulated. }

. 1915a. Das System der Seefedern. Zoologischer Anzeiger 45(6):284-287.

. 1915b. Pennatularia. Das Tierreich. Verlag von R. Friedlander und Sohn, Berlin. 43:1—132.

. 1921. Versuch eines natiirlichen Systems der Octokorallen. Sitzungberichte der preussischen Akademie
der Wissenschaften Mathematisch-Naturwissenschaftliche Klasse 1921(4):82—102.

. 1923-25. Octocorallia. Jn Handbuch der Zoologie, Thilo Krumbach, ed. 1:690—769.

KUKENTHAL, W. AND H. BROCH. 1910. System und Stammesgeschichte der Seefedern. Zoologischer Anzeiger
36:222—230. [Classification and phylogeny of Pennatulacea. ]

. 1911. Pennatulacea. Wissenschaftliche Ergebnisse der deutschen Tiefsee-Expedition “Valdivia” 13(1)
Lieferung 2:113—576. [Actinoptilum molle, Amphiacme abyssorum, Anthoptilum grandiflorum, Cavernu-
laria chuni, Cavernularia clavata, Cavernularia elegans, Cavernularia glans, Cavernularia habereri,
Cavernularia Liitkeni, Cavernularia obesa, Cavernularia pusilla, Cavernulina cylindrica, Chunella grac-
illima, Chunella quadriflora, Echinoptilum echinatum, Funiculina armata, Funiculina quadrangularis,
Kophobelemnon affine, Kophobelemnon heterospinosum, Leioptilum sinuosum, Leioptilum Verrilli, Pa-
vonaria finmarchica, Pennatula aculeata, Pennatula fimbriata, Pennatula inflata, Pennatula Murrayi,
Pennatula naresi, Pennatula pearceyi, Pennatula phosphorea, Pennatula phosphorea forma candida,
Pennatula phosphorea forma variegata, Pennatula phosphorea forma rebella, Pennatula phosphorea
forma antarctica, Pennatula rubra, Pennatula sp., Protoptilum cyaneum, Pteroeides bankanense,
Pteroeides breviradiatum, Pteroeides durum, Pteroeides esperi, Pteroeides griseum, Pteroeides Jung-
erseni, Pteroeides Lacazei, Pteroeides latissimum, Pteroeides lusitanicum, Pteroeides sagamiense,
Pteroeides Sparmanni, Pteroeides sp. aff. Diibeni, Pteroeides tenerum, Renilla amethystina, Renilla
edwardsi, Renilla Miilleri, Renilla reniformis, Sarcophyllum grande, Sclerobelemnon schmeltzi, Sclerop-
tilum grandiflorum, Scytalium Martensi, Scytalium sarsi, Stachyptilumsuperbum, Struthiopteron caledoni-
can, Stvlatula elegans, Stylatula elongata, Stylatula sp. aff. darwini, Umbellula Huxleyi, Umbellula
antarctica, Umbellula pellucida, Umbellula rigida, Umbellula spec., Umbellula spicata, Umbellula valdi-
viae, Umbellula Kéllikeri, Veretillum cynomorium, Virgularia affinis, Virgularia gustaviana, Virgularia
halsiceptrum, Virgularia juncea, Virgularia mirabilis, Virgularia Reinwardti, Virgularia Rumphii, Virgu-
laria Scultzei, Virgularia sp. aff. Bromleyi; plates 13-17 are perhaps the most accurate, beautifully detailed,
and realistic of any published color renderings of sea pens ever produced. |

KUMANO, M. 1937. Japanische Pennatuliden. Hakubutsugakuzasshi 35:246. [In Japanese. ]

KUMAR, S.,M. HARRYLOCK, K. A. WALSH, M. J. CORMIER, AND H. CHARBONNEAU. 1990. Amino acid sequence
of the Ca2+-triggered luciferan binding protein of Renilla reniformis. FEBS (Federation of European
Biochemical Societies) Letters 268(1):287—290.

LACAZE-DUTHIERS, H. DE. 1865. Des sexes chez les Alcyonaires. Comptes rendus hebdomadaires des Seances
de |’ Académie des sciences 60:840—843. [Sexual reproduction in Pennatula.]

WILLIAMS: INDEX PENNATULACEA 81

. 1887. Sur le développement des Pennatules (Pennatula grisea) et les bonnes conditions biologiques
que présente le laboratoire Arago pour les études zoologiques. Comptes rendus hebdomadaires des Séances
de 1’ Académie des sciences, Paris 104(8):463-469.

. 1891. Note sur la présence du Kophobelemnon dans les eaux de Banyuls. Comptes rendus hebdo-
madaires des Séances de |’ Académie des sciences, Paris 1 12(23):1294—1297.

LAM, C. N. H., J. O. T. JENSEN, AND D. F. ALDERDICE. 1982. Preliminary study of low gamete viability in adult
chum salmon (Obcorhyncus keta) held in sea pens at Deserted Creek, Hisnit Inlet, B. C. Canadian Technical
Report of Fisheries and Aquatic Sciences (no. 1133):1—49.

LAMARCK, J. B. P. A. 1801. Systeme des animaux sans vertebres 432 pp. Author, Paris.

. 1816. Histoire naturelle des animaux sans vertebres. Verdiere, Paris. 568 pp.

. 1836. Histoire naturelle des animaux sans vertebres . .. Deuxieme édition. Revue et augmentee . . . par
MM... P. Deshayes et H. Milne Edwards. Tome deuxieme. Histoire des polypes. J. B. Bailliere, Paris.
684 pp.

LANCASTER, CAPT. 1601. /n A general history and collection of voyages and travels arranged in systematic order,
forming a complete history of the origin and progress of navigation, discovery, and commerce, by sea and
land, from the earliest ages to the present time, 18 vol., R. Kerr, ed. 1824. W. Blackwood, Edinburgh. [In
volume 8, p. 119, is a description of behavior in intertidal sea pens at Sombrero Island, Indonesia—probably
a species of the genus Virgularia,; Darwin (1860:101) quotes Lancaster (in Kerr) as follows, “found a small
twig growing up like a young tree, and on offering to pluck it up it shrinks down to the ground, and sinks,
unless held very hard. On being plucked up, a great worm is found to be its root, and as the tree groweth in
greatness, so doth the worm diminish; and as soon as the worm is entirely turned into a tree it rooteth in the
earth, and so becomes great. This transformation is one of the strangest wonders that I saw in all my travels:
for if this tree is plucked up, while young, and the leaves and bark stripped off, it becomes a hard stone
when dry, much like white coral: thus is this worm twice transformed into different natures. Of these we
gathered and brought home many.”’}

LANGTON, R. W., E. W. LANGTON, R. B. THEROUX, AND J.R. UZMANN. 1990. Distribution, behavior and
abundance of sea pens, Pennatula aculeata, in the Gulf of Maine. Marine Biology 107(3):463—-469.
[Pennatula aculeata.]

LAUBIER, L. 1972. Lamippe (Lamippe) bouligandi sp. nov., copépode parasite d’octocoralliaire de la mer du
Labrador. Crustaceana 22(3):285—293. [Anthoptilum grandiflorum.]}

LEDGER, P. W. AND S. FRANC. 1978. Calcification of the collagenous axial skeleton of Veretillum cynomorium
Pall (Cnidaria: Pennatulacea). Cell and Tissue Research 192(2):249-266.

LENHOFF, H. M., L. MUSCATINE, AND L. V. DAvis. 1971. Experimental coelenterate biology. University of
Hawaii Press. [Pennatulacea: calcite skeleton of spicules with an axial skeleton in some species, p. 228.]

LEPECHIN, I. 1781. Novae Pennatulae et Sertulariae species descriptae. Acta Academiae scientiarum imperialis
petropolitanae, pro Anno 1778, Pars posterior 2:236—238.

LEUCKART, F. S. 1841. Einige Bemerkungen tiber die Familie der Halopteriden oder Seefedern, insbesondere
iiber des genus Veretillum, und eine von mir Veretillum calvatum benannte Art. /n Zoologische Bruchstiicke,
part 2. Rieger, Stuttgart. 130 pp.

LEUNIS, J. 1886. Synopsis der Thierkunde (Zweiter Band). Hannover. [Pennatulaceans, pp. 1057—1060. ]

LEWIN, R. 1984. Alien beings here on Earth. Science (6 Jan. 1984):39. [The taxonomic status of Precambrian
pennatulacean-like fossils. ]

LIGHT, S. F. 1921. Notes on Philippine Alcyonaria, Part VI: New Philippine Pennatularia (sea pens) of the genus
Lituaria. Philippine Journal of Science 19(2):247-255. [Lituaria kukenthali, Lituaria philippinensis,
Lituaria molle, Lituaria breve.]

LIGHTBOWN, C. M. 1918. The dorsal mesenteric filaments in the siphonozooids of Pennatulacea. Memoirs and
Proceedings of the Manchester Literary and Philosophical Society 62(1):(No.4):1—20.

LINDAHL, J. 1874a. Om Pennatulid-Slagtet Umbellula Cuv. Kungliga Svenska vetenskapsakademiens handlingar
13(3):1—22. (Preliminary accounts are given in Annals and Magazine of Natural History (4)13:258, and
Skandinaviske Naturforskeres Moede, Forhandlingar 11:377-379). [Crinillum seidenburgi, Umbellula
miniacea pallida.]|

. 1874b. Om twanne polypstockar af slagtet Umbellula Cuv. Skandinaviske Naturforskeres Moede,

Forhandlingar 11:377-379.

. 1874c. Umbellula from Greenland. Annals and Magazine of Natural History (4)13:258.

82 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 2

LINNAEUS, C. 1758. Systema naturae. Editio decima, reformata |:1—824. Salvii, Holmiae.

. 1767. Systema naturae. Editio duodecima, reformata 1(2):533—1327. Holmiae.

Liu, X. 1981. Metazoa fossils from Mashan group near Jixi, Heilongjiang. Bulletin of the Chinese Academy of
Geological Sciences 3(1):71—83. [In Chinese with English summary; Mashania new genus of pennatu-
lacean-like fossil from the Proterozoic of China. ]

. 1983. Some new materials of metazoan fossils from Mashan Group, Heilongjang Province. Bulletin of
the Shenyang Institite of Geology and Mineral Resources (No. 7):1—8. [Precambrian fossils resembling sea
pens. ]

LOpPEZ-GONZALEZ, P. J., J.-M. GILI, AND G. C. WILLIAMS. In press. On some veretillid pennatulaceans from the
eastern Atlantic and western Pacific Oceans (Anthozoa: Octocorallia), with a review of the genus Caver-
nularia Valenciennes and descriptions of the new taxa. Journal of Zoology.

LORENZ, W. W., R. O. MCCANN, M. LONGIARU, AND M. J. CORMIER. 1991. Isolation and expression of acDNA
encoding Renilla reniformis luciferase. Proceedings of the National Academy of Sciences of the United
States of America 88(10):4438-4442.

LUTHER, W. AND K. FIEDLER. 1961. Die Unterwasser fauna der Mittelmeerkésten. Ein Taschen buch fir Biologen
und Naturfreunde. Verlag Paul Parey, Hamburg and Berlin. 251 pp. [Coelenterata, pp. 203—222; Preroeides
griseum (a synonym of Pteroeides spinosum), and Veretillum cynomorium.]

Lutz, P. E. 1986. Invertebrate zoology. Addison Wesley Publishing Company, Reading, Mass. 734 pp.
[Ptilosarcus, p. 137]

LYDEKKER, R. n.d. The new natural history, Part 6. Merrill and Baker, New York. [Octocorals, pp. 511—517;
Umbellula.]|

LYKE, E. B. 1965. The histology of the sea pansies, Renilla reniformis (Pallas) and Renilla kéllikeri (Pfeffer),
with a note on the fine structure of the latter species. Ph.D. dissertation, Zoology, University of Wisconson,
Madison. 247 pp.

MACGINITIE, G. E. 1938. Notes on the natural history of some marine animals. The American Midland Naturalist
19:207-219 [Stylatula elongata, Renilla koellikeri.|

MACGINITIE, G. E. AND N. MACGINITIE. 1968. Natural history of marine animals, 2nd ed. McGraw-Hill, New
York. 523 pp. [Leioptilus guerneyi, Leioptilus quadrangularis, Leoioptilus undulatus, Renilla koéllikeri,
Stylatula elongata, Pteroeides sp.|

MACKIE, A. M. 1987. Preliminary studies on the chemical defenses of the British octocorals A/cyonium digitatum
and Pennatula phosphorea. Comparative Biochemistry and Physiology (A) 86(4):629-632.

MADSEN, F. J. 1948. The zoology of East Greenland. Octocorallia. Meddelelser om Grenland 122(2):1—22.
[Anthoptilum grandiflorum, Umbellula encrinus, Virgularia glacialis, Virgularia tuberculata. |

MAGNUS, D. B. D. 1966. Zur Okologie einer nachtaktiven Flachwasser Seefeder (Octocorallia, Pennatularia) im
Roten Meer. Ver6ffentlichungen Instituts fiir Meeresforschung in Bremerhaven, Sonderband 2:369-380.

MALARODA, R. 1951. Il Lattorfiano del Monteccio di Costozza (Colli Berici). Parte Prima: I macrofossili.
Memorie del Museo civico di Storia naturale di Verona 2 (for 1949—50):147-210. [Extinct sea pens:
Pavonaria? singularis, Graphularia incerta, Graphularia nigra from the Oligocene of Italy. ]

MALECKI, J. 1982 (1983). Bases of Upper Cretaceous octocorals from Poland. Acta palaeontologica polonica
27(1-4):65—75. [Octobasis; new genus of extinct Virgulariidae from the Upper Cretaceous of Poland. ]

MANGOLD, E. 1910. Die Produktion von Licht. Pp. 225-392 in Handbuch der vergleichende Physiologie 3 (2nd
half), H. winterstein, ed. Jena. [Bioluminescence. |

MANUEL, R. L. 1981. British Anthozoa. Synopses of the British fauna (new series) 18:1—241. [Pennatulacea;
key to species from British Isles. ]

Marion, A. F. 1906. Etude des Coelentérés atlantiques recueillis par la Commission des dragages de |’aviso
LeTravailleur druant les campagnes 1880-1881. Pp. 103-151 in Expédition scientifique du Travailleur et
du Talisman. Marson, Paris. [Bathypenna elegans, Umbellula ambigua.]

MaRISCAL, R. N. 1974. Nematocysts. Pp. 129-178 in Coelenterate Biology, Reviews and new perspectives, L.
Muscatine and H. M. Lenhoff, eds. Academic Press, New York.

——_—. 1979. The spined nematocysts of octocorals. American Zoologist 19(3):784.

MARISCAL, R. N. ANDC. H. BIGGER. 1977. Possible ecological significance of octocoral epithelial ultrastructure.
Preceedings Third International Coral Reef Symposium, Rosenstiel School of Marine and Atmopheric
Science, University of Miami 1977:127—133. [Renilla mulleri.]

WILLIAMS: INDEX PENNATULACEA 83

MARKS, M.H., R. S. BEAR, AND C. H. BLAKE. 1949. X-ray diffraction evidence of collagen-type protein fibers
in the Echinodermata, Coelenterata and Porifera. Journal of Experimental Zoology, Philadelphia
3(1):55—78. [Balticina: protein fibres. ]

MARSHALL, A. M. 1883a. On the polymorphism of the Alcyonaria. Report of the British Association for the
Advancement of Science 1883:529. [Abstract in the Journal of the Royal Microscopical Society (2)3(6):855
and Nature 29:580; Pennatula phosphorea and Umbellula gracilis: zooids bearing one tentacle. ]

. 1883b. Report on the Pennatulida dredged by H. M. S. Triton. Transactions of the Royal Society of
Edinburgh 32(1):119-152. [Virgularia tuberculata p. 129; Umbellula gracilis p. 142; listed as Marshall,
1887 by Bayer (1981a,1996) and Williams (1990, 1993b, 1997c); pp. 148-150, a discussion of the
phylogeny and affinities of deep-water sea pens. |

MARSHALL, A. M. AND G. H. FOWLER. 1887. Report on the Pennatulida dredged by H. M. S. ‘Porcupine.’
Transactions of the Royal Society of Edinburgh 33(2):453—464. [Pennatula phosphorea var. candida from
the Faeroe Channel, p. 456; Deutocaulon hystricis, from the Faeroe Channel, p. 461.]

. 1888. Report on the Pennatulida of the Mergui Archipelago, collected for the Trustees of the Indian
Museum, Calcutta, by Dr. John Anderson, F. R. S., Superintendent of the Museum. Journal of the Linnean
Society of London, Zoology 21(132):267-286. [Abstract in J. R. Microscopical Society 1889(4):529;
Virgularia prolifera, Policella tenuis, Pteroeides elegans, Pteroeides lacazii, Pteroeides chinense,
Pteroeides esperi, Virgularia rumphii, Virgularia prolifera, Cavernularia obesa, Lituaria phalloides,
Policella manillensis.]

MARSHALL, A. M. AND W. P. MARSHALL. 1882. Report on the Pennatulida collected in the Oban dredging
excursion of the Birmingham Natural History and Microscopical Society, July 1881. The Herald Press,
Birmingham. 81 pp.

MARSHALL, N. B. 1979. Developments in deep-sea biology. Blandford Press, Poole and Dorset. 566 pp.
[Octocorals, pp. 179-183; Umbellula, Chunella, Kophobelemnon, Pavonaria, Funiculina; the following
passage is noteworthy, “Umbellula is one of the classic animal types of the deep-sea floor, and so are the
stalked sea-lilies (crinoids), both converging in their palm-like forms—forms that look well designed for
passive suspension feeding. Bent over by currents, they look like palms in a breeze, though . . . the sea-lilies
are not altogether pliant . . . . Sea-pens luminesce when stimulated, the light (blue-green, yellowish or violet)
spreading over the colony from the site of the stimulus.”’]

MARSHALL, W. P. 1895. Virgularia mirabilis. Journal of the Marine Biological Association of the United
Kingdom (new series) 3(4):335—336.

MARSIGLI, L. F. 1725. Histoire physique de la Mer. Amsterdam. [French translation by Leclerc of the 1711 work
first published in Italian. ]

MATERN, U. 1984. Geschichte und Mechanismus der Biolumineszenz. Biologie in unserer Zeit 14(5):140—149.
[Renilla reniformis, history and mechanisms of bioluminescence. ]

MATHER, P. AND I. BENNET, eds. 1993. A coral reef handbook—a guide to the geology, flora and fauna of the
Great Barrier Reef, 3rd ed. Surrey Beatty and Sons Pty Limited, New South Wales. 264 pp. [Octocorals,
pp. 74-79; Cavernularia.]|

MATTHEws, J.C., K. Hori, AND M. J. CORMIER. 1977. Purification and properties of Renilla reniformis
luciferase. Biochemistry (American Chemical Society) 16(1):85—91. [Bioluminescence. ]

May, W. 1899. Alcyonarien. Jn Hamburger Magalhaensische Sammelreise 4:1—22. [Virgularia kophameli.]

. 1900. Die arktische, subarktische und subantarktische Alcyonaceen-fauna. Fauna Artica 1:381—408.

McCoy, F. 1890. Prodromus of the Zoology of Victoria, Zoophites. Publication of the Government of Victoria,
Melbourne. [Sarcoptilus grandis. |

McDONALD, G. R. AND J. W. NYBAKKEN. 1978. Additional notes on the food of some California nudibranchs
with a summary of known food habits of California species. The Veliger 21(1):110—119. [Ecological
associations, prey/predator interactions: Virgularia, Leioptilus, and Renilla—as food of the nudibranchs
Armina, Tritonia, Tochuina, and Hermissenda. |

MEGLITSCH, P. A. 1972. Invertebrate Zoology, 2nd ed. Oxford University Press, New York. 834 pp. [Octocorals,
pp. 129-140; Renilla, Renilla reniformis.]

MILNE, L. AND M. MILNE. n.d. Invertebrates of North America. A Chanticleer Press Edition, Doubleday and
Co., New York. 249 pp. [Anthozoa pp. 49-61; Stvlatula, Renilla kollikeri, Leioptilus guerneyi.]

84 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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MILNE EDWARDS, H. AND J. HAIME. 1850. A monograph of the British fossil corals, Part 1: Introduction; corals
from the Tertiary and Cretaceous formations. Palaeontographical Society, London. [New genus and species
Graphularia wetherelli. |

. 1857. Histoire naturelle des coralliaires ou polypes proprement dits. A la Librairie Encyclopedique de
Roret, Paris. 1:1—326.

MING, C. L. 1993. A guide to the dangerous marine animals of Singapore. Singapore Science Center, Singapore.
160 pp. [Pp. 61-63, color photographs of Virgularia sp. and Pteroeides sp.|

MIYAJIMA, K. 1897. Umi shaboten (Veretillum) no seitaiteki kansatsu. Dobutsugaku zasshi 9(107):367-371.
[Ecological observations on Veretillum.]

. 1900. Umi shaboten (Cavernularia obesa Val.). Dobutsugaku zasshi 12(142):280-287.

MopeeR, A. 1786. Slagtet Sjépenna (Pennatula). Kongl. Vetenskaps Academiens nya handlingar, Stockholm

7:267.

MOLANDER, A. R. 1929. South and West African Octactiniae in the Gothenburg Natural History Museum.

Goteborgs Kungliga vetenskaps- och vitterhetssamhalles handlingar (B)1(7):1—16.

Moore, A. R. 1926. On the nature of inhibition in Pennatula. American Journal of Physiology 76:112—115.

Moore, H. B. 1937. Marine fauna of the Isle of Man. Proceedings and Transactions of the Liverpool Biological

Society 50:38—57. [Virgularia mirabilis. |

Moore, R. C., ed. 1956. Treatise on Invertebrate Paleontology, Part F: Geological Society of America and

Unversity of Kansas Press, Lawrence, Kansas. 498 pp. [Pennatulacea by F. M. Bayer, pp. 224-230.]

MorI, S. 1943a. Daily rhythmic activity of the sea-pen, Cavernularia obesa Valenciennes. |. Observations in
nature. Dobutsugaku zasshi 55:285—291. [In Japanese. ]

. 1943b. Daily rhythmic activities of the sea-pen, Cavernularia obesa Valenciennes. 2. Activities under

constant darkness and constant illumination. Dobutsugaku zasshi 55:247—253. [In Japanese. ]

. 1944a. Daily rhythmic activities of the sea-pen, Cavernularia obesa Valenciennes. 3. Controlling of

the activity by light. Dobutsugaku zasshi 56:81—85. [In Japanese. ]

. 1944b. Daily rhythmic activities of the sea-pen, Cavernularia obesa Valenciennes. 4. Obvervations of

the activity in winter. Dobutsugaku zasshi 56:86—90. [In Japanese. ]

. 1944c. Daily rhythmic activities of the sea-pen, Cavernularia obesa Valenciennes. 5. Activities under

constant illumination and constant darkness in winter and influence of water temperature. Dobutsugaku

zasshi 56:91—95. [In Japanese. ]

. 1944d. Daily rhythmic activities of the sea-pen, Cavernularia obesa Valenciennes. 6. Analysis of the

endogenous rhythm. Dobutsugaku zasshi 56:96—100. [In Japanese. ]

. 1944e. Daily rhythmic activities of the sea-pen, Cavernularia obesa Valenciennes. 7. Overnight change

of reaction to light. Dobutsugaku zasshi 56:2 1—24. [In Japanese. ]

. 1945. Daily rhythmic activities of the sea-pen, Cavernularia obesa Valenciennes. 8. Endogenous daily

rhythmic activity. Kyodai Seiri Seitai 19:1—21. [In Japanese. ]

. 1947. Daily rhythmic activities of the sea-pen, Cavernularia obesa. 9. Activities when kept long under

constant darkness. Physiology and Ecology, University of Kyoto 1:8—14. [In Japanese. ]

. 1948. Daily rhythmic activities of the sea-pen, Cavernularia obesa. 10. Two kinds of stimulus which

control the activities. Physiology and Ecology, University of Kyoto 2:34-38. [In Japanese. ]

. 1949. Daily rhythmic activities of the sea-pen, Cavernularia obesa. 11. Controlling of the activity by

light (2). Physiology and Ecology, University of Kyoto 3:32—37 [In Japanese. ]

. 1950. Daily rhythmic activities of the sea-pen, Cavernularia obesa. 12. Conclusions—problems on

relations among environments, behaviors and internal physiological conditions. Physiology and Ecology,

University of Kyoto 4:14—20. [In Japanese. }

. 1960. Influence of environmental and physiological factors on the daily rhythmic activity of a sea-pen.
Cold Spring Harbor Symposia on Quantitative Biology 25:333-344.

———. 1994. Daily rhythmic activities of the sea-pen, Cavernularia obesa Valenciennes. 19. Further consid-
erations on the mechanism of the circadian rhythmic activity. Publications of the Seto Marine Biological
Laboratory 36(4):267—276.

Mort, S. AND Y. ONDO. 1957. Daily rhythmic activity of the sea-pen, Cavernularia obesa Valenciennes. 15.
Controlling the activity by light (3). Publications of the Seto Marine Biological Laboratory 6:79—98.

WILLIAMS: INDEX PENNATULACEA 85

MorI, S. AND H. TANASE. 1973. Studies on the daily rhythmic activity of the sea-pen, Cavernularia obesa
Valenciennes. XVIII. Ontogenetic development of the daily rhythmic activitiy. Publications of the Seto
Marine Biological Laboratory 20:455—467.

Morin, J. G. 1974. Coelenterate bioluminescence, Pp. 397-438 in Coelenterate biology: reviews and new
perspectives, L. Muscatine and H. M. Lenhoff, eds. Academic Press, New York. 501 pp. [Bioluminescence
in Renilla.]

. 1976. Probable functions of bioluminescence in the Pennatulacea (Cnidaria, Anthozoa), pp. 629-638,

In Mackie, G. O. (ed.), Coelenterate ecology and behavior. New York: Plenum. 744 pp. [Stvlatula elongata,

Renilla kollikeri, Ptilosarcus gurneyi, Acanthoptilum gracile, pennatulaceans; bioluminescence; function

and kinetics. ]

. 1998. Living light from cnidarians. Wings 21(2):14—17.

MorofF, T. 1902a. Aus der Miinchener Sammlung. 3. Einige neue Pennatuliden aus der Miinchener Sammlung.
Zoologische Anzeiger 25(678):579-S82. [Pteroeides sagamiense, Pteroeides rhomboidale, Pennatula
phosphorea var. longispinosa, Pennatula murrayi var. japonica, Pennatula americana, Ptilosarcus quad-
rangularis, Cavernularia habereri, Virgularia rigida Philippines. ]

. 1902b. Studien tiber Octocorallien. I. Ueber die Pennatulaceen des Miinchener Museums; II. Ueber
einige neue Gorgonaceen aus Japan. Zoologische Jahrbucher (Abteilung fiir Systematik) 17:363-410.
[Pteroeides sagamiense, Pteroeides rhomboidale, Pteroeides griseum, Pteroeides manillense, Pteroeides
breviradiatum, Pennatula phosphorea, Pennatula phosphorea var. longispinosa, Pennatula americana,
Pennatula murrayi var. japonica, Pennatula fimbriata, Ptilosarcus quadrangularis, Virgularia mirabilis,
Virgularia rigida, Pavonaria dofleini, Pavonaria finmarchica, Pavonaria californica, Acanthoptilum
scalpelifolium, Funiculina quadrangularis, Kophobelemnon stelliferum, Kophobelemnon leucharti, Renilla
reniformis, Veretillum cynomorium, Cavernularia habereri.]

Morton, S. G. 1830. Synopsis of the organic remains of the ferruginous sand formation of the United States;
with geological remarks. American Journal of Sciences, First Series 17:274-295. [Brief and unfigured
original description of Belemnites ambiguus, considered by Roemer (1880) to be a member of the extinct
sea pen genus Graphularia; see Shapiro and Ramsdell (1965). ]

. 1834. Synopsis of the organic remains of the Cretaceous group of the United States. Illustrated by
nineteen plates. To which is added an appendix, containing a tabular view of the Tertiary fossils hitherto
discovered in North America. Key and Biddle, Philadelphia. 88 pp. [A more complete and illustrated
description of Belemnites? ambiguus, so designated by Morton since he was at that time uncertain of its
generic placement; Roemer (1880) considered this taxon to be a member of the extinct sea pen genus
Graphularia; see Shapiro and Ramsdell (1965). ]

MOsELEY, H. N. 1872. On the substance exhibited at the British Association, Brighton, by Mr. P. L. Sclater, and
stated to be the ossified notochord of a fish. Nature 6:432. [Axis of Halipteris willemoesi. |

. 1877. On the coloring matter of various animals and especially of deep-sea forms dredged by H. M. S.
Challenger. Quarterly Journal of Microscopical Science 17:1—23. [Alcyonarian bioluminescence; a quote
from this work is as follows: “All of the Alcyonarians dredged by the Challenger in deep water were found
to be brilliantly phosphorescent when brought to the surface and their phosphorescence was found to agree
in the manner of exhibition with the same conditions as are observed in the case of shallow water relatives.” ]

Moss, E. L. 1873. Description of a virgularian actinozoon, from Burrard’s Inlet, British Columbia. Proceedings
of the Zoological Society of London 1873:730—732. [Pavonaria blakei.]

. 1878. On specimens of Osteocella septentrionalis. Journal of the Royal Dublin Society 1875:241.

MULLER, F. 1866. Ein Wort tiber die Gattung Herklotsia, J. E. Gray. Archiv fiir Naturgeschichte 30:352-358.
[Herklotsia, Renilla edwardsii.]

MULLER, O. F. 1776. Zoologiae Danicae Prodromus, seu animalium Daniae et Norvegiae indigenarum charac-
teres, nomina, et synonyma imprimis popularium. Havniae-Vermes, Cellulana.

MuscrAvVe, E. M. 1909. Experimental observations on the organs of circulation and the powers of locomotion
in Pennatulids. Quarterly Journal of Microscopical Science, London 54:443-48 1. [Pteroeides, Anthoptilum,
circulation, experimental observations. |

MyLilus, C. 1753. Beschreibung einer neuen Gronlandischen Thierpflanze. In einem Sendschreiben an . . . Hrn,
Albrecht von Haller. . .. Andreas Linde, London. 27 pp. [An early account of Umbellula.]

86 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 2

. 1755. An account of a new zoophyte, or animal plant, from Groenland. In a letter to Dr. Albert Haller,
President of the Royal Society of Sciences at Gottingen . . . now translated into English. London: Andreas
Linde, 27 pp. [An early account of Umbellula.]

NAKASONE, Y. AND H.-P. YU. 1987. Note on the Porcellanella triloba White (Crustacea: Decapoda: Porcellani-
dae) from Taiwan. Bulletin of the Institute of Zoology, Academia Sinica 26(1):107—111. [Pteroeides esperi,
commensalism with the crab Porcellanella triloba, commensal description, Taiwan. ]

Naumov, D. V. 1955. Tip Kishetsnopolostnie - Coelenterata, Ushakov, P. V. and others. /n Atlas Bespoz-
vonotsnix dal’ nevostotsnix morei S.S.S.R. Academiia Nauk SSSR, Zoologicheskii Institut, Trudy,
Leningrad 1955:51—68. [Pavonaria finmarchica]

NEALSON, K. H., A. C. ARNESON, AND M. E. HUBER. 1986. Identification of marine organisms using kinetic and
spectral properties of their bioluminescence. Marine Biology (Berlin) 91(1):77—83. [Renilla myriaster,
Stylatula elongata, \uminescence, kinetic and spectral properties use in identification. ]

NELLI, B. 1903. Fossili miocenici del Macigno di Porretta. Bollettino della Societa geologica Italiana
22:181—252. [Exinct sea pen Pennatulites manzonii from the Miocene of Italy. ]

NicoL, J. A.C. 1955a. Observations on luminescence in Renilla (Pennatulacea). Journal of Experimental
Biology 32:299-320.

. 1955b. Nervous regulation of luminescence in the sea pansy Renilla kollikeri. Journal of Experimental

Biology 32:619-635.

. 1955c. Physiological control of luminescence in animals. /n The luminescence of biological systems,

F. H. Johnson, ed. American Association for the Advancement of Science, Washington, D. C.

. 1958. Observations on the luminescence of Pennatula phosphorea, with a note on the luminescence of
Virgularia mirabilis. Journal of the Marine Biological Association of the United Kingdom 37:551—563.

NIEDERMEYER, A. 1911. Studien tiber den Bau von Preroides griseum (Bohadsch). Arbeiten aus den zoologischen
Institut der Universitat Wien und der zoologischen Station in Triest 19(1):99—-164. [Pteroeides griseum.]

_ 1912. Uber den Verschlussmechanismus der Stielporen bei Pennatula und Pteroeides. Zoologischer

Anzeiger 39:190.

. 1913. Uber einige histologische Befunde an Veretillum cynomorium. Zoologischer Anzeiger

43:263-270. [Veretillum cynomorium.]

. 1914. Beitrage zur Kenntnis des Histologischen Baues von Veretillum cynomorium (Pall.). Zeitschrift
fiir wissenschaftliche Zoologie 109:531—590. [Veretillum cynomorium.]

NIELSEN, K. 1914. Moltkia isis, Steenstrup og andre Octocorallia fra Danmarks Kridttidsaflejringer. (Moltkia
isis and some other Octocorallia in the Danish Cretaceous deposits). Mindeskrift i anledning af hundredaaret
for Japetus Steenstrups fodsel, Kobenhavn 18:1—20. [Extinct sea pens, Graphularia groenwalli, Graphu-
laria sulcata, Graphularia irregularis, from the Cretaceous of Denmark. ]

NISHIMURA, S. 1992. Guide to seashore animals of Japan with color pictures and keys, vol. 7. Hoikusha
Publishing, Osaka, Japan. 424 pp. [Color photographs of living animals: Cavernularia obesa, Cavernulina
orientalis, Echinoptilum macintoshi, Sclerobelemnon burgeri, Scytalium martensti, Virgularia gustaviana,
Virgularia halsiceptrum, Virgularia rumphii, Virgularia brochi, Struthiopteron caledonican, Pteroeides
sagamiense, Pteroeides sparmannii.]

Nosre, A. 1931. Contribuigoes para o estudo dos Coelenterados de Portugal. Instituto de Zoologia da
Universidade do Pérto 1931:1—82. [Funiculina quadrangularis, Pennatula phosphorea, Veretillum cyno-
morium, Stylobelemnon pusillum, Pennatula granulosa, Pteroeides griseum, Kophobelemnon stelliferum,
Umbellula thomsoni, Umbellula ambigua. |

NORDGAARD, O. 1905. Hydrographical and biological investigations in Norwegian Fiords. Bergens Museums
aarbog 156-159, 240-244. [Kophobelemnon stelliferum, Pennatula aculeata.]

NorMAN, A. M. 1867. Report of the committee appointed for the purpose of exploring the coasts of the Hebrides
by means of the dredge, Part II: On the Crustacea, Echinodermata, Polyzoa, Actinozoa, and Hydrozoa.
Report of the British Association for the Advancement of Science (Nottingham, 1866) Part 1:193—206.
[Report of Pennatula mollis Alder, 1867, from Scotland; considered a junior synonym of Pennatula
phosphorea by Cornelius and Garfath, 1980:274.]

NUTTING, C. C. 1908. Descriptions of the Alcyonaria collected by the U. S. Bureau of Fisheries steamer Albatross
in the vicinity of the Hawaiian Islands in 1902. Proceedings of the United States National Museum
34(1624):543-601. [Calibelemnon symmetricum, Cladiscus studeri, Pennatula sanguinea, Pennatula

WILLIAMS: INDEX PENNATULACEA 87

flava, Pennatula pallida, Protoptilum wrighti, Trichoptilum attenuatum, Umbellula jordani, Umbellula

gilberti.|

. 1909. Alcyonaria of the California Coast. Proceedings of the United States National Museum

35:68 1—727. [Acanthoptilum album, Acanthoptilum annulatum, Balticina pacifica, Halipteris contorta,

Halisceptrum cystiferum, Ptilosarcus quadrangularis, Stachyptilum quadridentatum, Umbellula loma.]

. 1912. Descriptions of the Alcyonaria collected by the U. S. Bureau of Fisheries steamer “Albatross.”
mainly in Japanese waters, during 1906. Proceedings of the United States National Museum
43(1923):1—-104. [Halisceptrum album, Helicoptilum rigidum, Kophobelemnon hispidum, Pennatula
longistvla, Pennatula rubescens, Pennatula brevipenna, Pennatula inermis, Protoptilum orientale,
Ptilosarcus brevicaulis, Trichoptilum spinosum, Umbellula eloisa.|

OKUTANI, T. 1969. Synopsis of bathyal and abyssal megalo-invertebrates from Sagami Bay and the south off
Boso Peninsula trawled by the R/V Soyo-Maru. Bulletin of Tokai Regional Fisheries Research Laboratory
No. 57:1—61. [Umbellula.]

Omorl, M. 1991. Studies on some relict animals. 6 - Pennatula. Aquabiology (Tokyo) 13(5):345-347. [In
Japanese. |

PALLAS, P. S. 1766. Elenchus zoophytorum sistens generum adumbrationes generaliores et specierum cogni-
tarum succinctas descriptiones cum selectis auctorum synonymis. Hagae Comitum.

. 1787. Charakteristik der Thierpflanzen ... aus dem Lateinischen tibersetzt, und mit Anmerkungen
versehen, von C, F. Wilkens, und nach seinem Tode herausgegeben von Johann Wilhelm Herbst. Niinberg,
Verlegt von der Raspischen Buchhandlung. Zweiter Theil.

PANCERI, P. 1870. Intorno ad una forma non per anco notata negli zooidi delle pennatule. Rendiconto
dell’ Accademia della scienze, Napoli 2:1—S.

. 187 1a. Intorno a due pennatularii, l’uno non per anco trovato nel Mediterraneo, |’altro nuovo nel nostro

Golfo. Rendiconto dell’ Accademia della scienze, Napoli 10(6):113—115.

. 1871b. Gli organi luminosi e la luce delle pennatule. Rendiconto dell’ Accademia della scienze, Napoli

10(10):204-211. [Also published in Archives de zoologie expérimentale et générale 1 :xxv—xxvi; and below

as Panceri, 1872a.]

. 1871c. Gli organi luminosi e la luce delle pennatule. Atti dell’Accademia della scienze, Napoli

5(10):1—46. [Pennatulacean bioluminescence. ]

. 1872a. The luminous organs and light of the Pennatulae. Quarterly Journal of Microscopical Science,

London (new series) 12:248—254. [English translation of Panceri, 1871b.]

. 1872b. Etudes sur la Phosphorescence des Animaux Marins. II. Du siege du mouvement lumineux dans

les Meéduses; III. Organes lumineux et lumiére des Pennatules; VI. Sur un Pennatulaire phosphorescent

encore inconnu dans les environs de Naples (Cavernularia pusilla); 1X. Des organes lumineux et de la
lumiere des Béroidiens. Annales des Sciences Naturelles, sér. 5 (Zoologie)16 (8):1—66. [Bioluminescence
in Pennatula phosphorea and Cavernularia pusilla. |

. 1872c. Intorno ad un pennatulario fosforescente non per anco rinvenuto presso Napoli. Rendiconto
dell’ Accademia della scienze, Napoli 1 1(4):88—90. [Pennatulacean biluminescence. ]

PANI, A. K. AND M. ANCTIL. 1994. Evidence for biosynthesis and catabolism of monoamines in the sea pansy
Renilla koellikeri (Cnidaria). Neurochemistry International 25(5):465—474.

PANTIN, C. F. A. 1950. Behaviour patterns in lower invertebrates. Symposia of the Society for Experimental
Biology 4:175—195.

PARKER, G. H. 1919. The organization of Renilla. Journal of Experimental Zoology 27:499-507.

. 1920a. Activities of colonial animals. 1. Circulation of water in Renilla. Journal of Experimental

Zoology 31:343-367.

. 1920b. Activities of colonial animals. 2. Neuromuscular movements and phosphorescence of Renilla.

Journal of Experimental Zoology 31:475—515.

. 1920c. The phosphorescence of Renilla. Proceedings of the American Philosophical Society
19:171-175.

PASTERNAK, F. A. 1960. The deep-sea Pennatularia from the Bering Sea and Kuril-Kamtschatka Trench. Trudy
Instituta okeanologii, Akademiya nauk SSSR 34:329-335. [In Russian. ]

. 1961a. Some new data on the specific composition and the distribution of deep-sea Pennatularia, genus

Kophobelemnon, in Northern-Pacific. Trudy Instituta okeanologii, Akademiya nauk SSSR 45:240-258. [In

Russian with English summary. ]

88 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 2

. 1961b. Pennatularia (Octocorallia) and Antipatharia (Hexacorallia) obtained by the Soviet Antarctic

Expedition in 1955—1958. Trudy Instituta okeanologii, Akademiya nauk SSSR 46:217—230. [In Russian. ]

. 1962. Pennatularia of the genus Umbellula Cuvier (Coelenterata, Octocorallia) from Antarctic and

Subantarctic Waters. Issledovaniya Fauny Morei 1:105—128 (In Russian). Translated into English in

Biological reports of the Soviet Antarctic Expedition (1955—1958), A. P Andriyashev and P. V. Ushakov,

eds. Vol. 1:107—130. Published for the National Science Foundation, Washington, D.C. by the Israel

Program for Scientific Translations. [Umbellula thomsoni, Umbellula lindahli, Umbellula magniflora.]

. 1964. The deep-sea pennatularians and antipatharians obtained by R/S ‘Vitjaz’ in the Indian Ocean and

the resemblance between the faunas of the pennatularians of the Indian Ocean and the Pacific. Trudy Instituta

okeanologii, Akademiya nauk SSSR 69:183—215. [In Russian with English summary. |

. 1966. On the finding of Kophobelemnon (Eucophobelemnon) stelliferum (O. F. Miiller) in the waters

of south-eastern Australia. Trudy Instituta okeanologii, Akademiya nauk SSSR 81:176—178. [In Russian

with English summary. ]

. 1970. Sea pens (Octocorallia, Pennatularia) of the hadal zone of the Kurile-Kamtaschatka Trench. Trudy

Instituta okeanologii. Akademiya nauk SSSR 86:236-248. (In Russian). Also in: Israel Program for

Scientific Translations, Jerusalem No. 600496:250—263; translated from Fauna of the Kurile-Kamchatka

trench and its environment, V. G. Bogorov, ed. Akademiya Nauk SSSr. Trudy Instituta Okeanologii, P. P.

Shirshova, Izdatel’stvo ‘Nauka’, Moskva, 1970.

. 1975a. Deep-sea pennatularian genus Umbellula from the Caribbean Sea and Puerto-Rican Trench. /n

Scientific studies Caribbean Sea, Gulf of Mexico and adjacent waters. Transactions of the P. P. Shirsov

Institute of Oceanology 100:160—173. [In Russian with English summary; Umbellula.]

. 1975b. New data on the specific composition and distribution of the deep-sea pennatularians (Octoco-

rallia, Pennatularia) of the Peru-Chile region and South Atlantic. Trudy Instituta okeanologii, Akademiya

nauk SSSR 103:101—118. [In Russian with English summary. ]

. 1980. Pennatularia Umbellula encrinus (L.) from the Canadian Basin of the northern Arctic Ocean. Pp.

236-239 in The biology of the Central Arctic Basin, M. E. Vinogradov and I. A. Melnikov, eds. Nauka,

Moscow. 251 pp.

. 1989. On the variability of sea-pens (Octocorallia: Pennatulacea) connected with the transition to the

deep-water existence. Trudy Instituta Okeanologii, Akademii Nauk SSSR 124:165—173. [In Russian with

English Summary. |

. 1993. Pennatularians, gorgonians and antipatharians collected in 43 cruises of R/V “Dimitri Mendelev”
in Argentine Basin, Orkney Trench and in the western part of African-Antarctic Basin. Trudy Instituta
Okeanologii, Akademii Nauk SSSR 127:82-88.

PATTERSON, G. L. J. 1986. Aspects of the zoogeography of some benthic animals in the Rockall Trough.
Proceedings of the Royal Society of Edinburgh (B) 88:316—318. [Umbellula lindahli, distribution and
habitat, ocean trough benthic zone, northeastern Atlantic Ocean. ]

PAVANS DE CECCATTY, M. AND B. BUISSON. 1964a. Le systeme nerveux intra-mésogléen dans les colonies de
Veretillum cynomorium Pall. (Cnidaire, Pennatulidae). Comptes Rendus hebdomadaires des séances de
1’ Académie des sciences, Paris 259(20):361 13613.

. 1964b. Quelques structures de type nerveux du sarcosome des octocoralliaires: Alcyonium digitatum L.

et Veretillum cynomorium Pall. Vie et Milieu 14(4):659—667.

. 1965. Reciprocal behavior of the rachis and peduncle in colonies of Veretillum cynomorium Pall.
American Zoologist 5:53 1—535.

PAVANS DE CEccatTTy, M., B. BUISSON, AND Y. M. GARGOUIL. 1963. Rythmes naturels et reactions motrices
chez Alcyonium digitatum Linn. et Veretillum cynomorium Cuv. Comptes Rendus des séances de la Societe
de biologie, Paris 157(3):616—-618.

PAX, F. 1936. Anthozoa (Teil II: Actiniaria, Octocorallia). Grimpe und Wagler. Die Tierwelt der Nord- und
Ostsee 30(3e):8 1-317. Leipzig. [Funiculina quadrangularis, Kophobelemnon stelliferum, Pavonaria chris-
tii, Pavonaria finmarchica, Pennatula grandis, Pennatula phosphorea subsp. variegata, Pennatula phos-
phorea subsp. candida, Pennatula aculeata, Protoptilum thomsoni, Stylatula elegans var. carnea, Stvlatula
elegans var. smaragdina, Virgularia tuberculata, Virgularia mirabilis.)

PAX, F. ANDI. MULLER. 1953. Die Anthozoen fauna von der Bucht von Kastela bei Split. Acta adriatica 5(1):1—35
[In German with Serbian summary; Pteroeides spinosum.]

WILLIAMS: INDEX PENNATULACEA 89

. 1954. Veretillum cynomorium an der Kiiste von Angola. Annales du Musée du Congo belge, Zoologie

1:23 1-237.

. 1955a. Gli antozoi del Golfo di Trieste. I. Lo Stato attuale della Ricerca e il Materiale di indagine

Esistente. Atti del Museo civico di storia naturelle di Trieste 20,2(6):49—102. [Veretillum cynomorium.|

. 1955b. Gli Antozoi del Museo Civico di Storia Naturale di Trieste. I. Parte: Antipatharia, Ceriantheria,

Zoantharia, Actiniaria, Alcyonaria e Pennatularia. Atti del Museo civico di storia naturelle di Trieste

20,3(7-8): 103-129. [Cavernularia pusilla, Funiculina quadrangularis, Kophobelemnon stelliferum, Pen-

natula phosphorea variegata, Pennatula phosphorea rubella, Pteroeides spinosum, Virgularia mirabilis
mirabilis. |

. 1955c. Die Korallentiere der Adria. Die Aquarien- und Terrarien Zeitschrift 8:10—-12. 39-40, 67-69.

[Funiculina quadrangularis, Pteroeides spinosum var. brevispinosa, Pteroeides spinosum var. longespi-

nosa.|

. 1959. The role of the Wroclaw Zoological Institute and Museum in the study of the fauna of Anthozoa

in the Adriatic Sea. Przeglad zoologiczny 3(1): 44—53 [In Polish with English summary; Preroeides

spinosum.|

. 1962. Die Anthozoen fauna der Adria. Fauna et Flora Adriatica 3:1-343. [In German with Serbian
resume. |

PEARSE, V., J. PEARSE, M. BUCHSBAUM, AND R. BUCHSBAUM. 1987. Living Invertebrates. Blackwell Scientific
Publications, Palo Alto. 848 pp. [Pennatulaceans, pp. 184, 186; Prilosarcus, Renilla.]

PERE, J. M. AND J. PICARD. 1958. Recherches sur les peuplements benthiques de la Méditerranée nord-orientale.
Bulletin de Institut Océanographie, Monaco 34:213-291.

PEREZ,C. D. 1996. Presencia de Renilla octodentata Zamponi y Pérez, 1995 (Cnidaria, Pennatulacea, Renillidae)
en la Bahia de Valparaiso (33°S — 72°W). Investigaciones Marinas, Valparaiso 24:145—147.

PERON, M. 1804. Précis, d’un Mémoire lu a I’Institut national, sur a température de la mer soit a sa surface, soit
a diverse profondeurs. Annales du Muséum D’Histoire Naturelle, Paris 5:123—148. [Contains a brief
description of octocoral bioluminescence; the following passage is quoted from p. 133, “Eh bien ! chaque
fois que nous retirions notre drague de la profondeur de 90 a 100 brasses, par laquelle nous naviguions
alors, elle étoit encombrée de zoophites de diverse espéces, particuliérement de rétipores, de sertulaires,
d’isis, de gorgones, d’alcyons et d’e¢ponges, mélés tous ensemble avec des fucus et des ulvas en grand
nombre. Presque tous ces objets étoient phosphoriques, et ce spectacle fut d’autant plus agréable, que notre
péche se faisoit au milieu des ténebres; . . . .”]

PERRIER, R. 1936. La Fauna de la France 1A: 8-64, 118, 229. Paris. [Pennatula phosphorea, Pennatula rubra,
Pteroeides griseum, Veretillum cynomorium. |

PEYSSONNEL, J. A. 1753. Traité du Corail. Philosophical Transactions of the Royal Society, London 47:445.

PFEFFER, G. 1886. Neue Pennatuliden des Hamburger Naturhistorischen Museums. Jahrbuch der Hamburgischen
wissenschaftlichen Anstalten 3:53-61.

PHILIPPI, R. A. 1835. Ueber Veretillum pusillum. Archiv fiir Naturgeschichte 1:277—280.

PIMENTEL, R. A. 1967. Invertebrate identification manual. D. van Nostrand Company, New York. 151 pp.
[Octocorals, pp. 43-46; Pennatulidae, Umbellulidae, Renillidae. ]

PLINIUS SECUNDUS, Caius (Pliny the Elder). 1469. Historia naturalis. Johannes de Spira, Venice. 356 lvs.

POCHE, F. 1914. Das System der Coelenterata. Archiv fiir Naturgeschichte (A) 80(5):47—128.

. 1915a. Zur terminologie der systematischen Kategorien und zur Benennung der suupergenerischen

Gruppen im allgemeinen und jener der Pennatulinea im besonderen. Zoologischer Anzeiger 45(11):

510-516. [“Pennatulinea” - nomenclatural discussion: Stephanoptilidae. ]

. 1915b. Uber das System der Anthozoa und einige allgemeine Fragen der zoologischen Systematik.
Zoologischer Anzeiger 46(1): 6-26.

POWELL, A. W. B. 1947. Native animals of New Zealand. Auckland. [Coelenterates, pp. 4-9; Sarcophyllum
bollonsi.]

PRATJE, A. 1923. Das Leuchten der Organismen. Eine Ubersicht iiber die neuere Literatur. Ergebnisse der
Physiologie 21:1—108. [Bioluminescence. ]

PRATT, E. M. 1909. Experimental observations on the organs of circulation and the powers of locomotion in
Pennatulids. Quarterly Journal of Microscopical Science 54:443.

90 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 2

Quoy, J. R. C. AND P. GAIMARD. 1827. Observations zoologiques faites a bord de |’ Astrolabe, en Mai 1826,
dans le Détroit de Gibraltar. Annales des Sciences Naturelles (1) 10:5—22, 172—193, 225-239. [Also
published in Oken’s /sis 21:col. 330-350, year 1927.]

RAMESH, A., B. LOGANATHAN, AND V. K. VENUGOPALAN. 1985. Commensal luminous bacteria of the coelen-
terate, Pteroeides sp. Current Science 54(12): 582-583. [Ecological interaction: Vibrio harvevi, bacteria
commensal in Preroeides sp., Bay of Bengal. ]

Rapp, M. W. 1827. Untersuchungen uber den Bau einiger Polypen des mittellandiscehn Meeres. Nova Acta
Physico-Medico Acad. Caesareae Leopoldino-Carolinae nat. curiosorum 14(2):643—658. [Biolumines-
cence. |

RHO, B.-O. AND J.-I. SONG. 1976. A study on the classification of the Korean Anthozoa 1. Gorgonacea and
Pennatulacea. Journal of the Korean Research Institute for Better Living 17:71—92.

. 1977. A study on the classification of the Korean Anthozoa 3. Alcyonacea and Pennatulacea. Journal
of the Korean Research Institute for Better Living 19:81—100.

Rice, A. L., P. A. TYLER, AND G. J. L. PATERSON. 1992. The pennatulid Kophobelemnon stelliferum (Cnidaria,
Octocorallia) in the Porcupine Seabight (North-East Atlantic Ocean). Journal of the Marine Biological
Association of the United Kingdom 72(2):417—434.

RICHIARDI, S. 1869. Monografia della famiglia dei Pennatularii. Archivio per la zoologia, |’anatomia e la
fisiologia (Turin) (2)1:1—150. [Reviewed by A. E. Verrill, American Journal of Science (2)49:426-427;
Pennatula targonii, p. 34 unknown locality; Pteroeides grayi, p. 54; Pteroeides vogtii, p. 55 Mediterranean;
Pteroeides cornaliae, p. 57 Adriatic; Pteroeides clausii, p. 58 Mediterranean; Pteroeides pancerii, p. 59
locality unknown; Virgularia leuckartii, p. 82 North Sea; Virgularia koellikeri, p. 83 Mozambique;
Cavernularia haimeii, p. 119 unknown locality; Cavernularia defilippii, p. 121 Mediterranean; Sceptoni-
dium new genus, p. 63; Sceptonidium mozambicanum sp. nov., p. 63 Mozambique. ]

RICKETTS, E. F., J. CALVIN, J. W. HEDGPETH, AND D. W. PHILLIPS. 1985. Between Pacific tides, Sth ed. Stanford
University Press, Stanford. 652 pp. [Sea pens: pp. 320-321; 324; 365-366; 513-514; 529; Acanthoptilum
gracile, Ptilosarcus gurneyi, Renilla amethystina, Renilla kollikeri, Stylatula elongata. |

RICHMOND, M. D., ed. 1997. A guide to the seashores of eastern Africa and the western Indian Ocean islands.
Sida/Department for Research Cooperation, SARE, Stockholm, Sweden. 448 pp. [Watercolor illustration
of Pteroeides sp. mislabeled as Virgularia gustaviana, p. 133.]

RICHTER, R. 1955. Die altesten Fossilien Siid-Afrikas. Senckenbergiana Lethaea 36(3—4):243—289. [Precambrian
fossils that resemble sea pens: Preridinium simplex and Rangea schneiderhoehni. |

RIDLEY, S. O. 1883. The coral fauna of Ceylon, with descriptions of new species. Annals and Magazine of Natural
History (5)11:250. [Pavonaria percarinata from Galle, p. 258.]

RIEDL, R. 1963. Fauna und Flora der Adria. Verlag Paul Parey, Hamburg und Berlin. 640 pp. [Sea pens, pp.
166—168.]

. 1983. Fauna und Flora des Mittelmeeres: ein systematischer Meeresfiihrer fiir Biologen und Naturfre-
unde, 3rd ed. P. Parey, Hamburg. 836 pp.

RITTSCHOF, D., I. R. HOOPER, AND J. D. COSTLOW. 1986. Barnacle settlement inhibitors from sea pansies, Renilla
reniformis. Bulletin of Marine Science 39(2):376—382. [Ecological interaction: Balanus amphitrite, larval
settlement and inhibitors characterization. |

. 1988. Settlement inhibition of marine invertebrate larvae: comparison of sensitivities of bryozoan and
barnacle larvae. Pp. 599-608 in Marine biodeterioration. Advanced techniques applicable to the Indian
Ocean, M.-F. Thompson, R. Sarojini, and R. Nagabhushanam, eds. A. A. Balkema, Rotterdam. [Renilla
reniformis, chemical composition, inhibitors of bryozoan and crustacean larval settlement, characteristics. ]

RIVEROS ZUNICA, F. 1948. Nuevos datos y redescripcion de Renilla chilensis Philippi. Revista de Biologia
Marine, Valparaiso 1(1):32—45.

ROBERTSON, D. 1887. Jottings from my notebook. On the local distribution of Pennatula phosphorea, Lin.,
Virgularia mirabilis, Lam., and Pavonaria quadrangularis, Pall. Proceedings and Transactions of the
Natural History Society of Glasgow 2:211—212.

ROEMER, F. 1880. Notiz tiber Belemnites ambiguus Morton aus der Kreide von New Jersey. Neues Jahrbuch fur
Mineralogie, Geologie und Palaeontologie, jahrgang 1880, 2:115—117. [Belemnites ambiguus is recognized
by Roemer as a member of the extinct sea pen genus Graphularia; see Shapiro and Ramsdell (1965:2).]

WILLIAMS: INDEX PENNATULACEA 9]

RONDELET, G. 1554-1555. Libri de piscibus marinis, in quibus verae piscium effigies expressae sunt. Lugduni,
apud Matthiam Bonhomme. [Translated from the Latin of 1554 - Lugduni, apud Matthiam Bonhomme
(1558) in two volumes. ]

Rossi, L. 1971. Guida a cnidari e ctenofori della fauna italiana. Quaderni della civica stazione idrobiologica di
Milano (No. 2):1—101. [Pennatula; key to the species; key to genera of Pennatulacea of Italy. ]

ROULE, L. 1905. Notice preliminaire sur les pennatulides recueillis par le “Travailleur” et le “Talisman,” dans
l’Océan Atlantique, au large du Maroc. Bulletin du Muséum national d’histoire naturelle, Paris 1 1:454—-458.
[Kophobelemnon bathyptiloides, Pteroeides echinatum, Stephanoptilum gen. nov. intermedium sp. nov.,
Umbellula crassiflora.|

. 1906. Une nouvelle famlie d’Anthozoaires. Bulletin du Muséum national d’histoire naturelle, Paris

12:120. [Stephanoptilidae fam. nov. ]

. 1907. Sur la morphologie comparée des colonies d’Alcyonaires. Comptes rendus hebdomadaires des

seances de |’ Académie des sciences, Paris 145:946—947. [Svavopsis gen. nov., S. elegans, morphology of

colonies. ]

. 1908. Alcyonaires d’Amboine. Revue suisse de zoologie 16(2):161—194. [Halisceptrum tenue, Svavop-

sis elegans.]

. 1932. Observations sur |’ontigenese du Véretille (Veretillum cynomorium Pall.). Archives de Zoologie
expérimentale et generale, Paris 74:233—247. [Development in Veretillum.]

Rowe, G. T. 1971. Observations on bottom currents and epibenthic populations in Hatteras submarine canyon.
Deep-Sea Research 18:569-581.

ROYAL SocIeTY (Great Britain). 1870. Preliminary report of the scientific exploration of the deep sea in H. M.
surveying-vessel ‘Porcupine,’ during the summer of 1869, conducted by Dr. Carpenter . . . J. Gwyn Jeffreys
...and Prof. Wyville Thomson. Proceedings of the Royal Society 121:492. [Observations regarding sea
pen bioluminescence]

RUMPHIUS, G. E. 1705 (recorded as 1740 or 1741 by most authors). D’Amboinsche Rariteitkamer, behelzende
eene Beschryvinge van allerhande zoo weeke als harde schaalvisschen, te weete raare Krabben, Kreeften,
en diergelyke Zeedieren, als mede allerhande Hoorntjes en Schulpen, die men in d’ Amboinsche Zee vindt:
Daar benevens zommige Mineraalen, Gesteenten, en soorten van Aarde, die ind’ Amboinsche, en zommige
omleggende Eilanden gevonden worden. T’Amsterdam, by Jan Roman de Jonge. 340 pp.

RUNNEGAR, B. 1992. Evolution of the earliest animals. /n Major events in the history of life, J. W. Schopf, ed.
Jones and Bartlett Publishers, Boston.190 pp. [A review of Vendian pennatulacean-like organisms. |
SACHS, E. 1913. Zur Kenntniss des feineren Baues von Echinoptilum. Jenaische Zeitschrift ftir Naturwissenschaft

50:839-847. [The genus Echinoptilum.]

SANCHEZ, M. J. A. 1994. Presencia de los octocorales Stvlatula diadema Bayer (Pennatulacea) y Carijoa riisei
(Duchassaing y Michelotti) (Telestacea) en la costa Caribe Colombiana. Anales del Instituto de Investi-
gaciones Marinas de Punta de Betin 23:137—147.

SANKOLLI, K. N. AND B. NEELAKANTAN. 1971. Animal associations along the west coast of India, 2: A porcellanid
crab (Decapoda, Anomura) on the coelenterate, Pennatula sp. Records of the Zoological Survery of India
2:51—S5S. [Ecological interaction: decapod and sea pen.]

SARS, M. 1846. Beschreibung der Pennatula borealis, einer neuen Seefeder. /n Fauna littoralis Norvegiae oder
Beschreibung und Abbildungen neuer oder wenig bekannten Seethiere, nebst Beobachtungen tiber die
Organisation. Lebensweise und Entwickelung derselben 1:1 7—19. Christiana, Druck und Verlag von Johann
Dahl. [Pennatula grandis.|

. 1851. Beretning on eni sommeren 1849 foretagen zoologisk reise i Lofoten og Finmarken. Nytt magasin
for naturvidenskapene 6:121—211.

SATTERLIE, R. A., P. A. V. ANDERSON, AND J. F. CASE. 1976. Morphology and electrophysiology of the
through-conducting systems in penntulid coelenterates. Pp. 619-627 in Coelenterate ecology and behav-
iour, G. O. Mackie, ed. Plenum Press, New York and London. 744 pp. [Nerve nets, conduction and colonial
organisation in pennatulaceans: Acanthoptilum, Ptilosarcus, Renilla, Stylatula, Virgularia.]

. 1980. Colonial coordination in anthozoans: Pennatulacea. Marine Behaviour and Physiology
7(1):25—46. [Pennatulacea: nerve nets, colonial coordination functions and electrophysiology. ]

SATTERLIE, R. A. AND J. F. CASE. 1979. Development of bioluminescence and other effector responses in the
pennatulid coelenterate Renilla koellikeri. Biological Bulletin (Woods Hole, Mass., Marine Biological

92 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 2

Laboratory) Boston 157(3):506—5S23. [Larval development of effector responses; metamorphosis and
settlement. ]

SCHAEFER, W. 1985. Phylum: Cnidaria. Nesseltiere. Pp. 162—1 195 in Lehrbuch der Zoologie, Teil 2:Systematick
(ed. 3), R. Siewing ed. Fischer, Stuttgart. 1107 pp. [Umbellula.]

SCHECHTER, V. 1959. Invertebrate zoology. Prentice-Hall, Inc., Englewood Cliffs, N. J. 530 pp. [Pennatulaceans,
pp. 164-169; Pennatula, Renilla.|

SCHOMANN, R. 1949. Die Welt der Tiere. Wien. 654 pp. (Coelenterates, pp. 366-385) [Pennatula phosphorea.|

SCHUHMACHER, H. AND J. HINTERKIRCHER. 1996. Niedere Meerestiere - Schwaémme, Korallen, Krebse,
Schnecken, Seesterne und andere - Rotes Meer, Indischer Ozean, Pazifik. Bestimmungsbuch fiir Taucher
und Schnorchler, Munchen. 320 pp. [Octocorals pp. 48-65; color photographs of Scytaliopsis djiboutiensis
(more probably Virgularia sp.), and Pteroeides sp.]

SCHULTZE, M. 1871. Ueber eine neue Species von Renilla: R. Miilleri. Sitzungsberichte der Niederrheinischen
Gesellschaft fiir Natur- und Heilkunde zu Bonn 28:36—37. [Renilla muelleri from Brazil.]

SCHULZE, F. E. 1875. Coelenteraten. Jahresbericht der Commission zur wissenschaftlichen Untersuchung der
deutschen Meere fiir die Jahre 1872—73:12—142. |Kophobelemnon leuckarti, Kophobelemnon stelliferum? |

SCLATER, P. L. 1872. Notice of a supposed new marine animal from Washington Territory north-west America.
Nature 6:436. [Probably represents the axis of Halipteris willemoesi.]

. 1873. The new marine animal from Washington Territory. Nature 8:487-488. [Probably represents the
axis of Halipteris willemoesi.|

SEILACHER, A. 1989. Vendozoa: organismic construction in the Proterozoic biosphere. Lethaia 22(3):229-239.
[Precambrian “sea pens” not recognized as pennatulaceans. |

SENUT, B. AND J.-M. FRANC. 1985. The collagenous component of Veretillum cynomorium (Cnidaria). NATO
ASI (Advanced Science Institutes) Series A (Life Sciences) 93:211—216. [Protein content, mesogleal
collagen and evidence for heterochain composition. ]

SHAPEERO, W. A. 1969. A positive chitosan test for spicules in the anthozoan order Pennatulacea. Pacific Science
23:261—163. [Ptilosarcus gurneyi referred to as Leioptilus guerneyi.|

SHAPIRO, E. A. AND R. C. RAMSDELL. 1965. The pennatulid species, Graphularia ambigua (Morton), from the
Upper Cretaceous and Lower Tertiary sediments of the Atlantic and Gulf Coastal Plain. Notulae Naturae
of the Academy of Natural Sciences of Philadelphia, no. 373:1—7. [Extinct sea pen Graphularia ambigua
from the Cretaceous/Tertiary of southeastern North America; a brief history of the literature pertaining to
this taxon is included. ]

SHAW, T. 173846. Travels, or observations relating to several parts of Barbary and the Levant, (1738). Oxford.
442 pp. [A supplement to the book entitled Travels, or observations, and etc., (1746). Oxford. 112 pp.; sea
pen bioluminescence. ]

SHEPHERD, S. A. 1983. The epifauna of megaripples: species adaptations and population responses to distur-
bance. Australian Journal of Ecology 8(1):3—8. [Virgularia mirabilis from South Australia. ]

SHIMEK, R. L. 1998. Subtidal splender in the northeastern Pacific. Wings 21(2):18-21.

SHIMOMURA, O. AND F. H. JOHNSON. 1975. Chemical nature of bioluminescence systems in coelenterates.
Proceedings of the National Academy of Sciences 72(4):1546—1549. [Renilla.]

——. 1979. Comparison of the amounts of key components in the bioluminescence systems of various
coelenterates. Comparative Biochemistry and Physiology (B) Comparative Biochemistry 64(1):105—107.
[Renilla and Cavernularia; luminescence; key chemical components; species comparison. |

SHORT, J. W. AND F. P. TROWER. 1986. Accumulation of butyltins in muscle tissue of chinook salmon reared in
sea pens treated with tri-n-butyltin. Marine Pollution Bulletin 17(12):542—545.

SIMONETTA, A. M. AND S. CONWAY Morkris, eds. 1991. The early evolution of Metazoa and the significance
of problematic taxa. Proceedings of an International Symposium held at the University of Camerino, 27-31
March 1989. Cambridge University Press, Cambridge. 296 pp.

Simpson, J.J. 1905. A new cavernularid from Ceylon. Annals and Magazine of Natural History
(7)15(90):561—S65. [Fusticularia herdmani.]|

SLOANE, H. 1707. A voyage to the islands Madera, Barbados, Nieves, S. Christophers and Jamaica, with the
natural history of the herbs and trees, four footed beasts, fishes, birds, insects, reptiles of the last of those
islands; ... Vol. 1, 1-264 + 156 pls. London.

SMIT, P. 1979. Hubrecht, Ambrosius Arnold Willem. /n Biografisch Woordenboek van Nederland, J. Charite.
Martinus Nijhoff, Gravenhage. 1:256—258. [Biographical sketch of A. A. W. Hubrecht. ]

WILLIAMS: INDEX PENNATULACEA 93

SMITH, R. I., ed. 1964. Keys to marine invertebrates of the Woods Hole region - a manual for the identification
of the more common marine invertebrates, compiled by Ralph I. Smith with the assistance of many other
contributors. Contribution No. 11 Systematics-Ecology Program, Marine Biological Laboratory Woods
Hole, Mass. 208 pp. [Octocorals, pp. 25—28; Pennatula aculeata.]

SMITH, R. I. AND J. T. CARLTON. 1975. Light’s Manual: Intertidal Invertebrates of the Central California Coast.
University of California Press, Berkeley. 716 pp. [Pennatulaceans, pp. 91 and 93; Acanthoptilum gracile,
Ptilosarcus gurneyi, Stvlatula elongata. |

SOARES, A. M. V. AND P. SAWAIA. 1975. Bioluminescénce de Renilla amethystina Verrill (Coelenterata -
Anthozoa). Influéncia do ion Mg. Ciéncias Cult., S. Paulo 23 (suppl.):392.

SPALLANZANI, L. 1784. Dissertazioni di fisica animale e vegetabile. English: Dissertations relative to the natural
history of animals and vegetables, tranlated from the Italian of the Abbe Spallanzani; to which are added
two letters from Mr. Bonnet to the author, two volumes. John Murray, London.

. 1796. Chimico esame degli esperimenti del sig. Gottling: professore a Jena, sopra la luce del fosforo di
Kunkel osservata nell’aria commune, ed in diversi fluidi aeriformi permanenti, nella qual occasione si
esaminano altri fosfori posti dentro ai medesimi fluidi, e si cerca se la luce solare guasti il gaz ossigeno,
siccome pretende questo chimco /, del cittadino Lazzaro Spallanzani. Presso la Societa tipografica, In
Modena. 171 pp. [Pennatulacean bioluminescence. |

SPHON, G. 1964. Sea pansies. Museum Talk, Santa Barbara Museum of Natural History 39(1):1—3. [Renilla
kollikeri.|

SPRUNG, J. AND J. C. DELBEEK. 1997. The reef aquarium, vol. 2: a comprehensive guide to the identification and
care of tropical marine invertebrates. Ricordea Publishing, Florida. 546 pp. [P. 15, color photograph of
Virgularia sp.; p. 256, color photograph of a zooxanthellate veretillid identified as Cavernularia obesa.]

SPURLOCK, B. O. AND M. J. CORMIER. 1975. A fine structure study of the anthocodium in Renilla muelleri.
Evidence for the existence of a bioluminescent organelle, the luminelle. Journal of Cell Biology
64(1):15—28.

SQUIRES, D. F. 1958. The Cretaceous and Tertiary corals of New Zealand. Paleontological Bulletin, New Zealand
29:1—107. [Extinct sea pen Graphularia longissima from New Zealand. |

STANDING, J. D., I. R. HOOPER, AND J. D. CoSTLOW. 1984. Inhibition and induction of barnacle settlement by
natural products present in octocorals. Journal of Chemical Ecology 10(6):823—834. [Renilla reniformis,
chemical composition, inhibitors and induces of crustacean larval settlement. |

STEARNS, R. E. C. 1873a. Remarks ona new alcyonoid polyp, from Burrard’s Inlet. Proceedings of the California
Academy of Sciences 5:7—12. [Abstract in American Naturalist 7:488, 633; pennatulacean axis. ]

. 1873b. Description of a new species of alcyonoid polyp. Mining and Scientific Press, San Francisco

27(6):88. [Pavonaria blakei.]

. 1873c. Description of a new genus and species of alcyonoid polyp. Proceedings of the California

Academy of Sciences 5:147—149. [Verrillia blakei new genus and species. }

. 1874. Remarks suggested by Dr. J. E. Gray’s paper on the “Stick Fish,” in “Nature,” Nov. 6th, 1873.

Proceedings of the California Academy of Sciences 5:283—285. [Probably axis of Halipteris willemoesi.]

. 1882. Verrillia blakei or Halipteris blakei. American Naturalist 16:55—56.

. 1883. Description of a new genus and species of alcyonid polyp from Japanese waters, with remarks
on the structure and habits of related forms, etc. Proceedings of the United States National Museum 6:96.
[Abstract in American Naturalist 17:1292; Verrillia.]

STEPHENS, J. 1909. Alcyonarian and madreporarian corals of the Irish coasts, by . .. , with description of a new
species of Stachyodes by Professor S.J. Hickson, F.R.S. Department of Agriculture and Technical
Instruction for Ireland; Fisheries Branch. Scientific Investigations 1907(5):1—28. [Benthoptilum sertum.]

STIASNY, G. 1937. Catalogue raisonné des alcyonides, gorgonides, zoanthides et pennatulides. Révision des
Collections H. Michelin. Bulletin du Muséum national d’histoire naturelle, Paris (2)9:391—397.

. 1938. Catalogue raisonné des alcyonides, gorgonides, zoanthides et pennatulides. Révision des Collec-
tions H. Michelin. Bulletin du Muséum national d’histoire naturelle, Paris (2)10:93—108. [Pennatula rubra,
Pennatula phosphorea candida.|

STIMPSON, W. 1855. Descriptions of some of the new marine invertebrata from the Chinese and Japanese Seas.
Proceedings of the Academy of Natural Sciences, Philadelphia 7:375-384. [Veretillum clavatum.]

STRAND, E. 1928. Miscellanea nomenclatorica zoologica et palaeontologica. Archiv fiir Naturgeschichte, Berlin
92 A.8:31—36. [Extinct sea pen Graphularia badenia.]

94 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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STRATHMANN, M. F. 1988. Reproduction and development of marine invertebrates of the northern Pacific
Coast—Data and methods of the study of eggs, embryos, and larvae. University of Washington Press,
Seattle. [Ptilosarcus gurneyi, reproduction and larval development, pp. 87, 90—91.]

STUDER, T. 1878. Uber die mit dem Schleppnetz angestellten Untersuchungen an der Westkiiste von Afrika
wahrend der Reise S. M. S. Gazelle. Sitzungsberichte der Gesellschaft naturforschender Freunde zu Berlin
1878:135—139.

. 1879. Ubersicht der Anthozoa Alcyonaria, welche wahrend der Reise S. M. S. Gazelle um die Erde

gesammelt wurden. Monatsbericht der K6niglichen Preussischen Akademie der Wissenschaften zu Berlin

1878:632—688. [ Pavonaria africana, Veretillum cynomorium, Cavernularia madeirensis, Renilla muelleri.|

. 1887a. Versuch eines Systemes der Alcyonarien. Archiv fiir Naturgeschichte, Berlin 53:1—74. [Classi-

fication of octocorals; Verrill’s 1865 subordinal name Pennatulacea is here corrected to ordinal status by

Studer. } ;
. 1887b. Systeme des Alcyonaires. Archives des Sciences physiques et naturelles, Geneve
18(11):43 1-432.

. 1887c. Ueber das System der Alcyonarien. Verhandlungen der schweizerischen Naturforschenden
Gesellschaft Actes de la Société helvetique des sciences naturelles. Atti della Societa elvetiva di scienze
naturali. Pp. 51—S3.

. 1887d. Ueber Bau und System der achtstrahligen Korallen. Mitteilungen naturforschenden Gesellschaft,

Bern 1886:xilI-xIv.

. 1891. Note préliminaire sur les Alcyonaires provenant des campagnes du yacht |’ Hirondelle 1886, 1887,

1888, Part 2: Alcyonacea and Pennatulacea. Mémoires de la Société Zoologique de France 4(2):86—95.

[Gyropohyllum hirondelli new genus and species, p. 94; at 1266 m depth between Pico and Sao Jorges,

Azores. |

. 1894. Reports on the dredging operations off the west coast of Central America to the Galapagos, to

the west coast of Mexico, and the Gulf of California, in charge of Alexander Agassiz, carried on by the

U.S. Fish Commission steamer ‘Albatross,’ during 1891. Lieut. Commaner Z.L. Tanner, U.S.N.,

commanding. X. Note préliminaire sur les alcyonaires. Bulletin of the Museum of Comparative Zoology

at Harvard College 25(5):53—69. [Abstracts in Journal of the Royal Microscopical Society 1894:350, 351;

and Zoologisches Centralblatt i: 314; Cladiscus agassizii, Distichoptilum verrillii, Kophobelemnon affine,

Pennatula alata, Pennatula koellikeri, Stachyptilumsuperbum, Umbellula geniculata. |

. 1901. Alcyonaires provenant de campagnes de |’Hirondelle (1886—1888). Résultats des Campagnes
scientifiques du Prince de Monaco 20: 1—64.

TAKADA, N. AND S. Mort. 1956. Daily rhythmic activities of the sea-pen, Cavernularia obesa Valenciennes. 13.
Rhythmic change of ammonium content in body fluid (1). Dobutsugaku zasshi 65:359-361. [In Japanese. ]

. 1957. Daily rhythmic activities of the sea-pen, Cavernularia obesa Valenciennes. 14. Rhythmic change
of ammonium content in body fluid (2). Dobutsugaku zasshi 66:284—288. [In Japanese. ]

TARDENT, P. AND R. TARDENT. 1980. Developmental and cellular biology of coelenterates - Proceedings of the
4th International Coelenterate Conference held in Interlaken, Switzerland, 4-8 September, 1979. El-
sevier/North-Holland Biomedical Press, Amsterdam. 499 pp. [Pennatulacaeans, pp. 43, 47; Pennatula
rubra, Veretillum cynomorium.|

THOMPSON, D’aArcy W. 1885. A bibliography of Protozoa, sponges, Coelenterata, and worms, including also
the Polyzoa, Brachiopoda and Tunicata, for the years 1861—1883. University Press, Cambridge. 284 pp.

THOMPSON, M.-F., R. SAROJINI, AND R. NAGABHUSHANAM, eds. 1988. Marine biodeterioration. Advanced
techniques applicable to the Indian Ocean. A. A. Balkema, Rotterdam. [Renilla reniformis: chemical
composition, inhibitors of bryozoan and crustacean larval settlement, characteristics. |

THOMSON, J. A. 1905. Appendix to the report on the Alcyonaria collected by Professor Herdman, at Ceylon, in
1902. In Report to the Government of Ceylon on the Pearl Oyster Fisheries of the Gulf of Manaar. Part 4,
supplementary report 28:167—186. [Virgularia loveni, Virgularia elegans, Virgularia calycina, Virgularia
indica. |

. 1927. Alcyonaires provenant des campagnes scientifiques du Prince Albert ler de Monaceo. Résultats

des campagnes scientifiques accomplies par le Prince Albert I 73:1—77. [Funiculina quadrangularis,

Kophobelemnon macrospinosum, Kophobelemnon stelliferum, Pennatula aculeata, Protoptilum carpen-

teri, Pteroeides griseum, Veretillum cynomorium, Virgularia cladiscus.|

WILLIAMS: INDEX PENNATULACEA 95

THOMSON, J. A. AND G. CRANE. 1909a. Alcyonarians of the Gulf of Cutch. Annals and Magazine of Natural
History (8)3:362-366.

. 1909b. The Alcyonarians of Okhamandal. Pp. 125—135 in Report Government Baroda Marine Zoology
of Okhamandal in Kattiawar, part 1, J. Hornell, ed. London. [Virgularia rumphii from India. ]

THOMSON, J. A. AND W. D. HENDERSON. 1905a. Report on the Alcyonaria collected by Professor Herdman, at
Ceylon, in 1902. /n Report to the Government of Ceylon on the Pearl Oyster Fisheries of the Gulf of Manaar.
Part 3, supplementary report 20:269-328. [Halisceptrum perivense, Halisceptrum gustavianum, Pteroeides
lacazei var. spinosum, Cavernularia obesa, Stylobelemnoides herdmani, Virgularia multiflora, Virgularia
tuberculata.]

. 1905b. Preliminary notice of the deep-sea Alcyonaria collected in the Indian Ocean. Annals and

Magazine of Natural History (7)15(90):547—557. [Junceoptilum alcocki p. 555, Kophobelemnon burgeri,

Microptilum willemoesi Andaman Sea p. 555, Pennatula murrayi p. 557, Protocaulon indicum, p. 554,

Protoptilum medium p. 555, Stachyptilum fuocum p. 557.]

. 1906a. The marine fauna of Zanzibar British East Africa, from collections made by Cyril Crossland,

M.A., V.Sc., F. Z.S., in the years 1901 and 1902. Alcyonaria. Proceedings of the Zoological Society of

London 1906(1):393-443. [Pteroeides brachycaulon, Pteroeides rigidum, Pteroeides pulchellum, Virgu-

laria mirabilis var. pedunculata, Virgularia UI.]

. 1906b. An account of the alcyonarians collected by the Royal Indian Marine Survey Ship Investigator
in the Indian Ocean, Part I: The alcyonarians of the deep sea. Trustees of the Indian Museum, Calcutta. 132
pp. [Anthoptilum decipiens, Bathyptilum indicum, Distichoptilum gracile, Funiculina gracilis, Ko-
phobelemnon burgeri var. indica, Pennatula indica, Pennatula veneris, Pennatula splendens, Pennatula
pendula, Protocaulon indicum, Protoptilum medium, Pteroéides triradiata, Sclerobelemnon kollikeri,
Stachyptilum maculatum, Thesioides inermis, Umbellula dura, Umbellula intermedia, Umbellula rosea,
Umbellula purpurea, Umbellula elongata, Umbellula kéllikeri, Umbellula radiata, Umbellula pendula,
Umbellula indica

THOMSON, J. A. AND D. L. MACKINNON. 1911. The alcyonarians of the “Thetis” Expedition. Australian Museum
Memoirs 4:661—695.

THOMSON, J. A. AND N. I. RENNET. 1927. Report on Japanese pennatulids. Journal of the Faculty of Science,
Imperial University of Tokyo 182(4, Zoology) 1(2):115—143. [Pennatula phosphorea constricta, Virgularia
kukenthali.|

. 1931. Alcyonaria, Madreporaria and Antipatharia. Scientific Reports, Australasian Antarctic Expedi-
tion, 1911—1914, Series C (Zoology and Botany) 9(3):1—46. [Umbellula carpenteri.|

THOMSON, J. A. AND J. RITCHIE. 1906. The Alcyonarians of the Scottish National Antarctic Expedition.
Transactions of the Royal Society of Edinburgh 41(3):851—860.

THOMSON, J. A. AND J. J. SIMPSON. 1909. An account of the alcyonarians collected by the Royal Indian Marine
Survey Ship Investigator in the Indian Ocean, II: The alcyonarians of the littoral area. Trustees of the Indian
Museum, Calcutta. 319 pp. [Cavernularia orientalis, Cavernularia andamanensis, Kophobelemnon inter-
medium, Lituaria hicksoni, Lituaria phalloides, Parabelemnon indicum, Pteroeides ilicifolium, Pteroeides
intermedium, Pteroeides robustum, Pteroeides andamanense, Pteroeides indicum, Pteroeides punctatum,
Virgularia ornata, Virgularia fusca.|

THOMSON, J. S. 1915. The Pennatulaceae of the Cape of Good Hope and Natal. Memoirs and Proceedings of the
Manchester Literary and Philosophical Society 59(1):1—26. [Actinoptilum molle, Pteroeides iscosceles,
Umbellula aciculifera.|

. 1917. The occurrence of Cavernularia Liitkenii KO6ll. in the Seas of Natal. Memoirs and Proceedings

of the Manchester Literary and Philosophical Society 62(7):1—S. [Cavernularia luetkenii.]

. 1924. Charts and comparisons of the distribution of South African Alcyonaria. Transactions of the Royal
Society of South Africa 11(1):45—84.

THURSTON, E., 1890. (Remarks as to the mode of life of the pennatulids, in a letter from . . . ). Proceedings of
the Zoological Society of London 1890:462-463.

TILEsIUS, A. VON. 1826. Naturhistorische abhandlungen und erlauterungen besonders die ptetrefactenkunde,
betreffend von Dr. A. von Tilesius. Bei Johann Christian Krieger und Compagnie, Cassel. 154 pp. [P. 85,
a reference to “Seechampagnon” as a possible synonym of Renilla muelleri, according to Zamponi and
Perez, 1995b:23, cited as Tilesius, 1812.]

96 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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TILESIUS VON TILENAU, W. G. 1819. Leuchten des Meers. Gilbert’s Ann. d. Phys. 61:36—44, 142—160, 161—176.
[Pennatulacean bioluminescence. |

TILLET-BARRET, E., J.-M. FRANC, S. FRANC, AND R. GARRONE. 1992. Characterization of heterotrimeric collagen
molecules in a sea-pen (Cnidaria, Octocorallia). European Journal of Biochemistry 203(1—2):179-184.
[Veretillum cynomorium. |

TITSCHACK, H. 1965. Untersuchungen uber das Leuchten der Seefeder Veretillum cynomorium (Pallas). Vie et
Milieu 15:547—563.

_ 1966. Uber die Lumineszenz und ihre Lokalisation bei Seefedern. Zoologischer Anzeiger, Supplement-

band 29, 1965(1966):120—131. [Pteroeides griseum, distribution of luminescent cells; Pennatula rubra and

Pennatula phosphorea, cytology of luminescent cells. ]

_ 1968. Uber das Nervensystem der Seefeder Veretillum cynomorium (Pallas). Zeitschrift fiir Zellfor-

schung und mikroskopische Anatomie 90:347-371.

. 1970. Histologische Untersuchung des Mesogloealen Nervenplexus der seefedern Pennatula rubra
(Ellis) und Preroides griseum (Bohadsch). Vie et Milieu (A)21:95—102.

TIXIER-DURIVAULT, A. 1954. Les octocoralliaires d’ Afrique du sud (II: Gorgonacea; III: Pennatulacea). Bulletin
du Muséum national d’histoire naturelle, Paris (2)26(5):624—631. [Actinoptilum molle, Anthoptilum grandi-
florum, Funiculina armata, Virgularia schultzei, Virgularia gustaviana, Pennatula phosphorea, Pteroeides
sp., plus the original description of Cavernularia dayi.]

. 1960. Les octocoralliaires de I’lle Inhaca. Bulletin du Muséum national d’histoire naturelle, Paris

(2)32(4):359-367. [Veretillum cynomorium, Veretillum leloupi, Actinoptilum molle, Virgularia gustavi-

ana.|

.1961a. Crassophyllum cristatum n. gen. et n. sp., type d’un genre de Pteroeididae (Pennatulacea).

Bulletin du Muséum national d’histoire naturelle, Paris (2)33(4):428—433. [Original descriptions of a new

genus and species in the family Pteroeididae. ]

. 1961b. Les octocoralliaires du Golfe de Guinée et des Iles du Cap-Vert (Alcyonacea, Pennatulacea).

Campagne de la ‘Calypso’: Golfe de Guinée. Annales de |’Institut Océanographique, Monaco

39(5):237-262. [Veretillum cynomorium, Cavernularia elegans, Cavernularia pusilla, Virgularia mirabi-

lis, Virgularia tuberculata, Pennatula rubra, plus the original description of Pteroeides morbosus.]

. 1963. Alcyonacea et Pennatulacea de I’ Afrique occidentale. Atlantide Report. Scientific Results of the

Danish Expedition to the Coasts of Tropical West Africa, 1945-1946 7:63—76. [Veretillum cynomorium,

Cavernularia pusilla, Funiculina quadrangularis, Virgularia mirabilis, Virgularia tuberculata, Pennatula

rubra, Crassophyllum cristatum, plus the original descriptions of Cavernularia mirifica and Pteroeides

hirsutus |

. 1965. Quelques octocoralliaires australiens. Bulletin du Muséum national d’histoire naturelle, Paris

(2)37(4):705—716. [Veretillum australe, Lituaria australasiae, Cavernularia obesa, Sclerobelemnon

schmeltzi, Virgularia gracillima, Virgularia gustaviana, Virgularia rumphi, Pteroeides bankanense,

Pteroeides oblongum, Sarcoptilus grandis, plus the original description of a new genus and species in the

family Kophobelemnidae - Malacobelemnon stephensoni.|

. 1966. Octocoralliaires de Madagascar et des Iles avoisinantes. Faune de Madagascar 21:1—456.

[Original descriptions of Virgularia densa, Pennatula delicata, Pteroeides crosnieri, Pteroeides acutum,

Pteroeides humesi, Pteroeides densum, and Pteroeides flexuosum. |

. 1970a. Octocoralliaires. Campagne de la “Calypso” au large des cOtes atlantiques de |’Amerique du

Sud (1961-1962). Annales de |’ Institut océanographique, Monaco 47:145—169. [Renilla muelleri, Renilla

reniformis, Stvlatula diadema.]

. 1970b. Les octocoralliaires de Nouvelle-Calédonie. L’Expédition frangaise sur les récifs coralliens de
la Nouvelle-Calédonie 4:171—350.

————. 1972. Nouvel rapport d’octocoralliaires de Madagascar et des Iles avoisinantes. Tethys, supplement
3:11-68. [Amphiacme abyssorum, Scytalium martensii, Virgularia densa, Virgularia helisceptrum, and

Virgularia juncea, plus the original descriptions of Pteroeides carnosum, Pteroeides spicatum, and
Pteroeides triangulum.]|

. 1987. Sous-Classe des Octocoralliaires. Pp. 3-185 in Traité de Zoologie, Anatomie, Systematique,

Biologie, P.-P. Grassé, ed. Masson, Paris. 3(3):1-859. [Pennatula phosphorea, Umbellula antarctica,

Anthoptilum grandiflorum, Veretillum cynomorium, Cavernuliaria subtilis, Echinoptilum echinatum,

WILLIAMS: INDEX PENNATULACEA 97

Renilla reniformis, Kophobelemnon stelliferum, Chunella gracillima, Funiculina quadrangularis, Virgu-
laria mirabilis, Pennatula rubra, Pteroeides morbosus, Distichoptilum gracile.|

TIXIER-DURIVAULT, A. AND M.-J. D’HONDT. 1974a. Nouvelles récoltes d’octocoralliaires a Madagascar. Téthys
5(2—3):25 1-266. [Actinoptilon molle, Anthoptilum grandiflorum, Funiculina quadrangularis, Amphiacme
abyssorum, Umbellula pellucida, Virgularia densa, Virgularia halsiceptrum, Virgularia juncea, Virgularia
mirabilis, Virgularia multicalycina, Scytalium martensii, Pennatula indica, Pennatula moseleyi, Pennatula
pearcevi, Pteroeides sp., and Gyrophyllum sibogae.|

. 1974b. Les octocoralliaires de la campagne Biagores. Bulletin du Muséum national d’histoire naturelle
Bulletin, Paris (Zool.) no. 174:1361—1433. [Anthoptilum murrayi, Funiculina quadrangularis, Protoptilum
carpenteri, Umbellula guentheri, Umbellula lindhali, Pennatula aculeata, Pennatula grandis, Pennatula
phosphorea, and Gyrophyllum hirondellei.}

TIXIER-DURIVAULT, A. AND F. LAFARGUE. 1968. Quelques octocoralliaires des cotes frangaises. Bulletin du
Muséum national d’Histoire Naturelle, Paris (2)40:621-629. [Original description of Cavernularia sub-
tilis.]

TIZARD, T. H., H. N. MOSELEY, H. Y. BUCHANAN, AND J. MARRAY. 1885. Narrative of the cruise of H. M. S.
Challenger with a general account of the scientific results of the expedition. Report on the Scientific Results
of the Voyage of the H. M. S. Challenger during the years 1873—76, Narrative, Vol. 1, (first part):1—S09.
[Pp. 49-51 on pennatulaceans, including an illustration of Umbellula thomsoni. The following passage is
worthy of quotation, “Many of the Pennatulida are known to be phosphorescent, and in this specimen of
Umbellula, when taken from the trawl, the polyps and the membrane covering the axis of the stem exhibited
a most brilliant phosphorescence. A like phenomenon was observed in the case of many other Alcyonarians
obtained from the deep sea, .. . Umbellula was long one of the rarest of zoological curiosities. The first
specimens ever described were obtained on the coast of Greenland, early in the last century, by Captain
Adriaanz, commander of the ‘Britannia,’ while on a whale-fishing expedition; on this occasion two
specimens were found adhering to the sounding line at a depth of 236 fathoms. These were described by
M. Christlob Mylius, and one of them was again described in the Philosophical Transactions for 1754, in
a letter from Mr. John Ellis to Mr. Peter Collinson, ‘Concerning a cluster-polyp found in the sea near the
coast of Greenland.’ Mr. Ellis compared it to the ‘Encrinos or Lilium lapideum...’ and indeed the
resemblance to a Crinoid is not a little striking. For more than a century the animal was not seen again, and
it is only a few years since two specimens were dredged in deep water during the cruise of the Swedish
ships ‘Ingegerd’ and ‘Gladan,’ in the Arctic Ocean. These were described in 1874 by J. Lindahl as two new
species,—Umbellula minacea and Umbellula pallida.]

TOMMASI, L. R., M. R. BIO, AND M. FUETA. 1972. Sobre a distribuicao de Renilla muelleri Kolliker, 1872 na
plataforma continental do Rio Grande do Sul (Anthozoa, Pennatulacea). Revista Brasileira de Biologia
32(1):55—58.

Torrey, H. B. 1901. Some facts concerning regeneration and regulation in Reni/la. Biological Bulletin (Woods
Hole, Mass., Marine Biological Laboratory), Boston 2(6):355—356. [Renilla: regeneration of colonies. ]

TRAUB, F. 1938. Geologische und palaontologische Bearbeitung der Kreide und des Tertiars im 6stlichen
Rupertuwinkel, nérdlich von Salzburg. Palaeontographica Stuttgart 88A(1—3):1—114. [Extinct sea pen
Graphularia salisburgensis from the Paleocene of Austria.]

TREMBLEY, A. 1744. Mémoires pour servir a I’histoire d’un genre de Polypes d’eau douce, a bras en forme de
cores. Abbé Trembley, Leiden.

TREMBLEY, P. 1941. Morphologie externe d’un fossile nouveau. Le Naturaliste Canadien, Québec 68(12):272.
[Possible fossil pennatulacean from Quebec. }

. 1942. Morphologie externe d’un fossile nouveau. Ann. Acfas, Montreal 8:85—86. [Possible fossil
pennatulacean from the Devonian of Canada. ]

TYLER, P. A., S. K. BRONSDON, C. M. YOUNG, AND A. L. RICE. 1995. Ecology and gametogenic biology of the
genus Umbellula (Pennatulacea) in the north Atlantic Ocean. Internationale Revue der Gesamten Hydro-
biologie 80(2):187—199.

TYLER, P. A. AND H. ZIBROWIUS. 1992. Submersible observations of the invertebrate fauna on the continental
slope southwest of Ireland (NE Atlantic Ocean). Oceanologica Acta 15(2):211—226. [Anthoptilum grandi-
florum.|

UmprIAco, D., M. ANCTIL, AND L. DESCARRIES. 1990. Serotonin-immunoreactive neurons in the cnidarian
Renilla koellikeri. Journal of Comparative Neurology 291(2):167—178. [Renilla koellikeri.]

98 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 2

UTINOMI, H. 1956a. Coloured illustrations of sea-shore animals of Japan. Fauna and Flora of Japan, No. 8
(2): 1-67. [In Japanese; coelenterates, pp. 3-28; Cavernularia obesa, Echinoptilum macintoshi, Pennatula
fimbriata, Pteroeides breviradiatum, Pteroeides esperi, Sclerobelemnon burgeri, Scytalium splendens,
Virgularia gustaviana.]|

. 1956b. On some alcyonarians from the West-Pacific islands (Palau, Panape and Bonins). Publications

of the Seto marine biological Laboratory 5(2):221—242. [Pennatula fimbriata. |

. 1958. On some octocorals from deep waters of Prov. Tosa, Sikoku. Publications of the Seto marine

biological Laboratory 7(1):89—110. [Kophobelemnon stelliferum, Pennatula murrayi, Scytalium splen-

dens.]|

. 1961. Noteworthy octocorals collected off the southwest coast of Kii Peninsula, Middle Japan. Part 2.

Telestacea, Gorgonacea and Pennatulacea. Publications of the Seto Marine Laboratory 9(1):197—228.

. 1964. Coloured illustrations of seashore animals of Japan. Hoikusha, Osaka. 168 pp. [Preroeides

sparmanni, Pteroeides esperi, Scytalium splendens, Virgularia gustaviana, Sclerobelemnon burgeri, Echi-

noptilum macintoshi, Cavernularia obesa, Leioptilus fimbriatus, Stachyptilum dofleini.]

. 1971. Port Phillip Bay Survey 2. Octocorallia. Memoirs of the National Museum of Victoria 32:7-18.
[Original description of Virgularia loveni.]

UTINOMI,, H. AND S. A. SHEPHERD. 1982. Seapens (order Pennatulacea). Pp. 207-211 in Handbook of the Flora
and Fauna of South Australia. Marine invertebrates of southern Australia. Part 1, S. A. Shepherd and I. M.
Thomas, eds. Handbooks Committee, South Australian Government, Adelaide. 491 pp.

VAFIDIS, D. AND A. KOUKOURAS. 1991. Crassophyllum thessalonicae sp. n. (Octocorallia, Pennatulacea), from
the Aegean Sea. Zoologica Scripta 20(3):201—205.

VarFIpDIS, D., A. KOUKOURAS, AND E. VOULTSIADOU-KOUKOURA. 1994. Octocoral fauna of the Aegean Sea with
a checklist of the Mediterranean species: new information, faunal comparisons. Annales de |’Institut
Oceanographique 70(2):217—229.

VALENCIENNES, A. 1850. Jn A monograph of the British fossil corals, Part I: Introduction; corals from the
Tertiary and Cretaceous formations, H. Milne Edwards and J. Haime, eds. Palaeontographical Society,
London.

VANDERAH, D. J. AND C. DJERASSI. 1977. Novel marine sterols with modified bile acid chain from the sea pen
Ptilosarcus gurneyi. Tetrahedron Letters 1977(8):683—686. [Steroid content; novel sterols, isolation and
structural determination. }

VAN Soest, R. W. M. 1977. A catalogue of the coelenterate type specimens of the Zoological Museum of
Amsterdam. III. Antipatharia, Pennatulacaea, Stolonifera, Telestacea, Alcyonacea. Beaufortia
26(332):77-98. [Veretillum malayense, Echinoptilum asperum, Echinoptilum elongatum, Echinoptilum
minimum, Echinoptilum roseum, Kophobelemnon pauciflorum, Sclerobelemnon elongatum, Sclerobelem-
non gravieri, Sclerobelemnon magniflorum, Anthoptilum malayense, Protoptilum celebense, Chunella
biflora, Umbellula weberi, Halisceptrum gustavianum var. parviflora, Halisceptrum gustavianum vat.
magnifolia, Virgularia rubra, Scytalium balssi, Pteroeides argenteum var. typicum, Pteroeides argenteum
var. durissimum, Pteroeides flavidum, Pteroeides griseum var. longespinosum, Pteroeides hystrix var.
angustifolium, Pteroeides hystrix var. latifolium, Pteroeides lacazii var. spinosum, Pteroeides malayense,
Pteroeides timorense, Pteroeides speciosum, Pteroeides tenerum, Pteroeides westermanni, Gyrophyllum
sibogae.|

VERRILL, A. E. 1864. List of the polyps and corals sent by the Museum of Comparative Zoology to other
institutions in exchange, with annotations. Bulletin of the Museum of Comparative Zoology at Harvard
College 1(3):29-60.

. 1865. Synopsis of the polyps and corals of the North Pacific Exploring Expedition, under Commodore

C. Ringgold and Captain John Rogers, U. S. N., from 1853 to 1856. Collected by Dr. W. Stimpson, naturalist

of the Expedition. With descriptions of some additional species from the west coast of North America.

Proceedings of the Essex Institute, Salem, Mass., Parts 2 and 3, Vol. 4:181—196, Vol. 5:17—50; Part 4, Vol.

5:315-333. [Pteroeides putnami, Funiculina forbesii = Pavonaria quadrangularis, Renilladanae = Renilla

americana, Renilla peltata, Renilla patula, Renilla amethystina, Ptilosarcus gurneyi, Veretillum stimpsont,

Kophobelemnon clavatum, Veretillum baculatum, Stylatula gracilis, Pteromorpha expansa, Leioptilum

undulatum, Virgularia pusilla, Virgularia gracilis, Virgularia elongata; introduction of the name Pennatu-

lacea as a suborder, with subsequent correction by Studer (1887a) as the ordinal name. ]

WILLIAMS: INDEX PENNATULACEA 99

. 1866. Review: Icones histiologicae, oder Atlas der vergleichenden Gewebelehre; zweite Abtheilung.
Der feinere Bau der hoheren Thiere. Erstes Heft. Die Bindesubstanz der Coelenteraten .... American
Journal of Science and Arts (2)42:283—284.

. 1868-1870. Notes on Radiata in the Museum of Yale College. No. 6. Review of the corals and polyps
of the west coast of America. Transactions of the Connecticut Academy of Arts and Sciences 1:377-422
(April 1868:377-390; June 1868:391-398; July 1868:399-414; December 1868:415—422); 423-502
(1869); 503-558 (1870). [Renilla amethystina, Stvlatula gracilis; Bayer (1996) remarks under his citation
of Verrill (1868—1870): “The regular edition up to p. 502 was destroyed by fire after distribution of the
author’s edition of 150 copies; the reprinted edition issued in 1869 contains nomenclatural changes marked
‘Reprint’ and thus constitutes a separate publication.” ]

. 1878. Notice of recent additions to the marine fauna of the eastern coast of North America, No. 2.

American Journal of Science and Arts (3)16:371-378.

. 1879. Notice of recent additions to the marine fauna of the eastern coast of North America, No. 3 and

5. American Journal of Science and Arts (3)17:239-243, 474. [Virgularia (?) grandiflora p. 239, Funiculina

armata p. 240.]

. 1882a. Notice of the remarkable marine fauna occupying the outer banks off the southern coast of New

England, No. 5. Brief contributions to zoology from the Musuem of Yale College: No. 51. American Journal

of Science and Arts (3)23:222—225, 309-316. [Pennatula aculeata var. alba, p. 310.]

. 1882b. Notice of the remarkable marine fauna occupying the outer banks off the southern coast of New

England, No. 7, and of some additions of the fauna of Vineyard Sound. American Journal of Science and

Arts (3)24:360-364. [Distichoptilum gracile, p. 362.]

. 1883. Report on the Anthozoa, and on some additional species dredged by the “Blake” in 1877-1879,

and by the U.S. Fish Commission steamer “Fish Hawk” in 1880-82. Bulletin of the Museum of

Comparative Zoology, Harvard 1|1:1—72.

. 1884a. Notice of the remarkable marine fauna occupying the outer banks off the southern coast of New

England, and of some additions to the fauna of Vineyard Sound. Annual Report of the United States

Commisioner of Fisheries for 1882:641—669. [Distichoptilum gracile.]

. 1884b. Notice of the remarkable marine fauna occupying the outer banks off the southern coast of New

England, No. 9. Brief contributions to zoology from the Museum of Yale College, No. 55. American Journal

of Science and Arts (3)28:213—220. [Umbellula bairdii, Kophobelemnon tenue, Scleroptilum gracile. |

. 1885a. Notice of the remarkable marine fauna occupying the outer banks of the southern coast of New

England, No. 11. Brief contributions to zoology from the Museum of Yale College, 57. American Journal

of Science and Arts (3)29:149—157. [Benthoptilum sertum, Stylatula sp.]

. 1885b. Results of the explorations made by the steamer Albatross off the northern coast of the United

States in 1883. Annual Report of the United States Commissioner of Fisheries for 1883:503—-699.

. 1922. The Alcyonaria of the Canadian Arctic expedition, 1913—1918, with a revision of some other
Canadian genera and species. Report of the Canadian Arctic Expedition, Ottawa 8G:1—87. [Pennatula
aculeata, Ptilosarcus gurneyi, Stylatula columbiana, Verrillia blakei.|

VIDAL, J. P., R. ESCALE, J. P. GIRARD, J. C. Rossi, J. M. CHANTRAINE, AND A. AUMELAS. 1992. Lituarines A,B,
and C: anew class of macrocyclic lactones from the New Caledonian sea pen Lituaria australasiae. Journal
of Organic Chemistry 57(22):5857—S860.

VINCENT, E. 1893. Sur la présence de Pennatuliens dans l’Eocéne belge. Annales de la Société malacologique
de Belgique 27:lviiix. [Extinct sea pen Graphularia belgica from the Eocene of Belgium. }

VoicT, E. 1958. Untersuchungen an Oktokorallen aus der oberen Kreide. Mitteilungen aus dem Geologisch-
Palaontologischen Institut der Universitat Hamburg 27:5—49. [Extinct sea pens Graphularia quadrata and
G. meijeri.|

Voss, G. L. 1976. Seashore life of Florida and the Caribbean—a guide to the common marine invertebrates and
plants of the Atlantic from Bermuda and the Bahamas to the West Indies and the Gulf of Mexico. Banyan
Books, Inc., Miami. 199 pp. [Octocorals, pp. 36-42; Renilla muelleri, Renilla reniformis.]

WAELE, J.-P. DE, M. ANCTIL, AND M. CARLBERG. 1987. Biogenic catecholamines in the cnidarian Renilla
kollikeri: radioenzymatic and chromatographic detection. Canadian Journal of Zoology 65(10):2458—2465.
[Renilla kollikeri, nervous transmitter substances. ]

100 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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WAMPLER, J. E., K. Hort, J. W. LEE, AND M. J. CORMIER. 1971. Structured bioluminescence. Two emitters
during both the in vitro and the in vivo bioluminescence of the sea pansy Renilla. Biochemistry (American
Chemical Society) 10:2903—2909.

WAMPLER, J. E., Y. D. KARKHANIS, J. G. MORIN, AND M. J. CORMIER. 1973. Similarities in the bioluminescence
from the Pennatulacea. Biochimica et biophysica acta 314(1):104—109. [Pennatulacea: bioluminescence,
comparative study. |

WARD, W. W. 1979. Energy transfer processes in bioluminescence. Photochemical and photobiological reviews
4:1—S7. [Renilla.]

WARD, W. W. AND Cormier, M. J. 1978a. Energy transfer via protein protein interaction in Reni//la biolumines-
cence. Photochemistry and Photobiology 27(4):389-396. [Renilla reniformis: luminescence; biolumines-
cence, energy transfer via protein protein interactions. |

. 1978b. Protein-protein interactions as measured by bioluminescence energy transfer in Renilla. Methods
in Enzymology 57:257-267. [Renilla reniformis; luminescence; energy transfer; protein interactions. ]

WATABE, N. AND DUNKELBERGER, D. G. 1979. Ultrastructural studies on calcification in various organisms.
Scanning Electron Microscopy. 1979(2):403-416. [Veretillum cynomorium; mineral content; calcification
ultrastructure of axial skeleton. ]

WATERMAN, T. H. 1950. Renilla kéllikeri. Pp. 131—136 in Selected Invertebrate Types, F. A. Brown, ed. John
Wiley, New York. [Renilla reniformis.]

WEIGUO, S. 1986. Late Precambrian pennatulids (sea pens) from the eastern Yangtze Gorge, China: Paracharnia
gen. nov. Precambrain Research 31(4):361-375. [Precambrian fossils that resemble sea pens: Charnia
dengyingensis Ding and Chen, 1981; designated type species for new genus Paracharnia of Charniidae,
Proterozoic. |

WEINBERG, S. 1996. Dé couvrir La Mer Rouge et |’océan Indien. Nathan, Paris. 415 pp. [Color photographs of
living animals: Veretillum sp., Virgularia/Pennatula sp., Pteroeides spp.]

WEKELL, J. C. 1974. Isolation and purification of a toxic component from the sea pen (Ptilosarcus quadrangu-
laris). Food-Drugs from the Sea Conference, Proceedings 1974:324-330.

. 1978. The isolation and characterization of a toxic diterpenoid compound from the sea pen, Prilosarcus
gurneyi (Gray). Dissertation Abstracts International (B)39(5):2210. [Lipid and fatty acid content. ]

WERNER, B. 1984. Stamm Cnidaria. Nesseltiere. Pp. 11-305 in Lehrbuch der speziellen Zoologie. Band
1:Werbellose Tiere, 2 Teil (Ed. 4), H.-E. Gruner, ed. Fischer, Jena. 621 pp. [Umbellula.]

WIEDEMANN, C. R. W. 1800. Cuvier’s Elementarischer Entwurf der Naturgeschichte der Thiere aus dem
Franzosischen tibersetzt und mit Anmerkungen versehen von C. R. W. Wiedemann, vol. 2, 599 Pauli,
Berlin. [Umbellula.]}

WIKTOR, J. 1974. Type-specimens of Anthozoa in the Zoological Museum of the Wroclaw University. Annales
Zoologici, Warszawa 32(3):29-37. [P. 35: Echinoptilum echinatum (two syntypes, Somalia); Actinoptilum
molle (one syntype, South Africa); Funiculina parkeri (fragment of the holotype or a syntype, California);
Pennatula inflata (one syntype, East Africa). ]

WILBUR, K. M. 1976. Recent studies of invertebrate mineralization. Belle W. Baruch Library in marine sciences.
5:79-108. [Renilla reniformis; skeleton; mineralization of sclerites, histochemical and ultrastructural
study. ]

WILKINS, P. AND J. BIRKHOLZ. 1986. Invertebrates—organ pipe, leather, and horny corals. Engelbert Pfriem
Publishing, Germany. 134 pp. [Color photograph of a specimen identified as Cavernularia obesa.]

WILLEMOES-SUHM, R. VON. 1875a. Notes on some young stages of Umbellularia, and on its geographical
distribution. Annals and Magazine of Natural History (4)15:312-316. [Umbellula.]

. 1875b. Distribution of the aleyonoid polyps of the Umbellularia group. American Journal of Sciences
and Arts (3)10:397—-398. [Umbellula.]

WILLIAMS, G. C. 1986. What are corals? Sagittarius (Natural History Magazine of the South African Museum)
1(2):11-15. [Reprinted in Underwater, Ihlane Publications, Natal, South Africa 6:26—27; illustrated general
account of corals including sea pens: Cavernularia sp., Actinoptilum molle; includes watercolor illustration
of color variation in A. molle.]

————. 1987. Systematics and zoogeography of southern African octocoral cnidarians. Ph. D. dissertation,
University of Cape Town, Cape Town, South Africa. 469 pp. [Pennatulacea of southern Africa, including
Namibia, South Africa, and southern Mozambique. |

WILLIAMS: INDEX PENNATULACEA 101

. 1989a. A provisional annotated list of octocorallian coelenterates occurring on the sublittoral coral reefs
at Sodwana Bay and Kosi Bay, northern Natal, with a key to the genera. South African Journal of Science
85(3):141—144. [Brief description: Cavernularia dayi.]

. 1989b. The pennatulacean genus Cavernularia Valenciennes (Octocorallia: Veretillidae). Zoological
Journal of the Linnean Society 95(4):285-310 [Synopsis of the genus, key to the species, original
descriptions of Cavernularia capitata and Cavernularia dedeckeri.|

. 1990. The Pennatulacea of southern Africa (Coelenterata, Anthozoa). Annals of the South African
Museum 99(4):31—119 [Regional fauna, synonymy, descriptions, key to the families and species, includes
illustrated descriptions of Cavernularia dayi, Cavernularia elegans, Echinoptilum macintoshii, Echinopti-
lum echinatum, Actinoptilum molle, Kophobelemnon stelliferum, Anthoptilum grandiflorum, Distichopti-
lum gracile, Halipteris africana, Chunella gracillima, Amphiacme abyssorum, Umbellula thomsoni,
Umbellula lindahli, Virgularia schultzei, Virgularia mirabilis, Virgularia gustaviana, Scytaliopsis djibou-
tiensis, Pennatula inflata, and Pteroeides isosceles; included are color photographs of living animals:
Actinoptilum molle, Virgularia schultzei, Echinoptilum echinatum, and Echinoptilum macintoshi, first
record of bioluminescence in Actinoptilum molle, p. 63.]

. 1992. Biogeography of the octocorallian coelenterate fauna of southern Africa. Biological Journal of
the Linnean Society 46(4):351—401. [Biogeographic affinities and bathymetry of southern African octoco-
rals including pennatulaceans. |

. 1993a. Coral reef octocorals—an illustrated guide to the soft corals, sea fans and sea pens inhabiting
the coral reefs of northern Natal. Durban Natural Science Museum. 64 pp. [Illustrated taxonomic descrip-
tion: Cavernularia dayi.]

. 1993b. Biotic diversity, biogeography, and phylogeny of pennatulacean octocorals associated with coral
reefs in the Indo-Pacific. Proceedings of the Seventh International Coral Reef Symposium 2:729—735.
[Biogeographic affinities of all pennatulacean genera and Indo-Pacific taxa in particular; phylogeny and
cladistics. The following passage from p. 734 on pennatulacean evolution is quoted here: “The Veretillidae
and the Echinoptilidae are the least derived of the extant sea pens. They are . . . concentrated in the relatively
shallow waters of the Indo-Pacific, while a variety of more derived forms are present worldwide and show
vast bathymetric ranges . . . . It is therefore postulated that the sea pens as a group initially differentiated in
the shallow waters of tropical oceans and have subsequently diversified and dispersed to all depths of the
temperate and polar regions, as well as the tropics.”’. ]

. 1993c. [See Branch and Williams (1993).]

. 1995a. Living genera of sea pens (Coelenterata: Octocorallia: Pennatulacea): illustrated key and
synopses. Zoological Journal of the Linnean Society 113:93—140. [Generic monograph, synonymy,
diagnoses, distributions, key to the families and genera, discussion of classification and phylogeny;
illustrated generic diagnoses include Lituaria, Cavernulina, Cavernularia, Veretillum, Actinoptilum, Echi-
noptilum, Renilla, Kophobelemnon, Sclerobelemnon, Malacabelemnon, Funiculina, Distichoptilum, Pro-
toptilum, Stachyptilum, Scleroptilum, Calibelemnon, Amphiacme, Chunella, Umbellula, Anthoptilum,
Halipteris, Stvlatula, Acanthoptilum, Scytalium, Virgularia, Scytaliopsis, Pennatula, Ptilosarcus, Gyro-
phyllum, Sarcoptilus, Crassophyllum, and Prteroeides.]

. 1995b. Preliminary assessment of the origin and phylogenetics of pennatulacean octocorals. Sixth
International Conference on Coelenterate Biology: Programme and Abstracts. P. 102. [Phylogenetic aspects
of the Pennatulacea and reevaluation of the Vendian frondlike fossils]

. 1995c. Revision of the pennatulacean genus Sarcoptilus (Coelenterata: Octocorallia) from southern
Australia and New Zealand, with descriptions of three new species. Records of the South Australian
Museum 28(1):13—32 [Taxonomic revision including three original descriptions; four species are consid-
ered valid: Sarcoptilus grandis, Sarcoptilus shaneparkeri, Sarcoptilus nullispiculatus, and Sarcoptilus
rigidus; with a discussion of the status of Sarcoptilus roseum and Sarcoptilus bollonsi.]

. 1995d. The enigmatic sea pen genus Gyrophyllum—a phylogenetic reassessment and description of G.
sibogae from Tasmanian waters (Coelenterata: Octocorallia). Proceedings of the California Academy of
Sciences 48(15):1—13. [Description, classification, cladistics: Gyrophyllum sibogae; family Pteroeididae
considered synonymous with Pennatulidae. |

. 1996. [See Gosliner et al.(1996); pp. 56-60. ]

. 1997a. A new genus and species of nepththeid soft coral (Octocorallia: Aleyonacea) from the western
Pacific Ocean, and a discussion of convergence with several deep-sea benthic organisms. Proceedings of

102 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 2

the California Academy of Sciences 49(12):423—437. [Umbellula: morphological convergence with three

other unrelated deep sea organisms. |

.1997b. Diversity and evolution of deep-sea pennatulacean octocorals. Eighth Deep-Sea Biology

Symposium, Abstracts. P. 138. [Evolutionary aspects of deep-sea pennatulaceans. }

.1997c. Preliminary assessment of the phylogeny of Penntulacaea (Anthozoa: Octocorallia), with a

reevalutation of Ediacaran frond-like fossils, and a synopsis of the history of evolutionary thought regarding

the sea pens. Proceedings of the Sixth International Conference on Coelenterate Biology. Pp. 497-509.

[Preliminary phylogeny of the Pennatulacea, assessment of the Vendian fossils that resemble sea pens, and

historical aspects of phylogeny. ]

. 1997d. Octocoral systematics and the Precambrian Vendian biota. Western Society of Naturalists (78th

annual meeting), Program and Abstracts. P. 57. [Comparison and contrast of morphology, living sea pens

vs. the Vendian frondlike fossils. ]

_1997e. Octocorallia. Jn Tree of Life World Wide Web Project, David and Wayne Maddison, eds.

University of Arizona, Tucson WEB SITE (http://ag.arizona.edu/tree/phylogeny.html). [Illustrated synop-

sis of the major taxa of Octocorallia, including the sea pens. ]

_ 1997f. Pennatulacea. /n Coelenterata (Cnidaria), J. van der Land, ed. Unesco-IOC Register of Marine

Organisms (UNESCO WEB SITE) (http://wwweti.eti.bio.uva.nl/database/urmo/default.shtml). [Compre-

hensive annotated listing of the species of Pennatulacea considered as valid. ]

. In press, a. [See Lopez-Gonzalez et al. (in press). ]

. In press, b. [See Fu et al. (in press). ]

WILLIAMS, G. C. AND ROGERS, J. 1989. Photographic evidence of bathyal octocorals from the Cape Basin. South
African Journal of Science 85(3):191—i92 [Deep-sea photographs and descriptions: Kophobelemnon sp..,
Umbellula sp.|

WILSON, E. B. 1880. Notes on some stages of Umbellularia, and on its geographical distribution. Annals and
Magazine of Natural History (4)15:312—316. [Umbellula.]

. 1881. The early stages of Renilla. American Journal of Science and Arts (3)20:446-449. [Renilla.]

. 1882a. Variation in the yolk-cleavage of Renilla. Zoologischer Anzeiger 5:545—548.

_ 1882b. Observations on the structure and development of Renilla and Leptogorgia. Johns Hopkins

University Circulars No. 17.

.1883a. The development of Renilla. Philosopical Transactions of the Royal Society of London

174:723-815. [Developmental biology of Renilla.]

. 1883b. The development of Renilla. Johns Hopkins University Circulars No. 22:78—79. [Developmental

biology of Renilla. |

_ 1884. The mesenterial filaments of the Alcyonaria. Mitteilungen aus der Zoologischen Station zu

Neapel; zugleich ein Repertorium fiir Mittelmeerkunde 5(1):1-27. [Observations on the anatomy of

mesenterial filaments. ]

. 1903. Notes on merogony and regeneration in Renilla. Biological Bulletin (Woods Hole, Mass., Marine
Biological Laboratory), Boston 4(5):215—226. [A summary is also published in Science (new series)
17:490-491; developmental biology of Reni/la.]

Wirtz, D. 1935. Die Korallen des norddeutschen Untermiozans. Jahrbucher Preussischen Geologischen Lan-
desantalt zu Berlin 55(1):84—-92. [Extinct sea pen Graphularia beyrichi from the Miocene of Germany. ]

WooDWARD, J. 1695. An essay towards a natural history of the earth, and terrestrial bodies, especially minerals:
as also of the sea, rivers, and springs. With an account of the universal deluge: and of the effects that it had
upon the Earth. Richard Wilkin, London.

WRATTEN, S. J., D. J. FAULKNER, K. HIROTSU, AND J. CLARDY. 1977. Stylatulide, a sea pen toxin. Journal of the
American Chemical Society 99(8):2824—2825. [Stvlatula sp.: toxins and venoms; structure and toxicity of
stylatulide to the crustacaean Tisbe. |

WRATTEN, S. J., W. FENICAL, D. J. FAULKNER, AND J. C. WEKELL. 1977b. Ptilosarcone, the toxin from the sea
pen Ptilosarcus gurneyi. Tetrahedron Letters 18:1559—1562.

WRIGHT, K. 1997. When life was odd. Discover—The World of Science 18(3):52—61. [ Vendian pennatulacean-
like organisms; includes watercolor life reconstructions of frondlike fossils that resemble sea pens]

WYVILLE THOMSON, C. 1874. The depths of the sea—an account of the general results of the dredging cruises
of H. M. SS. “Porcupine” and “Lightning” during the summers of 1868, 1869, and 1870, under the sicentific
direction of Dr. Carpenter, F. R. S., J. Gwyn Jeffreys, F. R. S., and Dr. Wyville Thomson, F. R. S., 2nd ed.

WILLIAMS: INDEX PENNATULACEA 103

Macmillan and Company, London. 527 pp. [Kophobelemnon miilleri (considered a synonym of K. stellif-
erum by Kiikenthal, 1915:29); Pavonaria quadrangularis (correctly referred to as Funiculina quadrangu-
laris, according to Kiikenthal, 1915). Two quotes are worthy of noting here; from p. 98, “In these dredgings
we got one or two very interesting alcyonarian zoophytes . .. . Many of the animals were most brilliantly
phophorescent .... In some places nearly everything brought up seemed full of luminous sparks. The
alcyonarians, the brittle-stars, and some annelids were the most brilliant. The Pennatulae, the Virgulariae,
and the Gorgoniae shone with a lambient white light, so bright that it showed quite distinctly the hour on
a watch; —and from p. 149, “We had another gorgeous display of luminosity during this cruise . . . the
dredge came up tangled with the long pink stems of the singular sea-pen Pavonaria quadrangularis. The
Pavonariae were resplendent with a pale lilac phosphorescence like the flame of cyanogen gas; not
scintillating ..., but almost constant, sometimes flashing out at one point more brightly and then dying
gradually into comparative dimness, but always sufficiently bright to make every portion of a stem caught
in the tangles or sticking to the ropes distinctly visible. From the number of specimens of Pavonaria brought
up at one haul we had evidently passed over a forest of them. The stems were a metre long, fringed with
hundreds of polyps.” ]

. 1878. The Voyage of the Challenger, Vol. 1, The Atlantic. Harper, New York. 391 pp.

YABE, H. AND T. SUGIYAMA. 1937. On a Graphularia-like fossil from the Pleistocene Tyoka beds of Tokyo.
Geological Society of Japan, Journal 44(531):1227—1230. [Possible extinct sea pen Graphularia? vamak-
awai from the Pleistocene of Japan; and placement of Corallium perplexum Gregorio, 1890, in the genus
Graphularia.|

YASHNOV, V. A. 1948. Anthozoa. Pp. 77-86 in Check list of the fauna and flora of the northern seas of the
U.S.S.R.,N. S. Gaevskoy, ed. Moscow. [In Russian; Funiculina quadrangularis, Pavonaria finmarchica,
Umbellula encrinus, Virgularia mirabilis]

ZAMPONI, M. O. AND C. D. PEREZ. 1995a. The family Renillidae Gray, 1860 (Cnidaria: Pennatulacea) from
sub-Antarctic region. Sixth International Conference on Coelenterate Biology, Programme and Abstracts.
Pe 105:

—. 1995b. Revision of the genus Renil/a Lamarck, 1816 (Octocorallia, Pennatulacea), with descriptions of

two new species from the Sub-Antarctic Region. Miscel-lania Zoologica (Journal of the Zoology Museum

of Barcelona) 18:21—32. [Taxonomic revision of Renilla including the original descriptions Renilla
octodentata and Renilla musaica; also included are descriptions and synonymies for Renilla reniformis and

Renilla muelleri; Renilla koellikeri is included in the key to the five recognized species, p. 28.]

1996. La presencia de la familia Virgulariidae Verrill, 1868 (Octocorallia, Pennatulacea) en aguas de
plataforma continental Argentina. Estratto dagli Annali del Museo Civico di Storia Naturale “G. Doria”
91:65—79.

ZAMPONI, M. O., C. D. PEREZ, AND R. CAPITOLI. 1997. El género Renilla Lamarck, 1816 (Anthozoa, Pennatu-
lacea) en aguas de Plataforma del Sur Brasilero. Estratto dagli Annali del Museo Civico di Storia Naturale
“G. Doria” 91:541—553.

ZIM, H. S. AND L. INGLE. 1955. Seashores—a guide to animals and plants along the beaches. Golden Press, New
York. 160 pp. [Watercolor illustration of Pennatula aculeata.]

ZIMMER, C, 1925. Willy Kiikenthal. Mitteilungen zoologischen Museum Berlin 1 1(2):169—179. [Biography of
Willy Kiikenthal, a major contributor regarding systematics of the Octocorallia. ]

ZUNIGA, F. R. 1948. Nuevos datos y redesrcipcion de Renilla chilensis Philippi, 1892 (Renillidae, Coelenterata).
Revista de biologia marina, Valparaiso 1(1):32—-45.

© CALIFORNIA ACADEMY OF SCIENCES, 1999
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San Francisco, California 94118

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PROCEEDINGS

OF THE

CALIFORNIA ACADEMY OF SCIENCES

Volume 51, No. 3, pp. 105—243, 58 figs., 7 tables.

July 20, 1999

Grenadiers (Pisces, Gadiformes) from the Continental Slope

of Western and Northwestern Australia

By

Tomio Iwamoto
Department of Ichthyology, California Academy of Sciences
Golden Gate Park, San Francisco, California 94118

and

Alan Williams
CSIRO Marine Research, Castray Esplanade,
Hobart, Tasmania 7000, Australia

CALIFORNIA
ACADEMY
SCIENCES @

GOLDEN GATE PARK

SAN FRANCISCO
PUBLISHED BY THE CALIFORNIA ACADEMY OF SCIENCES

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES

Volume 51, No. 3, pp. 105—243, 58 figs., 7 tables. July 20, 1999

Grenadiers (Pisces, Gadiformes) from the Continental Slope
of Western and Northwestern Australia

By

Tomio Iwamoto
Department of Ichthyology, California Academy of Sciences
Golden Gate Park, San Francisco, California 94118

and

Alan Williams
CSIRO Marine Research, Castray Esplanade,
Hobart, Tasmania 7000, Australia

This is the first taxonomic treatment of the grenadiers (Macrouridae and Bathygadidae)
from Australia’s western and northwestern continental slope. Collections made during two
trawl cruises by CSIRO vessels in 1984 and 1991 contributed most of the material for this
study. A remarkably diverse grenadier fauna was discovered with 63 species in 20 genera,
of which 18 species (about 29%) are here described as new. All species are treated, with
complete descriptions and illustrations for 37 species and more limited accounts for the 26
remaining species. Keys to genera and species are provided. Caelorinchus had by far the
most species represented (20), followed by Nezumia (7), Ventrifossa (6), Hymenocephalus
(5), Coryphaenoides (4), and Trachonurus (3). New species described include: Caelorinchus
amydrozosterus, C. charius, C. gaesorhynchus, C. goobala, C. lasti, C. mayiae, C. pardus, C.
thurla; Gadomus pepperi, Nezumia kapala, N. leucoura, N. merretti, N. soela, N. wularnia;
Trachonurus yiwardaus, Ventrifossa gomoni, V. paxtoni, V. sazonovi.

TABLE OF CONTENTS

PFUNTPOAINC TIO eae, i Sieh PS BAG hte AG san opin ee a RE Rd Re Lae ee 107
Memmogs-andyMatenialsiehcts ia ar. paniaes An Mahe Ons 1 fee Ih). nee aes 109
PEMA WAAL OMS pests elegy ar S), sahara tac MERE S UTE PSE. FAS Ce CE 110
fiom OmicaDESeriptons xe jas Sik) os, sas Wave aS Gi BU aes 5 ap ds ae ee I
Key ionGeneraion Westem-Australian Grenadiers® << 4.004. eeeee Aen meee 113
amaihyersathyaGdac. 3752 .0.4 Lc) = 4, PAAR ITSE a rl Ge a. aes eee eens 2, Bee ere we aS
Bamycaqus Gunther, US78) cco% gas ja vs So 544 Pa Aes Lee) eee: nS
Bathygadus:spongiceps Gilbert & Hubbs: 1920 ge 4cb 2. seeeisee Joe eee ee tS
Gadomus Resan, W903). 2s h00, se Sams 6 Ao so aces ae ee oe See ta?
Keystorspecies/of.Gadomus from: WestemmAusiraliayac <3 cence ao eee Lah,
Gadomusispxctacolle lives: fase las auainss fas cg EON ee Ly
Gadomiis pepperi NEWSPCCIESA ies 605.) ks es OSA Re eee 119
RamulyaViacroundacr « sa teehee ss ace lanees oh. hee, ee eee 22
SHptamuly Macrourinas. inac4 aeeeises bes/ca..ioe soba eee ee So eee 122
@aclomnchiis Giomia SiO sc Sarecnon Seo Ss OES Ses os ee eee 292
Keyatorspecies of Caclorinchus tromiWestemrAustraliay: 2a eee eee 122
Caelorinchusiacanthiger Barmatds 1925 —28 ek Oise ae eee 123

106 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 3

Saag eemcnusacuurosiis smith & Radcliffe, 1912 ........0.cce. ee ee nee eee eee 125
MTCIGRIICHUS QMYVAFOZOSTEFUuS NEW SPECIES ... 60... ee te ee eee 127
Gaclormenus argentatus Smith & Radcliffe, 1912 .... 0.2.62. .cce ewe wee cee e eee 129
DPSIEMICHIES CHOTIUS TIEW SPECIES 2.066. 6. eee cess on os ees no ee 132
Caclorinchus cacsorhynchus new Species ..... «05 6%... we cee es eee ee 136
WAEIOTINCHUS COOUGIADEW SPECIES. 20.5... re ee es ec ee eens ee os eee eee 138
@Gaclorinchus mnotabilis McCulloch? Y907 540.0. V0. ASRS 5 ee ee ee 140
WACIORINCRUS JASHENIEW SPECIES 22.22 oe be ci ct ee cee es eos dees ee 141
@aclorimehus macrorhynchus Smith & Radeliffe, 1912 .......22.......... 5 eee 145
@aclorinchus maculatus Gilbert & Hubbs, 1920) .........-..5-27 0001s os 147
Gaelorinchus matamua (McCann & McKnight, 1980)... see eo 2 148
Caelorinchus maurofasciatus McMillan & Paulin, 1993)... 4.2252... ase ee 150
Caclorinchus mayiae NéW SPECIES . . 2. 6 en ee ns tne ls ls oe is2
Caelorinchus mirus McCulloch, 1926: .... teem: « . 62.03.22 56@ wie caele nc) oo eee [55
Caelorinchus mycterismus McMillan & Paulin, 1993 .........9:....:..3.- eee Sy
Caclorinchus pardus New, SPeCieS 2: tenses D aes OR oo = [59
Caelorinchus smithi Gilbert.dqHubbSyl920 espera’: eatin... 2 sae cs 6 a ee 161
Caelorinchus.thurla new Species’ . .. .c. en he oo as Hs ob oe ee 2 ee 164
Caelorinchus trachycarus \wamoto, McMillan & Shcherbachev .................. 166
Cetonurichthys Sazonov & Shcherbachev; 1982 © 2 e522 22. SS Seas ee 167
Cetonurichthys subinflatus Sazonov & Shcherbachev, 1982 ..................... 167
Cetonurus Gunther, VSS7 esses ee fet oe ae oe ha See are elie aa a 169
Cetonurus globiceps (Vaillant, 1984)" 2. nu 6 ee es ne a ee 169
Coryphaenoides Gummer, 1165 « 6. open oa. 2g, +45 aati sae eens oes yeue 2. ety 1 gee eer 170
Key to Species of Coryphaenoides trom, Western Australia 9. 2.6. 6 .j.s..05 soe 170
Coryphaenoides.rudis:Gunther, 18738. nai. 2h saeco noel bak fas eae 170
CoryphaenoidesserrulatusGunthet, 1878. cant 24s 32 ees 2 Al
Conyphaenoides:striaturus Barnards1925) .¢..2eaoh Aes). Se ee eee 173
Coryphaenoides sp... PUR. 4 NEON, «| PAO ROS Fa ee 174
Hymenocephalus Gigholi, W884 ono Aires we ee a oe eee 175
Key to Species of Hymenocephalus from Western Australia .......-......-2--2s5e L75
Hymenocephalus adelscotti lwamoto & Merrett, 1997 \..............5.32-+ sce 176
Hymenocephalus longibarbis (Gunther, 1887) .... 23 04.5. s2 ane ese oe 2 ee w/a
Hymenocephalus nascens Gilbert & Hubbs, 1920. ....... ..525.+.5-+ 0 Ie
Hymenocephalus striatissimus aeger Gilbert & Hubbs, 1920 ...........---+.+--55- 178
FLYMENOCEPRAIUS SP & «2.04 sn dy sin so 55 dcr nd 9 Hee YS is 0 Nera 180
Kuroneziumia Twarmoto,, L974. os «sca a oo wr nw» I me ee 180
Key to Species of Kuronezumia from Western Australia ..............-2+++0+++000:- 181
Kuronezumia.leonis Barnatd, V925) 2... sacs anes oon se tee eee eee 181
Kuronezumia pallida Sazonov & Iwamoto, 1992 ..............0.0 ede cee eee eee: 181
Lepidorhynehus Richardson, 1846 3 5.5.45 50556 nn geo ce ao = eee 182
Lepidorhynchus denticulatus Richardson; 846.4207), S92.)...290 ae oe 182
Lucigadus.Gilbert & Hubbs; 1920. 2 95 4 snd ces ne ott aso Fo os tel rel ele 184
Lucigadus ori (Smith, 1968)) «2 x25 2055 a5 5s Js fete tel oe eee 184
Malacocephalus Gunther, 1862. «05.1 i455 545 3 ae yo 9 3 aa ee ee 186
Malacocephalus laevis (Lowe, 1843) 2.255.000 445005 045059) een 186
Matacocephalus:Betg, 1898 223 6.0000 aGes dain aa we oe ale 2s Sh ed te 187
Key to Species of Mataeocephalus from Western Australia ..............-+ eee eee 187

Mataeocephalus acipenserinus (Gilbert & Cramer, 1897) ..........0 002 eee ee eee 187

IWAMOTO AND WILLIAMS: GRENADIERS OF WESTERN AUSTRALIA 107

Wiataeoceniaiisispe 0). mente oe eta ements cre tee me ats een ene ener 189

IMCS OILS UD OSIOAWAITIOLO, IDM. 1. vn tnavert eka ox oe erat oem ete © eee ene ere ere 189
Kewatons peciestolVesODiUus! sana: 3. uc Sheen ae de ann eee fees ote a aieeee 189
Mesobiusaniipodum Hubbs é& Iwamoto, 1977 22.5.2... aes ee- se eee eee ee 189
Mesobias perryrt Lubbsiac Iwamoto, 197d enn. ao. oka sees See bialoe em ot ect 19]
INERTIA ONGATI NO Aue nee oe fac. ea de ORR ee ee Ae ION een re Etta oc Sn Manas 192
Meymoraustralianispecies Of Nezumiam = t.0.4. 0: 2-22 ene Uo oe eae ee ee 192
INE ZUTHIGURGIBELIEMIEW: SPECIES Pe ert es ce ee eh ee nee sen oe a 193

IN CZUMERICUCOUTE NOW SPOCICS# ei crs 2 nets nk a ee ea ae ae Oeste 195
INEZHMNLTHELTEHE NEW SPCCICS Set ee eee tee och iets Mer statstense we cteealomne so tienes 197,
INeEmnIG propingua(Gilbertoc Cramer, S97) ess oa. aw a eee eee ee ae eer 199

PN ZILINIG! SOCIO MCWASDECICSN © coe Meee 08 8 < See ee tf etinls ae serene eves ealtenay al cme eel 201
ezumiaspinosa(Gilbert-cckubbs, VONO)". 2 = 2. case eo eens so ome at ete 202
INCZUIMIG WIIATMIG MEW SPECIES ci)... ae ce et ne se einem Os oe ees al ee 205
ENenaonezimicnOKkamkalO7Ol Se secs os. oe aera me uate Make See ee leben eel seer 208
Pseudonezumia pusillus Sazonov & Shcherbachev, 1982 .....................00. 208
Naame MaerlnUs sr OW ler, TODS) 5 cvckwreyt eer) tse cece me eisrsin a etspage tie ere ere aerate 210
Sayacemacrurus pumiliceps (Alcock, 1894). 2.265022. ae sas Toe me Roe ie sees 210
aeRO ATT SAGUTGNET OIL os Le oe OLR Oe ee aD bare Ada oe OE Ee IE DA
Keyaoraustralian species of Trachonurus 2 2 mh oes Wee 2 5s ons ee D2
irachonurus cagates twamoto' é& McMillany 997 0509. 22202 oe eae oe eee oe eee ZA
iraechonurus senupellis Gilberts Cramer, 1897 ~ SUC NYE Te eee See ee coe 218
MrAChONUrUS yiwardaUus NeW SPECies 26 0. ks. Peek ons het ie eee eee ZANT
Venumossailben co ubbsnlO20 oe ea, Se Pe ee ene 219
MEN aIORAISthalian SPECIES OL VERITY OSSA .... 02. ws 2 oe ea eae ee es ee eee 29
VeniriOSSa COIMONENEW SPECIES. =... 22). 2 a.a's Pee eee Pe 220
Venmuossa jonnboborum iwamoto, 1982220.) Wok aa wa se ie ie le re 221
Ventuossa macropogon Marshall 197 3~, ie. 22 ne 2 a acie ne 2 ne see ae es ieee
Ventrijossamicrodorsals Gilbert & Hubbs, 1920: 22. 35 acess oe nee ss oe me eae 224

Venti OSSQ, PAXtOnl NEW:SPCCIES i.v..2 SSO. BLY Bae ee eee oe ee eee 228
VenirifosSa SAZOHOVE NEW: SPECIES» 4 3152-5 SAA oe ee Ae eee eo een eee oe 231
Submmillyeniacrouroldinae: =. 4.25 saet-2 oat? on 19 Rees eee see ee ra cosets, 0 eRe = 234
SoneOcaaus Gilbert: ccHUbbS, ONG y. 3 ais. NS ee de ed no eae eae ee eae 234
squalogadus modificatus Gilbert: & Hubbsy 1916) 225 eee ae ye eee eee 234
SUOMI MVaMlira chy TINCINAC: say nents epee ls Ls 4 © = en Ren eee Re a Er aetna 234
Keolopnonhynenus SaZonon, VOR seen Ws MOTE Een ee ee on a ees 2D
idiolophorhynchus andriashevi Sazonov; V8 leis. 5 452s) sane sales: oss a es tees 235
PARC EKO memes 2) AAMT A Sed A Neel ota cio tetceeh Minas La MA) Maa mern tard Meuoiory aie eieesaaeane eecce nme 237
ENROL GOINEN (Sterne aresase icra memes S1S8!, 21S fh aan ah ei thls nat eer ae meee 239
Bicrasre Cited: fnew sedate han kaa es A eh SUE chelate wia eesti tag a eee 239

In recent years the number of species of grenadiers (rattails or whiptails, families Macrouridae
and Bathygadidae) recorded from Australian waters has increased markedly as a result of exploratory
trawling for commercial fishery resources on the continental slope. Grenadiers are now recognized
as forming a large component of the Australian fish fauna: the 57 species recorded in Paxton et al.
(1989) ranked them the | 1th most speciose family. Despite this, they were poorly known, with 25 of
the 57 species either unidentified to species level or undescribed. In reality, closer to 100 grenadier
species in the family Macrouridae (sensu lato, including Bathygadidae) are represented in collections
from Australian waters (TI, unpublished data), ranking the group among the 10 most species-rich

108 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 3

families. A high number of these species remain either undescribed or unrecorded from Australia in
the published literature.

Collections of grenadiers from the extensive Australian western and northwestern slope region
came primarily from two exploratory fishing surveys undertaken by the Commonwealth Scientific
and Industrial Research Organisation (CSIRO) Marine Research. Records from the first of these
surveys, on the northwestern slope (Davis and Ward 1984), were included in the list compiled by
Paxton et al. (1989). The collection from the western slope, a region virtually unsampled until 1991,
elevated the number of undescribed species and added considerably to the species recorded from
Australian waters. Some material had been collected from this area during expeditions of the former
Soviet Union in the 1970s, but only a few grenadier species are treated in the published literature to
date (Sazonov and Shcherbachev 1982, 1985; Iwamoto and Shcherbachev 1991).

The purpose of this paper is.to provide the first regional taxonomic treatment of grenadiers from
Australia’s northwestern and western slope regions. It is also the first large-scale account of this rich
and abundant group of fishes from Australian waters. Included are descriptions for 37 species and
diagnoses only for the remaining 26 species. Keys are provided for genera with more than one species,
viz., Caelorinchus, Coryphaenoides, Gadomus, Hymenocephalus, Kuronezumia, Mataeocephalus,
Mesobius, Nezumia, Trachonurus, and Ventrifossa.

METHODS AND MATERIALS

Most of the material examined during this study was collected during CSIRO trawl surveys of
30 days duration off northwest Australia (1984) and western Australia (1991). However, many
additional specimens were provided by scientific observers aboard commercial fishing vessels from
both areas. Also, many reference specimens from eastern Australia were collected by the NSW FRV
Kapala.

The northwestern survey, using the FRV Soela, covered an area of the upper slope between
19°30'S, 115°30’E and 12°30’S, 123°03’E in depths from 300 to 500 m (Davis and Ward 1984). The
western survey, using the FRV Southern Surveyor, covered an area between 20°00’S, 114°00'E and
35°00'S, 115°15'E in depths from 200 to 1,400 m (Williams et al. 1996). In addition, one deeper trawl
(station SS1/91/12) was undertaken during the western survey; it started in 1,460 m but lost ground
contact somewhere between 1,500 m and 1,700 m over steeply sloping topography. (Plate 1)

Figure | shows the location of stations off Western Australia from which grenadiers were
collected. Appendix | provides station data for FRV Southern Surveyor, NSW Fisheries FRV Kapala,
and FRV Soela. (Note that coordinates and dates are not given in the Specimens Examined sections
where vessel stations are known.)

The northwestern survey, which was primarily targeted at crustacean resources, used an Engels
deep-sea lobster trawl with a 61 m ground rope. Mesh sizes ranged from 12 cm in the wings to 4 cm
in the codend. Ground gear was a combination of rubber discs weighted with chain and a tickler chain.
During the western survey, an Engels High-Lift fish trawl was used throughout. The trawl had a
headline length of 35.5 m, headline height of about 4 m, wingspread of 19 m, and was fitted with a 4
cm liner in the codend. The trawl was fitted with heavy rubber-bobbin ground gear and towed from
twin warps. Full details of net construction were given by May and Blaber (1989). A variety of gear
types was used by commercial vessels in both areas.

Fish specimens retained for museum collections were placed in 10% formaldehyde solution
aboard the research vessels. On commercial vessels specimens were frozen and preserved in the
laboratory at a later stage. Most of the material examined in this study are deposited in the I.S.R.
Munro Ichthyological Collection at CSIRO Marine Research Laboratories in Hobart (CSIRO), the
Australian Museum in Sydney (AMS), the Museum of Victoria in Melbourne (NMV), the Northern
Territory Museum in Darwin (NTM), and the Western Australian Museum in Perth (WAM). Other

IWAMOTO AND WILLIAMS: GRENADIERS OF WESTERN AUSTRALIA 109

PLATE Ib. Hauling in trawl aboard Southern Sur- PLATE lc. Fishery scientists from CSIRO Marine
veyor off Western Australia. Laboratories aboard Southern Surveyor examining
deep-water grenadiers from off Western Australia.

110 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 3

specimens were examined at the California Academy of Sciences, San Francisco (CAS); Queensland
Museum, Brisbane (QM); The Natural History Museum, London (BMNH); Zoological Museum,
Moscow State University, Moscow (ZMMGU); Muséum National d’Histoire Naturelle, Paris
(MNHN); the New Zealand Oceanographic Institute, Wellington (NZOI), now housed in the Museum
of New Zealand (NMNZ); and the National Museum of Natural History, Washington, DC (USNM).

We use the term grenadier to include bathygadids and macrourids, recognizing that they might
not be sister taxa and may actually belong in separate suborders (see Howes 1989; Howes and
Crimmen 1990).

Methods for taking counts and measurements follow Iwamoto and Sazonov (1988) and are given
in a condensed version in the Abbreviations section below. In the genus Caelorinchus, the fusion of
the lateral and medial nasal processes along the anterolateral margin of the snout is an important
character, described as “anterolateral margin fully supported by bone” or “... incompletely sup-
ported.” Whether or not the margin is fully or incompletely supported is easily determined by lightly
pressing along the margin with one’s fingernail. If there is a soft spot or some give to the pressure,
the margin is incompletely supported and there is a gap along the outer margin of the nasal bone; if
it feels solid, the margin is complete. Alternatively, the skin below the margin can be cut away to
view the bone. References to genera and type-species of genera are not provided, as Eschmeyer’s
(1990) Catalog of the Genera of Recent Fishes gives complete citations as well as type species
designation. (An updated Internet version of the Catalog of the Genera of Recent Fishes is available
at the WWW address http://www.calacademy.org.)

ABBREVIATIONS

Institutional abbreviations follow Leviton et al. (1985) and Leviton and Gibbs (1988). For
political entities, the following abbreviations are used: CSIRO — Commonwealth Scientific and
Industrial Research Organisation; GAB — Great Australian Bight; NSW — New South Wales; NT —
Northern Territory; QLD — Queensland; SA — South Australia; TAS — Tasmania; VIC — Victoria;
WA — Western Australia. Vessel abbreviations include: SS — FRV Southern Surveyor, SO — FRV
Soela; K—FRV Kapala.

Counts

1D. and 2D. — counts of the first and second dorsal fins; the Roman numerals “II” refer to the
two spinous rays of the first dorsal fin.

P. — count of pectoral fin rays; the lowercase “1” refers to the rudimentary uppermost ray.

V.—pelvic fin ray count.

A. —anal fin ray count.

GR-I, GR-II — gill raker counts of the first (outermost) and second arches. Counts of outer and
inner gill raker series are given and separated by a slash mark. Raker counts of the upper arm are
separated from those of the lower arm by a plus sign. :

Scales 1D., 2D., midbase 1D., lat.line — scale row counts, respectively, below the origin of the
first dorsal fin, second dorsal fin, midbase of first dorsal fin, and lateral line scales counted from the
anterior end over a distance equal to the predorsal length. Half-scale counts (0.5) are given when the
uppermost scale is considerably smaller than others of the series.

Caeca — pyloric caeca count.

Measurements

TL — total length; a plus sign following the length indicates that a portion of the tail tip was
missing.

HL — head length, taken from snout tip to upper posterior angle of opercle.

Head width — the greatest transverse measurement, usually across the opercles and including the
bony scutes of the suborbital ridge.

IWAMOTO AND WILLIAMS: GRENADIERS OF WESTERN AUSTRALIA

18°
Exmouth Plateau
20° t=
1500 m > | 2" Port Hedland
West Cape
ith |
24° INDIAN ) Jay ae
OCEAN vi | AUSTRALIA
° oe Bay
vy ui
JY
28° * \
¥ eer Geraldton
&, |
S.
e
32° M\
upon
Mandurah Vine
» or we eas ee pe,
oe 108° 112° LSS) yp aes ee 124°

FiGuRE 1. Station localities off Western Australia at which grenadiers were collected by the fishery research vessels Soe/a
and Southern Surveyor, and by other vessels.

112 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 3

Snout — length from tip of snout to anterior margin of orbit.

Preoral — median ventral length of snout from tip to upper margin of lips.

Internasal — least distance between supranarial ridges.

Interorb. — least distance between bony orbits.

Suborb. — least width of suborbital space.

Postorb. — distance from posterior margin of orbit to upper posterior angle of opercle.

Orb.-preop. — oblique measurement from posteroventral margin of orbit to posteroventral margin
of preopercle.

Up.jaw — length from anterior tip of premaxillary to posterior end of maxillary.

Pmx.—premaxillary; height measured from dorsal tip of anterior ascending process to anteroven-
tral margin of bone, not including teeth; length the greatest measure from anterior tip to posterior end;
gap measured between dentition bands of each premaxilla.

Barbel — length of free portion of barbel, from posterior insertion to distal tip.

Gill slit — greatest diameter of outermost gill slit.

Pre-A. — distance from tip of snout to origin of anal fin.

Pre-vent — distance from tip of snout to anal opening.

V.-A. — distance between base of outer pelvic ray to anal fin origin.

Isth.-A. — distance between anterior end of isthmus to anal fin origin.

Body depth; depth at A. - greatest body depth, usually below origin of first dorsal fin; depth at
origin of anal fin.

1D.-2D. — distance between posterior edge of | st dorsal fin and anterior origin of 2nd dorsal fin.

Ht. 1D., len. 1D. base, len. P., len. V. — height of first dorsal fin, length of base of first dorsal,
lengths of pectoral and pelvic fins, respectively.

Post. nostril — greatest diameter of posterior nostril.

Len. rictus — length of lateral opening of mouth measured from anterior tip of premaxillary to
posterior angle of mouth.

Other abbreviations: cf., to be compared with; coll., collector; compass directions are given as
n., ne., nw., etc., for north, northeast, northwest, etc.; collection dates are given as “10.11.1990"
indicating 10 February 1990 (day, month, year); est., estimate; fm, fathom(s); ht., height; I. (Island),
Is. (Islands); len., length; no., number; NW, North West [as part of a proper noun]; pmx., premaxilla;
Pt., Point; spec., specimen(s); sta., station; uncat., uncataloged.

TAXONOMIC DESCRIPTIONS

The remarkably diverse grenadier fauna of the western Australian region, the relative paucity of
solid taxonomic works on the species of that region, and the inadequate reference collections for many
species has made the current species treatments difficult, and in several instances, impossible to fully
resolve. Our treatments of the genera Hymenocephalus, Trachonurus, and Ventrifossa are of particular
concern in this respect, and they must be viewed as only tentative. We leave their full taxonomic
resolution to others who have more material and who are able to devote time and effort to more
extensive studies than we could. Many of the problems suit themselves to regional ichthyologists who
have access to fresh local material as well as comparative material from outside the area. In some
examples where an apparent species shows a broadly disjunct distribution (e.g., Caelorinchus
mycterismus, Trachonurus sentipellis, Ventrifossa macropogon), access to material from intervening
areas will be necessary to show whether or not the populations represent the same species. In several
species descriptions, we list specimens in an uncertain-variant category. Such specimens did not
properly fit our species concepts for one reason or another, but we could not justify recognizing them
as distinct taxa. Again, such problematic species and specimens will require a more thorough study
than we were able to give them.

IWAMOTO AND WILLIAMS: GRENADIERS OF WESTERN AUSTRALIA 113

We do not provide diagnoses for families and genera as this paper is not intended to be a
taxonomic review of the grenadiers. For these we refer the reader to recent publications by Howes
and Crimmen (1990), Iwamoto (1990), Iwamoto and Sazonov (1994), Sazonov and Iwamoto (1992),
Sazonov and Shcherbachev (1982, 1985), and Shcherbachev et al. (1992). Of the 63 species here
treated, full descriptions are given for 18 new taxa. Of the 45 remaining species, 16, that in our opinion
seemed to require detailed description, are also fully described; the other 29 species, because of being
so well known or already well described, are diagnosed only. Three undescribed species are diagnosed
but not named, two of these (Coryphaenoides sp. A and Mataeocephalus “shortsnout”) because others
are currently preparing descriptions for them, and a third species (Hymenocephalus sp. 1) because we
lacked adequate material. Synonymies are limited to the original description, a few principal
references, and to references germane to the area.

It is apparent from our studies that the number of grenadiers known from the western Australian
region will increase over time as the fauna becomes better studied, and as more effort is made to
collect over greater geographic areas and depths, and over a variety of terrain. As an example, that
our study material was limited to depths shallower than 1,700 m precluded the possibility of the
deeper-living Coryphaenoides species being represented. A broad expanse of shelf and slope north
and east of Port Hedland to the Arafura Sea was not sampled by CSIRO vessels. Valuable trawl
collections from the Timor and Arafura seas housed at the Northern Territory Museum include species
not collected by CSIRO vessels, suggesting that the area is ripe for more collecting.

We are a long way from knowing the full extent of the grenadier fauna of Western Australia, and
this paper is only a preliminary report with many species left out and many questions left unanswered.
The western Australian region holds the potential for many more stimulating and valuable research
projects.

KEY TO GENERA OF WESTERN AUSTRALIAN GRENADIERS
[Mostly based on Iwamoto (1990) and Iwamoto and Merrett (1997). Species and genera listed

in brackets have not been recorded from the area but may be expected. Note that the genera
Mataeocephalus and Ventrifossa are keyed out twice. ]

la. One continuous dorsal fin, anterior portion not elevated; orbit small, 10 or more times inhead length ........ 2
libs iwoidorsal fins, the firstelevated;orbit less than ]0jtimesinheadilength5 3. 7 se ee 3
PMMRe WICHINSSINAl ly TAY Sed — Ose eh cee cogs essen oy ch deste ceil Eee ee cial Sey Pee Squalogadus modificatus
DP DMMRE I ICAI NSHADSEN tee week eee Sie ora nick Settee eect eae Pr na Ne ee ie LUPO Hs) Uae [Macrouroides inflaticeps |
3a. Origin of second dorsal fin immediately behind first and equally or better developed than anal fin; outer gill arch free
mompeullkcoversoill-rakersjonminstrarchislendernotitubercwlan | pense etsy) ersten een re cite) enon 4
3b. A distinct gap between dorsal fins; anal fin usually much better developed than second dorsal fin; outer gill arch
restricted by folds of skin connecting upper and lower limbs to gill cover; gill rakers tubercular... ........ 7
4a. Snout long and sharply pointed; mouth inferior; heavy scutelike scales on body .............-.-+--. 5
4b. Snout bluntly rounded, not protruding; mouth essentially terminal; all scales thin, deciduous, lacking spinules ... 6
5a. Large platelike scutes in three or four longitudinal rows on body; body color black . . . /diolophorhynchus andriashevi

5b. Two rows of scutes on body, one along second dorsal fin, the other along anal fin; body color brownish
5.0 ee Ra MS SWer Syed adic Mae onRy ee, Oo cuchin tc ie ok ts art GOR, TLS Suen eas ue e ies ei aide [Trachyrincus |
6a. Chin barbel well developed; first dorsal, pectoral, and pelvic fins with a greatly elongated ray; pelvic fin rays 8 or 9

(SIAL Wt Nano) ciate Noy a er cueROnGniomno. 01 Ul omc. Culamis ia tcl eeate Nant Be US MPM NORE outiad eters cual oud cic 6 Gadomus
6b. No chin barbel; outer pelvic ray sometimes elongated, but other fins lacking a greatly elongated ray; pelvic fin rays

SeOMUstiallyp9) iw ees, chee eS ole nt eee cr ce TREE. Suusesty rch clase aes ro) oe av sna ste Chanel ete staapecmmraiens Bathygadus spongiceps
TE, SCOMG SOMOUS Hy OHS COMMON. oo op ames oenotonnodoodd ooo God oO os coo aa 8
7b. Second spinous ray of first dorsal fin serrated (weakly or much reduced insome) .................. 14
8a. Snout strongly pointed, armed with coarse spiny scales that form a stout continuous ridge extending from snout tip to

preopercleranslesanditerminatin cain alshanp Om taeem enna ttle ley (Ui prci sn itca cnn Ten cin aiei-ncin Caelorinchus

8b. Snout rounded to moderately pointed, no sharp ridge of modified scales extending from snout tip to preopercle
ANGICMEee Sal SR A Oe a oe A AAG DS Sen Goat Gee eee ee enna e Aone CER ERE Cmte nels 9

114 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 3

9a. Broad areas of fine (microscopic), parallel black lines (ventral striae) overlying silvery ground on ventral surfaces

OMchesmsnoulden cide, alongieachiside of isthmus; and'belly. .. .- 24-2 sera ss ss =) scene 10
Sin. IN® VERE SHES. c ono 6 GM Ole a 66 one ee ee eee ie en omen E NENG O° 11
10a. Ventral striae extends alongside anterior half or more of anal fin base; six branchiostegal rays; no lenslike light

MORPAMONIChesteaitaMs OSC lemee wa a a se ae ee a es bee fee Lepidorhynchus denticulatus
10b. Ventral striae rarely extends posterior to anus; seven branchiostegal rays; a small lens or light organ on chest;

Siallespeciesmusuallyzattainsslessythane2S Cnn Mis oa 6 ct de ctw wy ee ye eee ee ene Hymenocephalus
Pam bowemawawitonarses widely:spaceditecth in TOW... . «6 ee se wis ces ee) yee ey ee ee ee 12
MDMbOWwenjawawithiratnensmalliteethinmorettham Tow ...2 5... 50 05 6 7s ees 9 ce eee 13
12a. Canine teeth in | row in both jaws; grooved lateral line not developed beyond second dorsal fin; 10—11 pectoral fin

ray Syolbranchiostepalirayscolormostly blackish) 2%.) 5 40 - 2) © ele oe ronenee eee [Odontomacrurus murrayi}
12b. Teeth in 2 rows in upper jaw; lateral line complete to end of tail; 16 or more pectoral rays; 7 branchiostegal rays;

color usually grayish, somewhat silvery below ventral midline. ................. Malacocephalus laevis
13a. Pelvic rays 6-7; scales coarsely spinulated; broad, naked, black periproct region immediately before anal fin

TR ne et eee Seer ein Ree ar i nie Paice Ramil E Mas unm ac, iC LUIALS

13b. Pelvic rays 9-10; scales finely spinulated; periproct region separated from anal fin by many scale rows . . Ventrifossa
14a. Head massive, globose, soft; interopercle tightly adnate to preopercle; body scales along each side of second dorsal

Hin bAasepemlAnO eG pcre sek yes aes mien PS OY oe Sues a euL Vas oud uy Quel is: Sac, foie ales © voyuer SO eee IS)
14b. Head not especially massive or globose; interopercle not strongly adnate to preopercle; no enlarged scales along

secondidorsalitintbase "SPs a ee eee oe ites a toe Se a Ste ae len en rn 16
15a. Pelvic fin bases posterior to vertical through origins of first dorsal and pectoral fins; interorbital width 31-34% of

FAL eho coy ero li: ead wise bccn esas Shey kes Ra tiantt bade ec pe tors ore Sela as Cetonurichthys subinflatus
15b. Pelvic fin bases about at or anterior to vertical through origin of first dorsal and pectoral fins; interorbital width

SO—48 1/0, OLMIS er Mn crete ceo ct lee es bse er TA ig Gates Oe EA ceg oh Oe ea ene ne Cetonurus globiceps
16a. Scales of head elongated, with spinules longitudinally aligned to give striated pattern to head surfaces; chin barbel

absent. Sn Pde la oi et ee) oe GL eR OR Ee, VOIR, Lee ES ee ee ee oe Mesobius
llébyHeadiscalesmotelongated: nojstnated| pattem|toshead|surfacesi2 45) 2) ee 2 ee 17
17a. Snout low, narrow, rounded, devoid of scales; no ridges on head; scales on head and front of body without spinules

or ridges; maxilla reaches vertical to front marginoforbit ................ [Haplomacrourus nudirostris |

17b. Snout angular, completely naked to variously covered with scales; ridges usually present on head, sometimes
coarsely scaled; almost all scales covered with spinules or low ridges; maxilla usually extends well posterior to

irontotorbin (excepts phagemacrurusiandiiiiclsadus)s ee ena eee eee 18
LSA vB ranchiOSste Sal Ay ST se oso ewe Geer I Sas eyo RSM olemichas ey rysdesite yes etlue 2 ssh oesilan tase oy ORE mn 19
rSbsBranchiostegallirays ie sq a ee penn tris ae) os ch ke) SS TS eee Siete ae ot Soe) Eu? Gen en re 28
WSamBelvicsinuways=— tec ins cee Reet acces Rarer setae ence ea eee een ee Pseudonezumia pusilla
IObSPelvic-finiraysiS lA. sos Ge ees Gea oak leo See eke ae se, Ge ene 20
20a. Snout completely or almost completely naked; no tubercular scales at tip orlateralangles .............. 21
20b. Snout fully scaled or variously naked ventrally; tubercular scales present at tip and lateralangles .......... 23
AlavAnusiabutsranaltinstarnemoved from pelvichHinw qucme esc ie ciency cite eaeae ncn cnete ann 22
AllbwAnusiemovedttromanalitinsusually7clasermtolpelvic tinsess -ueeemensicentnralcr en cnencn cn amen ent nennmemanne [Kumba]
22a. Olfactory organ huge, length of posterior nostril about one-half diameter of orbit ........ [Macrosmia phalacra|
22b. Olfactory organ normal, posterior nostril much less than half diameter of orbit ..... . [Asthenomacrurus victoris |
28a. Anusicloserito:anal/omgini than to pelvic mSertions: = ms caussee) ci mel ceo) eestor cna 24
23b) Anusiclosemtoypelvic msertionsithan tojanalloneimi eae ener ieee cae) cane iene en 25
24b. Outer gill slit about 10% or less of head length; outer rakers of first arch rudimentary or absent; snout prominently
pPomted:dorsalitinibasemotelevated maesenrn- len nene nen wean nent nen nee Mataeocephalus acipenserinus
24b. Outer gill slit about 20% or more of head length; outer rakers of first arch tubercular and distinctly developed;
Snoutbluntyupturnedsdorsalifinibaseiclevated tean-seieu ee meine Cuenca nen nen Sphagemacrurus pumiliceps
25a. A stout suborbital shelf formed of 2 rows of coarsely spined scales; ventral edge of shelf forming a sharp ridge;
Snoutvanously makedionscalediventralliyarwenicm-m mente cit einen cn encmcmc tent) en nnn ace Nezumia
25b. Suborbital shelf covered with several rows of small, relatively unmodified scales, no sharp ridge developed;
undersideiof snoutimostly'scaled:- 3) 3 > @ sabes ee ea ew ee ee oh A ee 26
26a. Ventral region of body appearing to have swung far forward so that gill membranes unite below orbit, pelvic fins
belowioperclews Bic 2.2, sean dsl usy 3. Gr pie eee Te ROE un ee eRe Che co ee Lucigadus ori
26b. Ventral region of body more normal for a grenadier, with gill membranes uniting under interopercle, pelvic fins
below opercleor more posteriorly ses Ae) Eeyieyae ae ee ce oe tects aces ced cue Esa 27
Zjaxinner cillsrakersiof firstiarch Sirs sages fees mies eee Ls ee Ventrifossa
Zi palnnemsillerakersiofmirstarch) l2iorfewergsc. cucnacie mone neae in: neces Kuronezumia
28avAnUsnanmemoved trom/analeiinyclosemtOspelvichinlsaleaene mene nenenenenen ene) iene ncn nein Mataeocephalus sp.

ERO: ANUS MINEO NY AMET OEM 6 oo oo So oo OS DOE UBD OD ODS DOD OO Oo HS Coryphaenoides

IWAMOTO AND WILLIAMS: GRENADIERS OF WESTERN AUSTRALIA 115

FAMILY BATHYGADIDAE
Bathygadus Giinther, 1878

Although Howes and Crimmen (1990) recently reviewed the genus, it is obvious that a more
in-depth study using much more material is needed to resolve the many taxonomic problems within
this group. Iwamoto and Anderson (1994) and Iwamoto and Merrett (1997) discuss some of the
problems with Howes and Crimmen’s work.

Bathygadus spongiceps Gilbert and Hubbs, 1920
Fig. 2

Bathygadus spongiceps Gilbert and Hubbs, 1920:381-384, fig. 1 (Borneo; 1628 m). Howes and Crimmen
1990:189—190, table 4.
Bathygadus cottoides: Iwamoto and Merrett, 1997:7—8 (in part; New Caledonia).

DIAGNOsIS (for WA spec. only). — Barbel absent; first dorsal fin rays II,8—10; pectoral fin rays
114-117; pelvic fin rays 9 (occasionally 8 or 10); GR-I (outer) (4-6)+(19-21) = 23-27 total; caeca
19-27, length about equal to orbit diameter; orbit diameter 18-22% of HL, about 1.62.2 times into
interorbital, 2.4—2.8 into distance orbit to preopercle; interorbital width 32-40% of HL.

SPECIMENS EXAMINED. — WA: (Dark form): CSIRO H3017-08 (male, 67.2 HL, 325+ TL),
CSIRO H.3017-13 (male, 67.7 HL, 310+ TL), CSIRO H3017-14 (male, 78.2 HL, 385 TL); nw. of
Cape Leeuwin; 34°10'S, 114°16’E; 1,030 m; Akebono Maru No. 3, shot 17; coll. A. Williams;
24.XI1.89. CSIRO H2544-16 (2 females, 51.1-64.0 HL, 240+—315+ TL); Exmouth Plateau; 1,128
m; SS1/91/04. (Pale form): CSIRO H2615-05 (2 males, 79.0-85.5 HL, 425+—400 TL); w. of
Mandurah; 1,140 m; SS1/91/83. CSIRO H3017-12 (male, 84.2 HL, 435+ TL), CSIRO H3017-11
(female, 90 HL, 440+ TL), CSIRO H3017-07 (male, 79.4 HL, 440+ TL); nw. of Cape Leeuwin;
34°10'S, 114°16'E; 1,030 m; Akebono Maru No. 3, shot 17; coll. A. Williams; 24.XII.89.

FiGuRE 2. Bathygadus spongiceps Gilbert and Hubbs, 1920 (dark form). CSIRO H3017-13 (310+ mm TL) from continental
slope northwest of Cape Leeuwin, Western Australia, in 1,030 m. Fin rays partially reconstructed. Scale bar equals 25 mm.

116 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 3

COUNTS AND MEASUREMENTS (for WA spec. only).— 1D. II,8—10; GR-I (inner series)
3+(16—17), GR-II (2—3)+(15—16). Total length 240+—490+ mm; HL 51.5—107 mm. The following in
percent of HL: snout 29-32; postorb. 50-57; orb.-preop. 46-52; suborb. (fleshy) 14-18; up.jaw
52-56; ht. pmx. 15—17 (30-36% of len. pmx.); pmx. gap 9-19% of len. pmx.; len. outer gill rakers
11—14; len. gill filaments 6—10.

DESCRIPTION. — General features of fish best seen in Figure 2. Head bones and integument
relatively weak and readily damaged. Body scales highly deciduous, all lost in specimens examined;
scale pockets damaged in most specimens examined, often only faintly present. Head width about
one-half or more of head length; interorbital space broad, more than 1.6 times diameter of orbit, about
one-third or more of head length. Fin rays weak, most with tips broken off; outer pelvic ray in one
specimen thin, slightly prolonged, but not reaching to anus; spinous second ray of first dorsal thin,
flexible, slightly prolonged.

Color of dark form (see Comparisons) overall swarthy, region around belly, chest, head and gill
membranes black, fins dark dusky to blackish; oral, branchial, and abdominal cavities black; gill
rakers and arches dark, gill filaments pale. Pale form with body mostly pale, black regions of dark
form mostly grayish in pale form; fins dusky; chest and gular regions somewhat paler than in dark
form.

SIZE. — To at least 46 cm TL, possibly to 50 cm.

DISTRIBUTION. — Known from off the Kermadecs, New Caledonia region, New Zealand,
Australia, the Philippines, and Indonesia, usually in depths of approximately 900—1,500 m
(842—1,140 m off WA between latitudes 20°S and 35°S).

COMPARISONS AND REMARKS. — Bathygadus spongiceps appears to be a relatively widespread
species in the western Pacific and Australia. Iwamoto and Merrett (1997) erroneously thought their
specimens from New Caleodonia were B. cottoides Gunther, 1878, partly owing to the proximity of
their collection sites to the Kermadecs, the type locality for B. cottoides. However, most of their study
material (excluding CAS 90556 and CAS 90835) were B. spongiceps. They listed a number of
characters that differed significantly between their specimens and what has been called B. cottoides
from southern Africa (see Iwamoto and Anderson 1994). Subsequent examination (by TI) of
numerous specimens of B. cottoides from the Kermadecs and New Zealand (currently housed in
NMNZ) has revealed that B. cottoides is a small species, less than about 26 cm TL, with low pyloric
caeca counts (9-12), and few pectoral finrays (110-115, usually 112-114). This contrasts with B.
spongiceps, a large species attaining lengths of more than 46 cm, with 15—27 pyloric caeca, and
114-417 pectoral finrays. Gill raker lengths also differed, B. spongiceps having raker lengths of 9-13%
HL, compared with 14-19% in B. cottoides. We are uncertain of Howes and Crimmen’s (1990, table
4) counts of 6 and 12 pyloric caeca for paratypes of B. spongiceps. Gilbert and Hubbs (1920:383) in
their original description gave counts of 21, 16, and 17 for three paratypes, and Iwamoto and Merrett
(1997:8) recorded 15 each in two recently collected specimens from off eastern Luzon, Philippines.

Our specimens agreed well with specimens from the New Caledonia region (see Iwamoto and
Merrett 1997). We found in our WA collections two “forms” that we differentiated by color: a dark
form, with more prominent blackish scale pockets on the body giving a darker overall cast, most of
head, gill membranes, chest and abdomen black, fins blackish; and a pale form, with relatively pale
body and fins, the head, gular and gill membranes gray to blackish, the chest and abdomen light brown
to grayish. The overall physiognomy of the two forms seemed to differ, but we were unable to quantify
any specific differences. These differences are not sexual, as we have found both sexes represented
in each form, and the forms are sympatric, some of each having been taken in the same haul off Cape
Leeuwin (CSIRO H3017). We have been unable to determine whether these forms represent simple
color variants of a single species or are actually distinct species. More specimens and study are needed
before we can confidently state that either one or two species are represented in our WA material.

IWAMOTO AND WILLIAMS: GRENADIERS OF WESTERN AUSTRALIA LAG,

McMillan (in Gomon et al. 1994:344, fig. 305) recorded B. cottoides from the Great Australian
Bight. His description agrees well with that species, especially in pyloric caeca counts (7—11), low
pectoral ray counts (14-15), and small size (“at least 24 cm”). The presence of that species in the
Great Australian Bight suggests that it is likely to also occur off Western Australia at appropriate
depths.

Gadomus Regan, 1903

A world review of the genus (Howes and Crimmen 1990) included a key to eight species from
the Indo-Pacific region. Two species of the genus are represented in Western Australian waters. They
each differ slightly from known species, but because of inadequate descriptions and insufficient
comparative material, we had difficulty ascribing them with confidence to any known species. One
was sufficiently different that we have described it as new. The other we relate to a known species
but with qualifications. The only account of Gadomus from Australian waters is that of McMillan (in
Gomon et al. 1994:357, fig. 317), who provides description of an unnamed species from off southern
Australia that is similar to our new species. There is need of thorough revision of the genus based on
recent collections available in different museums. In the Comparisons sections for the two species,
data for other species are from the literature, as well as from personal observations of one of us (TI).
Peter McMillan (National Institute of Water and Atmosphere, Wellington, New Zealand) provided
valuable information on G. aoteanus, for which we are grateful.

KEY TO SPECIES OF GADOMUS FROM WESTERN AUSTRALIA

la. Total outer gill rakers on first arch 25—28; rakers relatively short, about equal to or slightly longer than longest gill fila-

ment; barbel 65-91% of HL; pyloric caeca 100—136; mouth, tongue, and gill cavity pale ...... G. sp. cf. colletti
1b. Total outer gill rakers on first arch 30-33; rakers relatively long, more than twice longest gill filament; barbel 48-56%
OH: pyloric cacca about 75: mouth: tongue; andgill'cavitydark ....%-:....-.2......- G. pepperi n.sp.

Gadomus sp. cf. colletti
Fig. 3

Gadomus sp. A: Williams et al., 1996:149 (listed).

DIAGNOSIS. — Gill rakers on first arch 25—28 total; snout long, 23-29% HL; head short,
postorbital length 45-50% HL and 47-51% preanal distance; barbel more than three times orbit
diameter, extending beyond end of jaws; about 100—136 pyloric caeca.

SPECIMENS EXAMINED (15 spec.). — WA: CSIRO H2596-03 (61.9 mm HL, 275+ mm TL);
wnw. of Green Head; 760-770 m; SS1/91/62. CSIRO H2550-09 (58.6 HL, 302 TL); 1,100—1,158 m;
SS1/91/11. CSIRO H3156-35 (5, 41.9-56.5 HL, 203+—268+ TL); n. of Monte Bello I.; 18°53’S,
115°59’E; 550 m; Surefire; coll. D. Evans; 16.1II.1992. AMS I.22809-010 (2, 46.5—53.9 HL, 224 290
TL); off Rowley Shoals; 492-584 m; SO2/82/19-21. AMS 1.22810-005 (59.2 HL, 287 TL); NW
Shelf; 736 m; SO2/82/22-24. LACM 43620-1 (51.2 HL, 240 TL); approx. 95 km wsw. of Rowley
Shoals; coll. N. Sinclair and P. Berry; 22. VIII.1983. NSW: AMS I.19862-006 (53.5 HL, 285 TL), se.
of Sydney; 768 m; K76-23-01. AMS 1.24659-001 (56.9 HL, 285 TL); off Broken Bay; 915 m;
K84-08-04. AMS 1.24979-011 (33.3 HL, 166 TL); e. of Broken Bay; 759 m; K84-16-04. AMS
1.29813-006 (38.3 HL, 195 TL); e. of Budgewoi; 722—768 m; K89-06-05.

COUNTS AND MEASUREMENTS (based on 10 spec.). — 1D. I,10—11; P. (116) 11720; V. 8; GR-I
(outer) 46+2 1—23; scales 1D. 6-7, 2D. 7—7.5; about 100 and 136 pyloric caeca in two specimens.

Total length 203+—302+ mm; HL 41.9-61.9 mm. The following in percent of HL: postrostral
75—79; internasal 16—18; interorb. 15—19; orb. 24-28; suborb. 10—14; orb.-preop. 45—49; up.jaw

118 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 3

FiGureE 3. Gadomus sp. cf. colletti, CSIRO H2596-03 (275+ mm TL), from west-northwest of Green Head, WA, in 760-770
m, Southern Surveyor station SS1/91/62. Drawn by Georgina L. Davis.

53-57; pmx. 49-54; post. nostril 4-7. barbel 65—91; pre-A. 190—215; pre-vent 171—191; V.-A.
78-104; isth.-A. 154-178; body depth 72-85; depth at A. 59-66; 1D.-2D. 7-8; ht. 1D. (88)137—171;
len. P. (143)179—-191; len. V. 86-97. The following in percent of pmx. length: pmx. ascending process
20.9-27.8; distance across midline between pmx. dentigerous surfaces 3.4—5.3. The following in
percent of pre-A.: HL 47-51; len. P. 87-95.

DESCRIPTION. — Head and body moderately compressed; tail slender (but damaged and short-
ened in most specimens). Snout blunt, rostral cartilage forming distinct knob at snout tip. Dorsal
profile rather straight over head; largest fish rather humped anterior to first dorsal fin where body
depth is greatest. Head ridges poorly developed and lacking modified body scales or scutes; bones of
operculum and snout particularly weak. Membranes of head thin (usually damaged). Orbit relatively
large, interorbital narrow. Mouth large and terminal, extending beyond posterior margin of orbit;
premaxillary ascending process well developed. Chin barbel prominent, fleshy, long, tapering to a
fine point, extending well beyond posterior margin of jaws.

Teeth minute, villiform, arranged in numerous rows, forming bands on both premaxilla and
dentary. Dentigerous bands of premaxilla broad, widest below anterior margin of orbit, separated by
a narrow gap at symphysis; dentigerous band of dentary relatively narrow, of uniform width.

Gill opening wide, first arch unrestricted; outer gill rakers on first arch well developed, slender,
flattened with tiny conical spines; inner rakers relatively short, about 2.0—2.5 into length of outer
rakers, clublike, slightly expanded distally with short curved spines; longest gill filaments of first gill
arch about 1.0—1.3 into longest outer gill rakers.

Scales large, thin, and deciduous. Gular and branchiostegal membranes unscaled. Light organ
absent.

All fins well developed; rays of second dorsal considerably longer than those of anal fin; second
ray of first dorsal and outer ray of pectoral (= second element) greatly extended; outer ray of pelvic
fin moderately extended, just reaching anal origin; extended rays of pectoral broad and fleshy; dorsal
fins separated by a narrow, rather variable space; pelvic fin bases broad and widely separated.

IWAMOTO AND WILLIAMS: GRENADIERS OF WESTERN AUSTRALIA it)

Body color pale pinkish on dorsal surface; pale silvery below midline from level of dorsal margin
of opercle; abdominal region between pelvic fin bases and vent bluish. Entire body speckled with
melanophores; these most dense on underside of head, gular membranes, around nostrils, along
posterior margin of opercle, and over ventral region anterior to pelvic fins. Buccal cavity, tongue, and
opercular cavity pale. Dorsal fin dark; pectoral and pelvic fins dusky.

SIZE. — To about 30 cm TL.

DISTRIBUTION. — Collected from the upper and midslope region (320-1,158 m) off the central
west and northwest coasts of Australia (latitudes 10°S—32°S) and off NSW (722-768 m).

COMPARISONS AND REMARKS. — The Indo-Pacific species of Gadomus can be separated into
two broad groups based on the number of lower-limb gill rakers on the first arch (Howes and Crimmen
1990). (Note: The “‘ceratobranchial” counts of Howes and Crimmen are actually counts for rakers on
the ceratobranchial + hypobranchial bones and are equivalent to our “lower-limb” gill raker counts.)
The low count (21—23) of G. pepperi differentiates it from four species with high counts (25-29): G.
aoteanus McCann and McKnight, 1980, G. capensis (Gilchrist and von Bonde, 1924), G. melanop-
terus Gilbert, 1905, and G. multifilis (Ginther, 1887).

Four species comprise the Indo-Pacific group with low gill rakers counts (17—23): G. colletti
Jordan and Gilbert, 1904 from the Pacific coast of Japan and the Kyushu-Palau Ridge, and G.
magnifilis Gilbert and Hubbs, 1920, G. denticulatus Gilbert and Hubbs, 1920, and G. introniger
Gilbert and Hubbs, 1920 from the Philippines and New Caledonia. Gadomus sp. cf. colletti is most
similar to G. colletti in having substantial overlap with the characters detailed by Howes and Crimmen
(1990) and Okamura (1970). Notably, it shares a high pectoral fin ray count (il 7120) that differen-
tiates G. colletti from the three other similar species, and it has strikingly similar external coloration
when fresh (see Okamura 1982, pl. 84). It differs from G. colletti in having a pale buccal cavity—lack-
ing the blackish pigmentation described by Okamura (1970, 1982). This character may be geographi-
cally variable and unimportant, but it is a distinct difference between the two western Australian
species of Gadomus and can be reliably used to separate them in the field. Because of this uncertainty,
and because G. colletti is known only from waters around Japan, we refer this material to G. sp. cf.
colletti pending a more detailed study. Gadomus sp. cf. colletti differs from G. introniger in its
relatively narrower interorbital (15-19% HL cf. 21-23% in our specimens and 20-22% as recorded
by Gilbert and Hubbs (1920) for the type specimens from the Philippines and Indonesia), and from
G. denticulatus in having more lower-limb gill rakers on the first arch (21—23 cf. 17-20). Gadomus
magnifilis is distinct among this group in having an extremely elongate second pectoral fin ray
(141—143% of preanal length cf. 87-95% in Gadomus sp. cf colletti) and a longer head (52-55% of
preanal distance cf. 47-51%).

Gadomus sp. cf. colletti can be most easily differentiated from the other Australian species, G.
pepperi by the lack of dark pigmentation of the buccal cavity and tongue, the relatively short outer
gill rakers on the first arch (similar or only marginally longer than the longest filaments cf. twice as
long), and the lower number of rakers on the first gill arch (25—28 total cf. 30-33).

Gadomus pepperi new species
Fig. 4

Gadomus sp. B: Williams et al., 1996:149 (listed).

DIAGNOSIS. — Gill rakers on first arch 30—33 total; a narrow space separating the premaxillary
dentigerous bands (5.3—6.6% of premaxillary length); first dorsal rays II,9Q—10; pectoral rays 116-119;
barbel less than three times orbit diameter, not extending beyond jaws. About 75 pyloric caeca.

SPECIMENS EXAMINED. — WA: HOLOTYPE: CSIRO H2541-04 (50.7 mm HL, 275 mm TL);
Exmouth Plateau; 913-914 m; SS1/91/2. PARATYPES: CSIRO H3008-12 (2, 56.1-66.0 HL, 280-365

120 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 3

iy RS ee Da Pe

Sis
Ayo cas
Sg?

FIGURE 4. Gadomus pepperi n.sp. Holotype, CSIRO H2541-04 (275 mm TL), from Exmouth Plateau, WA, in 913-914 m,
Southern Surveyor station SS1/91/02. Drawn by Georgina L. Davis.

TL); sw. of Albany; 35°25’S, 117°21'E; 842 m; Akebono Maru No. 3, shot 8; coll. A. Williams;
22.X11.1989. CSIRO H3017-01 (2, 53.0-60.0 HL, 290-306 TL); nw. of Cape Leeuwin; 34°10’S,
114°16’E; 1,030 m; Akebono Maru No. 3, shot 17; coll. A. Williams; 24.X1I.89 . CSIRO H2563-05
(3, 51.0-67.5 HL, 268-345 TL); w. of Quobba Pt.; 895—901 m; SS1/91/25. NSW: AMS I.24059-002
(4 of 7: 45.1-62.0 HL, 238+—383 TL); off Norah Head; 942-978 m; K83-09-02. QLD: AMS
1.20920-004 (54.7 HL, 290 TL); 10—11 miles ne. of Raine I.; 11°32'S, 144°10’E; 1,000 m.

COUNTS AND MEASUREMENTS (based on 8 spec.). — V. 8; GR-I (outer) 5—6(7)+24—26; about
75 pyloric caeca counted in one QLD specimen.

Total lengths 268—365+ mm; HL 50.7—-67.5 mm. The following in percent of HL: postrostral
73-76; snout 25—29; internasal 18—20; interorb. 18—23; orb. 23—26; suborb. 9-12; postorb. 49-53;
orb.-preop. 4449; barbel 48—56; pre-A. 175—193; pre-vent 163—180; V.-A. 63-81; isth.-A. 133-154;
body depth 72—83; depth at A. 57-64; 1D—2D. 3-6; ht. 1D. 157-208; len. P. 193-253; len. V.
132-175; post. nostril 3—6; up.jaw 57-62; pmx. 54-57. The following in percent of pmx.: pmx.
ascending process 19.8—24.3; distance across midline between pmx. dentigerous surfaces 5.3—6.6.
The following in percent pre-A.: HL 52—S7; len. P. 40—S2.

DESCRIPTION. — Head and body moderately compressed; tail slender (but damaged and short-
ened in most specimens). Snout blunt, rostral cartilage forming distinct knob at tip. Dorsal profile
rather straight over head; slightly humped at level of posterior margin of orbit. Bones of operculum
and snout particularly weak. Membranes of head thin, usually damaged. Orbit small to moderate;
interorbital relatively wide. Mouth extending beyond posterior margin of orbit; premaxillary ascend-
ing process well developed. Chin barbel prominent, fleshy and long, tapering to a fine point, not
extending beyond posterior margin of jaws.

Teeth minute, villiform, arranged in numerous rows forming bands on both premaxilla and
dentary. Dentigerous bands of premaxilla relatively narrow, of near-uniform width, separated by a
narrow gap at symphysis; dentigerous band of dentary relatively narrow and of even width.

Outer gill rakers on first arch long, slender, flattened, with tiny conical spines; inner rakers
medium length, about twice into length of outer rakers, clublike, expanded distally and armed with

IWAMOTO AND WILLIAMS: GRENADIERS OF WESTERN AUSTRALIA 121

medium-length, upward-pointing, curved spines; longest gill filaments of first arch about twice into
length of longest outer rakers.

Gular and branchiostegal membranes unscaled.

Second ray of first dorsal and upper ray of pectoral (= second element) greatly extended; outer
ray of pelvic fin extending well beyond anal origin; extended rays of pectoral slender. Pelvic fin bases
broad and widely separated.

Body color creamy yellow to sandy brown; color uniform above and below lateral midline;
abdominal region between pelvic fin bases and vent blue. Dusky pigmentation on anterior region of
jaws and lips, underside of head posterior to barbel, gular region, and around nostrils. Buccal cavity,
tongue, distal portion of branchiostegal membranes, and opercular cavity dark to black. Dorsal fin
dark to black; pectoral and pelvic fins dusky.

SIZE. — To more than 38 cm TL.

DISTRIBUTION. — Collected at upper- and mid-slope depths (817—1,500 m) off the west coast of
Australia (latitudes 20°S—35°S). Range extends eastwards to western Bass Strait (southeastern
Australia), NSW, and QLD.

ETYMOLOGY. — The new species is named to acknowledge Roger Pepper, the fishing master of
FRV Southern Surveyor and FRV Soela, for his contribution to many scientific fishing expeditions,
including those that provided much of the material for this study.

COMPARISONS AND REMARKS. — The outer, lower-limb gill raker count (24-26) of Gadomus
pepperi is similar to those of four Indo-Pacific species with a high raker count (25—29): G. aoteanus,
G. capensis, G. melanopterus, and G. multifilis (see Howes and Crimmen 1990, and note under
Gadomus sp. cf. colletti).

Gadomus pepper' is closest to G. multifilis from the northern and western Indian Ocean, sharing
a range of characters including a relatively longer head than in other species of Gadomus. However,
based on data from the holotype and [presumably] eight other specimens of G. multifilis provided by
Howes and Crimmen (1990:196, table 13), the new species has more rays in the first dorsal fin (11,9-10
cf. II,8(9) and pectoral fin (116-119 cf. “15—16”), and a narrower separation of the premaxillary
dentigerous areas (5.3-6.6% of premaxilla length cf. 8.2—13.1%). (Note that Howes and Crimmen
use premaxillary length, which should not be confused with our use of upper jaw length. The latter
is significantly longer as it includes the lengths of both premaxilla and maxilla.) Interestingly,
Okamura (1970) recorded II,9 dorsal rays and 19 pectoral rays in his description of a single specimen
of G. multifilis from the Philippines. However, the orbit diameter of that specimen is the same as the
interorbital width; in contrast, in the Australian species and in G. multifilis from the Indian Ocean,
the orbit is larger than the interorbital width.

Gadomus aoteanus from New Zealand is easily separated from G. pepperi by its relatively short
barbel (19-27% HL cf. 48-56%) and fewer pectoral fin rays (114-117 cf. 116-119). (Data from 12
specimens of G. aoteanus as provided by Peter McMillan, NIWA, Nov. 1996.) It should be noted that
the mensural characters for G. aoteanus in table 14 of Howes and Crimmen’s (1990) article are, for
the most part, wrong. The two authors used McCann and McKnight’s (1980:21) measurements
thinking they were in millimeters, but the measurements were actually expressed as a percentage of
the snout-to-anus length, viz, McCann and McKnight’s “standard length.” The pelvic fin ray count
that McCann and McKnight gave as eight is actually nine, from TI’s examination of the holotype,
NZOI 189, now deposited in NMNZ. Howes and Crimmen also gave the pyloric caeca count as
14-100+, numbers they obviously obtained from that given by McCann and McKnight for the entire
genus. McMillan has counted 18—25 caeca in 10 specimens, and 33 in an | 1th specimen.

Gadomus capensis from the Indian Ocean is also distinguished from the new species by its much
shorter barbel (2-7% HL cf. 48-56%). The high count of lower-limb gill rakers of first arch (27-29)
of the Hawaiian G. melanopterus separates it from G. pepperi (24-26 rakers) and the other species

122 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 3

in the “high-count” group. Gadomus melanopterus also has a shorter premaxillary ascending process
than the new species (18—20% of premaxillary length cf. 20-24%).

Gadomus pepperi can be most easily differentiated from the other Australian species, Gadomus
sp. cf. colletti, by the dark pigmentation of the buccal cavity and tongue (cf. unpigmented), the
relatively long outer rakers on the first gill arch (twice as long as the longest filament cf. similar length
or marginally longer), and the higher number of rakers on the first gill arch (total 30-33 cf. 25-28).
A possible third Australian species has been recorded as ““Gadomus sp.” by McMillan in Gomon et
al. (1994:357, fig. 317). That species is similar in many ways to Gadomus sp. cf. colletti, including
length of elongated fin rays, lengths of barbel, orbit, upper jaw, and snout, width of interorbital, and
counts of fin rays and gill rakers. However, it differs significantly in numbers of pyloric caeca (18-33
vs. about 75). A close comparison of more specimens of the two 1s in order.

Considerably broader collections from the Indo-Pacific are probably required before the identity
and distributions of Gadomus species from this region can be fully determined.

FAMILY MACROURIDAE
SUBFAMILY MACROURINAE
Caelorinchus Giorna, 1810

This genus, the largest in the family, contains more than 100 species, many of which are still to
be described. The Western Australian collections included representatives of at least 20 species, eight
of which are new.

KEY TO SPECIES OF CAELORINCHUS FROM WESTERN AUSTRALIA

(Names followed by an asterisk are species that have yet to be recorded from Western Australia,
but could be expected.)

lassnoutshortabluntausuallyaessiihamonroit diameter ss saeeue cnn cura cneinen ncn ners neice) en ee a 2
libsesnoutcacntely poimtedlonger thanjorbitidiameterss seen n nce ce cacti erie ccna 6
2a. Anal fin black on anterior 1/3 to 1/2, pale posteriorly; trunk completely encircled by a broad dark band; underside

oftheadiullyiscaled ams wee ete eee eee renee eee leet teen Gre cele, <n C. matamua
2b. Anal fin light dusky to pale or with blackish stripe extending to end of tail; trunk not encircled by a dark band;

underside of head naked or with a few scales posterior to mouth (in C. mirus) 2... 0... 2 ee ee 3
3a. Anterior dermal window of light organ extends forward to or beyond line connecting origins of pelvic fins (Fig.

20a) sr adistinct blackispotion pelViewins: 5 eiesnas ord 215 ee) kc tn oe ec eae cae ee Cee C. mirus
3b. Anterior dermal window falls short of line connecting pelvic fin bases (Fig. 18c); no distinct spot on pelvic fins .. 4

4a. Snout lacking naked areas dorsally behind leading edge (Fig. 18b); saddle marks on body prominent, dark
PP POR CR ARE OY, Cage e ae CS Ree eee tee eR ate ee POC ESEY Orel adous toes Occ eee eaeceS Estee we eee ee eG C. maurofasciatus
4b. Snout with naked, translucent areas dorsally behind leading edge (Fig. 20b); saddle marks on body rather faint or

alMOStlACKIM Oyo waxes cats ce Sies cet, ar lum a-aipweemsa ssp eran eee eaaiiStt ys (eakghh ut: Sytner 5
5a. Saddle marks obscure on trunk and nape; pale interspaces posteriorly on tail spotlike, especially viewed dorsally;
MOE MESA ly CCG <AMeOMIy 25 g00nponcnddomnoudonosdoconlbooongove C. parvifasciatus *
5b. Saddle marks faint but distinct along trunk and nape, with pale interspaces not spotlike and directed diagonally
downward and forward; median nasal process without dark pigmentation ........... C. amydrozosterus n.sp.
6a. Underside:ofiheadinaked (osx sere si oes 0 Poi aes) (on 5) 8 SES Chace ens page. ee er i
Gbs Underside‘of headiscaled@atleastipostenionlya-memens neuen eccrine Caen nent eee ear 14
7a. Light organ long, extends from anus to just behind isthmus; anus immediately before anal fin (Fig. 19a’) ....... 8
7b. Light organ extends to just forward of pelvic fin bases at most, but usually well behind; anus slightly removed
fromanalfiniorigin(Figs N6cs1'8c)) aes taekticotseebteRy ee bec) Gea) Cem 12
8a. Anterolateral margins of snout completely supported by bone; body scales small, 7—8.5 below origin of second dor-
SAUDI a ac ste oh, each cit. yo pent het ogc sae agers re eR ene a eg ae ee ee C. acutirostris

8b. Anterolateral margins of snout incompletely supported by bone; body scales moderate, 4.5—5.5 below origin of sec-
ondidorsalsfin™ t.55 Wes aie A ae pe a, OE oT ral nS atte ee ee 9

Ww

IWAMOTO AND WILLIAMS: GRENADIERS OF WESTERN AUSTRALIA 12

Ya. Spinules on body scales in discrete parallel rows (Fig. 8b); dorsal aspects of trunk covered with small to large spots or
elongated blotches; dorsal stripe absent or poorly developed ......-------.--5--555: C. pardus n.sp.

9b. Spinules on body scales in widely divergent to quincunx pattern (Fig. 8a, c); trunk either lacking spots, or large spots
and blotches in series forming broken longitudinal streaks; dorsal stripe present from near origin of second dorsal to

CEUN Pe loco ges akan rhe eR Ad AM RRO. 8 > oc Sc aU Ree a ee a Ts 10
10a. Upper jaw 22-24% of head length; snout length 49-50% ...........--2++555, C. gaesorhynchus n.sp.
10b. Upper jaw 30-36% of head length; snout length about 40-46% .. 2... 2... 2-2 eee ee ee eee 11
| 1a. Pectoral fin rays i13—i15; dark color on dorsum from nape to second dorsal essentially continuous, not broken into

multiple spots; gular membrane uniformly peppered with fine melanophores ..........--.. C. mayiae n.sp.

11b. Pectoral fin rays il5S—i18; dark color pattern of dorsum broken into multiple spots or blotches from below first dorsal
fin to origin of second dorsal fin; melanophores on gular membrane forming striated or reticulated pattern
Pee a 8 es, Sone cs oe eee ce eS es ae Gl elas el es Se gies BiG C. argentatus

12a. A large prominent pectoral spot (usually above and behind pectoral fin base) present atall sizes .......... 13
i2babectoralispotabsentes weer) ene ht eae em a ew rer eae tno h ery t-nshstn- lect) Toate C. goobala n.sp.
13a. Pectoral spot close to pectoral fin base; spot higher than wide ..........-.--.-5+5--55:5 C. maculatus
13b. Pectoral spot removed from pectoral fin base by 4 or 5 scale rows; spot wider thanhigh .........- C. thurla n.sp.
14a. Second dorsal fin about as high anteriorly as anal fin; spinules on body scales fine, needlelike, in 10 or more parallel
HOWIS “Ss dca ae SEO. SON ne eesti NS: crocu cet idiln. Wo. /6. oem mony Oo RcmOmoMoapecmr coms mciRA icant C. innotabilis
14b. Second dorsal fin much lower than anal fin anteriorly; spinules on body scales not fine and needlelike, in fewer than 10
TONY SHR ec catete MES, Dearest, RT Seat MBS Cage Ia aoeigs Jel oP eh te yn Remit ck, Rule Pee Ma st Iowa 15
15a. Anterolateral margin of snout completely supported by bone; a short dermal window of light organ present before
anus anus sichtlyiremoved fromyanal fin.ongini a5) 14 seerrnciegs © ase ees) 2) es | es Sek 16
15b. Anterolateral margin of snout incompletely supported by bone; no dermal window of light organ visible; anus
mamediatelyDelore analfinic.. -.- = 4 = Men eneery meme ge fe. Se sg eee so mien reel soe een 18
16a. Orbit 2.0—2.5 into snout length; 8-10 gillrakers on first arch, 9-10 rakers on inner side of second arch; nasal fossa
Almost Eman SeMeel 55 go Gaon cone Soo eHo dos bond noosnadoduaaaogac C. macrorhynchus
16b. Orbit 1.3—2.1 into snout length; 7-8 gillrakers on first arch, 7-9 rakers on inner side of second arch; nasal fossa naked
toysparselyascaled anteniOnlyanGaven tral emer ene gwen ewes tie mcwe renter eee efi) f=) ees eo- 17
17a. Color overall dark, swarthy to chocolate-brown in adults, grayish in juveniles; pectoral fin rays 116417... C. smithi
17b. Color dorsally light brownish gray, ventrally pale to whitish; pectoral finraysil7a19 ......... C. charius n.sp.
18a. Free neuromasts on head prominent, appearing as series of black dots; nasal fossa naked; triangular area bordered by
nasal MOssavonbitandsuborbitalemd Sena kc teem meni mene eer de mee A C. trachycarus
18b. Free neuromasts on head not conspicuous; nasal fossa partially or wholly scaled; area bordered by nasal fossa, orbit,
amel swlyonoall mtlementsel 5G alo bobo bod ec ood soe be ecuprcoocos goods odhaaance 19
19a. Color light brown to grayish; mouth, lips, gums pale; most trunk scales with 2 or more incomplete spinule rows, not
extending postenionytoiedge qiscalem 0: \Uae ce eee Ses Sr ele eGe «soe ss eck C. acanthiger
19b. Color overall dark brown to black; mouth, lips, gums dark-dusky to black; most trunk scales with all spinule rows com-
plastoculamirecciy 4 glgimogob om ooo ace oon obo ebb odo Mame Oe pomp sao 6 C. lasti n.sp.

Caelorinchus acanthiger Barnard, 1925
Fig. 5

Coelorhynchus acanthiger Barnard, 1925:502 (off Cape Point, South Africa; 841 m).

Coelorhynchus pseudoparallelus Trunov, 1983:895 (Namibia; Walvis Ridge, Discovery Tablemount;
800—1,800 m).

Caelorinchus acanthiger; McMillan in Gomon et al. 1994:350, fig. 310 (southern Australia). Williams et al.
1996:148 (WA).

DIAGNOSIS. — Snout of moderate length, about 40-43% HL, with dorsal profile straight to
slightly concave and tipped with a sharp, narrow scute; anterolateral margin not fully supported by
bone; orbit diameter 27-30% HL; upper jaw extends to below midorbit. A small, inconspicuous, black
scaleless fossa (length much less than posterior nostril) immediately anterior to anus. Underside of
head covered with small, fine, embedded scales having short, erect clusters of spinules; nasal fossa
usually entirely naked; body scales with a median keellike row of broad-based, enlarged spinules
flanked by two to four short rows of smaller spinules. Pyloric caeca 9-12.

SPECIMENS EXAMINED. — WA: AMS 1.31181-013 (2, 86.9-91.3 mm HL, 340-341 mm TL);
sw. of Shoal Point; 853-854 m; SS1/91/41. AMS 1.3177-009 (83.1 HL, 300+ TL); off Shark Bay;

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Ficure 5. Caelorinchus acanthiger Barnard, 1925. Lectotype, SAM 12632 (390+ mm TL), from off Cape Point, South Africa, in 460 fathoms. Fins and scales partially

reconstructed. Scale bar represents 25 mm.

IWAMOTO AND WILLIAMS: GRENADIERS OF WESTERN AUSTRALIA 125

996—1,009 m; SS1/91/44. AMS I.31180-012 (112.6 HL, 390+ TL); nw. of Shoal Point; 945-960 m;
SS1/91/48. NMV uncat. (3, 86.2-92.9 HL, 290-347+ TL); s. of Cape Leeuwin; 870-920 m;
SS1/91/96. CSIRO H3002-03 (130 HL, 450+ TL); se. of Albany; 35°23’S, 118°27'E; 1,030 m;
Akebono Maru No. 3, shot 2; coll. A. Williams; 21.XII.1989. CSIRO H3008-08 (4, 88.5—-108 HL,
335-355+ TL); sw. of Albany; 35°25'S, 117°21'E; 842 m; Akebono Maru No. 3, shot 8; coll. A.
Williams; 22.XII.1989.

COUNTS AND MEASUREMENTS (for 12 WA spec.). — 1D. II,8—9; P. (117) 118120; GR-I (inner)
2+6; GR-II (outer/inner) 0+6 / 2+(6—7); scales 1D. 7 or 8, midbase 1D. 5—7, 2D. 6.5—7.5, lat.line
33-39 (43).

Total lengths 290-450+ mm; HL 83-130 mm. The following in percent of HL: preoral 3742;
internasal 19-20; interorb. 22—24; suborb. 15—17; postorb. 30-32; orb.-preop. 32—36; up.jaw 23-26;
barbel 7-11; gill slit 11—12 (4 spec.); body depth 42—58; 1D.-2D. 18-31; ht. 1D. 36—50; len. P. 32-37;
len. V. 31—38; len. post. nostril (6) 7—8 (9).

SIZE. — Attains at least 50 cm.

DISTRIBUTION. — Widespread from southern Africa across Indian Ocean to Australia and New
Zealand. Relatively common in temperate waters of Australia, from NSW, VIC, TAS, SA, WA.

COMPARISONS AND REMARKS. — Caelorinchus acanthiger is most likely to be mistaken for C.
trachycarus from southern Australia and New Zealand, but can be distinguished by its smaller body
scales and finer scale spinulation that imparts a generally smoother texture to the head and body, the
lack of notably enlarged and coarsened scales on the head exclusive of the ridges, fewer pyloric caeca,
and paler body and fin color lacking violet/purplish tinge on naked areas. Caelorinchus mycterismus
is also closely similar but has a longer snout with a pronounced upturned tip (snout profile relatively
straight in C. acanthiger). In this regard, Western Australian specimens of C. acanthiger have a
slightly upturned snout compared to specimens from NSW, but agree well in all other features. We
attribute this slight difference to geographic variation. Compared to C. acanthiger, C. lasti has a darker
overall color, broader spinules on scales of snout, scales present ventrally on nasal fossa, and a number
of different proportions in head measurements, which are compared in Table 1.

Caelorinchus acutirostris Smith and Radcliffe, 1912
Fig. 6a

Coelorhynchus acutirostris Smith and Radcliffe in Radcliffe, 1912:134—136, pl. 30, fig. 2, text-fig. 10 (Philip-
pines between Cebu and Bohol, 291 m; Albatross sta. 5418; holotype, USNM 72947). Gilbert and Hubbs,
1920:512—514 (descr.; Cebu, Bohol, Mindanao; 291—320 m).

Coelorinchus sp. 1: Arai in Gloerfelt-Tarp and Kailola, 1984:85, fig. p. 84, and photograph of specimen without
caption on p. 82 (s. Indonesia and nw. Australia).

Caelorinchus acutirostris: Iwamoto and Merrett, 1997:485—486, fig. 6a (specimen from New Caledonia,
Queensland, Philippines).

DIAGNOSIS. — Snout notably slender, attenuate, and sharply tipped, its length 47-58% of HL:
anterolateral margin fully supported by bone; orbit diameter 20-23% of HL; upper jaw short, length
17-22% HL, restricted laterally, extending to below posterior one-third of orbit; barbel short, fine,
5—7% of HL. Premaxillary teeth band short, broad; mandibular band longer and more narrow;
subopercle lacking a prolonged narrow flap. Light organ a long, blackish, scale-covered streak
extending from just behind isthmus to front of anus. Underside of head completely naked; nasal fossa
usually naked, but small scattered scales in some specimens; body scales small, their exposed field
covered with conical erect spinules in 5—7 slightly divergent rows, 7—8.5 scale rows below origin of
second dorsal fin, 42—58 lateral line scales over a distance equal to predorsal length. A dark streak
(sometimes faint) on belly extending horizontally from pectoral fin base to above anal fin origin; a
curved dark streak (often faint) passing below base of first dorsal fin from nape to second dorsal;

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TABLE |. Comparison of characters of Caelorinchus acanthiger, C. trachycarus, C. mycterismus, and C.

lasti.

C. acanthiger C. trachycarus
nasal fossa scales naked or sparse naked
snout length (%HL) 40-43 39-49
orbit (Y%HL) 2730 22-30
snout/orbit 1.3—1.6 1.4-1.9
upper jaw len (%HL) 23-26 17-24
barbel (“oHL) 7-11 4-8
preoral len. 37-42 36-48
internasal width (%HL) 19-20 18-23
interorbital width (Y%HL) _ 22-24 20-27
gill slit 11-12 —-
pyloric caeca 9-12 7-9

C. mycterismus*

sparse-extensive
45-51
24-29
1.82.0
15-26
4-10
43-5]
16-20
19-24
9-11
8-11

C. lasti

sparse
37-38

* Data for C. mycterismus from McMillan and Paulin (1993) and current specimens examined.

FIGURE 6. (a) Caelorinchus acutirostris Smith and Radcliffe, 1912. MNHN 1996-961 (44.6 mm HL), from off New
Caledonia in 415—435 m. (From Iwamoto and Merrett 1997, fig. 6a.) (b) Caelorinchus argentatus BMNH 1996.7.19.2 (61 mm
HL), from off New Caledonia in 415435 m. (From Iwamoto and Merrett 1997, fig. 6b.)

IWAMOTO AND WILLIAMS: GRENADIERS OF WESTERN AUSTRALIA 127

chest and vent areas darkly punctulate to black; anterior 10—15 rays of anal fin blackish; suproccipital
scute well developed; second dorsal fin moderately well developed for genus.

SPECIMENS EXAMINED. — WA: CSIRO CA405; NW Shelf, Mermaid Reef, Rowley Shoals;
17°24'S, 119°54’E; 305-322 m; FRV Courageous; 10.V1.1978. CSIRO CA344; NW Shelf; 1978 (no
other data). CSIRO CA345 (76 HL, 270+ TL); NW Shelf; FRV Courageous; V.1978. AMS
1.23423-011 (59.0 HL, 216+ TL); NW Shelf; 376 m; SO4/82/leg1. AMS 1.22825-13 (4, 61+—79.5
HL, 200+—252 TL); NW Shelf; 300-326 m; SO2/82/43.44. WAM P.26209-003 (3, 45.8-67.0 mm
HL, 142—202+ mm TL); 225 km nnw. of Port Hedland; 297-330 m. Philippines: USNM 72947
(holotype, 205 mm TL); Philippines between Cebu and Bohol; Albatross sta. 5418; 291 m. (Also,
specimens cited in Iwamoto and Merrett 1997)

COUNTS AND MEASUREMENTS. — 1D. II,(7)8—9(10); P. 115-117; total GR-I (inner) (1—2)+(5—6),
6-8, GR-II 5—6/(1—2)+(5-6), 6-8; scales 1D. 7—9, midbase 1D. 5—7.

Total length 110+—238+ mm; HL 38.3—69.2 mm. The following in percent of HL: preoral (40)
4352: internasal 13—20; interorb. 18—22; suborb. 1 |—13; postorb. 23—30; orb.-preop. 27—32; gill slit
8-12 (13); pre-A. 126-153; V.-A. 26-44; isth.-A. 48-68; body depth 36-57; 1D.-2D. 8-13; ht. 1D.
32-42; len. P. 28-41; len. V. 23—34; post. nostril 4-9.

SIZE. — Largest specimen known is 24 cm TL.

DISTRIBUTION. — Known from the Philippines, off WA and QLD, and off New Caledonia;
probably occurs throughout Indonesia.

COMPARISONS AND REMARKS. — Caelorinchus acutirostris is a peculiar species among the
Caelorinchus of Iwamoto’s Group IV (Iwamoto 1990) in that it has a long light organ typical of the
group and a complete bony support of the anterolateral snout margin. All other Group IV macrourids
have the anterolateral snout margin incompletely supported by bone. The small scales and distinctive
body markings in fresh specimens coupled with other diagnostic features make the species unlikely
to be mistaken for any other.

Caelorinchus amydrozosterus new species
Fig. 7

Caelorinchus sp. 2: McMillan in Gomon et al., 1994:348, fig. 308 (descr.; VIC, 300-600 m).
Caelorinchus sp. A: Williams et al., 1996:148 (WA).

DIAGNOSIS. — Snout short, anterolateral margin incompletely supported by bone; orbits longer
than snout and postorbital. Naked fossa of ventral light organ (ADW) (Fig. 7c) relatively large,
anterior end reaches about to line connecting pelvic fin insertions. Underside of head and nasal fossa
naked; broad, naked, translucent areas dorsally behind leading edges of snout (Fig. 7b). A series of
scales on midline of nape with median spinule row elevated into low crests, forming weak but distinct
ridge from supraoccipital scute to dorsal fin. A series of 12—13 faint saddle marks, the anterior four
directed anteriorly downwards. Pyloric caeca 15—16.

SPECIMENS EXAMINED. — HOLOTYPE: NMV A3405 (52.5 mm HL, 200+ TL); 42 km sw. of
Portland; 297-334 m. PARATYPES: WA: NMV A7108 (5 of 7, 41.1-48.5 mm HL, 185+—221+ mm
TL); 100 km se. of Esperance; 34°41.1'S, 122°27.0'E; 717-710 m; Saxon Progress; field no. RP-1;
14. VIII.1988. CAS 79576 (formerly NMV A6191) (4, 41.3-50.2 HL, 194-226 TL); 80 km ssw. Of
Esperance; 34°34’36"S, 121°32'48”E; 504-477 m; Saxon Progress; coll. RJP; 15.VIII.1988. CSIRO
H2604-02 (39.4 HL, 180+ TL); sw. of Ledge Pt.; 512 m; SS1/91/70. CSIRO H2605-03 (37.9 HL,
183 TL); nw. of Rottnest I.; 485 m; SS1/91/71. CSIRO H2607-01 (41.1 HL, 197+ TL); Rottnest
Canyon; 550 m; SS1/91/73. CSIRO H2618-04 (39.7 HL, 193 TL); 430 m; SS1/91/87. AMS E3583
(51.6 HL, 220+ TL), E3585 (52.0 HL, 245+ TL), E3586 (48.9 HL, 195+ TL); Great Australian Bight
se. from Eucla; Endeavour; 6.V.1913. VIC: NMV A3405 (2, 43.3-52.5 HL, 200+—210 TL); 42 km

128 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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Ce

A f,
OAAN,

PII IGA

C

FiGuRE 7. Caelorinchus amydrozosterus n.sp. Holotoype (NMV A3405, 200+ mm TL) from 42 km southwest of Portland,
VIC, in 297-334 m. (a) Lateral view (fins and scales partially reconstructed); (b) dorsal view of head to show snout shape and
naked areas behind leading edge of snout; (c) ventral view of abdomen to show shape and location of light organ and anus.
Scale bar represents 25 mm.

sw. of Portland; 297-334 m. NMV A803 (2, 44.2-46.8 HL, 198+—225+ TL); Bass Strait off Portland;
38°50’S, 141°46’E; 549 m; coll. M. Gomon (MFG-15); 6.III.1980. NSW: AMS 1.15975-036 (42.9
HL, 200+ TL); 35 miles se. of Newcastle; 33°11'S, 152°23’E; K71-08-05. SA: AMS 1.871 1-006 (3,
40.7-44.6 HL, 182+—214 TL); “off southern Australia,” Dmitry Mendeleev; 28.11.1976.

COUNTS AND MEASUREMENTS. — ID. II,9-10, P. (115) 116-118; GR-I (inner) (1—2)+7, GR-II
(outer/inner) 7—8 total /2+(7—8); scales 1D. 5—6, midbase 1D. 3.5—4.5, 2D. 3.5—4.5, lat.line 23—27;
caeca 15-16.

Total length 177-245 mm; HL 37.9-52.0 mm. The following in percent of HL: snout 29-34;
preoral 27-35; internasal 21—26; interorb. 17—22; orb. 38-43; suborb. 15—18; postorb. 30-33;
orb.-preop. 30-34; up.jaw 26-30; barbel 8—14; gill slit 13-17; pre-A. 139-159; V.-A. 66-81; body
depth 56—70; 1D.-2D. 18-28; ht. 1D. 63—81; len. base 1D. 22—33; len. P. 54-61; len. V. 46—S1; post.
nostril 6—13 (13); len. ADW 14-21.

DESCRIPTION. — General features of fish seen in Figure 7. Snout short, acute in lateral profile,
but blunt with broadly convex sides in dorsal view; terminal scute small, broad; subopercle forming
a short, acute flap ventrally; mouth small, upper jaw extends posteriorly to below midorbit or beyond.
Chin barbel short, length less than least width of interorbital space.

Scales typical of other members of clade (C. fasciatus, C. maurofasciatus, C. parvifasciatus, C.
mirus, etc.); those on body with 8-12 (more in larger specimens) close, parallel rows of short,
recumbent spinules. A modified median series on nape (see Diagnosis).

IWAMOTO AND WILLIAMS: GRENADIERS OF WESTERN AUSTRALIA 129

Fins well developed, typical of genus; high first dorsal lacking prolonged spinous ray; second
dorsal rays low throughout. Pelvic fin with slightly prolonged outer ray.

Naked fossa of ventral light organ (Fig. 7c) separated from periproct region by a gap of several
scale rows, extending to line connecting pelvic fin insertions (black streak extends farther forward,
however); anus at anal fin origin.

Anterior four saddle marks on body directed anteriorly downwards, Ist saddle over nape,
separated by pale interspace 2 rows wide running from origin of first dorsal through hind margin of
occipital region, terminating at upper margin of opercle; 2nd oblique pale interspace originating under
midbase of first dorsal; 3rd pale interspace originating under origin of second dorsal; saddle bands
more pronounced posteriorly on tail, 8th saddle extending to anal base, pale interspaces not “spotlike”;
underside of head, including jaws and gill membranes, and most of body pale creamish to white, trunk
dark only around ventral light organ; opercle with a small black blotch; first dorsal fin dark dusky
with pale base and posteriormost rays; pectoral fin clear to light dusky; pelvic fin with blackish blotch;
anal fin dusky with fine, scattered speckling; anterior rim of orbits distinctly black; lips and gums
cream, mouth cavity dark.

SIZE. — To about 25 cm.

DISTRIBUTION. — Broadly distributed from WA, SA, VIC, and NSW. So far not known from
TAS. Depth distribution from about 297 m to 717 m.

ETYMOLOGY. — From the Greek amydros, indistinct, obscure, and zosteros, belt, girdle, in
reference to the faint bands on the body.

COMPARISONS AND REMARKS. — Caelorinchus amydrozosterus is another species of the C.
fasciatus group. It is most likely to be confused with C. parvifasciatus because of its small size, similar
general appearance, and overlapping distribution in Australia. The two are readily distinguished,
however, by C. amydrozosterus having a larger ADW; more pronounced banding pattern on body,
with pale interspaces not spotlike and angled diagonally down and forward (at least in first four);
fewer pyloric caeca (15—16 cf. 22—28, fide McMillan and Paulin 1993:827); a small dark blotch on
opercle, not extending onto subopercle and preopercle; and a series of modified scales along midline
of nape. Caelorinchus maurofasciatus differs in having the dorsal surface of the snout entirely scaled;
a smaller light organ; more pyloric caeca (22-32 fide McMillan and Paulin 1993:826); more
prominent saddles, the anteriormost saddle ending at origin of the first dorsal fin and the pale
interspaces directed ventrally and posteriorly; and a dark marginal stripe on the anal fin. Caelorinchus
fasciatus differs in having a smaller ADW; larger, more prominent scutelike scales on nape that form
a high, sharp, median ridge anterior to first dorsal fin; and a completely scaled dorsal snout surface.
Caelorinchus cookianus from New Zealand differs in having darker saddles, no modified scales on
the nape, and 19-27 pyloric caeca (McMillan and Paulin 1993:827). Caelorinchus mystax and C.
bollonsi from New Zealand waters differ from C. amydrozosterus in having many more pyloric caeca,
among other differences (see McMillan and Paulin 1993:827).

Caelorinchus argentatus Smith and Radcliffe, 1912
Fig. 6b

Coelorhynchus argentatus Smith and Radcliffe, in Radcliffe 1912:137, 138 (holotype USNM 72949, Philippines,
vicinity of Jolo, 582 m).

DIAGNOSIS. — Snout moderately long, 40-44% of HL, with a slight humplike rise in dorsal
profile over nostril; anterolateral margins not fully supported by bone; snout scales anterolaterally
overlap onto ventral surfaces; orbit diameter 22-27% of HL; upper jaw about one-third of HL,
extending posteriorly to below hind margin of orbit; barbel 6—-14% of HL; subopercle without a
slender projecting tab. Broad, median-ventral streak of light organ extends from anus onto chest just

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behind isthmus. Underside of head completely naked (except for overlapping scales on snout); nasal
fossa naked, naked area extending posteriorly onto suborbital shelf below anterior half of orbit; body
scales thin, spinules short, fine, conical, in slightly divergent rows to somewhat quincunx pattern.
Second spinous ray of first dorsal fin blackish over distal three-quarters; median nasal bone blackish;
body markings variable, often faint, but in most specimens, an elongate, horizontal blotch behind
occipital region over anterior end of lateral line, area below dorsal interspace with small blotches, a
narrow tapered stripe from below origin of second dorsal to end of tail, with anterior end somewhat
club-shaped; gular membrane with finely striated or reticulated pattern.

SPECIMENS EXAMINED. — WA: NTM S.12728-028 (83.1 mm HL, 275+ mm TL); sw. of Rowley
Shoals; 420 m. NTM S.12727-007 (79.5 HL, 250+ TL); sw. of Rowley Shoals; 410m. AMS
I.22808-037 (2, 71.6-78.0 HL, 224+—270 TL); NW Shelf, 220 km n. of Port Hedland; 420 m;
SO2/82/17-18. AMS I.22821-010 (3, 63.8-76.2 HL, 229-265+ TL); NW Shelf, ne. of Port Hedland;
298-320 m; SO2/82/36-38. AMS I.23425-015 (4, 66.6—93.4 HL, 250—326+ TL); NW Shelf; 298-320
m; SO4/82/leg1. NT: NT S.12458-002 (2, 64.6-76.3 HL, 242-250 TL); Arafura Sea, se. of
Tanimbar I.; 9°17’S, 131°08'E; 297 m; 5.VII.1988. NTM S.13146-017 (63.5 HL, 222+ TL); Arafura
Sea, n. of Bathurst I.; 9°45’S, 130°14’E; 265 m; 6.11.1990. Timor Sea: AMS 1I.21804-002 (est. 66
HL, 235+ TL); 190 m; 1979. QLD: AMS 1.22469-001 (93 HL, 300 TL); Great Barrier Reef, off No
Name Reef; 14°39’S, 145°43’E; Samarai; 30.X.1981. AMS 1.25814-003 (37 HL, 135 TL); n. of
Townsville; 260-264 m; SO1/86/29. AMS I.25817-002 (38 HL, 140 TL); n. of Townsville; 296 m;
SO1/86/35. AMS 1.25826-004 (42.1 HL, 147 TL); n. of Townsville; 300 m; SO1/86/53. CSIRO
H691-02 (79.0 HL, 279 TL); Queensland Trough, e. of Hinchinbrook I.; 402 m; SO6/85/47. Philip-
pines: USNM 72949 (holotype, 93.2 mm HL, 365 mm TL), vicinity of Jolo; A/batross sta. 5172; 582
m. CAS 34436 (82.7 HL, 300 TL); Batangas Prov., Balayan Bay, se. of San Pedrina Pt.; 144-170 m;
coll. J. E. Norton; 22. VII.1966. CAS 34857 (59.3 HL, 228 TL); se of Pagapas Bay; 326-362 m; coll.
J. E. Norton; 4. VITI.1966.

COUNTS AND MEASUREMENTS (see Tables 3—5 for additional measurements). — ID. II,8—9 (10);
P. i15—i17 (i18); total GR-I (inner) 7-9, GR-II (outer/inner) 5—7/8—9; scales 1D. 6-8, midbase 1D.
3.5—5.5, 2D. 4.5—5.5(7.5), lat.line 35—48; caeca 9-13.

Total lengths 135-365 mm; HL 42.1—93.2 mm. The following in percent of HL: preoral 27—37;
internasal 18—23; interorb. 22—27; suborb. 11—14; up.jaw 30-36; gill slit 13-18; pre-A. 137-167;
V.-A. 39-60; isth.-A. 71-96; body depth 43-60; 1D.-2D. 14-29; base 1D. 17-21; len. P. 32-43; post.
nostril 49,

DESCRIPTION. — Iwamoto and Merrett (1997) provided a detailed description of C. argentatus.
General proportions and shapes of fins, head, and body much as in C. mayiae and C. pardus.

Body scales of C. argentatus relatively deciduous; few scales remain in specimens examined
except on head, chest, and over dorsal aspects of trunk. Scales dorsally below origin of second dorsal
fin (Fig. 8a) having short, conical, slightly recurved spinules, arranged in irregularly divergent rows
to quinqunx pattern. Posteriormost spinules in each row scarcely extend beyond edge of scale. Scales
ventrally on chest with broader, flatter spinules, more triangular in shape; few remaining scales over
abdomen and above anal fin origin with short triangular spinules, similar to those illustrated for C.
kamoharai Matsubara, 1943 by Okamura (1970, fig. 64). Scales over snout and over interorbital space
rather thin, but abundantly covered with short, conical spinules. Underside of snout with tiny black
punctations and small, black, hairlike papillae, as also found in C. kamoharai. Body markings rather
similar to those of C. mayiae, but more spotted and not as prominent. Blotchy markings dorsally on
nape and trunk fairly extensive but blotches broken below dorsal interspace. Elongated horizontal
blotch behind occipital region present in most specimens, but faint in others. Thin, tapering dark stripe
beginning below origin of second dorsal fin extends length of tail. Anterior end of dorsal stripe
expanded and directed ventrally, but fading before joining darkened area of periproct. Lateral aspects
of abdomen peppered with large dots under a thin silvery surface covering; ventrally on abdomen,

IWAMOTO AND WILLIAMS: GRENADIERS OF WESTERN AUSTRALIA 131

FiGURE 8. Scanning electron micrographs of scales of seven species of Caelorinchus: (a) C. argentatus; (b) C. pardus; (c)
C. mayiae; (d) and (e) C. charius; (f) C. goobala; (g) C. maculatus; (h) and (i) C. mycterismus. All scales except (e) and (h)
are from the region below or near the interspace of the first and second dorsal fins; (e) and (h) are from the underside of the
snout.

132 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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from periproct region to anterior end of pelvic fin bases, dots sparse to moderate. Pelvic fin bases and
pelvic girdle whitish, with scattered black punctations. Dark chest strongly contrasts with pale belly.
Pelvic fins dark at base, tip, and along outer ray, but pale in middle. Segmented first dorsal rays with
faint trace of broad darkish stripe across middle. Anal fin somewhat blackish on anterior few rays,
but otherwise clear to light dusky posteriorly, with no blackened margin. Barbel dark to dusky, paler
distally. Maxillary and lips densely pigmented.

SIZE. — To about 37 cm TL.

DISTRIBUTION. — Widespread in warm-water regions of Australia, from Northwest Shelf (about
118°E) to Arafura Sea (to 133°39'W) to QLD, in 260-420 m. Also known from Philippines,
Indonesia, and New Caledonia.

COMPARISONS AND REMARKS. — Caelorinchus argentatus is so similar to C. mayiae that
meristic and morphometric differences are minimal. Selected characters of the two species are
compared in Tables 2-4. The generally fewer pectoral fin rays (Table 3) are helpful in distinguishing
C. argentatus from C. mayiae. The scales atop the snout in C. mayiae are much weaker and sparser
than in C. argentatus, giving the snout a more transparent and smoother appearance. Color patterns
appear to be the best features for distinguishing the two species. In C. mayiae the dorsum from nape
to second dorsal fin is continuously dark, whereas in C. argentatus the dark color is broken into smaller
blotches, especially under the space between the dorsal fins and around the base of the first dorsal
fin. (The dorsal color patterns are faint or almost absent in Philippines and some large Queensland
specimens of C. argentatus.) In C. argentatus the pigmentation on the gular membrane of adults is
in the form of fine striations or reticulations, in contrast to the uniformly peppered appearance in C.
mayiae. The pelvic fin base and pelvic girdle of C. mayiae are almost entirely without pigmentation
except along the edges, whereas those areas in C. argentatus have large scattered melanophores. In
specimens of C. argentatus, the underside of snout is densely covered with short, minute, hairlike
papillae (in addition to short, small, paired papillae), whereas in C. mayiae, these hairlike papillae are
sparse or almost entirely absent.

Caelorinchus argentatus differs from C. pardus in scale spinulation, color pattern on dorsum,
and in having a rather dense covering of papillae on the underside of head.

Caelorinchus denticulatus Barnard, 1925 from the western Indian Ocean, and four species from
southern Japan and the East China Sea (C. formosanus Okamura, 1963; C. longissimus Matsubara,
1943; C. kamoharai Matsubara, 1943; and C. multispinulosus Katayama, 1942) also belong in this
species clade. They are all very similar in general features to C. argentatus but can be distinguished
from C. argentatus by their gular membrane being uniformly punctate and not striate or reticulate.
Caelorinchus denticulatus is further distinguished by having somewhat fewer pectoral fin rays
(114-115); C. formosanus by scales on the underside of head posterior to the lower jaw; C. /ongissimus
by more pyloric caeca (16-23) and somewhat more pectoral fin rays (16-20); C. kamoharai by
somewhat more pectoral rays (16—20) and more gill rakers ({1—2]+[8—10] on inner side of first arch);
and C. multispinulosus by its much different pigmentation pattern on the body.

Caelorinchus charius new species
Fig. 9

Caelorinchus sp. E: Williams et al. 1996:148 (WA).

DIAGNOSIS. — Snout moderate in adults to long in young, sharply pointed, 38-50% of HL;
anterolateral margin fully supported by bone; orbit diameter slightly less than postorbital length of
head (in young about equal to or slightly greater than), 24-30% of HL, 1.3—2.1 times into snout length;
upper jaw extends posteriorly to below hind margin of pupil; subopercle produced into a short, narrow,
pointed flap. Short, naked black fossa of light organ immediately anterior to anus; anus removed from

IWAMOTO AND WILLIAMS: GRENADIERS OF WESTERN AUSTRALIA

TABLE 2. Selected measurements (%HL) compared for Caelorinchus argentatus and C. mayiae; specimens
of the former species separated by regions (P. I. = Philippines, WA = Northwest Shelf of Western Australia;

A&T = Arafura and Timor seas, QLD = Queensland).

Snout
Preoral
Internas.
Interorb.
Postorb.
Orb.-preop.
Barbel

Ht 1D

V. len.
Palen

Caelorinchus argentatus

* Range for four specimens.
** Range for six specimens.

Psi.

13
42-93
36-43
27-35
18=23
23-27
31-42
35—42

Sale)
39-50
28-38
33-36

WA

A&T
3
64-76
41-42
33-34
18-20
24-25
253i)
39-4]
11-13
40-45
27-33
34-37

C. mayiae

A&T
10
46-54
43—46
34-37
20-21
25-26
33-34
36-38
135
46-53
34-48
39-43

TABLE 3. Pectoral fin ray counts compared for Caelorinchus argentatus and C. mayiae, specimens of the
former species separated by regions (P. I. = Philippines, WA = Northwest Shelf of Western Australia, QLD

= Queensland, A&T = Arafura and Timor seas).

C. mayiae

C. argentatus
Pr
WA
QLD
A&T

Pectoral fin rays (excluding uppermost rudiment)

13 14 15 16 17 18 N x

l 16 a = — = 24 14.3
— — 12 9 2 l 24 Si?
= = l 9 9 = 19 16.4
= = 2 2 2 — 6 16.0
= _ 2 4 — — 6 17

anal fin by one or two scale rows. Underside of head covered with small, finely spinulated scales,
except along border of mouth, ventral and posterior margins of preopercle, and anteriorly on lower
jaw; nasal fossa with small scattered scales, most confined ventrally; body scales with short,
overlapping spinules arranged in 4—7 divergent rows, none especially broad-based or enlarged. No

prominent markings on fins.

SPECIMENS EXAMINED. — WA: HOLOTYPE: AMS I.31166-005 (79.5 mm HL, 265+ mm TL):
off Shark Bay; 610-612 m; SS1/91/31. PARATYPES: AMS I.31166-005 (7, 72.4-98.5 HL, 265+—370+
TL); same data as for holotype. CSIRO H1492-14 (51.7 HL, 170 TL)); NW Shelf; 420 m;
SO/05/88/70. CSIRO H1514-34 (5, 57.1-87 HL, 185-296 TL); NW Shelf; 582 m; SO5/88/190.
CSIRO H2554-10 (74.0 HL, 225+ TL); sw. of Point Cloates; 544 m; SS1/91/15. CSIRO H2557-05
(3, 63.1-103 HL, 180+—370 TL); w. of Cape Farquhar; 620 m; SS1/91/19. CSIRO H2573-16 (4,

134 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 3

TABLE 4. Comparison of selected measurements (in %HL) of Caelorinchus argentatus, C. maviae, C.
gaesorhynchus, and C. pardus.

C. argentatus C. mayiae C. gaesorhynchus C. pardus

Number of spec. 13 10 2 22

200 42-93 47-83 70-76 36-56
Snout 36-44 40-46 49-50 43-49
Preoral 27-37 33-37 44-47 37-43
Postorb. 31-42 34-37 27-28 27-34
Orb.-preop. 35-42 38-42 31-34 3390
Up.jaw 30-36 30-35 22-24 24-27
Barbel 6-14 11-16 7 9-16
Gill slit 13-18 14-18 10-11 10-18
Palen: 32-37 34-45 31— ? 34-46
V. len. 23-38 27-40 26-28 27-42

81.8—98 HL, 330-357+ TL); w. of Steep Point; 691 m;SS1/91/36. NTM S.12591-017 (60.0 HL, 168+
TL); NW Shelf; 17°35’S, 118°43’E; 445 m; coll. W. Houston; 3.XI.1985.

COUNTS AND MEASUREMENTS. — 1D. II,8—9; P. 117-119; GR-I (inner) 2+6, GR-II (outer/inner)
0+(6—7)/(1—2)+(6-7), 7—9 total; scales 1D. 5—7, midbase 1D. 3.5—4.5 (5), 2D. (3.5)4.5—7.5, lat.line
30-42; caeca 24-28 (3 spec.).

Total length 168+—370 mm; HL 51.7—103 mm. The following in percent of HL: snout 38—S0;
preoral 33—47; internasal 16-20; interorb. 19-25; orb. 24-30; suborb. 11—14; postorb. 25—34;
orb.-preop. 29-36; up.jaw 21—29; barbel 6—12; gill slit 9-12; pre-A. 121-169; V.-A. 27—52; isth.-A.
47-79; body depth 41-62; 1D.-2D. 18-39; ht. 1D. 32-48; len. base 1D. 14—23; len. P. 28-49; len. V.
25-48; post. nostril 6-11.

DESCRIPTION. — General features of fish seen in Figure 9. Snout of moderate length in adults,
decidedly longer (about twice orbit diameter) and more attenuated in young, sharply pointed and
tipped with a diamond-shaped terminal scute. Orbits slightly less than postorbital length in adults,
but slightly more in smallest specimens (51.7 mm and 63.1 mm HL). Mouth relatively small, rictus
extending to below midorbit, restricted posteriorly by lip folds; maxillary extending to below hind
margin of pupil; length upper jaw slightly less than postorbital length of head. Barbel short, about
equal to or less than length posterior nostril. Posteroventral corner of preopercle produced into a
moderate lobe; subopercle forming a short, slender, pointed ventral flap. Gill membranes broadly
attached to isthmus, restricting gill openings.

Premaxillary teeth in broad, short bands; mandibular teeth in narrow bands; all teeth small.

Height of first dorsal fin much less than postrostral length of head; second dorsal fin poorly
developed over most of length. No fin with notably elongated rays, but outer pelvic ray slightly
produced.

Underside of head mostly covered with small scales, except for broad naked margin along upper
jaws extending posteriorly to ventral and posterior margins of preopercle. Lower jaw sparsely scaled
posteriorly; naked anteriorly. Nasal fossa sparsely scaled, mostly along ventral portions in adults,
more broadly scaled in young. Scales on underside of snout small, relatively deciduous, and
nonimbricate; each armed with tiny, small, erect, conical to bladelike spinules arranged in a small
cluster or short rows. Spinules on body scales somewhat bladelike, overlapping along edges, and
aligned in sharp, thin, saw-toothed rows that increase in height posteriorly; each spinule relatively
erect, the most posterior on each row scarcely overlapping posterior margin of scale. In adults, midrow
of spinules on each scale with 6—7 spinules; those in lateral rows similar in height but usually
somewhat shorter, but all rows extend to margin of scale. Spinules on scales atop head and snout in

IWAMOTO AND WILLIAMS: GRENADIERS OF WESTERN AUSTRALIA 135

FIGURE 9. Caelorinchus charius n.sp. (a) Lateral view of holotype (AMS 1.3166-005, 370+ mm TL) (scales and fins partially
reconstructed) from off Shark Bay, WA, in 610—612 m; (a’) dorsal view of snout of holotype; (a") diagrammatic ventral view
of periproct and light organ; (b) lateral view of small paratype (CSIRO H1514-34, 270+ mm TL) from North West Shelf in
582 m. Scale bar represents 25 mm.

divergent rows, scales over interorbital with bladelike spinules; supraoccipital scute rather weakly
developed.

Light organ short, but naked fossa anterior to anus prominent, its length usually shorter than
greatest diameter of posterior nostril. Pyloric caeca long and slender, longest about 40 mm in 98.5
mm female from AMS 1.31166-005.

Color overall light brownish gray dorsally; faintly bluish over abdomen; ventrally over most of
body pale with ivory to silvery reflections. Underside of head whitish except anteriorly along snout
margin slightly dusky. Median nasal bone dark in darker specimens. Lips, gums, barbel, anterior
margin of mouth cavity cream colored, but otherwise dark; gullet and gill-cavity linings dark. Dorsal,
pectoral, and anal fins lightly dusky; pelvic fins dark dusky but outer ray distally white. In smaller
individuals from CSIRO H1514-34 and NTM S.12591-017, faint saddle bands noticeable on tail,
these bands less distinct in darker, fully scaled specimens.

SIZE. — To more than 37 cm TL.

DISTRIBUTION. — Known only from off WA, from west of Steep Point (26°S) to the North West
Shelf north of Port Hedland (18°S), and off Rowley Shoals (17°S). Depth range 420-691 m.

ETYMOLOGY. — From the Greek charieis, graceful.

COMPARISONS AND REMARKS. — This common new species shows considerable ontogenetic
change in snout proportions, with the snout notably longer and sharper in the young than in the adults.
These differences can be seen by comparison of the figures for young (Fig. 9b) and adult specimens
(Fig. 9a). The snout appears to actually diminish in absolute length after a certain size, such that a

136 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 3

small individual may have a longer snout than another much larger individual. This difference affects
most proportional values, as well as the lateral-line scale count.

Relationships of the new species seem to lie closest to C. smithi, but that species is distinctly
darker in overall color than C. charius; it has a sharper, thinner snout in lateral profile and somewhat
fewer pectoral fin rays (116-117 cf. 117-119). Another closely similar species is C. commutabilis Smith
and Radcliffe, 1912, but C. commutabilis has notably different spinulation on scales covering the
underside of the head; they are broadly bladelike in ridged rows, whereas in C. charius they are
needle-shaped (but usually 3-sided in cross section) and arranged in clusters.

Caelorinchus charius can be distinguished from most other members of the genus by the
combination of anterolateral margin completely supported by bone, characteristic light organ and
scale spinulation, mostly scaled underside of head, and sparsely scaled nasal fossa.

Caelorinchus gaesorhynchus new species
Fig. 10

DIAGNOSIS. — Snout 49-50% of HL, anterolateral margin of snout not completely supported by
bone; characteristic scales overlap onto anteroventral snout surface; orbit diameter 23-24% of HL;
upper jaw short, 22-24% HL, extends to below posterior one-third of orbit; subopercle lacking a
prolonged narrow flap; barbel short, about 7% of HL. Light organ extends from isthmus to anus.
Underside of head completely naked (except as noted above); nasal fossa naked; body scales thin,
spinules short, fine, in somewhat quincunx pattern. First dorsal fin dusky overall, but edges of base
blackish; no bold body markings (at least in adults); median nasal process blackish; mouth cream
colored, but lips and premaxillary processes blackish.

SPECIMENS EXAMINED. — WA: HOLOTYPE: CSIRO H2548-13 (76.3 mm HL, 245 mm TL); nw.
of NW Cape; 290 m; SS1/91/08. PARATYPE: CSIRO H2548-16 (70.6 HL, 235 TL); same data as for
holotype.

COUNTS AND MEASUREMENTS (holotype data first, followed by paratype data if different). —
1D. 11,9; P. 115/115, 115/114; GR-I (inner) 2+6; GR-II (outer/inner) 0+6/2+6; scales 1D. 5, 6, midbase
1D. 4, 2D. 5, lat.line 46, 43.

Total lengths 245 mm, 235 mm; HL 76.3 mm, 70.6 mm. The following in percent of HL: snout
49, 50; preoral 44, 47; internasal 18; interorb. 23, 21; orb. 24, 23; suborb. 11; postorb. 28, 27;
orb.-preop. 34, 31; up.jaw 24, 22; gill slit 10, 11; pre-A. 149, 139; V.-A. 41, 35; isth.-A. 69, 59; body
depth 41, 45; 1D.-2D. 19, 14; len. P. 31, —; len. V. 26, 28; len. base 1D. 17, 16; post. nostril 6, 8.

DESCRIPTION. — General features of fish seen in Figure 10. Snout long and attenuated, with a
sharply pointed tip. Orbits oblong, horizontal diameter about equal to postorbital length of head,
much shorter than snout length. Mouth relatively small, upper jaw less than orbit diameter, the rictus
extending posteriorly to below mid-orbit, the maxillary to below hind one-third of orbit. Suborbital
shelf broad, with a series of strongly adherent, somewhat platelike scales covered with short spinules
forming a sharp angular ridge separating upper and lower halves, scales above this ridge series thin
and deciduous in region under nostrils and orbit; posterior end of ridge extends almost to posteroven-
tral margin of preopercle bone. Preopercle somewhat prolonged posteriorly into a rather sharp lobe.
Subopercle bone closely adhered to small opercle, posteroventral corner forming short nib, not
prolonged into flap. Narrow naked cleftlike region along each side of median nasal ridge; nasal fossa
completely naked. Underside of head completely naked except for characteristic overlapping scales
along anterolateral margins. Barbel short, fine. Gill openings relatively wide, extending forward
ventrally almost to hind end of orbits, gill membranes narrowly attached to isthmus.

Teeth small, in moderately-wide tapered bands in both jaws, premaxillary band extending almost
to posterior end of rictus, mandibular band to or beyond end of rictus. No enlarged outer series of
teeth.

IWAMOTO AND WILLIAMS: GRENADIERS OF WESTERN AUSTRALIA 137

Keres

TILE MOSSE
io

FiGureE 10. Caelorinchus gaesorhynchus n.sp. Holotype (CSIRO H2548-13, 245 mm TL) from northwest of North West
Cape in 290 m. Fins and scales partially reconstructed. Scale bar represents 25 mm.

Fin size typical of other members of C. argentatus clade. Height of first dorsal fin less than
postrostral length of head; second dorsal fin poorly developed; pectoral and pelvic fins of about equal
length, outer pelvic ray slightly prolonged.

Light organ characteristic of Iwamoto’s (1990) Group IV, externally characterized by a long
median-ventral black streak extending from fossa just behind isthmus to immediately anterior to anus,
the streak expanded at each end and entirely scale covered.

Body overall pale. Gums cream colored, upper jaws anteriorly blackish, lower lips black; lower
jaw margins blackish; gill membranes dark. Opercle and subopercle dark. Gill cavity dark overall,
but pale over hyoid bones. Pectoral fins dusky overall; pelvic fins densely covered with large
melanophores; anal fin anteriorly dark, but dusky overall; medial surfaces of pectoral fin base black;
base of first dorsal outlined with thin black margin.

SIZE. — Attains at least 25 cm TL.

DISTRIBUTION. — So far known only from off Northwest Cape and Northwest Shelf at 290 m
depth.

ETYMOLOGY. — From the Greek, gaison, gaisos, a spear, javelin, and rhynchos, snout, in
reference to the notably long, sharp snout.

COMPARISONS AND REMARKS. — We initially confused specimens of C. gaesorhynchus with C.
argentatus and C. mayiae, but comparison with representatives of those species revealed notable
differences in snout length, mouth size, and pigmentation patterns. In C. argentatus the snout equals
about 40-44% of HL, in C. mayiae it equals 43-46%, but in C. gaesorhynchus it is 49-50% of HL.
Caelorinchus argentatus also has small black punctations and hairlike papillae on the ventral surfaces
of the head, and the spinules on scales posteriorly and ventrally on the trunk are triangular in shape
along the leading margin of the exposed field. The upper jaw length in C. argentatus equals about
30-35% of HL, in C. mayiae 31-33%, whereas it is 22-24% in C. gaesorhynchus (this character also
distinguishes the species from C. pardus). Caelorinchus kamoharai differs from the new species in
its longer upper jaw, and in having more rays in the first dorsal fin and more gill rakers. Caelorinchus
gaesorhynchus has pelvic fins heavily pigmented throughout, and the first dorsal fin is dusky. In
contrast, the other species of this group have pelvic fins with scattered punctations at the base, tip,
and sometimes the outer ray, but not in the middle; the black pigment on the first dorsal fin lies between
the second spinous and first segmented rays, with the proximal portion completely pale.

138 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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Caelorinchus goobala new species
Fig. b1

DIAGNOSIS. — Snout long, slender and sharply pointed in young, shorter, blunter in adults,
36-49% HL; anterolateral margin fully supported by bone; orbit 23-31% HL, 1.2 to almost 2.0 into
snout length. A short, poorly defined light organ in front of anus. Underside of head and nasal fossa
completely naked; body scales with conical spinules, aligned in about 5—11 divergent rows, the
individual spinules separate and not overlapping onto each other and without a transversely broadened
base; spinules on scales of interorbital space small, fine, conical, in divergent rows; a relatively broad,
translucent, unscaled area dorsally on each side of snout behind anterolateral margins. First dorsal fin
with distal tip black; mouth pale.

SPECIMENS EXAMINED. — WA: HOLOTYPE: NTM S.12631-004 (85.4 HL, 260+ TL); NW Shelf
off Rowley Shoals; 17°37’S, 118°40’E; 400 m; coll. W. Houston (WH85-21); 4.X1.1985. PARATYPES:
AMS 1.22808-042 (8, 63.7-94.0 mm HL, 183+—345 TL); NW Shelf, 220 km n. of Port Hedland; 420
m; SO2/82/17-18. CSIRO H1514-31 (6 of 11 spec., 59.7—77.2 HL, 206-290 TL); NW Shelf; 582 m;
SO5/88/190. AMS 1.23423-014 (63.5 HL, 211 TL); NW Shelf; 376 m; SO4/82/leg1. AMS 1.23425-
019 (54.8 HL, 200 TL) and CAS 200229 (61.4 HL, 200 TL); NW Shelf; 400 m; SO4/82/leg1. NTM
S.12588-023 (98.5 HL, 380+ TL); NW Shelf off Rowley Shoals; 17°22'S, 118°38’E; 403 m; coll. W.
Houston (WH85-15); 2.1.1985. NTM S.12614-023 (2, 57.4-82.9 HL, 180+—302 TL); NW Shelf off
Rowley Shoals; 17°34’S, 118°38’E; 410 m; coll. W. Houston (WH85-33); 7.X1.1985. NTM S.13115-
002 (57.5 HL, 150+ TL); n. of Cape Leveque; 14°07’S, 122°06’E; 423 m; coll. D. Evans; 24.1.1990.
NT: Timor Sea: ZMMGU uncat. (72.5 HL, 291 TL); 9°07.5’S, 131°14.9’E; 340 m; Akademik Berg;
26.V.1967. ZMMGU uncat. (75.2 HL, 291 TL); 9°00.0’S, 130°39.8’E; 445-520 m; Akademik Berg;
30.X.1968.

COUNTS AND MEASUREMENTS (21 spec.). — 1D. II,(7)8—9; P. 114417; GR-I (inner) (1—2)+6;
GR-II (outer/inner) 0+(5—6)/2+(5—6); scales 1D. 4.5—5.5, midbase 1D. 3.0-3.5, 2D. 4.0-5.5, lat.line
26-36 (38); caeca 30—44 (7 spec.).

Total length 180+—380+ mm; HL 57.4—-98.5 mm. The following in percent of HL: preoral 25-44;
internasal 16—19; interorb. 19-24; suborb. 10—15; postorb. 27-39; orb.-preop. 31-43; up.jaw 23-29;
barbel 7—14; gill slit 8-15; pre-A. 136-164 (171); V.-A. 31-57; isth.-A. 47-90; body depth 32-56;
1D.-2D. 15—39; ht. 1D. 38-53; len. P. 32-52; len. V. 33-44; post. nostril 7—10.

DESCRIPTION. — General features of adult seen in Figure 1 1a. Snout of moderate length in adults,
slightly longer than orbit diameter, but much longer and more attenuated in young (Fig. 11b) and
almost twice orbit in some juveniles; tip sharply pointed and armed with a slender terminal scute.
Orbit somewhat less than postorbital length of head. Mouth small, upper jaw length about equal to
orbit diameter; maxillary extends about to below hind margin of pupil; rictus to about front margin
of pupil. Barbel short, tapering to a fine tip; usually about equal to or less than suborbital width.
Posteroventral corner of subopercle forming a slender flap. Nasal fossa and underside of head
completely naked. A translucent naked area on each side of snout medial to anterolateral margins.
Gill membranes broadly attached to isthmus, moderately restricting gill openings.

Premaxillary teeth in broad short bands not extending to end of rictus; mandibular teeth in long
narrow band extending to end of rictus. No teeth enlarged.

Height of first dorsal fin less than postrostral length of head. Outer pelvic ray slightly prolonged.
Other fins without produced rays. Second dorsal fin poorly developed over most of length, the
interspace between first and second dorsal fins more than (1.2—1.7 times) length base of first dorsal
fin.

Body scales (Fig. 8f) covered with slightly divergent rows of slender spinules that are sharp and
narrow distally, but larger spinules often have bases broadened longitudinally in line with spinule
row. Spinule rows on larger body scales number about 5—8 in specimens less than about 70 mm HL

IWAMOTO AND WILLIAMS: GRENADIERS OF WESTERN AUSTRALIA 139

FiGureE 11. Caelorinchus goobala n.sp. (a and a’) An adult paratype (ZMMGU uncat., 291 mm TL) from the Timor Sea,
NT, in 445-520 m. (b and b’) A juvenile paratype (AMS I.12808-42, 183+ mm TL) from the North West Shelf, WA, in
420 m. Fins and scales in figure a partially reconstructed. Scale bar represents 25 mm.

and about 8-11 in specimens larger than about 70 mm HL. Supraoccipital scute weakly developed.
A slightly thickened, elongated scale at posterior end of occipital region above origin of lateral line.

Light organ short, slightly longer than posterior nostril, poorly defined and immediately preced-
ing anus, which is slightly removed by 2 or 3 scales from anal fin origin.

Color in alcohol light brownish; underside of head mostly creamish, extending onto gill
membranes; a light dusting of fine spots at tip of snout; mouth, lips and gums cream colored; inner
wall of gill chamber light grayish, outer walls pale. Abdomen darker, somewhat violet tinged, the
darker color extending posteriorly and ventrally onto tail. First dorsal fin dusky but tipped with black,
the blackish color more extensive in some smaller specimens, base usually pale; pectorals overall
dusky but with upper edge dark and base pale; pelvics creamy white with faint speckling in some

140 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 3

specimens, darker in others; anal fin creamy anteriorly but darker posteriorly along middle of fin, and
paler near end.

SIZE. — To about 35 cm TL.

DISTRIBUTION. — Known from off northern Australia in the Timor Sea and the North West Shelf,
in 300—S82 m.

ETYMOLOGY. — From the Australian Bardi language goobala, meaning star, in reference to the
spinules on scales of the median nasal ridge, which are, as in C. asteroides, arranged in rows radiating
from a central point; to be used as a noun in apposition.

COMPARISONS AND REMARKS. — Caelorinchus goobala is closely similar to C. asteroides
Okamura, 1970 and agrees rather well with the descriptions of that species provided by Okamura
(1970) and Yatou (in Okamura and Kitajima 1984). Two paratypes of C. asteroides (FAKU 21513,
19115) were kindly loaned to us by Dr. Okamura for our comparisons. Some distinctive squamation
features differentiate the two species, most notably the spinulation of body scales. In C. asteroides
the spinules are wide-bladed, keellike structures, recumbent and overlapping onto one another, and
forming sharp, narrow ridges, which are aligned in slightly divergent rows (see Okamura 1970,
text-fig. 82 A, A’). In C. goobala the spinules are slender, conical, individually separated from each
other and narrowly based in a transverse plane (but large spinules on or near midline of scales have
longitudinally broadened bases), and in rather widely divergent rows. Furthermore, Okamura
(1970:192) gave the ratio of the interspace between the dorsal fins and the length of the first dorsal
base as between 1.6 and 2.2; in the new species the ratio is 1.2—1.6. The pyloric caeca counts in seven
specimens were somewhat lower (30, 31, 36, 39, 41, 42, and 44) compared with the 40—S0 given by
Okamura (48 in two paratypes we examined). Okamura (1970:192) described the “lining of branchial

[sic, probably meaning buccal] cavity greyish, that of branchial cavity blackish brown ....” In our
specimens, the buccal cavity is pale, the branchial cavity somewhat dusky but paler along outer
margins.

According to Okamura (1970:192), C. asteroides is most closely related to C. radcliffei Gilbert
and Hubbs, 1920. The wide difference in the relative snout lengths of young and adults of that species
(as discussed by Gilbert and Hubbs 1920: 499) compares with that seen in C. goobala. In this regard,
we found that in our specimens, the snout of small specimens is proportionally elongated compared
with that of large adults. This distorts the values of the proportional measurements given above
because of our use of the head length for comparisons. Our largest specimen of 94 mm HL had head
proportions that were generally greater than those of smaller specimens, and body and fin proportions
that were shorter. Caelorinchus radcliffei differs from C. goobala chiefly in lacking a prolonged
subopercular flap, having darkly pigmented branchial and buccal linings, and having coarser,
bladelike scale spinules that broadly overlap each other.

Caelorinchus innotabilis McCulloch, 1907
Fig. 12

Coelorhynchus innotabilis McCulloch, 1907:348, 349, pl. 63, figs. 2, 2a (e. of Sydney; 1,463 m).
Caelorinchus innotabilis: Last et al., 1983:241, fig. 21.14 (TAS, VIC, NSW). Gomon et al., 1994:352, fig. 312
(NSW to SA). Williams et al. 1996:148 (WA distr.).

DIAGNOSIS. — Snout long, sharply pointed, anterolateral margin completely supported by bone.
Bony ridges of head generally narrow; bony scales of ridges with short, stout, posteriorly directed
spines. Light organ small, indistinct, closer to pelvic fin bases than to anal fin origin. Underside of
head mostly naked; scales moderately deciduous; scale pockets outlined by dark margins. Dark
coloration of trunk restricted to an area ventral to pectoral fin bases and extending from isthmus to
Just beyond anal fin origin.

IWAMOTO AND WILLIAMS: GRENADIERS OF WESTERN AUSTRALIA 141

FiGure 12. Caelorinchus innotabilis McCulloch. CSIRO H3007-10, from south of Albany, WA (35°26’S, 117°25’E), in
843 m.

SPECIMENS EXAMINED. — WA: CSIRO H3007-10 (4, 60.7-63.3 mm HL, 242-270 mm TL);
35°26'S, 117°25’'E; 843 m; Akebono Maru No. 3, sta. 7; coll. A. Williams; 22.XII.1989.

COUNTS AND MEASUREMENTS (based on 4 spec.). — Total GR-I (inner) 6—7, GR-II (outer/inner)
7—8/5—7; scales 1D. 7.5, midbase 1D. 6.5, 2D. 6.5—8.5.

Total length 242-270 mm; HL 60.7—-63.3 mm. The following in percent of HL: postrostral 57—59;
snout 41—44; preoral 39-42; internasal 20—22; interorb. 19-20; orb. 29-30; suborb. 13—14; postorb.
28-29; orb.-preop. 27—29; up.jaw 21; barbel 7-10; pre-A. 145—155; pre-vent 136—142; V.-A. 47-57;
isth.-A. 60—72; body depth 40—46; depth at A. 33-41; 1D.-2D. 13—14; ht. 1D. 43-49; len. P. 36-41;
len. V. 34-39; post. nostril 9-10; rictus len. 9.

DESCRIPTION. — Snout 2.3—2.6 in HL, a well-developed, elongate, sharp scute at tip. Dorsal
profile of snout shallowly concave. Mouth small, inferior.

Teeth small, in broad bands in both jaws.

Underside of head naked except for few scattered cycloid scales posterior to rictus. Body scales
semi-deciduous, mostly missing in our specimens; scale pockets conspicuous.

Light organ small, oval, indistinct, connected to periproct region by faint median extension.

Body color pinkish brown; region from isthmus to pelvic fin bases intensely blue. Dorsal,
pectoral, and anal fins dusky; pelvic fins dark; mouth and gill cavities black.

SIZE. — To at least 32 cm TL.

DISTRIBUTION. — Southern Australia and New Zealand, “common inclusion in trawl catches
from depths between 600 and 1000 metres” (Last et al. 1983:241).

COMPARISONS AND REMARKS. — In Western Australia, C. innotabilis was taken only south of
30°S. In southern Australian waters it most closely resembles C. kaivomaru Arai and Iwamoto, 1979.
The two species may be readily separated in the field by the distribution of the dark bluish black
abdominal tissue, which extends dorsally encircling the entire trunk in C. kaiyomaru, but only to the
level of the pectoral fins in C. innotabilis.

Caelorinchus lasti new species
Fig. 13, 14

Caelorinchus sp. G: Williams et al., 1996:149 (WA distr.).

DIAGNOSIS. — Snout moderately pointed, 1.4—1.6 times orbit diameter; anterolateral margin
incompletely supported by bone; upper jaw slightly less than 1/3 HL; orbit 1.5—1.8 into postorbital
length; subopercle produced into slender point ventrally, the tip protruding slightly beyond angle of
preopercle. Color overall dark chocolate brown. Nasal fossa with small scales on anteroventral corner
but otherwise naked. Spinules on body scales below dorsal interspace imbricate, broadly trihedral,
with wide lateral buttresses, in 3—5 saw-toothed, divergent rows. Anus immediately anterior to anal
fin; no external sign of light organ.

142 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 3

25mm

FiGurE 13. Caelorinchus lasti n.sp. Holotype (CSIRO H2615-02, 620+ mm TL) from west of Mandurah, WA, in 1,140 m. Fins and scales somewhat reconstructed. Scale bar

represents 25 mm.

143

IWAMOTO AND WILLIAMS: GRENADIERS OF WESTERN AUSTRALIA

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144 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 3

SPECIMENS EXAMINED. — WA: HOLOTYPE: CSIRO H2615-02 (163 mm HL, 620+ mm TL); w.
of Mandurah; 1,140 m; SS1/91/83. TAS: PARATYPE: CSIRO H1925-01 (142 HL, 455+ TL); w. of
Temma; 41°15'S, 143°58’E; 959—1,021 m; Petuna Endeavour; 18.11.1989.

COUNTS AND MEASUREMENTS. — ID. II,8; P. 117; GR-I (inner) 2+6, GR-II (outer/inner)
0+6/2+6; scales 1D. 6—7, midbase 1D. 4.5—5.5, 2D. 5.5—6.0, lat.line 32-34. The following in mm,
percent of HL in parentheses, holotype listed first: snout 52, 60 (37, 38); preoral 42, 52 (30, 32);
internasal 28, 30 (20, 18); interorb. 40, 40 (28, 25); orb. 38, 38 (26, 32); suborb. 21, 24 (14, 15);
postorb. 58, 67 (41, 41); orb.-preop. 60, 69 (42, 42); up.jaw 46, 52 (32, 32); barbel 14, 15 (10, 9); gill
slit 20, 23 (14, 14); pre-A. 210, 247 (148, 152); body depth 89, 102 (63, 63); 1D.-2D. 31, 35 (22, 21);
ht. 1D. 70, 59 (43, 42); 1D. base 24, 28 (17, 17); len. P. 64, 54 (39, 38); len. V. 61, 48 (37, 34); post.
nostril 12, 11 (9, 7).

DESCRIPTION. — See Figure 13 for general shape and proportions. Head large, more than 3.5
into TL; greatest width about equal to or more than postorbital length of head. Mouth moderately
large, rictus extends to midorbit, posterior edge of maxillary to below hind edge of orbit in holotype,
to below hind edge of pupil in paratype. Barbel short, less than 1/2 orbit diameter, thin, hairfine
distally. Body relatively deep, greatest depth below origin of first dorsal about equal to postrostral
length of head. Free neuromasts on head numerous, black (but not particularly noticeable because of
dark ground color).

Teeth in moderately broad bands in both jaws, about 3—6 teeth across widest part of premaxilla,
34 teeth wide across widest part of dentary; premaxillary band extending about 2/3 length of rictus,
mandibular band to end of rictus.

Paired fins relatively short, none extending to vertical of anal fin origin; outer pelvic ray slightly
prolonged but falling short of anus. Longest ray of first dorsal fin about equal to postorbital length of
head; second dorsal fin poorly developed except near terminal end. Anal fin well developed to end
of tail.

Scales (Fig. 14d) large, spinules on trunk scales in 3—5 divergent rows, the middle row slightly
to notably higher than lateral rows; 6—7 broadly trihedral spinules per row, the erect spinules
overlapping but free along most of their distal mesial edges; spinule rows complete in all trunk scales
of paratype, but in holotype, many scales with only 3 rows complete and | or 2 small spinules at
anterior edge of field in other rows. Scales on tail with more variable number of rows, the rows more
parallel and often incomplete to edge, and middle row usually notably higher. Most of scale spinules
on underside of head stoutly bladelike, aligned in high, narrow, crestlike or ridgelike rows, usually
]—2 rows per scale. Large broad scales of median nasal ridge row with short spinules in radiating
rows, posterior rows longest; in scales running mesial to supranarial ridges, spinule rows aligned to
some degree in wavy lines or striations. Occipital and cheek scales with widely divergent, sharp,
spinule rows. Scales atop head between ridges with 1—3 widely divergent spinule rows, the middle
row usually much longer than lateral rows.

Overall color dark brown; spinules on scales black; mouth, barbel, gill membranes, gill arches
and rakers dark brownish to blackish; lips, jaws blackish; gums pale in holotype, blackish in paratype;
all fins black.

SIZE. — To at least 62 cm TL.

DISTRIBUTION. — Known only from the holotype taken at about 33°S off WA, in 1,140 m, and
the single paratype taken at about 43°S off the west coast of TAS.

ETYMOLOGY. — Named after Peter Last of CSIRO in tribute to his contributions to Australian
ichthyology.

COMPARISONS AND REMARKS. — Caelorinchus lasti is distinctive among Iwamoto’s (1990)
Group I species of Cae/orinchus that are characterized by having underside of head scaled and snout
length less than twice orbit diameter (about 1.4~1.6 times orbit). It differs from other species of the
group in having the combination of incomplete support of anterolateral margins of snout, a dark

IWAMOTO AND WILLIAMS: GRENADIERS OF WESTERN AUSTRALIA 145

chocolate brown color, nasal fossa naked or only sparsely scaled anteroventrally, divergent rows of
spinules on scales of trunk and top of head, middle row of spinules on trunk scales higher than lateral
rows with rows complete to edge of scale in most trunk scales, and moderate-sized mouth (almost
one-third HL).

Of the Western Australian species, C. /asti appears most similar to C. mycterismus, with which
it has been confused, but it can be distinguished by its darker color, shorter snout (37-38% HL cf.
45-51%), longer upper jaw (32% HL cf. 15-26%), usually fewer scales on nasal fossa, and fewer,
more divergent and complete spinule rows on scales of trunk and tail. It can be distinguished from C.
acanthiger by the complete spinule rows on most scales of trunk (in C. acanthiger all body scales
with some rows incomplete, not reaching posterior edge of scale), and the very dark color of the head
and body.

Caelorinchus macrorhynchus Smith and Radcliffe, 1912
Bigal5

Coelorhynchus macrorhynchus Smith and Radcliffe, 1912:127, 128, pl. 29, fig. 1 (holotype USNM 72944, 56
cm long, Verde I. Passage, Luzon, Philippines; 180 fm [329 m]; A/batross sta. 5367; 9 paratypes, Philippines
and Indonesia, 432—759 m). Gilbert and Hubbs 1920:511, 512 (data on 5 paratypes).

Caelorinchus sp. D: Williams et al., 1996:148 (WA distr.).

DIAGNOSIS. — Snout about 2 times into HL, anterolateral margins completely supported by bone,
orbit diameter 4-5 into HL; upper jaw 4—S into HL, extends posteriorly to below hind 1/4 of orbit,
rictus to below midorbit; barbel small, less than half orbit diameter. Light organ very short, less than
greatest diameter of posterior nostril, externally a black streak extending forward from anus about
midway to pelvic bases. Nasal fossa covered with small scales except immediately anterior to nostrils;
underside of head uniformly scaled except for narrow margin above upper jaws; body scales with 5-8
divergent rows of broad-based, overlapping spinules; scales on top and underside of snout with few,
relatively broad-bladed, adnate spinules in one or few serrated, crestlike rows. Overall body color
swarthy to black; fins, mouth, and gill cavities blackish.

FiGure 15. Caelorinchus macrorhynchus Smith and Radcliffe. AMS 1.29600—01 (395+ mm TL) from off Sydney, NSW,
in 785-975 m. Fins and scales partially reconstructed. Scale bar represents 25 mm.

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SPECIMENS EXAMINED (9 spec.). — WA: CSIRO H2549-04 (112.5 mm HL, 365+ mm TL); w.
of NW Cape; 650 m; SS1/91/10. AMS I.22810-041 (112 HL, 382 TL); NW Shelf; 736 m; SO2/82/22-
24. QLD: AMS 1.20967-016 (138 HL, 407+ TL); e. of Hinchinbrook I.; 17°56'S, 147°14’E; 878 m;
27.11.1979. NSW: AMS 1I.29738-01 (85.5 HL, 280 TL); e. of Harrington; 485—925 m; K87-24-0S.
AMS 1.29825-01 (98.7 HL, 298 TL); off Terrigal; 741—768 m; K89-09-09. AMS 1I.24625-005 (440+
TL); off Broken Bay; 1,107 m; K84-06-06. AMS I.21722-07 (440+ TL); off Broken Bay; 822 m.
AMS 1I.29600-01 (127 HL, 395+ TL); off Sydney; 785-975 m; K87-24-01. AMS 1.27609-01 (55.6
HL, 162 TL); off Shoalhaven; K87-23-02.

COUNTS AND MEASUREMENTS. — ID. II,8—9; P. 116-119; GR-I (inner) (1—2)+(6—8) 8—10 total,
total GR-II (outer/inner) 6—8/9—10; scales 1D. 6-8, midbase 1D. 3.5—5.5 (usually 4.5), 2D. 5.5—6.5,
lat.line 38-55.

Total length 162-440+ mm; HL 55.6—138 mm. The following in percent of HL: snout 47—54;
preoral 3952; internasal 15—17; interorb. 17—20; orb. 21-25; suborb. 9—12; postorb. 24-31; orb.-
preop. 26-34; up.jaw 20-25; barbel 6-9; gill slit 9-13; pre-A. 131-143; V.-A. 24-36; isth.-A. 46-64;
body depth 33-48; 1D.-2D. 16-21; ht. 1D. 33-37; len. P. 29-33; len. V. 23-31; post. nostril 6—7.

DESCRIPTION. — General features of fish seen in Figure 15. Snout long, forming in lateral view
a slender pointed cone with shallow dorsal concavity, somewhat broader cone in dorsal view with
gently convex sides; diamond-shaped scute at tip. Orbit broadly oblate, 2.0—2.3 into snout. Suborbital
ridge stoutly developed along entire length, strongly spinulated along preopercular section. Preopercle
forming lobelike extension posteriorly; subopercle posteroventrally angular, but not forming long
slender flap. Chin barbel small and slender, length much shorter than least width of suborbital.

Premaxillary teeth in broad band extending almost 2/3 length of rictus; no enlarged teeth present.
Mandibular teeth in long narrow band, scarcely more than 3 or 4 teeth wide anteriorly, narrowing to
single row posteriorly; teeth extending to or beyond end of rictus.

Fins lacking any notably produced rays. First dorsal fin low, much less than postrostral length
of head; second dorsal fin low and inconspicuous over most of length, but somewhat higher along
posterior half. Outer pelvic ray slightly longer than mesial 6 rays.

Body scales (Fig. 14a) covered with ridgelike rows of broad, butressed spinules, each spinule at
a 45-60° angle from horizontal and closely adpressed to one another to form a saw-toothed ridge;
every row complete to margin of scale. Scales cover entire underside of head except narrow margins
above jaws, along ventral border of preopercle, and along lower jaw rami. Head scales highly variable
in size and extent of spinulation; those over occipital, opercle, and along posterior margin of
preopercle large, with many widely divergent spinule rows; most other head scales, aside from those
on ridges, small, with few spinule rows. Scales atop snout between ridges small, essentially nonim-
bricate, each with one or few short rows of broad-bladed spinules forming high, serrated crest.

Anus immediately anterior to anal fin origin.

Gums, gill arches, and rakers dark or blackish, peritoneum black. Most spinules of head and body
scales black.

SIZE. — To more than 44 cm TL.

DISTRIBUTION. — Known from the Philippines and Australia, including WA, NSW, and QLD;
depth range in Australia 485—1,107 m.

COMPARISONS AND REMARKS. — Caelorinchus macrorhynchus was originally described from
nine specimens taken in the Philippines in 329—750 m. Our Australian specimens agreed in most
details with the holotype (USNM 72944), but differed slightly in having a longer preoral (39-52%
HL cf. 37%), wider internasal (15—16% HL cf. 14%), larger orbit (21—25% HL cf. 19%), shorter
postorbital (24-31% HL cf. 34%), shorter upper jaw (20-25% HL cf. 29%), and shorter barbel (6—9%
HL cf. 11%). Furthermore, the snout-to-orbit ratios in the Australian specimens ranged 2.04—2.33, as
compared with 2.47 in the holotype of C. macrorhynchus and 2.2—2.3 in five smaller paratypes (data

IWAMOTO AND WILLIAMS: GRENADIERS OF WESTERN AUSTRALIA 147

from Gilbert and Hubbs 1920:512). The substantially larger size of the holotype (179 mm HL)
probably accounts for these proportional differences.

Caelorinchus smithi is similar to C. macrorhynchus, but there are numerous differences in
proportions, including a shorter snout, larger orbit, and broader interorbital and internasal widths.

The dark, swarthy, overall color of C. macrorhynchus, the scaled nasal fossa, the 5—8 divergent
rows of narrow-based spinules that are complete to the posterior margin of the scales, and the
distinctive spinulation on the dorsal and ventral snout surfaces are diagnostic for this species among
Iwamoto’s (1990) Group I Caelorinchus, all of which have a long, fully scaled snout, two or more
times orbit diameter, and a completely-supported, anterolateral snout margin.

Coelorhynchus macrorhynchus Weber, 1913 is a homonym, but Gilbert and Hubbs (1920:516)
questionably identified Weber’s species as representing their new species C. spinifer.

Caelorinchus maculatus Gilbert and Hubbs, 1920
Fig. 16

Coelorhynchus maculatus Gilbert and Hubbs, 1920:446-452, fig. 9 (Moluccas between Gillolo and Makyan Is.,
545 m; 10 paratypes from Moluccas and s. Luzon; 491-545 m). Arai in Gloerfelt-Tarp and Kailola 1984:83,
fig.

DIAGNOSIS. — Snout relatively short, acutely pointed in lateral view, broadly convex sides in
dorsal view, a small terminal scute; anterolateral margin completely supported by bone; orbits less
than snout length and postorbital length of head in specimens larger than about 170 mm, more than
in smaller specimens; mouth small, rictus extends to below midorbit, upper jaw to below hind
one-third of orbit; subopercle forms a short protruding tip. Ventral light organ (externally seen as a
black streak) extends forward from periproct to line connecting bases of outer pelvic fins or slightly
beyond, the slightly expanded anterior end forms shallow scale-covered fossa; anus removed from
anal fin origin by several scale rows. Underside of head completely naked; broad, naked translucent
areas dorsally behind leading edges of snout; nasal fossa naked; body scales (Fig. 8g) large, thin,
rather deciduous, exposed surfaces covered with small, sharp spinules aligned in 3—6 ridgelike rows,
scale pockets on dorsal surfaces darkly marked along edges; scales of head ridges relatively weak and
moderately spinulated (relatively coarser in juveniles). Ventral aspects of chest and abdomen dark,
bluish black, but sides silvery to whitish; a small to moderately large, vertically elongated dark blotch
above and immediately behind base of pectoral fin; numerous faint saddle marks posteriorly; most
fins light dusky, but elongated spinous dorsal ray blackish, and pelvic fin dusky to blackish with outer
ray paler; median nasal process dark. Anterior rays of second dorsal fin relatively well developed,
length of rays 34 into orbit diameter.

SPECIMENS EXAMINED. — WA: CSIRO H1701-02 (63.4 HL, 210+ TL), H1701-06 (62.6 HL,
248+ TL), and H1701-07(69.6 HL, 260+ TL); Rowley Shoals area; 16°59’S, 120°10’E; 411 m;
Striker; coll. S. Morris; 3.1V.1989. NTM S.12727-015 (52.2 HL, 215+ TL); sw. of Rowley Shoals;
17°52'S, 118°28'E; 410 m; coll. D. Evans; 9.11.1990. NTM S.12606-002 (36.1 HL, 125+ TL); off
Rowley Shoals; 18°00’S, 118°16'E; 430 m; coll. W. Houston (WH85-28); 6.X1.1985. NTM S.12609-
001 (33.6 HL, 132+ TL); off Rowley Shoals; 17°39’S, 118°40'E; 420 m; coll. W. Houston (WH85-
31). NTM S.12716-007 (56.1 HL, 247+ TL); nw. of Montebello I.; 20°15’S, 114°50’E; 500-600 m;
coll. D. Evans; 22.11.1990.

COUNTS AND MEASUREMENTS (6 WA spec.). — 1D. II,8—9; P. 114-416; total GR-I (inner) 7—8,
GR-II (outer/inner) 5—6/7—8; scales 1D. 6-7, midbase 1D. 3.54.5, lat.line 26-32.

Total length 125+—260+ mm; HL 33.6-69.6 mm. The following in percent of HL: snout 37-43;
preoral 28-35; internasal 19-22; interorb. 21—25; orb. 28-29; suborb. 13—16; postorb. 30-37;
orb.-preop. 35—43; up.jaw 26-30; barbel 6—12; gill slit 13-14; pre-A. 134-161; V.-A. 29-51; isth.-A.

148 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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57-80; body depth 47-62; 1D.-2D. 19-31; ht. 1D. 60-71; len. P. 43-56; len. V. 39-50; post. nostril
9-12.

SIZE. — To more than 26 cm TL.

DISTRIBUTION. — From southern Luzon (Philippines) in the South China Sea, to the Moluccas,
and off northwestern Australia, in 390-600 m.

COMPARISONS AND REMARKS. — Our six specimens from Western Australia agree well with
Gilbert and Hubbs’s (1920) original description of C. maculatus. The overall whitish color of larger
individuals is notable, particularly on the underside where the immaculate underside of head contrasts
sharply with the dark bluish purple to black of the chest and abdomen. The sides of the trunk are
generally whitish, with the dark color of the ventral surfaces not intruding onto these surfaces. These
color features appear to distinguish the species from other related species (see Gilbert and Hubbs
1920). The relatively long, spinous dorsal fin ray (about two-thirds head length in our specimens) is
somewhat shorter than that of other species in this group, where most have dorsal fin heights equal
to or greater than the head length. The anterior dermal window extends farther forward in our
specimens (to beyond the origin of the pelvic fins) than occurs in other species of the group, which
may cause confusion when using the key to species of Caelorinchus in lwamoto’s (1990) FAO
Catalogue. The species is normally assigned to Iwamoto’s Group II based on the position of the dermal
window at or behind the pelvic bases.

Caelorinchus matamua (McCann and McKnight, 1980)
Fig. 17

Mahia matamua McCann and McKnight, 1980:53—56 (New Zealand).

Coelorinchus matamua: Sazonov and Shcherbachev, 1982a:42—47, figs. 1, 2 (se. Atlantic, s. Indian Ocean). Last
et al. 1983:241—242, fig. 21.15 (Tasmania).

Caelorinchus matamua: Williams et al., 1996:148 (WA distr.).

DIAGNOSIS. — Snout short, blunt, anterolateral margin incompletely supported by bone; mouth
large, upper jaw usually longer than snout; orbit diameter equal to or longer than snout length. Light
organ small, indistinct. Underside of head including gular membranes scaled. Trunk completely
encircled by prominent dark broad band; anterior one-third to one-half of anal fin black, becoming
abruptly pale posteriorly.

SPECIMENS EXAMINED. — WA: CSIRO H3008-09 (93.5 mm HL, 380+ mm TL); sw of Albany;
35°25'S, 117°21'E; 842 m; Akebono Maru No. 3, shot 8; coll. A. Williams; 22.XI11.1989.

COUNTS AND MEASUREMENTS (based on | spec.). 1D. 11,9; P. 116; GR-I (inner) 12 total, GR-II
(outer/inner) 12 total/12 total; scales 1D. 10.0—10.5, midbase 1D. 9.5—10.5.

Total length 380+ mm; HL 93.5 mm. The following in percent of HL: postrostral 73; snout 30;
preoral 21; internasal 24; interorb. 25; orb. 37; suborb. 16; postorb. 37; orb.-preop. 44; up.jaw 29;
barbel 16; outer gill slit 22; pre-A. 143; pre-vent 140; V-A. 47; isth.-A. 74; body depth 76; depth at
A... 613 1D)-2D. 33; ht. ID466; leno PR S8:4en: V2.5); post. nostril, 7: rictus lena83:

DESCRIPTION. — Head large, snout about 2.9-3.3 in HL, with well-developed terminal scute;
dorsal profile gently rounded. Orbit very large, 34-37% HL, usually longer than snout. Subopercle
forming narrow, tablike flap at ventral end.

Teeth small, forming long tapered bands in both jaws, premaxilla bearing outer row of enlarged,
widely spaced teeth.

Dorsal fins well separated, second spinous ray of first dorsal fin not prolonged, its height about
50% HL; pectoral fin long, reaching to about 10th anal fin ray; anal fin well developed.

Scales of head mostly adherent; scales often missing on body. Suborbital ridge well developed;
large, ornate, spinulated scales of lower margin forming thorny ridges. Head ridges well defined,

IWAMOTO AND WILLIAMS: GRENADIERS OF WESTERN AUSTRALIA 149

Bie aietessactcaanaace:
i ecanueatengs
eS Ay Wire

FIGURE 16. Caelorinchus maculatus Gilbert and Hubbs. CSIRO H1701-02 (210+ mm TL) from Western Australia. Fins and
scales partially reconstructed. (a) lateral view; (b) dorsal view of head; (c) ventral view of abdomen to show extent of light
organ and dark pigmentation. Scale bar represents 25 mm.

FiGurRE 17. Caelorinchus matamua McCann and McKnight. CSIRO H3008-09, from south of Albany, WA (35°25’S,
117°21'E), in 842 m. Scale bar represents 10 cm.

150 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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scales enlarged but mostly lacking well-developed spinules; scales of anterior dorsal midline of head
largest, with rows of low ridges radiating from centers; scales in adjacent rows also large, but with
ridges forming a scalloped pattern.

Body color brown to grayish pink; head brown; distinct dark band around entire midsection of
body, dark blue over abdomen to level of at least 12th anal fin ray and to level of pectoral fin base,
less intense dorsally. All fins and branchiostegal membranes dark to black. Buccal cavity pale, lips
distinctly black.

SIZE. — To at least 65 cm TL.

DISTRIBUTION. — Southern Australia, southern Africa, and New Zealand, in depths of 450—
1085 m.

COMPARISONS AND REMARKS. — A distinctive species of Caelorinchus easily recognized in the
field by its bulky body shape, darker pigmentation of the anterior part of the anal fin, dark trunk region,
and the pockets of missing scales over the opercular region, which are reflective and clearly defined.
A rare species off the Australian west coast but common in other parts of its range. Nakamura’s
(1986:132, 133) record from Chilean waters is apparently in error, as the specimen illustrated is not
a member of this species.

Caelorinchus maurofasciatus McMillan and Paulin, 1993
Fig. 18

Caelorinchus maurofasciatus McMillan and Paulin, 1993:826, 827, fig. 5 (holotype NMNZ P.23003; Challenger
Plateau off New Zealand; 846-875 m; also n. and cent. NZ, 330-928 m; NSW, TAS, to GAB). Williams et
al. 1996:148 (distr.)

Caelorinchus sp. 1: McMillan in Gomon et al., 1996:347, fig. 307 (“false banded whiptail”; descr.; NSW to
GAB, TAS, and New Zealand, 400—800 m).

DIAGNOSIS. — Snout short, with broadly convex sides in dorsal view; terminal scute small,
broad, blunt; anterolateral margins incompletely supported by bone; orbits longer than snout length
and postorbital length of head; mouth small, maxillary extends to below about midorbit; subopercle
weakly produced ventrally into a short flap. Lens-shaped naked fossa of light organ developed anterior
to anus, about midway between anal and pelvic fins; anus immediately adjacent to origin of anal fin.
Underside of head completely naked; nasal fossa usually partially scaled; no broad naked areas
dorsally behind leading edge of snout; body scales covered with short, small spinules aligned in many
(to 11 or more depending on size of fish) parallel to slightly divergent rows; weak median-dorsal
ridge on nape from supraoccipital scute to origin of first dorsal, formed by modified scales with
somewhat higher and stouter median spinule rows. About 9—11 prominent saddle bands on body, the
third and fifth usually darker than others, the first beginning at about supraoccipital scute and
terminating below spinous rays of first dorsal fin, pale interspaces between bands angled posteroven-
trally from dorsal median line; first dorsal fin blackish distally but pale near base; anal fin often with
a dark distal stripe extending length of fin; oral and branchial cavities dark.

SPECIMENS EXAMINED. — WA: AMS1.31171-009 (45.3—51.6 HL, 197+210 TL); off Shark Bay;
500-508 m; SS1/91/37. NMV A9606 (6, 36.9-57.6 HL, 160-260 TL); 90 km sw. of Geraldton;
490-505 m; SS1/91/57. NMV A9660 (6, 38.9-63.3 HL, 168-282 TL); 50 km w. of Green Head;
480-490 m; SS1/91/65. CSIRO H2604-09 (2, 215-230 mm TL); sw. of Ledge Pt.; 512 m; SS1/91/70.
NMV A9639 (5, 49.0-53.3 HL, 210-225 TL); 50 km w. of Rottnest I.; 640-670 m; SS1/91/72. NSW:
AMS 1.24854-003 (33.8 mm HL, 150 mm TL); Jervis Bay off Wattamolla; 505 m; K84-14-01. AMS
1.23470-006 (5, 33.1-65.4 HL, 110-290 TL); off Sydney; 765 m; K83-17-01. NMV A2460 (32.2
HL, 147 TL); e. of Sydney; 33°04’S, 151°49’E; 421-494 m. TAS: NMV A2460 (46.6 HL, 197 TL);
70 km w. of Richardson; 41°13.4’S, 144°02.8’E; 528-600 m. NMV A4604 (58.6 HL, 252 TL); 21

151

IWAMOTO AND WILLIAMS: GRENADIERS OF WESTERN AUSTRALIA

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152 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 3

km e. of Maria I.; 42°41.7'S, 148°24.7’E; 428 m; FRV Soela. NMV A3831 (39.0 HL, 183 TL); 95
km ne. of Flinders I.; 39°02.9’S, 148°40.5’E; 636-660 m. NMV A8822 (69.7 HL, 302 TL); 80 km
ne. of Flinders I.; 39°28.5’S, 148°18.4'E; 430 m. CSIRO T1261 (4, 68.2—85.0 HL, 285+—320 TL);
South Tasman Rise; Kaiyo Maru; 22.X1.1975. VIC: NMV A768 (3, 69.8-83.5 HL, 285-310 TL);
off Portland; 38°50’S, 141°46’E; 549 m; Halycon; field no. MFG-7; 6.11.1980. NMV A8836 (60.0
HL, 267 TL); Tasman Sea, 40 km ese. of Gabo I.; 37°44.1'S, 150°16.7’E; 650 m; 3.VII.1984. AMS
1.18711-006 (2, 53.1-53.3 HL, 204+—206+ TL); “off southern Australia”; Dmitry Mendeleev;
28.11.1976.

COUNTS AND MEASUREMENTS. — ID. II,9—10; P. (116) 11720; GR-I (inner) (1—2)+(6—7), 7-9
total, GR-II (outer/inner) 0+(6—7)/(1—2)+(6—7); scales 1D. 4.5—6.0, midbase 1D. 3.5—4.5, 2D. 4.0-4.5
(5.5), lat.line 22-31; caeca 18—34.

Total length 110-320 mm; HL 32.0-83.5 mm. The following in percent of HL: snout 30-35;
preoral 28-34; internasal 21—25; interorb. 18—23; orb. 37-46; suborb. 15—17 (18); postorb. 28-33;
orb.-preop. 29-35; up.jaw 25-30; barbel 9-14 (16); gill slit 12-16; pre-A. 136-164; V.-A. 3449;
isth.-A. 62-84; body depth 59-73; 1D.-2D. (13) 15—25; ht. 1D. 57—80; len. P. 45-61; len. V. 48-59
(63); nostril 6—9; len. dermal window of light organ 6—13.

SIZE. — To at least 50 cm TL.

DISTRIBUTION. — From Shark Bay, WA, around Cape Leeuwin to VIC, TAS, and north to off
Sydney, NSW. Also off northern and central coasts of New Zealand. Depth range 330—930 m.

COMPARISONS AND REMARKS. — Caelorinchus maurofasciatus is strikingly similar to C. fas-
ciatus (Giinther, 1878) and was confused with that species before McMillan and Paulin (1993)
distinguished the two. They noted that the banding pattern is the same in the two species, but in C.
fasciatus the scales on the midline of the nape have a higher, stouter central spinule row, especially
the two or three nearest the dorsal fin; the body scales are generally more deciduous; the anal fin lacks
a distinct dark stripe (more uniformly dusky to black); the first dorsal fin lacks the pale proximal
portion that contrasts sharply with the blackish distal portion; and the pyloric caeca are generally
fewer. Our specimens showed considerable variation in the extent of dark pigmentation on the first
dorsal and anal fins; usually, we could not rely on those features alone to confidently identify our
material but had to couple them with other characters. The more-deciduous nature of body scales in
C. fasciatus was a useful clue, as representatives of that species in our collections were mostly denuded
of scales. Caelorinchus fasciatus has a more southerly distribution than C. maurofasciatus; off New
Zealand the latter species is abundant in northern waters, less so in central waters, and absent in
southern waters. Caelorinchus parvifasciatus and C. amydrozosterus are readily distinguished from
C. maurofasciatus by their fainter bands (which are oriented differently), usually paler body and fin
pigmentation, and broad naked areas on dorsal snout surfaces behind the leading edges.

Caelorinchus mayiae new species
Fig. 19b

DIAGNOSIS. — Snout 40-46% of HL, with a slight humplike rise in dorsal profile over nostril;
anterolateral margins not fully supported by bone; scales anterolaterally overlap onto ventral surfaces;
orbit diameter 22-25% of HL; upper jaw about 1/3 of HL, extends posteriorly to below hind margin
of orbit; barbel 11-16% of HL; subopercle without a slender projection. Light organ extends from
anus forward to just behind isthmus. Underside of head completely naked; nasal fossa naked. Spinous
dorsal ray blackish distally; median nasal bone blackish. Body markings prominent in most specimens
(see Description below). Head covering transparent; mouth lining white; lips and most of skin
covering premaxilla blackish; maxilla darkly pigmented; belly and pelvic bases almost entirely
without melanophores; vent and chest region densely pigmented.

IWAMOTO AND WILLIAMS: GRENADIERS OF WESTERN AUSTRALIA 153

FiGurE 19. (a) Caelorinchus pardus n.sp. Paratype (NTM S.12902-027, 232 mm TL) from northeast of Cape Don, NT, in
143 m. (b) Caelorinchus mayiae n.sp. Holotype, NTM S.13580—001 (170+ mm TL) from Arafura Sea, NT, in 179-205 m.
Fins and scales in (a) and (b) partially reconstructed. Scale bar represent 25 mm.

SPECIMENS EXAMINED. — NT (Arafura Sea): HOLOTYPE: NTM S.13577-001 (53.0 mm HL,
219 mm TL); 9°05.1'S, 133°39.7’E; 165-176 m; coll. R. Williams [RW92-66]; 20.X.1992. PAR-
ATYPES: NTM S.13580-001 (2, 53.0-71 HL, 171+—270 mm TL); 9°04.7’S, 133°04.7’'E; 179-205 m;
coll. R. Williams [RW92-68]; 20.X.1992. NTM S.13577-024 (6, 46.6-55.4 HL, 181-225 TL), QM
1.31225 (52.7 HL, 170+ TL), and CAS 200226 (2, 47-50 HL, 184-196 TL); same data as for holotype.
OTHER SPECIMENS (not paratypes): NTM S.13313-013 (3, 106-167 TL); Arafura Sea; 9°18'S,
133°12’E; 153 m;6.X1.1990.

COUNTS AND MEASUREMENTS (see Table 2-4 for additional counts and measurements). — 1D.
II,8-9; P. 114-416; total GR-I (inner) 8-10, GR-II (outer/inner) 6—8/8—10; scales 1D. 6-8, midbase
1D. 3.54.5 (5.5), 2D. 4.5—5.5, lat.line 37-42; caeca 10-11.

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Total length 153—275+ mm; HL 46.6—83.1 mm. The following in percent of HL: suborb. 12—14;
pre-A. 145-159; V.-A. 43-58; isth.-A. 73-91; body depth 45—54; 1D.-2D. 16-27; base 1D. 18-23;
ht. 1D. 40-53; len. P. 3445; len. V. 27-40; post. nostril 6-8.

DESCRIPTION. — A complete description of C. maviae would be redundant in that it is so similar
in most respects to C. argentatus and C. pardus, the latter of which has been described in detail.
Therefore, the following will emphasize only those features that differ between the species.

Body scales of C. mayiae (Fig. 8c) deciduous, most lost in all specimens examined. (This in
contrast to C. pardus, where all individuals still retain most of their scales.) Scales dorsally below
origin of first dorsal fin with spinules short, conical, little recurved, usually arranged in 4—6 somewhat
irregularly divergent rows to somewhat quincunx pattern. Posteriormost spinules in each row scarcely
or do not extend beyond edge of scale. Scales on chest with broader, flatter spinules, more triangular
in shape. Scales over snout notably thin, head covering here thin and completely transparent; scales
have few, short, weak spinules, even more reduced than in C. pardus, especially over median nasal
ridge. Most scales between occipital ridges with single median row of erect spinules. Scales atop head
sparse, spinulation thin, reduced (this contrasts with condition in C. pardus and C. argentatus, which
have dense covering of short erect spinules in rather random pattern).

Underside of snout with scattered tiny black, usually paired, punctations (neuromasts) similar to
those in C. argentatus and C. kamoharai but less dense; minute, black, hairlike papillae few or
virtually absent over ventral head surfaces.

Body markings already described in part in description of C. argentatus. Markings rather similar
to those of C. pardus, but generally not as spotty. Dorsally on nape and trunk, dark blotches fairly
extensive and continuous. Anterior end of dorsal stripe (below origin of second dorsal fin) forming
clublike or hockey-sticklike head directed anteroventrally and joining faint band that merges with
dark periproct region. Lateral aspects of abdomen heavily punctate, but ventral surfaces almost
completely devoid of pigmentation from periproct region to anterior end of pelvic fin bases, which
are white with scattered black punctations along base of fins. Chest sharply demarcated from pale
belly by dark pigmentation. Pelvic fins dark at base of rays, at tip, and along outer ray, but pale in
middle. First dorsal fin with second spinous ray and membrane following black on distal two-thirds
but completely pale proximally. Segmented rays with faint trace of broad darkish band across middle,
but pale proximally. Base of dorsal rays rather prominently marked with black. Anal fin somewhat
blackish on anterior few rays, but otherwise clear to light dusky posteriorly, with no blackened margin.
Lips dark, barbel dusky to pale, maxillary densely pigmented; lower jaw with prominent dark pigment
bands on inner surface of bones visible through transparent integument. Opercle and most of
subopercle black except pale along posterior margin; branchiostegal membrane dark dusky but pale
along posterior margin; gular membrane finely peppered posteriorly, with melanophores larger and
fewer toward anterior end, and very sparse around chin barbel. Fresh specimens probably silvery over
sides of head, ventral half of trunk, and midlaterally on tail.

SIZE. — To about 28 cm TL.

DISTRIBUTION. — Known only from the Arafura Sea, NT, in about 150—200 m.

ETYMOLOGY. — From the Australian Yindjibarndi language, mayi, meaning younger sister, in
reference to the putative sister-species relationship with C. argentatus.

COMPARISONS AND REMARKS. — Caelorinchus mayiae is so similar to C. argentatus that no
meristic or morphometric differences were found. The barbel appeared to be somewhat longer in C.
mayiae, but there is considerable overlap (11-16% HL cf. 6-14%). Spinulation of scales (see
Description) nonetheless distinguishes the two species. In C. argentatus the scales of the snout and
head are densely spinulated, in contrast to C. mayiae where they are thin and sparsely spinulated, and
the integument on the snout is not as thin and transparent in C. argentatus as in the new species. The
spinulation on body scales is somewhat denser in C. argentatus, and spinules along the anterior edge
of the scale field often have broadened bases aligned side by side on scales below the lateral line and

IWAMOTO AND WILLIAMS: GRENADIERS OF WESTERN AUSTRALIA 155

caudally on the dorsum beyond the trunk. Depth distribution shows some separation between the two
species, with C. mayiae taken in somewhat shallower depths (150-200 m cf. 260-420 m). In this
respect, C. maviae and C. pardus show more overlap, and the two species have been taken in the same
trawls.

Caelorinchus denticulatus Barnard, 1925 differs from C. mayiae in much the same way as the
latter differs from C. argentatus. In addition, the blotched pattern on the body is more broken in the
new species and there is no evidence of bands on the tail that we have noted in specimens of C.
denticulatus smaller than about 50 mm HL. Dentition appears to be slightly larger in C. mayiae, and
the scattered, paired punctations on the underside of the head in C. mayiae are not present in C.
denticulatus specimens.

Caelorinchus mayiae is also close to C. multispinulosus and C. pardus, but those species have
notably different pigmentation patterns, as well as a somewhat longer snout, proportionally shorter
upper jaw, postorbital, and orbit-to-preopercle lengths. Other measurements of C. pardus are
compared in Table 4. Caelorinchus pardus also has spinules on body scales in parallel rows.
Differences between C. mayiae and other members of this species complex are discussed in the
description of C. argentatus.

The surprising discovery of this new species, thought originally to be C. argentatus, serves to
illustrate the distinctiveness of the Australian grenadier fauna compared with those from Indonesia
and the Philippines. The small and subtle differences between C. mayiae and C. argentatus and
between C. mu/tispinulosus and C. pardus, however, suggest a fairly recent divergence between these
species pairs.

Caelorinchus mirus McCulloch, 1926
Fig. 20

Coelorhynchus (Paramacrurus) mirus McCulloch, 1926:178—180, pl. 46 (e. of Sydney off Montague I., NSW,
and e. edge of Bass Strait; 128-402 m).

Garichthys mirus: Whitley, 1968:38.

Coelorinchus mirus: Last et al., 1983:242, 243, fig. 21.16 (TAS, VIC, NSW, SA, WA).

Caelorinchus mirus: McMillan in Gomon et al., 1994:354, fig. 314 (deser.; distr. QLD to TAS, GAB). Williams
et al. 1996:148 (distr.).

DIAGNOSIS. — Snout pointed but short, much less than orbit diameter, tipped with a small spiny
scute; anterolateral margin not fully supported by bone; orbit large, more than postorbital length of
head; upper jaw extends to below posterior one-third of orbit, rictus to about below midorbits;
suborbital forming a short, angular flap at posteroventral corner. Black scaleless fossa of ventral light
organ large, extending forward from anus to line connecting origins of pelvic fins; anus removed from
anal fin origin by | or 2 scales at the most. Underside of snout and most of suborbital naked, but lower
jaw and underside of preopercle scaled posteriorly; nasal fossa and area immediately behind dorsal
leading edges of snout on both sides naked; body scales with minute spinules closely aligned in
numerous parallel rows (9-20 rows per scale in 37-57 mm HL specimens, larger individuals with
more rows on each scale). Color overall light brownish to grayish dorsally, and whitish ventrally,
with large, prominent, scattered peppering over gill membranes, posteriorly on underside of suborbital
and preopercle, and ventral surfaces of abdomen; pelvic fin whitish with prominent black blotch in
middle of fin; pectoral fin with small black spot at ventral corner; anal fin pale anteriorly, blackish
distal margins posteriorly.

SPECIMENS EXAMINED. — WA: ZMMGU uncat. (54.1 HL, 237+ TL); 30°30'S, 114°33.6’E;
320-340 m; Orlik; trawl 33; 8.X.1962. CSIRO C.2011 (1 spec.); Great Australian Bight; 1951.
CSIRO T1889-01(1 spec.); Great Australian Bight; III.1980 (no other data). NMVV A9658 (5, 195-285

156 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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FIGURE 20. Caelorinchus mirus McCulloch. (a) Adult specimen (ZMMGU P.17226, 237+ mm TL) from off Perth, WA, in
320-340 m; (a’) dorsal view of head to show naked regions behind leading edges of snout; (a”) ventral view of abdomen to
show light organ and anus. (b) Juvenile (AMS IB.4360-2, 75 mm TL) from east of Tuggerah Lakes, NSW, in 260 m. Fins and
scales in (a) partially reconstructed. Scale bar represents 25 mm.

TL); 50 km w. of Green Head; 30°00.1'S, 114°27.8’E; 380 m. NMV A10279 (3, 155-225 TL); 90
km sw. of Geraldton; 29°20.50’S, 113°56.80'E; 490-505 m. SA: CSIRO H1676-01 (1 spec.); 1980.
CSIRO T1917-01 (1 spec.) and CSIRO T1917-02 (1 spec.); Great Australian Bight; 33°21'S,
128°35’E; 322-325 m; Denebola; 18.1I1.1980. VIC: NMV A3044 (1 spec.); Bass Strait, s. of Point
Hicks; 38°14.89'S, 149°06.99’E; 212-240 m. NMV A8839 (6, 115-232 TL); Bass Strait, 40 km s.
of Point Hicks; 38°13.5'S, 149°12.8’E; 232 m. TAS: AMS JA.1379 (1 spec.), IA.1380 (1 spec.); e.
edge of Bass Strait; 39°S, 148°E. CSIRO CA3191 (1 spec.), CSIRO CA3192 (1 spec.), and CSIRO
CA3193 (1 spec.); Bass Strait, s. of Gabo I.; 276 m; SO1/72/12. NMV A3676 (290 TL); South Tasman
Rise; 47°32.00'S, 148°16.00’E. NMV A3715 (580 TL); South Tasman Rise, 47°29.00’S,
148°29.00'E. NSW: AMS I.15247 (holotype, 51.4 HL, 230+ TL); off Sydney; Endeavour. AMS
1.15246 (paratype, 57 HL, 273 TL), AMS 1.15037 (paratype, 52.5 HL, 245+ TL), AMS 1.15038
(paratype, 52.3 HL, 240+ TL), AMS 1.15248 (paratype, 37.6 HL, 191 TL); same data as for holotype.
AMS 1.15037 (1 spec.), 1.15038 (1 spec.), 115246 (1 spec.), 1.15247 (1 spec.), 1.15248 (1 spec.); e. of
Sydney; 33°S, 151°E; 1920. CSIRO CA77 (1 spec.); se. Australia; 400 m; 1976. CSIRO CA95 (1

IWAMOTO AND WILLIAMS: GRENADIERS OF WESTERN AUSTRALIA 157

spec.) and CSIRO CA96 (1 spec.); e. of Broken Bay; 33°33’S, 151°57'E; 265-305 m; F'RV Coura-
geous; 2.11.1977. CSIRO CA495 (1 spec.); e. of Wollongong; 34°32'S, 151°16’E; 198-202 m; FRV
Courageous; 17.X.1978. CSIRO C4799 (1 spec.); e. of Wollongong; 34°33’S, 151°15’E; 160-300
m; FRV Courageous; 8.V1.1976. AMS 1.19205-004 (4, 90-235 TL); e. of Broken Bay; K76-07-01.
AMS 1I.26221-002 (3, 230-250 TL); se. of Sugarloaf Point; K85-20-10. AMS 1.25932-006 (180 TL);
e. of Port Jackson; 1985. AMS IA.407 (1 spec.); off Montague I.; 36°S, 150°E. AMS IB.4336 (1
spec.), IB.4337 (1 spec.); IB.4338 (1 spec.), IB.4339 (1 spec.), IB.4340 (1 spec.); 35-40 miles ne. of
Broken Bay; 33°S, 151°E; 258 m; 1959. AMS IB.4360 (3, 72-80 TL), IB.4361 (1 spec.), IB.4362 (1
spec.); e. of Tuggerah Lakes; 33°S, 151°E; 260 m; 1959. AMS IB.4391 (1 spec.), IB.4392 (1 spec.);
e. of Newcastle; 32°S, 151°E; 293 m; 1959. AMS IB.8269 (5, 213—238 TL); 20 miles off Wollongong;
34°S, 151°E; 1968. NMV A2459 (3, 112-128 TL), e. of Sydney; 33°46’S, 151°49’E; 420 m. NMV
A6912 (14, 98-150 TL); 52 km ene. of Nowra; 34°43.55’S, 151°13.16’E; 450-345 m. QLD: AMS
1.15519-001 (2, 134-152 TL); off Brisbane; 26°17’S, 153°15’E; 188 m; 1968. AMS I.15528-003 (88
TL); off Brisbane; 26°31'S, 153°53’E; 373 m; 1968. AMS I.15550-004 (169 TL); off Brisbane,
267 31S153°S0'E; 329 m; 1968:

COUNTS AND MEASUREMENTS (6 spec.). — 1D. I,9—10; P. 117-118; GR-I (inner) (1—2)+6, GR-II
(outer/inner) 0+6/(2)+(6—7); scales 1D. 5.0—-S.5, midbase 1D. 3.5, 2D. 4.0—S.5, lat.line 23-26; caeca
about 40 in one spec.

Total length 191-273 mm; HL 37.6—57.0 mm. The following in percent of HL: snout 29-32;
preoral 29-31; internasal 22—25; interorb. 22—26; orb. 36-40; suborb. 16-17; postorb. 32-38;
orb.-preop. 37-40; up.jaw 28-33; barbel 15—20; gill slit 1S—19; pre-A. 154-167; V.-A. 4454; isth.-A.
74-96; body depth 69-86; 1D.-2D. 47-59; ht. 1D. 74-80; len. P. 51-58; len. V. 49-53; len. post.
nostril 4.9—9.0.

SIZE. — To about 30 cm TL.

DISTRIBUTION. — Widespread and common across southern Australia, north on west coast to
southwest of Geraldton, WA (about 30°S), and on east coast to off Brisbane, QLD (about 27°S), in
about 200-500 m. Off WA, most numerous between 300 and 400 m.

COMPARISONS AND REMARKS. — Caelorinchus mirus 1s a common species along the southern
part of the continent. It is easily distinguished from other similar species in the “C. fasciatus” group
by the large ventral fossa that extends forward of the vent region to between the pelvic fin bases, the
large black blotch on the pelvic fins, the small black spot ventrally at the base of the pectoral fin, the
scaled underside of the head posteriorly from above the angle of the jaws, and the absence of saddle
marks on the body of adults. Juveniles (Fig. 20b) as small as 72 mm TL (AMS IB.4360-2) were readily
identified by the pectoral and pelvic spots; saddle bands on the body are prominent in juveniles but
lost in adults.

|?|Caelorinchus mycterismus McMillan and Paulin, 1993
Fig. 21

Caelorinchus mycterismus McMillan and Paulin, 1993:827—-829, fig. 6 (New Zealand; 833—1,150 m).
Caelorinchus sp. F: Williams et al., 1997:148 (WA distr.).

DIAGNOSIS.—Snout slender, sharply pointed and upturned, 1.7—2.4 times orbit diameter, antero-
lateral margin incompletely supported by bone. Postorbital length of head 23-31% of HL, distance
orbit to preopercle angle 26-32%; orbit into postorbital 0.9—1.3 times. Light organ short, not
externally apparent; anus immediately before anal fin. Nasal fossa sparsely to (usually) extensively
covered with tiny scales; body scales with 4~9 parallel rows of spinules, the median row larger, lateral
rows often incomplete and not reaching posterior margin of scale.

158 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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SPECIMENS EXAMINED. — WA: AMS 1.22810-009 (4, 71.4-77.7 HL, 225—236+ TL); NW Shelf,
250 km nw. of Port Hedland; 736 m; SO2/82/22-24. CSIRO H2549-03 (74.5 HL, 228+ TL); w. of
NW Cape; 650m; SS1/91/10. NSW: AMS 1.29750-001 (90 HL, 278 TL); e. of Crowdy Bay;
1,024—1,052 m; K89-17-04.

COUNTS AND MEASUREMENTS (WA specimens only). — 1D. II,8—9; P. 116417; total GR-I
(inner) 7—9, total GR-II (outer/inner) 6—7/7—9; scales 1D. 5—7, midbase 1D. 4.5, 2D. 4.5—S.5, lat.line
3640; caeca 10 (2 spec.).

Total length 225—235 mm; HL 71.4-77.2 mm. The following in percent of HL: snout 45-48;
preoral 43-46; internasal 17—18; interorb. 20-23; orb. 24-26; suborb. 13—15; postorb. 29-31;
orb.-preop. 3 1—32; up.jaw 20—22; barbel 7-8; gill slit 10-11; pre-A. 132—145; V.-A. 30-38; isth.-A.
51-56; body depth 35—42; 1D.-2D. 13—17; ht. 1D. 32—36; len. P. 29-32; len. V. 31—33; post. nostril
6-7.
DESCRIPTION. — General features seen in Figure 21. Snout long, slender, slightly upturned with
concave dorsal profile, tipped with a large, sharp, diamond-shaped terminal scute. Mouth gape
restricted laterally by lip folds; rictus extends posteriorly to under midorbit. Barbel short, slightly
longer than vertical diameter of posterior nostril. Preopercle forming a lobe posteroventrally;
subopercle produced at ventral margin into a thin, narrow flap.

Premaxillary teeth in broad short band; mandibular teeth in narrow, more elongated band that
extends to posterior end of rictus.

First dorsal fin relatively low, height about one-third of head length, slightly more than postorbital
length; pectoral and pelvic fins similarly short, outer pelvic ray barely extends to vent; anal fin well
developed, second dorsal fin low to end of tail.

Scales completely cover head and snout; nasal fossa with tiny, finely spinulated scales scattered
only on ventral surfaces in most specimens, but over most of fossa in others. Underside of head
completely covered with small, mostly nonimbricate scales (Fig. 8h) armed with I—3 short rows of
broad, bladelike spinules, each spinule closely adjoined proximally to form a high, toothed crest, 2—5
spinules per row; spinules on scales dorsally atop snout and head similar, in |—4 rows. Supraoccipital
and occipital scutes not well differentiated; scales on head ridges thickened and armed with coarse,
strong spinules. Body scales (Fig. 81) with 4~7 parallel to slightly divergent rows of sharp spinules,
the median row largest and highest, formed of 4—9 broad-based spinules, the spinules rapidly increase
in height posteriorly to edge of scale; other scale rows shorter, those adjacent to median row often
not complete to edge of scale and only one or two spinules long. Scales posterior to origin of second
dorsal fin with more spinule rows than those on trunk.

Light organ short type, like those of Iwamoto’s (1990) Group I, without visible external fossa.

Color in alcohol overall tawny to grayish brown; first dorsal fin dark dusky to blackish; paired
fins and anal fin usually light dusky; abdomen with bluish tinge, but dark color not extending onto
chest area, region surrounding anus somewhat darker than adjacent areas; mouth and gill cavities
black, gums dusky to pale, upper and lower lips narrowly blackish, but usually pale laterally; barbel
pale; branchiostegal membrane blackish posteriorly and dorsally, but paler ventrally and over gular
membrane.

SIZE. — To about 50 cm TL.

DISTRIBUTION. — Known from WA only off the NW Shelf and NW Cape; also found off NSW;
abundant off New Zealand in 833—1,150 m (McMillan and Paulin 1993:829).

COMPARISONS AND REMARKS. — We tentatively identify the small WA specimens with C.
mycterismus, although scales on top and on the underside of the snout differ from one specimen we
have examined from New South Wales and others from New Zealand. These scales (Fig. 21c) had
broad spinules adjoined basally forming a short, high crest. Scales on the same areas of the head in
other specimens had spinules that were more slender and not closely adjoined. We could find no other

IWAMOTO AND WILLIAMS: GRENADIERS OF WESTERN AUSTRALIA 159

difference, however, to support recognizing them as distinct, but examination of additional specimens
covering a greater size range may change our view.

Caelorinchus mycterismus closely resembles C. kermadecus Jordan and Gilbert, 1904 and we
initially confounded the two species. However, Peter McMillan (pers. comm., Nov. 1997) advised us
as to characters that can be used to distinguish the two. According to McMillan and Paulin (1993),
C. mycterismus has a longer, more slender and upturned snout (45-55% HL in C. mycterismus cf.
41-47% in C. kermadecus), a lower first dorsal fin (28-36% HL cf. 36-43%), slightly shorter pectoral
fin (26-33% HL cf. 31-37%), shorter pelvic fin (21-31% HL cf. 29-39%), shorter upper jaw (15-26%
HL cf. 25-30%), shorter postorbital (25-30% cf. 34-36% [our data]), shorter orbit-preopercle
(25-30% cf. 34-36% [our data]). Additionally, the anal fin of C. mycterismus is pale posteriorly but
dark dusky to blackish in C. kermadecus, and the spinule rows on body scales are more numerous,
especially posteriorly on the tail (as many as 9 rows compared with about 7 as a maximum in C.
kermadecus), and the spinules are broader based, especially those lateral to the median row.

The apparently disjunct distribution of this species on opposite sides of the continent is peculiar
and unexpected. No collections of the species were made along the southern shores of Australia nor
off Queensland, despite fairly extensive sampling at appropriate depths.

Caelorinchus pardus new species
Fig. 19a

Coelorinchus sp. 4: Arai, in Gloerfelt-Tarp and Kailola, 1984:85, fig.

DIAGNOSIS. — Snout long, 43-49% of HL, anterolateral margin not completely supported by
bone; characteristic scales overlap anterolaterally onto ventral snout surface; orbit diameter 22-25%
of HL, 1.8—2.2 times into snout length; upper jaw 24-27% of HL; chin barbel short, about equal to
length of first gill slit; subopercle lacking a prolonged narrow flap. Light organ extends from behind
isthmus to front of anus. Underside of head naked; nasal fossa naked; body scales thin, spinules short,
fine, aligned in close parallel rows. Conspicuous body markings consisting of blotches, spots and
occasional ring on dorsum, becoming a narrow dorsal stripe to end of tail; belly between periproct
and pelvic fin bases lacking melanophores; maxillary ramus immaculate (rarely with 2-3 melano-
phores). First dorsal fin with membrane between second spinous ray and first segmented ray black;
anterior end of anal fin blackish distally. Pyloric caeca 8-11, usually 10 or 11; P. 114-115, rarely 116.

SPECIMENS EXAMINED. — NT (Arafura Sea): HOLOTYPE: NTM S.13551-001 (57.0 mm HL,
213 mm TL); 9°39.3'S, 133°04.4’E; 123 m; coll. R. Williams (RW92-38); 3.X.1992. PARATYPES:
NTM S.10774-001 (3, 170-185 TL); 9°32’S, 133°07’E; 126-130 m; FRV Soela sta. ASO2/81/112;
2.VII.1981. NTM S.12902-027 (12, 32-58 HL, 136-232 TL); ne. of Cape Don; 9°19'S, 132°42'E;
143 m; coll. R. Williams (RW90-3); 7.X1I.1990. NTM S.12903 (4, 180-200 TL); ne. of Cape Don;
9°19'S, 133°12'E; 152 m; coll. R. Williams (RW90-4); 7.X1.1990. NTM S.12923-003 (6, 36.444. 1
HL, 145-165 TL) and QM I.31224 (3, 136-143 TL); Arafura Sea (no other locality given); 112 m;
coll. R. Williams (RW90-24). NTM S.13313-013 (36.1 HL, 137 TL) and NTM S.13313-028 (12,
196-215 TL); 9°18’S, 133°12’E; 153 m; coll. R. Williams; 6.X1.1990. NTM S.13547-001 (11,
115-186 TL), AMS uncat. (2, 164-187 TL), BPBM 38460 (2, 161-172 TL), and BSKU uncat. (2,
116+-187 TL); 9°42.4’S, 133°23.2'E; 107m; coll. R. Williams (RW92-34); 3.X.1992. NTM
S.13548-001 (8, 128-195 TL) and CAS 200227 (4, 170-200 TL); 9°42.3’S, 133°17.2'E; 112-116 m;
coll. R. Williams (RW92-35); 3.X.1992. NTM S.13550 (5, 46.2-50.4 HL, 146+—199 TL); 9°30.7'S,
132°56.3'E; 136 m; coll. R. Williams (RW92-37). NTM S.13551-001 (5, 43.1-59.2 HL, 161+—192
TL); same) data as for holotype. NIM S.13577 (2, 51.3-52.8 HL, 200-200+ TL);9°05.1’S,

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133°39.7'E; 165-176 m; coll. R. Williams (RW92-65); 20.X.1992. NTM S.13578-001 (6, 178-208
TL); 8°55.3’S, 133°41.1'E; 179-187 m; coll. R. Williams (RW92-66); 20.X.1992.

COUNTS AND MEASUREMENTS (22 spec.). — 1D. I,7—10, usually 8—9; total GR-I (inner) 8—9
total, GR-II (outer/inner) 6—7/8—9; scales 1D. 4.5—7.5, midbase 1D. 3.5—5.5, 2D. 4.5—5.5, lat.line
36-43.

Total length 145-203 mm; HL 36.4-56.4 mm. The following in percent of HL: preoral 37-43;
internasal 1 8—23; interorb. 21—27; suborb. 1 1—14; postorb. 27—34; orb.-preop. 33-38%; barbel 9—16;
gill slit 10-18; pre-A. 137-159; V.-A. 35-58; isth.-A. 66-91; body depth 45—55; 1D.-2D. 14-27; ht.
1D. 40—S9; len. P. 3446; len. V. 27-42; post. nostril 5.1—9.0.

DESCRIPTION. — General features of fish seen in Figure 19a. Snout attenuated, armed with a
slender, sharp terminal scute. Orbit oblong, diameter about two or less in snout length, about equal
to interorbital width. Rictus extends posteriorly to below middle of orbit, maxillary to end of orbit.
Suborbital ridge forms a sharp edge separating upper and lower parts of head; ridge scales not
especially coarse. Preopercle margin sharply angular, forming a moderate lobe. Broad, translucent
naked areas dorsally behind leading edges of snout. Gill opening relatively wide, extending forward
to level of angle of lower jaws; gill membranes form a narrow free fold across isthmus.

Teeth small, fine, in a moderately wide band in upper jaw and extending to end of rictus; in a
narrow band in lower jaw extending slightly beyond end of rictus; none of teeth enlarged, although
in largest specimens, teeth overall more prominent.

Fins rather typical of most members of genus. First dorsal fin height less than postrostral length
of head; pectoral and pelvic fins of about equal length; outer pelvic ray barely reaching anal fin origin.

Body scales small (Fig. 8c), evenly covered with very small, conical, spinules. In smallest
specimens examined (about 140 mm TL), spinule rows in largest scales below interspace of dorsal
fins with 6-9 parallel rows, in larger specimens, number of rows ranged 8—12. Scales absent from
nasal fossa and suborbital shelf below anterior half of orbit. Scales on head with small erect conical
spinules, those on ridges not especially thickened nor coarsely spinulated; wide area behind antero-
lateral margin completely naked.

Light organ like those in Iwamoto’s (1990) Caelorinchus Group IV, with ventral streak expanded
at both ends.

Body markings prominent, characterized by series of blotches, vermiculations, and spots dorsally
that become long, narrow, dark streak below the second dorsal fin base extending to end of tail.
Usually, dusky color of postorbital region extends posteriorly as large, elongated blotch onto trunk
above pectoral fin, connected dorsally to dark surround of first dorsal fin. A large, elongated blotch
above midlateral septum about where lateral line levels off, beginning at or beyond level of depressed
tip of first dorsal and ending well beyond that tip. Ventral half of head and trunk probably silvery in
life. Chest region and broad area around vent darkly pigmented, but ventral surfaces of abdomen
relatively pale; light organ long extending as blackish streak from vent almost to isthmus. Top of head
and snout mostly pale; suborbital region and gill cover peppered with prominent melanophores.
Median nasal bone black. Opercle mostly black, but outer margins pale. Mouth cavity entirely pale,
lips darkly punctated. Lower jaw, gular membrane, and branchiostegal membrane generally dark;
underside of head otherwise mostly pale. Second spinous ray of first dorsal fin and membrane behind
black; base of fin with dark margin, especially noticeable at anterior edge. Pectoral fins pale to light
dusky. Pelvic fin whitish at base, pale with scattered punctations proximally and medially, blackish
along outer edge and distal tip. Anal fin with blackish distal margin at anterior end, becoming pale to
light dusky posteriorly, dark margined near end of tail in some.

SIZE. — A small species probably not exceeding 30 cm TL.

DISTRIBUTION. — Confined to the Arafura Sea in NT, in 107-187 m.

ETYMOLOGY. — From the Greek pardus, the pard, leopard, in reference to the leopardlike spots
characterizing the new species.

IWAMOTO AND WILLIAMS: GRENADIERS OF WESTERN AUSTRALIA 161

COMPARISONS AND REMARKS. — This species was recognized by Arai (in Gloerfelt-Tarp and
Kailola 1984) as an undescribed species, which he labeled as “sp. 4.” It appears to be a common
species at the edge of the continental shelf and upper slope, and it is one of the shallowest-living
grenadiers in Australian waters.

Caelorinchus pardus bears close resemblance to C. multispinulosus, a species known from off
Japan south to the Philippines and the Formosa Straits. The two are very similar in most characters,
but C. multispinulosus has generally more lateral line scales over a distance equal to predorsal length
(4146 cf. usually 33-41), smaller belly scales (usually 7-9 scale rows between inner bases of pelvic
fins, cf. 56 rows); spinules on body scales in quincunx pattern at all sizes, and more pyloric caeca
(11—14 cf. 8-11). Pigmentation pattern on the body also differs. In C. multispinulosus the formation
of three dark, lateral body stripes interspersed with three pale stripes is pronounced, although the dark
stripes are variously broken and the upper two are often interdigitated. In C. pardus only the uppermost
dark stripe is well developed, the second stripe being a series of elongate blotches or spots that end
rather abruptly in an elongated blotch slightly beyond a point below the tip of the depressed first dorsal
fin. Considerable variation is shown in these markings in both species, but on the whole, the
development of multiple stripes in C. multispinulosus is pronounced, whereas there is only a hint of
multiple stripes in the C. pardus. Certain proportional measurements show some differences between
the two species, but there is considerable overlap in ranges: for example, preoral length (somewhat
shorter at 32-36% HL cf. 33-43, but complete overlap in specimens larger than 60 mm HL), and
outer pelvic fin ray (slightly shorter, 30-33% cf. 27-42%).

Caelorinchus argentatus is also similar to C. pardus but has a somewhat shorter snout (40-44%
of HL), longer upper jaw (30-36%), spinules on body scales 1n quincunx pattern, and different body
markings. Caelorinchus gaesorhynchus lacks prominent body markings, lacks a black spinous dorsal
fin ray and black distal edge of pelvic fin, has a slightly shorter chin barbel (7% of HL) and upper
jaw (22-24%), and a longer preoral (44-47%). Caelorinchus mayiae, with which C. pardus has been
taken in the same trawl, has a distinctly different color pattern and spinule arrangement on scales.
Slight differences also exist in preoral and upper jaw lengths.

Caelorinchus smithi Gilbert and Hubbs, 1920
Fig. 22

Coelorhynchus smithi Gilbert and Hubbs, 1920:493—498, fig. 20 (holotype: USNM 78212; Philippines between
Gillolo and Makyan islands; 545 m; A/batross sta. 5621).

DIAGNOSIS. — Snout about 2.0—2.3 times into HL; anterolateral margin completely supported
by bone; orbit diameter 1.47—1.68 into snout length in adults, 1.7—2.0 in young; upper jaw 3.54.0
into HL in adults, 4.8—5.1 in juveniles, extends posteriorly to below middle one-third of orbit; barbel
short, less than 1/4 orbit diameter; subopercle produced into a short, angular flap directed posteriorly.
Light organ short, less than greatest diameter of posterior nostril. Nasal fossa mostly naked or covered
with small scales anteriorly and ventrally; underside of head uniformly scaled with small, nonimbri-
cate scales; body scales with 5—8 divergent rows of bladelike, imbricate spinules; scales atop snout
with spinules much like those on underside but in longer ridge-rows.

SPECIMENS EXAMINED (10 spec.). — Java: ZMMGU uncat. (89.6 mm HL, 300+ mm TL) and
CAS 202083 (94.6 HL, 338+ TL); offs. coast; 8°39’S, 111°55'E; 560-600 m; Prof: Mesiatzev cr. 7,
tr. 46; 27.1V.1979. NT: NTM S.12457-006 (95.7 HL, 312+ TL); Arafura Sea, se. of Tanimbar I.;
8°36'S, 132°00’'E; 525-540 m; coll. M. Sachse; 1.VII.1988. ZMMGU uncat. (4, 94-102 HL,
278+—305+ TL); Timor Sea; 13°22'S, 128°38.3'E; 525 m; Raduga tr. 1; 27. VIII.1966. QLD: AMS
1.32433-004 (2, 33.7-64.2 HL, 112—195+ TL); Papua Plateau, e. of Cape York; 10°29.8’S,
144°01.4’E; 596-603 m; R/V Franklin; 20.V1.1993. QM 1.23633 (88.6 HL, 270 TL); off Euston

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FIGURE 21. Caelorinchus mycterismus McMillan and Paulin. Juvenile (AMS 1.22810-009, 236+ mm TL) from 250 km
northwest of Port Hedland, WA, in 736 m. (a) lateral view, fins and scales partially reconstructed; (b) ventral view of head;
(c) scales from underside of head (not to scale). Scale bar represents 25 mm.

FIGURE 22. Caelorinchus smithi Gilbert and Smith. Adult (ZMMGU uncat., 300+ mm TL) from south coast of Java in
560-600 m. Scale bar represents 25 mm.

IWAMOTO AND WILLIAMS: GRENADIERS OF WESTERN AUSTRALIA 163

Reef; 16°40'S, 146°13’E; 600 m; 2.XII.1986. AMS 1.21796-012 (58.5 HL, 176+ TL); ne. of Danger
Pt.; 731 m; K78-23-08.

COUNTS AND MEASUREMENTS (10 spec.). — 1D. II,9; P. 116-117; total GR-I (inner) 7-8, total
GR-II (outer/ inner); 5—7/7—9; scale 1D. 5.5—6, midbase 1D. 3.5—4.5, 2D. 4.5—4, lat.line 29-37.

Total length 1 12—338+ mm; HL 89.6—102+ mm. The following in percent of HL: snout 41—S2;
preoral 35—49; internasal 17—19; interorbital 19-23; orb. 24-29; suborb. 10—14; postorb. 24-34;
orb.-preop. 28-38; up.jaw 20-29; barbel 6—9; gill slit 9-14; pre-A. 131—154; V.-A. 22-51; isth.-A.
46-80; depth 36—59; 1D.-2D. 16—36; ht. 1D. 33-40 (5 spec.); base 1D. 15—19; len. P. 30-44 (7 spec.);
len. V. 30—SO; post. nostril 6—10.

DESCRIPTION. — General features of fish seen in Figure 22. Snout moderately long, more so in
juveniles, sharply pointed in lateral view, dorsal profile slightly concave; viewed dorsally sides of
snout gently convex; a sharp, slender, diamond-shaped scute at tip; anterolateral margin sharp. Orbits
large, elliptic. Mouth relatively broad, slightly restricted at corners; chin barbel less than 10 percent
of head length. Suborbital ridge sharp, armed with stoutly modified scales; preopercle somewhat
lobelike at posteroventral corner; subopercle narrows to a short, pointed flap directed posteriorly to
posteroventrally. Scales cover underside of head except along narrow ventral margin of preopercle,
anteriorly on lower jaw, and over gill and gular membranes.

Premaxillary teeth in broad uniform band extending about three-fifths length of rictus; no
enlarged teeth. Mandibular teeth all uniformly small, in long, narrow band extending to posterior end
of rictus.

Dorsal and pectoral fins without produced rays; outer pelvic ray slightly produced, falling short
of, or scarcely extending beyond, anal origin. First dorsal fin low, height much less than postrostral
length, about equal to snout length. First dorsal and pelvic fins about on same vertical; pectoral fin
origin slightly in advance of that vertical. Anal fin origin far posterior, below interspace of first and
second dorsal fins, closer to origin of second. Base of first dorsal fin about 1.2—2.0 times into
interspace between dorsals.

Body scales large, covered with sharp, bladelike spinules arranged in divergent ridge-rows, the
rows complete to posterior margin of scale (Fig. 14c); posteriormost spinules in middle row often
with broad lateral buttresses, producing trihedral cross-section with each leg almost equal in width;
spinules in lateral rows often broadly triangular, and more or less aligned side by side, rather than
back to front. Head scales large on opercle and posteriorly and dorsally on preopercle, covered with
divergent ridge rows of low spinules; scales ventrally on head and above suborbital ridge and top of
snout generally small; an abrupt change in scale size between upper and lower portions of preopercle.
Scales atop snout and interorbital with 2—5 divergent rows of narrow, bladelike spinules of increasing
height; those on underside of head nonimbricate, with erect rows of bladelike, slightly recurved
spinules in one or two rows (Fig. 14b). Supraoccipital scute weakly developed.

Anus slightly removed by two or three scale rows from anal fin origin. A short, truncated light
organ anterior to anus, its length less than greatest diameter of posterior nostril; a narrow black
midventral streak anterior to periproct extending midway to pelvic bases, the light organ forming
posterior end. In NTM S.12457-006, 26 long (to about 30 mm), slender, pyloric caeca.

Overall color dark brown to swarthy in adults, more grayish in young, belly with bluish to violet
tinge; fins, mouth, gill cavities, and branchiostegal membrane black. Color of lips, external faces of
premaxillary and maxillary white to dusky; gums pale; barbel pale to dusky.

SIZE. — To more than 35 cm TL.

DISTRIBUTION. — Philippines, South China Sea, Borneo, Arafura Sea, Timor Sea (NT), off south
coast of Java, northern QLD, and northern NSW in 402-731 m.

COMPARISONS AND REMARKS. — The specimens from the Arafura Sea and Timor Sea represent
the only records of C. smithi from Western Australian waters, although the species is apparently rather
broadly distributed in tropical waters to the north. Our specimens agree well in all characters with the

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original description of C. smithi and with numerous CAS paratypes. However, a slight difference in
spinulation of the scales was noted. In NTM S.1247-006 the scale spinules are for the most part
narrowly bladelike, without a broadly truncated posterior margin, and the posteriormost spinule in
the middle row of body scales is not much different in size from the immediately anterior two or three,
and the spinule is not broadly trihedral. In contrast, in two large paratypes (CAS-SU 23993) the
posteriormost median spinule in most body scales is broadly trihedral, with a recurved, broad-bladed
shape in lateral view, and a broad-leafed shape in dorsal view. Also, the snout shape in dorsal view
is somewhat more convex in the Arafura Sea specimen. These are minor differences, however, and
probably attributable to individual variation.

Caelorinchus smithi is closely similar to C. charius but the two are distinguishable primarily by
the much paler overall color of C. charius and its slightly longer upper jaw, which ends below the
posterior one-third of the orbits, whereas in C. smithi the upper jaw ends below the middle one-third.
Caelorinchus charius also has a mostly pale mouth, whereas in C. smithi the mouth is wholly dark.
In comparable-sized specimens, the scale spinules on scales below the first dorsal fin of C. charius
are slightly more numerous in each row, more uniformly graduated in size posteriorly in each row,
and the spinules lack strong, broad lateral buttresses.

Caelorinchus smithi resembles C. macrorhynchus in many features, but differs in its much
shorter, somewhat broader snout (snout length 1.47—2.05 times orbit diameter cf. 2.04-2.47 in C.
macrorhynchus), somewhat fewer gill rakers (inner GR-I 7-8 cf. 8-10, inner GR-II 6~7 cf. 9-10),
fewer scale rows below the first dorsal (S—6 cf. 6-8), fewer lateral line scales over distance equal to
predorsal length (29-45 cf. 38-55), and a number of proportional measurements including internasal
width, interorbital width, and orbit diameter.

Caelorinchus thurla new species
Rigs 23

Coelorinchus sp. 5: Arai, in Gloerfelt-Tarp and Kailola, 1984:85, fig.

DIAGNOSIS. — Snout 37-45% of HL, anterolateral margin of snout completely supported by
bone; orbit diameter 26-29% of HL; upper jaw 22—26% of HL; subopercle prolonged into a short,
narrow flap; barbel short, 6—-10% of HL. Light organ a narrow black fossa extending forward from
periproct region to point at same level as insertions of pelvic fins; anus slightly removed from anal
fin origin. Underside of head completely naked; nasal fossa naked or with few tiny scales; body scales
covered with short fine spinules arranged in 6—1 | parallel rows. Prominent elongated black spot below
first dorsal fin, subtending 4-8 scale rows horizontally, 4 or 5 rows vertically, and removed from
pectoral fin base by about 4 scale rows; diagonal saddle mark below second dorsal fin.

SPECIMENS EXAMINED. — WA: HOLOTYPE: WAM P.21638-43 (49.0 mm HL, 210 mm TL);
NW Shelf; 13°45’04"S, 123°30'00"E; Umitaka-Maru. PARATYPES: WAM P.21638-43 (4, 29.5 HL,
105+—187 TL); same data as for holotype. WAM P.26270-016 (2, 47.4-48.6 HL, 196-204 TL);
15°30'S, 120°58’E; 280-320 m; Courageous sta. 0751; 28.V1.1978. NT: NTM S.13147-003 (45.1
HL, 173+ TL); Arafura Sea; 9°47'S, 130°26’E; 255 m; coll. D. Evans; 9.XII.1990. NTM S.13580-002
(4, 36.0-47.9 HL, 136+—206 TL); Arafura Sea; 9°04.7'S, 133°04.7’E; 179-205 m:; coll. R. Williams;
20.X.1992.

COUNTS AND MEASUREMENTS. — ID. II,8—9, 2D. 75—88; P. 114-416; total GR-I (inner) 6-8,
GR-II (outer/inner) S—6/6—7; scales 1D. 5—7, midbase 1 D. 4.5—5, 2D. 5.5—7, lat.line 30-37; 23 caeca
(1 spec.).

Total length 105+—210 mm; HL 29.5—49.0 mm. The following in percent of HL: preoral 32-40;
internasal 16—19; interorb. 18-21; suborb. 12-15; postorb. 29-35; orb.-preop. 33-38; gill slit 11—14;

IWAMOTO AND WILLIAMS: GRENADIERS OF WESTERN AUSTRALIA 165

FIGURE 23. Caelorinchus thurla n.sp. Holotype (WAM P21638-43, 210 mm TL); off North West Shelf, WA: (a) lateral
view, scales and fins partially reconstructed; (b) dorsal view of head; (c) paratype (187 mm TL), same locality as holotype,
ventral view of abdomen showing shape and position of light organ and anus. Fins and scales in (a) partially reconstructed.
Scale bar represents 25 mm.

pre-A. 140-161; V.-A. 29-41; isth.-A. 54-71; body depth 52—59; 1D.-2D. 24-34; ht. 1D. 58-70; len.
P. 43-49; len. V. 41-49; len. base 1D. 18—25; post. nostril 8—11.

DESCRIPTION. — General features of fish seen in Figure 23. Snout of moderate length for genus,
much longer than orbit diameter, which goes about |.5—1.6 times into length of snout; snout sharply
pointed in lateral profile, more broadly pointed in dorsal view, tipped with sharp spiny scute, which
in smaller specimens more elongated than in larger representatives. Orbits oblong, about equal to
postorbital length of head. Mouth small, upper jaw length less than orbit diameter, posterior margin
of maxilla extending to below middle of orbit or slightly beyond. Barbel fairly thick proximally,
rapidly tapering to fine tip. Suborbital ridge well developed, formed of series of stout spiny scales.
Posteroventral corner of preopercle somewhat angular, not especially produced. Gill membranes
rather broadly attached to isthmus, restricting opening of gill covers.

Premaxillary teeth in short broad band that falls well short of posterior end of rictus; mandibular
teeth in narrow tapering band that extends to beyond rictus. No enlarged outer row of teeth.

Except for outer pelvic ray, none of fins with produced rays. Second dorsal fin poorly developed
over most of length; other fins typical for genus.

Body scales (Fig. 14e) covered with needlelike spinules in parallel to slightly diverging rows.
Supraoccipital scute elongated and relatively well developed, with strong median row of enlarged
spinules. A weakly scutelike scale above origin of lateral line.

Light organ typical of Iwamoto’s (1990) Group II, externally visible as short, narrow black fossa
between pelvic fins (Fig. 23c). Anus removed from anal fin origin by three or four scale rows.

Body color brownish overall, somewhat creamy white on underside of head, lips, barbel, and
gums. Gular and branchiostegal membranes all whitish except dorsally on branchiostegal somewhat
blackish from underlying dark lining of gill cavity. Gill chamber linings dark, but paler along outer
ventral walls; gill arches and rakers dusky, filaments paler. Mouth and gullet mostly dark but paler

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near mouth opening; gums and lips cream colored. Abdomen with bluish tinge; dark area ventrally
extends onto all of chest, but not much dorsally onto trunk. A prominent occellated spot overlying
anterior end of lateral line. Fainter, elongated saddle mark, shaped more like a diagonal band in some
specimens, below second dorsal and separated from pectoral spot by distance about equal to snout
length or length of snout and orbit. First dorsal fin dusky but darker distally; pectoral fin lightly dusky;
pelvic fins blackish, densely covered with large melanophores; anal fin with larger melanophores
anteriorly and distally, giving a dark appearance, but fin pales posteriorly to dusky, becoming
completely pale near distal end.

SIZE. — To about 21 cm TL.

DISTRIBUTION. — So far known only from the Arafura Sea in Northern Territory and the North
West Shelf of Western Australia, in 179-320 m.

ETYMOLOGY. — From the ‘Australian Yindjibarndi word thurla, meaning eye, in reference to the
characteristic shoulder spot and to the name of its sister species, C. argus. It is treated as a noun in
apposition.

COMPARISONS AND REMARKS. — Caelorinchus thurla is similar to C. argus in most features,
but differs in having a shallower and more posteriorly placed pectoral spot. In C. argus the spot is
distinctly rounded, about 6 rows deep, and closely placed near the pectoral fin, being removed by
only 2 or 3 scale rows from the pectoral fin base, with the anterior margin below the origin of the first
dorsal fin. In C. thurla the spot is normally oblong, only 4 or 5 rows deep and removed from the
pectoral fin base by 4 or 5 scale rows. Furthermore, the scales in C. thurla are somewhat smaller,
about 4.5—5.0 rows below the midbase of the first dorsal (compared with 4.04.5 in C. argus) and
5.5—7.0 rows below the origin of the second dorsal fin (compared with 4.5—6.0). The preoral length
is also somewhat shorter (34-39% cf. 37-45%), the internasal somewhat wider (16-19% cf. 14-17%),
and the suborbital somewhat broader (13—15% cf. 11-13%).

Caelorinchus trachycarus lwamoto, McMillan, and Shcherbachey, 1999
Fig. 24

Caelorinchus trachycarus lwamoto, McMillan, and Shcherbachev, 1999:49—54.

DIAGNOSIS.—A species of Caelorinchus in lwamoto’s (1990) Group I, with underside of head
mostly scaled; anterolateral snout margin incompletely supported by bone; nasal fossa and triangular
area bordered by fossa, orbit, and suborbital ridges naked; orbit 1.4—1.9 into snout length; head ridges
especially spiny; broad margins of naked skin on preopercle and lower jaw; color dark swarthy with
violet to purple tinge, fin rays black; body scales with usually 7-10 rows of spinules, the middle row
highest; neuromasts on head prominently black; anus immediately before anal fin, without external
trace of light organ; pyloric caeca 7—9.

SPECIMENS EXAMINED (10 WA specimens from type series) WA: NMV A6189 (2 paratypes,
93.4-121 mm HL), NMV A6190 (1 paratype, 77.5 HL), sw. of Esperance, 1,015—1,030 m. AMS
1.18712-015 (1 paratype, 99 HL), ZMMGU uncat. (6 paratypes, 96—-117.7 HL), 1,080—1,100 m.

COUNTS AND MEASUREMENTS (from original description). — 1D. II,7—9; P. 115-118 (120, 121);
scales 1D. 4.5—7 (9), midbase 1D. 3.5—6, 2D. 4.5—6.5, lat.line 28-37 (42); caeca 7-9.

The following in percent of HL: snout length 39-49; preoral 36—48; internasal 1 8—23; interorb.
20-27; orb. 22-30; suborb. 14-17; postorb. 26-34; orb.-preop. 26-37; up.jaw 17—24; barbel 4-9;
pre-A. 141—171; body depth 39-58; 1D.-2D. 18-33; ht. 1D. 37—59; len. P. 32-41; len. V. 28-46.

SIZE—To about 51 cm TL.

DISTRIBUTION.—Southern Australia to New Zealand, in 1,015—1,234 m.

COMPARISONS AND REMARKS.—This recently described species is most similar to C. acanthiger
but the two can be readily separated because C. trachycarus has relatively much coarser, more spiny

IWAMOTO AND WILLIAMS: GRENADIERS OF WESTERN AUSTRALIA 167

scales of the head, more extensive naked areas, especially on the preopercle and anteriorly on the
suborbital shelf behind the nasal fossa, darker overall color, especially the black fins, and violet to
purplish tinge on naked areas. Scales are somewhat smaller in C. acanthiger, although there is overlap
in scale row counts (scale rows below 1D. usually 5.5—7 in C. trachycarus cf. 7-10 in C. acanthiger,
below midbase 1D. 3.5—6 cf. 5.5—7, and lateral line scales 28—37 cf. 3449).

Cetonurichthys Sazonov and Shcherbachey, 1982

The generic diagnosis provided by Sazonov and Shcherbachev (1982) for Cetonurichthys is very
similar to that for Cetonurus, as given in their later paper (Sazonov and Shcherbachev 1985). The
following additional characters may be used to distinguish Cetonurichthys: pelvic fin distinctly
posterior to line through first dorsal and pectoral fin bases; lateral line interdigitated, a series of
elongate pores covered by a transparent membrane, each about 2.0—2.5 scales in length; pores
separated by about 2.0—2.5 scales anteriorly, about 7 scales posteriorly; one pair of neuromasts located
medially on border of each pore; margin of pore pigmented, about 6—7 rows of teeth at symphysis of
upper and lower jaws; tooth patches broad, together forming crescent-shaped band in each jaw; lips
fleshy, well developed at angles of mouth, with conspicuous development of branched papillae; mouth
noticeably protrusible.

Only the single species known.

Cetonurichthys subinflatus Sazonov and Shcherbachev, 1982
Fig, 25

Cetonurichthys subinflatus Sazonov and Shcherbachev, 1982b:8—11 (holotype ZMMGU P2996, off nw. coast of
Australia; 16°55’S, 114°53’E, 1,600—1,700 m). Williams et al. 1996:149 (WA). Grandperrin et al. 1997:117
(New Caledonia off Norfolk Ridge).

DIAGNOSIS. — Head broad, shallow, interorbital 31-34% of HL; snout 36-42% HL, with paired
tubercular scales at tip; suborbital 17-20% HL, ridge lacking stout scutelike scales; jaws short, upper
Jaw 24-28% HL, inferior; tooth rows short, crescent-shaped, outer premaxillary series enlarged.
Cephalic sensory canal moderately dilated, lacking open pores; lateral line canal with modified scales,
interrupted in places. Trunk scales small (11—13 rows between origin of first dorsal fin and lateral
line), bearing erect spinules in no particular arrangement; scales along dorsal fins larger than
surrounding ones; head completely scaled, including patches on gular and branchiostegal membranes.
Spinous second ray of first dorsal fin with small serrations; pelvic fins with 8—9 rays, outer ray
elongated. Anus midway between bases of anal and pelvic fins; periproct narrow; no dermal window
of light organ. Eleven trunk vertebrae. (After Sazonov and Shcherbachev 1982:8).

SPECIMENS EXAMINED. — WA: CSIRO H2551-16 (64.5 mm HL, 363+ mm TL) and CSIRO
H2551-13 (79.0 HL, 395+ TL)); off NW Cape; 1,460—1,500 m; SS1/91/12.

COUNTS AND MEASUREMENTS (based on 2 spec.). — 1D. II,9—10; P. 115-117; V. 8; total GR-I
(inner) 8-9, GR-II (outer/inner) 8 /7—8; scales 1D. 11.5—13.0, midbase 1D. 8.5—12.0, 2D. 10.0—11.5.

Total length 363—395+ mm; HL 64.5—79.0. The following in percent of HL: postrostral 62-63;
snout 39-40; preoral 29-31; internasal 27—30; interorb. 30; orb. 23; suborb. 18—19; postorb. 39-41;
orb.-preop. 35-37; up.jaw 26-27; barbel 8; outer gill slit 10; pre-A. 154-160; body depth 76-83;
pre-vent 141—151; isth.-A. 67-68; 1D.-2D. 33-37; ht. 1D. 91-94; len. V. 45—56; post. nostril 8-10.

SIZE. — To at least 40 cm TL.

DISTRIBUTION. — Known from only five specimens, including our two, from two locations north
and east of the Exmouth Plateau, North West Cape, WA. Depth range 1,316—1,700 m.

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FiGurE 24. Caelorinchus trachycarus lwamoto, McMillan, and Shcherbachev, 1999. Paratype (100 mm HL, 350+ mm TL),
AMS 1.18712-015, from off southern Australia. Fins and scales partially reconstructed. Scale bar equals 25 mm.

FIGURE 25. Cetonurichthys subinflatus Sazonov and Shcherbachev. CSIRO H2551-16 (395+ mm TL) from off North West
Cape, WA, in 1,460—1,500 m. Drawn by Georgina L. Davis.

COMPARISONS AND REMARKS. — Our counts and proportional measurements slightly extended
the ranges given by Sazonov and Shcherbachev (1982). This is not surprising considering that our
specimens are the largest two of the five known. One noteworthy difference was the greater number
of tooth rows in our specimens: six or seven rows at the symphysis of both jaws, compared to three
reported in the type specimens.

In our specimens the pelvic fin origin was distinctly posterior to a vertical line through the
posterior margin of the pectoral fin base. Sazonov and Shcherbachev (1982) described the position
as “on the line or slightly posterior to it.” In the larger of our specimens, a well-developed elongate
scale patch on the midline of the gular membrane extended posteriorly to a vertical through the

IWAMOTO AND WILLIAMS: GRENADIERS OF WESTERN AUSTRALIA 169

posterior margin of the orbit; on each side of the midline, a large oval scale patch (about 0.5 of orbit)
was present on the anterior portion of the lower branchiostegal membranes; scattered scales linked
these patches with the gular patch.

Cetonurus Gunther, 1887

See Sazonov and Shcherbachev (1985) for diagnostic characters and a comprehensive discussion
of this genus. Two species recognized, only one of which was found in our study material.

Cetonurus globiceps (Vaillant, 1884)
Fig. 26

Macrurus globiceps Vaillant in Filhol, 1884:183, fig. 2 (name and figure; name attributed to Vaillant).
Cetonurus globiceps: McMillan in Gomon et al. 1994:345, fig. 306 (NSW, VIC, GAB, 950—1,200 m). Williams
et al. 1996:149 (WA)

DIAGNOSIS. — Orbit 24-28% of HL; scale rows below origins of first and second dorsal fins,
respectively, 16—18 and 10-19 (usually 14—15); 34 rows of closely spaced teeth on premaxilla.

SPECIMENS EXAMINED. — WA: CSIRO H2581-04 (77.0 mm HL, 366+ mm TL, CSIRO H2581-
03 (84.0 HL, 375+ TL), CSIRO H2581-05 (79.5 HL, 364+ TL), and CSIRO H2581-08 (76.0 HL,
358+ TL); ssw. of Shark Bay; 996—1,009 m; SS1/91/44. CSIRO H2593-02 (84.0 HL, 275+ TL); w.
of Leander Pt.; 942-970 m; SS1/91/58. CSIRO H3017-05 (68.1 HL, 298+ TL); off Cape Leeuwin;
34°10’S, 114°16’E; 1,030 m; Akebono Maru No. 3, shot 17; coll. A. Williams; 24.XII.1989. TAS:
CSIRO H1579-49 (12, 44.0-77.0 HL, 170+—310+ TL), w. coast; 792-830 m; SO3/86/37.

Also examined for comparison: holotype of Cefonurus crassiceps, BMNH 87.12.7.108 (66.3
HL, 272 TL); n. of Kermadec Is.; 650 fathoms [1189 m]; Challenger sta. 170; 14.V1.1874.

COUNTS AND MEASUREMENTS (based on 10 spec.). — ID. II,9; P. 115-117; total GR-I (inner)
11—13, GR-II (outer/inner) 10—12/10—12; scales 1D. 18-19, midbase 1D. 15—17, 2D. 14-16.

Total length 275+—375+ mm; HL 68.8—84.0 mm. The following in percent of HL: snout 36-42;
preoral 32-41; internasal 40—44; interorb. 41-45; suborb. 24-27; postorb. 43—S0; orb.-preop. 42-53;
up.jaw 28-33; pre-A. 129-142; pre-vent 128—133; V.-A. 46-52; isth.-A. 48-58; body depth 86-98;
1D.-2D. 28-39; ht. 1D. 47-53; len. P. 41; post. nostril 4-6.

SIZE. — To at least 38 cm TL.

DISTRIBUTION. — Circumglobal, recorded from the North and South Atlantic oceans, off the
Indo-Malayan Archipelago, and Japan; western, central, and eastern Indian Ocean; southern Australia
and New Zealand (Sazonov and Shcherbachev 1985). Widely distributed down the western coast of
Australia in depth from 740 m to 1,500 m.

COMPARISONS AND REMARKS. — Some confusion exists regarding the distribution of Cetonurus
species in Australian waters. Cetonurus globiceps was reported from southern Australia by Sazonov
and Shcherbachev (1985), although only C. crassiceps is recorded by Paxton et al. (1989). More
recently, Gomon, Glover, and Kuiter(1994) identified C. globiceps as the only species occurring in
southern Australian waters, an observation consistent with the material examined during this study.
The distribution of C. crassiceps appears to be disjunct, with populations in the central and southern
Atlantic Ocean and the mid-central and southern Pacific Ocean.

Several characters may be used to differentiate C. globiceps from C. crassiceps. Comparison of
data for C. crassiceps provided by Sazonov and Shcherbachev (1985) shows C. globiceps off WA
have a distinctly larger orbit (24-28% HL cf. 18-23%) and smaller scales (18-19 rows below 1D.
and 14-16 rows below 2D., cf. 11-14 and 9-13, respectively). Additionally, three to four rows of
closely spaced teeth occur on the premaxilla, compared with two rows of sparse teeth in C. crassiceps.

170 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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Sazonov and Shcherbachev (1985) also use the presence of a band of naked skin adjacent to the anal
fin in C. crassiceps and its absence in C. globiceps to differentiate the two species. However, this is
a variable character in the BMNH type specimens of C. crassiceps and may not be definitive.
Cetonurus globiceps was one of the most abundant grenadiers on the mid-continental slope region
off Western Australia, representing about 21% of the total grenadier biomass (AW, personal data).

Coryphaenoides Gunner, 1765

For more recent treatments and diagnoses of this genus see Iwamoto and Sazonov (1988),
Iwamoto (1990), Iwamoto and Shcherbachev (1991), and Shcherbachev and Iwamoto (1995).
Representatives of this diverse genus were relatively sparse in collections from Western Australia,
probably because only a small percentage of trawls were made at depths exceeding 1,000 m. The
primary vertical distribution of the genus appears to be at midslope to lower-slope depths, between
roughly 800 and 2,000 m. Many of the species, especially of the subgenera Chalinura and Lionurus,
are deeper living, commonly found at depths between 2,000 and 4,000 m. One species, C. yaquinae
Iwamoto and Stein, 1974 of the North Pacific, is known from greater than 6,000 m. Undoubtedly,
future work in deeper waters off WA will result in a much-expanded list of Coryphaenoides species.
Notably absent from our list were the common, widespread abyssal species C. armatus Hector, 1875
and C. filicaudus (Ginther, 1878). In addition, C. mcmillani Iwamoto and Shcherbachev, 1991 and
C. murrayi Ginther, 1878, both known from off VIC, should be expected off the southern coast of
WA.

More than 60 species, only four so far known from Western Australia.

KEY TO SPECIES OF CORYPHAENOIDES FROM WESTERN AUSTRALIA

la. Pelvic fin rays 7; large, conspicuous, tubercular scutes at tip and lateral angles of snout; spinules on body scales

leaf=shapediomlanceolate:® tg: Sy. duc towed 2 eee BAL SEY ERLE ee ee C. serrulatus
lb. Pelvic fin rays 8-12; snout tipped with rather inconspicuous tubercular scales; spinules on body scales conical . . . . 2
Das Pelvic taySshl al 2. Books oP Ss cs. sig sts. deeeet Sue We poke Wb easy s Gye cus, co cael ane C. striaturus
ODE PelviciaysS—llO! oe need wee 4S ag fe ys IS .4 Se SY Oy Bien we eo a eh Pe Ge 0g ee ee 3
San Outer cillishiigreatlvnestrictedno—9 clone: outer eillirakers) 30%) eee cn een C. rudis
SbeOuternscillishii7—24%vofldesoutercillirakersi7—9 26. -bad Geen lone) ane cei cote oe ee ee C. sp.

Coryphaenoides rudis Gunther, 1878
Bige27,

Coryphaenoides rudis Giinther, 1878:24 (n. of Kermadec Is., 1,152 m, Challenger sta. 170A). Williams et al.
1996:149 (WA).

Macrurus (Macrurus) rudis: Ginther, 1887:131.

Nematonurus macrocephalus Maul, 1951:17 (Madeira).

Macrourus paradoxus Smith and Radcliffe in Radcliffe 1912:115—116, pl. 25, fig. 1 (Philippines).

Coryvphaenoides paradoxus: Iwamoto and Sazonov, 1988:72—75, figs. 3c, 24, 28 (central and se. Pacific).

DIAGNOSIS. — Pelvic fin rays 9-10 (rarely 8 or 11); GR-I about 3 or 4 in outer, 9 or 10 in inner
series; premaxillary teeth in narrow band, with an enlarged outer series; mandibular teeth in about 3
irregular series anteriorly, becoming uniserial posteriorly; orbit diameter 17-20% of HL, preoral
length 10-12%, barbel length 15-20%, length outer gill slit 69%; body scales relatively adherent,
densely covered with small, short, conical spinules in an irregularly quincunx or divergent-row
pattern.

SPECIMENS EXAMINED. — WA: CSIRO H2551-14(600 mm TL); w. of NW Cape; 1 ,460—1,500+
m; SS1/91/12. CSIRO H2552-13 (540 TL); nw. of Point Cloates; 1,305 m; SS1/91/13. CSIRO

IWAMOTO AND WILLIAMS: GRENADIERS OF WESTERN AUSTRALIA 171

H2561-03 (1 spec.); w. of Cape Cuvier; 1,320 m; SS1/91/23. NSW: AMS 1.29340-001 (144 HL, 710
TL); off Sydney; 1,120—1,170 m; K89-13-02.

COUNTS AND MEASUREMENTS (data compiled mostly from Marshall and Iwamoto 1973;
Iwamoto and Sazonov 1988; and Shcherbachev and Iwamoto 1995). — 1D. II,9—11; P.119-4121 (rarely
i17 or i118); GR-II (outer/inner) 8—10 /9; scales 1D. 7—9, midbase 1D. 5.5—7.5, 2D. 5.5—6.5, lat.line
30-47; caeca 12-16.

The following in percent of HL: snout 23—29; preoral 10—12; internasal 1 7—19; interorb. 26-35;
suborb. 14—15; postorb. 57—S9; orb.-preop. 51—57; up.jaw 37-43; pre-A. 156—168; V.-A. 48-55; body
depth 80—100; ht. 1D. 43—73; len. P. 45—63; len. V. 50-111 (usually 50—70).

SIZE. — A large species, to at least 120 cm TL.

DISTRIBUTION. — Widespread in Atlantic, Pacific, and Indian oceans. In Australia, known from
NSW and WA (nw. coast), in 1,120—1,700 m.

COMPARISONS AND REMARKS. — Coryphaenoides rudis is unlikely to be confused with the other
three species of the genus from WA because of its pelvic fin ray count and low gill raker numbers.
The other three species have more and much better-developed gill rakers, especially on the outer series
of the first arch. Iwamoto and Sazonov (1988) treated the species (as C. paradoxus) in detail:
Shcherbachev and Iwamoto (1995) synonymized C. paradoxus and C. macrocephalus with C. rudis.

Coryphaenoides serrulatus Ginther, 1878
Fig. 28

Coryphaenoides serrulatus Ginther, 1878:26 (ne. of New Zealand; 1,280 m). McMillan in Gomon et al.
1994:356, fig. 316 (deser.; NSW, VIC, TAS, and New Zealand; 600—1,000 m). Williams et al. 1996:149
(WA).

Coryphaenoides s. serrulatus: Iwamoto and Shcherbachev, 1991:218-221 (NSW, TAS, VIC, SA and New
Zealand; 540—1,100 m).

DIAGNOSIS. — Pelvic fin rays 7; GR-I 7—9 in outer, 11—14 in inner series; premaxillary with
moderately wide band of small inner teeth flanked by enlarged teeth with arrowhead-shaped tips;
mandibular teeth in one row except about two at symphysis; orbit diameter 32-37% of HL, preoral
length 12-18%, barbel 23-30%, outer gill slit 16-23%; body scales covered with small, greatly
reclined, lanceolate spinules aligned in somewhat converging rows.

SPECIMENS EXAMINED. — WA: NMV A6192 (2, 405-425 mm TL); 95 km sw. of Esperance;
34°35.5’S, 121°19’E; 1,015—1,030 m. NSW: NMV A17 (285 TL) and NMV A9077 (2, 358-442 TL);
56 km ene. of Nowra; 34°43.95'S, 151°14.74’E; 1,009-817 m. NMV A5783 (336 TL); off Nowra;
35°00.0'S, 151°16.3’E; 1,100 m. VIC: NMV A3400 (5 spec.); 38 km sw. of Cape Bridgewater;
38°38'S, 141°04’E; 990-1,100 m. NMV A3412 (2 spec.); 45 km sw. of Cape Bridgewater; 38°36'S,
140°59’E; 1,040—1,170 m. NMV A6937 (2, 356-391 TL); s. of Point Hicks; 38°21.9’S, 149°20.0'E;
1,000 m. SA: NMV A5881 (2, 319-404 TL); 110 km w. of Robe; 37°19.04'S, 138°31.05’E;
990—1,000 m. CSIRO T1877-02 (1 spec.); w. of Cape Banks; 37°47'S, 139°39’E; 500-520 m;
Margaret Philippa; 6.X11.1984. TAS: CSIRO T359 (1 spec.), T381 (1 spec.); e. of Bicheno; 41°48’S,
148°38'E; 950—1,099 m; Challenger; coll. M. Wilson; 21.1V.1982. CSIRO T362 (1 spec.); e. of Long
Pt, 41°46'S, 148°37'E; 830-850 m; Bluefin; coll. R. Green; 18.V.1982. CSIRO T399 (1 spec.); nw.
of Cape Sorell; 42°25'S, 144°43’E; 956-835 m; Petuna Endeavour; coll. K. Evans; 7.VI.1982.
CSIRO T519 (476 TL); w. of Sandy Cape; 41°19’S, 144°03’E; 864-943 m; Challenger; coll. M.
Wilson; 15.XII.1981. CSIRO T527 (1 spec.); sw. of King I.; 40°44’S, 143°30'E; 920-927 m:;
Challenger, coll. M. Wilson; 16.XII.1981. CSIRO T570 (360 TL), T578 (378 TL), T599 (343 TL),
T601 (357 TL), T618 (321 TL), T807 (298 TL); sw. of King I.; 40°42’S, 143°29’E: 942-1,058 m;

172 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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28 cm

FIGURE 27. Coryphaenoides rudis Giinther. CSIRO H2561-03, from west of Cape Cuvier, WA, in 1,320 m.

-——_— — —

FIGURE 28. Coryphaenides serrulatus Giinther. CSIRO H3008-13, from southwest of Albany, WA, 842 m.

IWAMOTO AND WILLIAMS: GRENADIERS OF WESTERN AUSTRALIA 173

Challenger; coll. M. Wilson; 9.1.1982. CSIRO T591 (1 spec.); sw. of King I.; 40°44’S, 143°30’E;
920-927 m; Challenger; coll. M. Wilson; 16.X1I.1981. CSIRO T658 (328 TL); ne. of Flinders I.;
39°05'S, 148°43’E; 947 m; 1983. Southern Ocean, South Tasman Rise: NMV A3689 (270 TL):
47°42.0'S, 148°10.0’E; 1,200 m. NMV A3711 (320 TL); 47°29.0'S, 148°29.0‘'E. NMV A3719 (310
TL) and NMV A3721 (230 TL); 47°29.0'S, 148°30.0’E.

COUNTS AND MEASUREMENTS. — ID. II,9—11; P. 118-4124; (outer/inner) GR-II] 10—14/10—14;
scales 1D. 5.5—7.5, midbase 1D. 5.5—7.5, 2D. 7—10, lat.line 35-41; caeca 14-23.

The following in percent of HL: snout 26—31; internasal 16—19; interorb. 20—24; suborb. | 1—14;
postorb. 43—S0; orb.-preop. 37-45; up.jaw 39-44; pre-A. 148-186; V.-A. 44-67; isth.-A. 86—1 16:
body depth 74-108; 1D.-2D. 57-107; ht. 1D. 76—102; len. P. 50-74; len. V. 50-80.

SIZE. — To more than 48 cm TL.

DISTRIBUTION. — Off New Zealand, southern Australia (NSW, TAS, VIC, SA, and WA), and
oceanic elevations of the Indian Ocean, in 540—1,255 m.

REMARKS. — Iwamoto and Shcherbachev (1991) found geographic differences in a number of
characters and accordingly designated the oceanic population in the Indian Ocean as a distinct
subspecies. Specimens from the Great Australian Bight differed slightly from those captured off New
Zealand and in the Tasman Sea (including off the east coast of Tasmania), but the differences were
insufficient to warrant recognition of another subspecies. Among the 4 species of the genus from WA,
C. serrulatus can be easily distinguished by the presence of large terminal and lateral snout scutes, 7
pelvic fin rays, and lanceolate scale spinules. The closely similar C. mcmillani Iwamoto and
Shcherbachev, 1991 has a smaller, more-upturned terminal snout scute, more pelvic fin rays (8 or 9),
a rudimentary barbel, and needlelike scale spinules.

Coryphaenoides striaturus Barnard, 1925
Fig. 29

Coryphaenoides (Chalinura) striatura Barnard, 1925:500, 501 (off Cape Point, South Africa, in 823—1,737 m).
Coryphaenoides striaturus: lwamoto and Shcherbachev, 1991:214—217 (s. hemisphere distr., from se. Atlantic
to New Zealand; 823—2,010 m). Williams et al. 1996:149 (WA).

DIAGNOSIS. — Pelvic fin rays 11—12 (usually 12); GR-I 7-11 in outer, 12—16 in inner series;
premaxillary teeth in broad cardiform band flanked by an outer row of enlarged, slender, widely
spaced canines; mandibular teeth in one irregular row; orbit diameter 18—24% of HL, preoral length
10-16%; barbel length 18-26%, outer gill slit 16-23%; body scales rather deciduous, covered with
short, slender spinules aligned in 9—14 low parallel rows. (Adapted from Iwamoto and Shcherbachev
199 18)

SPECIMENS EXAMINED. — WA: CSIRO H2617-09 (460 mm TL); w. of Bunbury; 33°15.8’S,
114°11.1’E; 982 m; SS1/91/85. CSIRO H3017-06 (1 spec.); nw. of Cape Leeuwin; 34°10’S,
114°16’E; 1,030 m; Akebono Maru No. 3, shot 17; A. Williams; 24.XII.89. CSIRO T298 (1 spec.);
Great Australian Bight; 33°27'S, 128°36’E; 1,027—1,044 m; Margaret Philippa; coll. K. Evans;
24.X1.1984. VIC: CSIRO H1519-06 (1 spec.); s. of Warrnambool; 39°12.3’S, 142°30.0’E; 1988.
TAS: CSIRO H1260-19 (1 spec.); se. of St. Patricks Head; 1,195—1,185 m; SO2/88/87. CSIRO
H1523-02 (1 spec.); w. of Low Rocky Point, 42°54.5'S, 144°45.9’E; 1,785 m; 1988.

COUNTS AND MEASUREMENTS (from Iwamoto and Shcherbachev 1991).— 1D. II,8—10; P.
118-124; GR-I (outer/inner) 11—14/11—15; scales 1D. 7.5-10, midbase 1D. 5—7.5, 2D. 8-9, lat.line
30-36; caeca 9-12.

Total length 120-550 mm; HL 21-97 mm. The following in percent of HL: snout 26-30;
internasal 20—28; interorb. 23—30; suborb. 1 1—15; postorb. 51-57; orb.-preop. 46-53; up.jaw 38-46;

174 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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pre-A. 152—180; V.-A. 42-74; body depth 81—103; ht. 1D. 81-109; len. P. 56-68; len. V. 65-97; post.
nostril 3—7.

SIZE. — To about 55 cm TL.

DISTRIBUTION. — From the se. Atlantic, s. Africa, Indian Ocean between about 25°S to 35°S, s.
Australia including WA, SA, VIC, TAS, NSW, Tasman Sea, and New Zealand, in 823—2,010 m (data
mostly from Iwamoto and Shcherbachev 1991).

COMPARISONS AND REMARKS. — The completely scaled underside of snout, pelvic ray count,
relatively numerous gill rakers, and features of the dentition and body scales together serve to
distinguish C. striaturus from its congeners.

The species is widespread in southern temperate waters of the eastern hemisphere at depths
usually between 1,000 m and 1,400 m. It appears to be relatively abundant in places, but off WA the
species was uncommon and not captured north of Cape Leeuwin (about 35°S).

Coryphaenoides sp.
Fig. 30

Coryphaenoides (Coryphaenoides) sp. indet. Iwamoto, 1986:355 (s. Africa, Mozambique; 696—960 m).

Coryphaenoides (Coryphaenoides) rudis (nec Giinther, 1887): McCann and McKnight, 1980:35, figs. 16, 17,
18 (part) (New Zealand; 2,993 m).

Coryphaenoides sp.: Okamura in Amaoka et al., 1990:184, fig. 126 (New Zealand; 100-763 m). McMillan in
Gomon et al. 1994:355, fig. 315 (“Humpback Whiptail”; descr.; NSW, VIC, TAS, and New Zealand; “in
depths in excess of 800 m”).

Coryphaenoides sp. B: Paulin et al., 1989:129, fig. 56.30b (in key). Williams et al. 1996:149 (WA).

Coryphaenoides sp. 1: Shcherbachev and Iwamoto, 1995:309-312, fig. 8 (distr. New Zealand, Tasman Sea,
Coral Sea, Indian Ocean, se. Atlantic; 696—1,600 m).

DIAGNOSIS. — Pelvic fin rays 8 (rarely 9); naked areas on underside of head confined to a median
swath below snout tip, a narrow strip along ventral margins of snout and suborbital, and anterior end
of mandible; snout low, barely protruding in adults; premaxillary teeth in a broad band flanked by a
row of moderately enlarged and spaced conical teeth; mandibular teeth in 3 or 4 irregular rows; orbit
diameter 19-23% of HL; preoral length 7—11%; barbel length 21—34%; outer gill slit 17-24%; body
scales adherent, with numerous, small, reclined, closely packed spinules in subparallel to slightly
convergent rows.

SPECIMENS EXAMINED. — (Numerous specimens including those listed by Shcherbachev and
Iwamoto 1995).

COUNTS AND MEASUREMENTS (compiled mostly from Shcherbachev and Iwamoto 1995). — 1D.
11,9—10; P. 117-121; GR-I (outer/inner) 7—9/11—13; GR-II 9-12/11—13; scales 1D. 9.5—11, midbase
1D. 7-10, 2D. 7-12, lat.line 38-48; caeca 10-18.

Total length 140-770 mm; HL 24-170 mm. The following in percent of HL: snout 25—29;
interorb. 14-20; suborb. 9-13; postorb. 48-57; orb.-preop. 44-50; up.jaw 40-45; pre-A. 143-179;
V.-A. 43-76; body depth 62-89; ht. 1D. 65—84; len. P. 43-57; len. V. 43—70; post. nostril 4-8.

SIZE. — To at least 77 cm TE.

DISTRIBUTION. — Widespread throughout southwestern Pacific Ocean, Indian Ocean, and se.
Atlantic Ocean, in about 700—1,600 m (Okamura in Amaoka et al 1990:184, has recorded it from
“100-763 m’”’). Found in Australia off NSW, TAS, VIC, SA, and WA.

COMPARISONS AND REMARKS. — This large species is extensively treated in Shcherbachev and
Iwamoto (1995) and will be described soon by Peter McMillan of New Zealand (in litt. to TI, July
1998). It is unlikely to be confused with any other member of the genus currently known from
Australia because of its peculiar physiognomy of low, scarcely protruding snout, shallow head, high

IWAMOTO AND WILLIAMS: GRENADIERS OF WESTERN AUSTRALIA 175

FIGURE 29. Coryphaenides striaturus Barnard. CSIRO H3017-06 from northwest of Cape Leeuwin,WA, 1,030 m.

FIGURE 30. Coryphaenides sp. (“longbeard”). CSIRO H3010-09 from south of Cape Leeuwin, 945 m.

(“hump-backed”) nape, combined with usually eight pelvic fin rays and distinctive squamation of
head and body.

Hymenocephalus Giglioli, 1884

We use the generic name in the broadest sense to include Hymenogadus and Spicomacrurus as
subgenera.

KEY TO SPECIES OF HYMENOCEPHALUS FROM WESTERN AUSTRALIA

Li, [SINGH EYES Ryne td ere, | SACRA area ear Ook a er ee eae OR re, AR We Pees Pens a, Bo 2
J8, REMC TaN EC Ree Se ae ea ne en ee ee eee See meme Pangea he. kan Penn ic 5
2a. Inner gill rakers on first arch 19 or more; no broad, horizontal projections at snout tip ................. 3

2b. Inner gill rakers on first arch 1 1-12; three broad, horizontal projections forming tip and lateral angles of snout
CRC cian BN al eet Reon cae Got nce Moy od aaa Shoals oe wa nae Modus sage else see salem lo Ina Oe H. adelscotti

176 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 3

3a. Chin barbel long, greater than least suborbital width; melanophores present to end of tail. ........ H. longibarbis
3b. Chin barbel short, less than half suborbital width; melanophores absent posterior to belly region

Pe EER os Gem eG De Bw a ale Wha a eae se W Gem hos eae [?]H. striatissimus aeger
4a. Pelvic fin rays 12—14; orbits 23-29% of head length; interorbital greater than orbit diameter (about 0.8—0.9 into

Groin) yorpbleaboutley—2slsinto)distance orbit-preopercle = 4 = 92.2 mares. 4 2 ee ee ee H. sp.
4b. Pelvic fin rays 11—12; orbits 31-38% of head length; interorbital less than orbit diameter (1.2—1.4 into orbit), orbit
ASE amin Ox StancevOnrDit=preoperncle) <0. ns )ster se cw GG S = pee es, se H. nascens

Hymenocephalus adelscotti }wamoto and Merrett, 1997
Fig. 3la

Hymenocephalus adelscotti wamoto and Merrett, 1997:514—516, fig. 19 (holotype MNHN 1994-882, 162 mm
TL; n. of Fiji, 600 m).

DIAGNOSIS. — Pelvic fin rays 9, outer ray slightly expanded distally; nasal bones forming 3
horizontal platelike processes; gill rakers on outer arch 11; head bones relatively firm; barbel 3.8 into
orbit.

SPECIMEN EXAMINED. — WA: CSIRO H2573-14 (37.0 mm HL, 166+ mm TL); w. of Shark
Bay; 680-691 m; SS1/91/36.

COUNTS AND MEASUREMENTS. — ID. II,11; P. 119-120; V. 9; total GR-I (inner) 12, GR-II
(outer/inner) 11/12; scales 1D. 3, 2D. 4.

Total length 166+ mm; 37.0 mm HL. The following in percent of HL: postrostral 74; snout 31;
preoral 17; internasal 19; interorb. 18; orb. 32; suborb. 10; postorb. 44; orb.-preop. 36; up.jaw 36;
barbel 8; gill slit 24; pre-A. 158; pre-vent 154; V.-A. 66; isth.-A. 104; body depth 50; depth at A. 45;
1D.-2D. 59; ht. 1D. 63; len. P. 62; len. V. 49; post. nostril 12.

DESCRIPTION. — Body slender, subcylindrical; head bones relatively firm (compared to most
other Hymenocephalus species); crests on nasal, frontal, and occipital low; snout depressed, with
median and lateral processes of nasal bones forming three blunt, horizontal platelike processes; lateral
margins of snout concave posterior to processes. Underside of head flat; mouth moderately large,
nearly horizontal. Teeth small, slender, conical. Gill rakers short, tubercular, bearing few spines.
Scales large, deciduous (none remaining on our specimen). Second spine of first dorsal fin smooth.
Elongate outer ray of pelvic fin slightly expanded and fleshy. Light organ long, luminous tissue joining
a small lens anterior to vent, another lens just anterior to pelvic fin bases. Ventral striae well developed.
Color dusky brown, paler on ventral surface. Abdominal region and isthmus black, underside of head
dark. Lip of upper jaw black, surrounding tissue pale.

SIZE. — Our single specimen was about 17cm TL, but had a damaged tail and developed
pseudocaudal.

DISTRIBUTION. — Known only from the holotype captured north of Fiji and our single WA
specimen; capture depths 600-691 m.

COMPARISONS AND REMARKS. — Our specimen showed great similarity to Hymenocephalus
kuronumai (Kamohara, 1938), the only other species of the subgenus Spicomacrurus, in overall
morphology, counts and proportional measurements. However, it differs in having longer platelike
median and lateral processes of the nasal bone than figured for H. kuronumai by Okamura (1970).
This accounts for the shorter snout-to-head ratio (3.3 cf. 3.64.5) in our specimen. The WA specimen
also has a shorter barbel (3.8 into orbit diameter cf. 1.5—2.4) and smaller scales (4 rows from midbase
of first dorsal fin to lateral line, cf. 3 rows, and 3 rows from the origin of second dorsal fin, cf. 2).

IWAMOTO AND WILLIAMS: GRENADIERS OF WESTERN AUSTRALIA 177

Hymenocephalus longibarbis (Gunther, 1887)
Fig. 31b

Macrurus (Mystaconurus) longibarbis Ginther, 1887:139, 140, pl. 18, fig. C (Fiji Islands, Challenger sta. 173;
576 m).

Hymenocephalus sp.: Gomon et al., 1994:358, fig. 318 (GAB; 366 m).

Hymenocephalus longibarbis: lwamoto and Merrett, 1997:520—521, fig. 21b (New Caledonia region).

DIAGNOSIS (for WA specimens only).—Pelvic fin rays 8; chin barbel long, 0.55—0.63 into orbit
diameter; gillrakers | 7—21 on inner series of first arch; interorbital narrow, |.4—2.1 into orbit, 16-20%
HL; suborbital narrow, 2.3—2.9 into orbit, 11—13% HL; midlateral stripe becoming faint or obscure
posteriorly on tail, no prominent black marks on dorsal surfaces of trunk.

MATERIAL EXAMINED (9 spec.)—WA: AMS 1.22808-029 (5, 23.5—35.2 mm HL, 142-192 mm
TL); NW Shelf; SO2/82/17-18. AMS I.22826-006 (5, 29.4-40.9 HL, 167+—200+ TL); NW Shelf;
396-406 m; SO1/86/53.

SIZE. — To more than 23 cm TL.

DISTRIBUTION. — Australia (WA, NSW, QLD, ?SA), Fiji, and New Caledonia in 366-620 m.

COMPARISONS AND REMARKS. — This large member of the genus can be readily differentiated
from its congeners by the combination of long barbel, slender body, low, non-protruding snout,
relatively low gill raker counts, and eight pelvic fin rays. In having a notably long barbel, H.
longibarbis is most similar to H. longiceps Smith and Radcliffe, 1912 from the Philippines, but the
barbel in H. longiceps goes 1.0—1.4 times into the orbit diameter (cf. 0.55—0.63) and the suborbital
width goes 2.0—2.3 (cf. 2.32.9) times into the orbit.

The Western Australian specimens differed slightly from specimens recorded by Iwamoto and
Merrett (1997) from New Caledonia. The most striking differences were related to proportions of the
barbel, interorbital, and suborbital. Thus, the WA specimens had for these characters, respectively,
values of 0.55—0.63, 1.4—1.7, and 2.32.9 times into orbit diameter, in contrast to 0.7—1.0, 1.7—2.1,
and 3.14.6 in the New Caledonian specimens. The WA specimens were also much paler overall,
without bold body and head markings, but such differences are often attributable to differences in
preservation techniques. The paucity of specimens and the lack of a more thorough examination of
material from other areas prevents a decision as to whether or not they represent a new taxon.

That only two small WA collections were made of this species is somewhat surprising, in that it
(or its sister taxon) is apparently rather abundant off New Caledonia and the east coast of Australia.

Hymenocephalus nascens Gilbert and Hubbs, 1920
Fig. 32a

Hymenocephalus nascens Gilbert and Hubbs, 1920:535—539, fig. 30 (holotype USNM 78229; Borneo, vicinity
of Sibuko Bay, 759 m). Iwamoto and Merrett, 1997:523—525, fig. 22b (New Caledonia region, 600-855 m).

DIAGNOSIS. — Chin barbel absent or rudimentary, much less than half suborbital width; V.
11-12; snout length 25-30% HL, orbit broadly oval, 31-37% HL, slightly greater than interorbital
width, 25-28% HL; suborbital width 13-14% HL. Prominent midlateral stripe, ventrolateral surfaces
of tail finely speckled. Scale margins dorsally well marked; preopercle mostly pale; dark areas of
belly and chest with purplish tinge. Anterior lens of light organ small, about half size of posterior
lens.

SPECIMENS EXAMINED. — WA: CSIRO H1492-19 (2, 23.5-28.6 mm HL, 120—-120+ mm TL);
w. of Barrow I.; 420 m; SO5/88/70. CSIRO H2573-24 (24.0 HL, 123 TL); w. of Steep Point; 691 m;
SS1/91/36. CSIRO H2580-09 (25.5 HL, 120+ TL); sw. of Shark Bay; 491 m; SS1/91/43.

178 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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a

ae Be eh RRs og te Nt SE
HS ES DP ODL . ee bd ee
LALA PALATINE SG gO

Pe ae
lena

FIGURE 31. (a) Hymenocephalus adelscotti |wamoto and Merrett. Holotype, MNHN 1994-882 (33 mm HL), from near Wallis
and Futuna islands, southwestern Pacific, in 600 m. (From Iwamoto and Merrett 1997, fig. 19.) (b) Hymenocephalus longibarbis
(Giinther). NMNZ P.29239 (35.5 mm HL) from south of New Caledonia in 550—920 m. (From Iwamoto and Merrett 1997,
fig. 21b.)

COUNTS AND MEASUREMENTS. — ID. II,9; P. 114-4115; V. 11-12; GR-I (outer/inner) 1 7—18/24,
GR-II 23—24/23-—24. The following in percent of HL: postorb. 43-45; orb.-preop. 43-45; up.jaw
50-56; body depth 68—76; body width 44-47; head width 51—S5; len. V. 82—86.

COMPARISONS AND REMARKS. — These four Western Australian specimens agree well with the
detailed original description of the species by Gilbert and Hubbs (1920) and that of New Caledonian
specimens by Iwamoto and Merrett (1997), with the exception of the pelvic fin ray count of 11—12,
which agrees more with that of H. /ethonemus. The rudimentary barbel in three of the four specimens
is a bit disconcerting, but not especially surprising, in that a similar situation was recorded in
specimens of H. striatulus Gilbert, 1905 from Hawaii and the Sala y Gomez Ridge (Sazonov and
Iwamoto 1992).

|?|Hymenocephalus striatissimus aeger Gilbert and Hubbs, 1920
Fig. 32b

Hymenocephalus striatissimus aeger Gilbert and Hubbs, 1920:531—534, fig. 32 (holotype, USNM 78228,
Molucca Sea off Halmahera, 545 m).

IWAMOTO AND WILLIAMS: GRENADIERS OF WESTERN AUSTRALIA 179

Spade?

sietane teen

FIGURE 32. (a) Hymenocephalus nascens Gilbert and Hubbs. NMNZ P.29101 (26.2 mm HL), from south of New Caledonia
in 550-690 m. (From Iwamoto and Merrett 1997, fig. 22b.) (b) Hymenocephalus striatissimus aeger Gilbert and Hubbs.
Holotype (USNM 78228, 158 mm TL) from East Indies “between Gillolo and Makyan Islands” in 542 m. (From Gilbert and
Hubbs 1920, fig. 32.)

DIAGNOSIS. — Pelvic fin rays 8 [7 rays in | of 15 spec.]; total gill rakers on first arch 12-18
(lateral series), 20-23 on inner series; barbel rudimentary 24% HL; orbits 33-45; dorsolateral stripe
on trunk fades out completely on tail.

SPECIMENS EXAMINED. — WA: CSIRO H1514-35 (11, about 24-26 mm HL, 108+—153 mm
TL); NW Shelf; 582 m; SO5/88/190. CSIRO H3144-07 (5, 24.5-28 HL, 95+—-142+ TL); n. of
Dampier Archipelago; 18°51'S, 116°02’E; 550 m; Surefire, shot 3; coll. D. Evans; 16.11.1992. AMS
1.22816-003 (11, about 25.5-27 HL, 115+—262+ TL); NW Shelf, 220 km nw. of Port Hedland;
594-612 m; SO2/82/28-30.

COUNTS AND MEASUREMENTS. — 1D. II,7—9; P. i10-i13; total GR-II (outer/inner) 19—23/19-21.

Total lengths 95—-262+ mm, HL 24-28 mm. The following in percent HL: snout 26-29 (2 spec.);
interorb. 35—40; suborb. 14-18; postorb. 41—50; orb.-preop. 45—54; up.jaw 48-59; body depth 64-82;
body width 35-47; head width 48-55; ht. 1D. 68-92; len. P. 54-85; len. V. 48-69.

SIZE. — To at least 26 cm TL.

DISTRIBUTION. — Only three collections were made of the species off WA in 550-612 m.

COMPARISONS AND REMARKS. — This species is easily recognized among the Western Austra-
lian members of Hymenocephalus by the low pelvic and pectoral ray counts, rudimentary chin barbel,
and the complete absence on the tail of a dorsolateral stripe or speckling of any sort.

Our Western Australian specimens (all of which are damaged and in poor condition) seem on
the whole to agree with H. striatissimus aeger. The small, rudimentary barbel is much smaller,
however, than that reported for the subspecies (much less than half pupil diameter, cf. 2/3 to more

180 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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than pupil diameter in H. s. aeger). The pigmentation on the flanks posterior to the abdominal cavity
is wholly absent in our specimens, but appears to be present in H. s. aeger, as judged from the original
illustration of the subspecies. Some badly faded type specimens (CAS-SU 25463) lack pigmentation
posterior to the abdomen, but their absence may be an artifact of long preservation. Additionally, the
orbits appear more round in outline and not as deeply oblique as in H. s. aeger.

Compared with H. megalops from the New Caledonia region and QLD, H. s. aeger has a shorter
barbel (8-22% HL in H. megalops), fewer pectoral fin rays (19-113 cf. 113-115), a somewhat longer
orbit-to-preopercle distance (45-54% HL cf. 36-47%), a somewhat broader suborbital (14-18% HL
cf. 8-16%), and a broader interorbital space (35-40% HL cf. 24-36%). The eight pelvic fin rays, the
large orbit, and the absence of pigmentation on the tail are, however, characteristic of both species.

Hymenocephalus (Hymenocephalus) sp.

DIAGNOSIS. — Pelvic fin rays 12—14; barbel absent; orbits about 23—29% HL, less than interor-
bital width; sides of trunk and tail heavily speckled; dorsolateral stripe prominent, extending
posteriorly from behind head to end of tail; anterior lens of light organ tiny, inconspicuous.

SPECIMENS EXAMINED. — WA: CSIRO H2553-11 (33.5 mm HL, 153+ mm TL); w. of Pt.
Cloates; 910 m; SS1/91/14. CSIRO H2583-13 (29.1 mm HL, 108+ mm TL); nw. of Shoal Pt.;
945-960 m; SS1/91/48. CSIRO H2584-14 (27.4 HL, 132+ TL) and CSIRO H2584-15 (29.6 HL, 140+
TL); w. of Shoal Pt.; 853 m; SS1/91/49.

COUNTS AND MEASUREMENTS. — ID. II,9-11; P. 112-115; V. 12—14; total GR-I (outer/inner)
17—19/24-26, GR-II 24—25/23—24.

Measurements in percent of HL: snout 33-39; orb. 23—29; suborb. 13—16; interorb. 27-31;
postorb. 46—52; orb.-preop. 46—52; up.jaw 45—54; body depth 46-59; head width 4246; ht. 1D.
64—69; len. P. 47-55; len. V. 81-82.

DESCRIPTION. — This species resembles H. nascens rather closely; thus, only major character
differences will be described. Snout relatively high, the thin, slender, flexible tip elevated above mouth
a distance of about two lens diameters. Suborbital region relatively broad, least width more than lens
diameter, almost entirely black along inner walls. Opercular region behind and below preopercle
ridge, chest, and gill membranes black.

COMPARISONS AND REMARKS. — These specimens do not readily key out to any species in the
keys provided by Gilbert and Hubbs (1920) and Iwamoto (1990). They follow the description of 7.
nascens most closely, but the larger orbit diameter and higher pelvic fin ray count differentiate the
specimens from H. nascens. They are also darker, with heavier pigmentation, especially on the
opercular and suborbital bones, and ventrally on the tail below the dorsolateral stripe, where large
melanophores are aligned in a V-shaped pattern (fine, more uniform arrangement of melanophores
in H. nascens). There is some similarity to H. striatulus, but that species has a much more robust and
thicker body, larger orbits, and stouter head bones.

We have not described this species as new because only five specimens, all in relatively poor
condition, were present in our examined collections.

Kuronezumia lwamoto, 1974
See Sazonov and Iwamoto (1992) for a diagnosis of the genus. Seven species currently recognized

(Shcherbachev et al. 1992); only two species found in WA, both rare in the area, but K. /eonis is
relatively common in TAS, SA, and NSW.

IWAMOTO AND WILLIAMS: GRENADIERS OF WESTERN AUSTRALIA 181

KEY TO SPECIES OF KURONEZUMIA FROM WESTERN AUSTRALIA

la. Pelvic rays 8-9 (rarely 10); snout relatively pointed, tipped with a large tubercular scale; upper jaw 28-35% HL
a 0 Br Bete tal lees Fees ae bee came eg ee OG ae ta ee i ed oo" K. leonis
lb. Pelvic rays 10-11; snout bluntly rounded, not tipped with large tubercular scale; upper jaw 36-42% HL . . K. pallida

Kuronezumia leonis (Barnard, 1925)
Figs 33

Lionurus leonis Barnard, 1925:503 (off Cape Point, South Africa).

Macruroplus leonis: Smith, 1949:134.

Nezumia leonis: Krefft, 1969:36 (Uruguay).

Nezumia sp. A: Okamura, in Amaoka et al., 1990:181 (New Zealand).

Kuronezumia leonis: Shcherbachev et al., 1992:101, 102 (deser.; distr.; Indian Ocean).

Nezumia sp.: McMillan, in Gomon et al., 1994:361, 362, fig. 322 (“snubnose whiptail”; Australia [VIC, TAS],
New Zealand).

DIAGNOSIS. — Snout relatively pointed, tipped with enlarged, bony, tubercular scale; V. 8-10,
usually 8 or 9.

SPECIMENS EXAMINED. — WA. CSIRO H3008-10 (1 spec.); sw. of Albany; 35°25’S, 117°21'E;
842 m; Akebono Maru No. 3, shot 8; coll. A. Williams; 22.X1I.1989

COUNTS AND MEASUREMENTS (of 1 WA spec.)—1D. II,9; P. 120-421; total GR-I (outer/inner)
9/10, GR-II 9/10; scales 1D. 15—15.5, 2D. 11—12.

Total length 317+ mm; HL 50.5 mm. The following in percent of HL: postrostral 73; snout 30;
preoral 19; internasal 24; interorb. 29; orb. 29; suborb. 17; postorb. 45; orb.-preop. 37; up.jaw 28;
barbel 26; gill slit 15; pre-A. 150; pre-vent 130; V.-A. 40; isth.-A. 81; body depth 80; depth at A. 72;
1D.-2D. 42; len. P. 65; len. V. 41; post. nostril 8.

SIZE. — Attains about 50 cm TL.

DISTRIBUTION. — Widely distributed in southern Africa, off Uruguay (not verified by current
authors), southern Australia, New Zealand, and southern Indian Ocean on the Kerguelen Plateau.

COMPARISONS AND REMARKS. — This wide-ranging species has been adequately described in
the recent works of Shcherbachev et al. (1992) and Iwamoto and Anderson (1994). The former work
offers a key to the seven recognized species in the genus.

Kuronezumia leonis differs most obviously from K. pallida, the second species of the genus
recorded off the west coast of Australia, in the presence of a large terminal scute on the snout. The
pointed snout and relatively slender body of K. /eonis also easily distinguished this species from K.
pallida.

Kuronezumia pallida Sazonov and Iwamoto, 1992
Fig. 34

Kuronezumia pallida Sazonov and Iwamoto, 1992:65—67 (Sala y Gomez Ridge, se. Pacific, 540-800 m).

DIAGNOSIS. — No large lateral line pores on head; length upper jaw 36-42% HL; inner GR-I
9-10; scales below 1D. 11-13; V. 10; light organ a small lenslike structure between pelvic fin bases,
no bulbous swelling; color pale brown, slightly lighter on ventral surface; fins dark, abdominal area
bluish.

SPECIMENS EXAMINED. — WA: CSIRO H3041-14 (106 mm HL, 495+ mm TL); wsw of Shark
Bay; 26°36’S, 112°09’'E; 760 m; Akebono Maru No. 3 sta. 41; coll. A. Williams; 28.XII.1989. CSIRO
H3192-02 (66.9 HL, 340 TL); n. of Dampier Archipelago; 18°46’S, 116°13'E; 550 m; Surefire, shot

182 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 3

2; coll. V. Wadley; 10.11.1992. CSIRO H3086-01 (93.5 HL, 505+ TL); n. of Port Hedland; 17°49’S,
118°17'E; 725 m; Courageous; coll. D. Evans; 7.11.1990.

COUNTS AND MEASUREMENTS (based on 3 spec.).—1D. II,9—10; P. 119-122; total GR-I (outer/in-
ner) 7—-8/9-10, GR-II 9/910; scales 1D. 11—13, 2D. 10, midbase 1D. 8-9.

Total length 340+—S05+ mm, HL 66.9—106.0 mm. The following in percent of HL: postrostral
77-19; snout 26-27; preoral 16—19; internasal 18-19; interorb. 24—25; orb. 27—28; suborb. 17—18;
postorb. 51—54; orb.-preop. 4446; up.jaw 36-42; barbel 24-26; gill slit 17-20; pre-A. 141—149;
pre-vent 105—122; V.-A. 43; isth.-A. 78-91; body depth 89-98; depth at A. 77-82; 1D.-2D. 3446;
len. P. 58-60; len. V. 50—S9; post. nostril 4-8.

DESCRIPTION. — Large, deep-bodied species with deep, compressed head; snout high, blunt,
with vertical anterior profile. Mouth large, jaws subterminal; lips and adjacent areas of buccal caviy
with dense covering of fleshy, branched papillae that almost cover teeth. Suborbital region deep,
vertical, without modified scales.

Body scales covered with radiating series of long, densely packed conical spinules.

Light organ manifested externally as a small, black, scaleless anterior dermal window between
pelvic fin bases.

Color when fresh, light brown with slightly darker area on dorsal posterior region of head, lighter
on ventral surface, abdominal area bluish; fins dark.

SIZE. — To about 5] cm TL.

DISTRIBUTION. — Only three specimens were taken from the west and northwestern coast of
Australia. Kuronezumia pallida was known previously only from the Sala y Gomez Ridge in the
subtropical South Pacific.

COMPARISONS AND REMARKS. — Our specimens closely agree with the original description of
K. pallida in most counts and proportional measurements, and particularly in the physiognomy of the
head, which is deep and compressed with a high, blunt snout. They match Sazonov and Iwamoto’s
(1992) diagnosis of that species precisely; however, compared to data from the large holotype, the
orbit of the Australian material is larger (27-28% HL cf. 24%), the suborbital width narrower
(17-18% HL cf. 20%), pectoral rays fewer (119-122 cf. 124-125), and pelvic rays fewer (10, cf. 11).
In addition, the scales appear more densely spinulated. We attribute these differences to natural
variation among the few known specimens, and to geographical variation in a widely distributed
species.

Kuronezumia pallida may be separated from K. /eonis by the physiognomy of its head, which is
deep and compressed, and its high blunt snout, which lacks a terminal bony scute. It differs from K.
dara (Gilbert and Hubbs, 1916) from Japanese waters in having a humped dorsal profile over the
posterior region of the head, a shorter snout (26-27% HL cf. 28-29%), and an overall paler color.

Lepidorhynchus Richardson, 1846
A monotypic genus whose relationships are obscure.

Lepidorhynchus denticulatus Richardson, 1846.
Fig. 35

Macrourus denticulatus Richardson, 1846:53 (“coast of South Australia’).
Coryphaenoides denticulatus: Ginther, 1862:396.

Macrurus (Optonurus) denticulatus: Giinther, 1887:147.

“Optonurus ” denticulatus: Gilbert and Hubbs, 1916:144. McCulloch, 1926:33.
Lepidorhynchus denticulatus: McCann and McKnight, 1980:50—S2.

IWAMOTO AND WILLIAMS: GRENADIERS OF WESTERN AUSTRALIA 183

FiGURE 33. Kuronezumia leonis (Barnard). (From Shcherbachev et al. 1992, fig. 3)

FiGuRE 34. Kuronezumia pallida Sazonov and Iwamoto. CSIRO H3041-14 (106 mm HL, 495 mm TL) from off Shark Bay,
WA, in 760 m.

FiGuRE 35. Lepidorhynchus denticulatus Richardson. CSIRO unregistered, from off southern coast of Australia.

DIAGNOSIS. — Bony ridges of head poorly developed and lacking modified scales. Mouth large,
terminal; teeth of premaxilla in broad bands with large, widely spaced canines in outer row; teeth of
mandible well developed, closer set than those of premaxilla. Ventral striae conspicuous over
abdominal region from level of pectoral fin, extending along ventral region from isthmus to midpoint
of anal fin.

SPECIMENS EXAMINED. — WA: CSIRO H2023-03 (86.5 mm HL, 527 mm TL); w. of Dongara

and Leander Pt.; 29°14’S, 113°52’E; 556 m; South Passage; coll. D. Wright; 28.1.1989.

184 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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COUNTS AND MEASUREMENTS (based on | spec.).—1 D. II, 11; P.116; V. 9; total GR-I (outer/inner
11/17, GR-II 16/16.

Total length 527 mm; HL 86.5 mm. The following in percent of HL: postrostral 75; snout 24;
preoral 12; internasal 16; interorb. 22; orb. 34; suborb. 7; postorb. 42; orb.-preop. 43; up.jaw 49;
barbel 9; gill slit 26; pre-A. 164; pre-vent 157; V.-A. 57; isth.-A. 112; body depth 84; depth at A. 71;
1D.-2D. 83; len. 42; post. nostril 6; len. rictus 40.

DESCRIPTION. — This species is quite distinctive and has been adequately described in recent
works. McCann and McKnight (1980), Last et al. (1980), and Gomon et al. (1994) provide accounts
of material from southern Australia.

SIZE. — To about 55 cm TL.

DISTRIBUTION. — Southern Australia and New Zealand. Reported from a wide depth range
(60—1,000 m), but predominantly an upper-slope species in 270-450 m.

COMPARISONS AND REMARKS. — The well-developed teeth and distinctive reflective, silvery
coloration of L. denticulatus make it easily recognizable. Extremely abundant off southeastern
Australia, where it forms a substantial bycatch to the demersal trawl fishery for blue grenadier (or
hoki, Macruronus novaezelandiae). It was caught only occasionally off the southern coast of Western
Australia.

Lucigadus Gilbert and Hubbs, 1920

Recent treatments of species of Lucigadus include Iwamoto and Anderson (1994) and Iwamoto
and Merrett (1997). Only one species so far known from WA.

Lucigadus ori (Smith, 1968)
Fig. 36

Macruroplus ori Smith, 1968:4—6, pl. 2 (off Durban, South Africa).
Ventrifossa ori: Iwamoto, 1979:153.
Lucigadus ori: lwamoto and Anderson, 1994:16.

DIAGNOSIS. — Mouth small, upper jaw about 31% HL; pelvic fin rays 15—18; pectoral fin rays
il 8-23; body and anal fin lacking banding pattern; branchiostegal membranes with two small scale
patches.

SPECIMENS EXAMINED. — WA: CSIRO H2579-02 (31.1 mm HL, 230 mm TL); sw. of Shark
Bay; 666-688 m; SS1/91/42.

Comparative material of Lucigadus nigromaculatus: NSW: AMS 1.7894 (holotype, 32.1 HL,
222 TL) and AMS I.7895, 1.7896, 1.7897 (paratypes, 15.3-32.4 HL, 112-207 TL); off Sydney;
33°52'S, 151°48’E; 1,461 m. AMS 1.15974-008 (32.3-36.7 HL); 55 n.mi. [102 km] off Newcastle;
366 m; K71-08-03. AMS 1.15975-032 (32.3—38.3 HL); 35 n.mi. [65 km] se. of Newcastle; 567 m;
K71-08-05. VIC: AMS E.5499 (28.4 HL); sse. of Genoa Point; 37°S, 149°E; 365 m; 1914. SA: AMS
1.18711-005 (31.8-37.9 HL); Great Australian Bight; 33°29’S, 127°15’'E; 1976. AMS IA.1367 (40.1
HL); Great Australian Bight, s. of Eucla; 31°S, 128°E; 730 m; 1913.

COUNTS AND MEASUREMENTS (1 WA spec.).—ID. II,10; P. 120421; V. 14-15; total GR-I
(outer/inner) 9—10/13, GR-II 12/12; scales 1D. 17—18, midbase 1D. 13—14, 2D. 8-9.

Total length 230 mm; HL 31.1 mm. The following in percent of HL: postrostral 78; snout 28;
preoral 23; internasal 20; interorb. 24; orb. 38; suborb. 16; postorb. 39; orb.-preop. 32; up.jaw 31;
barbel 15; gill slit 18; pre-A. 163; pre-vent 137; V.-A. 46; isth.-A. 95; body depth 109; depth at A.
92; 1D.-2D. 65; len. 71; len. V. 47; post. nostril 6.

IWAMOTO AND WILLIAMS: GRENADIERS OF WESTERN AUSTRALIA 185

DESCRIPTION. — Body short, deep, and compressed, tapering from anal fin origin to end of long,
compressed tail. Head short, compressed, lacking distinctive ridges; dorsal profile nearly straight
(Madagascar specimens examined have a distinct hump), rising steeply to first dorsal fin origin. Snout
short, rounded, barely protruding beyond level of mouth, about 1.4 into orbit. Mouth short, oblique;
upper jaw extends posteriorly to vertical about one-third distance across orbit. Orbits relatively large,
about 1.5 into interorbital width. Suborbital ridges flat, lacking enlarged or bony scales.

Teeth of premaxilla and mandible conical, forming broad, tapering bands; 4-5 rows of teeth at
symphysis, outer row in each jaw slightly enlarged.

Gill rakers bearing clusters of elongate, straight to slightly curved spines; dorsal rakers on first
arch well developed, ventral rakers a series of raised tubercles.

Upper surfaces of head completely scaled; branchiostegal membranes with a small scale patch
on either side at level of isthmus; gular membranes and underside of lower jaw unscaled in WA
specimen (but in specimens from Madagascar, area with a broad median patch, and posterior end of
lower jaws scaled). Scales on body posterior of pectoral fin moderately deciduous; scales of dorsal
flanks bearing small, slender spinules in 8—9 near-parallel rows.

Light organ keyhole-shaped; anterior dermal window situated between posterior margin of pelvic
fin bases, connected to periproct by an isthmus of naked black skin; posterior dermal window
relatively large, immediately anterior to vent.

First dorsal fin high, second spinous ray with numerous serrations along leading edge; pelvic fins
broad, close together, outer ray only slightly produced; anal fin well developed along entire length.
Pelvic fin origin at vertical through posterior margin of opercle; pectoral fin origin just posterior to
pelvic fin origin; first dorsal fin origin set further back, posterior margin of base in line with origin
of anal fin.

Body light brown, speckled with small melanophores; dense clusters of melanophores on gular
membrane, on margin of branchiostegal membranes, and on pelvic fins; base of first dorsal, pectoral
and pelvic fins, opercle, and ventral abdominal region dark; sides of abdomen to level of pectoral fin
base with dark bluish pigmentation; periproct region black. Large dark patch over upper two-thirds
of first dorsal fin between 2nd and 8th elements; anal fin with distinct dark distal margin over anterior
15-20% of its length; no evidence of banding over trunk or anal fin.

SIZE. — To about 23 cm TL.

DISTRIBUTION. — From Namibia to Natal coast (South Africa), off Madagascar, and at 28°S off
west coast of Australia.

COMPARISONS AND REMARKS. — Lucigadus ori is very similar to L. nigromaculatus (McCul-
loch, 1907), which is widely distributed across temperate Australia and is also reported from New
Zealand (McCann and McKnight 1980), the southeastern Pacific (Iwamoto 1979), and the southeast-
ern Atlantic (Trunov and Konstantinov 1986). However, L. ori has a relatively small mouth (upper
jaw 31% HL cf. 41-47%), a more restricted first gill slit (18% HL cf. 25-31%), a greater preoral
length (23% HL cf. 17-21%), and a greater postorbital length (39% HL cf. 34-37%).

Our specimen of L. ori is most noticeably differentiated from L. acrolophus (Iwamoto and
Merrett, 1997) by a higher pelvic fin ray count (14-15 cf. 9-11), shorter upper jaw (31% HL cf.
35-41%), and more inner gill rakers (GR-I 13 cf. 9-11; GR-II 12 cf. 8-11). It lacks the banded
coloration over the body and anal fin of L. microlepis (Ginther, 1878) and differs from L. nigromar-
ginatus (Smith and Radcliffe, 1912) by its higher pelvic fin ray count (14-15 cf. 10-11) and several
proportional measurements including, among others, orbit diameter (38% of HL cf. 33-36%) and
barbel length (15% cf. 20-26%).

It is a bit curious to find a single representative of this predominantly southern African species
in Western Australia but none of the other two members of the genus (L. nigromaculatus and L.
microlepis) that have been recorded from Australia. The species has not been collected, so far as we
know, in other parts of the Indian Ocean. That only a single specimen was collected suggests that its

186 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 3

Ficure 36. Lucigadus ori (Smith). RUSI 4323 (20 cm TL) from off Natal, South Africa. (From Iwamoto 1986, fig. 93.32.)

occurrence in Australia is rare. The more cool-temperate species L. nigromaculatus appears to be
relatively common in waters off Tasmania, Victoria, and New South Wales, as well as off New
Zealand. It will probably be found along the south coast of Western Australia with more extensive
collecting.

Malacocephalus Gunther, 1862

See Iwamoto (1990) for generic diagnosis. Several nominal species, but probably only four or
five valid. One wide-ranging, worldwide species found off Australia.

Malacocephalus laevis (Lowe, 1843)
Rigas

Malacocephalus laevis: Ginther, 1862:397.
Macrurus (Malacocephalus) laevis: Ginther, 1887:148, pl. 39, fig. B).

DIAGNOSIS (after Iwamoto 1990).—Snout 26-31% HL; distance from orbit to angle of preopercle
41-47% HL. Premaxillary teeth in 2 distinct rows; 1 row of widely spaced canine teeth on mandible.
Dorsal fin spine smooth; first dorsal fin rays I1,9—13; pectoral fin rays 115-121; pelvic fin rays 8—9
(usually 9).

SPECIMENS EXAMINED. — WA: CSIRO H2573-18 (4, 56.5—82.4 mm HL, 298-450 mm TL) and
CSIRO H2573 (72.5 HL, 380 TL); off Shark Bay; 690-691 m; SS1/91/36.

COUNTS AND MEASUREMENTS (based on 5 WA spec.).—1D. II,10—12; P. 118; V. 8-9; total GR-I
(outer/inner) 6-9/10—13, GR-II 8—11/8—11; scales 1D. 15.5—18.5, midbase 1D. 13.5—16.5, 2D.
15.5—16.5.

IWAMOTO AND WILLIAMS: GRENADIERS OF WESTERN AUSTRALIA 187

FiGURE 37. Malacocephalus laevis (Lowe). CSIRO unregistered, from southern Australia.

Total length 298-450 mm; HL 56.5—82.4 mm. The following in percent of HL: postrostral 75—78;
snout 26-28; preoral 16-18; internasal 18-21; interorb. 26-32; orb. 32—37; suborb. 12—13; postorb.
42-45: orb.-preop. 41-43; up.jaw 46-48; barbel 16—20; gill slit 23-27; pre-A. 123-140; pre-vent
114-132; V.-A. 22-27; isth.-A. 55-62; body depth 79-88; depth at A. 75—87; 1D.-2D. 47-63; ht. 1D.
52-60; len. 46-59; len. V. 27-35; post. nostril 6-8; rictus len. 39-41.

DESCRIPTION. — Gomon, Glover, and Kuiter (1994) give a recent description of material
collected from southern Australia.

SIZE. — To about 60 cm TL.

DISTRIBUTION. — Widespread, probably cosmopolitan in warm waters of the Atlantic Ocean,
Indian Ocean, and Pacific Ocean. Reported off Australia from WA, NSW, and QLD, but probably
distributed across entire southern coast.

COMPARISONS AND REMARKS. — Malacocephalus laevis is the only member of the genus thus
far recorded from Australian waters. The genus comprises seven nominal species, although the close
similarity of M. hawaiiensis Gilbert, 1905 and M. luzonensis Gilbert and Hubbs, 1920 to M. laevis
casts some doubt on their validity. Small differences in scale counts between Atlantic and Indian
Ocean populations and fish from the Pacific have been noted (Iwamoto 1990), but there appear to be
no other reliable characters for differentiating the species. Malacocephalus nipponensis Gilbert and
Hubbs, 1916 apparently lacks the scale patch on the gular membrane present in (well-preserved)
specimens of M. laevis. The three remaining species, M. occidentalis Goode and Bean, 1885, M.
okamurai Iwamoto and Arai, 1987, and M. boretzi Sazonov, 1985, are distinct. A key to species was
provided by Iwamoto (1990:240, 241).

Mataeocephalus Berg, 1898

The genus includes five nominal species, but two or more undescribed ones are known, including
a species with six branchiostegal rays reported by Iwamoto and Merrett (1997) from New Caledonia
and now recorded from Australia. A revision of the genus is planned by Y. I. Sazonov (ZMMGU)
and Y. N. Shcherbachev (IOAN).

KEY TO SPECIES OF M4ATAEOCEPHALUS FROM WESTERN AUSTRALIA

la. Branchiostegal rays 7; underside of head naked; pelvic fin rays usually 8; anus closer to anal fin than to pelvic
AMISCHULON SI Wm seas gedteg ee (h8 Saat et EON OY TES Ee aE) ae ae ee tee tanec ts Comme M. acipenserinus

lb. Branchiostegal rays 6; underside of head mostly scaled; pelvic fin rays 7; anus closer to pelvic fin insertions than to
LGV NUL Me MmecnoREN on SRY PEM cinen ek SRR ene Ie seen ee er eee er aS ss EO MIEE ONS Gur o Bee Ss M. sp.

188 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 3

Mataeocephalus acipenserinus (Gilbert and Cramer, 1897)
Fig. 38

Coelocephalus acipenserinus Gilbert and Cramer, 1897:422, 423 (Hawaii).

Mataeocephalus acipenserinus: Berg, 1898:41 (replacement name for Coelocephalus, preoccupied in Coleop-
tera). Sazonov and Iwamoto, 1992:70, 71 (Sala y Gomez Ridge). Iwamoto and Merrett, 1997:533, 534, fig.
26 (New Caledonian region).

DIAGNOSIS. — Underside of head mostly naked. Serrations on second dorsal fin spine developed
and widely spaced. Jaw teeth in short bands. Anus at posterior end of broad periproct region situated
close to anal fin. Pectoral fin rays 119-125; pelvic fin rays 8.

SPECIMENS EXAMINED. — WA: CSIRO H2542-30 (3, 48.5-49.2 mm HL, 206-240 mm TL),
Exmouth Plateau; 854-868 m; SS1/91/3. CSIRO H2549-14 (43.6 HL, 178 TL); w. of NW Cape;
SS1/91/10; 650-685 m. CSIRO H2572-12 (3, 44.5-49.1 HL, 185-297 TL); w. of Dirk Hartog I.;
874-882 m; SS1/91/35. CSIRO H2580-08 (7, 39.2-46.0 HL, 170-190 TL) and CSIRO H2580-07
(44.2 HL, 175 TL); sw. of Shark Bay; 713—714 m; SS1/91/43.

COUNTS AND MEASUREMENTS (based on 12 spec.).—1D. II,7—9; P. 119-124; V. 8; total GR-I
(inner) 5—7, GR-II (outer/inner) 6—7 (usually 7)/6—8; scales 1D. 5—6, 2D. 4.5—-6.

Total length 170-245 mm; HL 39.2-49.2 mm. The following in % HL: postrostral 59-69; snout
36-41; preoral 30-41; internasal 20—24; interorb. 20—23; orb. 25—32; suborb. 13—16; postorb. 3 1—36;
orb.-preop. 29-34; up.jaw | 8—23; barbel 4-5; pre-A. 126-147; pre-vent 116—137; body depth 53-59;
depth at A. 41-57; 1D.-2D. 26-37; len. P. 40-46; len. V. 40-46; post. nostril 6-13; rictus len. 14-17.

DESCRIPTION. — Snout long, moderately depressed; suborbital ridge with a row of well-devel-
oped, coarsely spinulated scales; snout forming a broad, triangular leading edge with prominent bifid
terminal scute. Mouth small, inferior, extending approximately between levels of anterior margin and
midline of orbit. Upper jaw length less than orbit diameter.

Premaxillary teeth in short, broad, oval-shaped bands; mandibular teeth in crescent-shaped band.
Teeth long, fine; none enlarged.

Opercular openings relatively small; gill filaments well developed; gill rakers on first arch blunt,
tubercular, with a few small distal spines.

Serrations on second spinous ray of first dorsal fin poorly developed and widely spaced; second
dorsal fin poorly developed. Pectoral and pelvic fins of similar length; outer rays in both fins
moderately prolonged.

Body lacking prominent markings; overall coloration variable, ranging from pale to dusky.
Mouth region bluish in dusky specimens, cream colored in pale specimens. Upper jaw blackish
anteriorly, lower jaw dusky to dark; median nasal processes dusky to dark; buccal cavity dark;
posterior region of opercular cavity blue-black. Fin color variable pale to dark; pelvic bases dark.

SIZE. — To about 25 cm TL.

DISTRIBUTION. — Recorded in the Pacific from the Hawaiian Islands (type locality) and Sala y
Gomez Ridge (Sazonov and Iwamoto 1992), and believed to be widespread in Pacific and Indian
Oceans (Y. I. Sazonov, pers. comm. with AW, 1995). We took numerous specimens off WA between
20°S and 29°S, in 650-945 m depth.

COMPARISONS AND REMARKS. — Mataeocephalus specimens collected off WA were nearly
inseparable from M. acipenserinus based on comparison with type material and the accounts of Gilbert
and Cramer (1897), Gilbert (1905), Iwamoto (1990), and Sazonov and Iwamoto (1992). The only
notable difference was reduced development of the terminal scute on the snout in the WA specimens
compared to the type material. Of note was the high degree of variation in color in our specimens.
The pelvic and anal fins, branchiostegal membranes, and flanks ranged from darkly pigmented to pale
or virtually unpigmented. Some color variation may be due to variable abrasion of skin and fins in
the trawl, since M. acipenserinus has highly deciduous scales and rather delicate skin. This 1s borne

IWAMOTO AND WILLIAMS: GRENADIERS OF WESTERN AUSTRALIA 189

out by the absence of scale pockets in many light-colored fish. There were no differences in counts
or measurements between dark and lightly pigmented fish and no correlation with depth, latitude, or
station. Coloration differences were also noted by Gilbert (1905) in specimens from Hawaii.

Mataeocephalus sp.
Fig. 39

Mataeocephalus sp. Sazonov and Shcherbachev MS: Iwamoto and Merrett, 1997:534, 535, fig. 27 (New
Caledonian region).

DIAGNOSIS.— Branchiostegal rays 6; underside of head mostly scaled; pelvic rays 7; periproct
removed from anal fin, anus closer to pelvic fin insertions than to anal fin origin.

SPECIMEN EXAMINED.—- WA: CSIRO H2549-13 (1 spec.); w. of NW Cape; 650-685 m;
551/917 10:

SIZE— To at least 23 cm TL.

DISTRIBUTION.—-WA and New Caledonian region, 412—970 m.

COMPARISONS AND REMARKS.— This specimen represents an undescribed species that has been
recorded from the New Caledonian region by Iwamoto and Merrett (1997) and is known from other
areas in Australia (personal records, TI). Its six branchiostegal rays, as opposed to seven in all other
members of the genus, place it in a questionable position in the genus and obscures the differences
between the genera Mataeocephalus and Hyomacrurus Gilbert and Hubbs, 1920.

Mesobius Hubbs and Iwamoto, 1977

The two widely distributed species in the genus are known to be at least partially bathypelagic
(based on trawls fished in midwaters) and have peculiar, distinctive squamation on the head. The
scales there are elongate and aligned, each bearing |—3 rows of spinules that form low, sharp ridges.
For a complete diagnosis see the original description or Iwamoto (1979).

KEY TO SPECIES OF MESOBIUS

lawePelvictininays}o—7innerpillirakers.onisecondiarch) IZ-ll69 3 3s eee ee M. antipodum
lbweRelvicninmays)/—9sinneneillirakers.onisecondarchw O12) cea 2 fom ici cmemte cet ele tet eee M. berryi

Mesobius antipodum Hubbs and Iwamoto, 1977
Fig. 40a

Mesobius antipodum Hubbs and Iwamoto, 1977:245, 246 (New Zealand, e. of South I.; 995—1,110 m). Arai,
1979:286-289 (12 spec., New Zealand off Chatham Rise and Pukaki Rise; 860—1,050 m). Shcherbachev et
al., 1979:19-22 (Indian Ocean, Atlantic Ocean; comparison of M. berryvi and M. antipodum).

DIAGNOSIS. — Posttemporal region with differentiated squamation extending posteriorly of
vertical through origin of pectoral fin base. Outer margin of gill cover notably incised at subopercle.
Total inner rakers on first gill arch 12—15, on second arch 12-16.

SPECIMENS EXAMINED. — WA: CSIRO H3010-03 (2, 107-110 mm HL, 510-580 mm TL);
35°07'S, 115°O1'E, 945 m; Akebono Maru No. 3, shot 10, 23.12.1989, coll. A. Williams. CSIRO
H3008-02 (131.6 HL, 670+ TL); sw. of Albany; 35°25’S, 117°21'E, 842 m, Akebono Maru No. 3,
shot 8; coll. A. Williams; 22.12.1989.

COUNTS AND MEASUREMENTS (based on 3 spec.). — 1D. II,9—10; P. 113-114; V. 7; total GR-I
(outer/inner) 7—10/13, GR-II 13—14/13; scales 1D. 12-13, midbase 1D. 6—7, 2D. 9.5—10.5.

190 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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FiGuRE 38. Mataeocephalus acipenserinus (Gilbert and Cramer). CSIRO H5272-12 (CSIRO H2572-12 (45 mm HL, 187
mm TL) from off Shark Bay, WA, in 874-882 m. Drawn by Georgina L. Davis.

FIGURE 39. Mataeocephalus sp. (From Iwamoto and Merrett 1997, fig. 27.)

Total length 510-670 mm; HL 107.0—-131.6 mm. The following in percent of HL: postrostral
74-77; snout 30-31; preoral 18-21; internasal 26-29; interorb. 33-34; orb. 26-30; suborb. 14-16;
postorb. 48-49; orb.-preop.47—-49; up.jaw 44-47; gill slit 15—19; pre-A. 104-113; pre-vent 96-103;
V.-A. 15-19; isth.-A. 32-37; body depth 79-87; depth at A. 78-87; 1D.-2D. 58-61; ht. 1D. 47 (1
spec.); post. nostril 4 (1 spec.); rictus len. 34-38.

DESCRIPTION. — Adequate descriptions are provided in the original description and the more
recent accounts of Arai (1979), Shcherbachev et al. (1979), and Gomon et al. (1994).

SIZE. — To 67 cm TL.

DISTRIBUTION. — Widely distributed in the southern hemisphere: southern Atlantic Ocean off
South Africa, Indian Ocean, Southern Ocean off southern Australia, southern Pacific Ocean off New
Zealand.

COMPARISONS AND REMARKS. — The two species of Mesobius may be differentiated by counts
of pelvic fin rays and gill rakers: for M. antipodum and M. berryi, respectively, pelvic fin rays 6—7

IWAMOTO AND WILLIAMS: GRENADIERS OF WESTERN AUSTRALIA 19]

cf. 7-9, and inner rakers on second gill arch 12—16 vs. 10-12. Overall body size and development of
the head may also be useful for field identifications. The largest reported size of M. antipodum (67
cm TL) is considerably greater than the size attained by M. berryi (42 cm TL); M. antipodum has a
relatively well-developed and bulbous head, whereas the head of M. berryi is relatively slender.

Mesobius antipodum is possibly a pelagic species (see also comments under M. berryi), but thus
far most specimens have been taken by demersal trawls.

Mesobius berryi Hubbs and Iwamoto, 1977
Fig. 40b

DIAGNOSIS. — Posttemporal region with differentiated squamation not extending posteriorly of
vertical through origin of pectoral fin base. Outer margin of gill cover not notably incised at
subopercle. Total inner rakers on first gill arch 10—13; inner rakers on second arch 10—12.

SPECIMENS EXAMINED. — WA: CSIRO H2623-02 (76.0 mm HL, 375 mm TL); w. of Cape
Freycinet; 1,225—1,240 m; SS1/91/92.

COUNTS AND MEASUREMENTS (based on 1 WA spec.). — 1D. II,10; P. 113; V. 8—9; total GR-I
(outer/inner) 8/11, GR-II 10/10.

(UMTE A
MMMM MMM ize

FiGuRE 40. (a) Mesobius antipodum Hubbs and Iwamoto. CSIRO H3010-03, from south of Cape Leeuwin, WA, in 945 m.
(b) Mesobius berryi Hubbs and Iwamoto. CSIRO H2623-02 (76 mm HL, 375 mm TL) from off Perth, WA, in 1225-1240 m.
Drawn by Georgina L. Davis.

192 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 3

The following in percent of HL: postrostral 59; snout 20; preoral 12; internasal 17; interorb. 22;
orb. 24; suborb. 10; postorb. 37; orb.-preop. 37; up.jaw 37; gill slit 14; pre-A. 75; pre-vent 72; V.-A.
10; isth.-A. 20; body depth 63; depth at A. 58; 1D.-2D. 34; len. V. 23; post. nostril 5; rictus len. 39.

DESCRIPTION. — Adequate descriptions are provided in the original description and by Iwamoto
(1979).

SIZE. — To 41 cm TL.

DISTRIBUTION. — Indian Ocean, and central and northeastern Pacific Ocean.

COMPARISONS AND REMARKS. — Although our specimen and those of Shcherbachev et al.
(1979) were taken by demersal trawling, the species is thought to be primarily bathypelagic (Hubbs
and Iwamoto 1977). For comparison of Mesobius species, see remarks for M. antipodum.

Nezumia Jordan, 1904

This genus includes more than 40 species, some of which when more closely studied are likely
to be assigned to other genera. Shcherbachev et al. (1992) and Iwamoto and Sazonov (1994) recently
revised, respectively, the genera Kuronezumia and Kumba to include several species previously
considered as belonging to Nezumia. Many genera within this group of seven-branchiostegal-rayed
grenadiers are inadequately diagnosed, and the distinctions between genera are often hazy and based
on subjective or plesiomorphic characters. A thorough phylogenetic analysis of grenadiers with seven
branchiostegal rays is needed to resolve the proper status of the included taxa.

The genus Nezumia is represented in waters off Western Australia by seven species. Nezumia
namatahi McCann and McKnight, 1980 and Nezumia coheni lwamoto and Merrett, 1997 are included
in the key to species even though they have yet to be recorded from Western Australia. The former
is acommon species off New Zealand and the Australian east coast, but its farthest westward capture
is the eastern part of South Australia. The latter species is common in New South Wales. The
likelihood of the known ranges of these species extending into Western Australian waters and the
possibility of specimens of the two species being confused with other species known from Western
Australia prompt their inclusion in the key.

KEY TO AUSTRALIAN SPECIES OF NEZUMIA

layRelvicitinirays sion 9 si 9snodarke bandiencirclingytrunke esis ey eine ee ee) enn ene eee 2
lib} Pelvic finirays 9=l'8 i195 trunkiencircled bya broadidark band) 5 5 5 9.52). 4.5 24 2] one ae 3
2a. Scale rows below origin of 1D. 10-13, below 2D. 7.5—11; pores on underside of head prominent; lip margins black-
ishvantenronlyapalestowardSyjaweangle ss. <5 <: mcets vate e calen cutciven tc cm CMR necy ton tet cr teu nlm ee mr amo N. spinosa
2b. Scale rows below origin of 1D. 7—7.5, below 2D. 6.5—7.5; pores on underside of head inconspicuous; lip margins
blackitoijaw an@les-as <, moehenaite oF tpty waive cy tee Gi eee eels) Rech cole oR mnt rotor etree me N. merretti n.sp.
Savy Peélvictinirays T4 a7) 2 a Boies. ed, dia ietn eee Ree ccc ete Cae et N. propinqua
3b: Pelvic finirays:; 912. ve a oe ee ee eae 2S a as Be ee oe ee ne ee 4

4a. Tip of tail posteriorly from about 90th anal fin ray completely pale; scales below origin of second dorsal fin 10—11
SCR TE Ee Oe eT eee ee aoe een ee ras Chih Wicts SMe eof tehGro-o < N. leucoura n.sp.

4b. Tail lacking prominent pale tip; scales below origin of second dorsal fin5.5-9.5 .................... 5)
5a. Underside of snout completely scaled; orbit to preopercle distance 38-44% of HL .......... N. wularnia n.sp.
5b. Underside of snout broadly naked; orbit to preopercle distance 33-44% of HL ..................... 6
6a. ADW about on line with pelvic fin insertions; dorsum rather uniformly pigmented, although area above abdomen
oftentslightly darker...) soc i. 5.5 ine ele see) ee nn eects) Cour eee anton een 7
6b. ADW well posterior to line connecting pelvic fin insertions; dorsal aspects of trunk more darkly pigmented than
comparable areas on tail (and usually nape), forming complete dark band around trunk ................ 8
7a. Total GR-I (outer/inner) 9—12/11—13; orbit to preopercle distance 33-38% of HL... .......... N. soela n.sp.
7b. Total GR-I (outer/inner) 6—9/9—1 1; orbit to preopercle distance 38-44% of HL ................ N. coheni
8a. Pelvic fin rays 9-10; sensory pores on underside of head prominent ..................... N. namatahi

8b. Pelvic fin rays 11-12; sensory pores on underside of headsmall....................-. N. kapala n.sp.

IWAMOTO AND WILLIAMS: GRENADIERS OF WESTERN AUSTRALIA 193

Nezumia kapala new species
Fig. 41

No literature applies to this species.

DIAGNOSIS. — Pelvic fin rays | 1—12; height first dorsal fin 86—109% of head length; underside
of snout naked posteriorly to below about mid-orbit, pores of lateralis system small, inconspicuous;
body scales covered with slender, conical to lanceolate spinules in slightly convergent rows; 7.5—9.5
scale rows below origin of second dorsal fin, 33—38 lateral line scales over distance equal to predorsal
length; trunk encircled by broad dark band, sometimes poorly defined; first dorsal fin all black to
black anteriorly and dusky posteriorly; anterior dermal window of light organ about on or well
posterior to line connecting pelvic fin insertions; distance orbit to angle of preopercle 34-39% of head
length; length upper jaw 26-33%; gill rakers on inner side of first arch 8-10.

SPECIMENS EXAMINED. — NSW: HOLOTYPE: AMS 1.24993-008 (52.6 mm HL, 335 mm TL);
off Broken Bay; 1,043—1,070 m; K84-16-14. PARATYPES: AMS I.24059-006 (4 of 6 spec., 48.1—5S4.2
HL, 318-358+ TL); off Broken Bay; 933-969 m; K8003-09-02. AMS 1.24993-008 (57.1 HL, 367+
TL); same data as for holotype. CAS 200228 (formerly AMS 1.24150-006) (51.1 HL, 343 TL); off
Broken Bay; 988—1,020 m; K83-13-01. AMS 1.25265-002 (4, 45.5—60.0 HL, 277+—402+ TL); se. of
Broken Bay; 1,024—1,245 m; K84-19-04. AMS 1.21724-005 (52.8 HL, 320 TL); e. of Broken Bay;
1,006 m; K79-20-15. AMS 1.24060 (45.2 HL, 290+ TL); off Sydney; 942-960 m; K83-09-01. AMS
1.24057-001 (4, 39.4-61.7 HL, 212+—394+ TL); se. of Kiama; 951-978 m; K83-09-04. TAS: CSIRO
H2865-04 (43.7 HL, 255 TL); Pedra Branca area; 44°22.6'S, 147°08.6’E; 1,060—1,170 m; Belinda;
12.11.1992. WA: NMV A6195 (56.5 HL, 366 TL); 34°35.5’S, 121°14’S; 1,015—1,030 m; Saxon
Progress; 17.VIII.1988. CSIRO H3008-11 (48.2 HL, 320 TL); sw. of Albany; 35°25’S, 117°21’E;
842 m; Akebono Maru No. 3, shot 8; 22.XII.1989. CSIRO H.2584-21 (30.9 HL, 148+ TL); w. of
Shoal Point; 853 m; SS1/91/49.

COUNTS AND MEASUREMENTS (holotype first, followed by range in 24 paratypes). — 1D. II,9,
I1,8—10 (11); P. 120/120, 118-122; total GR-I (outer/inner) 9/10, 6—-8/8—10, GR-II 9/11, 8—-9/8—11;
scales 1D. 10, 8-12, midbase 1D. 7.5, 5.5—7.5 8.5, 2D. 9.5, 7.59.5, lat.line 36, 33-38.

Total length 335 mm, 148+—410+ mm; HL 52.6 mm, 30.9-67.0 mm. The following in percent
of HL: snout 33, 27—34; preoral 27, 23-31 (33); internasal 25, 21—25 (27); interorb. 22, 18—25; orb.
30, 29-33 (35); suborb. 15, 14-17; postorb. 41, 40-45; orb.-preop. 35, 34-39; up.jaw 27, 26-31 (33);
barbel 14, 11—20; gill slit 12, 10-14; pre-A. 150, 146-167; V.-A. 44, 35-52; body depth 77, 69-92;
1D.-2D. 54, 28-60; ht. 1D. 87, 86-109; len. 1P. 55, 52-64; len. V. 51, 45-67; post. nostril 8, 4-9.

DESCRIPTION (of holotype, with comments on paratypes in parentheses). — General features of
fish seen in Figure 41. A slender fish, head more than 6 times, greatest body depth more than 7 times,
into total length. Snout conically pointed in lateral view; bluntly angular in dorsal view; about equal
to orbit diameter; tip armed with a large, broad, bifid scutelike scale; lateral angles also studded with
stout spiny scales. Mouth subinferior, jaws short, less than one-third head length; posterior end of
maxilla falls about under mid-orbit. Chin barbel well developed but short, least width about half orbit
diameter. Suborbital shelf formed of 2 rows of stoutly modified scales; the lower row forming a rather
sharp angular ridge. A broad median swath of naked skin under snout, but almost all of suborbital,
preopercle, and lower jaw scaled. Exposed posterior tip of subopercle scaled. Preopercle broadly
rounded, chord of posterior margin vertical.

Exposed fields of body scales covered with sharp, slender, conical to narrowly lanceolate spinules
aligned in 11-15 slightly convergent rows (number of rows size dependent).

Teeth in broad tapered bands in both jaws; upper jaw teeth somewhat larger than those of lower
jaw, outer series enlarged.

194 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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FiGuRE 41. Nezumia kapala n.sp. Holotype, AMS 1.24993-008 (335 mm TL), from off Broken Bay, NSW, in 1,043—1,070
m. Fins and scales partially reconstructed. Scale bar represents 25 mm.

First dorsal fin high, less than head length in holotype (more than head length in several
paratypes); second spinous ray heavy, slightly recurved, leading edge armed with about 22 small sharp
denticles. Second dorsal fin rudimentary over anterior half or so, rather low over posterior half. Outer
pelvic fin ray slightly prolonged, extending posteriorly to about 4th or Sth anal ray. Pectoral and pelvic
origins about on same vertical; origin of first dorsal behind that vertical. Anal fin well developed
throughout.

Periproct region moderately developed; anus about midway between pelvic insertions and anal
origin; ADW small, lying well posterior to line connecting pelvic insertions (in one paratype, AMS
1.24060-023, ADW about at level of line).

Ground color pale to dark brown overall, with entire trunk region encircled by broad dark band.
Ventral aspects of chest, opercle, and all of abdomen (to above 12th or so anal ray) black to bluish
black; somewhat paler on dorsum and nape but still distinctly darker than head and posteriorly on
tail. First dorsal fin all black (in most specimens, somewhat paler posteriorly in some). Pectoral,
pelvic, and anteriormost rays of anal fins black; anal fin dark dusky posteriorly. Gular and bran-
chiostegal membranes black; mouth and lips dark; gill chamber black; gill arches and rakers dark;
barbel pale.

SIZE. — To about 41 cm TL.

DISTRIBUTION. — Australia, from NSW off Norah Head (33°32'S) south to TAS, and west to
WA, west of Shoal Point (28°S) in 842—1,243 m.

ETYMOLOGY. — Named for the former New South Wales Fisheries research vessel, the Kapala,
on which the holotype and many other grenadiers from New South Wales were collected.

COMPARISONS AND REMARKS. — Nezumia kapala is very similar in overall morphology to N.
namatahi, a species formerly known only from off New Zealand but known from our records from
off the southeast coast of Australia. The two species are sympatric, with individuals of both species
having been captured together (and confused) in numerous trawls. The most readily apparent
difference between the two lies in the pelvic fin ray count: 11 or 12 in the new species, 9 or 10 in N.

IWAMOTO AND WILLIAMS: GRENADIERS OF WESTERN AUSTRALIA 195

namatahi (see Table 5). Other differences include the broadly lanceolate to shield-shaped spinules in
N. namatahi and the large pores on the underside of head, which are absent or small in the NV. kapala.

Nezumia kapala is also closely similar to N. soela, but the two can be distinguished by the latter
having a more anterior position of the periproct and ADW, absence of a completely encircling dark
band on trunk, slightly higher gill raker counts on the first arch (9-12 cf. 6-8 in outer series, 11—13
cf. 8-10 in inner series), and slightly shorter postorbital length (37-40% HL cf. 40-45%). Another
species from NSW and New Caledonia, NV. coheni Iwamoto and Merrett, 1997, differs from N. kapala
in lacking a complete, dark trunk girdle, ADW at or slightly anterior to line connecting pelvic
insertions, and a somewhat greater orbit-preopercle distance (38-44% HL cf. 34 40%).

Nezumia leucoura new species
Fig. 42a

DIAGNOSIS. — Pelvic fin rays | |—12; height first dorsal less than head length; underside of head
including lower jaw naked, without prominent sensory pores; scales covered with tiny spinules in
parallel to slightly convergent rows; 10—11 scale rows below origin of second dorsal fin, about 50
lateral line scale rows from anterior origin over distance equal to predorsal length; dark color of trunk
confined to abdomen, not extending onto dorsum; first dorsal fin pale with blackish tip; anterior
dermal window of light organ small, situated between pelvic fin bases; tip of tail distinctly pale over
posterior 1/4—1/Sth of fin, from about 90th (88th—1 10th) anal ray.

SPECIMENS EXAMINED. — WA: HOLOTYPE: AMS 1.31181-012 (38.4 mm HL, 208+ mm TL);
sw. of Shoal Point; 853-854 m; SS1/91/49. PARATYPES (32 spec.): AMS I.131181-012 (3, 33.0-42.0
HL, 163+—223+ TL), AMS 1I.31170-005 (4, 18.7-26.8 HL, 103+—170 TL) and CAS 200230 (35.3
HL, 210+ TL); same data as for holotype. NMV A9543 (7, 32.6-38.0 HL, 155-216 TL); CSIRO
2573-12 a( 7 Oe RL), 2573-13 (6, 120-170, TL); w. of Steep Pt; 691 mz SS1/91/36: ‘CSIRO
H2579-03 (190 TL); sw. of Shark Bay; 688 m; SS1/91/42. CSIRO H2596-05 (215 TL), H2596-06
(5, 200-220 TL); wnw. of Green Head; 760 m; SS1/91/62.

COUNTS AND MEASUREMENTS (holotype first, followed by range in 13 paratypes). — 1D. II,13,
II,11—-13; P. i21, 120423; V. 12, 11-12; total GR-I (outer/inner) 12/10, 8—12/9-11, GR-II 9/10,
810/911; scales 1D. 16, 14-15, midbase 1D. ? , 8.5—10.5, 2D. ? , 10-11, lat.line 51, 45—S3.

Total length 208 mm, 103+—223+ mm; HL 38.4 mm, 18.7-42.0 mm. The following in percent
of HL: snout 27, 25—32; preoral 22, 17-24; internasal 23, 21—28; interorb. 21, 20—25; orb. 34, 32-43;
suborb. 14, 14—16; postorb. 42, 40-45; orb.-preop. 38, 3340; up.jaw 36, 33-39; barbel 21, 19-26;
gill slit 14, 13-20; pre-A. 156, 141-158; V.-A. 44, 36-52; body depth 94, 82-97; 1D.-2D. 40, 36-76;
ht. 1D. 81, 80-97; len. P. 57, 52—58; len. V. 57, 52-69; post. nostril 3, 3-6.

DESCRIPTION. — General features of holotype shown in Figure 42a. Snout short, vertical profile
relatively steep, length less than diameter of large orbits, protruding little beyond mouth. Jaws
relatively large for genus, upper jaw usually more than one-third head length, extending posteriorly
to below mid-orbit or slightly beyond. Suborbital shelf stout, formed of two series of modified scales,
the lower series with notably stout and coarse scales, the upper series of small and rather weak scales;
ventral edge of lower series forming a sharp ridge. Underside of snout, suborbital, and mandibular
rami naked; sensory pores of head rather well developed. Preopercle broadly rounded, posterior
margin almost vertical. Subopercle scaled and exposed only at posterior end. Chin barbel slender but
well developed, length about equal to interorbital width. Gill arches restricted by membranes across
lower extent.

Body scales small, rather deciduous; exposed fields covered with tiny conical spinules in 1 1-14
parallel to slightly convergent rows in holotype.

Teeth small, in moderately wide bands in both jaws; outer series in upper jaw slightly enlarged.

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES

196

Volume 51, No. 3

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IWAMOTO AND WILLIAMS: GRENADIERS OF WESTERN AUSTRALIA 197

Height first dorsal fin less than head length, leading edge of second spinous ray armed with small,
widely spaced denticulations; second dorsal fin rays rudimentary almost entire length of fin. Pectoral
and pelvic fins in advance of first dorsal fin; pectoral fin slightly more than half head length; pelvic
fin short, prolonged outer ray scarcely extends to origin of anal fin in holotype, to as far as 8th anal
ray in paratypes.

Periproct region large, well removed from anal fin, close between pelvic fins; anterior dermal
window small, between pelvic fin bases and anterior to line connecting insertions of fins. Pyloric
caeca of four paratypes short (slightly longer than least suborbital width), mostly branched into pairs,
24, 28, 30, and 31 in number.

Ground color somewhat grayish to tawny, swarthy on underside of head but whitish under snout,
blackish over operculum and ventrally around chest, pelvic fins, and anus; bluish over abdomen; no
broad band of dark pigment encircling trunk; tip of tail posteriorly from about 90th anal ray (88th to
110th in paratypes) pale, lacking normal pigmentation of tail. First dorsal fin black on membrane
between second spinous ray and first branched ray, the tips of next few rays black; pectoral fin dusky
at base, pale over most of rays, lightly dusky towards tip, narrow dorsal edge dark; pelvic fins mostly
black, but outer ray and distal tips paler; anal fin pale or faintly speckled anteriorly, becoming dusky
posteriorly. A broad horizontal streak from upper lip onto membrane between premaxilla and maxilla.
Chin barbel pale. Mouth lining dark except pale at angles of jaw; gums pale. Gill rakers and arches
dark.

SIZE. — To about 23 cm TL.

DISTRIBUTION. — Known only from WA, from about 26°S to about 30°S, in 688-854 m.

ETYMOLOGY. — From the Greek, /eukos, white, and oura, tail.

COMPARISONS AND REMARKS. — This small species appears to be confined to a short stretch of
continental slope off Western Australia. In its morphology it is a fairly typical member of the genus,
especially those found in shallow upper-slope waters of tropical seas. It is unique, however, in having
a pale tail tip, which distinguishes it from all other known species of Nezumia. It is most likely to be
confused with N. propinqua and N. evides, with which it shares a small adult size, black-tipped first
dorsal fin, relatively high pelvic fin ray count, short snout, small scales covered with short, conical
spinules, and the absence of a dark girdle around the trunk. The pale tail and counts of pelvic rays
and scale rows below the first dorsal origin, however, readily distinguish N. Jeucoura from N.
propinqua and N. evides.

Nezumia merretti new species
Fig. 42b

Nezumia sp. B: Williams et al., 1996:149 (in part; WA list).

DIAGNOSIS. — Pelvic fin rays 9; underside of head almost completely naked, pocketed with
prominent pores; body scales covered with moderately long, needlelike spinules in mostly parallel
(to slightly convergent) rows, spinules overlap posterior scale margin slightly; 6.5—7.5 scale rows
below origin of second dorsal fin; lower jaw teeth in broad, short band; abdomen light bluish, dorsally
on trunk dirty straw-colored or tawny; first dorsal fin dusky overall; anterior dermal window of light
organ between pelvic fin bases.

SPECIMENS EXAMINED. — WA: HOLOTYPE: CSIRO H2584-20 (54.2 mm HL, 245+ mm TL):
w. of Shoal Point; 853 m; SS1/91/49. QLD: PARATYPE: CSIRO H1966-01 (39.9 HL, 202+ TL);
Marian Plateau, e. of Townsville; 879-886 m; SO6/85/36.

COUNTS AND MEASUREMENTS (holotype first, followed by paratype). — 1D. II,10, 11,10; P.
122/121, 122/121; total GR-I (outer/inner) 10/12, 11/12; total GR-II 11, 11/13, 11/12; scales 1D. 7.5,
7.0, midbase 1D. 5.5, 6.5, 2D. 7.5, 6.5, lat.line 35, 39.

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198

Volume 51, No. 3

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IWAMOTO AND WILLIAMS: GRENADIERS OF WESTERN AUSTRALIA 199

The following in percent of HL: snout 33, 36; preoral 24, 31; internasal 19, —; interorb. 21, —;
orb. 30, 30; suborb. 13, 13; postorb. 40, 38; orb.-preop. 36, 33; up.jaw 31, 28; barbel 15, 8; gill slit
16, 14; pre-A. 157, 145; V.-A. 41, 30; body depth 81, 73; 1D.-2D. 39, 30; len. 1P. 54, 53; len. V. 76,
55; post. nostril 7, 11.

DESCRIPTION. — General features shown in Figure 42b. Body and head laterally compressed;
dorsal profile rises smoothly from snout tip to first dorsal fin. Snout broadly triangular in dorsal view,
tipped with large, blunt bifid scute, tubercular scales at lateral angles not prominent. Suborbital shelf
composed of two rows of stout, coarsely spinulated, modified scales. Underside of snout, most of
suborbital, and mandible naked; lower margin of preopercle narrowly naked; interopercle scaled at
exposed posterior end; pores on underside of head relatively large. Upper jaw extends to below
posterior one-third of orbit. Preopercle broadly rounded, although somewhat angular at posteroventral
corner.

Body scales relatively large, densely covered with small, needlelike spinules aligned in 16-18
parallel to slightly convergent rows in holotype (fewer rows in smaller paratype). Spinules reclined
at about 35—45° angle, those along posterior margin of scale slightly overlapping edge.

Teeth in both jaws in broad bands, lower jaw band broader, shorter, and more abruptly tapered
laterally. Outer teeth series in upper jaw slightly enlarged; lower jaw teeth uniformly small.

Tips of first dorsal fin rays in holotype and paratype broken off; leading edge of second spinous
ray armed with small, widely spaced denticles. Most pelvic fin rays fall short of, but outer ray
prolonged well beyond, anal fin origin.

Anterior dermal window of light organ prominently developed, situated between pelvic fin bases.
Periproct region large, far removed from anal fin origin.

Color in alcohol dirty-straw overall, abdomen light bluish, but blackish in vicinity of periproct
and ADW. Gill cover and gill membranes blackish. Lips and membrane connecting maxilla and
premaxilla blackish; gill and oral cavities dark gray to blackish. First dorsal fin dusky overall; pectoral
fin light dusky with black basal segment; pelvic fin black; anal fin dark dusky anteriorly, paler
posteriorly. Chin barbel pale.

SIZE. — To about 25 cm TL.

DISTRIBUTION. — Known only from two specimens from WA and QLD, in 853—860 m.

ETYMOLOGY. — Named in honor of our friend and colleague, Nigel R. Merrett, in recognition
of his contributions to deep-sea biology.

COMPARISONS AND REMARKS. — Nezumia merretti 1s closely similar in most features to Nezu-
mia spinosa. They can be separated by the larger scales of the new species, color of the first dorsal
(overall dusky in the new species, blackish anteriorly in N. spinosa), spinules on body scales (shorter
and do not overlap posterior margin as extensively and in more tightly parallel rows in the new
species), and head pores (more prominent in N. spinosa). Characters given in the key readily separate
the new species from other congeners from WA. Nezumia holocentra (Gilbert and Cramer, 1897) and
N. burragei (Gilbert, 1905) from Hawaiian waters also share many characters with the new species
and N. spinosa, including similar teeth bands, scale spinulation, terminal snout scute, naked underside
of head, and high first dorsal fin. The two Hawaiian species, however, have a shorter, blunter snout;
in N. holocentra the barbel is shorter, and in N. burragei the scales are smaller (about 15 below 1D.,
13 below 2D., 46 lateral line scales over predorsal distance).

Nezumia propinqua (Gilbert and Cramer, 1897)
Fig. 43a

Macrourus propinquus Gilbert and Cramer, 1897:424, 425, pl. XLII, fig. 2 (Hawaii).
Nezumia propinqua: Sazonov and Iwamoto, 1992:72—74, figs. 27, 28a (Sala y Gomez Ridge; extended
synonymy). Iwamoto and Merrett, 1997:541, 542, fig, 29a (New Caledonia region).

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DIAGNOSIS. — Pelvic fin rays 13—17; height first dorsal fin 97-1 13% of head length; underside
of snout, suborbital, and lower jaw naked; sensory pores in naked areas small but readily discernible;
body scales covered with needlelike spinules in as many as 10—12 parallel to slightly convergent rows;
8.5—10 scale rows below origin of second dorsal fin, 36—42 lateral line scales over distance equal to
predorsal length; no dark band encircling trunk; distal 1/3 to 3/4 of first dorsal black; anterior dermal
window of light organ slightly in advance of line connecting pelvic fin insertions.

SPECIMENS EXAMINED. — WA: CSIRO H2541-14 (31.4mm HL, 160+ mm TL); Exmouth
Plateau; 914 m; SS1/91/02. CSIRO H2549-17 (2, 30.0-31.5 HL, 160+—190 TL); w. of NW Cape;
650-685 m; SS1/91/10. CSIRO H2553-14 (2, 2832.8 HL, 120+—173+ TL); w. of Pt. Cloates; 910
m; SS1/91/14. CSIRO H2563-02 (7, 21.5—19.6 HL, 94+—170 TL); w. of Quobba Pt.; 901 m;
SS1/91/25. CSIRO H2572-10 (2, 29.6—32.7 HL, 158+—164+ TL); w. of Dirk Hartog I.; 874-882 m;
SS1/91/35. ZMMSU uncat. (27.6—27.7 HL, 150-160 TL); off Cape Cuvier; 23°57.9’S, 112°14.2’E;
831-834 m; Vityaz’ sta. 4564; 1.XII.1959.

COUNTS AND MEASUREMENTS (14 WA specimens). — 1D. II,10—12; P. 119-122; total GR-I
(outer/inner) 8—10/8—10, GR-II 7—9/9—10; scales 1D. (9.5) 11-13, midbase 1D. 6.5—8.5; caeca 21, 28
(2 spec.).

Total length 128+—190 mm; HL 21.3—32.8 mm. The following in percent of HL: snout 29-34;
preoral 18—24; internasal 19-26; interorb. 20-25; orb. 30-34; suborb. 14—18; postorb. 41-45;
orb.-preop. 30-39; up.jaw 31-35; barbel 15—23; gill-slit 13-17; pre-A. 134-166; V.-A. 38-61;
anus-A. 13—26; body depth 79—98; 1D.-2D. 23-40; len. 1P. 54-67; len. V. 76—105; nostril 5.4—10.0.

DESCRIPTION. — General features of fish seen in Figure 43a. Snout short, blunt, length about
equal to greatest orbit diameter, tipped with small but prominent, usually bifid, tubercular scales.
Mouth rather short, about one-third of head length; upper jaw extends to below midorbit. Suborbital
shelf formed by 2 rows of stoutly modified scales, ventral row larger with relatively sharp ventral
edge. Underside of snout, suborbital, ventral edge of preopercle, and mandibular rami naked; pores
of sensory system small but readily discernible. Preopercle broadly rounded, posterior margin almost
vertical. Interopercle exposed only along posteroventral margin. Chin barbel well developed, rather
stout throughout, not forming filamentous tip. Gill arches restricted ventrally by narrow fold of skin.

Body scales small, exposed fields covered with slender, needlelike spinules arranged in 10—12
parallel to slightly convergent rows in largest specimens examined. Spinules greatly reclined on body
scales, posteriormost ones extend well beyond posterior margin of exposed field.

Teeth small in both jaws, arrayed in short, narrow, tapered bands; those in lower jaw in bands 3
or 4 teeth wide; in upper jaw outer series slightly enlarged.

Height first dorsal fin about equal to or more than head length; second spinous ray armed along
leading edge with 6—12 wide-spaced, slender spikes. Outer pelvic fin ray thickened and elongated,
extending well beyond anal fin origin to base of 10th to 16th anal ray in some specimens. Pelvic fin
origin below opening of operculum, well anterior to origin of pectoral fin, which is anterior to that of
first dorsal fin.

Periproct region large, situated within middle one-third of distance between pelvic fin insertions
and anal fin origin, usually somewhat closer to pelvic fin insertions. Anterior dermal window of light
organ small, often inconspicuous between pelvic fin bases, well removed from periproct. Pyloric
caeca of 2 specimens well developed, rather thick, some bifidly branched from near base.

Ground color dark brown to swarthy overall, with abdominal region somewhat violet; gill cover,
gill membranes, and region around periproct black. All of head, including snout and underside,
swarthy. First dorsal fin blackish over distal 2/3 to 3/4 but paler near base. Pectoral fin dusky; pelvic
fin black; anal fin blackish anteriorly to dark dusky posteriorly. Proximal 1/2 of barbel dark; distal
half pale. Oral and gill cavity black, although latter ventrally pale; gill arches and rakers dark. Lips
blackish; skin connecting maxillary and premaxillary dark except near jaw angle.

SIZE. — To about 19 cm TL.

IWAMOTO AND WILLIAMS: GRENADIERS OF WESTERN AUSTRALIA 201

DISTRIBUTION. — Hawaii, southeastern Pacific (Sala y Gomez Ridge), Kyushu-Palau Ridge,
New Caledonia, Western Australia, and across Indian Ocean to southern Africa, in 523-914 m.

COMPARISONS AND REMARKS. — Nezumia propinqua shares in common with N. evides Gilbert
and Hubbs, 1920, and N. condylura Jordan and Gilbert, 1904, an unusually high (13—19) pelvic fin
ray count. The upper range of these counts, from 16 to 19, is higher than in any other species of
Macrouridae. The three species also share in common a relatively blunt snout, naked underside of
head, black-tipped first dorsal fin, and similar scale spinulation (needlelike, in 4—12 parallel to slightly
convergent rows). The species in this clade have not been adequately characterized, although
representatives have been recorded from the Indian Ocean, throughout the tropical western Pacific,
Hawaii, and the southeastern Pacific. Most characters used to distinguish the three species show
overlap, leading to the question of whether they are valid species. A more thorough study of this clade
using specimens from the entire distributional range will be necessary to resolve this question.

Nezumia soela new species
Fig. 44b

Nezumia sp. C: Williams et al., 1996:1949 (WA list).

DIAGNOSIS. — Pelvic fin rays 10—11; first dorsal fin with 10 or 11 segmented rays (rarely 12);
underside of snout with broad naked median swath, pores of lateralis system small, inconspicuous;
body scales covered with narrow, lanceolate spinules in slightly convergent rows; 7—9.5 scale rows
below origin of second dorsal fin, 36-41 lateral line scales over distance equal to predorsal length;
trunk lacking prominent broad dark girdle; first dorsal fin all black; anterior dermal window of light
organ more or less on transverse line connecting pelvic fin insertions; upper jaws 29-33% of HL;
orbit to preopercle distance 33-38% of HL; inner gill rakers on first arch 10-13.

SPECIMENS EXAMINED. — WA: HOLOTYPE: CSIRO H3017-04 (65.3 mm HL, 350+ mm TL);
nw. of Cape Leeuwin; 34°10’S, 114°16'E; 1,030 m; Akebono Maru No. 3, shot 17; coll. A. Williams;
24.XII.1989. PARATYPES (17 spec.): CSIRO H2551-17 (46.6 HL, 250+ TL); w. of NW Cape; 1,500+
m; SS1/91/12. CSIRO H3002-04 (54.5 HL, 290+ TL); se. of Albany; 35°23.5’S, 118°27’E; 1,030 m;
Akebono Maru No. 3, shot 2; coll. A. Williams; 21.XII.1989. CSIRO T292 (51.1 HL, 293+ TL),
CSIRO 1297 (53.0 HL, 388+ TL); 1299 61.4 HL, 285+ TL); GAB, s: of Eucla; 33°27'S; 128°36'E;
1,027—1,044 m; Margaret Philippa; coll. K. Evans; 24.X1.1984. AMS 1.18712.-014 (51.6 HL, 290+
TL); GAB, s. of Maduras; 33°49’S, 127°00.9'E; 1,080—-1,100 m; Dmitry Mendeleev, coll. J. R.
Paxton; 28.1.1976. SA: CSIRO H2877-06 (50.9 HL, 290+ TL); 33°59’S, 131°07’E; 1,062 m; 1992.
TAS: CSIRO H1579-48 (5 of 19 spec., 27.9-55.1 HL, 124+—296 TL); w. coast; 830 m; SO3/86/37.
CSIRO T831 (56.6 HL, 315+ TL); sw. of King I.; 40°33’S, 143°18’E; 1,250—-1,290 m; Margaret
Philippa; coll. K. Evans; 18.X.1983. CSIRO T358 (45.0 HL, 240+ TL); sw. of King I.; 40°33’S,
143°18'E; 1,250—1,290 m; Margaret Philippa; coll. K. Evans; 18.X.1983.

COUNTS AND MEASUREMENTS (holotype first, followed by range in 19 paratypes). — 1D. II,12,
II,10—-12; P. 120/120, 118-422; total GR-I (outer/inner) 9/13, 9-12/11—13, GR-II 11/12, 9—11/10-13;
scales 1D. 10, 8-11, midbase 1D. 7, 5.5—7.5(8.5), 2D. 8.5, 7.0-9.5, lat.line 40, 36-41.

Total length 350+ mm, 148+—410+ mm; HL 65.3 mm, 30.0-65.3 mm. The following in percent
of HL: snout 33, 30-36; preoral 25, 24-31; internasal 22, 21—26; interorb. 21, 20—26; orb. 33, 31—36:
suborb. 15, 14-17; postorb. 39, 37-40; orb.-preop. 38, 33-38; up.jaw 32, 29-33 ; barbel 17,
11—17(21); gill slit 14, 11-16; pre-A. 156, 137-159; V.-A. 47, 31-47; body depth 84, 63-81; 1D.-2D.
49, 29-62; ht. 1D. 93, 75—102; len. 1P. 51, 44-61; len. V. 46, 46-66; post. nostril 8, 6-12.

DESCRIPTION. — General features of fish seen in Figure 44b. Head 5—6.5 times into total length,
greatest body depth more than six times into total length. Snout conically pointed in lateral view;

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angular in dorsal view; about equal to orbit diameter; tip armed with two large, closely adjoined,
tubercular scales; lateral angles also studded with stout spiny scales. Mouth subinferior, upper jaws
about one-third or less of head length; posterior end of maxilla falls about under midorbit or slightly
beyond. Chin barbel short, length less than half orbit diameter. Lower row of modified scales on
suborbital shelf forming a rather sharp angular ridge. Preopercle broadly rounded, chord of posterior
margin inclined slightly forward.

Exposed fields of body scales covered with sharp, slender, lanceolate spinules aligned in 10-17
convergent rows (number of rows size dependent).

Teeth in broad tapered bands in both jaws; upper jaw teeth larger than those of lower jaw, outer
series slender, slightly enlarged.

First dorsal fin high, less than head length in holotype, but more than head in a few paratypes;
second spinous ray heavy, slightly recurved, leading edge armed with about 33 small, low, spaced,
non-overlapping denticles. Outer pelvic fin ray slightly produced into hair-fine filament, extending
posteriorly to just beyond anal fin origin.

Periproct region moderately developed; ADW small.

Ground color medium brown, with trunk dorsally slightly darker but without a prominent dark
encircling band (in contrast to N. kapala and N. namatahi). Ventral aspects of chest, all of abdomen
(to above 10th or so anal ray), and opercle black to bluish black; medium brown on dorsum and nape.
First dorsal, pectoral, pelvic, and anteriormost rays of anal fins black; anal fin dark dusky posteriorly.
Gular and branchiostegal membranes black; mouth and lips dark; gill cavity black except over hyoid
bones and isthmus, which are pale; gill arches and rakers dark; barbel dusky near base, pale distally.

SIZE. — To more than 35 cm TL.

DISTRIBUTION. — Australia, from west coast TAS to North West Cape, WA; in 830 to about
1,500 m.

ETYMOLOGY. — Named for the former CSIRO fisheries research vessel Soe/a, on which were
collected many of the specimens used in this report.

COMPARISONS AND REMARKS. — Specimens from the west coast of Australia north of Cape
Leeuwin appeared to be slightly paler and more brownish over the tail than those from southern
Australia. Furthermore, their snout appeared somewhat more pointed and angular, and the underside
of the snout had a slightly broader naked area. These differences were too slight to consider
recognizing two separate populations.

Nezumia soela is very similar in overall morphology to N. coheni in having a tiny anterior dermal
window lying at, or about on a line with, the pelvic fin insertions, 11 pelvic fin rays, no prominent
dark girdle encircling trunk, and generally similar proportions. The two species differ, however, in
gill raker counts of the first arch (9-12 inner rakers, 1 1—13, rarely 10, rakers on outer arch in N. soela
cf. 6-9 and 9-11, respectively, in NV. coheni), distance orbit to preopercle angle (33-38% HL in N.
soela cf. 38-44%), and size of outer series of premaxillary teeth (larger, thicker in N. soe/a). Pores
of the lateralis system on the lower jaw are larger with raised rims in N. soe/a (cf. smaller, no rims),
the sensory papillae on the underside of the snout are more prominent and numerous in N. soe/a, and
the snout is blunter and narrower. Several features of the two species are compared in Table 5.

The higher gill raker counts, the absence of a prominent dark trunk girdle, and the more anteriorly
placed anterior dermal window distinguish the species from N. namatahi and N. kapala. Nezumia
kapala and N. soela appear to be sympatric, although N. kapala occurs farther to the east and north
(to NSW).

Nezumia spinosa (Gilbert and Hubbs, 1916)
Fig. 43b

Lionurus spinosus Gilbert and Hubbs, 1916:199, pl. 10, fig. 2 (Japan); 1920:554 (4 spec., off Luzon, Philippines).

IWAMOTO AND WILLIAMS: GRENADIERS OF WESTERN AUSTRALIA 203

FiGure 43. (a) Nezumia propinqua (Gilbert and Cramer). (From Iwamoto and Merrett 1997, fig. 29a.) (b) Nezumia spinosa
(Gilbert and Hubbs). Holotype, USNM 76868 (280 mm TL), from off Japan in 777 m. From Gilbert and Hubbs 1916, pl. 10,
fig. 2.) Scale bars represent 25 mm.

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Nezumia spinosa: |wamoto and Anderson, 1994:18, 19 (22 spec., Natal, South Africa and Mozambique).
Williams et al., 1996:149 (WA list). Iwamoto and Merrett, 1997:542—545, fig. 29b (New Caledonia region).

DIAGNOSIS. — Pelvic rays 8 or 9; height first dorsal fin usually much greater than head length;
underside of head almost completely naked, pocketed with rather prominent pores along mandible
and ventral margin of suborbital region; scales densely covered with long needlelike spinules in
parallel to convergent rows, posteriormost spinules on scales extend most of their length beyond scale
margin; 7.5 to 11 scale rows below origin of second dorsal fin; teeth in lower jaw in notably broad
bands, 6 or more teeth wide across anterior end; dark color of trunk confined to abdomen, not
extending onto dorsum; first dorsal fin blackish anteriorly and proximally, paler near distal tips and
posterior margin; anterior dermal window of light organ prominent, situated between pelvic fin bases.

SPECIMENS EXAMINED. — WA: AMS I.22809-046 [formerly 014] (33.4 mm HL, 260+ mm
TL); NW Shelf, 250 km nw. of Port Hedland; 584-592 m; SO2/82/19-21. AMS 1.22810-045 (36.4
HL, 140+ TL); NW Shelf, 250 km nw. of Port Hedland; 694-736 m; SO2/82/22-24. CSIRO
H1492-25 (33.3 HL, 155+ TL); NW Shelf; 420 m; SOS/88/70. CSIRO H2542-29 (38.5 HL, 185 TL);
Exmouth Plateau; 854-868 m; SS1/91/03. CSIRO H2549-10 (49.1 HL, 255+ TL); w. of NW Cape;
650-685 m; SS1/91/10. CSIRO H2557-09 (33.3 HL, 165+ TL); w. of Cape Farquhar; 620 m;
SS1/91/19. CSIRO H2580-04 (47.7 HL, 245+ TL); sw. of Shark Point; 713-714 m; SS1/91/43.
CSIRO H2584-03 (51.1 HL, 220+ TL); w. of Shoal Point; 853 m; SS1/91/49. (Also material listed
by Iwamoto and Anderson (1994) from southern Africa and Iwamoto and Merrett (1997) from New
Caledonia, East China Sea, and South China Sea.)

COUNTS AND MEASUREMENTS (of Australian material only; see Tables 5 and 6). — P. 118-22;
total GR-II (outer/inner) 9—1 1/9—12; scales 1D. 10-13, midbase 1 D. 6.5—9; 2D. 7.5—11; lat.line 35—42.

Total lengths 155+—255+ mm; HL 32.0—-51.1 mm. The following in percent of HL: snout 28-32;
preoral 18-29; internasal 17—21; interorb. 18-24; orb. 28-31; suborb. 12-14; postorb. 43-46;
orb.-preop. 34-40; up.jaw 27-33; barbel 9-18; gill slit 11—14; pre-A. 141-155; V.-A. 34-47; body
depth 78-81; 1D.-2D. 34-52; ht. 1D. 93-156; len. 1P. 48-57; len. V. 61—84; post. nostril 7-11.

SIZE. — To at least 26 cm TL.

DISTRIBUTION. — Japan, South and East China seas, Philippines, Australia (QLD, WA), south-
ern Africa, in 420-900 m (and possibly to 1,258 m, based on field data of AW).

COMPARISONS AND REMARKS. — Nezumia spinosa 1s a distinctive species of the genus owing
to its low pelvic fin ray count, high first dorsal fin with leading edge of prolonged second spinous ray
beset with widely spaced teeth, almost completely naked underside of head pocketed with pores of
sensory lateralis system, notably broad band of dentary teeth, and small body scales covered with
long, needlelike, greatly reclined spinules arranged in somewhat convergent rows. Nezumia spinosa
is likely to be confused only with N. merretti, which it closely resembles but differs in having longer
scale spinules that greatly overlap the posterior scale margin (overlap slightly in N. merretti), smaller
body scales (rows below 1D. 7—7.5 in N. merretti, compared with 10—13 in N. spinosa), and darker
first dorsal fin (overall dusky in N. merretti).

Comparison of Australian specimens of N. spinosa with those reported by Iwamoto and Anderson
(1994) from southern Africa revealed some notable differences in frequency distributions of first
dorsal rays, preoral length, postorbital length and upper jaw length. These are compared in Table 6.
These differences and the disjunct distribution of the species suggest that isolation has resulted in
significant divergence between the populations on each side of the Indian Ocean. The extent to which
the African population has diverged from the eastern population may argue for the recognition of a
second species. Continental drift may have accounted for the disjunct distribution. This species is one
of few Western Australian grenadiers also found off Queensland.

IWAMOTO AND WILLIAMS: GRENADIERS OF WESTERN AUSTRALIA 205

Nezumia wularnia new species
Fig. 44a

Nezumia sp. D: Williams et al., 1996:149 (WA).

DIAGNOSIS. — Pelvic fin rays 11—12; height first dorsal fin about equal to or more than head
length; underside of head scaled, but lacking prominent pores of sensory lateralis system; body scales
with dense covering of broadly shield-shaped spinules; 5.5—7.5 scale rows below origin of second
dorsal fin, 36—42 lateral line scales over distance equal to predorsal length; trunk completely encircled
by broad dark band, although sometimes rather faint; first dorsal fin uniformly blackish; anterior
dermal window of light organ scarcely developed, usually apparent only as triangular anterior
protrusion of naked periproct, situated behind line connecting insertions of pelvic fins; orbit to
preopercle distance 38-44% of head length; upper jaw 29-34%); gill rakers on inner side of first arch
9-10.

SPECIMENS EXAMINED (all from WA). — HOLOTYPE: AMS 1.31157-011 (53.0 mm HL, 245+
mm TL); off Cape Cuvier; 1,293—1,320 m; SS1/91/23. PARATYPES: AMS 1.31157-011 (4, 48.2-55.0
HL, 215+—263+ TL); same data as for holotype. NTM S.12716-007 (76.1 HL, 247+ TL); nw. of
Monte Bello I.; 20°15’S, 114°50’E; 500-600 m; coll. D. Evans; 22.11.1990. CSIRO H2545-01 (2,
53.6-55.2 HL, 240+—280 TL); over Exmouth Plateau; 1,023 m; SS1/91/05. CSIRO H2544-14 (68.8
HL, 340+ TL) and H.2544-12 (64.1 HL, 330+ TL); over Exmouth Plateau; 1,128 m; SS1/91/04.
CSIRO H2553-09 (205+ TL); w. of Point Cloates; 910 m;SS1/91/14. AMS 1.31159-005 (6, 57.0—73.4
HL, 234-370+ TL); off Cape Cuvier; 1,293—1,320; SS1/91/23. CSIRO H2581-15 (4, 49.4-55.2 HL,
230+—276+ TL); sw. of Shark Bay; 996 m; SS1/91/44. NMV A9583 (3, 50.0-56.0 HL, 240-265 TL):
95 km sw. of Geraldton; 29°21.8’S, 113°46.6’E; 942-970 m. CSIRO H2615-07 (61.8 HL, 315+ TL),
H2615-08 (40.8 HL, 240 TL); and NMV A9523 (67.2 HL, 365 TL); w. of Mandurah; 1,030—1,140
m; SS1/91/83. CSIRO H2616-07, H2616-08, H2616-09; w. of Mandurah; 960 m; SS1/91/84. CSIRO
H3017-03 (370 TL); nw. of Cape Leeuwin, 34°10’S, 114°16’E; 1,030 m; Akebono Maru No. 3, shot
17; coll. A. Williams; 22.XI1I.1989. CSIRO H3022-03 (2 spec.); w. of Bunbury; 33°17’S, 114°13’E;
976 m; Akebono Maru No. 3, shot 22; coll. A. Williams; 25.XII.1989.

COUNTS AND MEASUREMENTS (holotype first, followed by range in 19 paratypes). — 1D. II,11,
II,10—12; P. 123/121, 118-123; total GR-I (outer/inner) ? / ?, 6—-8/9-10, GR-II ? /?, 8—9/9—10; scales
1D. 8, 7—7, midbase 1D. 5.5, 4.5—6.5, 2D. 6.5, 5.5—7.5, lat.line 37, 36-42.

Total length 215+—370+ mm; HL 40.8—73.4 mm. The following in percent of HL: snout 32,
27-32; preoral 24, 19-24; internasal 24, 20-25; interorb. 21, 18-23; orb. 30, 28-35; suborb. 16,
15—17; postorb. 44, 40-46; orb.-preop. 39, 38-44; up.jaw 34, 29-34; barbel 15, 11—18; gill slit ? ,
12-17; pre-A. 168, 148-168; V.-A. 45, 28-46; body depth 84, 74-89; 1D.-2D. 41, 35-59; ht. 1D. 82,
82—113; len. P. 58, 46-60; len. V. 65, 43—70; post. nostril 5, 3—7.

DESCRIPTION. — General features shown in Figure 44a. Snout conical in lateral profile, tipped
with large, but rather inconspicuous, tubercular scale; scales at lateral angles similarly large and
inconspicuous. Suborbital shelf strong, composed of two distinct rows of modified scales; ridge
formed by ventral margin of lower row not sharply angular, ventral aspects of suborbital rather
smoothly blend in with shelf dorsally. Ventral surfaces of snout, suborbital, preopercle, and mandible
completely and uniformly scaled; scales on underside of snout fairly deciduous and often lost. Free
neuromast of cephalic lateralis system prominent in pale specimens. Body scales densely covered
with broadly lanceolate to shield-shaped spinules.

Small fine teeth in bands in both jaws; outer premaxillary teeth slightly enlarged.

Long spinous ray of first dorsal fin relatively slender, especially prolonged distal portion:
serrations along leading edge fine and widely spaced. Other fins without notably prolonged rays; outer
pelvic fin ray slightly prolonged, extending at best to bases of first few anal fin rays.

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TABLE 6. Comparison of six characters of Nezumia spinosa by area: southern Africa and Australia.

Segmented 1D rays

8 9 10 11 12 N x
S. Africa -_ = 8 14 1 23 10.70
Australia ] 4 3 _ _ 8 9.25

Pelvic fin rays

8 9 N x
S. Africa 46 ber 46 46.00
Australia 12 4 16 8.25

Preoral length (“oHL)

Ie—18) WO—v)| Ae) WES 7) DS) BN =SBhil B= 8)2}) 34S) N 58
S. Africa = = — = — 5) 12 3 3 23 30.91
Australia 2) 2 2) = 1 1 = — = 8 22.38

Postorbital length (%HL)

37-38 39-40 41-42 43-44 45-46 N x
S. Africa 2 8 13 — _ 23 40.65
Australia — _ — 4 4 8 44.50

Length Outer Gill Slit (“oHL)

9 10 11 1 13 14 N X
S. Africa l 3 6 4 ) 19 11.89
Australia — = 1 2 D 2 7 12E al

Length Upper Jaw (%HL)

23 24 25 26 20 28 29 30 31 32 38 N 6

S. Africa ] — 2 3 6 6 2 ] 21 27.10
Australia — - — — 1 ] = 2 — 3 ] 8 30.50

Periproct region large, far removed from anal fin origin and closer to pelvic fin bases; minute,
inconspicuous anterior dermal window anterior to periproct and situated posterior to line connecting
insertions of pelvic fins.

Ground color of variable intensity; some specimens very dark brown, with nape, top of head, and
tail faintly paler; in others these areas distinctly paler. A broad, dark trunk girdle of variable intensity
(sometimes faint) present on all specimens. Abdomen bluish to black. Gular and branchiostegal
membranes and operculum, including hind margin of preopercle, black. Lips blackish, but premax-

IWAMOTO AND WILLIAMS: GRENADIERS OF WESTERN AUSTRALIA 207

FiGuRE 44. (a) Nezumia wularnia n.sp. Holotype, AMS 1.31157-011 (245+ mm TL), from off Cape Cuvier, WA, in
1,293—-1,320 m. (b) Nezumia soela n.sp. Holotype, CSIRO H3017-04 (350+ mm TL), from northwest of Cape Leeuwin, WA,
in 1030 m. Fins and scales partially reconstructed. Scale bars represent 25 mm.

illary anteriorly (over ascending limbs) and angle of jaws pale. Barbel dark at base, pale distally.
Paired and first dorsal fins black; anal fin blackish throughout in darker specimens, but in paler
specimens only anterior end of anal fin blackish, with dusky rays posteriorly over remainder of fin.

SIZE. — To more than 37 cm TL.

DISTRIBUTION. — Known only from off WA, from Exmouth Plateau (ca. 21°S) to west of
Mandurah (ca. 32°S), in 685—1,320 m.

ETYMOLOGY. — From the Yindjibarndi, wu/arni, from the west.

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COMPARISONS AND REMARKS. — Nezumia wularnia differs from most other members of Nezu-
mia in having the underside of the head completely scaled. (A few specimens of N. wularnia had a
medium swath of naked skin ventrally on the snout, but the scales in that area appeared to have been
sloughed off during capture or subsequent handling.) In other species of Nezumia from the region,
the underside of head is almost completely naked, or at the least, most of the underside of snout is
naked. Additionally, body scales in the new species are slightly larger, with scale-row counts below
the dorsal fins somewhat lower. Nezumia wularnia agrees in many respects with N. kapala but differs
in having slightly more first dorsal fin rays, a longer distance orbit to preopercle, and somewhat longer
upper jaw (see Tables 5 and 7).

Pseudonezumia Okamura, 1970

Pseudonezumia Okamura, 1970:38 (type species Pseudonezumia japonicus Okamura, 1970, by original desig-
nation).

Paracetonurus Marshall, 1973:615 (type species Macrourus parvipes Smith and Radcliffe, 1912, by original
designation).

See Iwamoto and Anderson (1994:21, 22) and Sazonov and Shcherbachev (1982b:11) for
diagnosis (as Paracetonurus). At least five species, including Pseudonezumia cetonuropsis (Gilbert
and Hubbs, 1920), P. flagellicauda (Koefoed, 1927), P. japonicus, P. parvipes, and P. pusilla Sazonov
and Shcherbachev, 1982b. Sazonov and Shcherbachev (1982b:11) included all of the above except
P. japonicus in the genus Paracetonurus.

Pseudonezumia pusilla (Sazonov and Shcherbachey, 1982)
Fig. 45

Paracetonurus pusillus Sazonov and Shcherbachev, 1982b:12—14, fig. 4 (Ninety East Ridge, central Indian
Ocean and Bismarck Sea off New Guinea; 1,380—2,000 m).

Pseudonezumia puscilla [sic]: Williams et al., 1996:149 (WA list).

Pseudonezumia pusilla: lwamoto and Merrett, 1997:547, 548, fig. 30b (New Caledonia off Loyalties).

DIAGNOSIS. — Chin barbel usually 14-23% HL, interorbital width 29-35%, upper jaw 30-39%,
body 59-87% (usually <75%). Adult size relatively small, about 250 mm TL or less.

SPECIMENS EXAMINED. — CSIRO H2551-20 (30.5 mm HL, 186+ mm TL); off WA, w. of NW
Cape; 1,460—1,500+ m; SS1/91/12.

COUNTS AND MEASUREMENTS (based on | WA spec.). — 1D. II,10; P. 115-116; V. 6; total GR-I
(outer/inner) 9/11, GR-II 10/11; scales midbase 1D. 13.

The following in percent of HL: postrostral 74; snout 31; preoral 22; internasal 26; interorb. 34;
orb. 33; suborb. 15; postorb. 43; orb.-preop. 36; isth.-A. 65; body depth 73; depth at A. 72; 1D.-2D.
41; post. nostril 19; up.jaw 35.

DESCRIPTION. — Body slender, compressed, tapering gradually. Head relatively large, dorsal
profile more or less flat; snout broad and high; orbits relatively large, the anterior margin adjacent to
posterior nostril developed as thin lateral flange; perimeter of nostril distinctly large, circular; mouth
long, narrow, subterminal. Gill rakers on outer side of first gill arch a series of rounded tubercles
lacking spination; those of inner side of first arch and all other arches well developed and armed with
series of dorsally directed, long, narrow, pointed spines.

Suborbital ridge and ridges of head poorly defined, their scales lacking enlargement or bony
development; head and underside of jaw scaled; branchiostegal and gular membranes naked. Scales
on head adherent, armed with long, slender, erect, and widely spaced spinules; body scales mostly
deciduous.

IWAMOTO AND WILLIAMS: GRENADIERS OF WESTERN AUSTRALIA 209

TABLE 7. Comparison of first dorsal fin ray counts, upper jaw length, and distance orbit to preopercle in
Nezumia wularnia and N. kapala.

Segmented First Dorsal Rays

8 9 10 11 12 iG
N. wularnia - _ 11 12 | 10.17
N. kapala 3 18 3 | — 9.08
Upper Jaw length (%HL)
26 Daf 28 29 30 31 32 33 34 x
AY elena E e = ib iy a5 5 5 4 4 asm
N. kapala 1 5 6 6 3 2 — ] -- 28.67

Orbit to Preopercle distance (Y%HL)

34 335) 36 37 38 39 40 4] 42 43 44 Xx
N. wularnia 3 3 7 8 ] 1 ] 40.33
N. kapala 2 7 7 3 p 3 36.21

Teeth slender, conical, forming tapering bands in both jaws; 2—3 indistinct rows at symphysis of
each jaw, those of outer row slightly enlarged.

Second bony element of first dorsal fin long, with numerous spinulations; pelvic fin bases close
together, situated well in advance of pectoral fin bases; all fins rather poorly developed.

Periproct region adjacent to anal fin origin, well developed, broadly oval in shape with a short
anterior extension; no external evidence of an accessory light organ.

Body color yellowish brown; head dark brown; operculum, branchiostegal membranes, mouth
and branchial cavities black; anterior section of anal fin base and dorsal midline darkish brown; fins
pale to dusky; periproct region black.

SIZE. — Sazonov and Shcherbachev (1982) reported sexually mature adult specimens reaching
a maximum length of around 25 cm TL—a relatively small size among members of the genus.

DISTRIBUTION. — Widespread in the tropical Indian Ocean and the western Pacific. Off Western
Australia, known from a single specimen taken in the deepest sample from which material was
examined (1,460 m to about 1,500 m).

COMPARISONS AND REMARKS. — Our specimen agreed well with P. pusilla, although it has a
well developed, broadly oval periproct with a short anterior extension (not “comparatively narrow,”
as described by Sazonov and Shcherbachev, 1982b). Those authors differentiated P. pusilla from the
two other Indo-West Pacific species of the genus, P. parvipes and P. cetonuropsis, on its greater orbit
diameter and longer barbel. Comparing their data with those of Gilbert and Hubbs (1916), the
respective differences of the three species are: orbit (29-38% of HL cf. 20-25% and 27%), and barbel
(14-20% HL cf. 4-9% and 6%). Additionally, the anal opening of P. pusilla is adjacent to the anal
fin origin, whereas in P. cetonuropsis it is separated by a distance about one-third distance between
anal and pelvic fins.

210 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 3

FIGURE 45. Pseudonezumia pusilla (Sazonov and Shcherbachev). Specimen CSIRO H2551-20 (30.5 mm HL, 186+ mm TL)
from off North West Cape, WA, in 1460-1500 m. Drawn by Georgina L. Davis.

\
if

pela
Baw yi,

FIGURE 46. Sphagemacrurus pumiliceps (Alcock). Specimen CSIRO H2551-22 (35.7 mm HL, 253 mm TL) from off North
West Cape, WA, in 1460-1500 m. Drawn by Georgina L. Davis.

Sphagemacrurus Fowler, 1925

The forward shift of the ventral aspects of the trunk, coupled with a blunt snout are convergent
developments in morphology with members of the genus Lucigadus. The location of the anus and
urogenital opening in a broad naked area extending about 2/3 distance between pelvic and anal fins
differs markedly from the condition in Lucigadus. Only one species of the genus was taken in WA,
but Sphagemacrurus richardi (Weber, 1913) should be expected in the tropical regions, as it has been
taken off Queensland and Indonesia.

Sphagemacrurus pumiliceps (Alcock, 1894)
Fig. 46

Macrurus pumiliceps Alcock, 1894:125—127 (Laccadive Sea; 1,315 m)

Lionurus pumiliceps: Gilbert and Hubbs, 1920:559, 560 (Philippine and East Indies; 732—1,646 m).

Sphagemacrurus pumiliceps: Marshall and Iwamoto in Marshall, 1973:627 (list; lectotype designation);
Iwamoto, 1986:339 (Mozambique; 1,510—1,600 m); Shcherbachev, 1987:41 (Indian Ocean distr.; 880—
1,880 m).

IWAMOTO AND WILLIAMS: GRENADIERS OF WESTERN AUSTRALIA 211

DIAGNOSIS. — Orbit longer than snout, about |.0—1.2 times into distance from orbit to angle of
preopercle, 33-37% of HL; underside of snout mostly covered with small, irregularly shaped scales;
a pale, naked area anterior to margin of upper lip; scales mostly deciduous except on nape and head;
teeth small, slender, with curved tips, 4—S rows at symphysis of premaxilla, 2—3 rows at symphysis
of mandible; pelvic fin rays 1 1—13; total inner gill rakers on first arch 8—1 1; body color variously pale
to dusky brown on nape and flanks, darker grayish brown on head.

SPECIMENS EXAMINED. — WA: CSIRO H2617-03 (34.5 mm HL, 231 mm TL); off Bunbury;
982 m; SS1/91/85. CSIRO H2551-22 (35.7 HL, 253 TL) and H2551-23 (6, 170-240 TL); off NW
Cape; 1,460—1,500+ m; SS1/91/12. CSIRO H2581-16 (2, 27.2-28.4 HL, 160-213 TL); 996—1,009
m; SS1/91/44. Madagascar: CAS 66504 (4, 18.0-32.3 HL, 147+—205 TL); off Cape St. Vincent;
22°27'S, 43°00’'E; 940-960 m; Vityaz’ sta. 2652; coll. M. E. Anderson; 3.XII.1988.

COUNTS AND MEASUREMENTS. — 1D. II,9—11; P. 119-122; total GR-II (outer/inner) 9—10/10—11;
scales 1D. 12—14, midbase 1D. 10-12, 2D. 7—10, lat.line 36-38.

Total length 160—253 mm; HL 27.2—38.0 mm. The following in percent of HL: snout 30-34;
preoral 16-21; internasal 24-29; interorb. 24-28; orb. 33-37; suborb. 15—18; postorb. 40-42;
orb.-preop. 34-38; up.jaw 34-38; barbel 16—21; gill silt 18-21; pre-A. 113—150; body depth 93-121;
1D.-2D. 41—S6; ht. 1D. 95 (1 spec.); len. P. 60-65; len. V. 52—71; post. nostril 6—12.

DESCRIPTION. — Body deep and strongly compressed; snout blunt, broad, with well-developed
bony scutes at tip and lateral angles; bony scutes with a crown of short, blunt, conical spinules.
Underside of snout mostly covered with small, irregularly shaped scales; naked area around margin
of upper lip. Scales easily lost through abrasion. Suborbital ridge well developed with short, blunt,
conical spinules present on lateral margins of ridge and lower margin of orbit; ridge extending beyond
midline of orbit. Jaws small, subterminal, oblique; upper jaw extending posteriorly to below midorbit.
Short fleshy barbel present. Gill rakers moderately developed, tubercular with stout spines on tip; gill
filaments well developed.

Body scales with about 5 near-parallel rows of short, sharp, slightly raised spinules; scales mostly
deciduous except on head and nape. Anus and urogenital opening ina broad naked periproct extending
about 2/3 distance between pelvic and anal fins.

Teeth small, slender, with curved tips; 4-5 rows at symphysis of premaxilla, 2—3 rows at
symphysis of mandible.

Base of first dorsal fin elevated, angled at about 45 degrees to long axis of body; second spine
long, slightly curved, leading edge with series of well-developed serrations becoming progressively
finer and more widely spaced towards tip; height of fin about equal to head. Pectoral fin poorly
developed; leading ray not prolonged; pelvic fin origin below gill cover, well in advance of pectoral
fin in, leading ray with moderately developed filamentous extension reaching to at least 10th anal fin
ray.

Body lacking prominent markings; nape and flanks variously pale to dusky brown; head darker
grayish brown. Buccal cavity and jaws black; lips dusky; margin of orbit dark. Underside of snout
brown, naked area anterior to margin of upper jaw pale; dark free neuromasts irregularly spaced over
head; opercle and preopercle, branchiostegal membranes, and abdomen black. Pelvic fins black at
base; pectoral, dorsal, and anal fins variable in color from pale to dusky to blackish.

SIZE. — A small species attaining about 26 cm TL.

DISTRIBUTION. — Widely distributed in the Indo-West Pacific. Taken off WA between 882 m
and about 1,880 m.

COMPARISONS AND REMARKS. — Our specimens generally agreed well with published data on
S. pumiliceps, however, they varied in having a relatively short postorbital distance (34-38% HL cf.
38-45%) and in being scaled on the underside of the snout. They also differed in several other
characters compared to S. pumiliceps from Madagascar, which have fewer pelvic fin rays (9-11 cf.
11-13) and a wider interorbital (30-37% HL cf. 24-28%). Scale spinulation was also markedly

22 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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coarser in the Madagascan specimens, particularly the degree of development on scales of the
suborbital ridge and those of the anterolateral margin of the snout. Body color in preserved material
was a uniform dark brown in Madagascan specimens compared to a variously pale to dusky brown
nape and flanks, with darker grayish brown head in the WA specimens. There may, therefore, be
unresolved problems with identification of Sphagemacrurus species in the Indian Ocean and west
Pacific regions, which will only be resolved by examination of material from a wide range of localities.

Sphagemacrurus pumiliceps is most similar to S. richardi from Indonesia, but can be differenti-
ated from that species by its higher pelvic fin ray count (11—13 cf. 8-10). It is also similar in many
counts and proportional measurements to S. hirundo (Collett, 1896) and S. grenadae (Parr, 1946),
known respectively from the northeastern and western Atlantic. However, in S. hirundo the snout and
interorbital dimensions are about equal (Marshall 1973), whereas in S. pumiliceps, the snout is
relatively long (30-34% HL cf. 24-27%). Marshall reported S. grenadae as being dark brown to
black, whereas S. pumiliceps is variously pale to dusky brown.

Trachonurus Gunther, 1887

See Sazonov and Iwamoto (1992:77) for diagnosis of genus. The number of species in this genus
is still uncertain. Two species have been recently described (Trachonurus gagates Iwamoto and
McMillan, 1997, and 7. robinsi Iwamoto, 1997). Trachonurus sentipellis Gilbert and Cramer, 1897,
which had been synonymized with T. villosus, was recognized by Iwamoto and McMillan (1997).
Three species are so far known from Western Australia, including a new species herein described.

KEY TO WESTERN AUSTRALIAN SPECIES OF TRACHONURUS

llaSGroovedilateralilineicompletelysabsent\ 1 yer =) ecient cee tet ey ecu T. gagates
lib: ‘Groovedilateralilinésprésent, = s-.5-0-4) 5 ce ee ee ee oS a eS 2
2a. Teeth all small, outer premaxillary series scarcely or not at all enlarged; 26—34 lateral line scales counted from an-
terior end over distance equal to predorsal length; body scales large, 8 or 9 between pelvic fin base and gill cavity,
about 8 from pelvic fin base to lower edge of pectoral finbase .................-....- T. sentipellis
2b. Outer series of premaxillary teeth distinctly enlarged, inner series of dentary teeth usually slightly enlarged; 39-50
lateral line scales over distance equal to predorsal length; body scales relatively small, 10 or 11 between pelvic fin
base and gill cavity, 12 or more between pelvic fin base and pectoral finbase ............ T. yiwardaus n.sp.

Trachonurus gagates |wamoto and McMillan, 1997
Fig. 47

Trachonurus sp. A: Williams et al., 1996:149 (WA).

Trachonurus gagates Iwamoto and McMillan, 1997:255—259, fig. 1 (holotype AMS 1.24059-009; Australia,
NSW off Norah Head, 978 m; paratypes from Australia [NSW, QLD, SA, TAS, VIC, WA], and New
Zealand; 435—1,240 m).

DIAGNOSIS. — Grooved lateral line absent; chin barbel 4—-8%, suborbital width 13—15%, post-
orbital 50-55% of head length; total gill rakers on second arch (outer/inner) 10—13/10—12; scale rows
from origin of second dorsal fin to anal fin 20—25; lateral scales over distance equal to predorsal length
usually 35-42; outer premaxillary teeth slightly enlarged; gular and branchiostegal membranes
usually heavily covered with scales.

SPECIMENS EXAMINED. — Type specimens listed in Iwamoto and McMillan (1997).

SIZE. — To about 48 cm TL.

DISTRIBUTION. — Widespread in Australia (NSW, QLD, SA, TAS, VIC, WA) and New Zealand
in 435—1,200 m.

IWAMOTO AND WILLIAMS: GRENADIERS OF WESTERN AUSTRALIA 213

COMPARISONS AND REMARKS. — Trachonurus gagates 1s closely similar to 7. vi/losus (Gunther,
1877) with the primary difference being the absence of a grooved lateral line in 7. gagates compared
with a well-developed one in 7. villosus. The two species are similar in having small scales,
compressed head, bluntly rounded snout profile, and relatively small outer premaxillary teeth.

Trachonurus sentipellis Gilbert and Cramer, 1897
Fig. 48

Trachonurus sentipellis Gilbert and Cramer, 1897:429, 430, pl. XLV, fig. 1 (Hawaiian Islands). lwamoto and
Merrett, 1997:79, 80, fig. 31b (New Caledonia region; 760—980 m).
Trachonurus sp. B: Williams et al., 1996:149 (WA).

DIAGNOSIS. — Grooved lateral line well developed; body scales relatively large, coarsely cov-
ered with stout, erect spinules, 26—34 lateral-line scales over distance equal to predorsal length, 4-6
below midbase | D., 5—7 below 2D. origin; small scale patch or none on gular membrane, few or no
scales at base of ventral-most branchiostegal rays. Teeth in both jaws all small, outer premaxillary
series scarcely enlarged. Total GR-II (outer) 10—14. Pyloric caeca short, thick, 9-13.

SPECIMENS EXAMINED. — WA: CSIRO H2541-15 (2, 53.1-59.5 mm HL, 258+—294+ mm TL);
Exmouth Plateau; 914 m; SS1/91/02. AMS 1.31159-006 (6, 39.9-62.8 HL, 200-300+ TL); off Cape
Cuvier; 1,060—1,064 m; SS1/91/21. CSIRO H2592-09 (2, 57.2-59.3 HL, 300+—290+ TL); w. of
Leander Pt.; 942-970 m; SS1/91/58. NMV A9653 (65.7 HL, 325 TL); w. of Leander Pt.; 1,132—1,136
m; SS1/91/61. NMV A9640 (2, 53.7-57.4 HL, 270-315 TL); ssw. of Geraldton; 770-760 m;
SS1/91/62. CSIRO H3041-13 (44.5 HL, 223+ TL); sw. of Shark Bay; 26°36’S, 112°09’E; 760 m;
Akebono Maru No. 3, shot 41; coll. A. Williams; 28.XII.1989. NSW: AMS 1.28100-003 (44.9 HL,
258 TL); e. of Taree; K88-08-06. AMS 1.24462-003 (51.3 HL, 248+ TL); off Cape Hawke; 980 m;
K 83-15-02.

UNCERTAIN VARIANTS.—NSW: AMS I[.27718-010 (51.9 HL, 295+ TL); NSW, off Brush I.;
1,190 m; 1988. AMS 1.20920-005 (47.6 HL, 263+ TL); QLD, ne. of Raine I.; 11°32'S, 144°10’E;
0-900 m; field no. FNQ79—33; 12.11.1979. WA: NTM S.12716-010 (43.1 HL, 247+ TL); nw. of
Monte Bello I.; 20°15’S, 114°50’E; 500-600 m; coll. D. Evans; 22.11.1990. NMV A11386 (54.6 HL,
312 TL); w. of Leander Pt.; 1,132—1,136 m; SS1/91/61.

COUNTS AND MEASUREMENTS (15 WA and 2 NSW specimens).— 1D. II,7—8(9); P. (11 1)i12—4115;
V (6)7; total GR-I (outer/inner) 2—8/(11)12—14, GR-II 10—14/12—14; scales 1D. 6-7, midbase 1D.
4-6 [total to A. 20-25], total 2D.-A. 14-21.

Total length 223+—325 mm; HL 44.5—67.5 mm. The following in percent of HL: snout 24-30;
preoral 14-19; internasal 18—22; interorb. 31—36; orb. 26-34; suborb. 10—12; postorb. 45—50;
orb.-preop. 31—37; up.jaw 30-36; barbel 9-14; gill-slit 12-17; pre-A. 134-165; body depth 69-87;
1D.-2D. 17—S2; ht. 1D. 49-61; len. P. 40-51 len. V. 30-47; nostril 5—11.

DESCRIPTION. — General features of fish seen in Figure 48. Head 5—6 times into total length;
greatest head width 1.7—1.8 into its length. Snout gently conical and protruding in lateral view, blunt
in dorsal view. Orbits relatively large, about equal to snout length and about 0.7 into broad, slightly
depressed interorbital space. Preopercle gently rounded, the chord of margin vertical. Jaws large,
extending posteriorly to below midorbit or slightly beyond. Chin barbel small, slender, length about
equal to least suborbital width.

Teeth fine, short, in narrow bands in both jaws, extending almost to end of rictus. Premaxillary
band about 4 or 5 teeth wide, tapering to 2 irregular series, then to | row near posterior end; outer
series scarcely, if at all, larger than inner teeth. Mandibular teeth about 3 or 4 teeth wide anteriorly,
becoming 2 irregular series, then | series near posterior end; inner teeth very slightly larger than outer.

214 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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FIGURE 47. Trachonurus gagates lwamoto and McMillan. (From Iwamoto and McMillan 1997, fig. 2).

FIGURE 48. Trachonurus sentipellis Gilbert and Cramer. (From Iwamoto and Merrett 1997, fig. 31b.) Scale bar represents
25 mm.

IWAMOTO AND WILLIAMS: GRENADIERS OF WESTERN AUSTRALIA 215

FiGURE 49. Body scales from dorsum below origin of second dorsal fin of (a) Trachonurus sentipellis and (b) T. yiwardaus.
Scale bars represent 1.0 mm.

216 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 3

Body scales large, coarsely covered with short, erect conical spinules (Fig. 49a). Enlarged,
vertically elongated scales along anal fin base; these armed with stouter spinules and interconnected
buttresses. Scales along each side of second dorsal also somewhat enlarged, with larger spinules
recurved and somewhat reclined. Scales over dorsal surfaces of preopercle and opercle platelike and
enlarged; those along posterior margin of orbit somewhat elongated, other scales of head much
smaller. Gular and branchiostegal membranes usually sparsely scaled or almost naked, but most with
small median patch of scales on gular membrane and isolated patches (or narrow files) of scales at
base of anteriormost branchiostegal rays. Scales over abdominal region large with spinule fields
sharply defined and with little overlap in scale margins. Scales over chest also large, usually 9 or
fewer rows between pelvic base and gill cavity, about 8 rows from pelvic base to lower edge of pectoral
base. Lateral line scales with prominent groove throughout and few or no interruptions.

Periproct region large, round to somewhat oval, with posterior margin abutting anal fin; entire
span between anal origin and inner rays of pelvic fins naked; anus about midway between or slightly
closer to anal fin.

Color in alcohol medium brown to gray overall; swarthy over abdomen, operculum, and laterally
and ventrally on head. Branchiostegal and gular membranes dark gray to blackish; branchial cavity
black, mouth, lips, jaws dark gray to black; barbel dark brown; peritoneal membranes black. Rays of
first and second dorsal fins dusky, pectoral and anal usually darker, pelvic fins blackish.

SIZE. — To at least 31 cm TL.

DISTRIBUTION. — So far known from the Hawaiian Islands, Australia (WA and NSW), and the
New Caledonian region, but probably more widespread. Capture depths off Australia varied from 500
mi cop 1 32m:

COMPARISONS AND REMARKS. — Our Western Australian specimens agree well, for the most
part, with Hawaiian and New Caledonian specimens of Trachonurus sentipellis. We were troubled
by four specimens, however, which we list as uncertain variants: one is from the North West Shelf,
another from southwest of Geraldton, WA, and two from NSW (off Brush Island and northeast of
Raine Island). The North West Shelf specimen (NTM S.127165-010) agreed well in almost all features
of the species except for the scale spinules, which were notably fine with flexible tips that gave a
velvety feel to the body surface. In contrast, all other specimens of the species had stout spinules that
gave a rough feel to the surface. The specimen from off Geraldton (NMV A1 1386) had low gill raker
counts (9 on inner series of first arch and 9 and 10, respectively, on the outer and inner series of the
second arch), low lateral line scale count (24), relatively greater snout length (30% HL), preoral length
(19%), internasal width (24%), interorbital width (37%), and height first dorsal (71%). Scales on the
branchiostegal rays were more extensive than on the other specimens. The Brush Island (AMS
1.27718-010) and Raine Island (AMS I.20920-005) specimens had relatively high lateral line scale
counts of 39 and 36, respectively. In addition, the Brush Island specimen had low gill raker counts
(8 total on inner series of first arch, 8 and 9 on outer and inner series of second arch) and a slightly
wider internasal distance (23% HL cf. 18—22%) than we found in the other specimens of 7. sentipellis.
Compared with a specimen of 7. sentipellis of almost identical size (AMS 1.27718-010) from Cape
Hawke, NSW, there was a notable difference in appearance. The Brush Island specimen appeared
slimmer, the snout more pointed, the head shallower and wider, the body scales larger, and the scale
spinules were somewhat coarser and more deciduous. The Raine Island specimen had low gill raker
counts (11 on inner side of first and second arches, but the counts were within the range found in other
specimens of 7. sentipellis), and a lateral line that was interrupted into long dashes behind the
abdominal cavity. Aside from the high lateral-line scale count, other meristic and morphometric
features fell well within the range of the species. A more thorough study of additional specimens is
needed to determine if these specimens are simply variants of 7. sentipellis or representatives of
different taxonomic entities.

IWAMOTO AND WILLIAMS: GRENADIERS OF WESTERN AUSTRALIA 217

Trachonurus yiwardaus new species
Fig. 50

Trachonurus sp. C: Williams et al., 1996:49 (WA).

DIAGNOSIS. — Grooved lateral line present; body scales relatively small, lateral line scales over
distance equal to predorsal length 39—S0, scale rows between pelvic fin base and operculum | 1—12,
below midbase of first dorsal fin to anal fin 26-30, below origin of second dorsal fin 6—9; scale
covering on gular and branchiostegal membranes extensive. Teeth in both jaws relatively small, but
with a distinctly enlarged outer series on premaxillary and a slightly enlarged inner series on dentary.
Total gill rakers on inner series of first arch 9-13, on inner and outer series of second arch 9-12.

SPECIMENS EXAMINED. — WA: HOLOTYPE: NMV A9643 (72.8 mm HL, 422 mm TL); w. of
Leander Pt.; 1,132—1,136 m; SS1/91/61. PARATYPES: NMV 19782 (formerly A9643) (3, 36.4—40.3
HL, 218—238 TL) and CAS 200231 (73.1 HL, 390+ TL); same data as for holotype. AMS I.3157-007
(2, 35.0-53.8 HL, 190-350 TL); off Cape Cuvier; 1,060—1,064 m; SS1/91/21. AMS 1.31180-010
(57.5 HL, 310+ TL); sw. of Shoal Pt.; 945-960 m; SS1/91/48. CSIRO H2592-10 (2, 55.5—64.5 HL,
270+—367+ TL); w. of Leander Pt.; 942-970 m; SS1/91/58. CSIRO H4783-01 (2, 50.4-60.8 HL,
265+—237+ TL); Western Australia; SS1/91 (no other data). NMV A11385 (78.5 HL, 383 TL) and
NMV A11386 (54.6 HL, 312 TL); 95 km sw. of Geraldton; 942-970 m; SS1/91/61. SA: CSIRO T321
(84.0 HL, 500+ TL); 33°58’S, 131°49’E; 930 m; Margaret Phillipa sta. 6/3; 10.X1.1984. CSIRO T317
(62.6 HL, 387+ TL); 34°28’S, 132°03’E; 1,175 m; Margaret Phillipa sta. 6/19; 14.11.1984. CSIRO
T320 (61.8 HL, 380+ TL); 34°10’S, 131°37’E; 1,115 m; Margaret Phillipa sta. 10/5; 14.V1.1985.

_

FIGURE 50. Trachonurus yiwardaus n.sp. Holotype, NMV A9643 (72.8 mm HL, 42 mm TL), from 115 km southwest of
Geraldton, WA, in 1,132—1,136 m. Fins partially reconstructed. Scale bar equals 25 mm.

218 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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UNCERTAIN VARIANT—WA: CSIRO H3002-05 (55.9 HL, 280+ TL); se. of Albany; 35°23’S,
118°27'E; 1,030 m; Akebono Maru No. 3, shot 2; coll. A. Williams; 21.X1I.1989. NTM S.12716-010
(71.5 HL, 350+ TL); nw. of Monte Bello I.; 20°15’S, 114°50’E; 500-600 m; coll. D. Evans;
22 A LO90.

COUNTS AND MEASUREMENTS (exceptional values are in parentheses)—1D. II,6—9; P. 112-118;
V. (6) 7; total GR-I (outer/inner) 3—8/9—13, GR-II 9—12/9—12; scales 1D. 7-10, midbase 1D. 4~7,
midbase 1D.-A. 26—30, 2D. 6-9, 2D.-A. 17—23, lat.line 39-50; caeca 9-13.

Total length 238—500+ mm; HL 35.0—84.0 mm. The following in percent of HL: snout 25-31;
preoral 14—19; internasal 20—25; interorb. 33—37(39); orb. 24—31; suborb. 12—15; postorb. 46—S2;
orb.-preop. 32—38; up.jaw 33-39; barbel 6—14; gill slit 13-17; pre-A. 139-162; V.-A. 28-42; body
depth 73—92; 1D.-2D. 21-30; len. 1D. 41—72; len. P. 37-67; len. V. 30-43; post. nostril 4—10.

DESCRIPTION. — General features of fish seen in Figure 50. Head about 6 in total length. Snout
rounded to bluntly pointed in profile, relatively high and not much protruding beyond mouth, length
less than interorbital width. Internasal about 1.6—1.7 into least interorbital width. Orbits round,
greatest diameter much less than snout and interorbital. Upper jaw extends posteriorly to below
posterior 1/3 to 1/4 of orbit.

Body scales relatively small, spinules short, slender, erect (Fig. 49b); scales dorsally along
median line with strongly recurved spinules, scales along second dorsal and anal fins somewhat larger
than adjacent scales.

Gular and branchiostegal membranes liberally covered with small scales forming a Y-shaped
patch, with the arms of the Y over the anteriormost branchiostegal rays. In some specimens, scale
patch broken up. Area between pelvic fins and periproct usually scaled, but completely naked in a
few specimens.

Premaxillary teeth short, slender, in moderately wide tapered band about 4 to 6 teeth across widest
portion near symphysis, tapering posteriorly to | or 2 rows; outer series slightly enlarged. Mandibular
teeth also in band slightly narrower than that of premaxillary, narrowing to 2 or 3 rows posteriorly,
with inner series slightly larger than outer.

Pyloric caeca variable from short and stubby to long and fingerlike, length from much less than
suborbital width to almost twice suborbital; 11-13 counted in 5 specimens.

Color in alcohol apparently variable, but holotype and most of paratypes uniformly dark gray to
brownish gray. Smaller paratypes (formerly NMV A9643) from same collection as holotype had
blackish abdomen, snout, opercle, and subopercle; CSIRO T316 similarly colored, but most of head
pale gray, with free neuromasts on head prominently marked (no other specimen had such prominent
free neuromasts). CSIRO T320 exceptional in more brownish than grayish coloration.

SIZE. — Attains at least 50 cm TL.

DISTRIBUTION. — Known only from WA and SA, from off Cape Cuvier, WA, south and east to
GAB (longitude 132°E), in 930—1,175 m (but 770—1,293 m based on field data of AW).

ETYMOLOGY.—From the Yindjibarndi word yiwarda, meaning ashes, in reference to the grayish
color of the species.

COMPARISONS AND REMARKS. — Trachonurus yiwardaus 1s readily distinguished from other
Western Australian Trachonurus by its relatively small scales, heavily scaled gular and branchiostegal
regions, bluntly rounded snout profile, enlarged outer premaxillary teeth, presence of a grooved lateral
line, and a gray overall color. The first four characters separate the species from 7. sentipellis, the last
two separate it from 7. gagates.

In the smaller of two specimens from off Cape Cuvier (AMS I.31157-007), the flexible spinous
dorsal fin ray had about nine small serrations along the leading edge. This is a most unusual
occurrence, as one of the primary diagnostic characters of the genus is the presence of a smooth
spinous dorsal fin ray. There is no question in our minds, however, that the specimen represents 7.
yiwardaus.

IWAMOTO AND WILLIAMS: GRENADIERS OF WESTERN AUSTRALIA 219

One specimen (CSIRO H3002-05) listed as an uncertain variant had only a small gular scale
patch and no scales on the branchiostegals, relatively high gill raker counts (13 on inner series of first
arch, 12 on both sides of second arch), narrow bands of jaw teeth (about three teeth wide at symphysis
in premaxillary and dentary), short barbel (6% of HL), and was quite dark (swarthy to blackish). It
was captured along with a specimen of 7. gagates, a species with which it agrees closely except for
its grooved lateral line. Another questionable specimen (from northwest of Monte Bello Point, NTM
S.12716-010) had finer spinules with flexible tips on body scales, fewer scales on the gular membrane,
slightly wider head (width equal to postorbital length plus half orbit diameter cf. slightly more than
postorbital length in 7. yiwardaus), posterior margin of preopercle inclined anteroventrally (cf. chord
of margin essentially vertical), and only seven nublike pyloric caeca.

Ventrifossa Gilbert and Hubbs, 1920

See Iwamoto and Merrett (1997), Sazonov and Iwamoto (1992), and Iwamoto (1990) for a
diagnosis of genus. The 20 or so species in the genus are readily allocated to this taxon, but
circumscribing the genus has been difficult because of the absence of any recognized synapomorphy.
A good field character that is not usually apparent in formalin-preserved specimens is the silvery
reflections along the sides of head, trunk, and tail in most species. This silvery pigmentation obscures
much of the black markings on the head (notably on suborbital and operculum) and trunk in fresh
specimens. The blackish dorsal leading margin of the snout coupled with the relatively thin head
covering are distinctive to Ventrifossa and Hymenocephalus.

All six Australian species of the genus are found in Western Australia. Ventrifossa nigrodorsalis
is widespread in the central western Pacific and appears to be common off the northern and eastern
shores of Australia, including Queensland and New South Wales. Ventrifossa johnboborum is found
off the east and west coasts of Australia and has been reported from the southeastern Pacific (Sazonov
and Iwamoto 1992). Ventrifossa macropogon Marshall, 1973 was first recorded from Australian
waters, although it has been recently reported from the New Caledonian region (Iwamoto and Merrett
1997). Ventrifossa sazonovi n.sp. 1s recorded from WA, QLD, and the South China Sea; V. paxtoni
n.sp. from NSW, QLD, WA, New Caledonia, and the Chesterfield and Bellona Plateau; and V. gomoni
n.sp. appears to be confined to WA.

KEY TO THE AUSTRALIAN SPECIES OF VENTRIFOSSA

[Fae SeCOnd SpinOUSimayO1Minstid OrsaliinesimOOth\eme sere eee lee ieee memes eaten rien enya: V. sazonovi n.sp.
Ib. Second spinous ray of first dorsal fin with finely serrated leading edge ...................---28. 2

2a. Lateral line scales over distance equal to predorsal length of head 65—75; suborbital shelf narrowly constricted

anteriorly; a small but distinct tubercle-like scale at apex ofsnout.................... V. johnboborum
2b. Lateral line scales less than 51 over distance equal to predorsal length; suborbital shelf not especially narrow at ante-
Monendanordistinct terminal: snoutitubercl eke, emewemrte meme ice el career ten cece came eee-eeenr-w eenS
3a. A prominent black blotch or streak across first dorsal fin; pelvic fin rays 8 or 9, usually 8; no dark longitudinal streak
over median nasal ridge; chin barbel moderate, 16-27% ofheadlength................. V. nigrodorsalis
3b. No prominent blotch or streak on first dorsal fin, but fin may be dark proximally, paler distally; pelvic fin rays 8-10;
dark streak over median nasal ridge present or absent; chin barbel 22-45% ofheadlength............... 4

4a. Outer premaxillary teeth notably enlarged, distinctly larger than teeth of inner band; first dorsal fin pale to light dusky,
darker proximally; a prominent dark median nasal streak; snout low, preoral length 9—14% of head length; preserved
specimens generally pale, with some blackish or dark regions on body and head, suborbital shelf with thin dark
margin, dark blotches on gill cover limited; pelvic fin rays 8-9, usually8................ V. gomoni n.sp.
4b. Outer premaxillary teeth scarcely larger than teeth of inner band; first dorsal fin pale to blackish; dark median nasal
streak present or absent; preoral length 11-19% of head length; overall color usually dark with substantial blackish ar-
eas, entire suborbital shelf blackish, gill cover extensively covered with dark markings; pelvic fin rays 8-10 .... . 5

220 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 3

5a. Chin barbel notably long, often thick, 34-45% of head length; first dorsal fin pale or dusky, sometimes proximal

Wao Siohantenorrourrays ang. Spines ark 245 5 bre eee odes 4 ess eae gap swage a nels) gees V. macropogon
5b. Chin barbel moderate length, usually thin, 25-38% of head length; first dorsal fin uniformly blackish, paler on
[HEISE WA SOMME 9 oo -o 6 BUS Jegiaeeeaae Mina aetalsiaet tps ak arma i Pear Pa erat Mat Bad a V. paxtoni ti.sp.

Ventrifossa gomoni new species
Fig. 51

No literature applies to this species.

DIAGNOSIS. — Pelvic fin rays 8—9; first dorsal fin dusky proximally, paler distally, no distinct
black blotch or streak; a dark streak along leading edge of snout, with a median extension atop snout;
barbel 22-35% of head length, about 75—130% of orbit diameter; orbit diameter 1.3—2.1 into
postorbital length of head; outer premaxillary teeth distinctly enlarged, mandibular teeth in two
irregular rows; ground color of body in preserved specimens light brown, dorsum of trunk and tail
not strongly demarcated from lateral and ventral surfaces; suborbital shelf broad, not constricted
anteriorly, blackish streak covers shelf broadly or narrowly, but confined along dorsal portions.

SPECIMENS EXAMINED. — HOLOTYPE: AMS I.23425-003 (61.1 HL, 300+ TL); WA, NW Shelf;
400 m; SO4/82/leg1. PARATYPES: WA: AMS 1.23423-012 (5, 34.1-50.1 mm HL, 130+—260+ mm
TL); NW Shelf; 376 m; SO4/82/leg1. AMS 1.23425-003 (4, 26.2-46.9 HL, 145+—222+ TL); same
data as for holotype. AMS 1.24449-001 (4, 45.1—about 53 HL, 245+—270 TL); NW Shelf; 450 m;
SO1/84/12. CAS 200225 (formerly NTM S.12588-016)(54.8 HL, 260+ TL); NW Shelf off Rowley
Shoals; 17°22'S, 118°38’E; 430 m; coll. W. Houston, WH-85-15; 2.X1.1985. NTM S.12590-010 (4,
44.3-48.6 HL, 226+—253+ TL); NW Shelf off Rowley Shoals; 17°23'E, 118°57’'E; 430 m; coll. W.
Houston; WH-85-17; 3.X1.1985. NTM S.12614-016 (3, 25.4—52.4 HL, 135—260+ TL); NW Shelf off
Rowley Shoals; 17°39’S, 118°38’E; 410 m; coll. W. Houston; WH 85-33; 7.X1.1985. NTM S.12727-
022 (2, 46.0-S3.0 HL, 197+—260 TL); sw. of Rowley Shoals; 17°52’S, 118°28’E; 410 m; 9.1I.1990.
CSIRO H1514-37 (5 spec.); nw. of Port Hedland; 18°08.8'S, 117°54.5’E; 582 m; SO5/88/190; 1988.

COUNTS AND MEASUREMENTS (24 spec.).—1D. II,9—11; P. (115) 118421; GR-I (outer/inner)
9—11/(13)14—15, GR-IT 13—15/13—14.

Total length 150+—270; HL 26.2—61.1 mm. The following in percent of HL: snout 25—31; preoral
9—14; internasal | 7—23; interorb. 23—28; orb. 24—34; suborb. 1 1—13; postorb. (43) 45—54; orb-preop.
38-47% (usually 40-44%); up.jaw 42-48; gill slit 21-28; pre-A. 124-156; V.-A. 27-42; body depth
76-89; 1D.-2D. 45-79; ht. 1D. 68-78; len. P. 49-61; len. V. 34-41; post. nostril 3—6.

DESCRIPTION. — General features of holotype shown in Figure 51. Snout low, short, pointed in
lateral view; blunt in dorsal view, width across lateral angles about equal to or slightly more than
interorbital width. Mouth large, upper jaw extends posteriorly to about vertical through hind margin
of orbit; barbel relatively thin and long.

Body scales highly deciduous, thin, covered with small, erect conical spinules in more or less
quincunx pattern. Spinuleless scales numerous (about 10—12) behind base of first dorsal fin, behind
pectoral fin, and along margin of gill cover.

Premaxillary teeth in broad band, with outer series large, recurved, and relatively widely spaced;
inner band teeth small. Mandibular teeth small, in about 2 irregular rows.

First dorsal fin origin well behind vertical through pectoral fin base, which in turn, lies behind
pelvic fin origin. Anal fin origin below about midbase of first dorsal fin. Outer pelvic fin ray scarcely
produced beyond other rays; barely extends to anal fin origin in some, to as much as 6th anal fin ray
in others.

Ground color in preserved specimens light to medium brown dorsally, tawny laterally and
ventrally; dark, somewhat violet colored over abdomen and chest; opercle and subopercle, lower

IWAMOTO AND WILLIAMS: GRENADIERS OF WESTERN AUSTRALIA 221

portion of preopercle, margins of ramus of lower jaw, gill and gular membranes dark; upper lips black,
lower lips generally dark but pale near angle of mouth; leading edge of snout with black margin, the
pigmentation extending onto suborbital shelf, but not as intensely black as in other species (e.g., V.
nigrodorsalis, V. macropogon) and in some specimens confined to upper part of shelf; fainter blackish
margin along supranarial ridges and median nasal ridge (darker and more prominent in some
paratypes); a spear-shaped darker area on nape pointing forward from origin of first dorsal and
narrowly connected to a pair of small, dark projections at anterior end of nape; a faint dusky band
across anterior end of interorbital region. Mouth cavity immaculate except blackish along edge of
upper oral valve and small patch behind symphysis of lower jaw teeth; most ventral surfaces of gill
cavity, gill filaments, arches, and rakers pale, outer and upper margins blackish, but narrow edge of
branchial membranes pale; chin barbel dark only on base, remainder pale. First dorsal fin dusky
proximally, pale distally; second dorsal and anal fins pale throughout; pectoral fin pale to lightly
dusky, but sometimes darker near base; pectoral fin base, small area over axil, and lunate naked area
immediately behind fin blackish; pelvic fin black proximally, pale distally.

SIZE. — Attains at least 27 cm TL.

DISTRIBUTION. — The species is known only from the North West Shelf of WA, in 376-450 m.

ETYMOLOGY.—Named for our ichthyological friend and colleague Martin F. Gomon, curator in
the Museum of Victoria, Melbourne.

COMPARISONS AND REMARKS. — This pale-colored species and the dark-colored V. sazonovi are
easily distinguished from other Australian congeners by their relatively large outer series of premax-
illary teeth. In the other four Australian species, the outer series is scarcely larger than the inner band
of teeth. Ventrifossa gomoni can be distinguished from V. sazonovi by its serrated (as opposed to
smooth) spinous dorsal ray, among other characters. In addition to the teeth character, V. gomoni
differs from: V. nigrodorsalis in lacking the distinct black streak or spot on the first dorsal fin, and
having a well-marked median nasal streak, and a lower snout (preoral length 9-14% HL compared
with 14-23%); V. macropogon in having a lower snout (preoral length 9-14% HL cf. 13-17%), a
shorter barbel (22-35% HL cf. 34-45%), a somewhat narrower internasal width (17-22% HL cf.
21-25%), and a paler overall color; from V. paxtoni in having a narrower suborbital (1 1—13% HL ef.
13-17%), lower gill raker counts on second arch (13—14 total on inner side cf. 15—18), and a long,
well-marked median nasal streak (cf. short and faint, or lacking). Three fairly recently described
species differ from V. gomoni in the following: V. /ongibarbata has a distinct black blotch on the first
dorsal fin, a higher snout, a somewhat larger orbit (31-38% HL cf. 24-34%), a shorter postorbital
length (39-44% HL cf. 43-54%), and more pectoral fin rays (“23—25” fide Okamura, 1982:157, cf.
118-121); V. rhipidodorsalis has a distinct black blotch on first dorsal fin, a black distal margin on
anal fin, no median nasal snout streak, and slightly higher gill-raker counts (inner GR-II 15—16 total
cf. 13-14 in the new species); V. saikaiensis has an overall dark first dorsal fin, lacks markedly
enlarged outer premaxillary teeth, lacks a blackish leading snout margin and median snout streak, has
a somewhat broader interorbital (27-31% HL cf. 23-28%), somewhat longer upper jaw (46-52% HL
cf. 42-48%), and prominently marked scale pockets on body.

Ventrifossa johnboborum Iwamoto, 1982
Figns2

Ventrifossa johnboborum Iwamoto, 1982:55—61, fig. 1 (Bismarck Sea). Williams et al., 1996:149 (WA list).

DIAGNOSIS. — Head broad, interorbital width 28-33% HL, less than orbit diameter; upper jaw
39-45% HL; barbel 13-19% HL; scales small, 65—71 lateral line scales from origin over distance
equal to predorsal length; spinous ray of first dorsal fin weakly serrated; V. 8-10; suborbital shelf
narrowly constricted at anterior end; mouth blackish.

222 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 3

ZZ
Z

a

Figure 51. Ventrifossa gomoni n.sp. Holotype, AMS 1.23425-003 (61.1 mm HL, 300+ mm TL), off North West Shelf, WA,
in 400 m. Scale bar represents 25 mm. (a) Lateral view. Fins partially reconstructed. (b) Dorsal view of snout showing extent
of dark pigmentation along ridges. Scale bar represents 25 mm.

FiGurE 52. Ventrifossa johnboborum |wamoto. (From Sazonov and Iwamoto 1992, fig. 30.)

SPECIMENS EXAMINED. — WA: CSIRO H1492-11 (1 spec.); NW Shelf, w. of Barrow I.; 420 m,;
SO5/88/70. CSIRO H2542-23 (1 spec.); Exmouth Plateau; 854-868 m; SS1/91/03. CSIRO H2553-07
(1 spec.); w. of Point Cloates; 910 m; SS1/91/14. CSIRO H2572-09 (1 spec.); w. of Dirk Hartog I.;
874 m; SS1/91/35. CSIRO H2573-22 (2, 140-185 mm TL)); off Shark Bay; 690-691 m; SS1/91/36.
CSIRO H2580-[11 through 16); NW Shelf, sw. of Shark Point; 420 m; SS1/91/43. CSIRO H2584-[9
through 13]; w. of Shoal Point; 853 m; SS1/91/49.

SIZE. — Attains about 48 cm TL.

i)
NO
Ww

IWAMOTO AND WILLIAMS: GRENADIERS OF WESTERN AUSTRALIA

DISTRIBUTION. — Widely distributed in tropical waters of the southern hemisphere, from the
Indian Ocean coast of southern Africa, east to Australia and into the Pacific, from Australia (WA,
TAS, NSW, QLD) and New Caledonia in the west, to the Sala y Gomez Ridge in the east. Off WA,
it has been taken between longitudes 21°49’S and 28°06’S. Depth range 540—850 m (Iwamoto and
Merrett 1997).

COMPARISONS AND REMARKS. — Ventrifossa johnboborum is a well-marked species that is
unlikely to be mistaken for any other grenadier in the West Australian region except as juveniles.
Ventrifossa fusca Okamura, 1982 and V. misakia (Jordan and Gilbert, 1904) are closely related, but
can be differentiated by characters provided in Iwamoto (1990:305) and Sazonov and Iwamoto (1992:
78-81). Iwamoto and Merrett (1997:85) noted differences in specimens from New Caledonia
compared with the holotype and material from the Sala y Gomez Ridge.

Ventrifossa macropogon Marshall, 1973
Fiss53

Ventrifossa macropogon Marshall, 1973:658-660, fig. 5 (holotype, USNM 198187, nw. Caribbean, 576 m).
Williams et al., 1996:149 (WA list). Iwamoto and Merrett, 1997:85—87, fig. 35 (New Caledonia region;
675-833 m).

DIAGNOSIS. — Pelvic fin rays 8—10 (usually 8 or 9); first dorsal fin lacking distinct black blotch;
a faint to prominent dark median nasal streak; chin barbel usually thick, long, 34-45% of head length;
preoral length 13-17% of head length; dorsal parts of trunk and tail darkly contrasted and sharply
demarcated from paler lateral and ventral parts of body; suborbital shelf broad with no anterior
constriction, entirely pigmented in black.

SPECIMENS EXAMINED. — WA: CSIRO H2549-09 (46.2 HL, 242+ TL); w. of NW Cape; 650
m; SS1/91/10. AMS 1.31166-009 (5 of 6, 26.8-39.1 HL, 150+—223+ mm TL); off Shark Bay; 610-612
m; SS1/91/31. AMS 1.31170-006 (5, 44.4—50.0 mm HL, 210-240 mm TL) and CSIRO H2573 (1 of
5, 48.5 HL, 240+ TL); off Shark Bay; 690-691 m; SS1/91/36. CSIRO H3034-03 (39.8 HL, 195+ TL)
and H3034-11 (46.6 HL, 247+ TL); w. of Shoal Point; 28°06’S, 113°27’E; 649 m; Akebono Maru
No. 3, shot 34: coll. A. Williams; 27.XII.1989.

COUNTS AND MEASUREMENTS (15 WA spec.).—1D. II,9—11; P. 118-124 (usually 120-121); total
GR-I (outer/inner) 9—12/13—15, GR-II 13—14(16)/12—15; scales 1D. about 8.5—1 1, midbase 1D. 6-8.5,
2D. 7.5—9.5, lat.line about 40-44.

Total length 150+—253+ mm; HL 26.8—50.0 mm. The following in percent of HL: snout 28-32;
internasal 21—25; interorb. 25-28; orb. 30-36; suborb. 12—15; postorb. 41—46; orb.-preop. 39-45;
up.jaw 4346; gill-slit 22-28; pre-A. 126-155; V.-A. 26-40; body depth 74-95; 1D.-2D. 23-58; ht.
1D. 62—87; len. P. 52-61; len. V. 36-45; post. nostril 3—7.

DESCRIPTION. — General features seen in Figure 53. Snout barely protruding beyond mouth,
broadly rounded in dorsal view; width across lateral angles more than interorbital width, about equal
to orbit diameter and snout length. Upper jaw extends to below posterior 1/4th of orbit; barbel usually
longer than orbit diameter.

Body scales densely covered with small, erect, conical spinules in more or less quincunx pattern;
spinules black. Spinuleless scales under margin of gill cover, beneath pectoral and pelvic fins, and a
small patch behind first dorsal fin.

Jaw teeth all small, in broad band 6—7 teeth wide in premaxilla with outer series slightly larger
than teeth of inner band; mandibular teeth small, conical in 24 rows (usually 3 irregular rows), none
enlarged.

224 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 3

Fin positions as shown in Figure 53. Leading edge of spinous second ray of first dorsal fin finely
serrated, the ray not produced beyond adjacent segmented rays. Outer pelvic fin ray slightly
prolonged, extending to 6th or 7th anal fin ray.

Specimens variable in overall darkness, some with swarthy or dark areas more extensive than in
others. Dark dorsum of trunk and tail sharply demarcated from paler lateral and ventral surfaces, the
division being stronger posteriorly toward dark-dusky tail tip, but sometimes faint in midbody region.
Dorsum of trunk often darker than adjacent anterior and posterior regions; abdomen dark with bluish
tinge. Ventral surfaces of chest and abdomen blackish; gular and gill membranes black; operculum
mostly black; a broad, diagonal, blackish streak from posteroventral corner of orbit to angle of
preopercle; leading edge of snout with blackish margin extending broadly onto entire suborbital shelf,
with dark streaks over median nasal and lateral nasal ridges. Lips sharply delineated in black; lower
jaw black along margins, dark along ramus; barbel with base narrowly black but overall pale. Mouth
cavity immaculate except for dusky dorsal oral valve and dusky to blackish ventral oral valve and
gum; roof of gullet dark. Gill chamber blackish except for pale hyoid regions and outer margins; gill
rakers and arches pale except for light peppering on rakers. First dorsal fin pale in smaller specimens
(less than ca. 40 mm HL) but dusky to dark in larger specimens; fin often paler over distal and posterior
margins. Pectoral fin with narrow black basal margin, and pale to dark dusky overall; pectoral axil
coal black, a lunate black margin behind base. Pelvic fin black at base and proximally, but distally
dusky or pale. Anal fin pale in smaller specimens, dusky to blackish (especially anteriorly) in larger
specimens.

SIZE. — The largest specimen examined from Australian waters was 25 cm; from New Caledo-
nian waters 32cm. In the western Atlantic, the species is known to exceed 45 cm (personal
observations, TT).

DISTRIBUTION. — In Australia known only from WA west of North West Cape (ca. 21° S) south
to about 33°30’S, in 320—760 m. Specimens have also been examined from off New Caledonia (see
Iwamoto and Merrett 1997). Otherwise known only from waters of temperate and tropical western
North Atlantic.

COMPARISONS AND REMARKS. — The seemingly disjunct distribution of V. macropogon is
perplexing, although the presence of the species in Australian waters is in itself not surprising. Merrett
and Haedrich (1997) cite examples of several deep-demersal fishes found in North Atlantic and
Australian waters, but their distributions are continuously through the South Atlantic and Indian
Ocean. (The orange roughy, Hoplostethus atlanticus Collett, 1889 is an example.)

Based on distribution patterns and morphological variability in material collected from New
Caledonia (Iwamoto and Merrett 1997), it is conceivable that more than one species is involved, as
several recognized species of Ventrifossa appear extremely similar, and it is possible that we were
unable to detect differences that actually exist between our specimens. On the other hand, we may be
dealing with a single, widespread, highly variable species that frustrates our attempts at circumscrip-
tion.

Ventrifossa nigrodorsalis Gilbert and Hubbs, 1920
Fig. 54

Ventrifossa nigrodorsalis Gilbert and Hubbs, 1920:546—549, fig. 36 (holotype, USNM 83627, off n. coast of
Mindanao, Philippines, 391 m; paratypes from Formosa, Philippines, and East Indies [Borneo, Celebes] in
290-686 m). Williams et al., 1996:149 (WA list) Iwamoto and Merrett, 1997:87—90, fig. 36, 37a (New
Caledonia region; 545-855 m).

DIAGNOSIS. — Pelvic fin rays 8—9 (usually 8), color black with distal and mesial margins paler;
first dorsal fin with distinct black blotch or streak across anterior part of fin; no dark streak over

IWAMOTO AND WILLIAMS: GRENADIERS OF WESTERN AUSTRALIA

No
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median nasal ridge; barbel 16-27% of head length, less than orbit diameter, pale with dark base; orbit
diameter |.1—1.6 into postorbital length of head; outer premaxillary teeth slightly enlarged; dark dorsal
parts of trunk and tail strongly contrast the pale or silvery lateral and ventral parts of body; suborbital
shelf broad with no anterior constriction.

SPECIMENS EXAMINED. — WA: AMS 1I.31154-013 (3, 49.7—56.2 mm HL, 246+—295+ mm TL),
CSIRO H2554-09 (42.2 HL, 220 TL), and CSIRO H2554-11 (4, 48.8-54.3 HL, 230—-260+ TL); sw.
of Point Cloates; 482-554 m; SS1/91/15. AMS 1.22817-012 (6, 30.7-43.6 HL, 168—231+ TL); NW
Sheli-520im:; SO2/82/31—35. CSIRO: H1492-13 (i of 12 spec., 31.5 HL, 169° TL) and CSIRO
H1492-21 (2 of 12 spec., 38.7-48.4 HL, 206+—245+ TL); NW Shelf, s. of Barrow I.; 420 m;
SO5/88/70. CSIRO H1514-11 (38.4 HL, 205+ TL); NW Shelf; 582 m; SOS/88/190. QLD: AMS
1.23613 (50.6 HL, 180+ TL); off Marion Reef; 19°06’S, 152°30’E; 650 m; 1987. AMS 1.20661 (3 of
8, 34.8-38.7 HL, 202+—223+ TL); e. of Murray Isles; 9°53’S, 144°23'E; 480 m; Queensland Fishery
Survey; 28.V.1983. AMS 1.20518-018 (2, 41.5-45.4 HL, 235+—250 TL); e. of Gold Coast; 548 m;
K78-09-05. AMS I.21795-005 (2, 36.0-43.1 HL, 201+—240+ TL); ne. of Danger Point; 731 m;
K78-23-08. AMS 1.20459-027 (4, 37.9-45.0 HL, 20+—228+ TL); e. of Danger Point; 548 m;
K78-17-10. AMS 1.32433-007 (46.9 HL, 251 TL); Papua Plateau, e. of tip of Cape York; 10°29.8’S,
144°01.4'E; 596-603 m; Franklin site 2; 20. VII.1992. NSW: AMS 1.15987-014 (31.1 HL, 203 TL);
30 miles ne. of Jervis Bay; 549 m; K71-11-09. AMS 1.21805-002 (5 of 25 spec., 28.0-38.8 HL,
175238 TL); se. of Newcastle; 586 m; K77-23-09. AMS 1.21669-004 (3, 35.9-37.0 HL, 167+—235+
TL); e. of Broken Bay; 603 m; K77-16-16. AMS 1.24852-009 (25.5 HL, 128+ TL); off Broken Bay;
512-530 m; K84-15-03. AMS 1.29535-005 (24.9 HL, 160 TL); e. of Sydney; 540 m; K79-15-01.

COUNTS AND MEASUREMENTS (41 spec.)—1D. I,9—11; P. 118-122 (123); GR-I (outer/inner)
8—12/13—16, total GR-II 13—15(16)/12—14; scales 1D. 7-10, midbase 1D. 5.5—7.5, 2D. 7.5—10, lat.line
39-42.

Total length 128+—295+ mm; HL 24.9-56.2 mm. The following in percent of HL: snout 26-33;
preoral 14-22; internasal 19-25; interorb. 24—30; orb. 29-35; orb.-preop. 38-45; suborb. 11—15;
postorb. 40-46; up.jaw 37-45; gill slit 20-27; pre-A. 125—162; V.-A. 27-40; body depth 75—95;
1D.-2D. 45—72; ht. 1D. 64—96; len. P. 46-77; len. V. 37-48; post. nostril 4-8.

DESCRIPTION. — General features shown in Figure 54. Snout short, conical in lateral profile,
barely protruding beyond large mouth; width across lateral angles more than interorbital width. Upper
jaw extends posteriorly to below middle of pupil or beyond; barbel moderately long and thin, length
less than orbit diameter.

Body scales thin, densely covered with small, erect, conical spinules in more or less quincunx
pattern; spinules often densely pigmented and blackish. Spinuleless scales under margin of gill cover,
beneath pectoral and pelvic fins, and behind first dorsal fin.

Small fine teeth in bands in both jaws; outer premaxillary teeth very slightly enlarged and not
prominent. Lower jaw band usually 3 or 4 teeth wide; premaxillary band 4-6 teeth wide.

Fin positions as shown in figure. Pectoral and pelvic fins originate about on same vertical; first
dorsal fin slightly behind that vertical; second dorsal fin beginning far behind first dorsal fin and
poorly developed throughout. Outer pelvic fin ray slightly prolonged, extending at most to bases of
first few anal fin rays.

Formalin-fixed specimens generally lack silvery guanine layer; dorsum dark, strongly contrasted
with pale lateral and ventral surfaces of body; color darkens, punctations increase toward tip of tail.
Leading edge of snout with broad black margin that joins black suborbital shelf; supranasal ridges
with some blackish margins, but only a trace over midorbit, a faint trace of a dark median nasal streak
in a few specimens, but absent in others. In fresh specimens silvery reflections obscure underlying
black pigmentation; silvery layer covers suborbital, gill covers, and lower 2/3 of trunk and tail. Lips
black; skin covering premaxillary and maxillary bones generally pale or lightly punctate; mouth lining
white; upper oral valve lightly peppered; gums of dentary dusky, that of premaxillary pale to dusky;

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226

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IWAMOTO AND WILLIAMS: GRENADIERS OF WESTERN AUSTRALIA

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228 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 3

gill arches and filaments generally pale, but rakers dark; chin barbel dark at immediate base but pale
distally. First dorsal fin spot or streak usually coal black and distinctive; spot extended as a broad
streak in many specimens, beginning on serrated spinous ray and fading beyond Sth or 6th ray. Pelvic
fins with inner distal portion pale or lightly dusky, with other parts black. Pectoral fin pale to dark
dusky overall, darker on base; a lunate area of naked black skin behind base. Anal fin clear to lightly
dusky but darker near anterior end.

SIZE.— To about 30 cm TL.

DISTRIBUTION. — Known from off Australia (WA, QLD, NSW), New Caledonian region, and
Indonesia north to the Philippines and South China Sea, in 290—759 m. Distribution in WA confined
to NW Shelf (about 18°S, 117°E) and coasts south to off Point Cloates (about 23°S).

COMPARISONS AND REMARKS. — Ventrifossa nigrodorsalis is one of five closely related species
that have a distinct black blotch in the middle of the first dorsal fin; these include V. petersoni (Alcock,
1891), V. longibarbata (Okamura, 1982), V. rhipidodorsalis Okamura, 1984, and V. ctenomelas
(Gilbert and Cramer, 1897). Ventrifossa petersoni appears to be most similar to V. nigrodorsalis but
differs in having a somewhat shorter preoral length (11-15% of HL cf. 14-22%), a smaller orbit
(25-28% HL cf. 29-35%), and a longer postorbital length (orbit |.7—2.0 into postorbital). Ventrifossa
rhipidodorsalis differs in having an anteriorly black-edged anal fin and on average more pelvic fin
rays (9 or 10). Ventrifossa longibarbata has a longer barbel (more than orbit diameter) and smaller
body scales (9 or 10 below midbase of first dorsal fin, compared with about 5.5—7.5 in V. nigrodor-
salis). Ventrifossa ctenomelas from Hawaii is readily distinguished by its larger outer premaxillary
teeth, more rays in first dorsal fin (II,12—13), and smaller scales (1 1—12 scale rows below first dorsal
fin origin, 50—S3 lateral line scales over distance equal to predorsal length).

This species complex deserves closer study, owing to the close similarity of the species and the
subtle characters that are used to differentiate one from another. Color patterns often differ among
individuals of the same species, and reliance on such characters to distinguish species may prove
unreliable. Variation in the relative intensity and the shape of the dark blotch on the first dorsal fin in
members of V. nigrodorsalis and V. rhipidodorsalis (see Okamura in Okamura and Kitajima, 1984)
lend an air of uncertainty in using such characters for species determinations.

Ventrifossa paxtoni new species
Figs 55

Ventrifossa species (NSW) Iwamoto and Merrett, 1997:92, 93, fig. 33b (New Caledonia region; 760—970 m).
Ventrifossa sp. B: Williams et al., 1996:149 (WA).

DIAGNOSIS. — Pelvic fin rays 9-10 (usually 9), fins all black; median nasal streak faint or absent;
chin barbel slender, moderately long, 24-35% (rarely to 38%) of head length; outer premaxillary teeth
small, scarcely enlarged; darker dorsum not strongly demarcated from paler lateral and ventral parts
of trunk and tail; suborbital shelf broad, without anterior constriction, almost entirely black.

SPECIMENS EXAMINED. — HOLOTYPE: AMS I[.27647-002 (58.4 mm HL, 317+ mm TL); NSW
off Sugarloaf Point; 880-950 m; K87-16-02. PARATYPES: NSW: AMS 1.27647-002 (57.3 HL, 306+
TL); same data as for holotype. AMS I.28189-005 (2, 49.2—-59.0 HL, 232+—235+ TL); e. of Newcastle;
960 m; K87-24-03. AMS 1.24990-001 (52.4 HL, 294+ TL); off Sydney; 927-955 m; K84-16-13.
AMS 1.29753-008 (4, 26.7-47.4 HL, 145+—255+ TL); e. of Crowdy Head; 878-933 m; K89-17-06.
AMS I.29805-001 (55.2 HL, 259+ TL); e. of Hawks Nest; 885-950 m; K87-24-04. AMS 1.2698 1-005
(5 of 6, 54.9-57.7 HL, 252-310 TL); e. of Wallis Lake; 900-960 m; K87-02-01. QLD: AMS 1.15822
(2, 62.8-68.6 HL, 295+—370+ TL); 10-11 miles ne. of Raine Island; 900 m; 12.11.1979. New
Caledonia: CAS 86476 (56.4 HL, 305+ TL); 23°07.26’S, 166°50.45’E; 850 m; BIOCAL exped., st.
CP31; R/V Jean-Charcot; 29.VIII.1985. Chesterfield and Bellona Plateau: MNHN 1994-1010 (6,

IWAMOTO AND WILLIAMS: GRENADIERS OF WESTERN AUSTRALIA

in)
N
\O

FiGure 55. Ventrifossa paxtoni n.sp. Paratype, CAS 86476 (305+ mm TL) from off New Caledonia in 850 m. From Iwamoto
and Merrett 1997, fig. 33b. Scale bar represents 25 mm.

64.8-82.1 HL, 325+—380+ TL); MUSORSTOM 5 exped., st. CP384; 19°42.40’S, 158°50.80'E;
772-156 m; R/V Coriolis; 21.X.1986. BMNH 1996.7.19:38, 46-58 (4 spec.) and MNHN 1997-668
(33.2 HL, 164+ TL); MUSORSTOM 5S exped., st. CC337; 19°53.80'S, 158°38.00’E; 412-430 m; R/V
Coriolis; 15.X.1986. OTHER SPECIMENS (not paratypes): WA: CSIRO H2621-01 (480 TL); w. of
Bunbury; 780 m; SS1/91/90. CSIRO H2549-09 (43.3 HL, 245 TL) and H2549-16 (305 TL); w. of
NW Cape; 650 m; SS1/91/10. CSIRO H3145-26 (4, 170-220 TL); Rowley Shoals area; 17°52’S,
HiSei6de: 550m; 1992:

COUNTS AND MEASUREMENTS (24 spec.; see also Diagnosis).—1 D. I,9—1 1 (usually I,10 or 11);
P. 120-125 (usually 120-123); total GR-I (outer/inner) 10—13/15—18, GR-II 14—18/15—18; scales 1D.
8—12, midbase 1D. 5.5—8.5, 2D. 7.5—9.0, lat.line 37—SO.

Total length 145+—380+ mm; HL 26.5—87.5 mm. The following in percent of HL (exceptional
measurements enclosed in parentheses): snout (24) 26-31; preoral 10—19; internasal 20—26 (29);
interorb. 24-30; orb. 27-34 (38); suborb. 13—17; orb.-preop. 42-48; postorb. 43-52; up.jaw 43—S0;
gill slit 24-30; pre-A. 122-164; V.-A. 27-45; body depth 77-96; 1 D.-2D. 38-61; ht. 1D. about 63-73;
len. P. about 50-65; len. V. about 40; post. nostril 3-6.

DESCRIPTION (of holotype, paratypes in parentheses unless otherwise stated)—Head and trunk
moderately compressed; head about 5 times in total length; body depth about 0.8 into head length,
more than postrostral length. Snout blunt in holotype, almost rounded in a few larger paratypes,
relatively lower in larger than in smaller specimens, especially those 60 mm or more in head length,
slightly extending beyond large subterminal mouth. Upper jaw extends posteriorly to below hind
margin of orbits. Orbit circular (to slightly oval), diameter slightly more than (about equal to) snout
length and interorbital width, 3.4 (2.73.7) into head length (size dependent, largest specimens have
proportionally smallest orbit). Suborbital region almost vertical, without strong or sharp ridge;
suborbital shelf broad, not narrowly constricted under anterior end of orbit. Opercular bones as for
most others of genus, as described for V. sazonovi.

Teeth all small, rather blunt with low flanges below tip; in broad band in premaxillary with outer
series scarcely enlarged; in 2 or 3 irregular series in mandible.

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Body scales covered with small, conical, sharp, reclined, slightly recurved spinules arranged in
subparallel (somewhat divergent or convergent) rows (about 15 or 16 rows, with posteriormost
spinules on exposed field somewhat larger than those preceding), rows short, about 4 to 6 spinules
per row. Spinules dorsally on trunk and tail, nape, and especially those remaining on chest all black
or black at base.

Height first dorsal fin less than postrostral length of head, spinous second ray finely serrated, not
prolonged beyond adjacent segmented rays. Second dorsal fin poorly developed throughout length;
anal fin well developed. Pelvic fin short, outer ray scarcely produced, barely, if at all, reaching anal
fin. First dorsal fin origin behind vertical through pectoral fin origin; origin pectoral fin slightly behind
that of pelvic fins. Anal fin origin below midbase of first dorsal fin.

Color in alcohol overall rather dark, swarthy in NSW and QLD specimens, but much paler in
WA specimens. In holotype and NSW specimens, entire head, chest, and abdomen swarthy, but WA
specimens more pale. Blackish margins of snout, supranarial ridges, suborbital shelf prominently
black in NSW specimens, much paler in WA specimens. Barbel blackish at and near base, pale distally.
Lips black, mouth cavity almost entirely pale but upper oral valves lightly peppered (blackish), and
gums dusky; roof of gullet dark. Gill chamber walls black, with contrasting white areas over hyoids,
gill filaments, and to some extent, isthmus. Gular and branchiostegal membranes dark to black. First
dorsal and pelvic fins black, pectoral fin black at base, blackish to dark dusky more distally. Anal fin
intensely blackish anteriorly at base, dark dusky over most of length, becoming swarthy to blackish
near end of tail (paler in these features in WA specimens).

SIZE. — To at least 43 cm TL.

DISTRIBUTION. — So far known only from Australia (NSW, QLD, WA), New Caledonia, and
the Chesterfield and Bellona Plateau, in 878—1,053 m.

ETYMOLOGY.—Named in honor of John R. Paxton, former curator at the Australian Museum,
Sydney, for his many contributions to ichthyology.

COMPARISONS AND REMARKS. — Of the Australian species, V. paxtoni appears most closely
similar to V. macropogon, but that species has a distinctly longer and thicker barbel (34-45% HL)
and its median nasal dark streak is much more prominent. In addition, gill-raker counts are notably
higher in V. paxtoni. Ventrifossa gomoni bears some resemblance to V. paxtoni, but it has a somewhat
narrower internasal (1 7—23% HL cf. 20-26%, exceptionally 29% in V. paxtoni), a narrower suborbital
(11-13% cf. 13-17%), generally fewer gill rakers, and a more prominent median nasal streak.

Among western North Pacific species, V. paxtoni appears to be most similar to V. saikaiensis
Okamura, 1984 in having a completely dark first dorsal fin, large mouth, small premaxillary teeth,
relatively numerous gill rakers, and similar proportions and counts. The chin barbel of V. saikaiensis,
which, according to Okamura (in Okamura and Kitajima 1984:209) is always more than orbit
diameter, appears to be somewhat longer than in V. paxtoni, in which the barbel is usually shorter
than the orbit, but varies from 0.84 to 1.29 into orbit. Four paratypes of V. saikaiensis were made
available to us by Dr. Okamura. In three of these four (BSKU 28005, 28007, 28008), the pelvic fin
ray counts were 8 on each fin, but in the fourth (BSKU 28006), the left side had 8, the right 9. Okamura
recorded 8 and 9 in the holotype, with a range of 8—9 in the paratypes. Ventrifossa paxtoni has 9 or
10 pelvic rays, usually 9. The outer gill raker counts on the first arch were slightly higher in V.
saikaiensis (12—15 cf. 10-13), and the count of scale rows below the second dorsal were slightly lower
(7.07.5 cf. 7.5—10). One notable pigmentation difference between the two species was the absence
in V. saikaiensis of dark border markings on the leading edge of the snout, over the supranarial ridges,
and on the median nasal ridge. These markings are prominent in V. paxtoni.

The Western Australian specimens are tentatively assigned to this species, although their overall
paleness and certain count and measurement differences suggest that they might not be conspecific
with V. paxtoni. In general, there was good overlap in most counts and measurements, but the WA
specimens often were at one end or another of the ranges and occasionally extended them marginally.

IWAMOTO AND WILLIAMS: GRENADIERS OF WESTERN AUSTRALIA We)

It is apparent that additional specimens from WA must be compared with material from other areas
of its range to confirm their identification.

Ventrifossa sazonovi new species
Fig. 56

Ventrifossa species 1: Iwamoto, 1990:293 (in key).
Ventrifossa species: Sazonov and Iwamoto, 1992:83, fig. 33 (in key and scatter diagram.)
Ventrifossa sp. A: Williams et al., 1996:149 (WA).

DIAGNOSIS. — Pelvic fin rays 9-10; second spinous ray of first dorsal fin fin with smooth leading
edge; a well-marked median nasal streak; chin barbel dark, length 17-24% of head length; outer
premaxillary teeth notably larger than teeth of inner band; dorsum not strongly demarcated in color
from rest of body in preserved specimens; suborbital shelf narrows anteriorly, somewhat constricted
at anterior end.

SPECIMENS EXAMINED. — HOLOTYPE: CAS 13564 (male, 63.6 mm HL, 297+ mm TL); South
China Sea off Vietnam; 15°48’N, 109°47'E; 479 m; MV Stranger sta. 60-63 (Naga 60-216; GVF
2079); 28.11.1960. PARATYPES: WA: AMS 1I.31166-006 (6, 38.2—71.5 mm HL, 200-330 mm TL);
off Shark Bay; 610-612 m; SS1/91/31. NT: NTM S.12457-003 (47.9 HL, 240+ TL); Arafura Sea,
se. of Tanimbar I.; 8°36’S, 132°00’E; 525-540 m; coll. M. Sachse; 1.VII.1988. QLD: CSIRO
H1973-01 (44.2 HL, 220+ TL); Marian Plateau, e. of Townsville; 879-886 m; SO6/85/36. SOUTH
CHINA SEA: CAS 99817 (7, 25.4—-57.7 HL, 136—274+ TL); data as for holotype.

COUNTS AND MEASUREMENTS (21 spec.)—1D. II,9—10; P. 119-125; total GR-I (outer/inner)
10—15/15—20, GR-II_ 14—18/15—19; scales midbase 1D. 7—8, 2D. ca. 7.5—8.5.

Total length 132+—-340+ mm; HL 21.9—71.5 mm. The following in percent of HL: snout 26-29;
preoral 6—12; internasal 19-25; interorb. 22—28; orb. 25—35; suborb. 11—14; postorb. 45—51; orb.-
preop. 43-46%; up.jaw 45—S0; gill slit 23-29; pre-A. 128-157; V.-A. 26-44; body depth 71-83;
1D.-2D. 41-60; ht. 1D. 56—74; len. P. 42-65; len. V. 32-39; post. nostril 2-6.

DESCRIPTION (of holotype, paratype data as for holotype or in parentheses).—Head and trunk
compressed, width over pectoral fin bases about 0.6, head about 0.7 greatest body depth. Greatest
depth somewhat less than postrostral length of head, 6.5 (6.0—6.5) in total length. Snout low, blunt,
scarcely protruding beyond large subterminal mouth, about equal to (slightly shorter than) interorbital
width. Orbit almost circular, diameter greater than snout length and interorbital width, 3.3 (2.8-3.6)
in head. Mouth large, upper jaw extends about to vertical through hind margin of orbit, length 2.1
(2.02.2) in head length. Barbel slender, tapering to fine tip, 0.58 (0.56—0.73) orbit diameter.
Interorbital region essentially flat. Suborbital region vertical, gently rounded, without strong or sharp
ridges; suborbital shelf slightly narrowed anteriorly. Preopercular ridge distinctly delineated; hind
margin of preopercle inclined slightly forward. Posterior and ventral margins of interopercle exposed
beyond preopercle. Ridges of gill cover distinctly incised at junction of opercle and subopercle;
subopercle terminating in a blunt, triangular point. Gill membranes unite under hind one-third of orbit,
narrowly attached to isthmus; a narrow free posterior fold.

Teeth in upper jaws in long, moderately broad band that tapers posteriorly to a narrow point at
end of rictus; outer series of enlarged wide-spaced recurved teeth with slightly flanged tips (arrow-
head-shaped); inner teeth minute. Mandibular teeth short, stoutly conical, in 2 irregular series, outer
series smaller than inner series, scarcely exposed above gum papillae.

Body scales deciduous, mostly missing in specimens examined. In holotype, those on dorsum
below second dorsal fin progressively beset with increasingly stout, long spinules (Fig. 56a’). Near
origin of second dorsal fin, spinules fine, conical, slightly reclined, and bespeckled with large
melanophores, an occasional midlateral spinule being larger than the remainder. Enlarged spinules

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232

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IWAMOTO AND WILLIAMS: GRENADIERS OF WESTERN AUSTRALIA 259

increase in size and number on more posterior scales. No scales remain on flanks. Nape scales covered
with uniformly small, conical, relatively erect spinules in typical (for genus) irregularly quincunx or
widely (and irregularly) divergent, V-shaped rows. Chest scales densely covered with spinules similar
to those on nape except that they are longer and more erect. No terminal or enlarged scutelike scales
on snout. Scales below second dorsal fin missing in paratypes smaller than about 40 mm HL; spinules
on these scales may not have been as enlarged as in larger individuals.

First dorsal fin relatively low, height much shorter than postrostral length of head, about equal
to or (usually) slightly longer than length pectoral fin. Second dorsal fin rudimentary to poorly
developed over most of length; 146 rays counted in 162 mm paratype with small rudimentary tail tip.
Anal fin of moderate height, well developed throughout; 140 rays counted in 162 mm paratype. Pelvic
fin short, outer ray barely, if at all, reaches anal fin. Origin of first dorsal fin behind vertical through
pectoral fin origin, which 1s behind that of pelvic fin. Anal fin origin below midbase of first dorsal
fin.

Color in alcohol light brownish overall, dark over most of head, abdomen, and chest. (South
China Sea specimens pale, possibly because they have been in alcohol for more than 30 years. Fresher
Australian paratypes generally darker.) Dorsum of trunk and tail slightly darker than surfaces
ventrally, but without sharp demarcation. Ridges of head mostly dark (notably so in Australian
specimens); leading margin of snout, suborbital shelf, median and lateral nasal ridges marked with
faint (prominent in many paratypes) blackish pigmentation. Lips, underside of head, operculum
mostly blackish; barbel pale (to dark); a faint (dark) oblique bar extending from posterior margin of
orbit towards angle of preopercle, but falling short of preopercle ridge. Distal surfaces of medial walls
of infraorbital bones blackish (visible through translucent skin). Mouth cavity white, roof of gullet
blackish; gums dark, upper oral valve blackish. Gill cavities dark with pale inner areas (as in other
species of genus); gill rakers and arches generally pale with tiny punctations (or dusky). First dorsal
fin dusky (darker proximally); pectoral fin dusky, axil blackish; pelvic fins black, with slight fading
near tips; anal fin dusky (anterior 1/5th or more dark dusky, turning paler posteriorly).

Periproct region as for others of subgenus. Anterior dermal window a small, circular, shallow
black fossa between pelvic fin bases, separated by a scaly bridge from periproct. Posterior dermal
window a black naked area immediately anterior to vent and forming anterior end of oval- to
teardrop-shaped periproct. About 74 slender, densely packed pyloric caeca, longest about equal to
length of barbel in 57.7 mm HL female paratype from South China Sea. Gonads in this specimen
large; eggs of various sizes, up to 1.3 mm in diameter.

SIZE.— To at least 34 cm TL.

DISTRIBUTION. — So far known only from WA, NT, and QLD in Australia, and the South China
Sea off Vietnam, in 420-612 m. Off WA it has been taken only off the NW Shelf (about 21°S) and
off Shark Bay (about 25°S).

ETYMOLOGY—-Named for our Russian colleague and fellow student of grenadiers, Yuri I.
Sazonoyv, curator of ichthyology in the Zoological Museum, Moscow State University.

COMPARISONS AND REMARKS.—VFentrifossa sazonovi 1s one of four members of the genus
having a smooth, second, spinous dorsal fin ray. The four species also share in common the character
of mandibular teeth in 1 or 2 irregular series, snout relatively low, gill raker counts relatively high,
and mouth large. The new species differs from the others in having enlarged spinules on scales below
the second dorsal fin in adults. In addition, V. atherodon (Gilbert and Cramer, 1897) from the
Hawaiian Islands differs in having somewhat larger orbits (32-40% HL cf. 25-35%); V. macrodon
Sazonov and Iwamoto, 1992 has a longer barbel (23-34% HL cf. 17-24%) and a somewhat shorter
postorbital length (39-47% cf. 45-51%); and V. macroptera Okamura, 1982 has longer pectoral fins
(3-15 in HU cf. 1.5-2.2).

234 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 3

SUBFAMILY MACROUROIDINAE

Squalogadus and Macrouroides Smith and Radcliffe, 1912, each with a single species, are the
only members of this peculiar group of grenadiers. There is some controversy as to whether or not
the group should be treated at the subfamily level or at the family level (see Cohen 1989), and it is
possible that the two species do not even belong within the Macrouroidei. Although Macrouroides
was not represented in WA, its presence there would not be surprising as it has a broad, generally
worldwide distribution (Philippines, Indian Ocean, South Atlantic, central western Atlantic).

Squalogadus Gilbert and Hubbs, 1916
Only one species, found circumglobal, except eastern Pacific.

Squalogadus modificatus Gilbert and Hubbs, 1916
Fig. 57

Squalogadus modificatus Gilbert and Hubbs, 1916:156 (Japan). Shcherbachev and Piotrovskiy, 1982:47 (world-
wide distr. plotted; 600—1,740 m). Shcherbachev, 1987:41 (Indian Ocean distr. including Mascarene Ridge,
West Australian Ridge, Exmouth Plateau; 800—1,740 m).

Squalogadus intermedius Grey, 1959:330 (Gulf of Mexico).

DIAGNOSIS. — Head massive, inflated, with rounded contours. Opercular cavity large, opercu-
lum extending well beyond gills; first gill slit unrestricted. Gill rakers on outer margin of first gill
arch numerous (20), styliform. Jaws relatively short; markedly inferior. One continuous dorsal fin,
anterior section not elevated. Pelvic fins small, with five rays.

SPECIMENS EXAMINED. — WA: CSIRO H2614-03 (110mm HL, 322+mm TL); 1280 m;
SS1/91/82. CSIRO H2544-01 (113 HL, 304+ TL); 1,128—1,139 m; SS1/91/04. CSIRO H2570 (77
HL, 198+ TL); 1,254—1,277 m; SS1/91/33.

COUNTS AND MEASUREMENTS (based on 3 spec.)—P. 122129; V. 5; total GR-I (outer/inner)
28-3 1/17—22, GR-II 16—19/17—19; scales 1D. 12-13.

Total length 198—322+ mm; HL 77-113 mm. The following in percent HL: postrostral 74-80;
snout 31—33; preoral 42-44; internasal 37-40; interorb. 38-46; orb. 11; suborb. 29-30; postorb.
64-68; orb.-preop. 62-64; up.jaw 26-33; premaxillary 24—27; pre-A. 121—125; pre-vent 119-122;
V.-A. 28-31; isth.-A. 41-43; body depth 64-67; depth over A. origin 42—S0; post. nostril 3; rictus
23-30.

SIZE.-— folat.least35 em TL.

DISTRIBUTION. — Widespread in tropical and southern temperate seas, but absent in eastern
Pacific.

COMPARISONS AND REMARKS.—An easily recognizable grenadier, separated from
Macrouroides inflaticeps, the only other representative of the subfamily, by the presence of pelvic
fins (absent in Macrouroides).

SUBFAMILY TRACHYRINCINAE
Two genera, Trachyrincus and Idiolophorhynchus. Only the latter has been found off WA, but

the former can also be expected as Gomon and others (1994) have recorded T. /ongirostris from off
Portland, VIC.

IWAMOTO AND WILLIAMS: GRENADIERS OF WESTERN AUSTRALIA

tN
i>)
Nn

Idiolophorhynchus Sazonov, 1981
Only one distinctive species, so far known only from off Australia and New Zealand.

Idiolophorhynchus andriashevi Sazonov, 1981
Fig. 58

Idiolophorhynchus andriashevi Sazonov, 1981:1,360—1,363, figs. 1, 2 (holotype ZMMSU P-15341; WA, e. of
Cape Leeuwin; 33°22’S, 113°59’E; 1,190—1,230 m). McMillan, 1995:84, 85, fig. 1 (descr.; 17 New Zealand
spec.; 1,030—1,484 m).

DIAGNOSIS. — Snout long, pointed, flattened dorsoventrally; mouth inferior; small teeth in bands
in both jaws; barbel absent. Gill rakers numerous; outer rakers on first arch long, tapering to fine tip;
remaining rakers relatively short, with broad rounded tips. Pelvic fin with 34 rays. Four longitudinal
rows of scutelike scales on each side; | dorsolaterally, 2 ventrolaterally, and 1 midlaterally; interven-
ing areas with weak, unspinulated scales or naked. Bony ridges of head smooth or with small
posteriorly directed spines. No temporal pit.

SPECIMENS EXAMINED. — WA: CSIRO H2623-01 (79.2 mm HL, 263 mm TL); off Cape Leeu-
win; 1,040—1,045 m; SS1/91/92.

COUNTS AND MEASUREMENTS (based on | spec.)—I1ID. 10; P. 117419; V. 3; total GR-I
(outer/inner) 26/17, GR-II 17/16; scales 1D. 8, 2D. 6.

Total length 263 mm; HL 79.2 mm. The following in percent of HL: snout 47; preoral 46;
internasal 26; interorb. 31; orb. 21; suborb.14; postorb. 35; orb.-preop. 29-30; up.jaw 15; outer gill
slit 9; pre-A. 150; pre-vent. 145; body depth 48; V.-A. 49; isth.-A. 67; len. P. 46; len. V. 53; post.
nostril 7.

DESCRIPTION. — Body elongate, tapering posteriorly to a point; greatest body depth at opercu-
lum. Snout long, sharply pointed, lacking terminal scute, flattened dorsoventrally; anterolateral
margins completely supported by bone; bony ridges of head smooth or with small, posteriorly directed
spines. Orbits small, more than twice into snout length; oblique, with notch in posteroventral margin.
Mouth small, inferior, rictus 2.3 times into preoral length, posterior margin of maxilla in line with
posterior margin of orbit. Barbel absent. Suborbital ridge poorly developed, narrow, bearing no
spinulated scales; a small cluster of scutelike scales where ridge extends to posterior margin of
operculum. Opercle and subopercle poorly developed; preopercle bearing a small, central bony keel.

Gill opening relatively wide, extending forward almost to posterior margin of orbit. Gill rakers
numerous; outer rakers on first arch long, flattened, and closely spaced, tapering to fine tip with fine
teeth on posterior margins; inner rakers on first arch and those on remaining arches relatively short,
stout, with broad, rounded tips bearing short, fine teeth.

Teeth small, in narrow bands in both jaws; tooth bands tapering gently to single row of teeth
posteriorly. Two dorsal fins of similar height separated by indistinct notch; second dorsal fin long,
rays more developed than those of anal fin. Pectoral fin moderately long, leading ray not prolonged.
Leading ray of pelvic fin fine, prolonged to origin of anal fin, inner ray fine and short.

Scales of 2 types: enlarged, adherent, scutelike with single, curved, central spine or small,
deciduous, weak, and unspinulated. Scutelike scales forming 4 longitudinal rows on each side; |
dorsolaterally, 2 ventrolaterally, and 1 midlaterally; additional row of scutelike scales curves between
pectoral fin base and isthmus; dorsolateral row extends from nape for about 50% TL; midlateral row
originates dorsal to pectoral fin base and about equal to postrostral length; long ventrolateral row
intersects pectoral/isthmus row just posterior to pelvic fin base and extends just posterior to dorso-
lateral row; short ventrolateral row extends from midpoint of isthmus and vent to anal fin origin; 1—2
scutes on inner margin of pelvic fin base. Areas between primary scale rows and entire head covered

236 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 3

FiGuRE 57. Squalogadus modificatus Gilbert and Hubbs. CSIRO H2614-03 (320 mm TL), from west of Rockingham, in
1,310 m.

FiGureE 58. /diolophorhynchus andriashevi Sazonov. CSIRO H2623-01 (79.2 mm HL, 263 mm TL) from off Cape Leeuwin,
WA, in 1040 m. Drawn by Georgina Davis.

with weak, unspinulated scales; these larger on head than on body; lost scales leave conspicuous and
distinctive pattern of scale-pocket margins on head and body; narrow naked margins adjacent to bony
head ridges.

Body color brown; skin in scale pockets pale; branchiostegal membranes dark brown; branchial
cavity and buccal cavity blackish brown. All fins light dusky brown.

SIZE.— To at least 26 cm TL.

DISTRIBUTION. — Southern Australia and New Zealand; 1,030—1,484 m.

COMPARISONS AND REMARKS. — /diolophorhynchus andriashevi is a distinctive grenadier and
is unlikely to be confused with other species. The species has been treated in a recent review of
trachyrincine grenadiers from New Zealand by McMillan (1995).

IWAMOTO AND WILLIAMS: GRENADIERS OF WESTERN AUSTRALIA

APPENDIX |. Station data for vessels Southern Surveyor, Kapala, and Soela.

FRV SOUTHERN SURVEYOR stations

SS1/91/02 [H2541]— WA:
SS1/91/03 [H2542] — WA:
SS1/91/04 [H2544] — WA:
SS1/91/05 [H2545] — WA:
SS1/91/08 [H2548] — WA:
SS 1/91/10 [H2549] — WA:
SS1/91/11 [H2550]— WA:
SS1/91/12 [H2551] — WA:
SS1/91/13 [H2552] — WA:
SS1/91/14 [H2553] — WA:
SS1/91/15 [H2554] — WA:
SS1/91/19 [H2557] — WA:
SS1/91/21 [H2559] — WA:
SS 1/91/23 [H2561]— WA:
SS1/91/25 [H2563] — WA:
SS1/91/31 [H2568] — WA:
SS1/91/33 [H2570] — WA:
SS1/91/35 [H2572] — WA:
SS 1/91/36 [H2573] — WA:
SS1/91/37 [H2574] — WA:
SS$1/91/41 [H2578] — WA:
SS$1/91/42 [H2579] — WA:
SS1/91/43 [H2580] — WA:
SS 1/91/44 [H2581] — WA:
SS 1/91/48 [H2583] — WA:
SS 1/91/49 [H2584] — WA:
SS1/91/57 [H2591] — WA:
SS 1/91/58 [H2592] — WA:
SS1/91/61 [H2595] — WA:
SS1/91/62 [H2596] — WA:
SS1/91/65 [H2599] — WA:
SS1/91/70 [H2604] — WA:
SS 1/91/71 [H2605]— WA:
SS1/91/72 [H2606] — WA:
SS1/91/73 [H2607] — WA:
SS1/91/82 [H2614] — WA:
SS1/91/83 [H2615]— WA:
SS1/91/84 [H2616] — WA:
SS1/91/85 [H2617] — WA:
SS1/91/87 [H2618] — WA:
SS1/91/90 [H2621] — WA:

SS1/91/92 [H2623] — WA
SS 1/91/96 [H2626] — WA

FRV KAPALA stations

Exmouth Plateau; 20°17.8’S, 113°12.3’E; 914 m; 1991.

Exmouth Plateau; 20°07.8'S, 112°56.3’E; 854-868 m; 1991.

Exmouth Plateau; 20°55.4’S, 112°51.5’E; 1128-1139 m; 23.1.1991.
Exmouth Plateau; 21°28.2'S, 113°38.6’E; 1022—1023 m; 24.1.1991.

w. of North West Cape; 21°44.7’S, 113°52.3’E; 320-290 m; 24.1.1991.
w. of North West Cape; 21°49.1’'S, 113°47.6'E; 650-685 m; 24.1.1991.
w. of NW Cape; 21°54.1'S, 113°40.7’E; 1,158—1,100 m; 24.1.1991.

w. of NW Cape; 21°58.7'S, 113°08.7’E; 1,460—1,500+ m; 1991.

nw. of Point Cloates; 22°26.2'S, 113°12.3’E; 1,305 m; 1991.

w. of Point Cloates; 22°45.8'S, 113°13.6’E; 910 m; 1991.

off Point Cloates; 22°57.6’S, 113°14.4'E; 482-554 m; 26:1:1991.

w. of Cape Farquhar; 23°42.9’S, 112°35.9’E; 620 m; 1991.

off Cape Cuvier; 24°00.4’S, 111°54.1'E; 1,060—1,064 m; 27.1.1991.

w. of Cape Cuvier; 24°07.5’S, 111°39.4’E; 1,320 m; 1991.

w. of Quobba Point; 24°30.6’S, 111°50.4’E; 895-901 m; 18.1.1991.
off Shark Bay; 25°19.1'S, 111°56.6’E; 610-612 m; 29.1.1991.

off Shark Bay; 25°52.1’S, 111°27.2’E; 1,254—1,277 m; 29.1.1991.

w. of Dirk Hartog I.; 26°04.4’S, 111°47.1'E; 874-882 m; 1991.

w. of Steep Point; 26°14.5’S, 112°03.2’E; 690-691 m; 30.1.1991.

off Shark Bay; 26°35.7’S, 112°29.0’E; 508—S00 m; 30.1.1991.

sw. of Shoal Point; 28°04.1'S, 112°42.6’E; 853-854 m; 1.1I.1991.

sw. of Shark Bay; 26°58.2'S, 112°21.4’E; 666-688 m; #1991.

NW Shelf, sw. of Shark Point; 20°40.8'S, 114°43.4’E; 713-714 m; 1991.
off Shark Bay; 27°22.1'S, 112°10.8’E; 996—1,009 m; 31.1.1991.

nw. of Shoal Point; 28°00’S, 112°41.0’E; 945—960 m; 1.11.1991.

w. of Shoal Point; 28°00.6’S, 112°41.9’E:; 853-854 m; 1991.

90 km sw. of Geraldton; 29°20.5'S, 113°58.3’E; 490-505 m; 6.11.1991.
w. of Leander Pt; 29°21.8’S, 113°46.6’E; 942-970 m; 6.1I.1991.

w. of Leander Pt.; 29°21.8’S, 113°46.6’E; 1,132—1,136 m; 6.11.1991.
wnw. of Green Head; 29°50.3’S, 114°10.9’E; 770—760 m; 7.11.1991.
50 km w. of Green Head; 30°00.0’S, 114°27.1'E; 480-490 m; 8.11.1991.
sw. of Ledge Point; 31°14.9’S, 114°52.3’E; 512 m; 1991.

nw. of Rottnest I.; 31°43.0’S, 114°58.7'E; 485 m; 1991.

Rottnest Canyon; 32°02.3’S, 114°54.5'E; 670-640 m; 10.2.1991.
Rottnest Canyon; 31°54.1'S, 115°04.6’E; 550 m; 1991.

32°19.8'S, 114°28.6’E; 1280 m; 1991.

w. of Mandurah, 32°33.0’S, 114°25.8’E; 1,140 m; 1991.

w. of Mandurah, 32°40.4’S, 114°28.2'E; 880-960 m; 14.11.1991.
33°17.9'S, 114°12.6’E; 982 m; 14.11.1991.

33°17.9'S, 114°30.8’E; 468-430 m; 15.11.1991.

33°25.5'S, 114°21.0’E; 817-780 m; 16.11.1991.

: w. of Cape Freycinet; 34°12.8’S, 114°07.7’E; 1,225—1,240 m; 1991.

: Ss. of Cape Leeuwin, 35°04.9’S, 114°59.7'E; 870-920 m; 18.11.1991.

K71-11-09 — NSW: 30 miles ne. of Jervis Bay; 34°50’S, 151°09’E; 549 m; 7.VII.1971.
K77-16-16 — NSW: e. of Broken Bay; 33°31’S, 152°04’E; 548-603 m; 28.1X.1977.
K78-09-05 — QLD: e. of Gold Coast; 28°02'S, 153°59’E; 548 m; 2.VI.1978.
K78-17-10— QLD: e. of Danger Point; 28°01'S, 154°00’E; 548 m; 17.VIII.1978.
K78-23-08 — QLD: ne. of Danger Point; 28°03’S, 154°04’E; 731 m; 6.X1.1978.

238

K79-15-01 —NSW
K79-20-15 —NSW

K83-09-01 — NSW:
K 83-09-02 — NSW:
K 83-09-04 —NSW:
K83-13-01 —NSW:
K83-15-02 —NSW:
K83-17-01 —NSW:
K84-06-06 —NSW:
K84-08-04 —NSW:
K84-14-01 —NSW:
K84-15-03 — NSW:
K84-16-04 — NSW:
K84-16-13 —NSW:
K84-16-14 —NSW:
K84-16-15 —NSW:
K84-18-06 —NSW:
K84-19-04 —NSW:
K87-02-01 — NSW:
K87-16-02 —NSW:
K87-24-01 —NSW:
K87-24-03 — NSW:
K87-24-04 —NSW:
K87-24-05 — NSW:
K89-06-05 — NSW:
K89-09-09 — NSW:
K89-13-02 —NSW:

K89-17-04 —NSW
K89-17-06 —NSW

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 3

: e. of Sydney; 33°48’S, 151°49’E; 440 m; 2.VII.1979.

: e. of Broken Bay; 33°39’S, 152°06’E; 1,006 m; 6.XII.1979.

off Sydney; 33°48’S, 151°27’E; 942-960 m; 22. VIII.1983.

off Broken Bay; 33°32’S, 152°09’E; 933-969 m; 23.VIII.1983.

se. of Kiama; 34°55’S, 151°13’E; 951-978 m; 24. VIII.1983.

off Broken Bay; 33°40’S, 152°04’E; 988—1,020 m; 17.X.1983.

off Cape Hawke; 32°02’S, 153°09’E; 980 m; 2.X1.1983.

off Sydney; 33°42’S, 151°54’E, 765 m; 12.X.1983.

off Broken Bay; 33°28’S, 152°13’E; 1,107 m; 1984.

On Broken bay; 5343) 9, 152-08 E: 915m: 1984:

Jervis Bay off Wattamolla; 34°19’S, 151°24’E; 505 m; 1984.

off Broken Bay; 33°32'S, 152°03’E; 512-530 m; 11.1X.1984.

e. of Broken Bay; 33°32’S, 152°05’E; 759 m; 25.1X.1984.

off Sydney; 33°45’S, 152°00'E; 927-955 m; 1984.

off Broken Bay; 33°43’S, 152°03’E; 1,043—1,070 m; 27.1X.1984.
off Broken Bay; 33°43’S, 152°03’E; 957-993 m; 27.1X.1984.

ne. of Sydney; 33°48’S, 151°56’E; 914-924 m; 11.X.1984.

e. of Broken Bay; 33°36’S, 152°07’E; 1,024—1,245 m; 16.X.1984.
e. of Wallis Lake; 32°24'S, 153°00’E; 900-960 m; 19.11.1987.

off Sugarloaf Point; 32°25’S, 153°01’E; 880-950 m; 27. VIII.1987.
off Sydney; 33°54’S, 151°48’E; 785-975 m; 7.XII.1987.

e. of Newcastle; 32°55’S, 152°44’E; 960 m; 8.XII.1987.

e. of Hawks Nest; 32°41'S, 152°53’E; 885—950 m; 8.XII.1987.

e. of Harrington; 31°56’S, 153°08’E; 485—925 m; 9. X11.1987.

e. of Budgewoi; 33°13’S, 152°22’E; 722-768 m; 12.1V.1989.

off Terrigal; 33°28’S, 152°09’E; 741—768 m; 18.V.1989.

off Sydney; 33°45'S, 152°03’E; 1,120—1,170 m; 30.VI.1989.

: e. of Crowdy Bay; 31°47’S, 153°18’E; 1,024—1,052 m; 16. VIII.1989.
: e. of Crowdy Head; 31°53’S, 153°16’E; 878-933 m; 16. VIII.1989.

FRV SOELA stations

SO1/82/12 — TAS:

38°06'S, 149°55’E; 250-276 m; 18.1.1982.

S05/84/70 — TAS: 42°41.7'S, 148°24.9’E; 444-448 m; 24.X.1984.

S06/85/36 — QLD: Marian Plateau, e. of Townsville; 17°59’S, 150°32'E; 879-886 m; 25.XI.1985.
SO6/85/47 — QLD: Queensland Trough, e. of Hinchinbrook I.; 17°55’S, 147°00’E; 402 m; 29.X1.1985.
S06/85/86 — Townsville Trough; 18°58.1'S. 150°29.6’E; 420 m; 1985.

S03/86/37 — TAS: w. coast; 42°25.8'S, 144°47.7'E; 792-830 m; 16.V.1986.

SO3/87/13 — TAS: e. of St. Helens; 41°21.2'S, 148°48.9’E; 1,060—1,260 m; 9.V.1987.

SO2/88/87 — VIC: Bass Strait, s. of Gabo I.; 38°06’S, 149°55’E; 250-276 m; 18.1.1982.

SO3/88/09 — VIC: s. of Warrnambool; 39°12.29’S, 142°29.99’E; 1,175 m; 1988.

SO5/88/70 — WA: NW Shelf, s. of Barrow I.; 20°40.8’S, 114°43.4’E; 420 m; 1988.

SO5/88/190 — WA: NW Shelf: 18°08.8’S, 117°54.5'E; 582 m; 12.X.1988.

IWAMOTO AND WILLIAMS: GRENADIERS OF WESTERN AUSTRALIA 239

ACKNOWLEDGMENTS

We take this opportunity to thank the many people who provided us assistance, information, or
resources in the course of this study. Support during the western slope trawl survey was given by the
officers and crew of the Southern Surveyor, and the scientific staff, in particular for providing
taxonomic assistance: Peter Last (CSIRO), John Paxton (AMS), and Martin Gomon (NMV). David
Wright and David Evans (CSIRO) took extensive collections of grenadiers from commercial fishing
vessels. Illustrations of several species were by Georgina L. Davis (GLD); photographs of others were
taken by Thor Carter (CSIRO) and images prepared by Ross Daley (CSIRO). Alan Leviton (CAS)
helped with digital images and provided sage advice on the manuscript. Scanning electron mi-
crographs were prepared by Darrell Ubick (CAS).

For loans of material and general curatorial assistance, we thank Peter Last, David Wright,
Alastair Graham, and Gordon Yearsley (CSIRO Ian Munro Ichthyological Collection); Doug Hoese,
John Paxton, Mark McGrouther, Kerry Parkinson, and other staff (AMS); Martin Gomon (NMV);
Clive Roberts and Andrew Stewart (NMNZ); Peter McMillan (National Institute of Water and
Atmosphere, Wellington, NZ); Barry Hutchins, Kevin Smith, and Sue Morrison (WAM); Helen
Larson and R. Williams (NTM); Nigel R. Merrett (BMNH); Jon Fong and David Catania (CAS);
Osamu Okamura (BSKU); Yuri I. Sazonov (ZMMGU); Yuri N. Shcherbachev (IOAN); Roland J.
McKay and Jeff Johnson (QM).

Tracey Bywaters of the Western Australian Education Service gave advice on Australian
aboriginal languages used in naming new species. The paper benefited from the careful reviews
provided by Alastair Graham and Gordon Yearsley (CSIRO) and two anonymous reviewers.

This work was part of a project supported by the Fisheries Research and Development Corpora-
tion, Grant No. 1988/74. The first author (TI) gratefully acknowledges the Australian Museum for a
visiting fellowship, which allowed a three-months’ visit to Australian museums; additional support
was provided by CSIRO Marine Research for a visit to Hobart, the New Zealand Foundation for
Research Science and Technology Contract MNZ603 to Dr. Clive Roberts (Programme Leader),
Museum of New Zealand Te Papa Tongawera, for a visit to Wellington, and the CAS In-House
Research Fund.

LITERATURE CITED

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. 1894. Natural history notes from H. M. Indian Marine Survey Steamer ‘Investigator,’ Commander

C. E. Oldham, R. N., commanding. Series 2, No. 11. An account of a recent collection of bathybial fishes

from the Bay of Bengal and from the Laccadive Sea. J. Asiat. Soc. Bengal 63 (pt. 2)(2):115—137, pls. 6—7.

. 1899. A descriptive catalogue of Indian deep-sea fishes in the Indian Museum, collected by the Royal
Indian Marine Survey Ship ‘Investigator.’ Indian Museum, Calcutta. 111 + 211 pp., 8 pls., 1 map.

AMAOKA, K., K. MATSUURA, T. INADA, M. TAKEDA, H. HATANAKA, AND K. OKADA, eds. 1990. Fishes collected
by the R/V Shinkai Maru around New Zealand. Japan Marine Fishery Resource Research Center. 410 pp.

ARAI, T. 1979. Additional information on a rare macrourid fish, Mesobius antipodum, from New Zealand.
Japanese J. Ichthyol. 25(4):286—289, figs. 1-3, table 1.

ARAI, T. AND T. IWAMOTO. 1979. A new species of the macrourid fish genus Coelorinchus from off Tasmania,
New Zealand, and the Falkland Islands. Japanese J. Ichthyol. 26(3):238—246.

BARNARD, K. H. 1925. Descriptions of new species of marine fishes from S. Africa. Ann. Mag. Nat. Hist., ser.
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BERG, C. 1989. Substitucion de nombres genéricos. Comm. Mus. Nac. Buenos Aires 1:41—43.

COHEN, D. M., ed. 1989. Papers on the systematics of gadiform fishes. Nat. Hist. Mus. Los Angeles County, Sci.
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COLLETT, R. 1896. Poissons provenant des campagnes du yacht I’Hirondelle (1885—1888). Rés. Camp. Sci.
Monaco 10:1—198, pls. 1-6.

240 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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DAVIS, T. L. O. AND T. J. WARD. 1984. CSIRO finds new scampi grounds off the North West Shelf. Austr. Fish.
43(8):41-45.

ESCHMEYER, W. N. 1990. Catalog of the genera of Recent fishes. Calif. Acad. Sci., San Francisco. 697 pp.

FOWLER, H. W. 1925. New taxonomic names of West African marine fishes. Amer. Mus. Novit. (162):1—S.

GILBERT, C. H. 1905. The deep-sea fishes of the Hawaiian Islands. Pp. 575—713, figs. 230-276, pls. 66-101 in
The aquatic resources of the Hawaiian Islands, D. S. Jordan and B. W. Evermann, eds. Bull. U.S. Fish
Comm. 1903, 22(pt. 2, sect. 2).

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IWAMOTO AND WILLIAMS: GRENADIERS OF WESTERN AUSTRALIA 243

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© CALIFORNIA ACADEMY OF SCIENCES, 1999
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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES

Volume 51, No. 4, pp. 245-257, 7 figs., 1 table. July 20, 1999

Two New Species of Gargamella (Mollusca, Nudibranchia)
from South Africa

by
Jocelyn B. Garovoy, Angel Valdés
and

Terrence M. Gosliner
Department of Invertebrate Zoology and Geology,
California Academy of Sciences
Golden Gate Park, San Francisco, CA 94118

This paper describes two new species of the genus Gargamella from South Africa. Gar-
gamella gravastella is characterized by a dark grey notum with white-ringed dark spots,
and a dense covering of caryophyllidia. Distinct reproductive features include a smooth
accessory gland, and a series of large penial hooks, each having a wide base and short cusp.
Gargamella bovina is characterized by a white notum covered with dark brown spots, and
short caryophyllidia. Its outermost lateral tooth is notably smaller than the other outer
lateral teeth. Distinct reproductive features include a large lobate accessory gland which
bears a muscular attachment to the female gland, and a series of large penial hooks, each
with a wide base and long cusp. Additional information on the caryophyllidia, reproduc-
tive, and radular characteristics is also provided.

In order to determine the systematic position of G. gravastella and G. bovina, a phylogenetic
analysis has been carried out, including representatives of the genera Baptodoris, Platydoris
and Gargamella. The phylogeny obtained supports the placement of these two new species
in the genus Gargamella. In addition, Gargamella, Platydoris and Baptodoris appear to be
distinct, monophyletic genera.

The genus Gargamella Bergh, 1894 presently contains only one valid species: Gargamella
immaculata Bergh, 1894 from South America. Miller (1996) synonymized Gargamella no-
vozealandica Eliot, 1907, from New Zealand, with Jorunna pantherina (Angas, 1864), and Schrédl
(1997) regarded Gargamella latior Odhner, 1926, another nominal species described from South
America, as a junior synonym of G. immaculata, based on observations of the internal anatomy and
geographical range.

Gosliner (1983) reported two additional, undescribed species of Gargamella from South Africa,
both distinguishable from the other described species. The present paper describes these two new
species based on specimens deposited in the Department of Marine Biology of the South African
Museum, Cape Town (SAM). Comparative material of other species discussed in this paper was
obtained from the Department of Invertebrate Zoology of the California Academy of Sciences
(CASIZ), the Department of Invertebrate Zoology of the Swedish Museum of Natural History
(SMNHI) and the Museo Nacional de Ciencias Naturales of Madrid, Spain (MNCN).

246 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 4

SPECIES DESCRIPTIONS

Gargamella gravastella sp. nov.
Pies, TA, 2,3

Gargamella sp. | Gosliner, 1987:70, fig. 94.

TYPE MATERIAL.— HOLOTYPE: Dale Brook, False Bay, Cape Province, South Africa (34°07'26"S,
18°27'33"E), 18 September, 1982, 11 mm preserved length, collected by T. M. Gosliner (SAM A35382).
PARATYPES: Dale Brook, False Bay, Cape Province, South Africa (34°07'26"S, 18°27'33”E), 18 September,
1982, one specimen 4 mm preserved length (SAM A54295); date unknown, one specimen 17 mm preserved
length, dissected, collected by T. M. Gosliner (SAM A35384); 10 September 1982, one specimen 24 mm
preserved length, dissected, collected by T. M. Gosliner (SAM A35383); 18 September, 1982, one specimen 10
mm preserved length, dissected, collected by T. M. Gosliner (SAM A35381).

ETYMOLOGY. — Gargamella gravastella is named for its external coloration and body texture.
The word gravastella is Latin for “grey headed fellow” and makes reference to the rough grey notum
of this caryophyllidia-bearing dorid nudibranch.

DISTRIBUTION. — Specimens of this species have been found only at Dale Brook, False Bay,
Cape Peninsula, Cape Province, South Africa.

EXTERNAL MORPHOLOGY. — The living animal (Fig. 1A) has a dark gray notum marked with
several dark gray spots, each surrounded by a white circle. The perfoliate rhinophores are pale
yellowish white in color. They consist of 8 transverse lamellae. The gill is also pale yellowish white,
the same color as the rhinophores, and is composed of 8 unipinnate branchial leaves. The anal papilla
lies within, and closes the circlet of the branchial leaves.

The body shape is oval with a slightly undulate mantle. The body is covered with many short
caryophyllidia tubercles measuring about 70 um. Each caryophyllidium consists of a rounded central
ciliated tubercle that occupies completely the space inside the surrounding ring of spicules. Several
(5—9) tall spicules project around the tubercle margin (Fig. 2D). The foot is bilabiate and notched.
(Fig. 3B) The foot is wide in relation to the mantle margin. The oral tentacles are well formed, and
appear conical in shape.

BUCCAL ARMATURE. — The jaws are smooth, with no rodlets. The radular formulae of two
specimens examined are 31 x 39.0.39 (SAM A35383) and 36 x 42.0.42 (SAM A35384). The inner
lateral teeth (Fig. 2A) are smooth, and gradually increase in size toward the mid-lateral teeth (Fig.
2B), which also lack denticles. The outer lateral teeth (Fig. 2C) are smooth as well, and gradually
decrease in size. The outermost lateral teeth are not extremely small relative to the other outer lateral
teeth.

REPRODUCTIVE SYSTEM. — The ampulla is tubular and simple. It narrows before branching into
the short oviduct, which enters the female gland, and the large prostate (Fig. 2A). The prostate is
nearly as wide as the female gland mass in the specimens examined. At the proximal end of the
prostate, there is a junction at which the long, convoluted deferent duct narrows and becomes the
tubular ejaculatory duct. The penial bulb widens at its proximal end, and is armed with four rows of
hooks. The hooks have a wide base approximately 0.1 mm wide, and a relatively short cusp
approximately 25 um long (Fig. 2C). The deferent duct and vaginal duct meet, forming an atrium
where there is a small spherical accessory gland, which lacks any armature. The proximal end of the
vaginal duct is very wide, wider than the deferent duct. The vagina narrows at the distal end before
entering the oval shaped bursa copulatrix. Another duct leads from the bursa copulatrix to the oval
shaped, stalked seminal receptacle. This duct, which is located directly adjacent to the junction of the
vaginal duct and bursa copulatrix, is the same width as the narrowest part of the vaginal duct and is

GAROVOY, VALDES, AND GOSLINER: NEW SPECIES OF GARGAMELLA 247

ee ee ae
wit de 2 . ts a - a
Ms } kee > =< .

, «
¢

FiGure |. Living animals. A.Gargamella gravastella sp. nov, specimen from Dale Brook on the western shore of False Bay,
South Africa, photograph by T. M. Gosliner. B. Gargamella bovina, sp. nov, specimen from Buffels Bay, on the False Bay

coast of Cape Point, South Africa, photograph by T. M. Gosliner.

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 4

FIGURE 2. Gargamella gravastella sp. nov. (SAM A35383), scanning electron micrographs. A. Inner lateral teeth, scale =
43 um. B. Mid lateral teeth, scale = 60 jum. C. Outer lateral teeth, scale = 43 xm. D. Caryophyllidia tubercles, scale = 100 pm.

long and convoluted. It joins the uterine duct. The uterine duct is short and enters the female gland
mass at the base of the stalk of the small seminal receptacle.

REMARKS. — Gargamella gravastella is the only South African caryophyllidia-bearing dorid
with a dark gray background color. Other distinctive features of this species are the dark gray spots,
circled in white, that cover the notum. In fact, the coloration of G. gravastella distinguishes it from

GAROVOY, VALDES, AND GOSLINER: NEW SPECIES OF GARGAMELLA 249

FIGURE 3. Gargamella gravastella sp. nov. (SAM A35381). A. Reproductive system, a = ampulla, be = bursa copulatrix,
d = deferent duct, f = female gland, g = accessory gland, p = prostate, rs = receptaculum seminis, scale bar = 0.5 mm.
B. Ventral view of the anterior border of foot, scale bar = | mm. C. Penial hooks, scale bar = 0.1 mm.

G. immaculata, with colors ranging from solid white, to yellow or orange with darker rhinophores
(Schréd! 1997). On the contrary, the pale yellow rhinophores of G. gravastella are much lighter than
the rest of the body, with the exception of the matching pale branchial leaves.

As with G. immaculata (see Schrédl 1997), G. gravastella is densely covered with caryophyllidia
tubercles, has penial hooks, a well-developed, large prostate, and an accessory gland. There are
notable differences in the buccal armature of this species as compared to G. immaculata. Both the
inner and mid lateral teeth of G. gravastella are wider and shorter than those of G. immaculata.
Moreover, G. gravastella has fewer rows of teeth, 31—36 in two specimen, than does G. immaculata,
which Schrédl (1997) reports as having from 60 to 91 rows.

The reproductive system of G. gravastella is also distinct. The overall shape of the bursa
copulatrix is oval, rather than spherical, and the junction of the vaginal duct with the bursa copulatrix
is directly adjacent to the duct connecting the bursa copulatrix with the receptaculum seminis, whereas
the two ducts on the bursa copulatrix of G. immaculata are much farther apart (Marcus 1959; Schrédl
1997). The receptaculum seminis is notably smaller, and the accessory gland is smooth in contrast to
the larger receptaculum seminis and lobate accessory gland of G. immaculata. Also, G. gravastella
lacks the large vestibular hooks described in G. immaculata (Schrédl, 1997).

Gargamella bovina sp. nov.
Figs 1B, 4, 5

Gargamella sp. 2 Gosliner, 1987:71, fig. 95.
TYPE MATERIAL. — HOLOTYPE: off Danger Point, Cape Province, South Africa (34°32'34"S, 19°15'48"E),

44 m depth, 16 April 1984, 20 mm preserved length, collected by W. R. Liltved (SAM A36013). PARATYPES:
off Danger Point, Cape Province, South Africa (34°32'34"S, 19°15'48”E), 44 m depth, 16 April 1984, one

250 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 4

specimen 20 mm preserved length, dissected, collected by W. R. Liltved (SAM A54296). Off Danger Point,
Cape Province, South Africa (34°32'34"S, 19°15'48”E), 34 m depth, 11 April 1984, two specimens 16 mm
preserved length, dissected, collected by W. R. Liltved (SAM A36010). Buffels Bay, False Bay, Cape Province,
South Africa (34°19'03"S, 18°27'42"E), July 1982, one specimen 14 mm preserved length, dissected, collected
by T. M. Gosliner (SAM A35385).

ETYMOLOGY. — Gargamella bovina is named for its external body coloration and pattern. The
word bovina is Latin for “cowlike” and makes reference to the pattern of dark brown spots on the
white notum that give this species an appearance similar to a Holstein dairy cow.

DISTRIBUTION. — Specimens of this species have been found off the coast of Cape Province,
South Africa, off Danger Point and Buffels Bay as well.

EXTERNAL MORPHOLOGY. — The body is oval, with a slightly undulate mantle. It is white in
color, with several dark brown irregular spots (Fig. 1B). The rhinophores and branchial leaves are
also white. The perfoliate rhinophores consist of 14 transverse lamellae. The bipinnate gill consists
of 7 branchial leaves. The anal papilla closes the circlet of branchial leaves. The notum of this animal
is covered with many low caryophyllidia, measuring about 70 pm. Each caryophyllidium bears a very
small ciliated tubercle, and is surrounded by 5—7 short projecting spicules (Fig. 4D). The foot is broad
with respect to the mantle margin. The anterior border of the foot is bilabiate and notched (Fig. 5B).
The oral tentacles are well formed, and digitiform in shape.

BUCCAL ARMATURE. — The jaws are smooth, with no rodlets. The radular formulae of two
specimens examined are 21 x 38.0.38 (SAM A36010) and 35 x 40.0.40 (SAM A35385). The inner
lateral teeth are smooth (Fig. 4A), and gradually increase in size toward the mid-lateral teeth, which
also lack denticles (Fig. 4B). The outer lateral teeth gradually decrease in size, the outermost lateral
tooth being much smaller than the other outer lateral teeth (Fig. 4C).

REPRODUCTIVE SYSTEM. — The ampulla of this species is tubular, simple, and small (Fig. 5A).
It branches and connects to a large, massive prostate and a short oviduct which connects to the female
gland mass. The distal end of the prostate connects to an elongate, convoluted deferent duct. The distal
end of the deferent duct is armed with four rows of penial hooks. The penial hooks have a broad base,
0.1 mm wide, anda long, straight hook, 37.5 zm long (Fig. SC). The vaginal duct is long, and narrower
than the deferent duct. The proximal end of the vagina opens into the round bursa copulatrix. The
duct connecting the bursa copulatrix and the sessile, spherical receptaculum seminis is long and
narrow. This duct connects to the bursa copulatrix at a point far from, and nearly opposite the junction
of the vaginal duct and bursa copulatrix (see Fig. 5A). The uterine duct is short and enters the female
gland mass. A large, lobed accessory gland is present at the widened opening of the deferent duct and
the vagina. The external surface of the gland bears a thin muscular connection to the female gland
mass. This accessory gland does not contain any armature.

REMARKS. — Gargamella bovina is externally distinguishable from G. gravastella by its white
background color, and dark brown spots. Its notal surface, though covered with caryophyllidia, has
lower tubercles, very different from those of G. gravastella. Also, its coloration distinguishes it from
G. immaculata which is solid white, yellow or orange (Schrédl 1997).

The buccal armature of G. bovina is different from G. immaculata in that there are fewer rows
of teeth, 21—35 in two specimens of G. bovina compared with 60—91 rows in G. immaculata (Schrédl
1997). The smooth teeth of G. bovina and G. immaculata are similar in shape, but the teeth of G.
bovina are considerably shorter than those of G. immaculata. The radula of G. bovina also differs
from that of G. gravastella in the shape of the lateral teeth that are more elongate in the former.

The reproductive system of G. bovina closely resembles G. immaculata, more so than G.
gravastella. Like G. immaculata, (see Marcus 1959; Schrédl 1997) the two ducts coming from the
bursa copulatrix are situated far apart from each other. Also, the accessory gland is lobate in both G.
immaculata and G. bovina. However, G. bovina bears a unique muscular attachment from the

GAROVOY, VALDES, AND GOSLINER: NEW SPECIES OF GARGAMELLA

FIGURE 4. Gargamella bovina sp. nov., scanning electron microcraphs. A. Inner lateral teeth (SAM A36010), scale = 60 tum.
B. Mid lateral teeth (SAM A36010), scale = 60 ym. C. Outer lateral teeth (SAM A36010), scale = 43 pum. D. Caryophyllidia
tubercles, (SAM A36013) scale = 60 um.

252 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 4

ee

FiGuRE 5. Gargamella bovina sp. nov. (SAM A36010). A. Reproductive system, a = ampulla, be = bursa cpulatrix, d=
deferent duct, f= female gland, g = accessory gland, p = prostate, rs = receptaculum seminis, scale bar = | mm. B. Ventral
view of anterior border of foot, (SAM A36013) scale bar = 1 mm. C. Penial hooks, scale bar = 0.1 mm.

accessory gland to the female gland mass. The receptaculum seminis is sessile, where that of G.
immaculata is stalked. Gargamella bovina also possesses penial hooks, a feature common to this
genus, but lacks the large vaginal hooks found in G. immaculata. The penial hooks of G. bovina have
the same wide base as G. gravastella, but the cusp is notably longer.

DISCUSSION

The genus Gargamella was introduced by Bergh (1894) based on Gargamella immaculata Bergh,
1894, the type species by monotypy. He included the following features in the diagnosis of this genus:
body somewhat depressed, dorsum silky, oral tentacles digitiform, branchial leaves tripinnate,
anterior border of the foot notched and grooved, jaws and rachidian radular teeth absent, lateral teeth
hook-shaped, prostate large, penis armed with hard spherical hooks, and vestibular gland present.

Ernest Marcus (1959) amplified the description of G. immaculata with anatomical information
based on newly collected material. The most outstanding new feature described by Ernest Marcus is
the presence of large copulatory spines in the atrium of G. immaculata and Gargamella latior Odhner,
1926, another nominal species described from South America. Schrédl (1997) redescribed G.
immaculata and synonymized G. /atior with it.

We have examined two specimens of G. immaculata collected from Seno Otway, near Punta
Prat, Chile (SMNHI 12520) for comparison. We have found large hooks in the penis, but the vestibular
hooks described by Ernest Marcus (1959) and Schrédl (1997) are actually situated in the opening of
the vagina (Fig. 6A). The vestibular gland of this species opens in the atrium, and therefore we called
it an accessory gland, to be distinguished from the vestibular glands of other dorids that open in a
different position. The vaginal hooks are very large, with the base reaching up to 500 ym, and are
arranged ina single row. The radula has hook-shaped, elongate teeth lacking denticles. The outermost
lateral teeth are large, almost the same size as the innermost. Externally this species is characterized

GAROVOY, VALDES, AND GOSLINER: NEW SPECIES OF GARGAMELLA 253

FiGuRE 6. Scanning electron micrographs. A. Open atrium of Gargamella immaculata (SMNHI 12520) showing the penial
and vaginal hooks, at = atrium, pe = penis, v = vagina, scale bar = 750 um. B. Open atrium of Baptodoris cinnabarina (MNCN
15.05/32381) showing the penial and atrial hooks, at = atrium, pe = penis, scale bar = 150 um.

by a convex shape of the body, a wide foot and the presence of large caryophyllidia tubercles, about
200 um long.

Other genera probably related to Gargamella are Baptodoris Bergh, 1884 and Platydoris Bergh,
1877. In fact, Bouchet (1977) considered that Gargamella and Baptodoris were synonyms of
Platydoris. The genus Baptodoris, type species Baptodoris cinnabarina 1884, is characterized by the
presence of a large hooks in the atrium besides a penial papilla armed with an additional set of hooks
(Fig. 6B), much smaller in size. It also has an accessory gland connected to the atrium. The lateral
radular teeth of Baptodoris have one or more denticles, and the outermost laterals have many tiny
denticles. Externally, Baptodoris is characterized by the presence of low caryophyllidia tubercles, a
flattened body and a narrow foot. This pattern is consistent in the only two species of the genus that
have been anatomically studied, B. cinnabarina and B. mimetica Gosliner, 1991. We have examined
one specimen of B. cinnabarina collected from the Mediterranean coast of Spain (MNCN
15.05/32381), and information on B. mimetica was obtained from Gosliner (1991). Other species
assigned to this genus are not sufficiently described to verify the presence of these characteristics.

The genus Platydoris, type species Platydoris argo Linnaeus, 1758, includes large species with
a ventrolaterally flattened body and a narrow foot. An anatomical study of P. argo based on specimens
collected from Northern Spain (CASIZ 115217), revealed the presence of several rows of large hooks
in the vagina and smaller hooks in the penis, about 50 um long, much smaller that those of G.
immaculata. A large accessory gland opens in the atrium. In addition, the prostate of Platydoris is
very small, the caryophyllidia tubercles are very low, and the outer lateral radular teeth are very small.
Other species of Platydoris, such as Platydoris formosa (Alder and Hancock 1866), may lack vaginal
hooks (see Kay and Young 1969), but this point needs confirmation.

Gargamella gravastella and G. bovina appear to be more closely related to G. immaculata than
to P. argo or B. cinnabarina. The species from South Africa differ from G. immaculata in lacking

254 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 4

vaginal hooks. However, the morphology of the penial hooks is very similar in these three species.
In all three the penial hooks have a broad base, 0.1 mm wide, and a long, straight cusp. G. immaculata,
G. gravastella, and G. bovina all have a large prostate and an accessory gland that opens in the atrium.
They also share similar radular morphology, with large, smooth marginal teeth, and mid-lateral teeth
with elongated cusps also lacking denticles. The caryophyllidia tubercles are also well-formed in these
three species.

PHYLOGENETIC ANALYSIS

In order to determine the phylogenetic relationships of Platydoris, Gargamella and Baptodoris,
and therefore the systematic position of the two new species from South Africa, we have carried out
a phylogenetic analysis. For this analysis, we have included the two new species from South Africa
described in this paper, as well as Gargamella immaculata, Baptodoris cinnabarina, Baptodoris
mimetica, Platydoris argo, Platydoris angustipes (Mérch, 1863), Platydoris stomascuta Bouchet,
1977 and Platydoris maculata Bouchet, 1977. Information regarding G. immaculata, B. cinnabarina
and P. argo was obtained from our own anatomical studies. Data on P. angustipes was extracted from
Ernest Marcus (1957), data on P. stomascuta and P. maculata from Bouchet (1977) and data on B.
mimetica from Gosliner (1991).

To polarize the morphological variability we have selected Diaulula sandiegensis (Cooper, 1862)
as the outgroup. Information on this species has been obtained from our own dissections of one
specimen collected from California (CASIZ 068277).

The following characters (Table 1) were considered in undertaking the phylogenetic analysis:

1. Body shape. Species of Baptodoris and Gargamella, including the two new species from
South Africa have a convex body. On the contrary, species of Platydoris have a flattened body. A
convex body is also present in D. sandiegensis and it is regarded as the plesiomorphic condition.

2. Foot width. Species of Platydoris and Baptodoris have a narrow foot, whereas Diaulula and
Gargamella, including G. bovina and G. gravastella, have a wide foot. A narrow foot appears to be
apomorphic.

3. Branchial sheath. In Platydoris argo and P. maculata the branchial sheath is lobate, whereas
in the other species of Platydoris, Gargamella, Baptodoris and Diaulula it is rounded. A lobate
branchial sheath is considered apomorphic.

4. Caryophyllidia. In species of Platydoris and Baptodoris, the caryophyllidia are very low and
difficult to recognize with an optical microscope. In Gargamella, including the species from South
Africa and Diaulula, the caryophyllidia are large and conspicuous. Small caryophyllidia are consid-
ered apomorphic.

5. Lateral radular cusp. The cusps of the radular teeth of G. immaculata, G. bovina and G.
gravastella are very elongate, longer than the base of the teeth. In Platydoris, Baptodoris, and
Diaulula the cusps are always shorter than the base of the teeth. An elongate radular cusp is considered
apomorphic.

6. Lateral radular teeth denticles. In species of Baptodoris the lateral radular teeth have one
or more denticles that are absent in Diaulula, Platydoris and Gargamella. The presence of denticles
on the lateral radular teeth is considered apomorphic.

7. Outermost radular cusp. The outermost lateral teeth of Platydoris and Baptodoris are short
plates (there is no information on P. stomascuta and P. maculata), whereas in Gargamella and
Diaulula they are typical hamate teeth similar in shape to the other radular teeth. The presence of
hamate outermost lateral teeth is regarded as plesiomorphic.

8. Outermost lateral teeth denticles. In Baptodoris, the outermost lateral teeth have a number
of tiny denticles that are not present in Platydoris, Gargamella and Diaulula. The presence of these
denticles is considered the apomorphic condition.

GAROVOY, VALDES, AND GOSLINER: NEW SPECIES OF GARGAMELLA 255

TABLE |. Character states for the taxa involved in the phylogenetic analysis. Data code: 0 = plesiomorphic
condition; 1, 2 = apomorphic condition; ? = missing data.

Ww
(oa)
—_
tN
Ww

D. sandiegensis 0
P. argo 1
P. angustipes ]
P. maculata l
P. stomascuta 1
G. immaculata 0
G. bovina 0
G. gravastella 0
B. cinabarina 0
B. mimetica 0

SII OS SS) & nN
SoS SSS SoS | iS

= OOO KK = = © i)
ooooococorek OS
pep OOO KH HK = = © as
IS SO oo Oo ©) & a
- KS COON Y HK = © ~
ISS Soo) &) &)
oooooocor-.o& Ne)
FB eENNNK KH Ho 1oO
or OreK OC OCC SO
RBS BH OO BR Be KE KH Oo
SSeS Storey ater oun

9. Prostate size. The prostate of P. argo and P. angustipes is very small compared to those of P.
stomascuta, P. maculata, Baptodoris, Gargamella and Diaulula. Presence of a small prostate is
considered apomorphic.

10. Penial hooks. In Diaulu/a there are no penial hooks. In species of Platydoris and Baptodoris
the hooks are small, never larger than 70 tm, whereas in Gargamella the penial hooks are very large,
over 120 um long. Absence of penial hooks is considered plesiomorphic. This character is treated as
unordered in the analysis.

11. Bursa copulatrix ducts. Two ducts lead from the bursa copulatrix of all species included in
the analysis. In species of Platydoris, G. gravastella, B. mimetica and D. sandiegensis these ducts are
immediately adjacent to each other. In the remaining species of Gargamella and B. cinnabarina, there
is some distance between the two ducts. Having the ducts separated by some distance is considered
apomorphic.

12. Vaginal and atrial hooks. In Baptodoris, Platydoris and G. immaculata there are very large
hooks situated in the atrium and in the opening of the vagina. These hooks are absent in Diaulula, G.
bovina and G. gravastella. The absence of vaginal or atrial hooks is considered plesiomorphic.

13. Vaginal hooks position. In G. immaculata, P. maculata, P. stomascuta and Baptodoris, the
atrial hooks are situated in the atrium or in the opening of the vagina, whereas in P. argo and P.
angustipes the hooks are situated along the vaginal duct. The latter position is considered apomorphic.

14. Accessory gland. In Baptodoris, Platydoris and Gargamella there is a lobate accesory gland
that opens in the atrium. This gland is absent in Diaulula. The absence of the atrial glands is considered
the plesiomorphic condition.

In order to develop phylogenetic hypotheses regarding these taxa, the above described characters
were placed in a data matrix (Table 1) and analyzed by means of Phylogenetic Analysis Using
Parsimony (PAUP), version 3.1.1, using the exhaustive algorithm. A single most parsimonious tree
was obtained (Fig. 7). This tree requires 17 steps, has a consistency index of 0.88 and a retention
index of 0.92.

The tree obtained clearly supports the placement of G. bovina and G. gravastella in the genus
Gargamella, which is the sister group of the clade formed by Platydoris and Baptodoris. It is also
evident from this phylogenetic hypothesis that Platydoris and Baptodoris form a monophyletic group,
supported by 3 synapomorphies: narrow foot (2), small caryophyllidia tubercles (4), and plate-shaped,
outermost radular teeth (7). Within the genus Gargamella, which is suported by a single synapomor-
phy, radular teeth with elongate cusps (5), G. gravastella is the sister group of the other two species.
Platydoris is also supported by a single synapomorphy, flattened body (1). Baptodoris is the sister

256 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 4

my ® w x ® 2
® Ss iS w g > c
Q © rs} = as} =) L o
= ~ Z o = 5 © om) =)
QD SY o Q i) ny & © D>
9 = a € % € € > os 2
i) 2 & S = = e 8 S 3
“a i o wo w & w w
a) 2 ae) Xn = = => => — Yo
S S S S Ss Sd & € & =
ae) 5) ss) ke) is) is) o o o S
= = = = 5 a 2) — ® =
AS) & AY Ay o ® G ® ny BY
Qo Q Qo Q jaa) ea) O © © Q
11 ie
3,9, 13 11
6, 8
1
Se 10)
CA ae:
10, 14

FIGURE 7. Hypothesis of phylogeny of Gargamella, Platydoris and Baptodoris. Numbers refer to characters listed within
the text.

group of Platydoris and it is supported by two synapomorphies, lateral teeth denticulate (6) and
outermost lateral teeth with minute denticles (8). Platydoris maculata and P. stomascuta appear to
be members of the genus Platydoris, but there is not enough resolution to determine their relatioships
with P. argo and P. angustipes.

BIOGEOGRAPHY

The genus Gargamella now includes three valid described species. Gargamella immaculata, the
type species, was described from Argentina and is also known from Chile (Schrédl 1997). With the
description of Gargamella gravastella and Gargamella bovina from South Africa, it is apparent that
members of this genus are restricted to southern oceans.

It is evident that the highly developed penial armature of G. immaculata is more derived than
that of the two South African species (see Phylogenetic Analysis), perhaps indicating a more recent
divergence from their common ancestor. Preliminary phylogenetic analysis of outgroups of Gar-

GAROVOY, VALDES, AND GOSLINER: NEW SPECIES OF GARGAMELLA 25

gamella indicates that they are more closely related to species of Platydoris and Baptodoris than to
any other caryophyllidia-bearing dorids. Species of Platydoris are found throughout tropical and
subtropical waters, with no cold water southern ocean representatives. The genus Baptodoris is also
absent in the southern ocean cold waters. Only one representative was decribed from the South Pacific,
Baptodoris fongosa Risbec, 1928 from the warm waters of New Caledonia. However, this species
was poorly described and its generic placement is uncertain.

These facts suggest that the southern distribution of Gargamella today is not a result of ancient
Gondwanan distributions, but is more likely a result of more recent vicariant and distributional events.
This is also a more likely scenario, owing to the fact that members of these genera are among the
more highly derived cryptobranch dorids.

ACKNOWLEDGMENTS

This paper was supported in part by the Summer Systematics Institute through the REU division
of the National Science Foundation, and the Ministerio de Educacion y Cultura of Spain (SEUI)
through its postdoctoral fellowships program.

The authors thank Dong Lin in the photography department of the California Academy of
Sciences for printing the scanning electron micrographs, and Darrell Ubick for helping with critical
point drying and operation of the SEM. Manuel Ballesteros, Jesus Ortea, and Anders Warén kindly
lent us specimens for examination. We also thank W. R. Liltved, who collected several specimens
used in the description of these species, the South African Museum for the support of original field
work and the use of preserved material, and Gary Williams for his lengthy discussions of southern
ocean biogeography.

LITERATURE CITED

BERGH, R. 1894. Die Opisthobranchen, 13. Reports of the dredging operations off the West coast of Central
America to the Galapagos, to the West coast of Mexico, and in the Gulf of California, in charge of Alexander
Agassiz, carried on by the U. S. Fish Commision Steamer “Albatros,” during 1891, Lieut. Commander Z.
L. Tanner, U.S. N., commanding. Bulletin of the Museum of Comparative Zoélogy 25:125—235, pls 1-12.

BOUCHET, P. 1977. Opisthobranches de profondeur de 1’Océan Atlantique: II— Notaspidea et Nudibranchia.
Journal of Molluscan Studies 43:28—66.

GOSLINER T. M. 1987. Nudibranchs of Southern Africa. A guide to Opisthobranch Molluscs of Southern Africa.
Sea Challengers, Monterey. 136 pp.

. 1991. Four new species and a new genus of opisthobranch gastropods from the Pacific Coast of North
America. The Veliger 34:272—290.

Kay, E. A. AND D. K. YOUNG. 1969. The Doridacea (Opisthobranchia; Mollusca) of the Hawaiian Islands.
Pacific Science 23:172—231.

MARCUS, ER. 1957. On Opisthobranchia from Brazil (2). Journal of the Linnean Society of London 43:390-486.

. 1959. Lamellaria und Opisthobranchia. Reports of the Lund University Chile Expedition 1948-49
36:1—133.

MILLER, M. C. 1996. The dorid nudibranch genus Jorunna Bergh, 1876 (Gastropoda: Opisthobranchia) in New
Zealand. Journal of Natural History 30:1095—1109.

SCHRODL, M. 1997. On the Magellanic nudibranch Gargamella immaculata Bergh, 1894, and its synonymy to
G. latior Odhner, 1926. Spixiana 20:8 1—92.

© CALIFORNIA ACADEMY OF SCIENCES, 1999
Golden Gate Park
San Francisco, California 94118

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES

Volume 51, No. 5, pp. 259-310, 21 figs., 7 tables. August 20, 1999

A Review of the Fish Genus Dibranchus with D i tions of New
Species and a New Genus, Solocisquama, >>

Ap A Sc rie
(Lophiiformes, Ogcocephalidae) Panog)§8bora

by

Margaret G. Bradbury!
Moss Landing Marine Laboratories ~~
P. O. Box 450, Moss Landing, CA 95039-0450

Dibranchus Peters, with 13 species, and Solocisquama, n. gen., with 3 species are reviewed. ~~
Three new species of Dibranchus are described, one each from the Atlantic, Indian, and
east Pacific oceans. Dibranchus scaber Garman 1899 is a junior synonym of D. hystrix
Garman 1899. Dibranchus alberti Nunez 1967 is a junior synonym of D. spinosus (Garman)
1899. Solocisquama includes 2 species formerly referred to Dibranchus and 1 species
described as new. A key to eastern Pacific species of Dibranchus and a key to species of
Solocisquama are presented.

Although Dibranchus atlanticus is known from hundreds of specimens, 5 species of Dibranchus
and the 3 species of Solocisquama are rare, known from one or a few specimens. All species are
uncommon. The material of Dibranchus from the eastern Pacific represents 9 species, many more
than from any other biogeographic region.

Like most ogcocephalids, members of the genus Dibranchus are bottom fishes with large,
strongly depressed heads reminiscent of skates. Present in the Atlantic, Pacific, and Indian oceans,
Dibranchus occurs at tropical and subtropical latitudes in depths from 200 to over 2200 meters. The
little available data suggest that the majority of species have characteristic bathymetric ranges; for
example, the two Atlantic species appear to be bathymetrically well separated, with D. atlanticus
distributed above 800—1000 m and D. tremendus below.

I have examined 407 specimens from the eastern Pacific representing 9 species, of which | is
new and 2 recently described (Bradbury et al. 1999), and 465 specimens from the Atlantic Ocean
representing 2 species. Most Atlantic specimens may be assigned to the type species of the genus,
Dibranchus atlanticus Peters, but 34 specimens represent a new Atlantic species from deeper water.
I have seen only a handful of specimens from the western Pacific and Indian Oceans, but of these, a
unique specimen from the Mozambique Channel is undescribed. One specimen of Dibranchus
japonicus Amaoka and Toyoshima 1981, described from the Japanese archipelago, is herein reported
from off Cape Town, South Africa, a significant range extension.

One result of this review has been the determination that several forms originally designated
Dibranchus do not agree with the generic diagnosis proposed by Bradbury (1967) and discussed
below. The two Hawaiian forms, Dibranchus erythrinus and D. stellulatus described by Gilbert
(1905), are here referred to a new genus, So/ocisquama, on the grounds of differences in the illicial
skeleton, the peculiar squamation, and the shape of the upper jaw. A unique specimen from Sala y
Gomez Ridge is found to be undescribed and is assigned to the new genus. The genus Dibranchus

'Research Associate, Department of Ichthyology, California Academy of Sciences, San Francisco, California 94118.

259

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was proposed by Peters (1876) to include ogcocephalids with only 2 gills on each side; that is, only
the second and third arches bear gill filaments, a unique character for lophiiform fishes. At the time,
only 2 other ogcocephalid genera were known to Peters, Halieutaea Valenciennes 1837 from the
Orient and Ogcocephalus Fischer 1813 (known to Peters as Malthe Cuvier 1816) from the New World;
both have gill filaments on the second and third arches plus the anterior side of the fourth arch, referred
to as gills 2%. [A third genus with gills 22, Halieutichthys Poey (in Gill 1863), had been described
about a dozen years before Peters proposed Dibranchus, but was evidently unknown to him.]

Fifteen years later Alcock (in Wood-Mason and Alcock 1891) proposed Malthopsis, another
genus with only 2 gills on each side. Malthopsis was said to resemble Ogcocephalus in having a
hornlike rostrum and the head depressed and formed as a “triangular wedge,” but these statements
about characters were too general or imprecise to separate Malthopsis from Dibranchus as new species
were discovered, with the result that Ma/thopsis had assigned to it species which later were reassigned
to Dibranchus on the basis of additional characters including the illicial skeleton and features of
squamation (Bradbury 1967). Eventually there were 11 nominal genera of ogcocephalids described,
7 of these characterized by having gills only on the second and third arches. One of these was
Halieutopsis Garman 1899, which also came to be confused with Dibranchus (summarized in
Bradbury 1967).

Attempting to redefine the limits of ogcocephalid genera, Bradbury (1967) showed that Di-
branchus differed from Malthopsis in having simple tubercles, not bucklers as seen in Malthopsis,
and in having the frontal bones of the cranium forming a groove for the illicitum instead of the tubular
canal seen in Malthopsis. Halieutopsis was shown to share with Dibranchus the character of tubercles,
but Halieutopsis has a flat cranial roof, no groove or canal. Halieutopsis was further shown to have
extra lateral line scales on the ventral surface of the body at or anterior to the anus, lacking in
Dibranchus, while Halieutopsis lacked the well-developed pads of teeth on the ceratobranchial V
bones seen in Dibranchus. Finally, the morphology of the illicial bone was found to be autapomorphic
for each genus, and the implication was that the illicial bone morphology, as far as it was studied,
established monophyly for each genus (Coelophrys excepted because no material of that genus had
been available for study).

Although not all species were available for study of the illicial morphology at that time, the illicial
bone in Dibranchus was shown to differ strongly from that in Halieutopsis in having two large
foramina located ventrally, one in each of the two lateral processes which form the articulation of the
illicial bone with the pterygiophore (Bradbury 1967, fig. 1). Further, Dibranchus lacks the median
dorsal process prominent in both Halieutopsis and Malthopsis. Employing these characters, the
following species originally described as belonging to Dibranchus were reassigned to Halieutopsis:
D. micropus Alcock 1891, D. stelliferus Smith and Radcliffe and D. simulus Smith and Radcliffe
(both in Radcliffe 1912) (see also Bradbury 1988). Similarly, the following species originally assigned
to Malthopsis by Garman 1899 were reassigned to Dibranchus: D. sparsa, D. erinacea, D. spinosa,
and D. spinulosa (spinulosa was treated as a junior synonym of spinosa). Finally, two monotypic
genera described by Garman 1899 were treated as junior synonyms of Dibranchus: Dibranchichthys,
with one species, D. nudivomer, and Dibranchopsis with one species, D. spongiosa (Bradbury 1967).
Characters of the illicial bone are further assessed herein because of availability of new material.

I have not been able to study all nominal species of Dibranchus. I herein place D. alberti Nunez
1967 in the synonymy of D. spinosus (Garman) 1899 on the basis of photographs published with the
original description. Listed below are 3 nominal species that may more correctly be assigned to
Halieutopsis and a fourth one that I cannot assign without examining the holotype.

(1) Il have had an opportunity to examine specimens at the Zoological Society of India including
ZS! 13028, the published number for the type specimen of Dibranchus nasutus Alcock (also listed
in Menon and Yazdani 1968). I found the remains of two specimens in the jar. The specimens were
largely disintegrated, but there was a fragment of cranium containing the esca of one, and frontal

BRADBURY: DIBRANCHUS REVIEW 261

bones and esca of the second, enough to establish that it is incorrect to assign them to Dibranchus.
The remains of the frontal bones showed them to be flat, not forming a median groove in the roof of
the cranium as in Dibranchus, and the escas had the two ventral lobes each spherical and separated
from one another, not joined as in Dibranchus. Both characters are consistent with Halieutopsis, but
I could not ascertain the species; the escas were most like H. vermicularis.

(2) Dibranchus nudiventer Lloyd, 1909 was described from a single specimen “75 mm in length”
from the Bay of Bengal. This specimen had the rostrum projecting beyond the mouth and the esca in
the form of “a pair of fleshy balls with a pair of filaments above and between them.” Neither of these
features agree with Dibranchus; they agree with Halieutopsis. The description of the esca suggests
H. vermicularis.

(3) Dibranchus infranudus de Beaufort, 1962 (in de Beaufort and Briggs 1962) is based upon a
single specimen, 80 mm, from the Flores Sea. Without having seen the holotype, I tentatively assigned
this name to the synonymy of Halieutopsis simulus (Smith and Radcliffe, 1912) in a review of
Halieutopsis (Bradbury 1988).

The holotype of Dibranchus infranudus de Beaufort is in the Institute of Taxonomic Zoology,
ZMA 101.877 (Nijssen et al. 1982).

(4) Dibranchus obscurus Brauer, 1908 was based on a single, 12-centimeter specimen from the
Gulf of Aden. Not having seen this specimen and lacking information on diagnostic characters, I am
unable to place it.

METHODS

Methods are those developed for a study of the related genus Ogcocephalus (Bradbury 1980)
and modified for a study of Halieutopsis (Bradbury 1988). Length of disk margin is the distance taken
on the right side from the front of the posterior swelling of the mandible just at the corner of the mouth
to the distal end of the subopercle excluding the subopercular spine. Skull length is the distance from
the upper lip to the slight pit that can be felt with the tip of the calipers between the cranium and the
anteriormost vertebra. Cranium width is the distance between the points of greatest concavity just
behind the orbits. Eye width is greatest width of orbit. Other measurements are standard measure-
ments: distance from jaw to anus, distance from jaw to anal fin, distance from snout to dorsal fin,
interorbital width, jaw length, and mouth width. Meristic characters studied include numbers of rays
in the pectoral and dorsal fins and numbers of neuromasts in certain lateral-line series. Neuromasts
are free neuromasts (lie on the surface of the skin) but are cupped in modified tubercles (seen in Fig.
Ac; Fig. 5c, d).

In Dibranchus, the lateral line is interrupted at the transition from dorsal surface of disk to lateral
sides of tail, resulting in 2 separate series, a disk lateral line and a tail lateral line. Disk lateral line
elements were too difficult to count accurately so were not used. The tail lateral line commences just
posterior to anus and runs onto the base of caudal fin. (A free neuromast without lateral-line scale
often occurs about half way out on the caudal fin, but is omitted from the count because it may be
difficult to see and its presence cannot be verified by the presence of the modified lateralis scale.) The
other lateral-line series used in this study are the preopercular series, the subopercular series, and the
dorsolateral branch of the subopercular series.

In lophiiform fishes, the first vertebra is fused to the skull; it is not included in the vertebral count
in this study. Counts begin with the first free vertebra and include the hypural fan. All vertebral counts
were made from radiographs.

In Dibranchus the surface sculpturing of tubercles and pattern of squamation are useful characters
for discriminating species, but to a large extent the characters are difficult to convey descriptively. It
was finally determined to restrict detailed description to the ventral surface of the tail, the portion of
the integument that shows in the least bewildering way the kinds of variation seen (Figs. 9, 10). The

262 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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ventral surface of the tail is relatively flat with a clear boundary row of tubercles on each lateral edge,
herein called principal rows.

Museum collections are indicated by standard symbolic codes (Leviton et al. 1985; Leviton and
Gibbs 1988).

Dibranchus Peters

Dibranchus Peters, 1876:736 [type species Dibranchus atlanticus Peters, 1876, by monotypy; gender mascu-
line]; Bradbury, 1967:413 [diagnosis].

DIAGNOSIS. — Gills 2; holobranchs present on 2nd and 3rd arches only. Scales in the form of
tubercles, not bucklers. Frontal bones of skull modified to form a median groove for the illicitum.
Illicial bone with each lateral process pierced by a large foramen; no long dorsal median process.
Tooth patches on ceratobranchial V bones contiguous with one another, meeting all along midline.
Tail lateral line commences posterior to anus; no lateral-line organs beside, or anterior to, the anus as
in Halieutopsis.

DESCRIPTION. — Disk variable in size and shape depending upon preservation, from oval or
bell-shaped when subopercular bones (which form lateral margins of disk) and pectoral pedicels
collapsed against trunk, to strongly triangular when subopercles and pectoral pedicels flared away
from trunk. Most species with prominent posteriorly directed, medially-curved spines at posterior
ends of subopercles, these subopercular spines multifid, often with largest spinelets curving dorsally.

Body depressed but cranium elevated above general surface of disk so eyes are directed laterally
and anteriorly. Rostrum composed of an array of closely-spaced tubercles forming a short shelf over
esca; no median horn, although median tubercle may be larger or longer than those to the sides. Illicial
cavity small, not cavernous as in some Halieutopsis. Esca (Fig. 1a) broader than high, consisting of
a median dorsal lobe continuous with 2 ventral lobes which are continuous with one another across
the midline and flared out laterally, subtended by a ridgelike edging, sometimes membranous. On
posterior side of esca, ventral lobes strongly separated, spherical (can be seen in Fig. 6a). Olfactory
organs sexually dimorphic: in females, organs relatively small and containing a few small lamellae,
the nostrils small and round; in males, olfactory organ large, swollen by enlarged lamellae within,
posterior nostril a wide gaping slit (see analysis below). Dorsal rays 5 or 6, rarely 4 or 7; pectoral rays
usually 13-15, range 12—16 (Table 1). Anal rays always 4, caudal rays always 9. Preopercular lateral
line series 14, subopercular series 4~7, dorsolateral branch of the subopercular series nearly
invariably 3. Lateral line interrupted just as it descends from the disk onto the tail. The lateral line
organs on the disk are difficult to see and count, so these were not studied. Tail lateral line commences
on the ventral side of tail just posterior to the anus, thence continuing along lateral sides of tail onto
base of caudal; tail lateral line 8—15 (Table 4).

Vertebral count | 7—20, usually 18 in D. atlanticus, usually 19 in other species (Table 3).

Teeth in oral and gill cavities simple, small, recurved, posteriorly hinged Type 4 teeth as described
by Fink (1981, table 1) for Dibranchus hystrix (his Dibranchus scaber) with a single exception. That
is the new species D. velutinus described herein as having bristlelike teeth (the true nature of teeth in
D. velutinus has not been studied for lack of material). Teeth in bands on jaws, visible when mouth
closed. Scanning electron micrographs of premaxillary teeth of 4 specimens of D. atlanticus varying
in size from 39.5 to 131.5 mm SL (Fig. 2) show that numbers of rows of teeth and numbers of teeth
within each row increase with size of fish; a similar pattern was seen in D. spongiosus and in D.
spinosus (not pictured). Palatal teeth present only in 5 species. Each ceratobranchial V with a broad
patch of teeth, the two patches always completely contiguous along midline.

Figures 3a and b show the general structure of gill rakers for most species of Dibranchus
(exceptions are velutinus and spongiosus). Pictured is D. atlanticus with relatively stout pedicels and

BRADBURY: DIBRANCHUS REVIEW 263

2mm

FiGurE 1. Frontal views of escas. (a) Dibranchus spinosus, 133.0 mm SL (ZMUC, GALATHEA Sta. 739). The esca in most
species of Dibranchus looks like this as long as the gland has not become distorted in the preservation process. The two ventral
lobes are less differentiated than in most other genera (although well-differentiated spherical lobes are visible from the posterior
view as seen in Fig. 6a). There is often a fringe or membrane along ventral margin. (b) Solocisquama stellulata (Gilbert),
holotype USNM 51595, 47.8 mm SL. (c) Solocisquama erythrina (Gilbert), holotype USNM 51642, 133.5 mm SL.

broad tooth plates with numerous teeth in a subspherical cluster. In some eastern Pacific species,
pedicels are slender with fewer teeth. In D. velutinus, teeth appear absent. In D. spongiosus (Figs. 3c,
d), the dome-shaped tooth plate appears to cover the pedicel and spreads out on the gill arch. The gill
rakers in the following species were studied only by gross examination — no SEM or clearing/stain-
ing: D. accinctus, D. cracens, D. discors, D. japonicus and D. velutinus.

Modified scales known as tubercles are, in Dibranchus, typically pyramid-shaped with 8—14
ridges radiating from a central spine and separating facetlike surfaces. Examples shown are at/anticus
(Figs. 4a, b) with pronounced ridges decorated by spinules, D. tremendus (Figs. 4c, d) with
barely-defined facets and thick terminal spines, and D. nudivomer (Fig. 5d). Lateralis scales are
crescent-shaped (Fig. 5c) with a central foramen for the protruding neuromast. An individual lateralis
scale usually lies between two good-sized tubercles, visible in the near background in both Figs. 4c
and Sd.

Illicial bone. In D. atlanticus, D. tremendus, D. hystrix, D. sparsus, D. spinosus, D. erinaceus,
D. nudivomer, D. cracens, and D. spongiosus, illicial bone with a large foramen on each side ventrally,
piercing on each side the lateral processes that articulate with pterygiophore (Fig. 6b). Lateral
processes separated by a broad high median space. Dorsal median process completely absent, but a
very short, blunt dorsal process on each side of central mass. Dibranchus discors shows this same
conformation but is compressed from side to side (Fig. 7a). (This feature was not studied in D.
accinctus, D. japonicus, and D. velutinus.)

Sexual Dimorphism in Olfactory Organs. Sexual dimorphism in olfactory organs has been
reported in lophiiform groups. Bertelsen (1951) reported that males in most ceratioid anglerfishes
have larger olfactory organs with more and larger lamellae than females. Bertelsen, Pietsch, and
Lavenberg (1981) later determined that metamorphosed males are macrosmotic in all ceratioids
except Ceratiidae and Neoceratiidae. Caruso (1975) reported for Lophiidae that Lophius has sexual
dimorphism in the olfactory organ, but slight and “easily overlooked,” while in Lophiodes and

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TABLE 1. Relationship of standard lengths and sex in Dibranchus atlanticus to numbers of olfactory lamellae.
Specimens with large flag-shaped lamellae always had testes. Specimes with small nub-like lamellae or no
identifiable lamellae (signified by —) always had ovaries. Specimens from the Gulf of Guinea are from 2 lots:
ZMUC Atlantidae Sta. 120, 18 examples, and ZMUC Atlantidae Sta. 135, 20 examples; total, 38 examples. Of
these the gonad was indeterminable in 12 (not shown in Table). Specimens from the Atlantic coast of NE South
America are from seven lots all from Oregon stations, as follows. (Numbers in parentheses are numbers of
examples.) CAS 42661 (3), Sta. 2005; CAS 42662 (10), Sta. 2007; CAS 42663 (3), Sta. 2008; CAS 42671 (4),
Sta. 2026; CAS 43673 (4), Sta. 2029; CAS 42674 (13), Sta. 2028; CAS 42690 (3), Sta. 2025; total, 40 examples.

Gulf of Guinea NE South America
Testes Present Ovaries Present Testes Present Ovaries Present
SL Lamellae S15) ber Lamellae SL Lamellae SL Lamellae

102.0 14 91.0 — 96.4 20 141.8 4
93.0 18 89.0 4 89.2 18 120.0 6
83.5 15 82.0 4 86.3 16 115.6 8
68.0 14 82.0 4 84.4 IS 109.3 6
S35) 11 81.5 4 83.0 7 103.0 =
AWS 6 81.0 3 82.0 14 102.7 —
80.5 4 75.8 7/ 93.4 =
80.5 ~ 74.0 15 82.1 —
80.0 _ 71.8 18 79.8 -
78.5 — 71.8 ws 76.8 _
78.0 4 alee 15 WY _
76.0 _ 70.0 7 70.0 =
75.0 4 69.1 14 68.0 —
Wd 0 64.0 12 65.7 —
WS 5 63.2 10 59.1 =
61.0 = 61.7 12 53.0 =
56.5 = 56.2 7 51.5 —

SIS — 49.2 0

45.1 0

Lophiomus, sexual dimorphism of olfactory organs is well developed. Bradbury (1988) reported that
males were macrosmotic in the ogcocephalid genus Halieutopsis. Since variation in size of olfactory
organs was noted in D. atlanticus and other species of Dibranchus, a test was made to determine
relationship between size variation and sex.

Two samples of D. atlanticus were chosen, 40 from off the Atlantic coast of northeastern South
America, and 38 from the Gulf of Guinea. Each sample was sorted into two groups: those with large
and those with small olfactory organs. The sample from the Gulf of Guinea yielded 6 specimens,
47.5—-102.0 mm SL, with large olfactory organs, and 32 specimens, 35.0—-91.0 mm SL, with small
olfactory organs. All specimens with large olfactory organs, upon dissection, proved to have testes.
Of the other group, 20 specimens, 51.5—-91.0 m SL, had ovaries, and 12, 35.0-59.0 mm SL, were
indeterminable by gross dissection. For the sample from off northeastern South America, 19
specimens, 56.2—-96.4 mm SL, had large olfactory organs as well as testes; 2 others, 45.1 and 49.2
mm SL, also had testes but did not yet show enlarged olfactory organs; 19 specimens, 51.5—141.8
mm SL, that had small olfactory organs had ovaries. We may conclude that mature males are
macrosmotic, as has been found in Lophiidae and most ceratioids (Caruso 1975; Bertelsen et al. 1981).

In large olfactory organs of males, the posterior nostrils are long horizontal slits, the lips of the
slit generally gaping so the opening appears large; within can be seen enormous, overlapping,
flag-shaped olfactory lamellae in the center of the organ, tapering to smaller leaf-shaped lamellae at

BRADBURY: DIBRANCHUS REVIEW 265

7 NSS, as «
Vays
YS . "s

FIGURE 2. Scanning electron micrographs showing that premaxillary teeth in Dibranchus atlanticus increase in numbers of
rows and numbers of teeth within rows with increasing standard length. Standard lengths as follows: (a) 39.5 mm; (b) 72.5
mm; (c) 81.0 mm, all from CAS 76446; (d) 131.5 mm, CAS 76452. Scale bars = | mm.

either end. The skin covering the olfactory capsule bulges to accommodate the enormous lamellae.
The olfactory lamellae were inspected and counted by simply looking into the posterior nares when
large, but, in small olfactory organs, the relatively small rounded posterior nares sometimes made it
difficult to count lamellae. The opening was enlarged by a cut in these cases, but even so, it could be
difficult to determine whether the material within was lamellar epithelium or mucous or other material.
Simple methods such as forcefully irrigating, scraping, or drying the structure seldom clarified
whether or not lamellae were present. It was clear, however, that none of the specimens with ovaries
has large, overlapping, flag-shaped olfactory lamellae as seen in males. Rather, the females had small
lamellae or only nubbins of epithelium.

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FIGURE 3. Scanning electron micrographs showing (a) second gill bar with gill rakers and (b) individual gill raker in
Dibranchus atlanticus, 131.5 mm SL, CAS 76452, and the same (c and d) for D. spongiosus, 115.0 mm SL, CAS 82252. Scale
bars for a and c = | mm, for b and d= 0.5 mm.

As can be seen in Table 1, not only do males have larger lamellae than females, they have many
more of them. Males show an increase in number of lamellae with increase in standard length; females
do not show this pattern although data for females is admittedly incomplete. In any event, although
small size fish 50-60 mm SL or smaller may not show it, mature males in Dibranchus atlanticus are
shown to have far more extensive nasal epithelia than females and seem to increase the area of these
epithelia with growth. Similar observations in species of Dibranchus of which large samples were
available (D. spinosus, D. nudivomer, D. tremendus, and D. spongiosus) showed the same pattern,
and all specimens with enlarged olfactory organs of other species proved to have testes with one
exception. The exception is the new species D. discors, in which males have enlarged olfactory organs,

BRADBURY: DIBRANCHUS REVIEW 267

FiGuRE 4. Scanning electron micrographs showing (a) tubercles from a section of skin posterior to eye and (b) principal
tubercles of the tail in Dibranchus atlanticus, 131.5 mm SL, CAS 76452, and the same (c and d) for D. tremendus, 155 mm
SL, USNM 320332. Tubercles are decorated with spinules in D. atlanticus but not in D. tremendus; spines of tubercles are far
longer in D. tremendus than in D. atlanticus. Scale bars= 1mm. In the near background in (c) is seen a specialized
crescent-shaped lateralis scale, placed as usual between two tubercles that mark the poles of the long axis of the scale. The
neuromast protrudes through a pore in the middle of the scale.

but much less pronounced than in congenors, so that it is difficult to be sure one has a male without
identifying testes.

Bioluminescence. No light organs have been observed in Dibranchus, but bioluminescence
nevertheless may play a role in the life of Dibranchus atlanticus. Crane (1968) reported that the skin
of the dorsal surface of freshly-caught specimens from 366 m off Brunswick, Georgia, showed
fluorescence, suggesting a luciferin might be present. Adrenalin hydrochloride injected into the heart
of one fish induced it to emit a low level glow over the entire dorsal surface, although neither the esca
nor ventral surface of the fish emitted light. Later, skin and escas from frozen fish were tested for
spontaneous light emission as well as stimulated by various reagents, resulting in the determination
that the esca and the skin from both the dorsal and ventral surface have bioluminescence capability.
The function of this feature is as yet obscure.

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FIGURE 5. Scanning electron micrographs. (a) second gill bar with gill rakers and (b) individual gill raker in Solocisquama
erythrina, 135.5 mm SL, BPBM 29270; in this species and in S. ste/lulata, the pedicels are stouter and longer that in Dibranchus.
(c) crescent-shaped lateralis scale with neuromast at center and (d) tubercles in skin of disk posterior to eye in Dibranchus
nudivomer, 98.5 mm SL, CAS 82261. A specialized lateralis scale in near background in (d) lies between its two associated
tubercles Scale bars for a, b, and d = | mm, for c = 0.5 mm.

DISTRIBUTION. — Two species in the Atlantic, both occurring in the eastern as well as western
Atlantic and Gulf of Mexico. Nine species in the eastern Pacific, from the coasts of Mexico to Peru
and west to the Galapagos; of the nine, two are known only from the Galapagos archipelago, and one
is trans-Pacific. One other species is known from the western Pacific, but its range is shown herein
to extend to the western Indian Ocean. One species, described herein from a single specimen, is known
only from the western Indian Ocean.

BRADBURY: DIBRANCHUS REVIEW 269

\)

lmm

FIGURE 6. (a) Postero-lateral view of cleared and stained illicium of Dibranchus tremendus, 96.5 mm SL, UF 25923. Internal
structures (stippled) are the elongate pterygiophore, left, and illicial bone, right. The dorsal lobe of the esca appears to the right
instead of dorsad because the esca has been bent down relative to the pterygiophore; note at its distal tip a large opening for
discharge of gland’s secretion. Scale bar = | mm. (b) Three views of distal end of pterygiophore and illicial bone in Dibranchus
(generalized; from Bradbury 1967) as follows: side view (upper left), dorsal view (lower left), and frontal view of illicial bone
(right). Note on the illicial bone the very short dorso-lateral process on each side and the large ventro-lateral lobe pierced by
a foramen on each side. Each ventro-lateral lobe extends laterally and posteriorly to form the articulating processes of the illicial
bone.

270 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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E
=

FiGurRE 7. Frontal views of illicial bones. (a) Dibranchus discors, 87.2 mm SL (CAS 47195); (b) Solocisquama stellulata,
76.0 mm SL(BPBM 17941). Bone spicules occasionally occur within the foramen of the lateral process in Dibranchus as seen
on the left in this specimen. On the left it can also be seen that the lateral process is malformed.

KEY TO ADULTS OF EASTERN PACIFIC SPECIES OF DIBRANCHUS

A key to only the eastern Pacific species of Dibranchus 1s offered here because the species from
other regions are few and easily separable. The two Atlantic species, D. atlanticus and D. tremendus,
are easily separated by their squamation (Fig. 4). The two species found in the western Pacific, D.
hystrix and D. japonicus (Fig. 9b, f), can also be distinguished from one another by their squamation,
as can D. accinctus and D. japonicus in the Indian Ocean (Fig. 9c, f).

laridicethupresent‘onipalat]e: 52.5 ae ea eee ls aepnesetae os wR ch ete copie fo ia, eal eee Sorcerer 2
ibsNotecthompalaten sa. ne oe Seach ORs Om Aa eo we terete ain SUA oe Clee oe pee © cee ee 6
AE, VFLINONOANAMES DUNNE ONVOINS? 5 6a cone cama oo ood no sce oss o de ose soo oe D. nudivomer
Zbwleethionibothpalatinesrand vomier at. 2s cseeMs. i oo cusaths Syiepeee = 5a cue = ce fe ieee eee 3
3a. Tubercles present in skin of eyeballs and on ventral surface of disk. Mouth width 14% ormoreofSL.......... 4
3b. No tubercles in skin of eyeballs, or on ventral surface of disk (except a few around pelvic bases). Mouth very nar-

TOM sIS WAGtHIIES/o1OrlessiOf Slew scenees. ay cueee)s. \-t an See ROS GME) ues eer ee D. discors
4a. Preopercular lateral-line count usually 2. A reticulate color pattern often discernable on dorsal surface (Fig. 15a). A

SmallispeciesmeachingalessithanySonmnnnts lms i- mene men alten mem arn ieee ee D. sparsus
4b. Preopercular lateral-line count usually 3. No reticulate color pattern on dorsal surface. Larger species, reaching

Meare SONI S Loe ce cap ou copuce se. ste ty n cP PAIR Gove) UE igs, onic gee en GE co) rey rena eect A 5
5a. Very small tubercles with slender spines covering body; enlarged tubercles are present, but these covered by the

small ones so effect is of uniformly small tubercles (Fig. 15c). Subopercular spine little developed... . . D. spinosus

5b. Tubercles moderate size interspersed with small ones (Fig 15b). Subopercular spine broad, moderately long D. erinaceus
6a. Tubercles so tiny that the texture of the skin resembles velvet (Fig. 14). Mouth very wide, the width 27.5% of SL.

Jawsicoverediwithipapillacs bunyineihie biSilelikeitec ties ment en ie emt eis aan ene eee D. velutinus
6b. Tubercles moderate to large size, skin not velvetlike. Mouth width less than 25% of SL. Small conical teeth in

bands visible-onyjawsr a. 9 as fo aes ke eee ae ee ae ee ates ae Ra OO, Sk ee yf
7a. Large tubercles decorated with spinules radiating in lines from central spine (Fig. 9e). Subopercular lateral-line

COUN OEY Pee ee acer ee Bee en Sete Cece ete ge nile tea tea ER Ce ete nl ee ee are re D. cracens
7b. Tubercles plain; no spinules. Subopercular lateral-line count usually5 ..................+-+--+---- 8
$a: 'Spines oftuberclesslono(Fig. 9b). ee wc ee ee eee ee Oe ee ee ge D. hystrix

SH¥Spines Oltubercles short (Ei SMO f) eee eee kent eee enn ere ese aa es ee nee eae D. spongiosus

BRADBURY: DIBRANCHUS REVIEW ant

Dibranchus atlanticus Peters, 1876
Figs. 2, 3a—b, 4a—-b, 8, 9d

Dibranchus atlanticus Peters, 1876:738 (original description, | specimen, the type, off West Africa; 675 m;
holotype not seen); Gunther, 1887:59 (W coast of Africa, 658 m): Vaillant, 1888:343 (off Cape Verde L.,
405 m); Goode and Bean, 1896:501; Beebe, 1937:207 (off Bermuda; pelagic specimens from depths to 600
m); Cadenat, 1937 (9°32'N, 16°25’'W, 250m); Amold, 1949:299 (off New England, 117m); Rees,
1963:1513 (continental slope of Canadian Atlantic); Crane, 1968:410 (off Georgia from 366 m; biolumi-
nescence); Hoese and Moore, 1977:144 (Gulf of Mexico off Texas and Louisiana; key; partial description);
Golovan, 1978:231 (northwestern Africa; benthic zonation); Pietsch, 1981:387 (osteology; phylogeny);
Uyeno et al., 1983:250 (off Surinam and French Guiana, 180-910 m; description; photograph); Matallanas,
1987:121 (off Guinea-Bissau, NW Africa).

Halieutaea senticosa Goode, 1881:467 (holotype USNM 26175; coast of southern New England, 412 m).

REMARKS. — Iam indebted to C. Karrer, Zoologisches Museum Hamburg, who informs me that
“nearly all the material of Peters is in the Zoologisches Museum, Berlin” (pers. commun. 1991).
Presumably the holotype of Dibranchus atlanticus is among this material. In any event, figures
accompanying the original description give an exceptionally clear picture of the species including the
diagnostic spinules on tubercles.

DIAGNOSIS. — Tubercles have ridges radiating from the apical spine; these ridges heavily edged
with spinules, especially the largest tubercles (Figs. 4, 9d); thus, tubercles in this species appear
multi-spined. Vertebral count usually 18 (Table 3). Subopercular lateral-line count usually 5 (Table
4). No palatal teeth.

DESCRIPTION. — Based on 432 specimens 30.0—-138.0 mm SL. Counts and body proportions
given in Tables 3—5. Skeleton firm; skin tough, studded with shagreenlike tubercles among which
protrude numerous enlarged tubercles with hard sharp spines. Rostrum short, median tubercle slightly
overhanging illicial cavity (juveniles have rostrum more elongated). Esca and nasal capsules as in
generic description.

Teeth as in generic description. No palatal teeth. Gill rakers as in generic description; usually 6
gill rakers on each side of second arch.

Tubercles mainly of 2 sizes; relatively small simple tubercles evenly distributed over entire body
so skin resembles shagreen. Interspersed are large tubercles featuring prominent facets, the ridges
between facets studded with rows of spinules forming a radiating pattern from the long central spine
(Fig. 4a). Largest of these occur on the sides of the disk, where terminal spines are bifid, and on the
dorsal and lateral sides of the tail. Spinules on edges of tubercle facets well developed in specimens
about 35 mm SL or more; in specimens about 30-35 mm SL, only largest tubercles have spinules
developed; tubercles in specimens under about 30 mm SL lack spinules. Subopercular spine promi-
nently long with 6-8 spinelets. Only small tubercles occur on the ventral surface of the disk, where
the skin is uniformly shagreenlike; this type of squamation also occurs in the skin on the eyeballs.

Ventral surface of tail with principal tubercles wide, low, covered with radiating spinules as
elsewhere on body, terminating in short stout recurved spines (compare Fig. 9d with others pictured
in Figs. 9 and 10). Similar tubercles lie between the principal rows in 2 longitudinal rows from the
anus to the anal fin base. Small tubercles thickly distributed between bases of large tubercles.

Fins, except dorsal, usually with tubercles on bases of rays.

Dermal cirri associated with channels for lateralis organs, especially around neuromasts.

Pectoral fins slender, sturdy relative to most other species; pelvic fins slender.

Color. Fresh specimens tinged with pink on light bluish gray or brown background. Fins very
pale to brilliant rose red. Pinks and reds disappear completely in preservative; preserved specimens
uniformly pale gray or light brown. Eastern Atlantic specimens frequently with a reticular pattern on
the dorsal surface of the disk that is darker than the ground color. Fins the same color as the body in

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 5

NO
—
tO

FIGURE 8. Dorsal views of similar-sized specimens of Dibranchus tremendus, 116.0 mm SL, USNM 320326 (left) and D.
atlanticus, 113.0 mm SL, USNM 186337 (right).

preserved specimens. Peritoneum black. Dark brown specimens occur, but rarely; in these, cirri
associated with lateral line organs on subopercle and tail dark brown or black, fins dark or with dark
blotches.

DISTRIBUTION. — Atlantic coasts of Canada and United States, Gulf of Mexico, Caribbean, and
coast of South America to Rio de Janiero, Brazil; in the eastern Atlantic, the Gulf of Guinea to Angola.
Bathymetric range: the vast majority of collections from 300—700 m. Two lots that I have seen came
from depths less than 100 m off French Guiana. A few lots have come from 700-1000 m, including
samples from the Gulf of Mexico, Straits of Florida, off Surinam, and off Angola; there is one record
from 1260 m from off Angola.

GEOGRAPHIC VARIATION. — Intraspecific variation occurs in this wide-ranging species. Color
differences between eastern and western Atlantic populations are observable: specimens from the
Gulf of Guinea often have reticulate markings on the dorsal surface of the body, usually vague,
occasionally strong, which are not seen in western Atlantic material.

BRADBURY: DIBRANCHUS REVIEW 23

There was also a difference in pectoral fin ray count between eastern and western Atlantic
populations (Table 2); note that the western Atlantic material was handled as five populations based
on geographic regions, as follows: Gulf of Guinea, Atlantic coast of the United States (US), Gulf of
Mexico, Caribbean, and coast of NE South America.

MATERIAL. — Numbers in parentheses give number of specimens; for lots used in morphometric
analyses, standard lengths follow the number of specimens. Method of capture, when known, is by
bottom trawls or dredges unless otherwise noted.

Atlantic Coast of United States: ARC 8600689 (1, 112.4), 43°07'N, 61°28’W, 500 m. ARC
8600690 (2, 65.0—125.3), 42°52'N, 62°35’W, 326 m. ARC 8600691 (1, 129.2), St. Pierre Bank. ARC
8600693 (1, 91.1), 42°47'N, 63°30'W, 600 m. ARC 8600694 (2, 94.0—97.7), 44°35'N, 58°44'W, 81
m. ARC 8600695 (1, 87.7), 42°46’'N, 63°30'W, 417 m. ARC 8600696 (1, 90.2), Nova Scotia Banks,
6’ Issacs Kidd. ARC 8600697 (1, 85.4), 43°56’N, 58°40'W, 373 m. CAS 3200 (5, 27.8-56.9), ISELIN
Cruise 7310, Sta. 45, 36°42.6'N, 74°38.2'W, 390 m. CAS-SU 9519 (4, 45.7—108.6), off Newport, R.
I. CAS 42667 (2, 76.0-92.0), COMBAT Sta. 362, 34°18'N, 75°51'W, 402 m. CAS 42669 (1, 71.0),
SILVER BAY Sta. 2070, 29°13’N, 79°59'W, 375 m. CAS 76452 (1, 131.5), DELAWARE Cr. 60-1,
Sta. 8, 41°54’N, 68°13'W-41°54'N, 68°17.5'W, 220-238 m. CAS 42670 (1, 77.5), PELICAN Sta.
58, 29°59'N, 80°10'W, 296-305 m. CAS 42700 (2, 45.6-82.0), SILVER BAY Sta. 469, 29°36'N,
80°10’ W, 348-366 m. CAS 42701 (1, 105.0), SILVER BAY Sta. 445, 28°03'N, 78°44'W, 914-951
m. CAS 42702 (1, 42.0), COMBAT Sta. 499, 29°50’N, 80°10'W, 366 m. CAS 42704 (3, 48.0—123.0),
DELAWARE Sta. 59-10, Tow 3, 39°57'N, 70°58’W, 320-366 m. CAS 42705 (1, 76.0), COMBAT
Sta. 445, 25°15’N, 79°13’W, 366 m. CAS 51206 (1, 90.0), COMBAT Sta. 436, 24°13'N, 81°42'W,
549 m. CAS 63251 (2, 68.7—-90.4), COMBAT Sta. 431, 29°57'N 80°08'W, 384 m. UF 25980 (2,
79.5—83.0), PELICAN P-70, 29°32'N, 80°08'’W, 274-348 m. UF 25981 (4, 54.0-106.0), SILVER
BAY Sta. 220, 29°29'N, 80°09'W, 329-348 m. USNM 186335 (1, 77.8), DELAWARE Cr. 59-7, Sta.
8B, 36°42'N, 74°41’W, 366 m. USNM 186337 (5, 75.3—113.0), DELAWARE Cr. 59-7, Sta. 4B,
36°05'N, 74°43'W, 366 m.

Gulf of Mexico: CAS-SU 17442 (4, 66.2—79.9), OREGON Sta. 501, 27°55.5'N, 91°32.5'W, 402
m. CAS 42668 (4, 80.2-92.1), OREGON Sta. 126, 29°02’N, 88°34.5'W, 357 m. CAS 42689 (1),
OREGON Sta. 1019, 24°16'N, 83°22'W, 686 m. CAS 42692 (1), OREGON Sta. 319, 29°20'N,
87°25'W, 576-622 m. CAS 42695 (4, 88.0-95.5), OREGON Sta. 270, 29°23’N, 87°25’W, 402 m.
CAS 42697 (13), OREGON Sta. 480-484, 28°56.5’-28°58'N, 88°39-88°42.5'W, 366-439 m. CAS
42698 (4, 37.0-79.5), SILVER BAY Sta. 1197, 24°15'N, 83°36’W, 914m. CAS 42699 (2,
76.5-87.0), SILVER BAY Sta. 1196, 24°11'N, 83°21.5’W, 732 m. CAS 42706 (1), OREGON Sta.
307, 29°00'N, 88°35'W, 402 m. CAS 76453 (1, 105.1), OREGON Sta. 279, 29°11'N, 86°11'W, 558
m. FMNH 45748 (1 ), OREGON Sta. 795, 29°15'N, 87°49'W, 421-549 m. FMNH 46730 (16,
51.7-97.0), OREGON Sta. 472-484, 29°06'N, 88°27'W—28°57'N, 88°42.5'W, 348-439 m. FMNH
46732 (2), OREGON Sta. 272, 29°16'N, 86°39'W, 395 m. FMNH 59915 (1), OREGON Sta. 726,
22°41.9'N, 86°41.2'W, 412 m. FMNH 64020 (1), OREGON Sta. 640, 29°O1'N, 88°24'W, 649-869
m. USNM 158230 (4) OREGON Sta. 489, 27°44'N, 85°09'W, 465 m. USNM 158536 (2), OREGON
Sta. 127, 29°02'N, 88°34'W, 424-472 m. USNM 158837 (3, 63.4-93.7), OREGON Sta. 1389,
29°00'N, 88°33'W, 408 m. USNM 158839 (1, 95.8), OREGON Sta. 1400, 29°30'N, 87°08'W, 384
m. USNM 159150 (1), OREGON Sta. 1283, 29°06'N, 88°19’'W, 476m. USNM 159160 (3,
45.6—71.0), OREGON Sta. 1282, 29°10'N, 88°03'W, 476 m. USNM 159181 (5, one measured 86.8),
OREGON Sta. 1238, 28°53'N, 88°48’W, 366 m. USNM 219120 (1, 165.0), OREGON II Sta. 11469,
28°S9’'N, 86°49’W, 521 m.

274 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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TABLE 2. Frequency distributions for pectoral fin ray counts in specimens of Dibranchus atlanticus from five
geographic regions of the Atlantic Ocean. Counts were made on both sides.

Pectoral Fin Ray Count

[2 13 14 iS 16 N x
Gulf of Guinea 1 93 Ty 3 — 168 135
Off NE South America = ] 47 50 1 99 AUS)
Carribean — 2 Sy 70 2 126 14.5
Gulf of Mexico = 1 54 60 1 116 14.5
Atlantic off United States - 5 46 49 = 100 14.4

TABLE 3. Frequency distributions for dorsal and pectoral fin ray counts and vertebral counts in species of
Dibranchus.

Dorsal fin Pectoral Fin Vertebrae

Species 4 5 6 7 11 12 13 14 SG 17 tse oe) 20
atlanticus _ 9 74 l — l NO LAK) 22392) 4 4 104 4 =
tremendus 1 11 19 3 -- 2 8 Qa - = — S 25 l
nudivomer — 5 11 1 — 14 20 3° 45 2
discors -- - 3 2 = 3 2 5 - ~ — 2 2 ]
sparsus — 12 Qos = 7 37 17 l
Spinosus = Dip g = fi 55 29 l - = = 42 7
erinaceus 1 ti 9 — i 2S 4 - = _ 3 8 -
velutinus _ — ] 2 l —
cracens - - 3 _ _ 2 4 2 1 ~
hystrix 1 14 IS - — 2 30 4 - = - 2 tO -
spongiosus — 8 16 — eG: 14 28 3 -— = — 6 18 —
Japonicus* = — l 8 Si 6 1 = — ] 7 =
accinctus - - l 2 1 _

Caribbean: CAS 42658 (2, 34.0-50.6), OREGON Sta. 2637, 17°37'N, 63°36'W, 512 m. CAS
42660 (12, 37.0-104.5), OREGON Sta. 3599, 9°00'N, 81°23'W, 457 m. CAS 42665 (6, 60.5—90.2),
OREGON Sta. 3616, 14°23'N, 81°45'W, 457 m. CAS 42666 (2, 36.3-66.6), OREGON Sta. 2353,
11°35'N, 62°41'W, 388-457 m. CAS 42693 (1, 48.7), OREGON Sta. 1911, 12°44'N, 82°14'W, 640
m. CAS 42696 (32, 31.5—-81.5), OREGON Sta. 3635, 16°58'N, 87°53'W, 457—732 m.

Atlantic Coast of South America: CAS 42661 (3, 80.0-105.0), OREGON Sta. 2005, 07°34'N,
54°50’W, 366 m. CAS 42662 (10, 49.0-81.5), OREGON Sta. 2007, 07°34'N, 54°49’W, 412 m. CAS
42663 (3, 52.6-85.2), OREGON Sta. 2008, 07°38’N, 54°43’W, 457 m. CAS 42664 (5, 67.5—90.9)
OREGON Sta. 2012, 07°34’'N, 54°19'W, 274 m. CAS 42671 (4, 63.5—115.5), OREGON Sta. 2026,
07°10'N, 53°07'W, 366 m. CAS 42672 (8, 65.1—95.7), OREGON Sta. 2320, 07°05'N, 52°47'W, 366
m. CAS 42673 (4, 95.0-143), OREGON Sta. 2029, 07°11'N, 52°56’W, 503 m. CAS 42674 (13,
61.5—111.0), OREGON Sta. 2028, 07°11'N, 52°58'W, 457 m. CAS 42685 (1, 95.0), OREGON Sta.
2006, 07°36'N, 54°42'W, 412 m. CAS 42686 (1, 95.0), OREGON Sta. 2009. 07°40'N, 54°47'W, 549
m. CAS 42687 (4, 71.5—85.5), OREGON Sta. 2010, 07°44'N, 54°40'W, 640 m. CAS 42688 (1, 79.5),
OREGON Sta. 2011, 07°46'N, 54°36’W, 732 m. CAS 42690 (3, 71.0—-76.5), OREGON Sta. 2025,
07°12'N, 53°11'W, 329 m. CAS 42691 (1, 57.3), 06°39'N, 52°53'W, 91 m. CAS 42694 (1, 37.0),

BRADBURY: DIBRANCHUS REVIEW 215

TABLE 4. Frequency distributions for cephalic and tail lateral-line counts of species of Dibranchus.

Cephalic Lateral-line Counts

Subopercular Preopercular Tail Lateral-line Counts

Species - 5 Gt l 2 3. 4 8 9 Oa 2 13 ei al
atlanticus 7 1 15 = | 78 2 - 2 25, 4, 24> 15) 43 7
tremendus 2 34 —- = ] 65 = = a Ii 236 3 2 — --
nudivomer — — 29° — = 36 5 5 tO 12 | l
discors — = 10 — — 10 = = — l 2 3 3 ] — -
Sparsus = f AQ? — 43 — = ie iba 7 3 l _
Spinosus = She OR l Sh Pad Sinan) l l sla F319 E20 eels 3
erinaceus — Ay 28.) 2 - Ae — — l Se ad gi l
velutinus = = Do = = 2 2 = —
cracens = = 6 = = 6 jy 2 2 2
hystrix — 36 4 - = 27 1 — | Ss isos si l —-— =
spongiosus 42 OS — 46 = = 8 lon Ly 2

Japonicus = 2 = = = 2 2

accinctus = Z = = — 2 Ys

OREGON Sta. 2084, 01°45'N, 46°46’W, 503 m. CAS 42703 (1), 23°01'S, 40°43'W, off Rio de
Janiero, Brazil.

Gulf of Guinea to Angola: CAS 76446 (101, 35.9-79.2), UNDAUNTED Cr. 6801, Sta. 111,
10°36’S, 13°12’E, 361 m. CAS 76454 (3, 51.0-72.0), GERONIMO Cr. 2, Sta. 221, 03°02’S, 09°16’E,
400 m. ZMMSU P14050 (2, 102.2—109.2), FIOLENT Cr. FAO-1, trawl 29, 03°30’S, 09°41’E, 1000
m. ZMMSU P14051 (1, 66.0), FIOLENT Cr. FAO-1, 08°33’S, 12°35’E, 1260 m. ZMMSU P14053
CeO) HIOLEND Cr FAO-1, trawl 79, 10°S4'S, 13°227E. 520m. ZMUC (22 79:2-85%6),
GALATHEA Sta. 110, 12°05’S, 13°08’E, 1000 m. ZMUC (1, 32.2), GALATHEA Sta. 137, 20°04’S,
11°56’E, 550-585 m. ZMUC (50, 33 measured 35.5—73.3) ATLANTIDE Sta. 120, 02°09'N, 09°27’E,
530-850 and 650-260 m. ZMUC (24, 35.0—102.0), ATLANTIDE Sta. 135, 07°55’S, 12°38’E,
440-360 and 460-235 m.

Dibranchus tremendus new species
Figs. 4c, d; 6a; 8; 9a; 11

Dibranchus atlanticus, Roule, 1916 (not of Peters, 1876):26 and 1919 (Cape Verde Islands, 1300 m; description;
lithographic drawings)
Probable reference: Haedrich and Horn, 1970:393 (New York Bight, 1280 m)

NOTES ON SYNONYMY. — Roule (1916, 1919) had two specimens from the Cape Verde Islands,
one taken at 1300 m and one at 875 m. Which was used for the fine illustrations is not given, but the
specimen drawn is the new species, Dibranchus tremendus, not D. atlanticus.

DIAGNOSIS. — A very large Dibranchus reaching over 190 mm SL. Tubercles without the
radiating rows of spinules seen in D. atlanticus, spines of tubercles extremely long and stout,
especially along sides of tail. Tubercles cover ventral surface of disk. Principal tubercles of ventral
surface of tail (Fig. 9a) very large and occupying entire ventral surface except for a few small tubercles
in the median portion between anus and anal fin base. No palatal teeth. Subopercular lateral-line count
usually 5 (Table 4).

276 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 5

FIGURE 9. Ventral views of tails of species of Dibranchus to show patterns of squamation. Pointers indicate lateral-line
channels on the sides of the tail. A principal row of tubercles is defined as forming the ventral edge of this channel. (a)
Dibranchus tremendus, 192.0 mm SL (holotype), UF 25923; (b) D. hystrix, 125.0 mm SL, CAS-SU 46657; (c) D. accinctus,
175.0 mm SL (holotype), CAS 82221; (d) D. atlanticus, 92.8 mm SL, CAS 42695; (e) D. cracens, 102.0 mm SL (holotype),

D. tremendus in the eastern and western Atlantic, D. hystrix in the eastern and western Pacific, and D. accinctus in the western
Indian Ocean. Two species have tubercles decorated with spinules: D. atlanticus (d) and D. cracens (e). (D. discors, not shown,
has tubercles decorated with spinules, but these are barely macroscopic and sparse.) The fine-tubercled species, D. japonicus
(f), has uniquely leathery skin and prominent stout spines on the principal tubercles.

BRADBURY: DIBRANCHUS REVIEW 77

FiGureE 10. Ventral views of tails of species of Dibranchus to show patterns of squamation. Pointers indicate lateral-line
channels on the sides of the tail. A principal row of tubercles is defined as forming the ventral edge of this channel. Note that
neuromasts may be seen in the channel posteriorly in specimen c. (a) D. sparsus, 81.2 mm SL, MCZ 28716; (b) D. spinosus,
91.0 mm SL, CAS 39921; (c) D. erinaceus, 114.0 mm SL (lectotype), MCZ 28712; (d) D. nudivomer, 101.2 mm SL, CAS
82225; (e) D. velutinus, 99.0 mm SL (holotype), CAS 82223; (f) D. spongiosus, 133.2 mm SL, CAS 82252. Dibranchus
sparsus, D. spinosus, D. erinaceus and D. nudivomer (a-d) are four of the five species in the genus with palatal teeth (the fifth,
Dibranchus discors, is not pictured). Dibranchus sparsus and D. spinosus are fine-tubercled species with principal tubercles
little enlarged, D. erinaceus has prominent principal tubercles, and D. nudivomer has uniformly moderate-sized tubercles.
Species lacking palatal teeth include D. velutinus (e), which has the smallest tubercles of any species in the genus, and D.
spongiosus (f), with approximately uniform moderate-sized but short-spined tubercles.

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES

278

Volume 51, No. 5

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BRADBURY: DIBRANCHUS REVIEW 279

DESCRIPTION. — Based on 34 specimens 36.2—192.0 mm SL. Counts and body proportions
given in Tables 3—5. Skeleton firm; skin pliant, especially on ventral surface of disk. Rostrum broadly
triangular viewed from above, slightly overhanging mouth. Esca and nasal capsules as in generic
description. Illictum of one specimen, FSM 25923, cleared and stained (Fig. 6a); illicial bone agrees
with Dibranchus as pictured in Figure 6b.

Teeth as in generic description. No palatal teeth. Gill rakers as in generic description, pedicels
relatively short; 5 gill rakers on each side of second arch.

Tubercles on dorsal surface of disk relatively widely spaced (Figs. 8, 11), medium to large in
size (no background of small tubercles forming a shagreen as in D. atlanticus). Tubercles on ventral
surface of disk similarly widely spaced but smaller, as are the tubercles on skin surrounding eyeballs.
Tubercles of edge of disk very large, closely spaced, with prominent bifid spines. Subopercular spine
conspicuously long in small specimens, becoming nearly the same size as other disk margin tubercles
in large specimens; usually 4 spinelets. A few enlarged tubercles along dorsal edge of each orbit. Very
large tubercles with extremely long sharp spines occur down the posterior midline of head, continuing
onto tail, and on lateral sides of tail.

Ventral surface of tail (Fig. 9a) with the huge heavily-spined tubercles of the principal rows
occupying most of the space. Between anus and base of anal fin occur 2 irregular rows of lesser
tubercles, with a sprinkling of small tubercles in between, but no small tubercles covering bases of
large tubercles as in D. accinctus.

Fins without tubercles except at the extreme bases of the pectorals and caudal.

Dermal cirri associated with channels of lateralis system, especially neuromasts.

Pectoral and pelvic fins relatively long and slender in small individuals, shorter and broader with
rays very fleshy in large ones.

Color. Ground color of body pale to medium brown; ventral surface in one specimen darker than
dorsal surface, but otherwise dorsal and ventral surfaces about the same. Cirri associated with
lateral-line scales and excavations of ventral surface often dark brown or black. In a few specimens,
each tubercle of dorsal surface of disk with skin at apex colored dark brown, giving effect of evenly
distributed tiny dark spots. Fins the same color as ground color of body, sometimes darker, or
sometimes fin membranes dark.

ETYMOLOGY. — The name tremendus means a Dibranchus to be trembled at, in reference to its
giant size and terrific spines.

DISTRIBUTION. — Caribbean Sea, Gulf of Mexico, western North Atlantic, eastern Atlantic from
Gulf of Guinea to Namibia. Bathymetric range from about 1000—2300 m, with one record 750-896
m; separated bathymetrically from D. atlanticus which usually occurs at depths less than 1000 m.

MATERIAL. — 34 specimens 36.2—192.0 mm SL. HOLOTYPE: UF 25923 (192.0), OREGON II
Sta. 11239, 9°32'N, 76°38'W, 1463 m. PARATYPES: Atlantic Coast United States: MCZ 37822 (3,
143.5-158.0), CAPT. BILL II Sta. 59, 39°59'N, 69°09'W, 750-896 m. VIMS 03504 (1, 174.0), slope
south of Hudson Canyon, 38°51'N, 72°44.1'W, 1333 m. Gulf of Mexico: USNM 320332 (1, 158.0),
OREGON II Sta. 10876, 28°55.2’N, 87°23'W, 1463 m. Caribbean: UF 25924 (3, 104.6—150.0),
OREGON II Sta. 11242, 10°10’N, 76°14’W, 1097 m. UF 25925 (5, 64.2—149.2), OREGON II Sta.
11240, 9°58’'N, 76°29'W, 1271 m. UF 104889 (1, 96.5) same data as for holotype. UF 229293 (1,
53.4), PILLSBURY Sta. 748, 11°24.8'-36'N, 67°10.1'"-06'W, 1865-1783 m. UF 232001 (2,
106.9—116.4), PILLSBURY Sta. 747, 11°46’-S4.7'N, 67°05.7'-05.0'W, 1174-1097 m. UF 232820
(1, 113.5), 12°55'-13°04'N, 71°46.5'-42.0'W, 1317-1298 m. USNM 320325 (1, 132.5), OREGON
Sta. 5639, 11°44’N, 68°43'W, 1006m. USNM 320326 (2, 43.3-116.0), OREGON Sta. 4449,
10°56'N, 67°38'W, 1079 m. USNM 320327 (1, 144.0), OREGON II Sta. 11242, 10°10’N, 76°14'W,
1097 m. Eastern Atlantic, Gulf of Guinea to Namibia: IOAN (1, 32.1), FIOLENT Cr. FAO-1,
Trawl 81, 10°56’S, 13°13’E, 960 m. UF 221621 (1, 82.7), PILLSBURY Sta. 314, 4°58’—52'N, 3°48’E,

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Dias 2342 me WF 221672 (6, 36.2—134.2), PILLSBURY Sta. 309, 4°15’—12'N, 4°27—23’E,
f2e0-1317m- WF 223274 (3, 49.1—-130.2), PILLSBURY Sta. 76, 4°32’-40’N, 9°42'49°W,
1555—1463 m.

One other paratype, UF 232665 (1, 111.0), GILLIS GS-21, cannot be assigned locality data.
Catalog data, specimen label, and GILLIS data sheets at the Florida Museum of Natural History all
agree on the following: 7°11'—7°12.5'N, 79°16'—79°18.5'W, erroneously placing the locality in the
Pacific rather than in the Caribbean where the ship was cruising at the time.

Dibranchus hystrix Garman, 1899
Figs. 9b, 12a

Dibranchus hystrix Garman, 1899:92 (original description, 3 specimens from eastern tropical Pacific, illustra-
tions); Bradbury 1962:2 (lectotype selected, MCZ 28776, from off northern Ecuador, 2196 m).

Dibranchus scaber Garman, 1899:94 (original description, 2 specimens from eastern tropical Pacific, illustra-
tion); Bradbury, 1962:2 (lectotype selected, MCZ 28724, from mouth of Gulf of California, 1820 m).

Dibranchus asper Garman, 1899:101 (original description, holotype from off Acapulco de Juarez, Mexico,
1207 m); Bradbury, 1962:2 (D. asper placed in synonymy of D. hystrix).

DIAGNOSIS. — Tubercles on sides of tail with extremely long spines (Fig. 9b). On ventral surface
of disk, tubercles small, widely spaced so skin seems naked, the effect intensifying with increase in
standard length; in larger specimens, these tubercles sometimes truly absent. Except for 2 short
longitudinal rows of tubercles anterior to anal fin base, principal tubercles occupy most of the ventral
surface of tail (Fig. 9b); no small tubercles in skin covering bases of principal tubercles. Subopercular
lateral-line count usually 5 (this count usually 6 in most eastern Pacific Dibranchus except D.
spongiosus (Table 4). No palatal teeth.

DESCRIPTION. — Based on 22 specimens 40.0-125.0 mm SL. Counts and body proportions
given in Tables 3—S. Skeleton pliant, body often flabby depending on preservation; subopercles
weakly calcified so lateral sides of disk have tendency to roll up in preservative. Skin thin, flabby,
especially soft on ventral surface of disk. Rostrum triangular, slightly overhanging mouth, relatively
small in small individuals, largest in largest specimens (usually in ogcocephalids the relationship is
an inverse one); a deep notch at either side of base of rostrum, which is the channel of the supraorbital
lateral line series. Esca and nasal capsules as in generic description.

Teeth as in generic description. No palatal teeth. Gill rakers as in generic description; usually 6
gill rakers on each side of second arch.

Tubercles widely spaced, varied in size but spines always relatively stout and long, bifid on edge
of disk, recurved on tail. Skin covering eyeballs with small widely-spaced tubercles. Largest tubercles
occur on face, edges of disk, and dorsal and lateral sides of tail. Elsewhere tubercles uneven in size,
not closely spaced so considerable bare skin in intervals, sprinkled over dorsal surface of disk and
pectoral fin pedicels. Subopercular spines extremely large with 4-5 spinelets. On ventral surface of
disk, small widely-spaced tubercles present in small specimens but becoming inconspicuous with
increase in body size, sometimes absent in large specimens; the skin thin and flaccid.

Ventral surface of tail (Fig. 9b) with principal tubercles large, close-set, terminating in long
recurved spines. Anterior to anal fin base, between the 2 rows of principal tubercles, lie 2 rows of
slightly smaller tubercles, also close-set; no small tubercles in intervening spaces or in skin covering
bases of large tubercles. Fins devoid of tubercles, except a few sometimes present at base of pectoral.

Dermal cirri relatively sparse, present around margins of disk and lateral sides of tail. Cirri
associated with neuromasts formed as fringed flaps.

Pectoral fins slender. Pelvic fins with thickened skin on rays.

BRADBURY: DIBRANCHUS REVIEW 281

Color. Garman [1899:91, 94-96] stated, “Color a rich dark chestnut to chocolate brown; fins
black.” For D. asper, (here placed in synonymy of D. hystrix) Garman said, “Blackish externally and
on the linings of the body cavity.” For D. scaber (here placed in the synoymy of D. hystrix) Garman
said, “Greyish black, apparently reddish or purplish in life, fins blackish; lower surface darker.” The
above specimens now faded. All specimens I have examined lack pigment externally, but several
have linings of buccal and gill cavities and the peritoneum covered with melanophores, giving a
medium brown effect.

DISTRIBUTION. — A disjunct distribution, with 11 localities recorded in the eastern Pacific and
one from the South China Sea. Occurs in eastern Pacific from mouth of Gulf of California to Gulf of
Guayaquil and west nearly to Galapagos Is. There are in addition two problematic lots, CAS-SU
46656 from about 150 mi ESE of Cocos I. and CAS-SU 57662 from the Galapagos archipelago. Hard
to identify because specimens are small and must have once dried out, they most likely represent D.
hystrix, and therefore provisionally are included here. Bathymetric range: 9142323 m.

MATERIAL. — 22 specimens 40.0—-125.0 mm SL. LECTOTYPE: MCZ 28726 (83.0 mm SL),
ALBATROSS sta. 3375, 2°34'N, 82°29'W, 2196 m.

CAS-SU 46287 (5, 41.0-75.0 mm SL) and CAS-SU 46657 (125.0 mm SL), ARCTURUS
74-D-1, 4°50’'N, 87°00’W, 1543 m. CAS 82242 (4, 41.1-83.1 mm SL), ANTON BRUUN Cr. 18B,
sta. 766, 4°10’S, 81°27'W, 1815—1860 m. CAS 82256 (43.3 mm SL), NAGA exp. 60-219, sta. 60-67,
South China Sea, 1234-1264 m,m.V. STRANGER. LACM 33588-5 (64.0 mm SL), VELERO
18932, Costa Rica, 14 mi off Punta Guiones. MCZ 28723 (40.0 mm SL, holotype of Dibranchus
asper Garman), ALBATROSS sta. 3418, 16°33’N, 99°52'30"W, 1207 m. MCZ 28724 (101.0 mm
SL, lectotype of Dibranchus scaber Garman), ALBATROSS Sta. 3431, 23°59’N, 108°40'W, 1820
m; MCZ 28725 (58.0 mm SL, lectoparatype of Dibranchus scaber Garman), ALBATROSS sta. 3364,
5°30/N, 86°08'30"W, 1650 m. MCZ 28727 (63.5 mm SL), ALBATROSS sta. 3392, 7°05’30"N,
79°40'W, 2323 m. MCZ 28728 (56.0 mm SL), ALBATROSS sta. 3362, 5°56’N, 85°10'30"W, 2149
m. USNM 135364 (2, 53.2-80.0 mm SL), 0°24'S, 89°06'W, 1485 m. ZMUC (2, 40.0-53.0 mm SL),
GALATHEA sta. 739, 7°22'N, 79°32'W, 938 m.

Two lots provisionally identified as D. hystrix: CAS-SU 46656 (1), ARCTURUS 61-T-4, 4°56'N,
84°35'W, 914 m, and CAS-SU 57662 (2), ARCTURUS 84-T-20, 0°17'S, 91°34'W, 914 m.

Dibranchus spongiosus (Gilbert), 1891
Figs: 3e,d: l0f; 12b

Halieutaea spongiosa Gilbert, 1891:124 (original description, holotype from eastern tropical Pacific, 841 m).
Dibranchopsis spongiosa, Garman, 1899:96 (description; illustrations; eastern tropical Pacific).
Dibranchus spongiosus, Bradbury, 1967:414 (diagnosis of Dibranchus).

DIAGNOSIS. — Body flabby, skin thin and translucent. Subopercular lateral-line count usually 5
(compared to 6 for most eastern Pacific Dibranchus, Table 4). Skull relatively long, 30 percent or
more of SL (Table 5). Lower lip with membranous fringe of fingerlike papillae. No palatal teeth. Gill
rakers in the form of broad tooth plates attached directly to gill arch; no pedicels evident (Fig. 3c, d).

DESCRIPTION. — Based on 30 specimens 43.6—-118.5 mm SL. Counts and body proportions
given in Tables 3—5. Body and skeleton flabby, bones easily bent, tubercles weakly ossified. Cranium
relatively high posteriorly, sloping sharply down to rostrum; rostrum very abbreviated, merely a
rounded, narrow shelf. Illicial cavity very shallow. Esca and nasal capsules as in generic description.
Fringe of fingerlike papillae on membrane hanging from lower lip; oral valves covered with small
short papillae.

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FiGurE 11. Dibranchus tremendus new species, holotype, 192.0 mm SL, UF 25923, from the Caribbean Ocean. (a) dorsal
view, (b) ventral view.

Teeth as in generic description. No palatal teeth. Gill rakers evidently without pedicels; low
dome-shaped tooth plates bearing tiny teeth extend broadly onto gill arch (Fig. 3c), 6 or 7 on each
side of second arch.

Tubercles on dorsal surface of body moderate and uniform in size with short spines, widely
spaced but evenly distributed. Tubercles on edges of disk and lateral sides of tail slightly larger,
sometimes bifid or trifid, but spines very short. Subopercular spines developed in small specimens,
in large specimens relatively very small and encroached upon by skin. Small tubercles evenly
distributed in skin covering eyeballs. Tubercles on ventral surface of disk smaller, widely but evenly
spaced except sparse on lateral sides disk.

Ventral surface of tail (Fig. 10f) with principal tubercles broad, low, terminating in very short
spines. Intervening space with small tubercles both anterior and posterior to anal fin base, these spaced
apart leaving bare skin between.

Pedicels of paired fins covered with tubercles but fins free of tubercles except for a few near
bases of pectoral and caudal fin rays.

Dermal cirri in the form of fringed flaps associated with neuromasts.

Fins moderate length, slender; membranes thin, transparent.

BRADBURY: DIBRANCHUS REVIEW 283

FiGuRE 12. Dorsal views of (a) Dibranchus hystrix, 125.0 mm SL, CAS-SU 46657, and (b) D. spongiosus, 133.2 mm SL,
CAS 82252, both from the eastern tropical Pacific Ocean.

Color. In original description, Gilbert (1891:125) wrote, “Uniform dusky, the tail sometimes
lighter; fins blackish, more or less edged with white.” In a black-and-white photograph of a fresh
specimen kindly sent by staff of Scripps Institution of Oceanography, the skin is translucent with dark
blotches over the top of the eyeballs, the posterior two-thirds of the disk, and on the tail; each tubercle
is darkly pigmented, and these contrast with the pale portions of skin around the face and tail.
Preserved specimens uniformly pale, including linings of gill cavities; peritoneum spotted with black.
One relatively fresh specimen with epithelium of gill bars and branchial cavities very dark, peritoneum
very dark.

DISTRIBUTION. — Recorded here from 10 localities off Mexico, from Bahia Magdalena to Gulf
of Tehuantepec, and | locality off Revillagigedo I. Bathymetric range: 700-1244 m.

MATERIAL. — 92 specimens, 39.2—133.2 mm SL. HOLOTYPE: USNM 44284 (89.5 mm SL).
PARATYPES: USNM 44392 (14, 49.0-75.5mm SL,), ALBATROSS sta. 2992, 18°17'30"N,
114°43'15"W, 841 m.

CAS 57834 (2, 76.1-78.1 mm SL), TE VEGA Cr. 19, Sta. 19, 17°24'N, 101°31"W, 940-1000
m. CAS. 82252 (St, 32.9=133:2imm’ SL); TE VEGA Cr. 19) sta. 17, 19°43 S¢N, 105°335.5W, 700—
900 m. MCZ 28720 (9, 51.0-118.5 mm SL), ALBATROSS sta. 3417, 16°32'N, 99°48’W, 902 m.
MCZ 28721 (2, 46.0-66.5 mm SL), ALBATROSS sta. 3425, 21°19’N, 106°24’W, 1244 m. MCZ
28722 (43.6 mm SL), ALBATROSS sta. 3418, 16°33'N, 99°52'30"W, 1207 m. SIO 58-419 (2,

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82.0-96.5 mm SL), Mexico, Salina Cruz, Gulf of Tehuantepec. SIO 59-266 (1 specimen), 23°55'N,
108°11,5'23°51'N, 108°07.5'W. SIO 64-14 (3, 48.8-67.1 mm SL), 24°59'N, 113°00.5’W—
24°49 .3'N, 113°14.2’'W. UAZ (6, 58.6-72.0 mm SL), Lugar 50 m SW Baluarte, Sinaloa, 800 m.

Dibranchus cracens Bradbury, McCosker, and Long, 1999
Figs. 9e, 13

DIAGNOSIS. — Large tubercles decorated with rows of bosslike spinules radiating out from the
central spine along the ridges between facets (only D. atlanticus has a similar feature). Jaw relatively
long compared to D. atlanticus, 14.6 percent of SL (Table 5). No palatal teeth. Illicial skeleton as in
Dibranchus as shown in cleared and stained illicium from CAS 88034.

DESCRIPTION. — Based on 3 specimens, all males, 102.0-128.3 mm SL. Counts and body
proportions given in Tables 3—5. Skeleton relatively firm, skin tough. Rostrum bluntly triangular,
short, scarcely overhanging mouth; when viewed from above, appears notched on either side near
base owing to excavations for supraorbital lateral-line. Esca with ventral lobes well separated at
midline. All specimens, which are males, with large swollen nasal capsules and the posterior nostril
a long vertical gaping slit.

Teeth as in generic description. No palatal teeth. Gill rakers as in generic description, 6 on
posterior side of second arch.

Dorsal surface of body sprinkled with large tubercles spaced apart, except those of dorsal surface
of tail in 2 longitudinal rows. Intervening spaces like shagreen, packed with tiny simple tubercles.
Large tubercles with short stout center spines and rows of bosslike spinules radiating out from center
spine. Subopercular spines moderate size. Ventral surface of disk and pectoral pedicels covered with
small tubercles.

Ventral surface of tail (Fig. 9e) with principal tubercles broad, low, their central spines short;
prominent spinules radiate from central spines. An orderly row of similar tubercles on each side of
midline anterior to anal base, with small tubercles packed in intervening spaces.

Skin covering eyeballs thickly covered with fine tubercles. Fine tubercles also present on fin rays
of dorsal and caudal fins and dorsal side of pectoral fins, running out for about half the length of the
fin ray; ventral surface of pelvic fin rays have tubercles near fin base.

Dermal cirri present but not abundant; those associated with neuromasts formed as flaps.

Paired fins relatively slender, fin membranes transparent.

Color. In life, holotype with dorsal surface of body salmon colored, tubercles white; dorsal,
caudal and pectoral fins salmon to orange (Bradbury et al. 1999). Faint dark saddle across tail at base
of dorsal fin. Ventral surface of body white, pelvic and anal fins pink. Lower jaw and tip of snout
white, esca red-orange, eyes blue. Paratype, CAS 88034, with dorsal surface brown, tubercles white;
orange around ventral body margin, tail, and fins; dorsal fin brown basally, orange distally; esca
brown, mouth and anus white. In preservative, one specimen (CAS 82222) faded white, but others
light brown with tubercles pale so they seem like pale spots. Olfactory capsule dark or pale. Dark
saddle across tail at base of dorsal fin. Dorsal dark basally, pale distally. Caudal dusky on dorsal half,
otherwise pale. Paired fins and anal fin pale. Ventral surface of body pale. Internal membranes
pigmentless except peritoneum, which is spotted with brown melanophores.

DISTRIBUTION. — All three specimens come from the Galapagos Islands.

ETYMOLOGY. — The name Dibranchus cracens means a neat or tidy Dibranchus, in reference
to the firm, trim body and regularity of size and distribution of tubercles.

MATERIAL. — 3 specimens 102.0-128.3 mm SL. HOLOTYPE: CAS 86516 (119.1 mm SL, male),
JSL 3947, off Cabo San Rosa, Isla Isabella, 01°04.7’S, 91°11.9'W, 152-480 m, 10 Nov. 1995, J. E.
McCosker and party. PARATYPES: CAS 82222 (102.0 mm SL, male), between Islas Santa Cruz and
Floreana, 350 m, dredge, collected by A. deRoy and J. deRoy, 1978. CAS 88034 (128.3 mm SL,

BRADBURY: D/JBRANCHUS REVIEW 285

view, (b) ventral view, (c) frontal view.

male), JSL 3957, off Cabo Douglas, Isla Fernandina, 00°17.5'S, 91°38.9'W, 354 m, 16 Nov. 1995,
J. E. McCosker and party.

Dibranchus velutinus new species
Figs. 10e, 14

DIAGNOSIS. — Entire body covered with extremely fine tubercles, giving skin a velvety appear-
ance. No palatal teeth. Jaw teeth fine, bristlelike, buried among papillae that cover the jaws. Mouth
very wide, 27.5 percent of SL (Table 5).

DESCRIPTION. — Based on a single specimen 99.0 mm SL. Counts and body proportions given
in Tables 3—S. Skeleton flabby (but specimen shown by radiographs to have been decalcified, probably
owing to having been held too long in formalin, so it is possible the skeleton is normally sturdier).

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FiGure 14. Dibranchus velutinus new species, holotype, 99.0 mm SL, CAS 82223, from the Peru-Chile Trench. (a) dorsal
view, (b) ventral view.

Anterior margin of disk blunt, so there is virtually no rostrum, the spines small but sharply upturned.
Esca as in generic description. Specimen a female with small nasal capsules and nostrils.

Pads of papillae prominent on jaws and within oral cavity including oral valves. On jaws and
each ceratobranchial V, slender, translucent teeth lie concealed among papillae, the papillae-teeth
pads of the jaws occurring as bands; each ceratobranchial V with suboval-shaped pad, the two
contiguous along midline their entire lengths. Papillae present on palate, but teeth are not.

Gill rakers finger-shaped, covered with relatively tiny papillae but no tooth plates evident; 5 gill
rakers on each side of second arch.

Skin soft, somewhat puffy, covered everywhere (including skin covering eyeballs) with ex-
tremely fine tubercles giving skin velvety appearance. Large tubercles occur on lateral edges of disk,
dorsal surface of trunk and dorsal and lateral surfaces of tail, and sprinkled over dorsal surface of disk
and pectoral pedicles, but always overlain by fine tubercles so velvety surface uniform throughout.
Large tubercles terminate in modest recurved spines, these bifid or trifid along disk margin.
Subopercular spines moderate size.

Ventral surface of tail (Fig. 10e) uniformly thickly covered with extremely fine tubercles,
tubercles of principal rows scarcely differentiated from the others except for slightly thicker spines.

Extremely fine tubercles in skin thickly covering paired fin pedicels, running out on the fin rays
for about half their length; fine tubercles also occur on fin rays of caudal.

Dermal cirri moderately developed, those associated with neuromasts flaplike.

Paired fins slender, weak; skin of fin rays slightly puffy but fin membranes thin, translucent.

Color. Body and fins tan color, somewhat darker on snout and anterior end of disk, contrasting
with pale lips and esca. Spines of tubercles on face and chin tipped with black, giving effect of skin
being peppered with fine black spots. Lining of gill cavities pale, peritoneum brown.

ETYMOLOGY. — The name velutinus is an adjective meaning velvety, in reference to the fine
tubercles that give the skin the look of velvet.

BRADBURY: DIBRANCHUS REVIEW 287

MATERIAL. — HOLOTYPE: CAS 82223 (99.0 mm SL), IMARPE, 4°59’S, 81°24.5'W. Depth
unknown, but D. sparsus taken in the same haul suggests D. velutinus will be found at depths at which
D. sparsus occurs, about 450-525 m.

Dibranchus sparsus (Garman), 1899
Figs. 10a, 15a

Malthopsis sparsa Garman, 1899:101 (original description, 19 specimens from eastern tropical Pacific):
Bradbury, 1962:3 (lectotype selected, MCZ 28717, from Gulf of Panama, 443 m).
Dibranchus sparsus, Bradbury, 1967:414 (diagnosis of Dibranchus).

DIAGNOSIS. — Teeth present on vomer and palatines. Subopercular lateral-line count usually 6,
preopercular lateral-line count usually 2 (Table 4). Tubercles simple, very small, thickly distributed
over entire body. Subopercular spines small, inconspicuous. A small species reaching just over 80
mm SL.

DESCRIPTION. — Based on 23 specimens 21.9-83.7 mm SL. Counts and body proportions given
in Tables 3—5. Skeleton relatively firm. Rostrum not developed, anterior end of disk rounded. Illicial
cavity very shallow. Esca and nasal capsules as in generic description.

Teeth as in generic description. Teeth present on vomer and palatines. Gill rakers as in generic
description; second gill arch with 6 or 7 gill rakers on each side.

Tubercles small with very short spines uniformly distributed over dorsal and ventral surfaces of
body including skin covering eyeballs. Larger tubercles in several longitudinal series dorsally on trunk
and tail, around edges of disk, and lateral sides of disk, their spines stronger but still short, some
multifid, particularly of edge of disk. Subopercular spines inconspicuous, short, 5—6 spinelets.

Ventral surface of tail (Fig. 10a) covered by the same small, close-packed, slender-spined
tubercles as the rest of body; principal tubercles scarcely larger than intervening ones, but some with
short blunt spines.

Pedicles of paired fins covered with small tubercles; paired fins and caudal with tubercles running
out along fin supports for 1/2 to 2/3 their length. Dorsal and anal sometimes with a few prickles,
especially near bases.

Dermal cirri well-developed on lateral margins of disk and tail; cirri associated with neuromasts
take the form of fringed flaps.

Fins moderate length, relatively stout.

Color. Garman (1899:102) stated: “Fresh specimens brownish, more or less gray, with rather
faint and ill defined spots of light brown somewhat closely placed over the disk; or in cases with white
vermiculations in the brown, or in others with whitish spots over a brownish surface; ventral surface
uniform light; orbits darker.” In one relatively fresh specimen from Peru-Chile Trench: a discernable
reticulum on a relatively darker background; cirri blackish, associated with the lateralis channels on
face and ventral surface of disk. Gill, buccal, and gut cavities with brownish linings; individual
melanophores present, giving wash effect. Some specimens have peritoneum darker brown. Fins
colorless.

DISTRIBUTION. — Recorded here from 6 localities in the eastern Pacific, 3 offthe Gulf of Panama
and 3 off the coast of Peru. Bathymetric range: 443-528 m.

MATERIAL. — 23 specimens 21.9-83.7 mm SL. LECTOTYPE: MCZ 28717 (71.5 mm SL) and
lectoparatypes MCZ 41595 (7, 53.0-70.5 mm SL), ALBATROSS sta. 3386, 7°33'12"N, 79°17'15"W,
443 m.

CAS 82227 (74.8 mm SL), IMARPE, 4°59’S, 81°24.5'W, 528 m, 2 Jul 1980. CAS 82240 (46.1
mm SL), IMARPE, 8°42’S, 78°49’W, 24 Nov 1970. CAS 82236 (2, 29.1-48.7 mm SL), IMARPE,
Bonco de Moncora, Dec 1970. MCZ 28715 (6, 52.4-68.7 mm SL), ALBATROSS sta. 3385, 7°32'N,

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FIGURE 15. Dorsal views of 4 species of Dibranchus with palatal teeth, all from the eastern tropical Pacific Ocean (the 5th
species with palatal teeth, D. discors, is pictured in Fig. 17). (a) D. sparsus, 74.8 mm SL, CAS 82227, showing the reticulated
color pattern apparently unique for Pacific species; (b) D. erinaceus, 110.0 mm SL, SIO 90-71; (c) D. spinosus, 87.9 mm SL,
CAS 82238; (d) D. nudivomer, 98.1 mm SL, CAS 82261. D. spinosus and D. nudivomer sometimes co-occur and, in sorting,
are easily confused, but absence of teeth from the vomer in D. nudivomer separates the two.

BRADBURY: DIBRANCHUS REVIEW 289

78°36'30"W, 474m. MCZ 28716 (4, 67.5-83.7 mm SL), ALBATROSS sta. 3385, 7°32'36"N,
TING We 523 ml.

Dibranchus spinosus (Garman), 1899
Figs. 10b, 15c

Malthopsis spinosa Garman, 1899:104 (original description, 2 specimens from Gulf of Panama, illustrations);
Bradbury, 1962:4 (lectotype selected, MCZ 28710, Gulf of Panama, 2323 m).

Dibranchus spinosus, Bradbury, 1967:414 (diagnosis of Dibranchus).

Malthopsis spinulosa Garman, 1899:106 (original description, 21 specimens from Gulf of Panama); Bradbury,
1962:4 (lectotype selected, MCZ 28709, 935 m; synonymized with D. spinosus).

Dibranchus alberti Nunez, 1967:43 (original description, 5 specimens from off Arica, Chile, 366 m; photo-
graphs; no museum or museum numbers given).

DIAGNOSIS. — Teeth on vomer and palatines. Subopercular lateral-line count usually 6, preop-
ercular lateral-line count usually 3 (Table 4). Tubercles simple, very small, thickly covering entire
body. Subopercular spines small, inconspicuous.

DESCRIPTION. — Based on 58 specimens 32.9-149.0 mm SL. Counts and body proportions
given in Tables 3—5. Skeleton somewhat rubbery, subopercles (lateral margins of disk) flabby, often
rolled inward in preserved specimens. Rostrum variable when viewed from above, from bluntly
rounded to triangular but always very short, appearing notched at the base owing to excavation of
supraorbital lateral line channel on either side. Similarly prominent notches formed by lateralis system
channels on margins of disk opposite posterior end of orbits and near base of subopercular spine. Esca
and nasal capsules as in generic description.

Teeth as in generic description. Pads of teeth on palatines, these contiguous with pad of teeth on
vomer. Gill rakers as in generic description, usually 6 rakers on each side of second arch.

Tubercles very small and slender with relatively long weak spines, about uniform in size in
smaller examples, but, in larger specimens, many tubercles on dorsal surface of body slightly enlarged,
these sprinkled thinly but evenly over dorsal surface of disk, denser along margins of disk; in either
case, tubercles thickly distributed over dorsal surface of body including skin covering eyeballs;
tubercles of edge of disk and lateral sides of tail with longer, stouter spines. Tubercles of edge of disk
and lateral sides of tail with slightly stouter longer spines. Subopercular spines large and prominent
in smallest specimens, but the larger the specimen, the smaller the spine, until it can hardly be noticed;
5—6 spinelets. Fine tubercles cover entire ventral surface of body.

Ventral surface of tail (Fig. 10b) covered with very small densely-packed fine-spined tubercles.
Principal tubercles slightly larger than intervening ones, the spines long and delicate.

Paired fins with tubercles covering pedicels and occurring along both dorsal and ventral surfaces
of fin rays. Caudal also with tubercles running out along fin rays, dorsal with a few prickles at base,
anal sometimes with prickles but usually none.

Dermal cirri numerous, conspicuous because of black color, on chin and lateral margins of disk
and tail. Cirri associated with neuromasts take the form of fringed flaps, also black.

Fins slender, relatively weak.

Color. Garman (1899:106, 107) wrote of D. spinosus: “Uniform dark brown; fins black” and of
D. spinulosa (here placed in synonymy of D. spinosus): “Translucent whitish to brown, with or
without cloudings or spots . . . . Traces of brilliant red colors appear on light colored individuals and
... some of the lateral lobes of the illicium are deep red, while the median lobe is of cream color; the
illicium varies from light color to dark brown.” Newer material has some specimens faded to
extremely pale, otherwise tan to brown, spines of tubercles appearing as points of pigment darker
than background. Nasal capsule blotched to entirely dark, esca variously pale or blotched, a tinge of
red-orange visible in some. Sometimes membranes around gill pores and/or anus-urogenital region

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relatively dark, contrasting with rest of body. Cirri associated with lateralis channels on face on ventral
surface of disk usually conspicuoulsy darker than body color. Fins about same color as body, often
darker distally; dorsal and caudal sometimes barred or blotched with darker color. In freshest
specimens, skin covering gill bars dark brown with gill rakers white, contrasting strikingly. Perito-
neum light to medium brown.

DISTRIBUTION. — Recorded here from numerous localities off Mexico and Central and South
America in the eastern Pacific, from the mouth of the Gulf of California to off the southern tip of
Peru. Bathymetric range: the majority from 528-1439 m, but 3 lots from 1815—2323 m, one of these
the lectotype. One lot with little data recorded as being from 100 m. One literature record from off
Arica, Chile, 19°08’09"S, 70°20'02"W, 366 m (Nunez, 1967).

MATERIAL. — 140 specimens 32.9—149.0 mm SL. LECTOTYPE: MCZ 28710 (129.5 mm SL),
ALBATROSS sta. 3392, 7°05'30"N, 79°40'W, 2323 m.

CAS-SU 25242 (14), ALBATROSS sta 2792, 0°37'S, 81°W, 733 m. CAS 39919 (2, 138.8-149.0
mm SL), ANTON BRUUN Cr. 18B, sta. 766, 4°10'S, 81°27’W, 1815-1860 m. CAS 39920 (17,
37.2-81.2 mm SL), ANTON BRUUN Cr. 18B, sta. 754, 7°49’S, 80°38’W, 605-735 m. CAS 39921
(16, 49.4-97.1 mm SL), ANTON BRUUN Cr. 18B, sta. 770, 3°15’S, 80°55'W, 945-960 m. CAS
42708 (46.2 mm SL), ANTON BRUUN Cr. 18B, sta. 755, 7°43’S, 80°43'W, 700-1110 m. CAS-SU
46656 (1 specimen, poor condition), ARCTURUS 61-T-4, 4°56'N, 84°35'W, 914 m. CAS 57853
(94.0 mm SL), TE VEGA Cr. 19, Sta. 144, 2°10’S, 81°13'W, 800-1000 m. CAS 57862 (3, 35.2—102.5
mm SL), TE VEGA Cr. 19, Sta. 148, 2°25’S, 81°10’W, 700-1000 m. CAS 82228 (109.2 mm SL),
IMARPE, 18°17.7’S, 71°11.3’W, 600 m. CAS 82229 (135.2 mm SL), IMARPE, 18°10’S, 71°29" W,
610 m. CAS 82230 (94.6 mm SL), IMARPE, 16°29’S, 73°33’'W, 1300 m. CAS 82231 (96.2 mm SL),
IMARPE, 16°30.5’S, 73°27.5'W. CAS 82232 (115.5 mm SL), IMARPE, 10°51.8’S, 78°30.7 W, 800
m. CAS 82233 (2, 117.8-122.8 mm SL), IMARPE, 18°19'S, 71°12'W, 810m. CAS 82234 (2,
68.2—-106.9 mm SL), IMARPE, 7°46’S, 80°31'W, 800 m. CAS 82235 (139.2 mm SL), IMARPE,
17°41'S, 71°42'W, 650 m. CAS 82237 (101.8 mm SL), IMARPE, 100 m, no other data. CAS 82238
(90.5 mm SL), IMARPE, 16°30.9'S, 73°26’W, 800 m. CAS 82239 (135.2 mm SL), IMARPE, 4°59’S,
81°24.5'W, 528 m. CAS 82241 (96.7 mm SL), IMARPE, 3°48.2’S, 81°22'W, 695 m. CAS 82260
(36, 34.5-96.9), ANTON BRUUN Cr. 18B, sta. 754, 7°49'S, 80°38’W, 605-735 m. GCRL (3,
83.3-109.0 mm SL), CANOPUS sta. 1291, 7°13'N, 79°18'W, 805-841 m. LACM 10098 (32.9 mm
SL), ELTANIN sta. 34, 7°47.5’S, 81°23'W, 677 m. LACM 31124-6 (1 specimen, damaged), VEL-
ERO 13770, 21°52'30"N, 106°47'36"W, Middle American Trench. LACM 33699-2 (2, 103.0-129.7
mm SL), VELERO 19128, off Cabo Matapalo, Costa Rica, 22 Jun 1973. MCZ 28709 (101.0 mm SL,
lectotype of Malthopsis spinulosa Garman) and MCZ 41600 (19, 87.0-97.5 mm SL), ALBATROSS
sta. 3394, 7°21'N, 79°35'W, 935 m. MCZ 28714 (4, 55.0-77.0 mm SL), ALBATROSS sta. 3418,
16°33'N, 99°52'30"W, 1207 m. MCZ 41597 (62.5 mm SL), ALBATROSS sta. 3354, 7°09’45"N,
80°50'W, 589 m. MCZ 41599 (130.5 mm SL), ALBATROSS sta. 3393, 7°15'N,79°36'W, 1865 m.
SIO 59-265 (4 specimens), 23°40.5'N, 107°38.5—23°37.0'N, 107°51.8’W, 1366-1439 m. USNM
135365 (131.0 mm SL), ALBATROSS sta. 3418, 16°33'N, 99°52’30"W, 1207 m. ZMUC (13;
52.5—134.0 mm SL), GALATHEA sta. 739, 7°22'N, 79°32’W, 938 m.

Dibranchus erinaceus (Garman), 1899
Figs. 10c, 15b

Malthopsis erinacea Garman, 1899:103 (original description, 8 specimens from eastern tropical Pacific);
Bradbury, 1962:3 (lectotype selected, MCZ 28712, from Galapagos Is., 770 m).

BRADBURY: DIBRANCHUS REVIEW 291

Dibranchus erinaceus, Bradbury, 1967:414 (diagnosis of Dibranchus); Bradbury, et al. 1999 (6 specimens from
Galapagos Is.).

DIAGNOSIS. — Teeth on vomer and palatines. Subopercular lateral-line count usually 6, preop-
ercular lateral-line count usually 3 (Table 4). Tubercles distributed over entire body, consisting of
moderate sized tubercles interspersed with very small tubercles (Fig. 10c and 15b). Subopercular
spines long.

DESCRIPTION. — Based on 18 specimens 37.0—-110.0 mm SL. Counts and body proportions
given in Tables 3—S. Skeleton relatively rigid. Rostrum prominent, triangular in shape when viewed
from above, overhanging mouth in small specimens, becoming relatively shorter in large specimens;
when viewed from above, a conspicuous notch present on either side at the base of the rostrum where
the supraorbital lateralis channel on each side passes from the face to the roof of the cranium. Esca
and nasal capsules as in generic description.

Teeth as in generic description. Teeth present on vomer and palatines. Gill rakers as in generic
description; usually 6 gill rakers on each side of second arch.

Tubercles on dorsal surface of disk moderate size with relatively short stout spines, fairly uniform
size except as noted below, evenly distributed. Larger tubercles occur in several longitudinal series
along dorsal side of trunk and tail; large tubercles also along edges of disk and sides of tail, these
often with extra spinules that are much smaller than terminal spines. Subopercular spines relatively
long, 6-8 spinelets. Tiny tubercles evenly distributed in skin covering eyeballs. Tubercles of ventral
surface of disk much smaller than those of dorsal surface, closely spaced, evenly distributed.

Principal tubercles of ventral surface of tail (Fig. 10c) moderate size with short spines. In the
intervening space just posterior to the anus is a short series of similar moderate-sized tubercles;
remainder of the intervening space filled by small tubercles both anterior and posterior to anal fin
base.

Paired fins and caudal with sparse tubercles running out on fin rays a short distance.

Dermal cirri usually present along disk margins and sides of tail; not abundant. Cirri in form of
fringed flaps associated with neuromasts.

Fins moderate size, pectorals slender, pelvic rays with thickened skin.

Color. Garman (1899:104) stated, “Brownish, with traces indicating a rose color in life; young
individuals blackish: fins blackish.” These specimens and most others at my disposal now faded pale.
One specimen, a male, (SIO 90-71) has fins, nasal capsules, and esca blotched with dark pigment;
cirri associated with lateralis channels on face and ventral surface of disk also blackish; lining of gill
cavities dark brown, epithelium of gill bars also dark brown, constrasting sharply with white gill
rakers; peritoneum blotched with dark spots.

DISTRIBUTION. — Recorded here from only 5 localities in the eastern Pacific, from the vicinities
of Galapagos Is., Cocos I., and Peninsula de Azuero, Panama. Bathymetric range: 700-1150 m.

MATERIAL. — 19 specimens 37.0—-110.0 mm SL. LECTOTYPE: MCZ 28712 (114.0 mm SL) and
lectoparatype MCZ 41598 (81.0 mm SL), ALBATROSS sta. 3402, 0°57'30"S, 89°03'30"W, 770 m.

CAS 60476 (8, 37.0-97.0 mm SL), ARCTURUS 74 OT-4, 60 mi. S of Cocos Id., 4°50'N,
87°00'W, 1143 m. CAS 86503 (3, 83.8—-105.2 mm SL), JSL 3977, James Bay, Isla San Salvadore,
914 m, J. E. McCosker and party. CAS 86529 (2, 98.0-105.8 mm SL), JSL 3949, off Cabo Rosa, Isla
Isabella, 744 m, J. E. McCosker and party. CAS 86533 (1, 105.5 mm SL), JSL 3976, James Bay, Isla
San Salvadore, 914 m, J. E. McCosker and party. MCZ 28711 (102.0 mm SL), ALBATROSS sta.
3358, 6°30'N, 81°44'W, 1015 m. SIO 90-71 (110.0 mm SL), Galapagos Is., 1°37.73’S, 90°10.7'W,
965-310 m. USNM 135579 (90.0 mm SL), ALBATROSS, 0°29'S, 89°54'30"W, 715 m.

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Dibranchus nudivomer (Garman), 1899
Figs. Sc, d; 10d, 15d

Dibranchichthys nudivomer Garman, 1899:99 (original description, 8 specimens from eastern tropical Pacific,
1271-1335 m); Bradbury, 1962:2 (lectotype selected, MCZ 28719, eastern tropical Pacific, 1271 m).
Dibranchus nudivomer, Bradbury, 1967:414 (diagnosis of Dibranchus); Castro-Aguirre and Moncayo-Lopez,

1976:307 (41 specimens from off Sinaloa, México; photograph, osteology, figures).

DIAGNOSIS. — Palatal dentition unique for family: teeth on palatines but none on vomer.
Subopercular lateral-line count usually 6, preopercular lateral-line count usually 2 (Table 4).

DESCRIPTION. — Based on 50 specimens 61.0—-102.5 mm SL. Counts and body proportions
given in Tables 3—5. Skeleton somewhat rubbery. Rostrum very short. Skin thin, semi-transparent,
soft. Esca and nasal capsules as in generic description.

Teeth as in generic description. Palatal dentition unique in having pads of teeth on each palatine
bone but none on vomer. Gill rakers as in generic description, relatively short; 6 gill rakers on each
side of second arch.

Tubercles moderately small with short spines, uniform in size, evenly distributed over dorsal
surface of disk and tail. Tubercles on edges of disk slightly larger, their spines sometimes bifid but
short; tubercles of lateral sides of tail also larger, the spines in these stout, relatively long, recurved.
Subopercular spine moderate, 5—6 spinelets. Smaller tubercles evenly distributed over ventral surface
of disk, on pedicels of paired fins, and on skin covering eyeballs.

Ventral surface of tail (Fig. 10d) evenly covered by tubercles, tubercles of principal rows only
slightly larger than intervening ones, their spines retrosely curved.

Caudal with tiny tubercles running out short distance along fin rays, pectorals with only a few
tubercles on dorsal surface at base, other fins devoid of tubercles.

Dermal cirri well developed on disk margins and sides of tail. Cirri associated with neuromasts
formed as fringed flaps.

Fins small, slender.

Color. — Garman (1899) stated that his specimens were brown to black. Material available to
me varied from pale tan to dark tan; in many, cirri associated with the lateralis excavations on face
and disk dark brown. Some specimens with dark pectorals, particularly tips. Linings of gill cavities
pigmented but light colored; peritoneum often medium brown.

DISTRIBUTION. — Present material is from the Gulf of Panama, Gulf of Guayaquil, and from off
Peru. Bathymetric range: 605—1400 m. Reported also from off the Rio Baluarte, Sinaloa, México,
900 m (Castro-Aguirre and Moncayo-Lopes 1976).

MATERIAL. — 102 specimens 36.5—112.0 mm SL. LECTOTYPE: MCZ 28719 (94.0 mm SL), and
lectoparatype MCZ 41595 (84.0 mm SL), ALBATROSS sta. 3353, 7°06'15"N, 80°34'W, 1271 m.

CAS 35339 (3, 78.4-112.0 mm SL), TE VEGA Cr. 19, Sta. 84, 2°20'S, 81°16’W, 850-1400 m.
CAS 82225 (31, 36.5—104.7 mm SL), ANTON BRUUN Cr. 18B, Sta. 770, 3°15’S, 80°55'W, 945-960
m. CAS 82261 (59, 47.0-99.1 mm SL), ANTON BRUUN Cr. 18B, sta. 754, 7°49’S, 80°38'W,
605—735 m. MCZ 28718 (5, 61.0-87.5 mm SL), ALBATROSS sta. 3395, 7°30'36"N, 78°39'W, 1335
m. USNM 57867 (75.0 mm SL), ALBATROSS sta. 3395, 7°30'36"N, 78°39’W, 1335 m.

Dibranchus discors Bradbury, McCosker, and Long, 1999
Figs. 7a, 16

DIAGNOSIS. — Teeth present on palatines and vomer. Interorbital and mouth both exceptionally
narrow (Table 5). No tubercles in skin covering eyeballs. No tubercles on ventral surface of disk
except a few near bases of pelvics. Conspicuous chalk-white oval marking on midline of lower jaw.

BRADBURY: DIBRANCHUS REVIEW 293

FIGURE 16. Dibranchus discors Bradbury, McCosker, and Long.. (a) dorsal view, and (b) ventral view of holotype, 80.2 mm
SL, SIO 90-71; (c) frontal-oblique view of paratype, 87.2 mm SL, CAS 47195. Both from the Galapagos Islands.

DESCRIPTION. — Based on 5 specimens 80.2—107.9 mm SL. Counts and body proportions given
in Tables 3—S. Skeleton relatively firm. Rostrum short, narrow, upturned, about even with jaws, not
overhanging mouth. Esca higher than wide, otherwise not differing from other Dibranchus; ventral
margin with membranous fringe. Illiclum of one specimen (CAS 47195) cleared and stained, agrees
with Dibranchus in lacking a long median dorsal process and in having the lateral processes each
with a foramen (Fig. 7a). Olfactory organs in males somewhat larger than in females, the posterior

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nostril slit-shaped, but this dimorphism less pronounced in D. discors than in other Dibranchus.
Females, as usual, show no swelling of the olfactory organ and have both nostrils small and round.
A unique chalk-white oval patch on midline of chin with surface appearing pitted or sculptured, that
appearance due to the presence lateral line organs and cirri that are all absolutely white but otherwise
normal. The patch does not appear to be a light organ; the skin is tough as elsewhere on the body, not
glandular.

Teeth as in generic description. Teeth present on vomer and palatines. Gill rakers as in generic
description, 5 on each side of second arch.

Tubercles relatively large, widely spaced, each with a short, stout spine, the largest with minute
spinules radiating from the central spine. Tubercles of edge of disk multifid. Prominent tubercles
present in longitudinal rows on dorsal side of trunk and dorsal and lateral sides of tail, and sprinkled
over dorsal surface of disk and pectoral pedicels with intervening skin bare of tubercles. No tubercles
on skin covering eyeballs. Subopercular spines long, thick, covered with rows of minute spinules.
Skin of ventral surface of disk devoid of tubercles except for patch of very small tubercles between
and anterior to pelvics.

Ventral surface of tail with large principal tubercles, their spines moderate, recurved. A few small
tubercles scattered among the bases of the principal tubercles.

Dermal cirri not developed.

Paired fins slender, moderately strong, fin membranes transparent; tips of pectoral rays fleshy.
No tubercles on fins.

Color. In life, dorsal surface light brown with pale tubercles making whitish spots (Bradbury et
al. 1999). Dark margins around anterior nostrils give effect of dark moustache; esca brown, fins pink
except dorsal brown. Brownish saddle over tail beneath dorsal fin. Ventral surface pale except for
chalk-white oval marking on midline of lower jaw which is ringed by black cirri. In preservative,
ALBATROSS specimen (CAS-SU 47195) faded white. Other specimens medium brown dorsally
with pale tubercles making pale spots. Dark margins around anterior nostrils give effect of dark
moustache. A dark saddle on tail beneath dorsal fin, the dark color continuing onto dorsal fin. Ventral
surface pale; pectorals, pelvics, and anal pale. A conspicuous patch on chin colored chalk-white, the
effect intensified by black edging composed of black cirri.

DISTRIBUTION. — All specimens taken in the Galapagos archipelago.

ETYMOLOGY. — The name discors means unlike or different, in reference to the aberrant shape
of the head and illicial bone in this Dibranchus.

MATERIAL. — Five specimens 80.2—107.9 mm SL. HOLOTYPE: CAS 86502 (107.9 mm SL,
female), JSL 3957, 00°17.5'S, 91°38.9'W, Isla Fernandina off Cabo Douglas, 340 m, 16 Nov. 1995,
J. E. McCosker and party. PARATYPES: CAS-SU 47195 (87.2 mm SL, female), ALBATROSS sta.
4642, 1°30'30"S, 89°35'W, 549 m, 7 Nov 1904. CAS 86542 (88.2 mm SL, male), JSL 3974, N shore
of Tower Island, 00°21.6'S, 89°58.3'W, 373 m, J. E. McCosker and party. SIO 90-71 (80.2 mm SL,
female), Galapagos Is., 1°37.73’S, 90°10.7'W, between 310—965 m, 1 Feb 1990. USNM 344535 (84.1
m SL, male), JSL 3957, 00°17.5'S, 91°38.9’W, Isla Fernandina off Cabo Douglas, 415 m, J. E.
McCosker and party.

Dibranchus japonicus Amaoka and Toyoshima, 1981
Fig. 19f

Dibranchus japonicus Amaoka and Toyoshima, 1981:115 (original description, holotype and 7 other specimens
from off Iwate Prefecture, Pacific coast of Japan, 1180-1230 m; photographs); Yamakawa (in Okamura and
Kitajima, ed. 1984:380 (Pacific coast of Japan, 900-1000 m).

BRADBURY: D/BRANCHUS REVIEW 295

DIAGNOSIS. — Skull relatively short, the length 24.2 percent of SL (Table 5) compared to 26
percent or more in other Dibranchus. Skin extremely thick and leathery, studded over entire body
with small tubercles; principal tubercles of ventral surface of tail very stout but relatively short (Fig.
9f). No palatal teeth.

DESCRIPTION. — Based on | male specimen 154.5 mm SL and on original description (Amaoka
and Toyoshima 1981). Counts and body proportions given in Tables 3—S. Skeleton firm. Skin very
thick, fairly tough, with tubercles forming a coarse leathery shagreen. Rostrum short and broad. Esca
wider than high, the ventral lobes in this specimen flared out and distended so that each appears
two-parted, but is essentially as in generic description. Nasal capsule with enlarged lamellae within,
and the posterior nostril a gaping slit.

Teeth as in generic description. No palatal teeth. Gill rakers as in generic description; 5 gill rakers
on each side of second arch.

Small tubercles abundantly scattered in the thick skin, forming a coarse leathery shagreen over
entire body including skin covering eyeballs. Large tubercles with strong short spines occur on dorsal
margins of orbits, along disk margins and lateral sides of tail, and sprinkled irregularly on dorsal
surface of disk. These large tubercles have their bases covered with skin bearing small tubercles.
Subopercular spine thick with 4—S spinelets. Ventral surface of disk covered with the same small
tubercles as seen on dorsal side. No tubercles on fin rays.

Ventral surface of tail with tubercles of principal rows somewhat larger than nearby tubercles
and terminating in strong short spines (Fig. 9f). A few large tubercles occur between the principal
rows, but the noticeable feature is the thick covering of numerous small tubercles between the
principal rows, encroaching on the bases of the large tubercles as described for the dorsal surface of
the body.

Pectoral and pelvic fins both relatively short, broad, the covering skin extremely fleshy, the
membranes thick.

Color. The specimen agrees with those in original description in its very dark coloration,
including fins, and blackish linings of oral, gill, and peritoneal cavities.

DISTRIBUTION. — Known from the Japanese archipelago from 620-1270 m (Amaoka and
Toyoshima 1981) and from off eastern Australia, from off Raine Is. (11°35’S) and off Sidney to
Shoalhaven (33°50'S—34°55’S), (Paxton et al. 1989:284). A disjunct locality reported here from off
Cape Town, South Africa at 855—900 m.

MATERIAL. — IOAN (154.5 mm SL), FIOLENT Cr. 3, trawl 305, 35°26'S, 18°45’E, 855—900
m, 21 Nov 1973.

Dibranchus accinctus new species
Enee, fen 1

Possible reference: Dibranchus obscurus, Norman, 1939:113 (3 specimens from Gulf of Aden and Indian Ocean
off Zanzibar, 1022—1789 m:; illustration).

DIAGNOSIS. — A large Dibranchus (175 mm SL) with tubercles covering ventral surface of disk.
Ventral surface of tail with principal tubercles occupying entire ventral surface, but with numerous
small tubercles in skin thickly covering bases of principal tubercles (Fig. 9c). Cranium relatively wide,
the width 26.7 percent of SL compared to a range of 16.7—25.6 for the genus (Table 5). No palatal
teeth.

DESCRIPTION. — Counts and body proportions given in Tables 3—5. Skeleton firm. Skin tough,
like coarse shagreen on dorsal surface of disk; on ventral side of disk, skin somewhat flabby, especially
on belly. Rostrum short, only slightly overhanging mouth. Esca as in generic description. The single
specimen a female, its nasal capsule small with small round nostrils.

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FIGURE 17. Dibranchus accinctus new species, holotype, 175.0 mm SL, CAS 82221, from the western Indian Ocean. (a)
dorsal view, (b) ventral view.

Teeth as in generic description. No palatal teeth. Gill rakers in the form of low pedicels each
capped by a tooth plate bearing a semi-spherical cluster of teeth. 5 gill rakers on each side of second
arch.

Tubercles on dorsal surface of disk grading in size from small on the face, interorbital, and skin
covering eyeballs, and sides of disk to moderate (and variable) in size posteriorly on the disk. Large
tubercles with long spines occur on dorsal margins of orbits, down the midline posteriorly on the disk
and onto the tail, and all around the disk margin, these last with bifid spines. Subopercular spine longer
than other spines of disk margin, with 4 spinelets. Ventral surface of disk covered with small tubercles.

Ventral surface of tail with tubercles of principle rows large, long-spined and close-set (Fig. 9c).
Small tubercles fairly evenly distributed between and over the bases of the large tubercles both anterior
and posterior to the anal fin base.

Paired fins with small, widely spaced tubercles running out in the thick skin of the rays nearly
2/3 the way to the fin tip. Caudal fin with a few tubercles along rays near their bases.

Dermal cirri abundant on face, ventral edges of disk margin, and lateral sides of tail, readily
visible because darkly pigmented; flaplike cirri associated with neuromasts. On dorsal surface of disk,
darkly-pigmented cirri associated with neuromasts which in this location are not housed in channels.

Paired fins relatively short, broad, fleshy; fin membranes translucent.

BRADBURY: DIBRANCHUS REVIEW 297)

Color. The specimen has ground color uniform tan. Tubercles of face, interorbital space, and to
some extent the pectoral pedicels and sides of disk, have dark pigment around tips of spines, giving
dorsal surface a dotted pattern. Dorsal fin dark, pectorals and caudal darkly blotched. Cirri dark brown
or black.

ETYMOLOGY. — The name accinctus means a well-armed Dibranchus, in reference to its long
tough spines.

MATERIAL. — HOLOTYPE: CAS 82221 (175.0 mm SL), ANTON BRUUN Cr. 6, sta. 399C,
21°16'S, 38°18'E, 1510-1600 m.

Solocisquama new genus

TYPE SPECIES. — Dibranchus stellulatus Gilbert, 1905, is here designated as the type species of
the new genus Solocisquama.

DIAGNOSIS. — Gills 2; holobranchs present on 2nd and 3rd arches only. Upper jaw with a marked
cleft in midline, unique for family. Scales in the form of tubercles, not bucklers, but spines highly
modified (Fig. 18). Tooth patches on fifth ceratobranchial bones relatively small, not meeting, or
barely meeting, in midline, unlike broad tooth patches meeting broadly in midline seen in Dibranchus.
Illicial bone (Fig. 7b) with two well-developed laterodorsal processes somewhat resembling those
found in Halicmetus (Bradbury 1967:401); no foramina in lateral processes as seen in Dibranchus.
Tail lateral line commences posterior to anus; unlike Halieutopsis, no lateral-line organs beside, or
anterior to, anus.

DESCRIPTION. — Disk triangular in outline unless pectoral pedicels collapsed against body,
which gives disk an oval or bell-shaped outline. Subopercular spines short, stout, multifid.

Body depressed but cranium elevated above general surface of disk so that eyes are directed
laterally and anteriorly. Rostrum composed of closely-spaced tubercles forming a short shelf above
esca; one species, S. stellulata, with a prominent median tubercle larger and longer than those to the
sides. Illicial cavity small, not cavernous as in Coelophrys or some Halieutopsis. Esca consisting of
a dorsal semi-leaf-shaped lobe with a median furrow which ventrally separates two semi-spherical
ventral lobes (Figs. 1b, c; 19). Olfactory organs sexually dimorphic in the one species (S. erythrina)
with sufficient material to assess: in females, olfactory organs relatively small and containing a few
small lamellae, the nostrils small and round; in males, olfactory organ large, swollen by numerous
enlarged lamellae within, posterior nostril a wide, gaping slit. Status of sexual dimorphism in olfactory
organ in S. ste/lulata and S. carinata unknown.

Vertebral count 18, dorsal rays 6-7, pectoral rays 13—16 (Table 6), anal rays always 4, caudal
rays always 9. As in Dibranchus, lateral line interrupted just as it descends from the disk onto the tail.
The lateral-line organs on the disk are too difficult to count, so these were not studied. Tail lateral
line commences on the ventral side of tail just posterior to anus, thence continuing along lateral sides
of tail onto base of caudal fin; tail lateral-line count 6—12 (Table 6). Subopercular lateral-line series
usually 5, range 4-5 (Table 6). The dorsolateral branch of the subopercular series shows no variation
in the specimens at hand, the count always 3. Likewise, the preopercular series does not vary, the
count always 2.

Teeth in oral and gill cavities small and recurved as described for Dibranchus. Teeth in bands
on jaws, visible when mouth closed. No palatal teeth. Patches of teeth on ceratobranchial V small,
not meeting, or barely meeting, in midline. Gill rakers in the form of pedicels capped by tooth plates
(Fig. 5a, b); pedicels relatively long and stout compared to Dibranchus. (Except for gross examination,
this feature in S. carinata was not studied.)

Dermal cirri appear to be absent in this genus.

Illicial Bone. Illicial bone known for S. ste//ulata (Fig.7b) and S. erythrina; the two agree in
lacking the median dorsal process seen in the majority of ogcocephalid genera and in having, instead,

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FiGureE 18. Sketches of tubercles from the dorsal surface of the disk in species of Solocisquama. (a) S. stellulata, 88.8 mm
SL, ZMMSU P.20352; (b) S.ervthrina, 135.5 mm SL, BPBM 29270; (c) S. carinata, 73.2 mm SL, ZMMSU P.20359, holotype.
The tubercles in S. ervthrina and S. carinata are very different from one another, and each is unique for the family. Multi-spined
tubercles seen in S. ste/lulata occur in many species just on the edges of the disk, but S. ste//u/ata is alone in having them nearly
all over the body.

two long dorsolateral processes, resembling those in Halicmetus (Bradbury 1967:401). They further
agree in lacking foramina in the lateral processes that articulate with the pterygiophore, unlike
Dibranchus. Since S. carinata is known only from the holotype, clearing and staining was not
performed on that specimen, so the nature of its illicial bone is unknown.
DISTRIBUTION. — From Sala y Gomez Ridge near Easter Island in the eastern Pacific, Hawaiian
archipelago, Philippine Sea, South China Sea, and western Indian Ocean off South Africa.
ETYMOLOGY. — Solocisquama means bristly or coarsely scaled.

KEY TO THE SPECIES OF SOLOCISQUAMA

la. When disk viewed from the front or side, dorsal surface of disk seen to be thickly covered by tubercles with simple

but relatively long spines (Fig. 19), true even for specimens as smallas37 mmSL............. S. stellulata
lbs Dorsalisuntace ohdiskicovered byslows tubercles withOut)lOnoSPIeCSas emi) eee aoe eure eae 2
2a. Tubercles along sides of tail (herein called principle tubercles) each with 2-4 long, irregularly sized, somewhat flar-

Ings spines (PIE 20CKA eS Au hee ver ae cee oey cick ci be ch ating RSmEE oh aerediny, Sata A Ee | chat aut lee?, uk’ Chee S. erythrina
2b. Principle tubercles shaped like toothed scutes (Fig. 20a), aligned in keellike series on each side of tail .. . . S. carinata

Solocisquama stellulata (Gilbert), 1905
Figs. 1b, 18a, 19a, 20b

Dibranchus stellulatus Gilbert, 1905:698 (original description, holotype from Hawaiian Is., 326-369 m);
Bradbury, 1967:414 (diagnosis of Dibranchus),; Mochizuki, in Okamura, Amaoka, and Mitani 1982:361
(Kyushu-Palau Ridge, Japan, 550 m): Bradbury, in Smith and Heemstra, 1986:371 (off Natal, South Africa,
475 m).

DIAGNOSIS. — Tubercles on dorsal surface of disk with simple spines (Fig. 19a), but elsewhere
on body tubercles multi-spined (Fig. 18a), these especially prominent on principal tubercles of tail

BRADBURY: D/JBRANCHUS REVIEW 299

TABLE 6. Frequency distribution for dorsal and pectoral fin ray counts, vertebral counts and subopercular
and tail lateral-line counts in species of Solocisquama.

Lateral-line Count

Dorsal Pectoral Fin Verte- Subopercular Tail
Fin brae
6 ii 13> el4g is. 6 18 = 5 6 7 8 9 OL pF pllicn le
erythrina 3 ] —- 2 5 ] 6 — 8 2 3 l
stellulata 4 — ] 3 = 8 ] 9 ) ~ ] 4 2 - =
carinata l — - — 2 — ] = 2 —-— = = l ] - =

(Fig. 20b). Rostrum with a large multifid median spine protruding beyond its neighbors, terminating
in 5 or more flaring spinelets.

DESCRIPTION. — Based on 7 specimens including holotype, USNM 51595; counts and body
proportions given in Tables 6 and 7. Skeleton firm, skin tough. Rostrum short but with a jaunty,
upturned, thick, median tubercle protruding further than its neighbors with 5 or more spinelets near
tip flaring outward. Upper jaw with median cleft. Esca composed of large central lobe with a median
furrow and two smaller ventral lobes, one on each side (Fig. 1b, 19a). Sexual dimorphism of nasal
capsules problematic; 2 specimens (including holotype) have small round posterior nostrils and also
have ovaries, but other specimens too small or too poorly preserved, or both, to see nature of capsules
and/or gonads.

Teeth in bands on jaws, visible when mouth closed. Small elongate tooth pad on each cerato-
branchial V, set at 90° angle to one another but not meeting at midline. (In the original description,
Gilbert recorded “tongue without teeth,” but this is an error.) No palatal teeth. (Mochizuki [Okamura
etal. 1982:361] stated that palatine teeth were present for 4 specimens from the Kyushi-Palau Ridege,
although no vomerine teeth. I have checked my material in this regard and can find no palatine teeth.)
Gill rakers as in generic description; 6 gill rakers on each side of second arch.

Tubercles with long, strong sharp spines evenly distributed over entire dorsal surface (in contrast
to S. erythrina, with far shorter spines (Fig. 19) and S. carinata with none. On ventral side of disk,
tubercles small, evenly distributed down center of disk but thinning laterally. Except for dorsal surface
of disk, most tubercles are the peculiar multi-spined tubercles (Fig. 18a), occurring on face, edges of
disk, pectoral pedicels, and all over tail, arranged so that small tubercles crowd among large ones.
Tubercles present in skin covering eyeballs. Subopercular spines moderate length but thick with 5—6
spinelets.

Ventral surface of tail covered with multi-spined tubercles, these especially large and prominent
on principal tubercles (Fig. 20b).

Fins without tubercles. Paired fins sturdy. Fin membranes translucent.

No dermal cirri.

Color. Gilbert recorded for the holotype, “light olive-brown above, white below; 2 small black
spots behind each eye, a second pair on each side the middle of disk, and a third pair above and in
front of each gill-opening; a vertical dark shade immediately behind dorsal fin; a faint dark bar on
terminal portion of dorsal and one on caudal.” A color photograph ofa specimen from Japan (Okamura
et al. 1982:196) shows ground color medium tan with orange-tinged blotches on pectorals and caudal;
the dark spots are as described by Gilbert for the holotype (above). Preserved specimens, including
holotype, pale creamy or pale brown; internal membranes colorless, or peritoneum sometimes with
a few scattered melanophores.

DISTRIBUTION. — Hawaiian archipelago, Philippine Sea, South China Sea, and western Indian
Ocean off South Africa. Bathymetric range: 274-530 (possibly to 900) m.

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FIGURE 19. Frontal views of species of Solocisquama to show contrast between tall single spines on dorsal surface of disk
in S. stellulata (a) and the low complex tubercles in S. erythrina (b) and S. carinata (c). (Note that the esca of the S. erythrina
is nearly gone; ordinarily it would be the same relative size as in the S. ste/lulata specimen.)

BRADBURY: DIBRANCHUS REVIEW 301

MATERIAL. —7 specimens 37.4-101.6 mm SL. HOLOTYPE: USNM 51595 (47.8 mm SL),
ALBATROSS sta. 4080, off N coast Maui, 326-369 m. BPBM 17941 (76.0 mm SL), Hawaiian Is.,
Molokai Channel, 21°02'N, 157°32'W. CAS 42709 (37.4 mm SL), TOWNSEND CROMWELL Cr.
33, sta. 34, 20°01.6’N, 156°54.2'W—20°02.7'N, 156°53.9'W, 344-356 m. CAS 42710 (48.9 mm SL),
TOWNSEND CROMWELL 40-92, 21°04'N, 156°29'W, 274-318 m. IOAN (1, 88.8 mm SL),
28°05'N, 134°40’E, 530-900 m. RUSI 14046 (77.8mm SL), 3°49’S, 40°00’E. ZMMSU
P15841(101.6 mm SL), PROFESSOR MESIATSEV Cr. 6, trawl 124, 4°43’S, 110°56’E, 4 00—
430 m.

Solocisquama erythrina (Gilbert), 1905
Figs. lc; 5a, b; 18b; 19b; 20c, d

Dibranchus erythrinus Gilbert, 1905:697 (original description, holotype from Hawaiian Is., 786-872 m);
Bradbury, 1967:414 (diagnosis of Dibranchus).

DIAGNOSIS. — Unique tubercles multi-spined, that is, with numerous spines clustered in center
of each tubercle instead of single spine (Fig. 18b). No large rostral spine as in S. ste//ulata, nor long
simple spines on dorsal surface of disk; no toothed scutes on tubercles as in S. carinata.

DESCRIPTION. — Based on 6 specimens including holotype, USNM 51642. Counts and body
proportions for the 4 largest specimens plus vertebral counts for all 6 specimens are given in Tables
6 and 7. Skeleton moderately firm. Rostrum short, rounded when viewed from above, slightly
overhanging jaws. Esca with dorsal median lobe and 2 ventral lateral lobes, a strong vertical median
furrow down the dorsal lobe and separating the lateral lobes (Fig. 1c). Lower jaw included in cleft
upper jaw. Nasal capsules exhibit sexual dimorphism: in females, capsule small with round nostrils;
in males, nasal capsules large, swollen because of enlarged lamellae within, with long vertical slitlike
posterior nostril.

Teeth as in generic description. Each ceratobranchial V with a relatively small leaf-shaped patch
of teeth, these patches set at 90° to one another so ensemble resembles a pair of wings in outline. (In
the original description, Gilbert recorded “tongue edentulous,” but this is an error.) No palatal teeth.
Gill rakers finger-shaped with relatively long pedicels, each with distal tooth plate bearing cluster of
tiny teeth. Second gill arch with 6 or 7 gill rakers on each side.

Skin thickly covered with tubercles. Tubercles unique; rather than each with a single central spine
as generally seen in this family, there are 2—6 or more spines clustered (conjoined) in center of each
tubercle (= multi-spined). In holotype and one other large specimen (BPBM 29270), each shows a
variety of sizes of tubercles: large tubercles in several longitudinal rows on dorsal surface of trunk
and fairly evenly distributed over rest of dorsal surface of body; much smaller tubercles fill intervening
spaces; largest tubercles of all, with relatively stronger, longer spines, occur along sides of disk and
tail, those of face modified somewhat to form toothed ridges over lips and excavations of the lateralis
system. Small specimens have many fewer tubercles, these about uniform in size; presumably these
increase in number and differentiate with growth. Skin covering eyeballs densely set with small
tubercles. Subopercular spines long in small specimens, but relatively short and far less prominent in
large specimens. Ventral surface of disk densely covered with small tubercles.

Ventral surface of tail, in small as well as large specimens, covered with multi-spined tubercles,
small and close-set except for principal tubercles, which are relatively large with long prominent
spines (Fig. 20c, d)

No dermal cirri.

Paired and caudal fins with tubercles at their bases; in largest specimens, tubercles occur along

proximal half of fin rays. Dorsal fin rays with a few tubercles, none on anal fin.

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES

302

Volume 51, No. 5

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BRADBURY: D/BRANCHUS REVIEW

304 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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Paired fins relatively long and broad, paddle-shaped, tips of pectoral rays fleshy, skin of pelvics
thick, somewhat puffy.

Color. Gilbert (1905:698) described the holotype as “light carmine-red” with the ventral surface
“suffused with purple” and the fins deeper red. He found the lining of the gill cavities and peritoneum
both “blackish.” In preserved specimens, body and fins completely pigmentless. Lining of gill
cavities, buccal cavity, pale; peritoneum black or brown or pale with black spots.

MATERIAL. — 6 specimens 28.0—135.5 mm SL, all from Hawaiian Is. HOLOTYPE: USNM 51642
(133.5 mm SL), ALBATROSS sta. 3985, vicinity of Kauai, 786-872 m. BPBM 24729 (3, 26.1-65.7
mm SL), TOWNSEND CROMWELL Cr. 59, sta. 20, 21°09'N, 156°11'W, 100-800 m. BPBM 29270
(135.5 mm SL), off N shore of Kauai, 622 m. CAS 42707 (74.4 mm SL), TOWNSEND CROMWELL
Cr. 61, sta. 66, 21°01'-05.7'N, 156°08.4—10.2'W, 786 m.

Solocisquama carinata new species
Figs.18c, 19c, 20a, 21

Dibranchus sp. nova Bradbury; Parin 1990:18 (Sala y Gomez Ridge).

DIAGNOSIS. — Tubercles unique in family, low, without spines, having only 3 or 4 facets, the
ridges between facet surfaces drawn out into scutes, each scute bordered by a comb of fine teeth (Figs.
18c, 20a). Disk margin short, 35% of SL (40% in S. erythrina and S. stellulata of equal size or larger).

DESCRIPTION. — Based on holotype, 73.2 mm SL. Counts and body proportions given in Tables
6 and 7. Skeleton rigid, integument relatively tough. Rostrum very short, only slightly overhanging
mouth, slightly upturned. Upper jaw with median cleft. Esca relatively wide, leaf-shaped median
dorsal lobe very short, two ventral lobes subspherical. This specimen a female with small nasal
capsules and nasal openings.

Teeth in bands on jaws; teeth visible on upper jaw when jaws closed; small oval pad of teeth
anteriorly placed on each ceratobranchial V, pads widely separated at midline. No palatal teeth. Gill
rakers finger-shaped, 7 on each side of second arch, pedicles relatively long as in S. erythrina, distal
end of each with tooth plate bearing cluster of tiny teeth.

Strong, extraordinarily shaped tubercles everywhere over body except fins, each with only 3 or
4 facets, the ridge between facet surfaces drawn out into scutes, each scute bordered with comb of
fine teeth. Tubercles of dorsal surface of disk fairly uniform in size, evenly spaced; tubercles of ventral
surface of disk smaller but uniform size, widely spaced. Tubercles on skin of eyeballs arranged around
iris in concentric rows. Subopercular spines prominent although relatively short. Scutes of tubercles
on edges of disk and sides of tail forming keeled edges for excavations housing lateralis organs.

Ventral surface of tail (Fig. 20a) covered with close-set tubercles uniform in size. Principal
tubercles somewhat enlarged, aligned so carinated scutes form strong keels.

No dermal cirri.

Paired fins relatively large, strong, paddle-shaped, pelvics with skin thickened and puffy,
membranes not emarginate, ray tips not free. Fin membranes translucent.

Color. Specimen totally devoid of pigment externally and in gill cavity; peritoneum with black
splotches.

ETYMOLOGY. — The specific name carinata is a Latin adjective meaning keeled, in reference to
the alignment of tubercles forming keels along the disk edge and sides of tail.

MATERIAL. — HOLOTYPE: ZMMSU P.20359 (female 73.2 mm SL), Sala y Gomez Ridge,
25°08.2'S, 99°25'W, 750-800 m, PROFESSOR SHTOKMAN sta. 1996, bottom otter-trawl, 5 May
1987.

BRADBURY: DIBRANCHUS REVIEW 305

FIGURE 21. Solocisquama carinata new species, holotype, 73.2 mm SL, ZMMSU P.20359, from Sala y Gomez Ridge. (a)
dorsal view; (b) ventral view.

DISCUSSION

The majority of species of Dibranchus reside in the eastern Pacific Ocean, a peculiar distribution
for ogcocephalid genera and uncommon in other fish groups. I adduce the argument that Dibranchus
evolved in the eastern Pacific and offer the following justifications.

Relationships Within the Family. In a survey of the cranium and illicium of many species of
Ogcocephalidae, Bradbury (1967:401, 405) implicitly identified in the Indo-Pacific genus Halieutop-
sis two characters plesiomorphic for the family. First, the frontal bones on the dorsal surface of the
cranium were found to be relatively flat and scalelike, not modified to form a groove for the illictum
(as seen in Dibranchus, Halieutaea, Halieutichthys, and Halicmetus) or a tube housing the illictum
(as seen in the remaining genera Malthopsis, Zalieutes, and Ogcocephalus). Second, the illicial bone
was found to be spinelike and relatively simple, not perforated, thickened and rounded, pitted, or split
as seen in other genera (conditions which, in fact, constituted autapomorphies defining these other
genera). Only one genus, Coelophrys, another Indo-Pacific genus, was omitted from the survey
because no material had been available for clearing and staining.

Subsequently, I examined a cleared and stained specimen of Coelophrys brevicaudata, BSKU
26847. It resembles Halieutopsis. The frontal bones of the cranium, although poorly ossified, show
no deformation to form a groove, and the illicial bone is elongate, slender, and sub-spinelike, not
perforated or split or thickened and rounded. If C. brevicaudata is representative of the genus, perhaps

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future work will indicate that Coelophrys and Halieutopsis ought to be combined in a single genus.
Meanwhile, it is fair to consider that the Coelophrys/Halieutopsis complex 1s the closest to an ancestral
ogcocephalid that we have, because it is difficult to think that it can be derived from taxa with grooved
or tube-bearing skulls. Coelophrys and Halieutopsis micropa differ markedly from other ogco-
cephalids in having globose or sub-globose bodies, and they, plus several other species of Halieutop-
sis, have markedly loose skin with subdermal gelatinous layers. I take the view (below) that these are
plesiomorphic characters within the Ogcocephalidae.

In two comprehensive papers, which were landmarks for lophiiform systematics, Pietsch (1981,
1994) presented two different hypotheses for sister-group relationships of the Ogcocephalidae. The
earlier hypothesis (1981) proposed that Chaunacidae and Ogcocephalidae were sister-groups, but this
was superceded (1994) by the hypothesis that the sister-group of Ogcocephalidae is the suborder
Ceratioidei (with Chaunacidae the sister-group of combined Ceratioidei and Ogcocephalidae; see the
Pietsch papers for discussion). In either case, the implicated taxa are pelagic forms with (usually)
globose or sub-globose bodies and, in the main, soft loose skin, sometimes with subdermal gelatinous
tissue. When scales occur in the skin, they are small, simple tubercles occurring over the entire body,
little or not at all differentiated on different parts of the body. Coelophrys and Halieutopsis micropa
appear to be benthopelagic and have globose or sub-globose bodies, soft loose skin, subdermal
gelatinous tissue and simple, undifferentiated tubercles completely covering the body. The pelagic
postlarval stages now known among ogcocephalids are all globose with loose skin, extensive
subdermal gelatinous tissue and (usualiy) simple, small tubercles covering the body. It is worth
considering that these are plesiomorphies for the family, that pelagic life is the ancestral condition
for the Ogcocephalidae, and that Coelophrys and similar species of Halieutopsis are paedomorphic.
Forms with benthic adults would, then, all be derived. These would include all the genera with frontal
bones forming a groove or tube, that is, all other known genera of Ogcocephalidae.

There is one more point of interest having to do with the condition of the lateral line canals and
neuromasts. Pelagic forms like Coelophrys have excavations in the skin which may be interpreted to
be open lateralis canals. Lying within the canals are large prominent free neuromasts, a condition
common in bathypelagics. All other species of ogcocephalids have versions of this condition even
though they are benthic forms, some of which live as far inshore as to be nearly subtidal. In benthic
deepwater forms like Halieutopsis and Dibranchus, the free neuromasts on the ventral surface of the
disk and lateral sides of the tail still lie in canals, but are sheltered by modified scales and cirri, the
cirri sometimes in thick mats. On the dorsal surface of the bodies in these forms, the neuromasts are
reduced in size and nestled in and among specialized scales, for the open canals have disappeared. In
forms like Ogcocephalus and Malthopsis, found at continental shelf depths where, presumably, waters
are more turbulent, all canals are lost, but the neuromasts remain free on the body surface, where they
are greatly reduced in size but still protected by specialized scales and cirri and situated in depressions
among strongly calcified bucklers. The loss of canals and tiny size of neuromasts are derived character
states, modifications for benthic life. The open lateralis canals and large free neuromasts as seen in
Coelophrys are characters suited to pelagic life, the hypothesized ancestral condition for ogco-
cephalids.

Of the genera with frontal bones forming a groove, Dibranchus is probably the one closest to
Coelophrys/Halieutopsis. | have found two synapomorphies. One is the reduction of gills to 2. The
other is the peculiarity of the lateral line such that the lateral line is interrupted just where it passes
from the disk to the tail and the anterior part of the tail lateral line runs instead onto the ventral surface
of the tail towards, or in advance of, the anus. Halicmetus shares these characters, but has been very
poorly studied; it is set apart by having scales in the form of bucklers, an apomorphy it shares with
all the genera with a cranial tube for the illicium. That is, Dibranchus is probably more closely related
to Halieutopsis than to Halicmetus.

BRADBURY: DIBRANCHUS REVIEW 307

Relationships Within the Genus Dibranchus. Most species of Dibranchus have tubercles
showing derived character states including enlargement of some tubercles with loss of others, strong
differentiation on different body parts, tubercles ankylosed to underlying skeletal parts, patterns of
spinules overlaying tubercles, small tubercles overlaying the bases of large tubercles, and so on. The
majority of dibranchids also have lost the palatal teeth, and most show loss in one or another of meristic
characters. The two Atlantic species and the three Indo-Western Pacific species have all lost the palatal
teeth and show other apomorphies, as follows. Dibranchus atlanticus has a radiating pattern of
spinules on the tubercles and has a reduced number of vertebrae, 18 instead of 19. Dibranchus
tremendus, D. hystrix, and D. accinctus all have tubercles very reduced in number, with many
remaining tubercles greatly enlarged and bearing enormous spines, especially on the tail. These three
species also have lost one neuromast from the subopercular series. Finally, D. japonicus, although
retaining simple tubercles all over the body, where they are relatively undifferentiated on different
parts of the body, has peculiar leathery skin, an autapomorphy.

All dibranchids that retain palatal teeth are eastern Pacific species: D. spinosus, D. sparsus, D.
erinaceus, and D. discors, but D. discors is removed from candidacy for being most like an ancestral
Dibranchus by these derived characters: loss of tubercles from portions of the body and minute
spinules on large tubercles. Also, its laterally compressed illicial bone, narrow mouth and interorbital,
and large eye, all unique for Dibranchus, make D. discors a highly derived Dibranchus.

Of the remaining three species with palatal teeth, the widespread D. spinosus has small simple
tubercles without giant spines, evenly distributed over the body, without much differentiation on
different parts of the body, in skin that is relatively somewhat soft and loose. It may be the closest we
have to an ancestral Dibranchus. The other two species, D. sparsa and D. erinaceus, are very similar
to D. spinosus, but D. sparsa has lost one neuromast from the preopercular lateral line series, and D.
erinaceus has moderately enlarged tubercles interspersed in a regular pattern among small simple
tubercles.

All these observations and guesses taken together suggest that Dibranchus, while descended from
Indo-Pacific forms, first radiated in the eastern Pacific, with descendants of that radiation populating
the Atlantic, western Pacific, and Indian Oceans.

Relationships of the Genus Solocisquama. With ogcocephalids still so poorly known, hypothe-
ses about relationships, of course, await revision, especially in the case of Solocisquama. Characters
so far known suggest two different hypotheses. One is that Halicmetus is the primitive sister-group
of Solocisquama. One suspected synapomorphy is the conformation of the illicial bone, with two
dorsolateral processes replacing (derived from?) the median dorsal process. The two genera share two
other derived characters, an interrupted lateral line with one neuromast on the ventral surface of the
tail on each side, near the anus, and loss of palatal teeth. However, these latter two characters are also
shared with Dibranchus. If Solocisquama instead shares a common ancestor with Dibranchus, it
would mean that an eastern Pacific genus has given rise to a distinctive eastern Pacific genus.

ACKNOWLEDGMENTS

I thank the curators and staff members of the numerous institutions that provided material for
this study; my colleagues T. Iwamoto and W. N. Eschmeyer of CAS were particularly generous. I
gratefully acknowledge the extensive technical assistance given me by D. Catania (CAS) and K. Hartel
(MCZ), the helpful information I received from G. Burgess, C. R. Robins, N. V. Parin, R. N. Lea, C.
Karrer, and J. L. Castro-Aguirre, and the important specimens from O. Okamura and T. Machida, N.
Chirichigno F., and N. V. Parin. Discussion that enlightened was the gift of M. E. Anderson and R.
N. Lea; also of J. E. McCosker, who prodded and encouraged. For painstaking reviews, I thank an
anonymous reviewer and D. M. Cohen, whose judicious comments decidedly sharpened this report.

308 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 5

I am grateful to the Department of Biology, San Francisco State University, San Francisco, for
use of their photographic facility and their electron microscope facility, and to G. Antipa and K. Letsch
for technical coaching.

APPENDIX

Specimens Used for Scanning Electron Micrography of Gill Rakers, Premaxillae, and
Tubercles. Dibranchus atlanticus, 131.5 mm SL, CAS 76452; the following all from CAS 76446:
39.5 mm SL, 72.5 mm SL, 81.0 mm SL. Dibranchus erinaceus, 94.0mm SL, CAS-SU 60476.
Dibranchus hystrix, 82.0 mm SL, CAS 82242. Dibranchus nudivomer, 83.5 mm SL, CAS 82261.
Dibranchus sparsus, 75mm SL, CAS 82227. Dibranchus spinosus, 139.0mm SL, ZMUC
GALATHEA sta. 739; 140.0 mm SL, CAS 82235. Dibranchus spongiosus, 71.0 mm SL and 115.0
mm SL, both CAS 82252. Dibranchus tremendus, 158.0 mm SL, USNM 320332. Solocisquama
erythrina, 135.0 mm SL, BPBM 29270.

Specimens Used for Clearing/Staining. Dibranchus cracens, 128.3 mm SL, CAS 88034
(illictum). Dibranchus discors, 87.2 mm SL, CAS-SU 47195 (illictum). Dibranchus erinaceus, 94.0
mm SL, CAS-SU 60476 (gill arch). Dibranchus hystrix, 82.0 mm SL, CAS 82242 (gill arch).
Dibranchus nudivomer, 82.3 mm SL, CAS 82261 (entire). Dibranchus spongiosus, 115.0 mm SL,
CAS 82252 (entire). Dibranchus tremendus, 96.5 mm SL, UF 25923 (illictum). Solocisquama
erythrina, 135.0mm SL, BPBM 29270 (gill arch), and 65.7 mm SL, BPBM 24729 (illicium).
Solocisquama stellulata, 76.0 mm SL, BPBM 17941 (illicum), and 101.6 mm SL, ZMMSU P.15841
(gill arch). Coelophrys brevicaudata, BSKU 26847 (entire).

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© CALIFORNIA ACADEMY OF SCIENCES, 1999
Golden Gate Park
San Francisco, California 94118

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES

Volume 51, No. 6, pp. 311-336, 8 figs., 5 tables. August 20, 1999

Systematics of the Garter Snake Thamnophis atratus at the
Southern End of Its ange». ne Biological Labora —

Orato
cc ANOgraphic
ry

by

S
Jeff Boundy BE 0 : 1999

Museum of Natural Science, Louisiana $tate Ui "We sity
Baton Rouge, Louisiana 70803 Gus AGle, Ma 02848

Variation in populations of Thamnophis atratus, south of the range of T. a. hydrophilus,
was examined through multivariate analysis of nineteen characters among seven geo-
graphic units. A phylogenetic analysis of five population groups was performed using 24
characters, resulting in a single, most parsimonious tree. Variation in scalation, color
pattern, tooth counts, and head morphology suggested the presence of two taxa south of
the range of 7. a. hydrophilus: typical T. a. atratus was restricted to California’s Santa Cruz
Mountains, while other central California Coast Range populations were described as a
new subspecies. Thamnophis a. aquaticus was found to represent populations intermediate
between 7. a. atratus and T. a. hydrophilus, and is synonymized as a representative of an
intergrading, non-diagnosable population. Ecological observations found differences in
habitat use and seasonal activity between coastal and inland populations. A biogeographi-
cal model was proposed for the derivation of the subspecies, and a new subspecies, 7. a.
zaxanthus, is defined.

Resolution of garter snake (Thamnophis) systematics in western North America has proven to
be a complicated process that is incomplete. Thamnophis atratus is a polymorphic species ranging
from southwestern Oregon to the coast of central California (Rossman et al. 1996). Several systematic
problems are unresolved within 7hamnophis atratus, including interpretation of clinal variation
within the species, status of Thamnophis atratus aquaticus (Rossman and Stewart 1987), and
taxonomic identity of the inland, three-striped morph of 7. a. atratus (Fox 1951). Fox (1951) analyzed
variation within the species and defined three subspecies based on north-south character shifts. He
included all three taxa within Fitch’s (1940) hydrophilus group (= aquatic morph; Thamnophis couchii
complex of Rossman and Stewart 1987 [7. couchii, T. hammondii, T. gigas, T. atratus]). Fitch (1940)
characterized the aquatic morph, including T. a. hydrophilus, as having reduced or indistinct striping,
an obscure, spotted pattern, subdued colors, an elongate, pointed muzzle, reduced size of posterior
supralabials reflecting reduced salivary glands, relatively large eye, small pupil, and pale iris. In
contrast, Fitch’s (1940) elegans and ordinoides groups (= terrestrial morph; 7. elegans, T. ordinoides,
T. a. atratus) were characterized by bold striping on a black background, bright yellow throat and
dorsal stripes, relatively blunt muzzle, increased height of posterior supralabials, relatively small eye,
large pupil, and dark iris. Thamnophis a. atratus has been thought to exhibit some characters
convergent with the terrestrial morph (Fox 1951; Bellemin and Stewart 1977). A third subspecies,
T. a. aquaticus (Fox 1951), occurs between T. a. atratus and T. a. hydrophilus immediately north of
San Francisco Bay, and displays characteristics intermediate between the other two subspecies. Fox
(1951) reported intergrades between adjoining subspecies. Rossman and Stewart (1987) considered

Sit

312 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 6

T. a. aquaticus to be weakly differentiated morphologically from 7. a. hydrophilus, and molecular
studies by Lawson and Dessauer (1979) reflected this conclusion.

Variation is exhibited within each subspecies of Thamnophis atratus. Fitch (1940), Fox (1951)
and Rossman (1979) detailed variation within 7. a. hydrophilus, and Rossman and Stewart (1987)
provided details for 7. a. hydrophilus and T. a. aquaticus. Rossman (1979) and Rossman and Stewart
(1987) showed that 7. a. hydrophilus is polymorphic with respect to color pattern, having distinctly
spotted and striped snakes occurring through much of the range. Fox (1951) detailed variation in 7.
a. atratus and found it to comprise two morphs: “blue-black,” single-striped snakes on the San
Francisco Peninsula, and three-striped snakes inland and from coastal Monterey County. He found
this variation “difficult to explain,” and did not define the two color morphs as subspecies. I observed
that these two color morphs exist on either side of the Santa Clara Valley, with intermediates occurring
at the southern end of the Santa Cruz Mountains. This observation prompted me to conduct a
multivariate analysis of morphological variation in 7. a. atratus and T. a. aquaticus to determine
whether or not the current delineation of these subspecies is accurate. I tested the hypothesis that
variation within 7. atratus reflects a partial shift from an aquatic to a terrestrial morphotype, using
sympatric Thamnophis elegans as the model for the terrestrial morph. Analyses of variation in T. a.
hydrophilus (Fitch 1940; Fox 1951; Rossman and Stewart 1987) precludes the necessity of reevalu-
ating northern populations of 7. atratus.

MATERIALS AND METHODS

A total of 1054 snakes, both live (including field observations) and preserved, was examined
from the southern half of the distribution of Thamnophis atratus (south of Mendocino, Lake, and
Colusa counties, California). Specimens were examined in the collections of the California Academy
of Sciences (CAS, CAS-SU), California Polytechnic State University, San Luis Obispo (CPS),
Louisiana State University Museum of Natural Science (LSUMZ), Museum of Vertebrate Zoology
(MVZ), San Jose State University (SJSU), University of California, Santa Barbara (UCSB), and
National Museum of Natural History (USNM; see Appendix A and B). From 1969 through 1994, 506
T. atratus were observed in the field. Coloration, pattern, and ecological notes were recorded for each,
and all but 63 snakes were released. Four gravid females, were kept alive until parturition. These,
their offspring, and the other retained snakes were preserved and deposited in SJSU or CAS.

Only snakes over 350 mm SVL, the approximate minimum adult snout-vent length (Rossman
1979), were used for the multivariate analysis to assure that ontogenetic growth of cephalic features
had approached the asymptote. Gape length was measured to the nearest 0.1 mm. Counts were made
of ventrals, subcaudals, preoculars, postoculars, supralabials, infralabials, and temporals. Anterior
scale rows were counted at the tenth ventral scale posterior to the chin shields. Midbody scale rows
were counted halfway between occiput and vent. Posterior scale rows were counted at the antepenul-
timate ventral.

Vertebral stripe width was measured in number of scale rows covered at the widest part of the
nuchal area (anterior stripe width), and at a point about one-third the distance from snout to vent
(midbody stripe width). A ranked value (stripe value) was derived from relative visible intensity of
the lateral stripe, if present, and intensity of ventral dark pigment. Because these characters were
correlated, only the single ranking was necessary. Dark snakes with only a vertebral stripe were given
the lowest value of | (Fig. 1f), while three-striped snakes with pale labials and venter received the
highest value, 5 (Fig. 1d, h). Color patterns shown in Fig. le and 1g were ranked 2 and 4, respectively.

Clinal trends in cephalic proportions were compared through the following six ratios: antepenul-
timate supralabial width/height (to eliminate contour curvature of the supralabials, the stratum
corneum was removed and measured on a flat surface); internasal median suture length/combined
internasal posterior width; left internasal-rostral contact/combined nasal-rostral contact (after Fitch

313

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BIWIOJILD “OD ZNID eyURS ‘apRASIOIUL sNYJUDXDZ ‘YJ x SNIDAID YD *] (D) (OF9L6] SVD) PIWofITRD “OD aye] URS ‘sNID.UD “D -] (4) (€76Lb ZAW) BIUIOJI[R) “OD Oa}RJ] URS
‘snjpajD “DJ jeseq (J) ‘(waned . snouonbp,, ‘prs 16 SVD) BlusojpeD “OD eden ‘snyjunxnz ‘p ‘| leseq (q) (L7STpe ZAW) BlwWoseD “OD Ayuiy ‘sapydospdy P -1 (D) (OFS 16]
SVD) PlWOf TR “OD esnjod ‘snjiydoipay ‘v7 (gq) (adAjodoy ‘¢ZpL | ZAW) UORAIQ “OD uosyore ‘snpiydoipay ‘D °] (W) :snypao siydouupyy jo susayed [esiog *| AYN

BOUNDY: SYSTEMATICS OF THAMNOPHIS ATRATUS

314 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 6

1940); anterior chin shield length/posterior chin shield length (greatest length of each shield); eye
diameter/gape length; interocular width/gape length. A seventh ratio compares tail length to total
length.

Specimens of 7. atratus examined from Lake and Colusa counties, and north of the Gualala River,
Sonoma County, were referrable to 7. a. hydrophilus as defined by Fox (1951) and Rossman and
Stewart (1987), and determined the northern limit of the area considered in the present paper. Southern
Thamnophis atratus populations were divided into seven geographic samples for geographic analysis
(Fig. 2). Five areas were selected because they were isolated from each other due to the presence of
San Francisco Bay or naturally uninhabited valleys (Sonoma, Santa Clara, and Salinas). Two
additional areas (C and E) are annectant to Area D, but were considered separately due to a priori
recognition of intermediate or sympatric color pattern morphs in those regions. Hartley’s Fmax test
was used to insure homogeneity of variances for each population and each character. Area A includes
all of Marin County, and Sonoma County west of the Russian River and Sonoma Valley. Area B
includes the Coast Range of Napa and western Solano counties. Area C includes the San Francisco
Peninsula north of Half Moon Bay and San Andreas Reservoir. Area D includes southern and central
San Mateo County and the Santa Cruz Mountains south to the San Lorenzo River and Guadalupe
Creek watersheds. Area E includes the southeastern flank of the Santa Cruz Mountains north to the
Alamitos Creek watershed. Area F includes the Diablo Range. Area G includes the Santa Lucia and
San Rafael mountains from Monterey to Santa Barbara counties. Areas A and B are currently referable
to T. a. aquaticus, and Areas C-G to T. a. atratus.

The morphological data were tested for concordance with geographical distribution. A multivari-
ate analysis was performed with the SYSTAT Discriminant Function program (Wilkinson 1988). One
hundred-eighteen adult males and 115 adult females were utilized for the MANOVA. Character
correlation was determined by Pearson correlation matrix using SYSTAT. The BIOSTAT t-test
application (Pimentel and Smith 1986) was used to determine significance of sexual dimorphism for
each character. Additional morphological analyses were performed with STAT VIEW II (Feldman et
al. 1986).

Phylogenetic relationships of populations in areas were evaluated with the branch-and-bound
option of PAUP version 3.1 (Swofford 1993), using ACCTRAN optimization. Thamnophis a.
hydrophilus was used as the outgroup because it is the sister taxon to southern populations of T. atratus
(Lawson and Dessauer 1979). The use of phylogenetic trees is applicable because I believe southern
populations of 7. atratus evolved in a hierarchical fashion (see Biogeography). However, Areas C
and E were left out of the phylogenetic analysis because they proved to be intermediate or mixed
populations that would not conform to terminal taxa in a phylogeny. To enhance phylogenetic tree
resolution several additional characters were used, including maxillary tooth count and the following
ratios: pupil diameter/eye diameter, posterior frontal width/anterior frontal width, ventral margin
length of sixth supralabial/length of seventh supralabial, combined internasal and prefrontal median
suture length/frontal length, frontal length/parietal length, dorsal/ventral length of loreal, and prefron-
tal median suture length/internasal median suture length. Two ranked characters were also examined:
ground color, ranked from medium gray or brown (= 3) to black (= 5), and, the projected angle formed
by the suture between the sixth and seventh SL (1 = through eye, 2 = to posterior margin of eye, 3 =
to posterior margin of postoculars, 4 = roughly vertical). Character means are provided in Table 1.
These data were obtained from specimens in the LSUMZ collection (Appendix B), and from data in
Rossman and Stewart (1987). Character states were ordered on a scale in proportion to the magnitude
of difference between character means (Thiele 1993): the smallest observed distance between any
pair of means was given a distance of one, and all other pairwise distances were coded relative to the
smallest distance. This method preserves evolutionary information from continuous variables, and
was analyzed using the step matrix procedure in PAUP.

BOUNDY: SYSTEMATICS OF THAMNOPHIS ATRATUS

FiGURE 2. Population samples of Thamnophis atratus used for
analyses (Areas A-G) in central California; lined area represents ap-

proximate distribution of Thamnophis a. hydrophilus.

iat

315

The test of convergence between
Thamnophis a. atratus and T. elegans ter-
restris was performed by alternatively con-
straining trees which would compare T. e.
terrestris (convergence) or 7. a. hydrophi-
lus (non-convergence) as sister taxon to
southern 7. atratus populations. The latter
arrangement is supported by allozyme and
DNA sequence analyses (de Queiroz and
Lawson 1994). Morphological data for 7.
elegans was obtained from LSUMZ speci-
mens (Appendix B) and from Bellemin and
Stewart (1977).

RESULTS

SEXUAL DIMORPHISM.—Prior _ to
ANOVA, sexual dimorphism was exam-
ined to ensure statistical homogeneity
(Thorpe 1989). Sixteen characters were
analyzed using Student’s t-test, eight of
which were significantly dimorphic (Table
2). Thirteen non-correlated characters
were examined for sexual dimorphism by
discriminant function analysis: predicted
vs. actual assignment resulted in 112
(97.4%) of 115 correct female assign-
ments, and 115 (97.5%) of 118 correct
male assignments (F = 63.506, df= 13,
219, p<0.001). Significant sexual dimor-
phism in half of the characters tested ne-
cessitated separate multivariate analyses
for males and females.

Female SVL increased at a greater rate
than that of males (Fig. 3; comparing larg-
est male with largest female and progres-
sively comparing pairs of smaller snakes).
The ratio of mean male/female snout-vent
length for each area is A = 0.96, B = 0.83,
C=0.92, D=0.88, F=0.88, and G=
0.86. Only three adult females were avail-
able from Area E.

TOOTH COUNTS.— Maxillary teeth,
per side, were not significantly different
between sexes: male range 23—27 (mean
25.3, n= 46), female range 21-28 (mean
25.2, n= 42). Tooth counts were signifi-
cantly different between areas (F = 5.64 at
5, 95 df, p<0.0001), but not between areas

316 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 6

TABLE 1. Character means for Thamnophis elegans and six population groups of 7. atratus used for
phylogenetic analyses. Abbreviations: ACS/PCS, anterior/posterior chin shield length; ASR, anterior scale
rows; ED, eye diameter; FL, frontal length; GL, gape length; INL, internasal median suture length; INR,
internasal-rostral contact length; IOW, interorbital width; LDL/VL, loreal dorsal/ventral lengths, M/F SVL,
male/female snout-vent length ratio; ML, muzzle length; MVSW, midbody vertebral stripe width; NR,
nasal-rostral contact length; PD, pupil diameter; PFW/AFW, posterior/anterior frontal width; PL, parietal
length; PrF, prefrontal median suture length; PSR, posterior scale rows; SC, subcaudals; SL6/SL7, ventral
lengths of supralabial 6/supralabial 7; SLW/H, sixth supralabial width/height; STR, stripe value; SVL,
snout-vent length; TL; tail length; TTL, total length; V, ventrals.

Six population groups of 7. atratus T. elegans

T. a. hydro- Area-A Area B Area D Area F Area G

philus
SVL (mm) 466.5 380.3 407.4 392.7 415.3 400.0 467.0
V 164.2 IS 7h 161.1 153.4 159.5 153.3 158.0
SC 86.6 83.8 82.7 79.4 80.6 WS 80.0
ASR 20.2 19.0 19.6 19.2 19.4 19.2 19.7
PSR 16.9 15.4 16.1 14.9 ISA 15.5) 17.0
IOW/GL 0.38 0.40 0.39 0.40 0.39 0.40 0.40
MVSW ila 1.9 1.9 2.4 Dep 2.0 1.8
STR 4.3 3.9 5.0 13) 5.0 5.0 5.0
SLW/H 0.99 1.10 1.04 1.04 1.11 1.08 0.92
INR/NR 0.82 0.93 0.93 0.86 0.83 0.83 1.21
ACS/PCS 0.83 0.79 0.80 0.76 0.80 0.78 0.94
WLE/AMOL, 0.26 0.27 O27 0.26 0.26 0.26 0.24
ED/GL 0), 7 0.18 0.19 0.19 0.19 0.19 0.17
Max. teeth 23.6 24.8 DoT 24.5 DGD 25.6 16.9
M/F SVL 0.86 0.91 OMS 0.81 0.78 0.81 0.77
PD/ED 0.45 0.49 0.43 0.47 0.43 0.45 0.60
PFW/AFW 0.83 0.80 0.76 0.75 0.81 0.84 0.75
SL6/SL7 1.29 1.38 1.26 123 1.18 1.42 1.47
ML/FL 0.85 0.67 0.67 0.70 0.69 0.65 0.94
FL/PL 0.92 1.02 1.02 0.97 1.09 1.16 0.96
LDL/VL 0.71 0.66 0.65 0.63 0.64 0.56 0.58
PrF/INL 1.08 0.85 aval 0.98 0.97 1.01 1.20
SL angle 3.4 3.4 35) 3.7 3.4 3.2 Sal]
Ground color 32 4.0 4.0 5.0 4.7 4.3 4.6

C and D, and areas B and G (Fischer’s Least Significant difference = 0.37 at 71 df).

CORRELATION.— Nineteen characters were initially analyzed by MANOVA (Table 3). Linear
measurements (snout-vent length, gape, interocular width, eye diameter) were significantly correlated
(r >0.608). Eye diameter was hypothesized to be affected by an aquatic-terrestrial morph shift, plus
had the lowest variance, and was retained. Relative tail length was correlated with subcaudal count
(r >0.571), and dropped from further analysis in favor of the non-ratio measurement. Relative eye
and interocular proportions were dropped from the discriminant function analysis, since the linear
components of each were included separately.

DISCRIMINANT FUNCTION ANALYSIS.— The remaining thirteen characters were statistically
analyzed for each sex. For males, Wilks’ lambda test resulted in F = 6.245, df=78, 551, p>0.0001,
among Areas A~G. Univariate F tests showed ten variables to be significant (p<0.05) based on squared
multiple correlations, eliminating anterior scale row count, relative internasal length, and relative chin
shield length (Table 4). The test of residual roots indicated that only Factors (= discriminant functions)

BOUNDY: SYSTEMATICS OF THAMNOPHIS ATRATUS ST

TABLE 2. Student’s t-test results for sexual dimorphism in Thamnophis atratus from area D (Santa
Cruz Mountains region); data presented are means, standard deviations and sample sizes.

Character Males Females (p)

Snout-Vent length 420.0 + 4.6 (25) 468.9 + 7.5 (25) 0.001
Ventrals 158.4 + 0.5 (25) 152.4 +0625) 0.001
Subcaudals 80.4 + 0.5 (25) 72.9 + 0.4 (25) 0.001
Anterior scale rows 19S 02025) L92ZDOI 25) n.s
Midbody scale rows 18.4 +0.1 (25) neyoves (D-IL (59) n.s
Posterior scale rows IS -Se= OEY), 1S 2ee Oss) n.s
Anterior vertebral stripe width 3.6+0.1 (20) 35 = 051120) n. s
Midbody vertebral stripe width 2.3 + 0.0 (20) 2.2 + 0.0 (20) n.s
Stripe value Leh Op. 0)) [.7 = 0.1 (20) n.s
6th Supralabial width/height 85.9 + 0.7 (35) 79.1 + 0.7 (35) 0.001
Internasal length/width 109.2 + 1.1 (19) 108.3 + 1.3 (19) n.s
Internasal-rostral/nasal-rostral 910+ £.3 25) 80.5 + 1.8 (25) 0.001
Anterior/posterior chin shield #93, £09 (19) 78.7 0.8 (19) n.s
Tail/total length 26.0 + 0.2 (18) 24.5 + 0.2 (18) 0.001
Eye diameter/gape length 18.8 + 0.2 (30) 18.0 + 0.1 (30) 0.05
Interorbital width/gape length 39.9 + 0.3 (16) 36.3 + 0.3 (16) 0.001

1-3 were significant (p<0.0001), with canonical correlations >0.500. For females, Wilks’ lambda test
also resulted with p<0.0001 (F = 6.407, df = 78, 535). The univariate F-test found 11 variables to be
significant (p<0.05), eliminating internasal-rostral contact and relative chin shield length. The test of
residual roots indicated that factors 14 were significant (p<0.01), with canonical correlations >0.482.

Canonical plots (Fig. 4) indicate the existence of several distinctive population groups for both
sexes. Factor | of males and Factor | of females separate Area D snakes from Area AB and FG snakes.
The chief distinguishing character of Area D snakes is the stripe value: 1.4 for Area D and 4.7 for
Areas A, B, F, and G. Ventral count is nearly identical for males between Areas D and G, but the Area
D mean (153.4) was lower than the mean for Areas A, B, and F (159.2). Posterior scale row counts
in Area D are lower than all others: 15.0 vs. 15.6. For Area D, midbody stripe width averages wider
(2.7 scale rows) than other Areas (combined average of 2.0 scale rows).

Factor 2 of males and factor 3 of females separate North and South Bay populations, although
supporting characters are not discretely different. Area F—G males are longer than Area A-B males
(mean snout-vent length = 409.7 mm vs. 392.9 mm), have larger eyes (mean eye diameter = 3.05
mm vs. 2.84 mm), shorter tails (tail length/total length = 0.259 vs. 0.267), fewer subcaudals (mean =
78.8 vs. 83.3), and more pointed internasals (internasal length/width = 0.83 vs. 0.93). Stripe value
differs (4.9 for Areas F, G, 4.3 for Areas A, B; t= 10.9 at 225 df, p<0.001), but is thought to be
influenced by intergradation with Area D snakes in coastal Marin County snakes. Females have
significantly taller 6th supralabials in Areas F—G (supralabial width/height = 0.80 vs. 0.87).

Factor 3 of males and Factor 2 of females separate between Areas A and B and Areas F and G
at a significant level (p<0.001), based on ventral count (mean = 148.8 vs. 154.6 for females), midbody
stripe width (mean = 1.8 vs. 1.9), and snout-vent length (mean = 394.5 mm vs. 493.9 mm for
females). Factor 1 of males also distinguishes Area A from B based on posterior scale row count
(mean 15.4 vs. 16.1) and stripe value (mean = 3.9 vs. 5.0).

Predicted vs. actual assignments based on discriminant scores shows homogeneity (at least 75%
correct assignment) for all but Areas C and F (Table 5). For Area C, 70.0% of the snakes are correctly
assigned, while an additional 13.3% are assigned to Area D and 3.3% to Area A. For Area F, 65.4%
of the snakes are correctly assigned, while 15.4% are assigned to Area B and 13.5% are assigned to
Area G.

318 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 6

TABLE 3. Summary of basic statistics (mean, standard deviation, range) for male and female Thamnophis. atratus:
abbreviations as in Table |.

Area N SVL Vv SC ASR MSR PSR G ED IOW AVSW

A 16 380.3427.5 157.144.0 83.8+4.1 19.0+0.0 19.0+0.0 15.4408 14.8411 2.7402 5940.2 3.2406
(350-430) (149-164) (76-93) (19) (19) (15-17) (13.0:16:8) (2:2:3:1)__ GS-64) (nleeay
B14 407.4425.3 161.143.8 82.743. 196409 19.0400 161+1.0 15.8+1.2 3.0402 61404 29404
(355-462) (156-168) (75-88) (19-21) (19) (15-17) (13.9-18.7) (2.6-3.4) (5.5-7.0) _(1.9-3.3)
C 15 406.7425.1 150.944.0 77.2440 19.3+40.7 185409 15.0+0.0 159412 29402 63404 3.7409
(365-450) (144-158) (71-84) = (19-21) (17-19) (15) (13.8-17.8) (2.5-3.4) (5.4-6.7) (2.8-6.4)
D 32 392.7430.2 153.4429 79.4+5.0 19.2405 181410 149404 156413 29402 61404 38408
(350-457) (146-159) (67-89) = (19-21) (17-19) (13-15) (12.8-16.0) (2.6-3.5) (5.5-7.0) —(2.9-6.5)
E 8 41264276 157.443.2 79.0442 19540.9 18.041.1 15.0+0.0 166+1.1 3.2403 65405 38+08
(370-452) (153-162) (73-88) (19-21) _—- (17-19) (15) (15.0-18.0) (2.7-3.6) (5.6-7.0) —(3.0-5.0)
F 21 415.3441.7 1595+4.9 806449 19.4408 185409 154408 160412 3.0403 62406 3.6406
(354-514) (149-168) (68-89) (19-21) (17-19) (15-17) (13.9-18.7) (2.5-3.8) (5.1-7.2) — (2.8-5.0)
G 12 400.0432.0 153.3+3.6 755446 19.2408 188406 155409 165+1.1 3.1402 66404 2840.6

(350-461) \-49-161) (65-82) (19-21) (17-19) (15-17) (14.8-18.5) (2.8-3.4) (6.1-7.6) — (1.5-3.5)

A 12 394'5:4:386 48:83:19 7472250 1932018 18i8+06 15:33:10 164+15 28203 6.205) 3307
(356-479) (144-156) (71-78) (19-21) (17-19) (15-17) (14.8-19.0) (2.5-3.4) (5.2-7.1) — (2.2-4.7)
B 10) 49319 75.2) 15416-1233) 76:1-b2-2) 19:4 10:8) 19:0 OO) GOT 20:25 3:2) 3S) 10l4 EEE Oe se le Onl
(385-609) (152-159) (74-79) (19-21) (19) (15-17) (15.2-25.0) (3.0-4.3) (5.7-8.5) — (2.1-4.6)
Cc TS R44207 4] Aol 3-6 ZONE Sale e905 Sel8:Se ILO) Sts OOF a1'S'SeE1E8) | e3aIt-E OSG: See OSes ey -tOrd
(366-525) (141-151) (65-75) (19-21) (17-19) (15-17) (15.1-21.5) (2.6-3.8) (6.1-7.9) — (2.0-4.6)
Dy sy) AVG HOVER I iilse sks) WOessiit NOEs Theses ilo) eyes) ieee “sysiaaQl pact) chil se 40
(351-582) (141-155) (64-79) (18-21) (17-19) (15-17) (14.2-23.3) (2.6-4.2) (5.1-8.2) — (2.7-6.3)
E gy Asan) Ose WOsieasks AlOes00) MWesily) WeMojss0)0) iss laf skasOy! PsjesOs7/ Si7/ a2 0G
(417-537) (146-149) (66-73) (21) (17-19) (15) (16.9-19.9) (2.9-3.7) (6.5-7.9) (3.1-4.2)
F Bi 47NES80:2) 154932 S134 7SiG 4a) 19 SEOL9) V81G)320/8) 15:3)-E018) 1827/5 257) 3-3) 51014 16:9 OS eS oe Oey
(359-701) (142-167) (66-83) (19-21) (17-19) (ISSN) (1520-2424) (255-481) 1S 25-Se7) 9a)
G 12 463.54+68.7 147.6+4.1 69.8+3.0 193408 18.7408 158+1.0 19.4423 33403 7.4408 3.0404

(368-627) (140-156) (63-74) (19-21) (17-19) (15-17) (15.8-23.0) (2.8-3.8) (6.0-8.5) (2.3-3.6)

BOUNDY: SYSTEMATICS OF THAMNOPHIS ATRATUS

319

TABLE 3. Summary of basic statistics (mean, standard deviation, range) for male and female Thamnophis atratus

(continued).

MVSW

1940.5
(1.2-2.9)
19403
(1.3-2.6)
2540.2
(2.2-3.0)
2.44 0.6
(1.6-4.0)
2340.2
(2.1-2.7)
2.24 0.2
(1.8-2.5)
2.0+0.2

(1.5-2.4)

1.7406
(1.2-2.8)
2.0+0.4
(1.2-2.4)
2.4403
(2.1-2.9)
2.9+0.9
(1.8-4.6)
2.2+0.4
(1.5-2.9)
2.2+0.4
(1.0-2.8)
2.0+0.2

(1.6-2.4)

STR

3.9414
(1.0-5.0)
5.0+0.0
(5.0)
28+1.2
(1.0-5.0)
1.3+0.7
(1.0-4.0)
3.1+1.0
(1.0-4.5)
5.0+0.0
(5.0)
5.0+0.1

(4.5-5.0)

3.8+1.0
(2.0-5.0)
45+0.4
(4.0-5.0)
25+08
(1.5-4.0)
1.4+0.7
(1.0-3.0)
27415
(1.0-4.0)
4.9+03
(3.0-5.0)
49+03

(4.0-5.0)

Rl

1.10+0.10
(0.94-1.32)
1.04 + 0.06
(0.93-1.17)
1.10 + 0.08
(0.98-1.23)
1.04 + 0.09
(0.76-1.24)
1.09 + 0.06
(0.98-1.18)
1.11 + 0.07
(0.99-1.21)
1.08 + 0.10

(0.91-1.26)

0.88+0.11
(0.75-1.20)
0.86 + 0.07
(0.72-0.95)
0.82 0.06
(0.70-0.90)
0.79 + 0.09
(0.60-0.97)
0.79 + 0.05
(0.74-0.84)
0.79 + 0.08
(0.59-0.96)
0.82 + 0.08

(0.69-0.98)

R2

0.94+0.21
(0.66-1.45)
0.93 + 0.17
(0.66-1.30)
0.95 + 0.20
(0.61-1.31)
0.85 +0.17
(0.45-1.09)
0.99 + 0.10
(0.84-1.12)
0.92 +0.14
(0.59-1.25)
0.92+0.17

(0.64-1.26)

1.08 + 0.14
(0.91-1.46)
1.05 +.0.09
(0.93-1.18)
1.13 £0.07
(0.99-1.25)
1.04 + 0.07
(0.88-1.16)
1.15+0.11
(1.05-1.27)
1.05 +0.10
(0.89-1.27)
1.09 + 0.09

(0.94-1.23)

R3

0.93 0.10
(0.81-1.16)
0.93 + 0.07
(0.82-1.07)
0.85 + 0.10
(0.68-1.00)
0.86 + 0.08
(0.75-1.00)
0.83 + 0.04
(0.78-0.89)
0.83 + 0.07
(0.67-0.96)
0.83 + 0.08

(0.62-0.95)

0.83 + 0.16
(0.56-1.15)
0.80 + 0.17
(0.57-1.08)
0.86 +0.14
(0.66-1.07)
0.83 + 0.13
(0.58-1.07)
0.77 £0.11
(0.69-0.90)
0.84 + 0.18
(0.49-1.19)
0.85 £0.15

(0.57-1.10)

R4

0.79 + 0.08
(0.66-0.98)
0.80 + 0.09
(0.66-1.00)
0.77 + 0.07
(0.65-0.90)
0.76 + 0.08
(0.53-0.96)
0.81 £0.07
(0.71-0.91)
0.80 + 0.09
(0.56-0.99)
0.78 + 0.06

(0.67-0.89)

0.79 + 0.08
(0.67-0.95)
0.76 + 0.06
(0.66-0.85)
0.77 + 0.07
(0.65-0.88)
0.74 + 0.07
(0.59-0.88)
0.77 £0.04
(0.73-0.81)
0.78 + 0.08
(0.62-1.03)
0.76 + 0.06

(0.69-0.84)

RS

0.27+0.01
(0.26-0.28)
9.27+0.01
(0.25-0.29)
0.27+0.01
(0.25-0.29)
0.26 + 0.01
(0.22-0.28)
0.26 + 0.01
(0.25-0.26)
0.26+0.01
(0.24-0.28)
0.26 + 0.01

(0.25-0.27)

0.25 + 0.01
(0.24-0.27)
0.25+0.01
(0.24-0.26)
0.24+0.01
(0.23-0.26)
0.24+0.01
(0.22-0.26)
0.24+0.01
(0.23-0.25)
0.24+0.01
(0.22-0.27)
0.24+0.01

(0.22-0.26)

R6

0.18
(0.15-0.21)
0.19
(0.15-0.20)
0.18
(0.17-0.21)
0.19
(0.16-0.22)
0.19
(0.18-0.20)
0.19
(0.16-0.22)
0.19

(0.17-0.21)

0.17
(0.15-0.21)
0.18
(0.15-0.20)
0.17
(0.15-0.19)
0.18
(0.15-0.20)
0.19
(0.17-0.22)
0.18
(0.15-0.21)
0.17

(0.15-0.18)

R7

0.40
(0.36-0.44)
0.39
(0.35-0.42)
0.39
(0.35-0.45)
0.40
(0.35-0.45)
0.39
(0.36-0.43)
0.39
(0.33-0.44)
0.40

(0.36-0.44)

0.39
(0.35-0.42)
0.36
(0.32-0.40)
0.39
(0.34-0.42)
0.37
(0.32-0.42)
0.38
(0.38-0.39)
0.37
(0.33-0.41)
0.38

(0.34-0.42)

320 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 6

600 PHYLOGENY.— The _ phylogenetic
analysis compared areas A, B, D, F and G
using 7. a. hydrophilus (Shasta County

“ie “a populations) as an outgroup. The parsi-
mony analysis resulted in a single shortest
400 tree of length = 103, CI = 0.848 (Fig. 5 ).
Southern populations of 7. atratus formed
an two main branches leading to areas A and

D, and to a clade containing areas (G (B,

F)). Character support and values for

200 branches are provided in Appendix C. The

following characters were homoplasious:

tae relative 6th SL height, internasal-rostral

100 200 300 400 500 600 700. g00 contact, relative tail length, posterior/ante-

rior frontal width, ventral width of 6th/7th
supralabial, and ground color.

FiGuRE 3. Relationship of male vs. female snout-vent length for TEST OF CONVERGENCE. — The non-
Thamnophis atratus. Individual male and female SVL are contrasted convergence tree (minus 7. e. terrestris)
in decreasing order of SVL beginning with largest male and female;
diagonal line represents a 1:1 ratio.

Snout-vent Length MALES -- MM

Snout-vent Length FEMALES -- MM

was best supported (length=41, Cl=
0.800), while the convergence hypothesis
tree (minus 7. a. hydrophilus) obtained
only modest support (length = 43, CI = 0.571). The non-convergence phylogeny supported the 7. a.
atratus-T. a. hydrophilus sister-taxa relationship by relative internasal width, pupil diameter and tail
length, and male/female snout-vent length. The convergence phylogeny supported a 7. elegans-T. a.
atratus relationship by relative interocular width, stripe value, and ground color. Character conver-
gence between southern Thamnophis atratus and T. elegans appears to be based only on color pattern,
a shorter muzzle and greater relative interocular width as compared to 7. a. hydrophilus.

DISCUSSION

SEXUAL DIFFERENCES. — Sexual differences in T. atratus are based on characters of proportion
and ventral-subcaudal counts. Smaller eyes and taller supralabials in females (p<0.001) may be the
result of ontogenetic scaling and larger female body size. In large adults, eye diameter increases with
respect to gape in males (r = 0.88) and decreases with respect to gape in females (r = -0.93).

Males of the 7. couchii complex average 82% of female snout-vent length (SVL) (Fitch 1984).
Fitch (1981) found the ratio of male-female SVL to increase from aquatic (73%) to terrestrial (88%)
Thamnophis. In southern T. atratus, values are within the range of “terrestrial” snakes (83-96%, mean
89%), as is the value for 7. a. hydrophilus (87%), while the value for 7. elegans (77%) approaches
the “aquatic” value. The male-female SVL ratio for 7. atratus appears to be correlated with mean
female SVL: (r = 0.91, F = 18.8, p<0.01), but not with male SVL (r = 0.60, F = 2.3, p<0.2). This ratio
can be explained by the differential growth rates of adults, in which females continue to increase SVL
at a greater rate than males (r = 0.99; Fig. 2). The conflict between these and Fitch’s results may be
due to a complexity of parameters affecting garter snake size that defy a “terrestrial” and “aquatic”
classification scheme.

LENGTH. — Snakes from the coastal fog belt average smaller than those occurring just outside
of this zone. Of 20 adult snakes from Dillon Beach, Marin County, only 3 (15%) exceeded the mean
SVL for area A. Fox (1951) suggested that the coastal snakes grew less rapidly than inland snakes,
causing them to reach maturity at a smaller size. Snakes from near the coast in Areas C and D are also
smaller, but less markedly so, than the means for their entire areas.

BOUNDY: SYSTEMATICS OF THAMNOPHIS ATRATUS 321

TABLE 4. Character weights and factor loadings for canonical plots between Areas for male and female
Thamnophis a. stratus. Only significant values contributing to canonical separation of factors are shown.
Abbreviations as in Table 1, except AVSW = anterior vertebral stripe width, INL/INW = internasal length/in-
ternasal width, MSR = midbody scale rows.

Males Females

Weight Fl F2 F3 Weight Fl F2 F3 F4
ACS/PCS n. S. n.s
ASR n.s 0.140 0.675
AVSW 0.247 0.225
ED 0.258 0.445 0.199 0.408
INL/INW n.s 0.121
INR/NR 0.201 0.427 nes
MSR 0.179 0.117
MVSW 0.241 0.214 0.320 0.199 0.437
PSR 0.255 0.219 0.138
SC 0.240 0.408 0.304 0.677
SLW/H 0.111 0.128 0.309
STR 0.794 0.838 0.306 0.837 0.770 0.435
V 0.457 0.286 0.728 0.419 0.235 0.563

EYE MORPHOLOGY. — Fitch defined his elegans group as possessing relatively smaller eyes,
larger pupils and darker irises than the Aydrophilus group. However, neither Bellemin and Stewart
(1977) nor Rossman (pers. comm.) found significant differences in eye diameter between T. elegans
and 7. atratus, and no significant difference in relative eye diameter was found between southern 7.
atratus populations. Small pupils in highly aquatic Thamnophis appear to be an adaptation to increase
underwater visual acuity (Schaeffel and de Queiroz 1990). Aquatic Thamnophis also possess pale
irises, whereas diurnal, terrestrial snakes such as Coluber, Masticophis, and Thamnophis elegans,
possess dark irises (pers. obs.), perhaps to reduce sun glare. Iris color follows a north-south cline
towards darker irises in 7. atratus, and Fox (1951) characterized T. a. atratus as having a darker iris
than 7. a. aquaticus. However, southern 7. atratus retain a pupil size characteristic of highly aquatic
snakes.

HEAD SHAPE. — Fitch (1940) characterized the hydrophilus group as possessing a relatively
pointed muzzle and shorter posterior supralabials than snakes of the e/egans group, and Fox (1951)
found T. elegans to possess a shorter gape relative to head length. Within southern 7. atratus, no
identifiable linear clines were found for supralabial height, relative internasal width, or internasal-
rostral contact. These snakes retain the relatively short posterior supralabials and pointed internasals
of aquatic snakes, but possess a wider snout characteristic of 7. elegans (Table 1). Southern 7. atratus
also retain the longer gape of the Aydrophilus group. Gape length is determined by length of
supratemporals, quadrates and mandibles (Varkey 1979).

COLOR PATTERN. — The presence of a predominantly spotted pattern in northern 7. atratus is
limited to populations north of the present study area, except near the mouth of the Russian River,
based on examination of MVZ and CAS specimens. Thus, all snakes utilized here are relatively dark
with bright stripes. Variation in stripe value converges on Area D: as the Santa Cruz Mtns. are
approached, vertebral stripe width and ventral darkening increase, and lateral stripe contrast decreases
(Fig. 1f). Parallels to this pattern occur in the Puget Sound region of Washington, where the three
species of Thamnophis occurring there exhibit increased melanism (Stebbins 1985), which is the
diagnostic character for two subspecies: 7. elegans nigrescens and T. sirtalis pickeringii. Two T.
atratus from coastal Sonoma County (MVZ 48013 and 48016) show ventral and lateral melanism,

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BOUNDY: SYSTEMATICS OF THAMNOPHIS ATRATUS 323

TABLE 5. Predicted vs. actual results of group membership in a discriminant function analysis of
Thamnophis atratus from Areas A-G.

Area | A B Cc D E F G Total

A | 21 3) 3 2) 28 A
B | 20 4 24 C
C 21 4 9) 2 30 T
D 7 57 64

E 1 10 11 "
F eae 8 | 34 7 52

G | 3 21 24 L
Total 28 30 31 62 13 43 30

PARSE De ED,

although the narrow vertebral stripe and cephalic proportions in these specimens are indicative of T-
a. hydrophilus. Other snakes from this area possess a paler, three-striped pattern.

Fox (1951) found Area A-B snakes to have a gray ground color, pale yellow vertebral stripe, and
salmon midventral suffusion (Fig. 1d). Area D—G snakes have abundant black suffusion on the dark
fields, a bright yellow to yellow-orange vertebral stripe, and yellow or yellow-orange midventral
suffusion (Fig. 1f-h). The latter two characters fade in preservative and were not quantified for the
present study. Area F snakes show an overall increase in the amount of yellow in the pattern,
paralleling another Diablo Range endemic, Masticophis lateralis euryxanthus (Riemer 1954), and
Thamnophis sirtalis fitchi in the southern Diablo and Santa Lucia ranges (pers. obs.).

CANONICAL SEPARATION AND PHYLOGENETIC IMPLICATIONS. — Specimens from Area D were
the most significantly differentiated from other Areas (mean Chi-square = 375.6; Factor 1). Area D
snakes possess a unique color pattern, which all young of several broods from this Area displayed.
The light, three-striped pattern is omnipresent in all broods examined from Areas A, B, and F, and in
broods from Areas C and E the patterns are mixed. Because of their unique color pattern, combination
of other character extremes, and phylogenetic position, restriction of the name Thamnophis a. atratus
to Area D is warranted (see Taxonomic Synopsis).

Despite the numerous characters that distinguish between populations north and south of San
Francisco Bay, the branching pattern of the phylogenetic tree indicates that Areas A and B are
divergent. This is supported by Factor 3 for males and Factor 2 for females, which separate Areas A
from B and Areas F from G. Thus, 7. a. aquaticus is polyphyletic.

The branching pattern of the phylogenetic tree indicates divergence between (A, D) and ((B, F)
G). Factor 3 of males and Factor 2 of females reflect this pattern, while factor 2 of males and factor
3 of females reflects Fox’s (1951) arrangement of T. a. atratus and T. a. aquaticus. Because Area A
and B snakes are geographically proximal to the outgroup (ancestral taxon), they hypothetically
represent basal populations to the clades formed by node 2, and are not as readily distinguishable
from each other as the more geographically distant populations (D and F,G). The phylogenetic tree
can alternately be viewed as containing 2 ([A, D] and [B, F, G]) or 3 (A, D, and [B,F, G]) terminal
taxa. The latter alternative would require recognition of 7. a. aquaticus in Area A, creating an
intermediate taxon that would be difficult to diagnose. The two-taxon alternative corresponds to a
bifurcation of coastal and inland populations near the southern end of the range of 7. a. hydrophilus.
This latter alternative is utilized here to avoid taxonomic recognition for populations that would be
based on intermediate characters within a north-south cline.

Populations composing both clades have evidently diverged on separate trajectories from T. a.
hydrophilus. The coastal clade includes 7. a. atratus in the restricted sense, while the inland clade

324 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 6

represents a distinct taxon described be-

A low. The populations in areas A and B,

formerly called 7. a. aquaticus, are here

5 referred to as “basal-southern” 7. atratus.

INTERGRADES. — Within the range of
southern 7. atratus are several areas of
morphological intergradation. Areas A and
C are composed of snakes intermediate in
color and pattern between the subspecies
QD T. a. hydrophilus and T. a. atratus (Fig.

Bp !e). The change is gradual from the Marin

Peninsula to the San Andreas Rift Lakes

(Fig. 6), although some snakes from Half

4 Moon Bay to Pescadero have faint lateral

stripes. This appears to be clinal intergra-

F dation, in which characters have shifted

3 along a geographic gradient from one ex-

treme to another. The ends of these ex-

tremes are homogeneous (at least

morphologically) groups of populations.

G Snakes from northwestern Sonoma

County have high ventral counts, occa-

sionally lack lateral stripes, and may have

dark spots in the dark fields. Although 7.

a. hydrophilus may be striped and/or spot-

H ted, the spotted pattern is unique to that

FiGurE 5. Phylogenetic relationships of five population groups of Subspecies, and is seen in snakes as far

Thamnophis atratus (Areas A, B, D, F, G) using T. a. hydrophilus (H) south as the Gualala River on the coast and

as the outgroup; length = 103, CI = 0.848; numbers identify nodes Cache Creek, Colusa County, inland. Fox
referred to in the text. : 3 :

(1951) considered intergradation between
the subspecies 7. a. hydrophilus and “T. a. aquaticus” to be abrupt, both morphologically and
geographically, ocurring at the Gualala River.

Fox (1951) stated, “the transition from aquaticus to atratus [striped morph] is gradual, . . . [have
arbitrarily divided the two races at the San Francisco Bay.” Fox’s concept of the subspecies T. a.
aquaticus included the variable populations ranging north to Trinity County. He cited the four Solano
County specimens as being intergrades because of enlarged supralabials and intermediate color. The
large size of the supralabials is due to the fact that two are huge females (with enlarged salivary
glands). One male has supralabials in proportion with other Area B snakes, and those of the other
could not be measured. All have the gray ground color typical of “7. a. aquaticus” but vertebral stripes
similar to the striped morph of 7. a. atratus. Snakes from Contra Costa County are typical of the
striped form of 7. a. atratus. Unfortunately the subtle color differences between “T. a. aquaticus” and
striped 7. a. atratus are not evident in preserved specimens, and no fresh specimens from Contra
Costa County were available for study. MVZ 48975, from Wildcat Canyon, Contra Costa County,
has a gray dorsum with a narrow vertebral stripe as in 7. a. aquaticus. However, two others collected
with this individual lack these characteristics (pers. obs.). No snakes were available from the
Carquinez Straits region between Suisun Creek and the Berkeley Hills. The southernmost area of
intergradation between T. a. hydrophilus and southern T. atratus remains unresolved as to whether it
is abrupt or gradual. A genetic study is needed to determine the nature of contact at points of parapatry.

BOUNDY: SYSTEMATICS OF THAMNOPHIS ATRATUS

Ww
i)
Nn

Area E contains a mixture of snakes bearing character extremes, and others which are interme-
diate. The dark form of 7. a. atratus occurs south to Uvas Canyon, while striped snakes overlap the
dark form north to the Los Gatos Creek drainage (Fig. 6). At Uvas Canyon and upper Guadalupe
Creek, both morphs and intergrades occur sympatrically. Of 26 snakes from Area E and Guadalupe
Creek, 38% are dark, 19% are striped, and 43% are intermediate. All snakes in the open oak woodland
below Uvas Canyon and south of the Santa Teresa Hills are the striped morph, and are identical in
color pattern to snakes east of the Santa Clara Valley. This distribution of color morphs and
intermediates suggests secondary contact in Area E (see Biogeography below).

The southwestern side of the Santa Cruz Mountains, from Soquel Creek east, contains popula-
tions of intergrade snakes having faint lateral stripes and slightly darkened venters (Fig. 1g). It is
possible that these snakes resulted from initial contact when the striped form reached the south end
of the Santa Cruz Mountains (see below). The intergrade zone may have broadened, creating a buffer
from constant influence from the striped form. Also, the heavily forested summit may prevent the
striped form from remaining in contact with most of these populations. There is some gradation, with
snakes from the geographical extremes (Soquel Creek and Hecker Pass) increasingly resembling the
dark and striped forms, respectively.

ECOLOGY. — At the northern end of the range of Thamnophis atratus, the subspecies T. a.
hydrophilus is highly aquatic (Fitch 1940), while snakes from the San Francisco Bay region are
semiterrestrial (Fox 1951, pers. obs.). This ecological shift is thought to be the result of change in
habitat and, possibly, competition with other species of Thamnophis.

Southwestern Oregon and northern California have an abundance of large rivers, and large
museum series of 7. atratus collected on single dates along these rivers indicate their abundance there.
The rivers are typically lotic all year, with rock, sand, or gravel beds, and snakes rarely venture far
from these or smaller bodies of water (Fitch 1940). There are few large rivers or streams in the San
Francisco Bay region and most are intermittent. Here, 7. atratus has adapted to a semiterrestrial life
style, and acquired some morphological characteristics of Fitch’s (1940) elegans group. South of the
Pajaro River drainage large rivers are again prevalent and, except on the outer coast, 7. atratus has
retained an aquatic life style (S. Sweet, pers. comm.). The morphological differences between
southern T. atratus and T. a. hydrophilus may be related to these differences in habitat.

Much of the range of 7. a. hydrophilus (southwestern Oregon and northern California) is forested.
Southward and inland, canopy vegetation is patchy in oak savannah-woodland, and is often restricted
to riparian strips. This is reflected in an increased coastward restriction of 7. atratus distribution from
north to south. In conjunction, the drab, spotted morph of T. a. hydrophilus has been lost in southern
T. atratus. The drab, spotted pattern provides a camouflage advantage in open, gravel-bottomed
streams (Fitch 1940). Snakes moving on the bottom of these streams are nearly invisible, and I have
lost sight of snakes, even though in plain view, as their pattern and sinuous movements blended with
the gravel and riffles. A boldly striped pattern might be detrimental to survival in such situations.
Thamnophis atratus in Coyote Creek were readily observed at distances >10 m due to their contrasting
pattern and bright colors. However, striping is advantageous to terrestrial snakes as both camouflage
in vegetation, and as an illusionary escape device. The dorsal stripe leaves the observer with a brief
after-image posterior to the retreating snake (Jackson et al. 1976), and I have found this to be effective
when trying to capture rapidly crawling snakes on land. Striping in snakes, possibly detrimental in
the water, may be offset by its terrestrial advantage, since seasonal drying of streams forces the snakes
to terrestriality during part of the active season. I believe that selection for a boldly striped pattern in
southern 7. atratus has positive survival value, and is related to the marked habitat differences between
northern and central California. However, 7. a. hvdrophilus exhibits an increase in number of striped
individuals inland, but does not show seasonal disuse of streams during the annual period of activity
(D. Rossman, pers. comm.).

326 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 6

FIGURE 6. Localities for Thamnophis atratus in the San Francisco Bay region. Circles darkened below represent basal
southern 7. atratus; open circles, T. a. zaxanthus; closed circles, T. a. atratus; circles darkened above, T. a. atratus x T. a.
zaxanthus intergrades.

BOUNDY: SYSTEMATICS OF THAMNOPHIS ATRATUS 327

Within Area D, both prey and understory vegetation are widely dispersed. In Area F, prey and
cover are restricted to riparian strips, which is reflected in habitat use by snakes: 52.5% (G = 0.718)
of Area D snakes were encountered within 20 m of water, compared to 88.2% (G = 10.352) of Area
F snakes. Increased cover and prey dispersal has enabled Area D snakes, and probably coastal Area
G snakes, to increase terrestriality over other 7. atratus. Area D snakes are less seasonally restricted
in activity, with 61.6% of snakes observed May-August, vs. 73.2% in Area F. May—August activity
is significantly greater than spring and fall activity in Area F snakes (t = 2.447 at df = 6, p<0.05), but
not for Area D snakes. Snakes from the southern extreme of the range are active from April to July
only (S. Sweet, pers. comm.).

FOOD. — Quantitative feeding studies for southern 7. atratus are unavailable, and Fitch’s (1940,
1941) concept of 7. atratus included T. elegans terrestris. For the present study, 7. atratus were not
routinely checked for prey. Observations revealed only fish and amphibians, and feeding was
observed only in the water. The presence of the plethodontid salamanders Aneides lugubris, Ba-
trachoseps attenuatus, and Ensatina eschscholtzii in four snakes from Area D indicates some
terrestrial feeding (pers. obs.). Otherwise, there is no indication of similarity with the food habits of
T. elegans (Fox 1952; Gregory et al. 1980; James et al. 1983).

BIOGEOGRAPHY. — Geographic patterns of variation within the 7hamnophis couchii complex,
combined with paleogeographic and molecular data, suggest the following hypotheses for observed
variation within 7. atratus. Thamnophis atratus occasionally hybridizes with sympatric members of
the 7. couchii complex at either end of its range (Rossman and Stewart 1987), but is non-convergent
in most characters with 7. couchii and T. hammondii at their respective contact zones. The ranges of
T. atratus and T. hammondii overlap broadly, and hybridization may be the result of secondary contact
(S. Sweet, pers. comm.), suggesting that derivation of one from the other is unlikely. The ranges of
T. atratus and T. couchii are largely parapatric, creating a narrow zone of hybridization, which implies
derivation of one species from the other. In addition, paleogeographical evidence supports the notion
that 7. atratus originated in the north, and dispersed southward. Scenarios in Peabody and Savage
(1958), Howard (1962), Morafka and Banta (1972), and Yanev (1980), indicate that territory currently
occupied by southern 7. atratus was submerged until 5 million years ago (mya). A southward dispersal
route for 7. aftratus was supplied by continuing uplift of the Coast Ranges during the next 4 million
years.

A scenario for the derivation of morphotypes of 7. atratus can be developed by superimposing
their hypothetical past distribution on maps provided in Yanev (1980). Between 8 and 5 mya the
northern Coast Ranges uplifted southward from the present-day vicinity of Lake County, forming a
peninsula toward the present-day Santa Cruz Mountains. The interface created by this uplift runs from
near the mouth of the Gualala River eastward in the region in which Lake County borders Sonoma
and Napa counties. The Sonoma Gap, represented by a low grassland region interrupting the otherwise
continuous band of coastal forest, runs east-west approximately 60 km to the south of this interface.
A number of taxa have differentiated genetically and/or morphologically near the interface: Dicamp-
todon (Good 1989), Aneides lugubris (Sessions and Kezer 1987), Ensatina (Stebbins 1949: Wake
1997), Rana aurora (Green 1985), Diadophis amabilis (Blanchard 1942), and Thamnophis atratus
(Fox 1951). During this period, it is conceivable that 7. atratus, or its ancestor, moved southward
along the peninsula.

Subsequent uplift and contact with the Diablo Range in the north formed two peninsulas, one
extending southward through the Santa Cruz Mountains, the other southward to the vicinity of
present-day Pacheco Pass in the Diablo Range. Once 7. atratus moved south of the Sonoma Gap, two
morphotypes may have diverged in response to habitat differences between inner and outer coast.
Dark-sided snakes of the outer-coastal forests would have extended southward through the Santa Cruz
Mountains, while the three-striped inland morph would have extended southward in the Diablo Range
in savannah-riparian strip habitat (Fig. 7a). This scenario is supported by node 2 and the resulting

328 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 6

branches of the phylogenetic tree (Fig. 5). This pattern is paralleled by Ensatina, in which an
inner-outer coast divergence occurs immediately north of San Francisco Bay, following through to
the Santa Cruz Mountains-Diablo Range (Stebbins 1949).

As the Pajaro outlet for the Merced Sea closed about | mya, the inland morph may have extended
southward through the Santa Lucia Range to the Transverse Ranges south of the Monterey Peninsula
(Fig. 8). Thamnophis atratus is relatively common on the outer coast, but is scarce inland. This may
be due to competition with the highly aquatic 7. hammondii, perhaps already established in the region,
by increased restriction of riparian habitats, and seasonality of water and food supplies. Initial closure
of the sea at the Diablo Range would have permitted the inland morph to enter these areas in advance
of the Santa Cruz Mountains morph. Subsequent elimination of contact between inner and outer Coast
Range populations by the Salinas Valley would explain node 3 of Fig. 5. The eventual shift of the
Central Valley outlet to the San Francisco Bay coincides with nodes 4 and 5 of Fig. 5.

Closure of the Santa Cruz-Diablo Range division at the Santa Clara Valley-San Francisco Bay
would have allowed secondary contact between populations already differentiated (Fig. 7b). The two
morphotypes meet along the southeast slope of the Santa Cruz Mountains, where both intergradation
and sympatry occur. Other taxa following a similar pattern include the scorpion Uroctonus mordax
(Hjelle 1972) and Lampropeltis zonata (pers. obs.). By presence-absence, Pinus sabiniana also
demonstrates this pattern (R. Myatt, pers. comm.). The preceding are woodland species, none of which
currently occur at the proposed Santa Clara Valley crossing, suggesting that such a crossover would
have taken place during more mesic conditions.

TAXONOMIC SYNOPSIS

Thamnophis atratus zaxanthus subsp. nov.
Fig. th

HOLOTYPE. — MVZ 207940, an adult female from 3 mi S of Gilroy Hot Springs, Santa Clara
County, California, collected 15 April 1986 by William P. Hutchins.

DIAGNOSIS. — Maximum total length 902 mm; ventrals, males 148—168 (mean 158.3, n= 71),
females 140—167 (mean 152.1, n = 86): subcaudals, males 65—89 (mean 80.8, n = 61), females 59-83
(mean 72.5, n = 73); anterior scale rows 19 (82%) or 21 (18%); midbody scale rows 19 (85%) or 17
(15%); posterior scale rows 15 (78%) or 17 (22%); vertebral stripe relatively broad, averaging 3.2
(range 1.5—5.0) scale rows on the nape, 2.1 (1.0-2.8) rows in the thoracic region; vertebral stripe
yellow to orange-yellow; lateral stripe conspicuous, pale green; dorsum dark gray to black; iris gray
or brown; top of head dark, olive black, with a prominent parietal spot; supralabial suture marks
narrow when present; demarcation between dorsal head color and pale supralabials distinct; chin
cream, becoming bright yellow on the throat, grading to pale green in the thoracic region; ventral
color darkening slightly posteriorly; midventral suffusion yellow to orange; dark markings absent
from transverse ventral sutures; eye moderate in size (eye diameter/gape length 0.15—0.22, mean =
(0.19 in males, 0.18 in females); tail moderately long, averaging 26.2% (24-29%) of total length in
males, 24.4% (22-27%) of total length in females. Color photographs of adults are provided by
Boundy (1990, frontispiece, holotype in life), Mara (1994:4, mislabeled 7. sirtalis fitchi) and Rossman
etal. (1996; ple).

DESCRIPTION OF HOLOTYPE. — (from life); Adult female, 448 mm SVL, 128+ mm tail length
(approx. 7 mm missing from tip); ventrals 149, subcaudals 62+; scale rows 19-19-15, all but first row
keeled; supralabials 8-8; infralabials 9-9 (Sth and 6th fused on each side); preoculars 1-1; postoculars
3-3; temporals 1+2+2 on each side; posterior chin shields longest; intergenials 3; anal plate single;
head distinctly wider than neck; top of head flat; snout moderately blunt; eyes and nostrils with a
slight dorsolateral tilt; pupil approximately 1/2 eye diameter; iris dark gray; eye diameter/gape length

BOUNDY: SYSTEMATICS OF THAMNOPHIS ATRATUS 329

FiGurE 7 (A) Hypothetical distribution of southern Thamnophis atratus | million years ago; crosshatching represents 7° a.
atratus; area of widely spaced lines represents 7. a. zaxanthus; SF = present day location of San Francisco; MP = Monterey
Peninsula. (B) Hypothetical distribution of 7. atratus 0.5 million years ago.

0.17; top of head dark olive gray, becoming lighter at preoculars; parietal spot present; labials and
side of neck to about one head-length posterior of jaw angle bright yellow, slightly lighter yellow on
ventrals to about two head-lengths posterior of throat; chin yellow cream; narrow black sutures present
between all but first two supralabials; upper half of supralabials dusky yellow; venter pale gray green,
gradually becoming darker posteriorly, with midventral yellow suffusion; olive-gray ventrolateral
stripe on first scale row and tips of ventrals, beginning at the 11th ventral; small black markings on
anterior edges of first 3 dorsal scale rows; lateral stripe on 2nd and 3rd dorsal scale rows, light olive
gray on yellow, the yellow fading posteriorly; vertebral stripe orange yellow, 3 plus two half scale
rows wide at nape, narrowing to | and two 1/2 rows posteriorly; stripe becoming duller, two halfrows
wide, on tail to tip; ground color dark olive brown, heavily suffused with black, giving an overall
black appearance (Fig. Ih).

VARIATION. — Juveniles are identical in coloration to adults, although they often have brighter,
more distinct markings. Intrapopulation variation is limited to shade and intensity of stripe color.
Vertebral stripes may be pale yellow to yellow orange. Lateral stripes have varying amounts of olive
gray overlaying the yellow, creating a range of pale gray-yellow to bright yellow stripes. Midventral
suffusion is yellow, orange or pale salmon.

Interpopulation variation between Diablo Range and Santa Lucia Range snakes consists of the
latter having lower ventral and subcaudal counts, a narrower vertebral stripe, and fewer individuals
with 20 or 21 anterior scale rows. Diablo Range snakes differ from Napa and Solano county snakes
in having a broader vertebral stripe, lower posterior scale row and subcaudal counts, and taller
subralabials. Intergrades with T. a. atratus exhibit loss of the lateral stripes to a varying degree, with
increased dark olive coloration on supralabials, lower sides, and venter (Fig. le).

330 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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DIAGNOSTIC COMPARISONS. — Thamnophis a. zaxanthus differs from other central California
Thamnophis as follows: from T. sirtalis it differs in having 8 rather than 7 supralabials, in lacking a
red head and red in the dark fields, and in having a yellow or orange midventral suffusion; from 7.
elegans it differs by always lacking red pigment, and by having a yellow midventral suffusion, the
posterior chin shields noticeably longer than the anterior, narrower contact between the internasals
and the rostral, the internasals usually longer than wide, the anterior scale rows more often 19 than
21, the posterior scale rows more often 15 than 17, and a smaller pupil relative to eye diameter
(Bellemin and Stewart 1977 provide a statistical analysis of these differences); from 7. gigas and
northern populations of 7. atratus hydrophilus it differs by usually having 19 (80%) rather than 21
or more anterior scale rows, an orange midventral suffusion, bright yellow supralabials and throat,
and a vertebral stripe 2 or more scale rows wide. Thamnophis a. zaxanthus also differs from T. a.
hydrophilus in lacking dark markings on the transverse ventral edges, in always lacking distinct dark
spots on the dorsum, and in having a darker iris; and from 7. a. atratus by having lateral stripes, a
strong demarcation between the dorsal head and pale supralabial color, and a lighter venter.

ETYMOLOGY. — Greek, za (intensive), and xanthus (yellow).

DISTRIBUTION. — Inner Coast Range from Napa and Solano to Santa Barbara counties, and the
Santa Lucia Range (Fig. 6). It occurs from hills south of the Carquinez Straits to Tres Pinos Creek in
the Diablo Range, and to the Pinnacles region in the Gabilan Mountains. It occurs on the coastal slope
of the Santa Lucia Range from the Monterey Peninsula south to the San Rafael Mountains, Santa
Barbara County (La Brea Creek and Birabent Canyon; S. Sweet, pers. comm.). It also occurs in the
foothills along the southeastern slope of the Santa Cruz Mountains, south of the Santa Teresa Hills.
It is absent from major valleys.

Thamnophis atratus atratus (Kennicott 1860)
Fig. lf

LECTOTYPE. — USNM 970a, designated by Fitch (1940:89), from San Francisco (Cochran
1961:181). Descriptions of the cotypes in Kennicott (1860:296) and Cope (1891:658), and examina-
tion of lectotype, allow correction of the type locality to Area D, that is, the Santa Cruz Mountains
or southern San Francisco Peninsula. The lectotype is an adult male (486 mm SVL) with the following
characters: tail/total length 0.260; supralabials 8-8; infralabials 10-10; oculars 1-3/1-3; temporals
1+1+2, 1+2+2:; scale rows 19-19-15; pupil/eye diameter 0.30; eye diameter/gape length 0.19;
interorbital width/gape length 0.38; posterior/anterior frontal width 0.76; frontal/parietal length 0.89;
muzzle/frontal length 0.73; internasal length/width 0.43; prefrontal/internasal length 0.79; internasal-
rostral/nasal-rostral 0.80; loreal dorsal/ventral length 0.80; 6th/7th SL angle 3.5; anterior/posterior
chin shield length 0.92; vertebral stripe width 2.9; coloration is typical of Area D snakes.

DIAGNOSIS. — Maximum total length 1016 mm; ventrals, males 146—165 (mean 154.2, n = 57),
females 141—155 (mean 147.0, n = 57); subcaudals, males 67-89 (mean 79.4, n = 49), females 64-79
(mean 70.3, n= 48); anterior scale rows 19, rarely 17 or 21; midbody scale rows 19 (65%) or 17
(35%); posterior scale rows 15, rarely 13 or 17; vertebral stripe yellow to orange yellow, and broad,
averaging 4.0 (range 2.76.5) scale rows in the nuchal area, 2.5 (1.64.6) rows in the thoracic region;
lateral stripes absent; dorsum olive black, grading to dark olive at the ventrals; dorsal black spots
obscure; iris dark brown; top of head dark olive brown to olive black, with a prominent parietal spot;
supralabial suture marks narrow when present; demarcation between dorsal head color and dusky
olive supralabials indistinct; chin cream, becoming deep yellow on the throat; venter abruptly
becoming olive gray in the thoracic area, continuously darkening posteriorly; prominent yellow-or-
ange midventral suffusion; dark markings absent from transverse ventral sutures; eye moderate in
diameter (eye diameter/gape length 0.15—0.22, mean = 0.19 in males, 0.18 in females); tail moderately

BOUNDY: SYSTEMATICS OF THAMNOPHIS ATRATUS B51

s

»
15 |

FiGuRE 8. Location of localities and geographic features mentioned in the text: 1) Alamitos Creek; 2) Coyote Creek; 3)
Guadalupe Creek; 4) Half Moon Bay; 5) Los Gatos Creek; 6) Pacheco Pass; 7) Pajaro River; 8) Pescadero Creek; 9) San
Andreas Lake; 10) San Lorenzo River; 11) Santa Teresa Hills; 12) Soquel Creek; 13) Suisun Creek; 14) Uvas Creek; 15)
Wildcat Canyon.

332 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 6

long, 26.2% (0.22—0.28%) of total length in males, 24.1% (0.22—0.26%) of total length in females.
Color photographs of adults are provided by Mara (1994:13) and Rossman et al. (1996, pl. 1).

DISTRIBUTION. — Santa Cruz Mountains and southern San Francisco Peninsula, from the San
Andreas rift lakes to the San Lorenzo River watershed and Uvas Canyon (Fig. 6).

Thamnophis atratus hydrophilus Fitch 1936
Fig. la—c

HOLOTYPE. — MVZ 18127, from Trail Creek, 6 mi from its mouth, Jackson County, Oregon.

DIAGNOSIS. — Maximum total length 916 mm; ventrals, males 151-171 (mean 163.6, n= 77),
females 148—168 (mean 159.0, n = 71); subcaudals, males 77—95 (mean 87.5, n = 62), females 65—82
(mean 76.3, n = 60); anterior scale rows 21 (73%; northern Humboldt County northward) or 19 (98%;
southern Humboldt County southward); midbody scale rows 19 (84%) or 21 (16%); posterior scale
rows 17 (46%) or 15 (54%); vertebral stripe narrow, less than | and two half scale rows wide,
occasionally absent; vertebral stripe cream to orange yellow; lateral stripes absent to prominent;
dorsum olive gray to dark olive, with alternating darks spots distinct except in individuals having dark
ground color; top of head olive to dark brown, with a distinct parietal spot; paired, dark neck blotches
often distinct; supralabial suture marks present, of variable width; demarcation between dorsal head
and pale or dusky supralabial color distinct to only slightly so; venter light dusky, with a pale
midventral orange to yellow wash; dark markings present on transverse ventral sutures; eye diameter
moderate (eye diameter/gape length 0.16—0.22, mean = 0.19 in males and females); tail relatively
long, 26.8% (24.1—28.2%) of total length in males, 25.0% (22.6—-26.8%) of total length in females.
Color illustrations of adults are provided by Smith and Brodie (1982:151) and Rossman et al. (1996,
ple b):

DISTRIBUTION. — Northern Coast Ranges of California and southwestern Oregon north of the
Gualala River and Lake County, California.

SUMMARY

Southern populations of Thamnophis atratus are well-differentiated from northern populations
(= T. a. hydrophilus), and differences appear abruptly along a zone running from near the mouth of
the Gualala River eastward to the ridge separating Lake and Napa counties. The southern populations,
formerly thought to be separable into subspecies north and south of San Francisco Bay, are found
instead to have differentiated on two north-south trajectories (coastal and inland). Populations of 7.
atratus immediately north of San Francisco Bay exhibit little differentiation between coastal and
inland populations, but there is increasing divergence in characters between coastal and inland
populations south of the Bay. Populations south of the Bay appear to have come into secondary contact
at the southeastern part of the Santa Cruz Mountains, where phenotypic hybridization occurs.
Elsewhere, the two morphotypes are geographically separated.

The phylogenetic tree produced by the present study, coupled with hypothetical Pleistocene
geography of the central California coast, recommends a revised taxonomic arrangement of southern
T. atratus populations. The north Bay populations, formerly called 7. a. aquaticus, are morphologi-
cally and phylogenetically basal to south Bay populations. Because they lack morphological and
phylogenetic uniqueness, they cannot be considered terminal taxa and are not taxonomically recog-
nizaeable. The south Bay populations have evolved on two separate trajectories from north Bay
populations. Populations comprizing the two trajectories are morphologically and phylogenetically
unique, and are therefore given taxonomic status. The coastal trajectory represents Thamnophis
atratus atratus in a restricted sense, while the inland trajectory is named T. a. zaxanthus.

BOUNDY: SYSTEMATICS OF THAMNOPHIS ATRATUS 333

ACKNOWLEDGMENTS

For loans/lab space I thank Fred Andoli (CPS), Thomas Balgooyen (SJSU), Robert Drewes and
Jens Vindum (CAS), Harry Greene (MVZ), Douglas Rossman (LSUMZ), Samuel Sweet and Mark
Holmgren (UCSB) and George Zug (USNM). For statistical assistance I thank Charles Bell and
William Bros. For helpful comments and criticisms I thank Alan de Queiroz, Robert Hassur, Vida
Kenk, Robin Lawson, Rod Myatt, Douglas Rossman, Joseph Slowinski, Samuel Sweet, and four
anonymous reviewers. For preparing photographic prints of the figures I thank Ron Bouchard. Thanks
are extended to Andy Ford, Tom Heath, Owen Holt, Bill Hutchins, Steve Laymon, Bill Meyers, Tony
Rice, Blair Wolf, Bernie Wone, and my family for field assistance, and to John Boundy for formatting
the manuscript.

APPENDIX A

Specimens examined for MANOVA (all California)——T7hamnophis atratus aquaticus (132).
Marin Co.: MVZ 18196, 18198, 36184, 40708, 47953, 47957-47966, 47969-47995, 48196 (holotype
of Thamnophis elegans aquaticus), 172251, 206938. Sonoma Co.: CAS 27939, 28019-28021, 28024,
28025, 28029, 149676, 156208; CAS-SU 4219, 4229-4232, 4256-4258, 4323, 10741-10743,
11038-11041, 11044; MVZ 4913, 4914, 48013, 48015, 48016, 48042, 69691-69693, 69695-69697,
125414.

Thamnophis atratus atratus - T. a. aquaticus intergrades (64). Marin Co.: CAS 93771; MVZ
47954-47956, 47967, 47969, 68194. San Francisco Co.: MVZ 19118-19120, 19122, 19123,
19125-19128, 19133, 19134, 37138. San Mateo Co.: CAS-SU 8157, 10706, 10712; MVZ
47883-47893, 47896-47925, 93805.

Thamnophis atratus atratus (187). “San Francisco”: USNM 970a (lectotype of Eutainia atrata
and Eutaenia infernalis vidua). San Mateo Co.: CAS 149585—149617, 149677, 191545, 191640,
191641, 191646; CAS-SU 1198, 1200-1205, 1654, 1655, 4149, 4155, 5180, 5183, 5184, 8105,
8109-8112, 10686-10688, 10691, 10692, 10694-10696, 10698, 10827, 10828, 10831, 10833, 10834,
10897-10899, 13781, 17867, 17906, 17907, 22949-22952, 25567, 25568; MVZ 9316, 14003, 16592,
32628, 47882, 47894, 47895, 4792647928, 92508, 187632—187636, 187654, 187655. Santa Clara
CorsONS 542255 191576; 191578, IOIS8I=N915839 191596; 191598, 191599" 1916025 19G09:
M9163, 191617, 191620, 191622, 191624, 191625, 191633, 191635; CAS-SU 4157, 4322,
6378-6380, 8207, 8208, 13780, 18160-18162; Santa Cruz Co.: CAS 81547, 191557,
191560—191563, 191565, 191567—191569; CAS-SU 4150-4153, 7636, 7637, 10817; MVZ 26988,
35545-35553, 39443, 47935-47941, 4897648979, 62106-62118, 100280, 149624; SJSU R4737.

Thamnophis atratus atratus - T. a. zaxanthus intergrades (25). Santa Clara Co.: CAS 191584,
191587; 1915917191595, 191597, 191600; 191603, 191621, 191623, 1911626, 191634, 191636,
191642, 191645, 191653. Santa Cruz Co.: CAS 191558, 191559, 191564, 191566, 191570—191572;
CAS-SU 1675, 1679, 4148.

Thamnophis atratus zaxanthus (169). Alameda Co.: CAS-SU 4161, 11913, 11914; MVZ 1629,
2457, 3757, 10524, 12857, 24897, 24898, 24905, 24906, 25200, 32244, 38951, 38952, 47943-47948,
48538, 111370, 187622—187626, 187649, 191323. Contra Costa Co.: CAS 43695; MVZ 4009, 19139,
32885, 47949-47952, 48973-48975, 85208, 85209, 111362. Merced Co.: MVZ 39350, 39351.
Monterey Co.: CAS 13764, 13765, 149553, 149554, 191541; CAS-SU 1685, 5143, 5147, 5149,
10327-10329, 11316; CPS 248; MVZ 7243, 26010, 44865, 66382, 125415. Napa Co.: CAS 191544;
CAS-SU 6134, 6310-6314; MVZ 42639-42644, 47887, 47888, 47999-48008, 48010-48012, 60234,
60235, 78049-78052; 78817, 85214, 149606—149609, 14962 1—149623, 149625, 187630. San Benito
Co.: CAS 49212, 191537—191540; MVZ 83647. San Joaquin Co.: CAS 191542; MVZ 28206, 28207.
San Luis Obispo Co.: CPS 120, 188, 189, 191, 2420, 2426, 2428, 2436, 2437, 3216-3220, 3224,

334 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 6

3225, 3227, 3228, plus two unnumbered specimens; UCSB 15552, 21997. Santa Barbara Co.: CPS
94; UCSB 9078, 9081-9083, 13707, 14213, 17805, 17584, 18762, 22203. Santa Clara Co.: CAS
39652, 39653, 41661, 41662, CAS-SU 4091, 5852, 6520, 22953, 22954, 191574, 191575, 191577,
191579, 191580, 191585, 191586, 191588, 191590, 191592-191594, 191604191608,
191610191612, 191614191616, 191618, 191619, 191628—-191632, 191637—191639, 191643,
191644, 191647—191652, 195846; MVZ 79576, 207940 (holotype of Thamnophis atratus zaxan-
thus); Solano Co.: MVZ 4006-4008, 48017. Stanislaus Co.: CAS 161384, 191547—-191551; MVZ
39349, 78816.

APPENDIX B

Supplemental specimens examined.— Thamnophis atratus aquaticus (16). Marin Co.: LSUMZ
8222-8225, 22205-22208, 34238, 34239. Sonoma Co.: LSUMZ 16531, 16638, 16644, 16648, 16649,
39069.

Thamnophis atratus atratus (20). San Mateo Co.: LSUMZ 16546, 24352-24356, 2891 128914,
2893 128933, 36936, 37905, 37907, 37908, 39599, 39600. Santa Cruz Co.: LSUMZ 16545.

Thamnophis atratus atratus - T. a. aquaticus intergrades (4). San Mateo Co.: LSUMZ 8305,
8306, 20359, 34255.

Thamnophis atratus hydrophilus (35). Shasta Co.: LSUMZ 16554, 16555, 16557, 34417, 34418,
35177, 36685, 36686, 36690-36701, 36705, 36747-36753, 39088, 39089, 40017, 40022, 40029,
46844, 46845.

Thamnophis atratus zaxanthus (23). Alameda Co.: LSUMZ 8129, 16532—16534, 16536, 16537,
16539, 16540, 34591-34593. Contra Costa Co.: LSUMZ 8131, 42815. Monterey Co.: LSUMZ
17023, 39097. San Luis Obispo Co.: LSUMZ 23907, 37189-37191, 37902. Stanislaus Co.: LSUMZ
44370, 44390, 44393.

Thamnophis elegans terrestris (42). Alameda Co.: LSUMZ 7908, 44780. Marin Co.: LSUMZ
7914. San Luis Obispo Co.: LSUMZ 55342-55346. San Mateo Co.: LSUMZ 7900-7904, 7907, 7911,
7912, 7916, 7918, 7920, 16502, 16504-16506, 16508, 16509, 19285-19291, 20356-20358, 34231,
34371, 34373, 40248. Santa Cruz Co.: LSUMZ 13536. Sonoma Co.: LSUMZ 16636, 34374.

APPENDIX C

Character support for the phylogenetic tree—— The ingroup clade, comprising southern popula-
tions of Thamnophis atratus, was supported by shorter SVL (<415.3 mm vs. 466.5 mm [mean values,
and hereafter]), fewer ventrals (<161.1 vs. 164.2), fewer subcaudals (<83.8 vs. 86.6), fewer ASR
(<19.6 vs. 20.2), fewer PSR (<16.1 vs. 16.9), broader interorbital width (>0.39 vs. 0.38), broader
MVSW (>1.9 vs. 1.7), STR value either greater than 4.9 or less than 3.9 (vs. 4.3), longer PCS (<0.80
vs. 0.83), greater eye diameter (>0.18 vs. 0.17), more maxillary teeth (>24.5 vs. 23.6), male/female
SVL ratio either greater than 0.81 or less than 0.91 (vs. 0.86), greater ML/FL ratio (<0.70 vs. 0.85),
lesser FL/PL ratio (>0.97 vs 0.92), greater LDL/LVL ratio (<0.66 vs. 0.71), and PRF/INL ratio either
greater than 1.01 or less than 1.11 (vs. 1.08).

The branch leading to A, D is supported by lower STR value (<3.9 vs. 5.0), and higher pupil/eye
ratio (0.47 vs. <0.45). The clade (G (B, F)) is supported by greater STR value (4.9 vs. <3.9) and more
maxillary teeth (25.6 vs. <24.8). The clade (B, F) is supported by more V (>159.5 vs. 153.3), narrow
interorbital (0.39 vs. 0.40), and lower pupil/eye ratio (0.43 vs. 0.45).

BOUNDY: SYSTEMATICS OF THAMNOPHIS ATRATUS 335

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© CALIFORNIA ACADEMY OF SCIENCES, 1999
Golden Gate Park
San Francisco, California 94118

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
ac Ea. AeA aa Lo peau Lc Ae ee ce

| OCT 21 1999

Volume 51, No. 7, pp. 337-364, 16 figs., 1 table. October 8, 1999

VW/O0G8 MAB

Neamecer®

Revisionary Systematics of the Western Pacific Soft Coral Genus
Minabea (Octocorallia: Alcyoniidae), with Descriptions of a
Related New Genus and Species from the Indo-Pacific

by

Gary C. Williams
Department of Invertebrate Zoology and Geology, California Academy of Sciences
Golden Gate Park, San Francisco, California 94118

and

Philip Alderslade
Museum and Art Gallery of the Northern Territory, GPO Box 4646
Darwin, Northern Territory, 0801, Australia

The dimorphic soft coral genus Minabea Utinomi, 1957, originally described from Japan,
is here recognized as a monotypic taxon and restricted to the type species, which is
designated as Minabea phalloides (Benham, 1928). Minabea ozakii Utinomi, 1957, pre-
viously recognized as the type species, cannot be morphologically differentiated from M.
phalloides, and is therefore considered a synonym. In light of the present situation in which
Benham’s and Utinomi’s type specimens cannot be located, and Benham’s type locality is
uncertain, recently collected material from New Zealand is here described in detail.

Eight other taxa, including one new species, which superficially resemble Minabea, are
allocated to a new genus, which is morphologically distinct and has an Indo-West Pacific
distribution. The new species has recently been discovered from Okinawa and the Philip-
pines. The Indian Ocean specimens upon which the species Alcyonium indicum Thomson
and Mackinnon, 1905, and Bellonella indica Thomson and Henderson, 1905, were based
have been reexamined, and it has been determined that they are conspecific and belong to
the new genus. These are compared with the new species. In addition, newly-collected
material of a recently described Melanesian species has allowed for an assessment of
phenotypic variability, which is the most marked of any species in the genus.

Revised keys and a table comparing the nine species in the two related genera, along
with a discussion of variability and distribution, are also included.

Several genera of alcyoniid soft corals with unbranched digitiform to capitate growth form and
dimorphic polyps are known. Included here are Acrophytum Hickson, 1900; Anthomastus Verrill,
1878; Malacacanthus J.S. Thomson, 1910; Minabea Utinomi, 1957; and Verseveldtia Williams,
1990 (see key to the dimorphic Alcyoniidae in the present paper).

Williams (1992b) monographed and revised the genus Minabea, in which five new species were
described, and nine species were recognized as valid for the genus. A subsequent comparison of
Minabea ozakii Utinomi, 1957 from Japan, with Minabea phalloides (Benham, 1928) from New
Zealand, has shown that the two are morphologically indistinguishable. Minabea phalloides is
therefore designated as the type species for the genus Minabea, and M. ozakii, previously accepted
as the type species of the genus, is necessarily relegated to the status of junior synonym. A

251)

338 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 7

redescription of Minabea phalloides (Benham, 1928) is included, from previously unavailable
material.

Comparative investigation has also shown that Minabea phalloides can be separated on the basis
of sclerite composition and siphonozooid morphology, from the seven other species previously
considered to belong to the genus Minabea. A new generic name is proposed to accommodate these
taxa. Included in this new genus is a recently discovered species from the western Pacific, which is
here described as new. Specimens of an Indian Ocean species, previously described as Bellonella
indica and Alcyonium indicum, are reexamined and compared with the new species. Generic revision
and the new taxon description makes a total of eight species known for the new genus in addition to
a single species in the genus Minabea.

An aspect of relevant interest is related here. One of these taxa, originally named Minabea
aldersladei by Williams (1992b), has been used in chemical products research. Ksebati and Schmitz
(1988) reported the discovery of diterpene lactones and minabeolides—a new group of withanolide
lactones—all isolated from this species, which was collected from Chuuk, Micronesia.

Recently collected material, referable to a species previously described as Minabea acrono-
cephala Williams, 1992b from the Solomon Islands, along with material from Fiji and the type locality
in Papua New Guinea, exemplifies a high degree of morphological variation with respect to growth
form, which is here assessed.

A brief review of the dimorphic genera of the Alcyoniidae is also provided, including revised
keys, a table of comparative characters for the one species of Minabea and eight species of the new
genus, and new distributional data for six taxa in the western Pacific and three in the Indian Ocean.

METHODS

Abbreviations used in the text are as follows: CAS (California Academy of Sciences, San
Francisco), CRRF (Coral Reef Research Foundation, Palau), NHM (The Natural History Museum,
London—formerly known as the British Museum of Natural History), NMNZ (National Museum of
New Zealand, Wellington—now known as Museum of New Zealand, with collection acronym
unchanged), NZOI (New Zealand Oceanographic Institute, Wellington—now known as National
Institute of Water and Atmospheric Research, with collection acronym unchanged), NTM (Northern
Territory Museum of Arts and Sciences, Darwin—now known as the Museum and Art Gallery of the
Northern Territory, with collection acronym unchanged).

Material was collected by SCUBA or bottom trawl, and preserved in 70% ethanol. Sodium
hypochlorite was used to isolate sclerites from soft coral tissues. An Olympus CH-2 compound
microscope with drawing tube was used to draw sclerites, while a Hitachi S-510 scanning electron
microscope was used to make scanning electron micrographs.

SYSTEMATIC ACCOUNT

KEY TO THE WORLD SPECIES OF MINABEA AND PARAMINABEA
(Revised from Williams 1992b:2)

1. Sclerites include spindles derived from radiates, or slender rodlike forms .................+.-.--. 2
==Spindlesiandirods‘absentyr ise) fo be. Gynt ete Se TS A STE fos St ea eee 3
2. Sclerites are radiates and slender, elongate, spiny rodlike forms which are densely distributed in the interior of the

colonys Coloniestelongateandifinserlikestapenine distallyaees- ieee neces nccnr an Minabea phalloides

—Sclerites are radiates and spindles derived from radiates. Colonies digitiform and cylindrical in shape

say UA a batch a id Bia Ade lend arial ae tree: Pogue Co aimee tre Panny Sansa Notes gba ue Paraminabea kosiensis
3iColonies lobate Gridigitate ung «ts... AO Os ee MES SESE DESPA Schr, Osteo ano a at oe fe 4
—Colonies digitifornijps). hrs. cysweks cect od cho teen ees ous Migs see seh ee od cio) yee aa ee 5
4. Colonies digitate with 4-6 fingerlike branches emanating laterally from a thicker main stem . . . Paraminabea arborea

WILLIAMS AND ALDERSLADE: REVISIONARY SYSTEMATICS OF MINABEA 339

—Colonies lobate with <3 robust lobes arising from the top of a basal stalk, or digitiform (fingerlike to clavate or

GOTTEAIEIEG)) go pcm Gao Bo old a a Oi oO Bl Smo omtOeorcly 9 .de pxttisto. 5 c. deomonce rane: Paraminabea indica
Seeoloniesidicititonmm dinpersshaped or cylindrical) <0.) a) as eee eee ee eee 6
—Colonies clavate to capitate (conical or dome-shaped to torch-shaped)..............-.....+....... 9

6. Surface of coenenchyme with two distinct types and sizes of sclerites—small radiates and large robust barrels
Se GANONG Sood oho MOOD. ty Ge. CRORES, o he CREED ayS C1 OSRER SO) akan een e Aare ge .. . Paraminabea cosmarioides

—Sclerites throughout the colony are not of conspicuously disparate sizes. 2... ee 7
7. Polyparium forms > 50% total colony length. Colonies from shallow water (<30m) ...........-....04. 8
—Polyparium forms < 50% total colony length. Colonies from deeper water (>70m).........- Paraminabea indica
Susclenites areeieht-radiatesjandiclongate barnrelSiey cae et teehee Paraminabea aldersladei
—Sclerites are radiates, robust barrels and tuberculate spheroids ................... Paraminabea robusta

9. Colonies conical, short (usually wider than long); stalk inconspicuous (< length of polyparium); sclerites include double
GUIS’ bo rod ach B ope ee ee RRS oo ABS eS. AS “oat OSG Sogtian cy okog Cer ceria ee Paraminabea goslineri

—Colonies dome-shaped or torch-shaped, (usually longer than wide); stalk variable in length (usually > length of
polyparium); sclerites are exclusively barrels and radiates .........-..--.--. Paraminabea acronocephala

Minabea Utinomi, 1957

Minabea Utinomi, 1957:139; Utinomi and Imahara, 1976:206 (in part). Tixier-Durivault, 1987:154. Williams,
1992:2 (in part).
Anthomastus (non Verrill, 1878): Benham, 1928:79.

TYPE SPECIES. — Anthomastus phalloides Benham, 1928 by subsequent designation (present
work); New Zealand.

REVISED GENERIC DIAGNOSIS. — Alcyoniid soft corals, digitiform (unbranched, finger-shaped
and tapering distally). Polyps dimorphic. Polyparium conspicuously longer than stalk. Polyps evenly
distributed over surface of polyparium, retractile, without calyces or sclerites. Siphonozooids crowded
between autozooids, numerous and conspicuous, 0.40.7 mm in diameter, 1/3—1/2 diameter of
retracted autozooids. Sclerites densely set: predominantly radiates in the surface layers, radiates and
short spindles in the stalk interior, and numerous spiny rods more-or-less longitudinally disposed in
the canal walls of the polypary.

DIVERSITY AND DISTRIBUTION. — A monotypic genus from central Japan (250-270 m) and New
Zealand (139-179 m).

Minabea phalloides (Benham, 1928)
Figs: I, 2,3, 4; 15; 16

Anthomastus phalloides Benham, 1928:79.
Minabea phalloides: Utinomi and Imahara, 1976:211. Williams, 1992b:19, 23.
Minabea ozakii Utinomi, 1957:139.

MATERIAL EXAMINED. — NMNZ Cn 807 (NZOI Stn R81), New Zealand, North Island, Bay of
Plenty, 37°35.9'37.6’S 176°59.5'—59.8’E, 139-179 m depth, 20 January 1979, collected on NZOI
cruise; three specimens total: two whole specimens and one partial specimen that is missing the stalk.

DESCRIPTION. — Growth form and size. The three specimens examined measure 43 mm and
73 mm in length (whole specimens), and 73 mm in length (partial specimen). The smallest specimen
is attached to the inside of a clam shell. The three specimens vary in minimum width from 5—7 mm,
and in maximum width from 16-18 mm. The specimens are elongate digitiform, slightly to distinctly
curved, and are conspicuously tapered toward the rounded distal terminous (Fig. 1A represents the
73 mm whole specimen). The polyparium and stalk are distinctly demarcated. The polyparium is
longer than the stalk. The stalk constitutes 17-38% of the total colony length (Fig. 1A).

340 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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FiGuRrE |. Minabea phalloides. A. Entire colony, 73 mm in length; B. Detail of the polyparium surface showing one autozooid
and numerous siphonozooids, diameter of diagram represents 2.7 mm; C. Portion of gastric cavity wall showing disposition
of sclerites of spiny rods, scale bar = 0.2 mm; D. A single rod from gastric cavity wall (polyparium interior), 0.21 mm in length.

WILLIAMS AND ALDERSLADE: REVISIONARY SYSTEMATICS OF MINABEA 34]

FiGuRE 2. Sclerites of Minabea phalloides. A. Sclerites from the surface of the polyparium; B. Sclerites from the interior of
the polyparium; C. Sclerites from the surface of the stalk; D. Sclerites from the interior of the stalk. Scale bar = 0.1 mm.

342 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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FiGURE 3. Minabea phalloides. Scanning electron micrographs of sclerites from the surface of the polyparium. A. 0.26 mm;
B. 0.18 mm; C. 0.16 mm; D. 0.06 mm; E. 0.04 mm; F. 0.05 mm; G. 0.05 mm.

WILLIAMS AND ALDERSLADE: REVISIONARY SYSTEMATICS OF MINABEA

FIGURE 4. Minabea phalloides. Scanning electron micrographs of sclerites from the surface of the stalk. A. 0.05 mm; B. 0.07
mm, C. 0.15 mm; D. 0.12 mm; E. 0.08 mm; F. 0.06 mm.

344 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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Polyps. The polyps are evenly distributed throughout the polyparium. The preserved expanded
anthocodiae average 2 mm in diameter, and have slender tentacles with 10—15 pinnules per side. The
siphonozooids are numerous and conspicuous, disposed between the autozooids, each measuring
approximately 0.4 mm in diameter (Fig. 1B).

Sclerites. The sclerites throughout the colonies are densely set and are predominantly radiates
and spiny rodlike forms. A few triradiate or cross-shaped forms are also present. The rods of the
polyparium are numerous and elongate, whereas any similar sclerites in the stalk are much fewer,
shorter and stouter (Fig. 2). Surface samples may also include a few subsurface rods between 0.13
and 0.27 mm. These rods (Fig 2A, 3A—C) are relatively spiny, with smooth, conical to somewhat
acute tubercles towards both ends, the middle portions often being devoid of prominences. Although
the interior of the polyparium includes a few radiates immediately below the surface (0.04—0.09 mm),
further within, the walls of the gastric cavities exclusively contain these spiny rods, numerous, often
clustered, more-or-less longitudinally disposed, and 0.14—0.23 mm long (Fig. 1C, D). The surface of
the stalk contains mostly six- and eight-radiates that are 0.06—0.12 mm long, and elongate forms,
0.12—0.15 mm long, with two girdles of prominences (Fig. 2C). The interior of the stalk has six- and
eight-radiates, 0.04—0.10 mm long, and robust, sparsely ornamented spindles, 0.15 and 0.18 mm in
length (Fig. 2D), which do not have a middle portion devoid of pronounced tubercles (Fig. 4C), as is
the case with the rods of the polyparium (Fig. 3A—C). In addition, the tubercles of the stalk spindles
are more pronounced than those of the polyparium rods (compare Fig. 2D with 2A).

Color in alcohol-preserved specimens. The color of the stalk is grey-white to cream white. The
polyparia are pale yellow-orange, sometimes lighter distally approaching grey-white to cream white.
The autozooids are white. The sclerites from the middle of the polyparium are pale yellow, while
those of the stalks are colorless.

DISTRIBUTION. — New Zealand, 139—179 meters in depth; and Honshu, Japan, 250-270 meters
in depth (Figs. 15C, 16).

REMARKS. — The holotype was not available for examination and cannot be located at present.
In addition, the type locality is uncertain. Benham (1928:79) states, “A single colony was obtained
by Hon. G. M. Thomson during the experimental fishery cruise in the G.S.Y. ‘Hinemoa,’ in 1915.
Unfortunately neither the exact station at which it was dredged nor the depth was noted, but it seems
probable that it was somewhere in Foveaux Strait.” All recently collected material is from the opposite
end of the country, in the Bay of Plenty (Fig. 15C).

The holotype of Minabea ozakii Utinomi, 1957, for which the genus Minabea was originally
proposed, cannot be located at present. The type locality is recorded as the Kii Strait, southwest of
the town of Minabe, southern Honshu, and Utinomi’s description is sufficiently detailed to show that
the Japanese specimen is morphologically indistinguishable from the New Zealand material of
Minabea phalloides. Because of the lack of differentiating morphological characters, we here consider
Minabea ozakii as ajunior synonym of Minabea phalloides, despite the fact that the taxon is comprised
of two remarkably disjunct populations.

Utinomi and Imahara (1976:211) considered Anthomastus agilis Tixier-Durivault, 1970 from
New Caledonia to be a possible synonym of Minabea phalloides. However, Alderslade (1994) has
shown the taxon of Tixier-Durivault to be included in the genus Sinularia.

Paraminabea gen. nov.

Minabea (non Utinom1, 1957:139): Utinomi and Imahara, 1976 (in part):206. Bayer, 1981:913. Tixier-Durivault,
1987:154. Williams, 1992b (in part):2.

Bellonella (non Gray, 1862):Thomson and Henderson, 1905:274; Bayer, 1974:261; Faulkner and Chesher,
1979267, pla 22:

Alcyonium (non Linnaeus, 1758): Thomson and Mackinnon, 1910:174.

WILLIAMS AND ALDERSLADE: REVISIONARY SYSTEMATICS OF MINABEA 345

DIAGNOSIS. — Alcyoniid soft corals with colonies commonly unbranched and dome-shaped or
hemispherical to digitiform, or rarely digitate/lobate with few branching processes. Polyps dimorphic.
The distal polyparium arises from a proximal stalk that is variable in length. Polyps evenly distributed
over surface of polyparium and capable of complete retraction, calyces and sclerites absent. Siphono-
zooids numerous to scarse, small to minute (< 0.3 mm in diameter; < 1/3 diameter of retracted
autozooids), distributed between the autozooids. Sclerites of surface and interior of colony are densely
set: mostly barrels and six- or eight-radiates; spindlelike forms derived from radiates, and tuberculate
spheroids may also be present; seven-radiates or double stars sometimes occurring. Color red, orange,
yellow or pinkish white to cream white.

DIVERSITY AND DISTRIBUTION. — Eight species in the genus are here considered valid (Table
1); with an Indo-West Pacific distribution from southeastern Africa, Seychelles, the Maldives, and
Sri Lanka to Japan, Guam, Chuuk and Fiji (Figs. 15, 16); 1-370 m in depth.

TYPE SPECIES. — Bellonella indica Thomson and Henderson, 1905, here designated.

Paraminabea acronocephala (Williams, 1992)
Figss SBE. 6, 7:8, 9; 15B16

Minabea acronocephala Williams, 1992b:3. Gosliner, Behrens, and Williams 1996:35.

MATERIAL. — CAS 101094, Solomon Islands, N. Russell Group, Karomolun Island, | 1 m depth,
14 November 1994, Gary C. Williams using SCUBA, one specimen in two pieces.

DIAGNOSIS. — Growth form clavate to torch-shaped with distal-most portion of stalk slightly
wider than polyparium. Hemispherical polyparium represents distal-most 1/3 of entire colony length.
Sclerites are six- to eight-radiates and barrels, 0.05—0.11 mm long. Color of polyparium in the living
animal as well as the wet-preserved specimen is deep yellow, with stalk pale orange.

DESCRIPTION. — Growth form and size. The specimen is 20 mm in length with a maximum
width of 10 mm, unbranched and clavate, lamp-shaped to torch-shaped (Fig. 6A). The stalk and the
polyparium are distinctly delimited by a somewhat undulating border. The polyparium is shorter than
the conspicuous stalk. The stalk is prominent and elongate, the length being approximately twice its
width. The proximal holdfast region of the stalk is spread. The distal-most portion of the stalk is
distinctly flared and slightly wider than the polyparium, representing the widest portion of the entire
colony. The polyparium is hemispherical and represents approximately 33% of the entire colony
length, while the stalk and holdfast comprise the remaining 67% (Fig. 6A).

Polyps. The polyps are all tightly retracted. The autozooids are represented by slightly indented
pits, evenly distributed over the surface of the polyparium. The siphonozooids are minute, indistinct,
and scattered between the autozooids (Fig. 6B).

Sclerites. The sclerites throughout the specimen are radiates and robust barrels. Sclerites of the
polyparium and stalk interior are generally larger and more robust than those of the surface
coenenchyme. Sclerites from the surface of the polyparium are radiates and relatively narrow barrels,
0.05—0.09 mm in length (Fig. 7A); some of the radiates are only very sparsely ornamented and may
be sclerites still in the process of developing. Sclerites from the interior of the polyparium are mostly
robust barrels, often with distinct medial waists. Most of these vary between 0.08 and 0.12 mm in
length (Fig. 7B). The surface of the stalk contains sclerites that are radiates and relatively small barrels,
0.05—0.08 mm in length (Fig. 7C). The interior of the stalk contains mostly robust and slightly larger
barrels, 0.07—0.10 mm in length (Fig. 7D).

Color in living animal and alcohol-preserved specimen. The upper portion of the stalk, and
the polyparium, are deep yellow, while the lower part of the stalk and holdfast are salmon orange.
The sclerites of the polyparium are yellow, while those from the middle of the stalk are pale
orange-yellow.

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 7

FIGURE 5. A. Paraminabea arborea sp. nov., underwater photograph of the paratype in life (NTM C-12559) from Zambales,
Luzon, Philippines (photograph by Patrick Colin, CRRF). B-E. Paraminabea acronocephala. B. Underwater photograph of a
yellow form in life, Banana Rock, Madang Barrier Reef, Papua New Guinea, 23-26 m depth, 19 June 1992 (photograph by
Gary Williams, CAS); C. Underwater photograph of voucher specimen in life (CAS 094306) from Madang, Papua New Guinea
(photograph by Gary Williams, CAS); D. Underwater photograph of a yellow and red form in life, Rasch Passage, Madang
Barrier Reef, Papua New Guinea, 15-21 m depth, 17 June 1992 (photograph by Gary Williams, CAS); E. Underwater
photograph of voucher specimen in life (CAS 111926) from Somosomo Straits, Fiji (photograph by Patrick Colin, CRRF).

WILLIAMS AND ALDERSLADE: REVISIONARY SYSTEMATICS OF MINABEA 347

FIGURE 6. Phenotypic variation in Paraminabea acronocephala. A. Specimen from Solomon Islands, 20 mm long x 10 mm
wide (CAS 101094); B. Detail from the polyparium in A showing a single autozooid surrounded by five siphonozooids, diameter
of figure 2.5 mm; C. Specimen from Fji, 16 mm long x 18 mm wide (CAS 111926). D-F. Specimens from Madang, Papua
New Guinea. D. 13 mm long x 7 mm wide (CAS 078417); E. 15 mm long x 17 mm wide (CAS 094306); F. 6 mm long x 8
mm wide (CAS 078417).

348 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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FIGURE 7. Sclerites of Paraminabea acronocephala. A. Sclerites from the surface of the polyparium; B. Sclerites from the
interior of the polyparium; C. Sclerites from the surface of the stalk; D. Sclerites from the interior of the stalk. Scale bar = 0.1
mm.

WILLIAMS AND ALDERSLADE: REVISIONARY SYSTEMATICS OF MINABEA

FIGURE 8. Paraminabea acronocephala. Scanning electron micrographs of sclerites from the surface of the polyparium of
specimen shown in Figure 6A. A. 0.09 mm; B. 0.07 mm; C. 0.08 mm; D. 0.07 mm; E. 0.06 mm; F. 0.06 mm.

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES

350

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WILLIAMS AND ALDERSLADE: REVISIONARY SYSTEMATICS OF MINABEA 3

FiGure 9. Paraminabea acronocephala. Scanning electron micrographs of sclerites from the surface of the stalk of specimen
shown in Figure 6A. A. 0.05 mm; B. 0.08 mm; C. 0.07 mm; D. 0.08 mm; E. 0.07 mm.

DISTRIBUTION. — The specimen is from the Russell Group (Solomon Islands). This variable
species 1s also known from the Madang region and the Louisiade Archipelago (Papua New Guinea),
as Well as Fiji (Williams 1992, and the present work) (Figs. 15B, 16).

REMARKS. — This specimen differs from all others known of the species by having a prominent,
elongated stalk and hemispherical polyparium (see discussion of phenotypic variability).

352 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 7

Paraminabea arborea sp. nov.
miesesan lO, i125 13, 15A, 16

MATERIAL. — HOLOTYPE: CAS 112114, Japan, Ryukyu Islands, Okinawa, Serigaki, 68 m depth,
12 February 1995, Robert F. Bolland using SCUBA, one whole specimen. PARATYPE: NTM C-12559,
Philippines, Luzon, Zambales, Luan Island near Masinloc Town (15° 29.06°N; 119°54.41°E), 10-12
meters depth, 1 May 1995, Pat Colin using SCUBA, one whole specimen.

DIAGNOSIS. — Growth form digitate, with 3-6 somewhat curved fingerlike processes, rounded
to slightly clavate distally, not pointed or tapered. Main trunk tapers distally. Stalk represents the
proximal-most one sixth of the total body length. Sclerites mostly robust barrels (0.06—0.10 mm long)
and six to eight radiates (0.04—0.06 mm long). Color in life as well as preserved is red-orange, with
yellow dots representing retracted autozooids.

DESCRIPTION. — Growth form and size. The holotype is 64 mm long while the paratype is 49
mm in length. The branched polyparium arises from a short robust stalk. The polyparium comprises
approximately 80% of the total length of the colony in the holotype, and approximately 87% in the
paratype. The proximal-most portion of the holotype is represented by a holdfast region that spreads,
and in the paratype it is slightly flared from the rest of the stalk. The holotype has six relatively narrow,
digitate processes that are curved throughout their length, and are rounded or slightly clavate distally
(Fig. 10). The paratype has four such processes. The digitate processes arise at different levels along
the length of the main trunk, which is elongate, upright, and tapers distally.

Polyps. Several autozooids with their white anthocodiae protrude from the polyp apertures in
the preserved holotype. Maximum diameter of the tentacle spread is 1.7 mm. Tentacles have about
18 pinnules arranged more or less in two rows down each edge. The siphonozooids are sparsely
arranged between the autozooids and are minute (ca. 0.1 mm in diameter) (Fig. 10C). They appear as
small yellow hemispherical protuberances on the surface of the polyparium (Fig. SA).

Sclerites. Sclerites of the surface of the polyparium are mostly barrels 0.07—0.09 mm long and
six- or eight-radiates 0.03—0.06 mm in length (Figs. 11A, 12), while those of the interior of the
polyparium are slightly larger and more robust barrels, 0.10—0.11 mm in length (Fig. 11B). The
surface of the stalk contains sclerites that are mostly six-radiates 0.04—0.08 mm long and robust barrels
0.09-0.11 mm in length (Figs. 11C, 13). The interior of the stalk contains mostly barrels 0.07—011
mm in length; some of these may approach tuberculate spheroids (Fig. 11D).

Color of living and wet-preserved material. The color of the coenenchyme is predominantly
red-orange approaching brick red. The expanded autozooids are white, and the numerous yellow spots
on the surface of the colony are the closed polyp apertures. The sclerites are very pale orange to
colorless and mostly transparent.

DISTRIBUTION. — Luzon (Philippines) and Okinawa (Ryukyu Islands, Japan); 10-68 m depth ~
(Figs. ISA, 16).

ETYMOLOGY. — The specific epithet is derived from the Latin, arbor (tree); in reference to the
sparsely branched growth form characteristic of this species.

REMARKS. — This species is the only member of the genus with a digitate/lobate growth form
besides Paraminabea indica (see description below). Paraminabea indica can be digitiform or lobate
with thick short lobes arising from a short common base at roughly the same level (Fig. 14), while P.
arborea is digitate with several narrow fingerlike processes arising from an elongate common trunk
at different levels (Fig. 10). All other described species are unbranched and digitiform to dome-shaped
(Table 1). Some species of the alcyoniid genus Eleutherobia may be lobate to digitate such as E.
somaliensis Verseveldt and Bayer, 1988, which superficially resembles Paraminabea arborea.
However, Eleutherobia differs from Paraminabea in having monomorphic polyps, not dimorphic
ones.

WILLIAMS AND ALDERSLADE: REVISIONARY SYSTEMATICS OF MINABEA 353

Ficure 10. Paraminabea arborea sp. nov. A. Holotype, 64 mm in length; B. Paratype, 49 mm in length; C. Detail of the
polyparium surface showing one autozooid surrounded by thirteen siphonozooids, diameter of diagram 2.7 mm.

354 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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FIGURE 11. Sclerites of Paraminabea arborea sp. noy. A. Sclerites from the surface of the polyparium; B. Sclerites from
the interior of the polyparium; C. Sclerites for the surface of the stalk; D. Sclerites from the interior of the stalk. Scale bar =
0.1 mm.

wy)
N
Nn

WILLIAMS AND ALDERSLADE: REVISIONARY SYSTEMATICS OF MINABEA

FiGurE 12. Paraminabea arborea sp. nov. Scanning electron micrographs of sclerites from the surface of the polyparium
of the holotype. A. 0.04 mm; B. 0.05 mm: C. 0.05 mm; D. 0.04 mm; E. 0.08 mm; F. 0.07 mm.

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 7

FIGURE 13. Paraminabea arborea sp. nov. Scanning electron micrographs of sclerites from the surface of the stalk of the
holotype. A. 0.07 mm; B. 0.10 mm; C. 0.06 mm; D. 0.08 mm; E. 0.09 mm; F. 0.08 mm.

WILLIAMS AND ALDERSLADE: REVISIONARY SYSTEMATICS OF MINABEA Soi

Paraminabea indica (Thomson and Henderson, 1905)
Figs. 14, 15

Bellonella indica Thomson and Henderson, 1905:274, pl. 6, fig.5. Non Bayer, 1974:261. Non Faulkner and
Chesher, 1979:267, pl. 22.

Alcyonium indicum Thomson and Mackinnon, 1910:174, pl. 12, fig. 7.

Minabea indica: Williams, 1992b:14, figs. 10, 11.

MATERIAL. — HOLOTYPE: NHM Reg. No. 1933.3 13/206 (labeled as Nidalia indica). Sri Lanka.
South of Galle: deep water, Sir J. A. Thomson Collections, one specimen in which part of the
polyparium has been cut away longitudinally and is missing.

NHM Reg. No. 1912.2.24.72. Seychelles, 39 fathoms; A/cyonium indicum specimen of Thomson
and Mackinnon.

REDESCRIPTION. — A redescription of the holotype was provided by Williams (1992), but the
sclerites are figured in the present paper (Fig. 14D—Q) for comparative purposes. A redescription of
the specimen of A/cyonium indicum described by Thomson and Mackinnon in 1910 follows.

Growth form and size. The specimen is 40 mm in height. The lobate polyparium arises from
the stout stalk (Fig. 14A). A portion of the surface of the stalk has been cut away. The stalk measures
15 mm at its widest diameter. The polyparium comprises no more than 50% of the length of the
specimen. The three lobes of the polyparium arise from the stalk at approximately the same level.
One of these three lobes has been cut away and is missing. The base of this lobe is 6 mm in diameter.
The remaining lobes are short, stout, and rounded at the tips, not tapering. The central one is 17 mm
long and 7 mm wide, while the lateral one is 16 mm long by 8 mm wide.

Polyps. The autozooids appear as small yellow spots distributed evenly over the surface of the
polyparium. Some of these form low, rounded protuberances (ca. 0.8 mm in diameter). The siphono-
zooids are apparently sparsely distributed, and are minute and difficult to observe in the tightly
retracted specimen. Some occur only as very minute pores between the autozooids.

Sclerites. The sclerites are robust barrels (many of which are only slightly longer than wide),
radiates that vary as six-, seven-, or eight-radiates, and some forms that approach double stars.
Sclerites from the polyparium surface and interior are similar, and are more variable in form than the
stalk sclerites. The polyparium has barrels, radiates, and double stars, while the surface and interior
of the stalk contains many robust barrels with some radiates (Fig. 14B, C). These sclerite characters
are similar to that of the holotype of Paraminabea indica (Fig. 14E-Q).

Color of alcohol-preserved specimen. The specimen is uniform reddish orange, with lemon
yellow dots representing the closed autozooid apertures.

DISTRIBUTION. — Indian Ocean: Sri Lanka (“deep water,” Thomson and Henderson 1905:275),
the Seychelles (71 m, Thomson and MacKinnon 1910:174), and possibly the Maldive Islands (ca. 10
m, Schuhmacher and Hinterkircher 1996:50)(Fig. 15).

REMARKS. — Both Paraminabea indica and P. arborea sp. nov. may appear branched (digitate
or lobate), the only two species in the genus observed to have this growth form. Because of this
superficial similarity, it was necessary to compare the specimens of P. indica with those of the new
species. (See Remarks for P. arborea sp. nov. for differentiation of these taxa).

The lobate specimen of A/cyonium indicum was compared with that of the digitiform holotype
of Paraminabea indica in order to ascertain whether the two are indeed conspecific or represent two
separate species. Comparative investigation shows them to be conspecific in our opinion. Sclerite
form and distribution are consistent in both specimens, as is the fact that the polyparia and the stalks
of both specimens each occupy approximately half of the total specimen length.

358 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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DISCUSSION

PHENOTYPIC VARIABILITY. — The proliferation of intraspecific phenotypic variability in seden-
tary marine animals has been noted or described by several authors, including Veron (1986:60;
1995:14) and Miller (1995) for scleractinian corals, and for octocorals Hickson (1907:25—36), J. S.
Thomson (1921:165, 166), Groot and Weinberg (1982:293), Verseveldt (1980, 1982, 1983), and
Williams (1986:255; 1992a:282; 1992c:383, 384). The high degree of variation in many soft corals
is at least partly attributable to localized isolation of populations, lack of mobility in adults, and
ephemeral larval stages with limited or short-range dispersal capabilities (Williams 1986:255).

Potts (1984:48) linked a potential cause of prolific variability in Indo-Pacific reef corals to chronic
environmental disturbances such as sea level fluctuations associated with glacial cycles. Veron
(1995:126) stated that glacio-eustatic changes have affected tropical western Pacific reef coral
diversity to a greater degree than in other areas of the world, due largely to the vast extent of
shallow-water habitat in the region. These pronounced environmental perturbations have produced a
complex mosaic of faunas over time, through the alternation of environmental regimes (marine vs.
terrestrial), the isolation of refugia, and the subsequent repopulation of previously non-marine areas
with the concomitant overlap of relictual forms from the various high diversity refugia.

Millard (1978:182), working with hydroids, concluded that at least two factors are indicators of
an active region of evolutionary radiation: the proliferation of intraspecific phenotypic variation and
a preponderance of endemic taxa. Regarding the octocoral fauna, Williams (1992c:384) suggested
that in the region of overlap between the Indo-Pacific and the Cape Endemic Province in southern
Africa, a high degree of endemism ts derived from older and well-established components of the
fauna, while the prolific phenotypic variability observed in these endemics is associated with a more
recent result of repetitive glacio-eustatic disturbances during the past several million years.

Related to this, regarding genotypic variation, Solé-Cava and Thorpe (1991:66, 69) provide
evidence for very high levels of genetic variation for marine sponges and coelenterates in general.

Figure 16 shows the ranges of the six presumably endemic species of Minabea and Paraminabea
in the western Pacific. Of these six, Paraminabea acronocephala exemplifies, by far, the greatest
range of phenotypic variation regarding growth form and color (Fig. 6). Williams (1992b:5, fig 2D)
illustrates variation in colony shape at the type locality of this species.

Recently collected material from throughout the range of the species shows a wide spectrum of
variation in growth form. The specimen from the Solomon Islands described above (CAS 101094)
has an elongate stalk that flares toward its distal end, which represents the widest portion of the entire
colony. The polyparium is hemispherical and is narrower than the distal part of the stalk. The stalk
represents two thirds while the polyparium 1s approximately one third of the total colony length (Fig.
6A). The specimen is bicolored, deep yellow and salmon orange.

A recently collected specimen from Fiji (CAS 111926) (Figs. SE, 6C) resembles the Solomon
Islands material except that the stalk 1s extremely short. The polyparium is hemispherical and narrower
than the smooth bandlike upper portion of the stalk. The holdfast and lower portion of the stalk are
<5 mm in length, and comprise < 5% of the total colony length. Color of the specimen is uniform
rust orange to reddish brown.

Somewhat similar to the Fiji material is a specimen from Papua New Guinea (CAS 078417) (Figs
5C, 6E). It also has a hemispherical polyparium atop a short stalk. The uppermost portion of the stalk
is smooth and bandlike and exceeds the width of the polyparium. The lower portion of the stalk tapers
proximally. The stalk and holdfast constitute approximately 50% of the length of the specimen. Color
is yellow-orange with retracted autozooids shown as red eight-pointed stars on the surface of the
polyparium.

Two other specimens, also from the north coast of Papua New Guinea (both CAS 078417) (Figs.
6D, F) differ significantly from those with hemispheric polyparia, by having relatively flattened or

2)

WILLIAMS AND ALDERSLADE: REVISIONARY SYSTEMATICS OF MINABEA

Figure 14. Paraminabea indica. A. Specimen of “Alcyonium indicum,” described by Thomson and Mackinnon (1910:174,
pl. 12, fig. 7) from the Seychelles, 40 mm in length; B. Sclerites from the surface of the stalk; C. Sclerites from the surface of
the polyparium, scale bar = 0.1 mm; D. Holotype of Bellonella indica Thomson and Henderson (1905:274, pl. 6, fig. 5) from
Sri Lanka, 20 mm in length. E—M. Sclerites from the surface of the polyparium. E. 0.50 mm; F. 0.06 mm; G. 0.05 mm; H. 0.05
mm; I. 0.05 mm; J. 0.07 mm; K. 0.05 mm; L 0.07 mm; M. 0.07 mm. N—Q. Sclerites from the surface of the stalk. N. 0.08 mm;

O. 0.08 mm; P. 0.08 mm; Q. 0.07 mm.

360 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 7

see inset A

> 6 iy be

@>.

SS ® ace ; ‘
=>... aoe a « ie
see inse

e rr 2 Se ¥ : ee
‘o-. e

see inset oF
9 &

P. cosmarioides

10°E
|

is

FIGURE 15. Maps of the Indo-West Pacific showing distribution of the genera Minabea (A) and Paraminabea (@). A. China
and Philippine Sea, showing collecting stations for Paraminabea arborea sp. nov.; B. Melanesia showing collecting stations
for Paraminabea acronocephala; C. New Zealand collecting stations for Minabea phalloides. Arrows show type localities.

WILLIAMS AND ALDERSLADE: REVISIONARY SYSTEMATICS OF MINABEA 361

FiGURE 16. Map of the western Pacific Ocean showing geographic ranges for five species of the genus Paraminabea and
the monotypic genus Minabea .

low rounded polyparia where the polyps are restricted to the distal-most extremity of the colony. The
first-mentioned specimen has an elongated stalk while the second 1s dome-shaped with a shorter stalk.
Both specimens show the distal-most portion of the stalk as the widest portion, a feature that is
characteristic for the species.

Color of Bismarck Sea material is variable, from uniformly yellow or orange-red to bicolored
yellow and orange with varying amounts of red-orange. Some specimens are yellow with red polyparia
or with retracted autozooids showing as red eight-pointed stars on the polyparium surface.

DISTRIBUTION OF THE GENERA MINABEA AND PARAMINABEA (See Table 1; Figs. 15, 16). — These
genera are recorded from the following localities, and range in depth from 1-370 m (Williams,
1992b:12, 17). First records are shown in parenthesis. Indian Ocean: Transkei and Natal, South
Africa (Williams 1992a:314 for Paraminabea cosmarioides and P. kosiensis); Maldive Islands
(Schuhmacher and Hinterkircher 1996:50 for Paraminabea sp. indet.); Seychelles (Thomson and
MacKinnon 1910:174 for Paraminabea indica as Alcyonium indicum); Sri Lanka (Thomson and
Henderson 1905:274 for Paraminabea indica as Bellonella indica); Rowley Shoals, Western Austra-
lia (Williams 1992b:7 for Paraminabea aldersladei). Western Pacific Ocean: Malaysian Borneo,
the Philippines, and the Solomon Islands (Gosliner, Behrens, and Williams, 1996:35 for Paraminabea
aldersladei and P. acronocephala); Great Barrier Reef, Papua New Guinea, Fiji, and Guam (Williams
1992b; 5, 13 for Paraminabea aldersladei, P. acronocephala, and P. goslineri); Palau (Bayer

362 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 7

1974:261 for Paraminabea aldersladei as Bellonella indica); New Zealand (Benham 1928:79 for
Minabea phalloides as Anthomastus phalloides), Okinawa (present work for Paraminabea arborea);
Honshu, Japan (Utinomi 1957:144 for Minabea phalloides as M. ozakii); Chuuk, Federated States of
Micronesia (present work, CAS No. 096509, Paraminabea aldersladei, ).

Of the six presumably endemic species in the western Pacific (Minabea phalloides, Paraminabea
acronocephala, P. aldersladei, P. arborea sp. nov., P. goslineri, and P. robusta), P. aldersladei shows
the greatest contiguous range of distribution, as well as some limited sympatry with P. arborea in the
northwest and P. acronocephala in the east. Minabea phalloides and Paraminabea robusta also show
overlapping distributions. Minabea phalloides is the only taxon with a distinct disjunct distribution
(Fig. 16).

Regarding bathymetrics, Minabea and Paraminabea show wide depth distribtutions, unlike
other genera of the Alcyoniidae such as Sinularia, Lobophytum, Sarcophytum, and Cladiella.
Minabea and Paraminabea are azooxanthellate taxa, differing in this regard from the other four
genera, which are zooxanthellate and restricted to the relatively shallow water of coral reefs. The lack
of zooxanthellae allows Minabea and Paraminabea to inhabit a wide range of depths, as well as a
greater variety of habitats such as caves and alcoves.

KEY TO THE GENERA OF ALCYONIIDAE WITH DIMORPHIC POLYPS
(Revised from Williams 1990:34)

A revised key is provided here, which utilizes new data and enables easy differentiation of the
various dimorphic alcyoniid taxa.

lS Goloniesiwaithitmmorphic polyps. esozooids present semen =) cy any) cic ace) tenn newcrene Heteropolypus
——Colomesswith dimorphicipolypsionly-mesozooids/absemtsy ss) yi) ie ate ee ee eee py
2. Polyparium capable of partial or complete retraction into the stalk ....................--..+--. 3
—Polyparium incapable of partial or complete retraction into the stalk .. 2... 1.2... eee ee ee ee es 4
3. Polyps disposed on several digitiform lobes that are capable of at least partial retraction into the stalk. Stalk not

COVEEGl Wine OTN CUI s eENS phate See t a ls SN Soale SS oS ote et eo cc Carotalcyvon
—Polyps disposed on a rounded capitulum that is capable of complete retraction into the stalk. Stalk covered with a

omnyicuticlewascs = Wy theses al ro Gene emey eee orca) we gin hepa et, Ge Sy eee ee rem Ce Malacacanthus
4. Colonies capitate with a hemispherical or spherical capitulum, digitiform with a tapering polyparium, or

digitate/lObate ie reece ere ce ara caer. uNeneoss eich Meee ed cs ia oteione sac casag fo reece 5
—Colonies with a broad polyparium either flared and discoidal often with deep open marginal folds, or surface with

numerous often crowded closed folds or lobes that vary from low and hemispherical to digitiform or ridgelike ... 9
SMCOlONIeS capitate mew pc se: mca Cats | sais Sea tge cobb ace Cache tans: Su icy Gate Ot its nS a ee a 6
—— Colomesidicititormiomdicitate/lobatelr ry cuencmemcms ci crc te icici citer cints atc N+ aC 7
6. Coenenchymal sclerites are double stars and spiny rods... ............ Anthomastus (including Bathyalcyon)
—Coenenchymal sclerites are eight radiates and/or double stars, barrels and tuberculated spheroids... .. . . Verseveldtia
(mSclenitesjaremumenousistoutsthonm Club Speyememenciraicmey Carn ntne ncn urs mein mnt iets ee 0 ee an Acrophytum
—Sclerites are mostly radiates, sometimes with spindles orspiny rods ...................-+.-...... 8
8. Sclerites are predominantly radiates and spiny rodlike forms. Siphonozooids are large (0.4—0.7 mm in diameter; or

WS=l/2itherdiametenofiretracted autoZOoids) ane cnceee tuum et eieca tone ecient ae ne Minabea
—Sclerites are predominantly radiates, sometimes barrels, double stars, or spindles derived from radiates. Siphono-

zooids are small to minute (< 0.4 mm in diameter; or < 1/3 the diameter of retracted autozooids) ... . Paraminabea
9. Polyparium is broadly flared and discoidal often with deep open marginal folds. Surface of polyparium smooth.

Coenenchymal sclerites are small clubs and strong spindles with irregularly arranged tubercles ..... . Sarcophyton

—Polyparium margin with numerous, often crowded, closed folds or lobes that vary from low and hemispherical to
digitiform. Surface of polyparium often with lobes or plications. Coenenchymal sclerites are spindles with tuber-
elesioltenbarrancedtinntrans Verse) cnc] Sire men meme ae mn meat eee Cec Lobophytum

WILLIAMS AND ALDERSLADE: REVISIONARY SYSTEMATICS OF MINABEA 363

ACKNOWLEDGMENTS

We are grateful to Pat and Lori Colin (Coral Reef Research Foundation, Palau), and Robert
Bolland (University of Maryland, Okinawa) for providing field collected material, and to Leen van
Ofwegen (Natural History Museum, Leiden) and Yehuda Benayahu (Tel Aviv University, Ramat
Aviv) for their helpful comments.

LITERATURE CITED

ALDERSLADE, P. 1994. A redescription of Anthomastus agilis Tixier-Durivault with a generic placement in
Sinularia (Coelenterata: Octocorallia), and some remarks on Sinularia ramosa Tixier-Durivault. The
Beagle, Records of the Museums and Art Galleries of the Northern Territory 11:133—139.

BAYER, F. M. 1974. A new species of Trichogorgia and records of two other octocorals new to the Palau Islands.
Micronesica 10:257—271.

_ 1981. Key to the genera of Octocorallia exclusive of the Pennatulacea (Coelenterata: Anthozoa), with
diagnoses of new taxa. Proceedings of the Biological Society of Washington 94:902—947.

BENHAM, W. B. 1928. On some alcyonarians from New Zealand waters. Transactions and Proceedings of the
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FAULKNER, D. AND R. CHESHER. 1979. Living corals. Clarkson N. Potter, Inc., New York. 311 pp.

GOSLINER, T. M., D. W. BEHRENS, AND G. C. WILLIAMS. 1996. Coral reef animals of the Indo-Pacific, animal
life from Africa to Hawai’i exclusive of the vertebrates. Sea Challengers, Monterey. 314 pp.

Groot, S. AND S. WEINBERG. 1982. Biogeography, taxonomical status and ecology of Alcyonium (Parerythropo-
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HICKSON, S. J. 1907. The differentiation of species of Coelenterata in shallow water seas. Report and Transactions
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KsEBATI, M. B. AND F. J. SCHMITZ. 1988. Minabeolides: a group of withanolides from a soft coral, Minabea sp.
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MILLARD, N. A. H. 1978. The geographic distribution of southern African hydroids. Annals of the South African
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MILLER, M. W. 1995. Growth ofa temperate coral: effects of temperature, light, depth, and heterotrophy. Marine
Ecology Progress Series 122:(1—3):217—225.

Potts, D.C. 1984. Generation times and the Quaternary evolution of reef-building corals. Paleobiology
10(1):48-S8.

SCHUHMACHER, H. AND J. HINTERKIRCHER. 1996. Niedere Meerestiere: Schwamme, Korallen, Krebse,
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Minchen. 320 pp.

SOLE-CAVA, A. M. AND J. P. THORPE. 1991. High levels of genetic variation in natural populations of marine
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THOMSON, J. A. AND W. D. HENDERSON. 1905. Report on the Alcyonaria collected by Professor Herdman, at
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Part 3, supplementary report 20:269-328.

THOMSON, J. A. AND D. L MACKINNON. 1910. Alcyonarians collected on the Percy Sladen Trust Expedition by
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VERON, J. E. N. 1986. Corals of Australia and the Indo-Pacific. 1986. Angus and Robertson, Publishers, North
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delingen 192:1—91.

. 1983. A revision of the genus Lobophytum von Marenzeller (Octocorallia, Alcyonacea). Zoologishe
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WILLIAMS, G. C. 1986. Morphology, sytematics, and variability of the southern African soft coral Alcyonium
variabile (J. Stuart Thomson, 1921) (Octocorallia, Alcyoniidae). Annals of the South African Museum
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. 1990. A new genus of dimorphic soft coral from the south-western fringe of the Indo-Pacific (Octoco-

rallia: Aleyoniidae). Journal of Zoology, London 221:21—35.

. 1992a. The Alcyonacea of southern Africa. Stoloniferous octocorals and soft corals. Annals of the South

African Museum 100 (3):249-358.

. 1992b. Revision of the soft coral genus Minabea (Octocorallia: Alcyoniidae) with new taxa from the

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© CALIFORNIA ACADEMY OF SCIENCES, 1999
Golden Gate Park
San Francisco, California 94118

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES

Volume 51, No. 8, pp. 365-383, 8 figs. | ale T 2 ih 1999 October 8, 1999

aoe |
Description of Three New Species of Halgerda from the
Western Indian Ocean with a Redescription of
Halgerda formosa, Bergh 1880

By
Shireen J. Fahey and Terrence M. Gosliner

Department of Invertebrate Zoology and Geology, California Academy of Sciences
Golden Gate Park, San Francisco, California 94118

Three new species of Halgerda are described from the tropical Indo-Pacific. Halgerda
dalanghita sp. nov. is known from South Africa, Papua New Guinea and the Philippines.
Halgerda dichromis sp. nov. is known from South Africa. Halgerda toliara sp. nov. is
described from Madagascar. Comparison of the three new species is made with the original
description and newly collected specimens of the type species, Halgerda formosa Bergh
1880 and other described members of the genus. The coloration, reproductive system and
the radular morphology of these new species differ significantly from H. formosa and other
previously described Halgerda species. Specimens of Halgerda formosa have been misiden-
tified as H. punctata Farran, 1905 in recent literature and are compared here. Although
these two species share some characteristics, they differ significantly in external and
radular morphology.

Bergh (1880) introduced the genus Halgerda with the type species Halgerda formosa, based on
a single specimen collected from La Reunion, and provided a description of the living animal based
on the field notes of Dr. Koerbl, the collector. According to Bergh, the external coloration of this
species 1s yellowish white with “orange-yellow stripes and black spots on its back, and black
rhinophores” (Bergh 1880). Bergh described the preserved animal as having ridges which branch
around the gills and rhinophores. His illustration of a second specimen from Mauritius (Bergh 1888,
pl. 77, fig. 10) shows distinct dorsal ridges.

Farran (1905) introduced Halgerda punctata collected from Ceylon by Professor Herdman in
1902. Both Farran’s description and illustration indicate large blunt tubercles arranged along the
dorsum in a reticulate pattern. He also stated that in a second specimen, the ridges were “almost
obsolete.” Farran cited other differences between H. punctata and H. formosa. These differences
included radular morphology, size and shape of both the foot and the anal papilla, and the number of
gill branchia. Recent workers (Rudman 1978; Gosliner 1987; Wells and Bryce 1993) have suggested
that H. punctata may be a synonym of H. formosa. The taxonomic relationships of these two nominal
species are revised in the present study.

This paper describes three new species of Halgerda, which like H. formosa, are found in the
Indian Ocean.

All material studied is deposited in The Natural History Museum of London (BMNH), the
Department of Invertebrate Zoology and Geology of the California Academy of Sciences (CASIZ)
and the South African Museum (SAM).

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SPECIES DESCRIPTIONS

Family Halgerdidae Odhner, 1926
Genus Halgerda Bergh, 1880

TYPE SPECIES. — Halgerda formosa Bergh, 1880, by monotypy.

REMARKS. — Odhner (1926) introduced the family name Halgerdidae for Ha/gerda, and Thiele
(1931) introduced Asteronotidae for Asteronotus. Odhner in Franc (1968) suggested that both family
names are synonymous and included the genera Aphelodoris, Halgerda, Sclerodoris and Asteronotus.
He employed Asteronotidae for the family despite the fact that Halgerdidae was used earlier and has
priority. Others (Willan and Coleman 1984; Wells and Bryce 1993; Rudman 1998) have united most
of the cryptobranch dorids into the single family Dorididae with or without employing a series of
subfamilies. This approach unites more than sixty genera into a single unwieldy family which does
not reflect phylogenetic relationships.

Since Halgerdidae is the older available name, we prefer to continue to place Halgerda,
Sclerodoris, Aphelodoris and Asteronotus together in Halgerdidae, pending a complete phylogenetic
analysis of the cryptobranch dorids.

Halgerda formosa Bergh, 1880
Figs. 1A—B, 2, 3A

Halgerda formosa Bergh, 1880:190—195, pl. 4, figs. 15—20; pl. 5, figs. 10-12. 1888:822—826, pl. 77, figs. 10,
11; pl. 81, figs. 13—17.

Halgerda punctata Rudman, 1978:67, figs. 3A,7,8. misidentification; not H. punctata Farran, 1905.

Halgerda punctata Gosliner, 1987:69, middle photograph. misidentification; not H. punctata Farran, 1905.

Halgerda punctata Wells and Bryce, 1993:106, top photograph. misidentification; not 1. punctata Farran, 1905.

MATERIAL EXAMINED. — BMNH Reg. No. 1998027, Acc. No. 2350, one specimen, dissected, Grand Baie,
Mauritius, 5 March 1990, H. Debelius. CASIZ 099340, one specimen, dissected, NE side Manahuanja Island,
Msimbati, Mtwara Region, Southern Tanzania, 2 m depth, 4 November 1994, T. M. Gosliner.

DISTRIBUTION. — Halgerda formosa was originally described from Reunion Island (Bergh
1880), and later reported from Mauritius (Bergh 1888). Recent collections of specimens indicate that
it is also found off Zanzibar (Rudman 1978), southern Tanzania (present study), Western Australia
(Wells and Bryce 1993) and Sodwana Bay, South Africa (Gosliner 1987).

EXTERNAL MORPHOLOGY. — The preserved animals studied (CASIZ 099340 and BMNH 2350)
measured 12 mm and 7 mm respectively. The body is firm and smooth, but rigid. The body profile
is high and the dorsum has a series of low, angled ridges arranged in a reticulate pattern. There are
no conical tubercles at the junctions of the ridges. The ground color of the dorsum and foot is whitish
with a gray tinge. The ridges are lined with yellow-orange, with shorter, thinner yellow-orange lines
in the depressions between the ridges on one specimen (CASIZ 099340) which are absent in the other.
The ridges closest to the mantle margins have no yellow-orange coloration on them. There are small
white tubercles along the mantle margin. The mantle edge has a thin, white line around the
circumference. Along the edge of the mantle and on the foot are irregularly spaced, dark brown to
black spots. The chocolate brown viscera can be seen through the translucent notum.

The upright rhinophores are tapered towards the tips, which have black coloration subapically.
The black coloration extends down the posterior side of the translucent white rhinophores as a
longitudinal band. There are 17 to 19 transverse rhinophoral lamellae on the animals included in the
present study.

FAHEY AND GOSLINER: NEW SPECIES OF HALGERDA 367

~~, £2

2 ; ee re

FiGure 1. A. Halgerda formosa Bergh 1880 (CASIZ 099340). Specimen from NE side of Manahuanja Island, Mtwara
Region, Tanzania, photograph by T. M. Gosliner. B. Halgerda formosa Bergh 1880. Specimen from Adlam’s Reef, Sodwana
Bay National Park, Natal, South Africa, photograph by T. M. Gosliner.

368 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 8

Figure 2. Halgerda formosa (CASIZ 099340). Scanning electron micrographs. A. Inner lateral teeth, scale = 20 pm; B.
Middle lateral teeth, scale = 43 jum; C. Outer lateral teeth, scale = 30 pm.

There are four tripinnate branchial leaves. The gills have black-lined branches and the pigment
encircles each branch tip. Within the gill rachis are numerous flattened, translucent structures that
appear glandular. The anal papilla is long and tubular with black coloration on both the posterior and
anterior sides. The oral tentacles are short and digitiform. Some black spots on the ventral side of the
foot are retained on the preserved animals.

BUCCAL ARMATURE. — The labial cuticle is smooth and devoid of rodlets. The radular sac is
elongate, and the radular formula of one of the two dissected specimens is: 36 x 40.0.40 (CASIZ
(99340). The inner 7-8 lateral teeth are hamate although the hook is much shorter than that of the

FAHEY AND GOSLINER: NEW SPECIES OF HALGERDA 369

middle lateral teeth (Fig. 2A). The middle lateral teeth are hamate with-a rounded point on the crest
(Fig. 2B). The 3 outer lateral teeth are modified with a series of fimbriate denticles (Fig. 2C).

REPRODUCTIVE SYSTEM. — The reproductive system is triaulic (Fig. 3A). The ampulla is long
and lies across the anterior of the female gland mass. The female gland mass is about the same size
as the prostate gland. The ampulla narrows very slightly as it enters the postampullary duct and
bifurcates into the vas deferens and oviduct. The short oviduct enters the female gland mass. The vas
deferens separates from the ampulla and widens into the large, glandular prostate. The muscular
portion of the vas deferens leaves the prostate in a long, single duct with one half-loop, then widens
into the expansive penial bulb. The short uterine duct emerges from the female gland mass and joins
the receptaculum seminis duct near its entry to the receptaculum. The receptaculum duct is long,
convoluted and joins the oval receptaculum seminis with the larger, spherical bursa copulatrix
adjacent to the vagina. The bursa is completely covered by the larger prostate. The vaginal duct, which
emerges from the base of the bursa copulatrix, is short. There is a muscular sphincter at the entry to
the vagina. The enlarged muscular region of the vagina constricts before its exit in the center of the
genital atrium which has several deep fingerlike folds. Both the penis and the vagina are unarmed.
The genital atrium is wide and large.

DISCUSSION. — Bergh (1880) described the 23-millimeter, type specimen of Halgerda formosa
as having a white, high, oval body with a grayish tinge. The ridges were joined at two or three spots
on the dorsum situated between the rhinophores and the gills. The ridges branched around the gills
and rhinophores and continued to the mantle edge (Fig. 1A). Bergh (1880) reported the branches of
the main ridges as yellowish in color, and in between the lateral ridges the bluish black coloration of
the viscera could be seen. There were brownish black spots along the mantle edge. The wide stalks
of the rhinophores were yellowish above the middle of the club, with coal black markings on the
posterior surface, whereas the anterior of the stalk is whitish. He counted 25 rhinophoral lamellae.
The small, tapering anal papilla was white with black markings on the inner surface, as were the stalks
of the gill. The three branches of the gill also had black lines along their inner surface. The foot as
described by Bergh (1880) and as found in the present study is fairly small and has black spots medially
down the length.

Bergh’s (1880) description of the radula included 51 rows of hooked teeth and 42-44 teeth per
half-row. His drawings of the teeth of H. formosa show two denticulate outer lateral teeth. In addition,
the radular sac as drawn (pl. 81, fig. 14) was long and curved.

Bergh presented a detailed description of the reproductive system. He reported that H. formosa
has a large female gland mass, a prostate that covers the upper portion of the bursa copulatrix, a
receptaculum seminis shaped like a compressed sphere and one-third to one-half the size of the bursa.
There was no vaginal or penial armature. The penis was cylindrical and three times as thick as the
sperm duct. The prostate had two distinct parts, one glandular and one long and snakelike that
terminates into the penis. The uterine duct was long and the genital vestibule had long, deep folds.

A comparison of the specimens dissected in the present study (BMNH 2350, CASIZ 099340) to
Bergh’s (1880) description indicates that both are Halgerda formosa due to the following similar
characters:

As with Bergh’s descriptions of Halgerda formosa, both specimens examined have a series of
low, angled ridges, with no tubercles at the junctions of the ridges. The ridges are lined with
yellow-orange. The body color of the two specimens is white with a grayish tinge. Along the edge of
the mantle and on the foot are irregularly spaced, dark brown to black spots. The rhinophores of the
specimens, like H. formosa are tapered and have black coloration subapically. The gills have
black-lined branches and the anal papilla is long and tubular with black coloration on both sides.

The specimen dissected (CASIZ 099349) has the same elongate radular sac and fimbriate (=
denticulate of Bergh) outer denticles as described by Bergh for Halgerda formosa.

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES

370

Volume 51, No. 8

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FAHEY AND GOSLINER: NEW SPECIES OF HALGERDA 37]

The reproductive system of both specimens (Fig. 3A) is as described by Bergh for Halgerda
formosa. The penial bulb is large, as 1s the genital atrium. The large, glandular prostate covers the
bursa copulatrix. The vaginal duct is short and emerges from the base of the bursa. Neither the vagina
nor the penis are armed.

Yonow and Hayward (1991) examined the external morphology of the specimen (BMNH 2350)
collected from Mauritius, and identified it as Halgerda formosa. Although no reproductive morphol-
ogy is described in Yonow and Hayward’s paper, the dissection was made in the present study, and
there is little doubt that the specimen is indeed H. formosa. The external morphology described by
Yonow and Hayward is nearly identical to Bergh’s (1880) description of Halgerda formosa. The
reproductive morphology of this specimen, examined for this study, is also identical to Bergh’s
description.

Farran (1905) placed Halgerda punctata in the genus Halgerda based on the following features:
a smooth, leathery mantle with raised tubercular ridges, a bi-lipped foot, small tentacles which in H.
punctata are tuberculate, a small number of branchial leaves, absence of jaws, a long curved radular
sac, outer lateral teeth not denticulate and innermost teeth smaller. There were 50 rows of hooked
teeth, with 50 teeth per each side of the row. There were no rachidian teeth and the penis and vagina
were unarmed.

Farran (1905) described the mantle of Halgerda punctata as being white with numerous tubercles,
which have a pinkish tinge on some of the apices. He noted that these tubercles were “more or less
connected by ridges in a triangular network.” Both the mantle and the ventral side of the animals he
studied had a few black spots. The notes sent with the specimens from Professor Herdman, the
collector, included mention of yellow papillae on the mantle and purple spots on the foot, with a few
spots of the same color on the mantle. Farran described the anal papilla as long and tubular, white
with a black scalloped margin.

Farran’s (1905) work comparing Bergh’s specimens of Halgerda formosa with H. punctata
confirmed Bergh’s original description of H. formosa that noted numerous dorsal ridges, crimson
spots and yellow lines. The above external differences, along with the denticulate outer lateral teeth
of H. formosa, as opposed to smooth ones in H. punctata, were the only differences Farran noted
between the two species. Farran’s illustration (pl. 3, fig. 4, ) of H. punctata is markedly different from
Bergh’s drawing of H. formosa in that Farran drew distinctive tubercles connected by low ridges
along the high body profile, whereas Bergh drew distinctive ridges with no tubercles at all on a low
body profile.

Rudman (1978) examined a specimen of Ha/gerda from Zanzibar and identified it as Halgerda
punctata Farran. The animal he described had distinct ridges with bright lemon yellow angles and
purple spots on the foot with few on the mantle. Rudman did not describe tubercles on this specimen,
although in his discussion he mentioned that his specimen was identical to Bergh’s H. formosa,
including the “yellow papillae” on the mantle (Rudman 1978). Rudman described the radula as having
smaller inner teeth and smooth, outer lateral teeth. Rudman’s illustration of the radular sac indicates
that it is long.

Rudman described the ejaculatory duct of the Zanzibar specimen as large, muscular and separate
from the vagina, and the vagina as a thin-walled bulb with a muscular sphincter at the opening of the
thin-walled vaginal duct as in our specimens of H. formosa.

Due to the external morphological similarities between Rudman’s Halgerda punctata and
Bergh’s description of H. formosa, it appears that Rudman’s (1978) specimen was actually H.
formosa. Although the outer lateral teeth in H. formosa are drawn and described as denticulate (Bergh
1880), and Rudman’s specimen are non-denticulate, radular variations within species are noted in
other recent studies of Halgerda (Bertsch and Johnson 1982; Gosliner and Fahey 1998).

Gosliner (1987) described a specimen from South Africa also as Halgerda punctata that was
similar to the Zanzibar specimen Rudman examined and identified as H. punctata. Gosliner described

B72 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 8

a translucent white color, plum colored spots on the notum, and yellow-orange ridges. Color slides
of animals (Figs. 1A, B) collected in Zanzibar and Tanzania show the spots to be arranged along the
mantle edge. While the specimen from South Africa has spots all along the mantle edge, the Tanzanian
specimen has only five spots arranged asymmetrically on its posterior end, four spots on the anterior
mantle edge, with two next to the rhinophores, and two on the extreme front. No tubercles are present
on the notum of any Halgerda collected by Gosliner from South Africa. Based on the consistent
characters between H. formosa and Gosliner’s (1987) specimen, we have determined that Gosliner’s
specimen is most likely H. formosa.

Wells and Bryce (1993) illustrated a Halgerda specimen collected from Bundegi Reef, North
West Cape of Australia, as Halgerda punctata. However, the animal pictured in that work has the
identical, external morphological characteristics of H. formosa as described by Bergh (1880). This
includes yellow-lined, low ridges, tubular anal papilla with dark pigment on the anterior side, black
markings on the rhinophores, notum and gills. The gills also have black lines along their inner surface
rather than black spots. No reproductive anatomy is described.

Based upon a comparison with Bergh’s (1880) original description of Halgerda formosa and
Farran’s (1905) original description of H. punctata, Rudman’s, Gosliner’s and Wells and Bryce’s
specimens should have all been identified as H. formosa, not H. punctata.

Debelius (1996:254, bottom photo) described an animal collected from Sri Lanka whose external
morphology is nearly identical to Halgerda punctata (Farran 1905). For example, the animal has
blunt, yellow tubercles, some with pink apices. There are dark spots on the notum, arranged somewhat
symmetrically. The rhinophores appear as described by Farran (1905): stout with black tips and thick
white bases. The four branchia also match Farran’s description as being thick with the anterior pair
larger than the posterior pair, and black on the interior, but white on the outside. Debelius did not
describe the reproductive or radular morphology of the specimen. We suggest that the specimen
pictured in Debelius (1996) is H. punctata. The animal was not collected and could not be examined
for the present study.

Rudman (1978) suggested that H. punctata may be asynonym of H. formosa. However, it appears
from the morphological differences noted in the present study, that these are two distinct species,
despite the fact that the reproductive anatomy of H. punctata has not been presented. Those
distinctions are:

¢ Halgerda formosa has distinct low ridges lined in yellow to yellow-orange. Halgerda punctata
has yellow tubercles. In H. punctata, the ridges are indistinct.

¢ Halgerda formosa has three, thick, main gill branches with the two lateral branches forked. The
black coloration wraps around the gill from approximately halfway up the rachis. Halgerda punctata
has four gill branches with the anterior two larger than the posterior pair. The gills have black margins
on the interior and are white on the outside. The black pigment covers the entire length of each gill
branch.

¢ The oral tentacles of H. formosa are short and digitiform. The tentacles of H. punctata are very
small and tubercular (Farran 1905).

¢ The anal papilla of H. formosa is squat and tapering with black marks on the inner surface. The
anal papilla of H. punctata is long and tubular with a black scalloped margin.

* Halgerda formosa has a thin foot; approximately one-fourth to one-third the width of the mantle.
Halgerda punctata has a moderately wide foot (Farran 1905).

¢ The radular formula of H. formosa is 51 x 42.0.42 (Bergh 1880), 65 x 64.0.64 (Rudman 1978)
and 36 x 40.0.40 (present study), with the outer two lateral teeth denticulate (Bergh 1880; present
study) or smooth (Rudman 1978). The radular formula of H. punctata is 50 x 50.0.50 and the three
outermost teeth are small and smooth.

FAHEY AND GOSLINER: NEW SPECIES OF HALGERDA 373

¢ The innermost teeth of Halgerda formosa have large developed hooks and the four outer teeth
are markedly smaller than the middle teeth. The 20 innermost teeth of H. punctata are very small and
gradually increase in size towards the middle lateral teeth.

Halgerda dalanghita sp. nov.
Figs. 3B, 4A, 5, 6

Sclerodoris sp. Gosliner, 1987:68, top photograph.

TYPE MATERIAL. — HoLotyPe: CASIZ 114775, Bethlehem, Maricaban Island, Batangas Province,
Luzon, Philippines, 15 m depth, 24 April 1997, T. M. Gosliner. PARATYPES: CASIZ 110373, two specimens,
one dissected, Bethlehem, Maricaban Island, Batangas Province, Luzon, Philippines, 15 m depth, 24 April 1997,
T. M. Gosliner. CASIZ 111301, two specimens, one dissected, Anemone Reef, Papua New Guinea, 20 m depth,
22 February 1992, T. M. Gosliner. SAM A35416, one specimen, dissected, Mbibi, South Africa, 5 m depth, 8
May 1982, T. M. Gosliner. SAM A35410, one specimen, dissected, Mbibi, South Africa, 5 m depth, 6 May 1981,
T. M. Gosliner. SAM A35415, one specimen, dissected, Adlam’s Reef, Sodwana Bay, Natal, South Africa. 2 m
depth, 9 May 1982. T. M. Gosliner.

DISTRIBUTION. — This species is known from Natal, South Africa, Papua New Guinea and the
Philippines (present study).

ETYMOLOGY. — Halgerda dalanghita, is named for the color of the animals. Dalanghita is the
Tagalog word for a small orange fruit.

NATURAL HISTORY. — Halgerda dalanghita is found crawling in the open in shallow water, on
near-shore reefs, rocky surfaces and also under coral rubble. It feeds on unidentified orange sponges
and has been found from 5—20 m in depth. The three specimens from the Philippines were almost
entirely embedded in a lobate orange sponge on which they had been feeding.

EXTERNAL MORPHOLOGY. — The preserved animals studied are 15-28 mm in length. The body
is firm and smooth, like gelatin, but rigid. The body profile is relatively low and the dorsum has a
series of angled ridges arranged in a reticulate pattern. There are no conical tubercles at the junctions
of the ridges, as is present in some other Halgerda species. The ground color of the dorsum and foot
is lemon yellow to orange. The ridges are lined in white, with additional short lines or dots of white
sprinkled on the dorsum among the ridges. Brown dots, which outline the ridges and are scattered
along the mantle edge, can be seen on some of the yellow to pale orange specimens. These brown
spots are not present on one of the specimens from the Philippines. The pinkish viscera can be seen
through the notum in the paler colored living animals.

The upright rhinophores are tapered towards the tip with brown coloration on the posterior side
near the base, black subapical coloration and a translucent white background color. There are 17-18
rhinophoral lamellae.

The six branchial leaves are highly pinnate. The gills have black-lined branches, and the
undersides are white. The pigment completely encircles each branch. Within the finely divided pinnate
portion of the gill rachis, near the apices, are numerous flattened, translucent structures that appear
glandular. The anal papilla is orange.

The oral tentacles are long and tapered outwards and posteriorly. Some light brown pigment spots
on the ventral side of the mantle, near the margins of the foot, are retained on the preserved animals.

BUCCAL ARMATURE. — The buccal mass is not pigmented. The radular sac is short. The radular
formulae of the five dissected specimens are: 42 x 33.0.33 (CASIZ 110373) (Fig. 5), 34 x 29.0.29
(SAM A35415), 35 x 25.0.25 (SAM A35410), 26 x 27.0.27 (SAM A35416) (Fig. 6) and 35 x 22.0.22
(CASIZ 111301). The outer and middle lateral teeth are extremely elongate, which 1s atypical of dorid
hamate teeth. It was difficult to confirm the radula formulae due to the teeth being extremely narrow,

374 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 8

numerous and irregularly arranged (Fig. 5B, D). The 5 or so inner lateral teeth are hamate (Fig. 6A,
B).

REPRODUCTIVE SYSTEM. — The reproductive system is triaulic (Fig. 3B). The wide ampulla is
flattened and lies tightly against the female gland mass. The female gland mass is much larger than
the prostate gland. The ampulla narrows into the postampullary duct, which bifurcates into the vas
deferens and oviduct. The short oviduct enters the female gland mass. The short vas deferens separates
from the ampulla and widens into the large, elongate glandular prostate. The prostate consists of two
distinct glandular types but they are not as well differentiated as in other members of Halgerda, but
are differentially pigmented on all material examined. The muscular portion of the vas deferens leaves
the prostate in a long, single duct with one half-loop, then widens slightly into the penial bulb. The
short uterine duct emerges from the female gland mass and joins the receptaculum seminis near its
base. The duct connecting the receptaculum and the bursa is moderately long. The pyriform
receptaculum seminis is equal to or much smaller than the spherical bursa copulatrix. The bursa is
not covered by the larger prostate. The receptaculum is partially hidden under the bursa and is also
slightly tucked under the prostate. The vaginal duct that emerges from the base of the bursa copulatrix
is long and thin. At its exit adjacent to the base of the male aperture, is an enlarged, but undivided
portion that is not obviously glandular. The genital aperture is wide and large.

DISCUSSION. — Gosliner (1987) placed this species in the genus Sc/erodoris Eliot, 1904.
However, the type species, S. tuberculata Eliot, 1904 possesses a notum with caryophyllidia and
broad outer teeth (Rudman 1978). The reproductive system of S. tuberculata also contains a distinct
vestibular gland. As in other species of Halgerda, the present species has a smooth notal texture,
narrow outer teeth and lacks a vestibular gland. Also, it has elongate rhinophores with black pigment,
ridges that are of a different color than the body and opaque white gill glands, as in other species of
Halgerda. Halgerda dalanghita is the only species of Halgerda with an orange general body color
with varied pigment on the notum. The middle and outer lateral teeth of the radula of H. dalanghita
are thin and elongate. The only other record of a Halgerda with similar radular teeth is that of Gosliner
and Fahey (1998). In this case, some specimens of H. elegans Bergh, 1905 have elongate teeth while
others have typical hamate teeth. All specimens of H. dalanghita examined here had elongate teeth.
The radula sac of H. dalanghita is unique in being short, although all other described species of
Halgerda have an extremely elongate sac.

The reproductive system of H. dalanghita has several distinctive features. The prostate does not
cover the bursa copulatrix and is far less obviously differentiated into two distinct portions. However,
the prostate does have two distinct gland types that are differentially pigmented in all preserved
material examined. The vaginal and ejaculatory ducts are elongate and thin throughout their length.

Halgerda dichromis sp. nov.
Figs. 3C, 4B, 7

Halgerda formosa Gosliner, 1987:69, top photograph, misidentification; not H. formosa Bergh, 1880.

Type MATERIAL. — HOLOTYPE: SAM A35413, one specimen, dissected, Vetchie’s Pier, Durban Harbor,
Natal, South Africa, 5 m depth, July 1980, W. R. Liltved.

—>

FiGuRE 4. A. Halgerda dalanghita sp. nov. (SAM A35415). Specimen from Adlam’s Reef, Sodwana Bay, Natal, South
Africa, photograph by T. M. Gosliner; B. Halgerda dichromis sp. nov. (SAM A35413). Specimen from Vetchie’s Pier, Durban
Harbor, South Africa, photograph by T. M. Gosliner; C. Halgerda toliara sp. nov. (CASIZ 111302). Specimen from 5 km SW
of Mora Mora, Madagascar, photograph by T. M. Gosliner.

FAHEY AND GOSLINER: NEW SPECIES OF HALGERDA

BS

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Volume 51, No. 8

FiGure 5. Halgerda dalanghita sp. nov. (CASIZ 110373) Bethlehem, Batangas Province, Philippines. Scanning electron
micrographs. A. Inner lateral teeth, scale = 60 tum; B. Outer lateral teeth, scale = 75 um; C. Close up of inner lateral teeth, scale
= 30 um; D. Outer lateral teeth, scale = 60 pm.

ETYMOLOGY. — Halgerda dichromis is named to distinguish its bicolored ridges as compared
to other Halgerda species, which have only one color on their ridges.

DISTRIBUTION. — The single specimen of Halgerda dichromis was found at the entrance to
Durban Harbor in South Africa.

NATURAL HISTORY. — Halgerda dichromis is found in shallow sub-tropical water.

SH,

FAHEY AND GOSLINER: NEW SPECIES OF HALGERDA

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378 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 8

EXTERNAL MORPHOLOGY. — The preserved animal is 16 mm in length. The living animal has
an oval body that is firm, smooth and rigid. Although the body profile is slightly arched, this species
does not have the high body form common in most other Halgerda. There are no tubercles at the
junctions of the low, insignificant ridges. The ground color of the dorsum is grayish white, and the
brown viscera can be seen through the skin of the notum. There is one thick orange line extending
longitudinally along the notal midline. This orange line begins in front of the rhinophores and ends
two-thirds of the way to the gill. On either side of this orange line are two heavy, equally-spaced black
lines, which begin at the base of the rhinophores. They converge behind the orange midline, and
continue as one black line up to and past the gill. There are shorter orange and black lines that form
a reticulate pattern on the notum. Small black spots are found at the mantle edge both posteriorly and
anteriorly.

The stocky rhinophores are angled posteriorly and have black coloration on their base posteriorly
and also at the tips where the color encircles the club. There is a small white patch on the club
approximately two-thirds of the way up its length.

The low, spreading gill has two lateral branches, each tripinnate. The two anterior branches are
larger and more finely divided than the two posterior branches. There is a black ridge line on top of
each branch. The underside of the gill is white. The same flattened, glandular structures reported
previously in other Halgerda (Gosliner and Fahey 1998; present study) can be seen inside each gill
rachis.

The anal papilla is short and stocky with black markings on the anterior side.

BUCCAL ARMATURE. — Since the radula is elongate, it is assumed that the radular sac was also,
but this was not verified as the buccal mass was not available for this study. The labial cuticle is
smooth and devoid of denticles. The radular formula of the holotype is 48 x 41.0.41 (Fig. 7). The first
twenty teeth are much smaller than the middle and outer lateral teeth, which gradually increase in size
towards the outer edge. The outer three teeth are fimbriate in some rows but simply pointed in others.

REPRODUCTIVE SYSTEM. — The reproductive system (Fig. 3C) is triaulic. The ampulla is long
and widens as it curves against the anterior of the female gland mass. It narrows and bifurcates into
the vas deferens and oviduct. The massive prostate has two distinct glandular parts. The female gland
mass is approximately the same size as the prostate gland. The muscular portion of the vas deferens
leaves the prostate in a long, single duct with two half-loops, then widens into the large penial bulb.
The short uterine duct emerges from the female gland mass and joins the receptaculum seminis near
its base. The duct joining the receptaculum and bursa is long and joins the pyriform receptaculum
seminis with the much larger, spherical bursa copulatrix adjacent to the vagina. The bursa is
completely covered by the much larger prostate. The vaginal duct, which emerges from the base of
the bursa copulatrix, is short relative to other species (Fig. 3). It narrows for most of its length but
widens into a short muscular portion and a larger glandular portion near its exit adjacent to the base
of the male aperture. Near the genital atrium, the vagina is large, muscular and bulbous, but not
obviously glandular. Both the penis and the vagina are unarmed. The genital atrium is wide, large and
elongate.

DISCUSSION. — Gosliner (1987) misidentified this specimen as Halgerda formosa, the type
species of the genus, due to certain similarities. Both animals have a grayish white background color
with yellow-orange ridges. Neither have tubercles at the ridge junctures and both have stocky
rhinophores with black pigmentation. There are small black spots at the mantle edge of both animals
and the gills are low and spreading with black pigmentation on the main branches.

Examination of the specimen from South Africa, and comparison to Bergh’s 1880 and 1888
descriptions of Halgerda formosa indicate that the South African specimen should be regarded as a
distinct species, H. dichromis.

FAHEY AND GOSLINER: NEW SPECIES OF HALGERDA 39)

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FiGure 7. Halgerda dichromis sp. nov. Radular morphology. A. Outer lateral teeth, scale = 40 jum; B. Inner and middle
lateral teeth, scale = 400 tm.

The following differences are noted between Halgerda formosa and H. dichromis:

1) Halgerda formosa has distinct ridges lined in yellow, while H. dichromis has indistinct ridges
and both black and yellow-orange lines on the notum.

2) The gills of Halgerda formosa are erect when fully extended, whereas the gills of H. dichromis
are spread to form a flat gill.

3) Halgerda formosa has a distinct muscular sphincter at the point where the vagina widens prior
to entering the genital atrium, whereas H. dichromis has no sphincter.

4) Halgerda formosa has a prostate that only covers the bursa copulatrix. The prostate of H.
dichromis completely envelops the bursa, the anterior portion of the receptaculum and the oviduct.

5) The ejaculatory duct of Halgerda formosa is long, slender and terminates into a bulbous penis.
The penis has a separate opening that terminates near the center of the wide genital atrium. The
ejaculatory duct of H. dichromis is wide, then narrows before it terminates into a large, bulbous penis
which itself terminates into a very long, wide genital atrium.

6) Halgerda formosa has a radular formula of 51 x 42.0.42 (Bergh 1880), 65 x 64.0.64 (Rudman
1978) and 36 x 40.0.40 (present study), whereas H. dichromis has a formula of 48 x 41.0.41.

7) Halgerda dichromis has the inner 20 teeth much smaller than subsequent ones. Halgerda
formosa has the inner 7—8 teeth smaller than the middle lateral teeth.

8) The gills of H. dichromis have a black medial line along the entire length of the gill as in H.
wasiniensis Eliot, 1904 (Gosliner 1987:68, bottom photo) while in H. formosa, black pigment is
restricted to the apices of the gill branches.

Although some of the external characteristics of Halgerda dichromis are similar to those of H.
formosa, the two are clearly different species. Since many of the characteristics of H. dichromis are
similar to other members of the Halgerda genus, a testable hypothesis of phylogeny is needed to
determine how closely the species are related.

380 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 8

Halgerda toliara sp. nov.
Figs. 3D, 4C, 8

TYPE MATERIAL. — HOLOTYPE: CASIZ 111302, one specimen, 5 kilometers west of Mora Mora, Mada-
gascar, 2 m depth, 7 April 1989, T. M. Gosliner. PARATYPES: CASIZ 073278, one specimen, off Soanambo
Hotel, Ile Saint Marie, Madagascar, 2 m depth, 4 April 1990, T. M. Gosliner. CASIZ 073272, 2 specimens, one
dissected, Patch Reef, Mora Mora, Madagascar, 2 m depth, 27 March 1990, T. M. Gosliner.

ETYMOLOGY. — Toliara is the Malagasy name for the Tulear region of Madagascar, where this
species was first collected.

DISTRIBUTION. — Halgerda toliara has been found on both the west and east coasts of Mada-
gascar (present study). :

NATURAL History. — Halgerda toliara was found crawling in the open in shallow water on
coral rubble.

EXTERNAL MORPHOLOGY. — The preserved animals are 9 mm (CASIZ 073272 and CASIZ
111300) and 5 mm (CASIZ 073278). The relatively low-profiled body is oval and the texture is
smooth and firm. The ground color of the living animal is white to pale yellow-white. The white
viscera can be seen through the notum of the living animals. There are low but distinct ridges arranged
in a reticulate pattern on the dorsum. There are no tubercles at the junctions of the orange-lined ridges.
Small yellow tubercles are scattered densely along the mantle margin. There is a thin white border
along the mantle edge.

The long rhinophores have a base that 1s narrower than the top one-third of the club. The black
coloration is found only at the top one-third, and it encircles the club. The bottom two-thirds of the
rhinophores 1s translucent white.

The large gill has four sparsely pinnate branches, with the posterior pair divided into two on the
paler colored animals. Black coloration is found on the top one-half to one-third of the gill branches,
with no pigmentation on the apices on the paler colored animals. The background color is translucent
white. Inside the gill rachis can be seen the same flat, glandular-like structures found in other Halgerda
species as noted in the present study and in previously described species (Gosliner and Fahey 1998).

The anal papilla is short and white in color. The wide foot, which has a white edge, extends
posteriorly beyond the mantle margin. The oral tentacles are short and digitiform.

BUCCAL ARMATURE. — The buccal mass is not pigmented, and the radular sac is elongate and
lies flat against the buccal bulb. The radular formula of the dissected specimen is 43 x 30.0.30 (CASIZ
073272) (Fig. 8). The inner four teeth are slightly smaller than the middle lateral teeth, and all teeth
are hamate. The outer three lateral teeth are fimbriate.

REPRODUCTIVE SYSTEM. — The reproductive system is triaulic (Fig. 3D). The wide ampulla is
flattened and lies against the female gland mass. The female gland mass is larger than the prostate
gland. The ampulla narrows into the postampullary duct, bifurcates into the vas deferens and oviduct.
The short oviduct enters the female gland mass. The short vas deferens separates from the ampulla
and widens into the large, two-part glandular prostate that folds onto itself. The muscular portion of
the vas deferens leaves the prostate in a long, single duct with one half-loop, then widens into the
short, wide penial bulb. The short uterine duct emerges from the female gland mass and joins the
receptaculum seminis its base. The receptaculum duct is moderately long and joins the round
receptaculum seminis with the much larger, spherical bursa copulatrix adjacent to the vagina. The
prostate lies alongside the bursa, which is approximately the same size. The top of the receptaculum
is partially hidden under the bursa and is also slightly tucked under the prostate. The vaginal duct,
which emerges from the base of the bursa copulatrix, is long and thin. It does not enlarge at its exit
adjacent to the base of the male aperture. The common genital atrium is not as wide as the penis.

FAHEY AND GOSLINER: NEW SPECIES OF HALGERDA 381

Ficure 8. Halgerda toliara sp. nov. (CASIZ 073272) Mora Mora, Madagascar. Scanning electron micrographs. A. Inner
lateral teeth, scale = 20 pm; B. Outer lateral teeth, scale = 15 um; C. Middle lateral teeth, scale = 25 um.

DISCUSSION. — Halgerda toliara shares some common characteristics with other species of the
genus. It has a firm, smooth, and rigid body with a pattern of low ridges on the mantle arranged ina
reticulate pattern. There are no large tubercles on the dorsum of Halgerda toliara as is common in
many other members of the genus. Its body profile is relatively low like H. dalanghita and H.
dichromis. There is a large gill with branchial leaves that appear to have internal glands. Halgerda

382 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 8

toliara has four sparsely pinnate branchia, unlike other species (e.g., H. formosa) with four highly
pinnate branchia.

As with all other members of the genus except Halgerda dalanghita, H. toliara has an elongate
radula sac. However, the sac does not protrude away from the buccal mass, but lies flat against It.

The rhinophores of most Halgerda are stocky at the base, and tapering at the tips, with the
lamellae extending the length of the stalk. In contrast, the stalks of the rhinophores of H. toliara have
a smooth, consistent diameter two-thirds of their length. The stalk then widens at the lamellae before
tapering to the apices. In addition, H. foliara is unique among described species in that the rhinophore
base is devoid of black pigment while the rhinophore club is entirely black. The anal papilla of some
Halgerda has black pigmentation on either or both the posterior and anterior sides. H.toliara has only
the white ground color.

The reproductive system of Halgerda toliara shares some characters, such as the round bursa
copulatrix, with other Halgerda (H. formosa, H. dalanghita, and H. dichromis). The long, thin vagina
is common to H. dalanghita and H. albocristata (Gosliner and Fahey 1998). The bulbous penis is
common to H. formosa and H. albocristata. The prostate of other Halgerda, including H. formosa
covers the bursa copulatrix (Willan and Brodie 1989; Carlson and Hoff 1993). The prostate of H.
toliara has two parts, 1s long and folded and lies next to the bursa, similar to H. dalanghita.

There are distinct differences between Halgerda toliara and other Halgerda species with
yellow-orange lined ridges. For example, there are significant reproductive anatomical differences
between H. toliara, H. wasinensis, and H. formosa. Halgerda toliara has a prostate which does not
cover the bursa copulatrix whereas H. formosa has a prostate which covers the bursa as in H.
wasinensis (Rudman, 1978). Halgerda toliara has a long, thin vaginal duct that is longer than the
ejaculatory duct. Halgerda formosa has a short vaginal duct that is much shorter than the ejaculatory
duct. Halgerda wasinensis has a thin vaginal duct that is the same length as the ejaculatory duct. There
is a muscular sphincter at the entry to the vagina in H. formosa, that is absent in H. toliara. Halgerda
wasinensis has a muscular vagina, but no sphincter is apparent in Rudman’s (1978) drawing of the
reproductive system. Halgerda formosa has a much larger, bulbous penis than does H. foliara.
Halgerda formosa also has long, fingerlike folds in the vaginal atrium, which H. toliara does not have
in the long, thin vagina.

In addition, the radular morphology differs between the species with yellow-orange lined ridges.
The inner four teeth of H. toliara are similar in size whereas the first 25 teeth of H. wasinensis are
similar in size and the seven inner teeth of H. formosa are similar.

The external morphology differs between these species with yellow-orange ridges as well. Both
H. formosa and H. wasinensis have dark markings on the notum and the foot, whereas H. toliara has
no dark spots. H. toliara has small yellow tubercles along the mantle edge, and H. formosa has white
tubercles. The gill branches of H. formosa and H. wasinensis are highly pinnate, but the gill branches
of H. toliara are sparsely pinnate.

A discussion of generic characters has been treated by previous authors (Rudman 1978; Willan
and Brodie 1989; Gosliner and Fahey 1998) and will not be repeated here.

From this study, it appears that many of the characteristics of Ha/lgerda toliara are similar to
other members of the Halgerda genus and a testable hypothesis of phylogeny is needed to determine
how closely these species are related.

ACKNOWLEDGMENTS

The authors are grateful to Heinke and Peter Schultz who kindly assisted with the translation of
Bergh’s original description of Halgerda formosa. Angel Valdés provided some of the scanning
electron micrographs (SEMs) for H. formosa. He also reviewed the manuscript and provided excellent
suggestions. Bill Liltved collected the specimen of H. dichromis. Dong Lin of the California Academy

FAHEY AND GOSLINER: NEW SPECIES OF HALGERDA 383

of Sciences photography department provided the final production of the black and white SEMs. The
two reviewers provided very thoughtful and helpful suggestions to improve the manuscript, and we
are grateful to them for their effort.

LITERATURE CITED

BERGH, R. 1880. Beitrage zur Kenntniss der japanischen nudibranchien. I. Verhandlungen der Koniglich-Kais-
erlich Zoologisch-botanischen Gesellschaft in Wien 30:155—200.

. 1888-9. Malacologische Untersuchungen, nudibranchien vom Meer der Insel Mauritius Reisen im
Archipel der Philippinen. Semper Section 2, 3(16):755—872.

. 1905. Die Opisthobranchia der Siboga Expedition. Siboga Expedition Reports 50:1—248.

BERTSCH, H. AND S. JOHNSON. 1982. Three new species of dorid nudibranchs (Gastropoda: Opisthobranchia)
from the Hawaiian Islands. The Veliger 24(3):214-218.

CARLSON, C. H. AND P. J. Horr. 1993. Three new Halgerda species. (Doridoidea: Nudibranchia: Opisthobran-
chia) from Guam. The Veliger 36(1):16—26.

DEBELIUS, H. 1996. Nudibranchs and sea slugs Indo-Pacific field guide. IKAN-Unterwasserarchiv, Frankfurt.
321 pp.

ELiot, C. 1904. On some nudibranchs from East Africa and Zanzibar. Part III. Proceedings of the Zoological
Society of London 2:371—374.

FARRAN, G. P. 1905. Report to the government of Ceylon on the pearl oyster fisheries of the gulf of Manaar
Supplementary Reports, Ceylon Pearl Oyster Fisheries No. 21:332—364.

FRANC, A. 1968. Sous-classe des Opisthobranchs. /n Traité de Zoologie. Anatomie, systématique, biologie:
Mollusques, gastéropodes et scaphopodes, Grasse, ed. 5(3):608-893. Masson et Cie, Paris.

GOSLINER, T. M. 1987. Nudibranchs of Southern Africa. Sea Challengers, Monterey, California; Jeff Hamann,
El Cajon, California and E. J. Brill, Leiden, Netherlands. 136 pp.

GOSLINER, T. M. AND S. J. FAHEY. 1998. Description of a new species of Halgerda from the Indo-Pacific with
a redescription of Halgerda elegans Bergh, 1905. Proceedings of the California Academy of Sciences
50(15):347-359.

ODHNER,N. H. J. 1926. Die Opisthobranchien. /n Further zoological results of the Swedish Antarctic Expedition
1901—1903 under direction of Dr. Otto Nordenskjold 2(1):1—100, pls. 1-3.

RUDMAN, W.B. 1978. The dorid opisthobranch genera Halgerda Bergh and Sclerodoris Eliot from the
Indo-West Pacific. Zoological Journal of the Linnean Society 68:59—-87.

. 1998. Suborder Doridina. Pp. 990-1001 in Mollusca: The southern synthesis. Fauna of Australia. Vol.
5, Part B viii. P. L. Beesley, G. J. B. Ross and A. Wells, eds. CSIRO Publishing, Melbourne.

THIELE, J. 1931. Handbuch der systematischen Weichtierkunde. Verlag. Teil 2. pp. 377-788.

WELLS, F. E. AND C. W. Bryce. 1993. Sea slugs and their relatives of Western Australia. Western Australian
Museum. 184 pp.

WILLAN, R. C. ANDG. A. BRODIE. 1989. The nudibranch Halgerda aurantiomaculata (Allan, 1932) (Doridoidea:
Dorididae) in Fijian Waters. The Veliger 32(1):69—80.

WILLAN, R. C. AND N. COLEMAN. 1984. Nudibranchs of Australasia. Australian Marine Photographic Index,
Sydney.

Yonow, N. AND P. J. HAYWARD. 1991. Opisthobranches de l’ile Maurice, avec la description de deux especes
nouvelles (Mollusca: Opisthobranchia). Revue Frangaise d’ Aquariologie 18(1):1—30.

© CALIFORNIA ACADEMY OF SCIENCES, 1999
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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES

Volume 51, No. 9, pp. 385-395, 6 figs. October 22, 1999

A Review of the Southeast Asian Catfish Genus Ceratoglanis
(Siluriformes: Siluridae), with the Description of a

New Species from Thailand
OCT 25 1999
by
Ng Heok Hee
Department of Biological Sciences, National University of Singapore
10 Kent Ridge Crescent, Singapore 119260

The poorly-known Southeast Asian silurid genus Ceratoglanis is reviewed. Previously
considered monotypic, a second species of Ceratoglanis from the southeastern Chao Phraya
and Mekong drainages in mainland Southeast Asia is described here as C. pachynema new
species. Previously unreported sexual dimorphism in the morphology of the pectoral spines
and maxillary barbels is described.

The diverse Eurasian catfish family Siluridae includes nearly 100 extant nominal species, and is
represented in Southeast Asia by putatively 18 genera (Bornbusch 1995), of which the genus
Ceratoglanis Myers, 1938 comprises moderately large silurid catfishes (attaining standard lengths of
up to 500 mm) found in large rivers. The genus has been recorded from mainland Southeast Asia,
Java, Borneo and Sumatra and only one species 1s currently recognized, viz. C. scleronema (Bleeker,
1862). In reviewing the genus, it was found that specimens of Ceratoglanis from the Chao Phraya
drainage, Thailand, previously assigned to C. scleronema represent a second, undescribed species
here described as C. pachynema new species. This study also documents previously unrecorded sexual
dimorphism in the shape of the maxillary barbels in Ceratoglanis species.

MATERIALS AND METHODS

Measurements were made point to point with dial calipers and data recorded to tenths of a
millimeter. Counts and measurements were made on the left side of specimens whenever possible.
Subunits of the head are presented as proportions of head length (HL). Head length and measurements
of body parts are given as proportions of standard length (SL). Measurements and counts were made
following Ng and Ng (1998). The ventral angle of the snout is taken as the angle between a vertical
line passing through the tip of the snout and the tangent of the ventral half of the upper jaw (Fig. 1).
This angle was measured from drawings of the lateral views of the heads.

Fin-ray counts were obtained under a binocular dissecting microscope using transmitted light.
Numbers in parentheses following a particular fin-ray, branchiostegal-ray or gill-raker count are the
numbers of examined specimens with that count. Vertebral counts were taken from radiographs
following the method of Roberts (1994). Numbers in parentheses following a particular vertebral
count are the numbers of radiographed specimens with that count.

Drawings of the specimens were made with a Nikon SMZ-10 microscopic camera lucida.
Institutional codes follow Eschmeyer (1998). A listing of the material examined for this study can be
found under the accounts of the appropriate species.

385

386 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 9

FiGure |. Lateral view of head of Ceratoglanis, showing how the ventral angle (V) of the snout is measured.

SPECIES DESCRIPTIONS

Ceratoglanis Myers, 1938

Ceratoglanis Myers 1938:98; Haig, 1952:65, 95; Roberts, 1989:144.
Hemisilurus (in part) Bleeker, 1862:93; 1863a:116; Weber and de Beaufort, 1913:210.

TYPE SPECIES. — Hemisilurus scleronema Bleeker 1862, by original designation and monotypy.

DIAGNOSIS. — Ceratoglanis 1s characterized by a combination of the following apomorphies
(after Bornbusch and Lundberg 1989): (1) snout enlarged and bulbous, extending beyond gape; mouth
subterminal; (2) posterior ramus of fourth transverse process broadly expanded by lamina that extends
onto anterior ramus, rami separated by small indentation; (3) mandibular sensory canal enlarged in
angularticular and dentary; and (4) maxillary barbel short, stiffened by cartilaginous rod, tip of barbel
retrorsely hooked.

NG HEOK HEE: REVIEW OF THE GENUS CERATOGLANIS 387

REMARKS. — Ceratoglanis and Hemisilurus are sister groups that comprise a monophyletic
group within the Siluridae (Bornbusch and Lundberg 1987; Bornbusch 1995) and possess the first
three synapomorphies listed above.

Ceratoglanis shows the typical sexual dimorphism in the pectoral spine and articulated segments
reported in the silurid genera Hemisilurus by Bornbusch and Lundberg (1989), Ompok by Inger and
Chin (1962), Pterocryptis by Bornbusch (1991), Silurichthys by Ng and Ng (1998) and Silurus by
Kobayakawa (1989). The males have distinct serrations on the pectoral spine and the articulated
segments; in females the serrations are either very small and indistinct, or are absent altogether. In
addition, Ceratoglanis also exhibits sexual dimorphism in the shape of the maxillary barbels
(previously only known in Hemisilurus, its sister taxon), which is reported here for the first time. The
males of Ceratoglanis have membranous extensions on the anterior and (in C. pachynema) the
posterior edges of the maxillary barbels, whereas in the females, these extensions are smaller (in C.
pachynema) or absent (in C. scleronema). The role this dimorphism plays in the biology of
Ceratoglanis is unclear.

The historical biogeography of the two Ceratoglanis species is probably similar to that proposed
for Hemisilurus moolenburghi + H. mekongensis lineage by Bornbusch and Lundberg (1989), 1.e.,
during the Pleistocene, the ancestral Ceratoglanis was dispersed throughout the North Sunda River
system (to occupy branches in Borneo and Sumatra) and the South Indo-China River system.
Post-Pleistocene isolations of the two river systems then led to the divergence of the two species.

Ceratoglanis scleronema (Bleeker, 1862)
Figs. 2, 5a, 6a—b

Hemisilurus scleronema Bleeker, 1862:93, pl. 101 (type locality: Krawang, Java); 1863b:370; 1863c:74; Weber
and de Beaufort, 1913:214, fig. 86; Hardenberg, 1934:306.

co

FIGURE 2. Ceratoglanis scleronema, ZRC 41896, 196.0 mm SL; Sumatra: Batang Hari basin.

388 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 9

Ceratoglanis scleronema Boulenger, 1894:247; Vaillant, 1902:49; Haig, 1952:65, 95; Tweedie, 1952:84; Imaki
et al., 1981:40, fig. 69; Zakaria-Ismail, 1984:25; Bornbusch and Lundberg, 1989:441 (in part); Roberts,
1989:144; Kottelat et al., 1993:68, pl. 32 (in part); Lim et al., 1993:7; Kottelat and Lim, 1995:239.

MATERIAL EXAMINED. — Java: RMNH 2918 (1, holotype, 352 mm SL) Krawang, Bleeker
collection. Borneo: CAS 49387 (3, 228.4-331.2 mm SL) Kalimantan Barat, Sintang market, 19 Jul
1976, T. R. Roberts and S. Wirjoatmodjo. ZRC 30810 (2, 270.0-286.4 mm SL) Sarawak, Marudi
market; 18—20 Jun 1994, M. Kottelat and T. Tan. ZRC 40037 (1, 440 mm SL) Kalimantan Selatan,
Muara Teweh, Pasar Pendopo (fish market), 3-4 Jun 1996, H. H. Ng and O. Chia. ZRC 42654 (4,
236.7-290.7 mm SL) Kalimantan Barat, Kabupaten Sanggau, market at Sanggau (0°7.29'N,
110°35.94’E), 21-27 Apr 1998, H. H. Tan et al. ZRC 42739 (1, 343 mm SL) Sarawak, market in
Kapit, 28 Feb-3 Mar 1998, H. H. Tan and R. Kerle. Sumatra: USNM 93287 (1, 356 mm SL) Riau,
Mandau river, Siak, 10 Jan 1934, W. C. Abbott. ZMA 121.819 (1,216.6 mm SL) Jambi, Batang Hari,
1909, P. H. Moolenburgh. ZRC 38981 (2, 158.1-379 mm SL) Jambi, Pasar Angso Duo (fish market),
Jun 1995, P. K. L. Ng et al. ZRC 41531 (4, 340-420 mm SL) Jambi, Pasar Angso Duo (fish market),
23-29 Jul 1997, H. H. Tan and H. H. Ng. ZRC 41896 (1, 196.0 mm SL) Jambi, Pasar Angso Duo
(fish market), 21-28 Nov 1996, H. H. Tan et al. Peninsular Malaysia: ZRC 11985 (1, 406 mm SL)
locality and collector unknown.

DIAGNOSIS. — A species of Ceratoglanis having a snout with a steep ventral profile (ventral
angle of snout 20°-26°), relatively more anal-fin rays (98-108), and smaller eyes (eye diameter
10.7—-14.0 %HL) that are set further apart (interorbital distance 35.4-40.9 %HL). Mature males of
this species have membranous extensions on only the anterior edge of the maxillary barbel.

DESCRIPTION. — Head and body laterally compressed. Dorsal profile slightly convex, rising
steeply from origin of epaxial muscle on skull roof to dorsal-fin pterygiophore. Snout prominent and
bulbous, with a steep ventral profile (ventral angle of snout 20°-26°).

Anterior pair of nostrils tubular and anteriomedial to base of maxillary barbel. Posterior pair of
nostrils located between anterior border of eye and base of maxillary barbel.

Mouth subterminal; gape horizontal. Well-developed rictal fold present, consisting of large and
fleshy upper lobe joined at corner of mouth with lower lobe, which is subtended by a submandibular
groove.

Teeth villiform. Dentary teeth in slightly curved, elongate bands narrowing posteriorly, reaching
from symphysis almost to mouth corners; premaxillary teeth in a single, slightly curved, rectangular
band formed from the fusion of two similarly-shaped patches on both sides and an anterior median
triangle patch. Vomerine teeth in two separate round patches.

Eyes small, subcutaneous; located in anterior half of head; visible dorsally, but not ventrally.

Gill membranes separate and overlapping, free from isthmus. Branchiostegal rays 9(9) or 10(4).
Gill rakers short, 2 + 10 = 12(1) or 2 + 11 = 13(1).

Maxillary barbel short, stiffened by cartilaginous rod and retrorsely hooked. Membranous
extensions on the anterior edge of the maxillary barbel present in males. Females with simple barbels
without such membranous extensions.

Distal margin of pectoral fin broadly convex, with I,13,i (2), I,14 (5), 1,14,1 (1), L15 (), en
(1) or 1,16 (3) rays. Proximal two-thirds of first pectoral-fin element co-ossified into a spine without
anterior and distal denticulations. Pectoral spine and articulated segments sexually dimorphic in
mature individuals. Males with spine broad and somewhat flattened dorsoventrally, with 11-16
distinct posterior serrae, increasing in size distally; proximal articulated segments with 9-10 well-de-
veloped posterior serrae; distal-most segments have smaller serrae. Females or juveniles with spine
slender and without serrae; proximal articulated segments with 6 minute posterior serrae, distal
segments without serrae. Distal margin of pelvic fin convex, with i,7 (13) rays. Distal margin of anal
fin straight, with 98(1), 99(1), 100(1), 101(1), 102(3), 104(2), 106 (2), 107(1) or 108(1) rays; posterior

NG HEOK HEE: REVIEW OF THE GENUS CERATOGLANIS 389

margin separate from caudal fin. Integument over anal fin thickened proximally for slightly more than
half of ray lengths; fin-ray erector muscles extending along anterior edges of anal-fin rays. Caudal
fin forked, lobes equal; principal rays 8/9 (13). Urogenital papillae of both sexes located immediately
posterior to insertions of pelvic fins.

In %SL: head length 14.4-17.0, head width 7.9-9.9, head depth 8.9—10.6, preanal length
26.3-30.2, prepelvic length 23.3—27.6, prepectoral length 15.2—18.0, body depth at anus 15.6—20.4,
depth of caudal peduncle 4.9—S.8, pectoral-spine length 6.9—9.1, pectoral-fin length 11.3—14.1,
pelvic-fin length 4.0—6.0, length of anal-fin base 62.5—71.2, caudal-fin length 12.3—15.5; in %HL:
snout length 27.2—31.9, interorbital distance 35.4-40.9, eye diameter 10.7—14.0. Vertebrae 12 + 50
= 62(1), 12 + 51 = 63(3), 12 + 52 = 64(3) or 12 + 53 = 65(1).

COLORATION. — Flanks and thickened integument over anal fin diffusely pigmented with
scattered melanophores; a thin line of melanophores more densely concentrated along the distal edge
of the thickened integument over anal fin. Dorsal surfaces and sides of head gray, with relatively
dense aggregations of melanophores. Melanophores less dense ventral to level of the eye; ventral
surfaces of head and belly unpigmented with scattered melanophores. Ventral surface of head slightly
orange in life. Maxillary barbels unpigmented.

Interradial membrane of pectoral fin diffusely pigmented with gray, with black distal margin.
Pelvic fin with interradial membrane diffusely pigmented with gray along margin. Anal fin with
interradial membrane diffusely pigmented with gray along distal two-thirds, becoming denser along
the distal edge. Interradial membranes of caudal-fin lobes diffusely pigmented with gray along distal
three-quarters; melanophores absent from proximal quarter of lobes and particularly dense at tips of
lobes.

DISTRIBUTION. — This species is known from the Citarum drainage in Java, the Batang Hari and
Siak drainages in Sumatra, the Pahang drainage in Peninsular Malaysia and the Baram, Barito,
Kapuas, and Rejang drainages in Borneo (Fig. 4).

REMARKS. — Ceratoglanis scleronema can be distinguished from C. pachynema in having a
snout with a steep (vs. gently sloping) ventral profile (ventral angle of snout 20°—-26° vs. 40°-42°,
Fig. 5). Mature males of C. scleronema have a small membranous extension of the maxillary barbel
which is present along the anterior edge of the barbel (vs. larger membranous extensions present along
both the anterior and posterior edges of the barbel in C. pachynema; Fig. 6).

The single specimen examined from the Malay Peninsula (406 mm SL) differs from all other
specimens examined from Sumatra, Java and Borneo in having the bony surface of the toothplates
covered with transverse plicae and minute peglike teeth embedded in the plicae, with the teeth rather
sparsely distributed throughout the toothplates (vs. bony surface of the toothplates exposed, with
many densely-packed rows of villiform teeth). Furthermore, the vomerine toothplate of the specimen
from Peninsular Malaysia is in the form of a single ovoid patch (vs. two, clearly separate, rounded
patches in other specimens examined).

The differences in the morphology of the toothplates seem to indicate that the specimen from the
Malay Peninsula is not conspecific with those from Sumatra, Java and Borneo. However, there are
no other significant morphological differences between the specimen from the Malay Peninsula and
those from Sumatra, Java and Borneo. Furthermore, no exact locality information exists for the
specimen in question and the large size of the specimen raises the possibility that the differences
observed might be due to ontogenetic changes. The largest specimen from Sumatra (ZRC 41531, 420
mm SL) lacks a plicate toothplate. We are not sure if all big specimens possess a plicate toothplate
because we lack a large series of big specimens to fully understand ontogenetic changes in this species.
Based on these arguments, it would be more prudent to tentatively consider the population from the
Malay Peninsula as being conspecific with those from Sumatra, Java and Borneo, until more
specimens from the Malay Peninsula are available.

390 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 9

Little is known about the biology of this species, but C. sc/eronema appears to be predatory in
nature, feeding on aquatic invertebrates as do most other medium-sized silurids. Crabs (Neorhyn-
choplax prima; family Hymenosomatidae), isopods, insect remains and remains of a small fish were
found in the stomach of a dissected specimen (ZRC 41531, 420 mm SL) from the Batang Hari
drainage.

Ceratoglanis pachynema n. sp.
Figs. 3, 5b, 6c—d

Ceratoglanis scleronema (non Bleeker) Smith, 1945:339; Suvatti, 1950:286; 1981:83; Bornbusch and Lundberg,
1989:441 (in part); Kottelat, 1989:14 (in part); Nabhitabhata, 1991:187; Kottelat et al., 1993:68 (in part);
Vidthayanon et al., 1997:44.

Hemisilurus scleronema (non Bleeker) Smith, 1931:181; Suvatti, 1936:71.

TYPE MATERIAL. — HOLOTYPE: CAS 96577 (1, 266.9 mm SL) Thailand: Prachinburi market,
28 May 1991, T. R. Roberts. PARATYPES: CAS 92882 (1, 203.0 mm SL) Thailand: Prachinburi
market, 8—10 Sep 1990, T. R. Roberts. CAS 94185 (1, 128.2 mm SL) Thailand: Prachinburi market,
4-7 Dec 1990, T. R. Roberts. CAS 96576 (2, 218.7—279.0 mm SL) Thailand: Prachinburi market,
8—10 Sep 1990, T. R. Roberts.

DIAGNOSIS. — A species of Ceratoglanis with a snout with a gently sloping ventral profile
(ventral angle of snout 40°-42°), relatively fewer anal-fin rays (90-101), and larger eyes (eye diameter
13.0-16.4 %HL) set more closely together (interorbital distance 33.7—-36.6 %HL). Mature males of
this species have membranous extensions on both the anterior and posterior edges of the maxillary
barbel.

IGURE 3. Ceratoglanis pachynema, CAS 96576, 218.7 mm SL; Thailand: Chao Phraya basin.

NG HEOK HEE: REVIEW OF THE GENUS CERATOGLANIS 39]

FiGuRE 4. Map showing distribution of Ceratoglanis species: C. scleronema (@) and C. pachynema (4). Open symbols
indicate records from literature.

392 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 9

DESCRIPTION. — Head and body later-
ally compressed. Dorsal profile slightly con-
vex, rising steeply from origin of epaxial
muscle on skull roof to dorsal-fin pterygio-
phore. Snout prominent and bulbous, with a
gently sloping ventral profile (ventral angle of
snout 40°-42°).

Anterior pair of nostrils tubular and ante-
riomedial to base of maxillary barbel. Poste-
rior pair of nostrils located between anterior
border of eye and base of maxillary barbel.

Mouth subterminal; gape horizontal.
Well-developed rictal fold present, consisting
of large and fleshy upper lobe joined at corner
of mouth with lower lobe, which is subtended
by a submandibular groove.

Teeth villiform. Dentary teeth in slightly
curved, elongate bands narrowing posteriorly,
reaching from symphysis almost to mouth
corners; premaxillary teeth ina single, slightly
curved, rectangular band formed from the fu-
sion of two similarly-shaped patches on both
sides and an anterior median triangle patch.
Vomerine teeth in two separate round patches.

Eyes small, subcutaneous; located in an-
terior half of head; visible dorsally, but not
ventrally.

IGURE af schematic illustration of the cae oe Gill membranes separate and overlap-
2) Cerra clerner, ZRC 28010, 2864 nm SL») ping, fre from isthmus. Branchiostegal rays
—— 9(1), 10(3) or 11(1). Gill rakers short, | + 10

= 11(1) or2 +11 = 13(1).

Maxillary barbel short, stiffened by cartilaginous rod and retrorsely hooked. Membranous
extensions on the anterior and posterior edges of the maxillary barbel present in males. Females with
simple barbels having smaller extensions on the anterior edges of the maxillary barbels.

Distal margin of pectoral fin broadly convex, with 1,14 (3), 1,15 (1), 1,16 (1); pelvic i,7 (5) rays.
Proximal two-thirds of first pectoral-fin element co-ossified into a spine without anterior and distal
denticulations. Pectoral spine and articulated segments sexually dimorphic in mature individuals.
Males with spine broad and somewhat flattened dorsoventrally, with 10 distinct posterior serrae,
increasing in size distally; proximal articulated segments with 17 well-developed posterior serrae;
distal-most segments have smaller serrae. Females or juveniles with spine slender and without serrae;
proximal articulated segments with 6 minute posterior serrae, distal segments with 5 minute serrae.
Distal margin of pelvic fin convex, with i,7 (5) rays. Distal margin of anal fin straight, with 90(1),
96(1), 97(1), 98(1) or 101(1) rays; separate from caudal fin. Integument over anal fin thickened
proximally for slightly more than half of ray lengths; fin-ray erector muscles extending along anterior
edges of anal-fin rays. Caudal fin forked, lobes equal; principal rays 8/8 (1) or 8/9 (4). Urogenital
papillae of both sexes located immediately posterior to insertions of pelvic fins.

In %SL: head length 15.5-18.8, head width 8.8-11.7, head depth 9.9-13.1, preanal length
27.2-34.7, prepelvic length 24.3-31.0, prepectoral length 16.0—-19.6, body depth at anus 16.4—22.3,
depth of caudal peduncle 4.1—6.1, pectoral-spine length 8.2—10.8, pectoral-fin length 12.6—-15.6,

NG HEOK HEE: REVIEW OF THE GENUS CERATOGLANIS 393

pelvic-fin length 4.5—6.9, length of anal-fin
base 63.7-68.3, caudal-fin length

a b
11.9-15.3; in %HL: snout length 30.7-31.5,
interorbital distance 33.7—36.6, eye diame-
ter 14.0-16.4. Vertebrae 12 + 51 = 63(1),
13 + 50 = 63(3) or 12 + 52 = 64(1).
COLORATION. — Flanks and thickened
integument over anal fin diffusely pig-
mented with scattered melanophores; a thin
line of melanophores more densely concen-
Cc \ d
“Y

trated along the distal edge of thickened
integument over anal fin. Dorsal surfaces
and sides of head gray, with relatively dense
aggregations of melanophores. Melanopho-
res less dense ventral to level of the eye;
ventral surfaces of head and belly unpig-
mented with scattered melanophores. Max-
illary barbels unpigmented.

Interradial membrane of pectoral fin
diffusely pigmented with gray, with a black

distal margin. Pelvic fin with interradial
FIGURE 6. Schematic illustration of the maxillary barbels of: a) : : :
nt Ta
Ceratoglanis scleronema, male, ZRC 41531, 378mm SL; b) C. membrane diffusely pigmented with) gray

scleronema, female, ZRC 41531, 336 mm SL; c) C. pachynema, male along margin. Anal fin with interradial
holotype, CAS 96577, 266.9mm SL; d) C. pachynema, female membrane diffusely pigmented with gray

paratype, CAS 96576, 218.7 mm SL. Scale bar = | mm. along distal two-thirds, becoming denser
along the distal edge. Interradial mem-

branes of caudal-fin lobes diffusely pigmented with gray along distal three-quarters; melanophores
absent from proximal quarter of lobes and particularly dense at tips of lobes.

DISTRIBUTION. — This species is known from the southeastern part of the Chao Phraya drainage,
and has been recorded from the Mekong drainage at Ubon Ratchathani by Nabhitabhata (1991)
(Fig. 4).

ETYMOLOGY. — From pachys, Greek for thick and nema, Greek for thread, in reference to the
thickened maxillary barbels of mature males.

REMARKS. — Ceratoglanis pachynema can be easily distinguished from C. sc/eronema in having
a snout with a gently sloping (vs. steep) ventral profile (ventral angle of snout 40°-42° vs. 20°-26°;
Fig. 5). Mature males of C. pachynema also have large membranous extensions of the maxillary barbel
present along both the anterior and posterior edges of the barbel (vs. a smaller membranous extension
present only along the anterior edge of the barbel in C. scleronema; Fig. 6).

ACKNOWLEDGMENTS

I thank Carl Ferraris for commenting on the manuscript, and the following for the loan of material
and permission to examine specimens under their care: William Eschmeyer and David Catania (CAS);
Lynne Parenti (USNM); Isaac Isbriicker (ZMA); Yang Chang Man and Kelvin Lim (ZRC). I also
thank Jon Fong (CAS) for providing radiographs of the specimens. Funding for this project was
provided by a short-term visitor grant to the author from the Smithsonian Institution and research
grant RP 960314 to Peter K. L. Ng from the National University of Singapore.

394 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 9

LITERATURE CITED

BLEEKER, P. 1862. Atlas Ichthyologique des Indes Orientales Néérlandaises. Tome 2. Siluroides, Chacoides et
Hétérobranchoides. Frederic Muller, Amsterdam. 112 pp., pls. 49-101.

. 1863a. Systema silurorum revisum. Ned. Tijd. Dierk. 1:77—122.

. 1863b. Description de trois especes nouvelles de Siluroides del’ Inde Archipélagique. Ned. Tijd. Dierk.

1:368—371.

. 1863c. Description de trois espéces nouvelles de Siluroides del’Inde Archipélagique. Versl. Akad.
Amsterdam 15:70—76.

BORNBUSCH, A. H. 1991. Redescription and reclassification of the silurid catfish Apodoglanis furnessi Fowler
(Siluriformes: Siluridae), with diagnoses of three intrafamilial silurid subgroups. Copeia
1991(4):1070—1084.

. 1995. Phylogenetic relationships within the Eurasian catfish family Siluridae (Pisces: Siluriformes),
with comments on generic validities and biogeography. Zool. J. Linn. Soc. 115:1—46.

BORNBUSCH, A. H. AND J. G. LUNDBERG. 1989. A new species of Hemisilurus (Siluriformes, Siluridae) from the
Mekong River, with comments on its relationships and historical biogeography. Copeia 1989(2):434 444.

ESCHMEYER, W. 1998. Catalog of Fishes. California Academy of Sciences, San Francisco. 2905 pp.

HAIG, J. 1952. Studies on the classification of the catfishes of the Oriental and Palaearctic family Siluridae. Rec.
Indian Mus. 48:59-1 16.

HARDENBERG, J. D. F. 1934. Additional notes to my paper “The fishfauna of the Rokan mouth.” Treubia
14:299-312.

IMAKI, A., A. KAWAMOTO, AND A. SUZUKI. 1981. Results of an ichthyological survey in the Kapuas river, west
Kalimantan, Indonesia. Res. Inst. Evol. Biol. Sci. Rep. 1:33—54.

INGER, R. F. AND P. K. CHIN. 1962. The freshwater fishes of North Borneo. Fieldiana: Zool. 45:1—268.

KOBAYAKAWA, M. 1989. Systematic revision of the catfish genus Si/urus, with description of a new species from
Thailand and Burma. Japanese J. Ichthyol. 36:151—186.

KOTTELAT, M. 1989. Zoogeography of the fishes from Indochinese inland waters with an annotated check-list.
Bull. Zo6l. Mus. 12:1—S5.

KOTTELAT, M. AND K. K. P. Lim. 1995. Freshwater fishes of Sarawak and Brunei Darussalam: a preliminary
annotated check-list. Sarawak Mus. J. 48:227-258.

KoTTELAT, M., A. J. WHITTEN, S. N. KARTIKASARI, AND S. WIRJOATMODJO. 1993. Freshwater fishes of western
Indonesia and Sulawesi. Periplus Editions, Hong Kong. 221 pp., 84 pls.

Lim, K. K. P., P. K. L. NG, M. KOTTELAT, AND M. ZAKARIA-ISMAIL. 1993. A preliminary working list of native
freshwater fishes of peninsular Malaysia. AWB Publication No. 94. Asian Wetlands Bureau, Kuala Lumpur.
11+ 10 pp.

Myers, G. S. 1938. Notes on Ansorgia, Clarisilurus, Wallago and Ceratoglanis, four genera of African and
Indo-Malayan catfishes. Copeia 1938(2):98.

NABHITABHATA, J. 1991. Endangered species and habitats of Thailand. Ecological Research Department,
Thailand Institute of Scientific and Technological Research. 241 pp.

Na, H. H. AND P. K. L. NG. 1998. A revision of the South-east Asian catfish genus Si/urichthys. J. Fish Biol.
52:29 1-333.

ROBERTS, T. R. 1989. The freshwater fishes of Western Borneo (Kalimantan Barat, Indonesia). Mem. Calif.
Acad. Sci. 14:1-210.

. 1993. Just another dammed river? Negative impacts of Pak Mun Dam on fishes of the Mekong basin.
Nat. Hist. Bull. Siam Soc. 41:105—133.

SMITH, H. M. 1931. Notes on Siamese fishes. J. Siam Soc. Nat. Hist. Suppl. 8:177—190.

————., 1945. The freshwater fishes of Siam or Thailand. Bull. U.S. Natn. Mus. 188:1—622.

SUVATTI, N. C. 1936. Index to the fishes of Siam. Bureau of Fisheries, Bangkok. 226 pp.

———. 1950. The fauna of Thailand. Department of Fisheries, Bangkok. 11 + 1100 pp.
———, 198]. Fishes of Thailand. The Royal Institute, Thailand. 380 pp. [in Thai].

Tweepir, M. W. F. 1952. Notes on Malayan fresh-water fishes. 3. The anabantoid fishes, 4. Some new and
interesting records, 5. Malay names. Bull. Raffles Mus. 24:63—95.

VAILLANT, L. 1902. Résultats zoologiques de l’expédition scientifique Neerlandaise au Bornéo Central. Notes
Leyden Mus. 24:1—166, pls. 1-2.

NG HEOK HEE: REVIEW OF THE GENUS CERATOGLANIS 395

VIDTHAYANON, C., J. KARNASUTA, AND J. NABHITABHATA. 1997. Diversity of freshwater fishes in Thailand.
Office of Environmental Policy and Planning, Bangkok. 102 pp.

WEBER, M. ANDL. F. DE BEAUFORT. 1913. The fishes of the Indo-Australian Archipelago. Vol. 2. Malacopterygil,
Myctophoidea, Ostariophysi: I. Siluroidea. E. J. Brill, Leiden. xx + 404 pp.

ZAKARIA-ISMAIL, M. 1984. Checklist of fishes of Taman Negara. Malayan Nat. 37:21—26.

© CALIFORNIA ACADEMY OF SCIENCES, 1999
Golden Gate Park
San Francisco, California 94118

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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES

Volume 51, No. 10, pp. 397-424, 8 figs., 7 tables. October 22, 1999

Revision of the South Asian Bagrid Catfish Genus Sperata,
with the Description of a New Species from Myanmar

'

by

Carl J. Ferraris, Jr. OCT 2 5) 1999
Department of Ichthyology, California Academy of Sciences
Golden Gate Park, San Francisco, California 94118

and

Kathryn E. Runge
289 Prince's Point Road
Yarmouth, Maine 04096

The South Asian bagrid catfish genus Sperata Holly, 1939, comprises a group of four species
distributed from Pakistan to Myanmar. All species of this genus are commercially impor-
tant food fishes and among the largest catfishes of South Asia. The generic name for this
group has undergone a series of replacements and corrections since the name Macrones
Duméril, 1856, was proposed for Bagrus lamarrii Valenciennes, 1840 |= Sperata seenghala
(Sykes, 1839)| and several other species. Most recently, the genus was known as Aorichthys
Wu, 1939, a name published several months after Sperata. Sperata aor (Hamilton, 1822)
and S. seenghala (Sykes, 1839) were found to be widely distributed in India and neighbor-
ing countries. Sperata aorella (Blyth, 1858), which has been included in the synonymy of
one or the other of these species since shortly after the name was first proposed, was instead
found to represent a valid species of the Ganges River delta and nearby areas. The species
of Sperata distributed widely in Myanmar is not any of the named forms and is herein
described as Sperata acicularis n. sp.

In 1822, Hamilton published a description and two figures of a new species of catfish from the
Ganges River that he named Pime/odus aor. Among the characteristics mentioned in the description
and seen in the illustrations were a prominent black spot on the posterior end of the adipose fin (Fig.
la), a strongly depressed head, and an ovoid bone on the nape situated on the dorsal midline, between
the tip of the supraoccipital process and the base of the dorsal fin (Fig. 1b).

Soon thereafter, several new species of fish were described as being similar to Hamilton’s
Pimelodus aor. Sykes (1839a, 1839b, 1841) stated that he considered his new species, Platvstoma
seenghala, from the Deccan region of peninsular India “closely allied” (Sykes 1841:372) to Hamil-
ton’s species. Valenciennes (in Cuvier and Valenciennes 1840) described his new Bagrus lamarrii
as similar in form to, and of the same coloration as, Pimelodus aor. The similarity to Hamilton’s
species was clearly implied by the choice of names given to Bagrus aorinus Valenciennes, 1840 (in
Jacquemont 1835—1844), Bagrus aorides Jerdon, 1849, and Bagrus aorellus Blyth, 1858.

At about the same time as these last species were being named, Duméril (1856) suggested that a
group of catfishes, including Valenciennes’ Bagrus lamarrii, belonged to a distinct genus for which
he proposed the name Macrones. That name was quickly adopted for a large and diverse group of
Asian bagrid catfishes (e.g., Giinther 1864; Day 1877). Within the broad genus Macrones, Giinther
(1864) recognized a small subgroup based on a comparatively short adipose fin and a separate

397,

398 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 10

interneural shield, to which he assigned Pimelodus aor, its purported synonyms, and Bagrus lamarrii.
Although not formally named, this appears to be the first recognition of a group that is called Sperata
herein. This group has, over time, been elevated to subgeneric rank [as Mystus (Osteobagrus), Jayaram
1954] or generic rank (e.g., Aorichthys, Jayaram 1968; Talwar and Jhingran 1991). As part of a broad
phylogenetic study of the catfish family Bagridae, Mo (1991) presented additional morphological
evidence that supported the idea that these fishes formed a natural group deserving of generic rank.

The purpose of this paper is to review the species of the group referred to in the recent literature
as Aorichthys or Osteobagrus in order to determine the number of, and correct names for, included
species and determine the valid generic name for the group.

METHODS AND MATERIALS

Specimen lengths are given as standard length. Measurements were taken from the left side of
the body as follows: anal-fin base—from anal-fin origin to posterior base of last anal-fin ray; body
depth at pectoral fin—measured perpendicular to long axis of body at pectoral-fin base; body width
at cleithrum—measured at anterior margin of pectoral-spine base; head length—from snout tip to
most posterior edge of fleshy operculum; interneural shield length—superficial, exposed part of
supraneural measured along dorsal midline; interorbital width—dorsal edge of membranous orbit of
one eye to that of other eye; orbit length—horizontal distance across eye to margins of free orbital
membrane; pectoral-spine length—measured from joint to tip of spinous portion, not including
filamentous extension; pelvic-fin length—from pelvic-fin origin to tip of longest ray of adpressed fin;
preadipose length—from snout tip to point where adipose fin starts to rise from body; preanal
length—distance from snout tip to anal-fin origin; predorsal length—from snout margin to base of
first dorsal-fin spine (= spinelet); prepelvic length—snout tip to base of first pelvic-fin ray; snout
length—from tip of snout to horizontal line through anterior margin of fleshy orbit; snout to
supraoccipital spine—from snout tip to posterior-most edge of supraoccipital spine visible on surface
of skin; supraoccipital spine length—from anterior-most portion of spinous process to end of visible
portion of spine.

Dorsal and pectoral-fin ray counts do not include spinous elements. Anal-fin ray counts are
reported as unbranched and branched rays. Anterior unbranched rays are embedded in thick skin and
difficult to count, except by radiograph. Usually, the first branched anal-fin ray occurs immediately
posterior to the longest ray of the fin, which is unbranched. However, in some individuals the longest
ray was branched near its tip and was therefore included in the branched ray count. Principal caudal-fin
ray counts include branched rays and one unbranched ray in each lobe of the fin. Gill raker counts
include all bony elements on the outer face of the first gill arch, including anterior rudiments. Total
gill rakers as well as differential raker counts (upper and lower) are provided as both meristics proved
useful in discriminating species. A gill raker situated at the angle of the arch was included in the count
of the lower arm.

Vertebral counts were taken from radiographs or dry skeletal preparations. Counts include four
vertebrae for the Weberian complex and one for the hypural complex. Vertebra 5 was recognized by
having an elongate hour-glass shaped centrum, which was sutured anteriorly to the Weberian
complex, and no rib. The precaudal vertebral count includes all vertebrae anterior to the centrum with
a complete hemal arch; the caudal vertebral count includes all remaining vertebrae. The preanal
vertebral count includes all vertebrae for which the hemal spine was anterior to first anal-fin
pterygiophore. Rib counts refer to the number of vertebrae possessing a rib, even if ribs are not
bilaterally paired.

Throughout the paper, we refer to the third edition of /nternational Code of Zoological Nomen-
clature (International Commission on Zoological Nomenclature 1985) as the Code.

Museum abbreviations used in this paper follow Leviton et al. (1985).

FERRARIS AND RUNGE: SOUTH ASIAN BAGRID CATFISH GENUS SPERATA

HY | | wm,

j Fis

aa 3 F A \\y J f f
l/h

FIGURE |. Sperata aor, illustration from Hamilton (1822, pl. 20, fig. 68).

399

400 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 10

SYSTEMATIC ACCOUNTS
Sperata Holly, 1939

Macrones Dumeril, 1856:484 (Type species: Bagrus lamarrii Valenciennes, 1840, by original designation).
Preoccupied in Coleoptera by Macrones Newman, 1841.

Aoria Jordan, 1919:341 (replacement for Macrones Dumeril, 1856, and therefore taking the same type species,
Bagrus lamarrii Valenciennes, 1840). Preoccupied in Coleoptera by Aoria Baly, 1863.

Sperata Holly, 1939:143 (replacement for Macrones Dumeril, 1856, and therefore taking the same type species,
Bagrus lamarrii Valenciennes, 1840 [but mistakenly listed with “Sperata vittatus (Bloch)” as type species]).

Aorichthys Wu, 1939:131 (replacement for Aoria Jordan, 1919, and therefore taking the same type species,
Bagrus lamarrii Valenciennes, 1840).

Macronichthys White and Moy-Thomas, 1940:505 (replacement for Aoria Jordan, 1919, and therefore taking
the same type species, Bagrus lamarrii Valenciennes, 1840).

Osteobagrus (subgenus of Mystus) Jayaram, 1954:529, 547 (Type species: Pimelodus aor Hamilton, 1822, by
original designation).

DIAGNOSIS. — Sperata is distinguished from all other bagrid catfishes in having the following
combination of derived characters: an elongate depressed snout, an interneural shield that is not
suturally attached to the exposed surface of the first dorsal-fin pterygiophore, a concavity on the
posterior surface of the posttemporal into which an anterior extension of the swimbladder rests
(Jayaram 1973), a swimbladder with a complete internal longitudinal septum (Jayaram 1954), an
elongate maxilla (Mo 1991), and a large round or ovoid dark spot near the posterior margin of the
adipose fin.

REMARKS. — Sperata is a readily recognizable group of South Asian catfishes. The elongate,
depressed head and snout that gives the appearance of a duck’s bill and the prominent dark adipose-fin
spot permit ready identification of these fishes. In addition, two species, Sperata aor (Fig. 2b) and S.
seenghala (Fig. 2d), have a large, rugose shield-shaped supraneural bone on the nape (called the
interneural shield by Jayaram [1954]), that is not sutured to the first dorsal-fin pterygiophore. The
supraneural bone in other bagrids is firmly united along its posterior margin to the first dorsal-fin
pterygiophore. The supraneurals of the other two species, Sperata acicularis (Fig. 2a) and S. aorella
(Fig. 2c), are not much expanded anteriorly or laterally, but are separate from the pterygiophore. These
latter two species have proportionally much longer supraoccipital spines than do the former species.
Together, the supraoccipital spine and superficial part of the supraneural traverse nearly the entire
middorsal extent of the nape in all four species.

As indicated in the synonymy of Sperata, the generic name for this group has undergone a number
of changes. Almost immediately after Macrones was proposed by Duméril (1856), it was adopted for
a large group of Asian bagrid catfishes, with Bagrus /amarrii as its type. Macrones was used widely
until Jordan (1919) noted that the name was preoccupied by Macrones Newman, 1841 (in Coleoptera).
Unfortunately the replacement he proposed, Aoria, was itself preoccupied by Aoria Baly, 1863 (in
Coleoptera). The prior use of Aoria was noted independently in Wu (1939) and White and Moy-
Thomas (1940), probably due to the recent publication of the first volume of Neave’s (1939)
Nomenclator Zoologicus, and a replacement name was proposed in each: Aorichthys Wu, 1939, and
Macronichthys White and Moy-Thomas, 1940. At about the same time, Holly (1939) also discovered
that Macrones was preoccupied, but he was apparently unaware of Jordan’s replacement name when
he proposed Sperata as a replacement. Jayaram (1954) knew of all of the generic names proposed
previously for this group and correctly noted that Macrones and Aoria were preoccupied. He was
apparently unaware, however, that Sperata, Aorichthys, and Macronichthys were each proposed as
replacements, each with Bagrus lamarrii as its type. Instead, he considered the taxon that included
the names Pimelodus aor and Bagrus lamarrii to be without an available generic-level name and

FERRARIS AND RUNGE: SOUTH ASIAN BAGRID CATFISH GENUS SPERATA 401

proposed the subgenus Osteobagrus, with P. aor as its type. He again supported this distinction in
reviewing the taxonomic status of the generic name Mystus (Jayaram 1962), into which he
synonymized Macrones, Aoria, Sperata, Aorichthys, and Macronichthys, but not Osteobagrus.
Subsequently, Jayaram (1968, 1973) recognized Aorichthys as the valid name for the taxon he
previously called Osteobagrus, which he then considered to be of generic rank. Since then, Aorichthys
has been widely regarded as the valid name for the taxon, as either a subgenus of Mystus (e.g., Misra
1976) or a genus (e.g., Jayaram 1981; Talwar and Jhingran 1991).

Although both Sperata and Aorichthys were proposed in 1939, the former predates the latter by
several months. Sperata was published in part 5/6 of volume 125 of the Zoologicher Anzeiger, the
cover page of which bears a publication date of February 15, 1939. Aorichthys was published in
volume 10 of Sinensia which was issued in its entirety in December, 1939. Thus, Aorichthys must be
considered a junior synonym of Sperata.

Holly (1939) specifically indicated that Sperata was proposed as a replacement for Macrones
Dumeril, 1856, which was shown to be preoccupied. According to article 67(h) of the Code, a
replacement name for the junior homonym must take the same type as the name it replaces “despite
any statement to the contrary.” Therefore, Sperata must take Bagrus lamarrii Valenciennes as its type
even though Holly, for whatever reason, listed Sperata vittata (Bloch) as the type of his new name.

Four species of Sperata are recognized herein. The species were found to differ in a variety of
meristic characters, summarized in Tables | through 6. In addition, the shape of several bones of the
head and nape also appear to diagnose the species. The small sample size of some species available
for study, combined with the wide range of sizes of specimens, made it difficult to find morphometric
trends among the species. A few morphometric distinctions are proposed (Table 7), but a thorough
study of the morphometry of the species was not possible on the basis of the available material.

Because we now recognize three Sperata species in the Ganges River basin, within which only
two species have been recognized previously, it is difficult to ascertain at this time which aspects of
the biology of these fishes, if any, are species specific. Therefore, a summary of the biology of Sperata
is presented here, rather than in the individual species accounts.

All species of Sperata are important food fishes (Prashad and Mukerji 1929; Talwar and Jhingran
1991) that grow to a large size, cited as reaching 1.8 m in various literature sources (e. g., Talwar and
Jhingran 1991), but reports of specimens greater than | m are rare. Some preliminary information of
the size at maturity is found in Sundara Raj (1962:194), within which the species were in the genus
Mystus. Therein, nest-guarding males identified as Mystus aor ranged in size from 342 to 460 mm,
while a single M. seenghala male was 685 mm. Saigal (1967:24) found that gonad maturation
commenced at about a total length of 800 mm in specimens identified as Mystus aor.

Food habits were reported for Sperata aor (Saigal 1967). Young individuals were found to eat
primarily insects off the substratum, whereas adults fed on fish and insects from both the water column
and on the bottom. A wide variety of fish were found in the gut contents of S. aor, and the relative
proportion of various types of food changed seasonally.

The reproductive biology of Sperata is incompletely known. Spawning is said to occur before
the beginning of the southwest monsoon season (Saigal 1967; Talwar and Jhingran 1991), and nests
were found in rocky-bottoms at Bhavani, India, during the month of May (Sundara Raj 1962:194).
Individuals were reported to spawn at irregular intervals throughout the reproductive season and
females were capable of producing more than 1.2 million ova (Saigal 1967:25). The most remarkable
aspect of the reproductive biology of Sperata was the suggestion made by Sundara Raj (1962:195)
that males may incubate eggs in highly vascularized, spongy skin on the ventral surface of their
abdomen, and that the same males may feed the newly hatched fry with a milky-white exudate from
the abdominal wall. Saigal (1967:29) reported that mature individuals had “almost empty guts” during
the breeding season, suggesting that they did not feed during that period. As mature Sperata are
primarily piscivorous, the cessation of feeding activity by the parents while caring for the young be

402 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 10

TABLE |. Precaudal and caudal vertebral counts for four species of Sperata.

Precaudal vertebrae Caudal vertebrae

lone ie 19> 20°" 21" 22° 23 23 29 © 30" “3 32 33 34, 335) scone

S. acicularis A vd. l D | 2
S. aor 4014 9 l pee le Roe pee

S. aorella 5 3 l l

S. seenghala Cm 2 De eee l Sy all

TABLE 2. Preanal and total vertebral counts for four species of Sperata.
Preanal vertebrae Total vertebrae

DO i ae Oe oa oe 47 48" "49 50" “S11 52 Same

S. acicularis 5 l 2 pd D
S. aor 3 lO) 4 | 6 9 D
S. aorella l 2 2 3 | |
S. seenghala 4 4 ] l 3 l 5 l
_ TABLE 3. Pectoral and branched anal-fin ray counts for four species of Sperata.
Pectoral-fin rays Branched anal-fin rays

8 9 10 1] 8 9 10 11 I
S. acicularis | 14 12 2 l
S. aor 21 5 2 iS 9
S. aorella 9 | 10
S. seenghala 2 15 Z 12 3

TABLE 4. Gill rakers on lower and upper arm of first gill arch of four species of Sperata. |
Lower gill rakers Upper gill rakers

11 12 13 14 15 16 Ig) 2 3 4 5 6
S. acicularis i 5 l 2 13 D l
S. aor 3 10 12 l | 6 18 l
S. aorella 2) 4 l 4 6
S. seenghala 15 2 wo} 13 2

a mechanism to avoid cannibalism. Although a wide range of parental behavior has been documented
for catfish (Blumer 1982; Ferraris 1991), parental feeding of fry is rare. Only one other species, Bagrus
meridionalis Giinther, has been reported to feed its young. McKay (1986) observed young B.
meridionalis feeding on unfertilized eggs expelled by the guarding adult female, and hypothesized
that this food source was supplemented by invertebrates brought to the nest by the male. Further
investigation of this report by LoVullo and Stauffer (1992) confirmed these observations. Although
the mode of parental feeding in B. meridionalis differs from that reported for Sperata, it is noteworthy
that these two genera were recently hypothesized to be sister groups (Mo 1991).

FERRARIS AND RUNGE: SOUTH ASIAN BAGRID CATFISH GENUS SPERATA 403

KEY TO THE SPECIES OF SPERATA

1. Interneural shield (exposed portion of supraneural) as long as, or longer than, supraoccipital spine; in large individuals,
IMtemMeunaleshieldmucosean GON O1GI (EIT See 02 Cl) rename ser Oro an 2
Interneural shield markedly shorter than supraoccipital spine and not rugose (Figs. 2a,2c) ............... 3
2. Interneural shield approximately as long as supraoccipital spine (Fig. 2b); pectoral-fin rays 10 or 11; snout rounded in
dorsal view; maxillary barbel typically extends to caudal fin; orbit extends across the middle of length of head; gill
rakersibypicallyalOon2 keen) eres A Sk cee eas, Sms as a ee Sperata aor (India, Bangladesh, Nepal)
Interneural shield longer than supraoccipital spine (Fig. 2d); pectoral-fin rays 8 or 9; snout distinctly truncate in dorsal
view; maxillary barbel typically extends no further than to middle of body, at least in large individuals; orbit entirely
in anterior half of head; gillrakers 13-15 ........... Sperata seenghala (India, Bangladesh, Nepal, Pakistan)
3. Supraoccipital spine long, slender and needlelike, tapering to a point posteriorly (Fig. 2a); pectoral-fin rays 10 (rarely 9);
branched anal-fin rays 10—12; gill rakers usually 14 or 15 (rarely 16-19)
RE fe ee ee ae eK OE ee nee Ctr e Te ts ek Sperata acicularis n. sp. (Myanmar, ? Thailand)
Supraoccipital spine long, distinctly narrower anteriorly and with rounded posterior end (Fig. 2c); pectoral-fin rays 9
(rarely 10); branched anal-fin rays 9; gill rakers 20 to22.......2... Sperata aorella (eastern India, Bangladesh)

Sperata acicularis new species
Figs. 2a, 3, 4a

Macrones aor (not Hamilton): Day 1873:cclxi (in part) (Burma); 1877:444 (Burma); 1889:149 (Burma).
Vinciguerra 1890:217 (Burma: Mandalay). Kyaw Win 1971:53, fig. 22-1 (Burma: Irrawaddy River at
Taung-tha-man “inn’).

Aoria aor (not Hamilton): Prashad and Mukerji 1929:178 (Burma: Indawgyi Lake).

Mystus (Aorichthys) aor (not Hamilton): Misra 1976:74 (in part) (upper Burma).

Aorichthys aor (not Hamilton): Jayaram, 1981:205 (in part, Burma). Talwar and Jhingran 1991:547 (in part)
(upper Burma).

?Macrones seenghala (not Sykes): Chaudhuri, 1911:20 (“Lake Tali Fu, Yunnan’).

TYPE MATERIAL. — HOLOTYPE: CAS 209024, 277 mm, Myanmar, Yangon Division, South
Oak-ka-lar-pa Market (eastern Yangon), 27—28 Oct. 1997, C. J. Ferraris, Mya Than Tun. PARATYPES:
BMNH 1894.5.24.23—24 (2, 274-295 mm), Myanmar, Sittaung River, E. W. Oates. CAS 209023 (2,
70-199 mm), Tenasserim River, upstream from Htee-tah, 1-16 March 1992, T. R. Roberts. CAS
67697 (1, 417 mm), Thailand, Chao Phraya Basin, Sing Buri Market. CAS(SU) 14491 (2, 258-283
mm), Myanmar, Pegu [= Bago], 1940, A. W. Herre. NRM 14980 (2, 150.0—-165.0 mm), Myanmar,
Mandalay, 1935. NRM 31056 (1, 214 mm), Myanmar, Kachin State, Ayeyarwaddy River, 10 March
1934, R. Malaise. NRM 31080 (2, 157-189 mm), Myanmar, Sagaing Division, Shweli River,
probably emptying into Ayeyarwaddy at Inywa, Feb 1935, Maung Lu Daw. NRM 40002 (1, 196 mm),
Myanmar, Kachin State, Myitkyina market, 30 March 1997, F. Fang, and A. Roos. NRM 40650 (3,
191-234 mm), Myanmar, Myitkyina, Lonton village market on Lake Indawgyi, 31 March 1998,
S. O. Kullander, and R. Britz.

DIAGNOSIS. — Sperata acicularis 1s readily distinguished from its congeners by having both a
long, slender supraoccipital spine that tapers posteriorly to a point and a slender interneural shield
that is shorter than the supraoccipital spine and no wider than the widest part of that spine.

This species can be further distinguished from each of the other species of Sperata by one or
more additional characters. Sperata acicularis differs from S. aorella in having fewer gill rakers
(14-19, typically 14 or 15, vs. 20-22), more precaudal vertebrae (17 or 18, vs. 16), more preanal
vertebrae (29 or 30, vs. 25—27), more total vertebrae (51—54 vs. 47-49), more ribs (15—17, vs. 13 or
14), more branched anal-fin rays (10 or more, vs. 9) and more pectoral-fin rays (10, rarely 9, vs. 9,
rarely 10). Sperata acicularis differs from S. aor in having modally fewer gill rakers (14 or 15, vs.
19 or 20) and generally more anal-fin rays (10-12, vs. 8-10) than S. aor, and a truncate, rather than
broadly rounded, snout. Sperata acicularis differs from S. seenghala in having fewer precaudal
vertebrae (17 or 18, vs. 21—23), more caudal vertebrae (33-27, vs. 28-32) more pectoral-fin rays (10,

404 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 10

TABLE 5. Total gill rakers on first gill arch of four species of Sperata.

Gill rakers
Spe sla. S568 16 Ly Sc | NOE e202

S. acicularis 6 6 l l |

S. aor l ] l 2 11 8 D.

S. aorella l 6 3
S. seenghala l 13 3

TABLE 6. Number of rib-bearing vertebrae in four species of Sperata.

Ribs
13 14 15 16 7/ 18 19 20
S. acicularis | 3 2
S. aor 10 7 et l
S. aorella 4 l |
S. seenghala 4 4 |

rarely 9, vs. 8 or 9), modally more branched anal-fin rays (10, vs. 9), and by the placement of the orbit
in the middle of the head, rather than entirely in the anterior half.

DESCRIPTION. — Body long, slender; deepest at dorsal-fin origin, tapering gradually both ante-
riorly and posteriorly. Ventral surface of head and body flat to anal-fin base. Body slightly compressed
and triangular in cross section across abdomen, compressed and ovoid across caudal region. Caudal
peduncle narrow, only slightly deeper than diameter of eye. Anus and urogenital openings located at
vertical through middle of adpressed pelvic fin; remote from anal-fin base. Skin smooth. Lateral line
complete, midlateral; canal curves slightly dorsally on caudal-fin base. In humeral region, lateral line
with numerous accessory canals and pores.

Head elongate, progressively depressed anteriorly. In lateral view, profile of head acutely
triangular, with ventral surface of head nearly horizontal. Gill opening wide, extending from exposed
surface of posttemporal to beyond isthmus. Gill membranes free from, and not attached across,
isthmus. Branchiostegal rays 12. Bony elements of dorsal surface of head covered with thin skin;
bones readily visible, ornamented with fine, irregular, radial grooves. Midline of cranium with
elongated fossa extending from posterior of snout nearly to base of supraoccipital spine. Within fossa,
broad epiphysial bar separates slender anterior and posterior medial fontanels. Lateral fontanel small,
shorter than orbital diameter. Supraoccipital spine elongate, slender, pointed at tip. Cephalic lateral-
line canal system with extensive branching of most canals; integument covering fontanels, cheeks,
and olfactory chamber thoroughly covered with canals and pores.

Barbels in four pairs. Maxillary barbel long, slender, without medial membrane; barbel extends
at least to middle of adipose fin, and reaches caudal peduncle in some specimens. Nasal barbel slender,
short; extending no further than anterior margin of orbit. Inner mental barbel remote from midline
and slightly thicker and longer than nasal barbel; extending past middle of eye. Outer mental barbel
originating about one eye diameter posterolateral of inner mental barbel; barbel thicker and longer
than inner mental barbel, extending to level of pectoral-fin origin.

Eye ovoid, horizontal axis longest; located entirely in dorsal half of head. Posterior margin of
orbit at middle of head length. Upper margin of orbit nearly reaching dorsal profile of head. Eye
diameter more than one-third of snout length and slightly less than interorbital width. Orbital margin
mee:

FERRARIS AND RUNGE: SOUTH ASIAN BAGRID CATFISH GENUS SPERATA

405

FiGure 2. Head and nape of four species of Sperata, in dorsal view. Maxillary barbels truncated; stippled areas indicate
bones that are exposed at surface. a) Sperata acicularis new species, CAS 209024, 277 mm; b) Sperata aor, CAS(SU) 14124,
278 mm; c) Sperata aorella, CAS(SU) 34852, 234 mm; d) Sperata seenghala, CAS 24247, 284 mm. Abbreviations: ss —

supraoccipital spine; in—interneural shield; dfp— exposed portion of first dorsal-fin pterygiophore. Illustration by Alison
Schroeer.

Volume 51, No. 10

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FERRARIS AND RUNGE: SOUTH ASIAN BAGRID CATFISH GENUS SPERATA 407

Snout long and depressed; lateral margins nearly parallel. Anterior margin of snout slightly
rounded, nearly truncate in some individuals. Mouth slightly subterminal; most premaxillary teeth
exposed when mouth closed. Oral teeth small, sharply pointed, in irregular rows on all tooth-bearing
surfaces. Premaxillary tooth band shallowly crescentic, width of band somewhat greater near midline.
Tooth band of dentary narrower than premaxillary band at symphysis, tapering laterally. Palatal tooth
patch unpaired, continuous across midline; smoothly arched along anterior margin, tapering laterally
to point that extends posteriorly well past level of premaxillary band; band width narrower than
premaxillary band at midline, widening laterally and then tapering to sharp point posterolaterally. Gill
rakers of first arch long, slender, and closely spaced. Gill rakers 11 to 14 on lower arm plus 3 (rarely
4 or 5) on upper arm. Rakers present only on outer face of first two arches, on both faces of next two
arches.

Dorsal fin located above middle of body; fifth or sixth dorsal-fin ray located at vertical through
middle of standard length. Dorsal-fin base shorter than length of first branched ray. Dorsal fin margin
straight, first branched ray longest, more than twice length of last ray. Last ray without posterior
membranous connection to body. Dorsal fin with spinelet, spine, and 7 branched rays. Dorsal-fin
spine long, straight and slender; spine margin smooth anteriorly and laterally, but with fine serrations
on distal half of posterior edge. Spine slightly shorter than first branched ray. Tip of adpressed spine
just reaches adipose-fin origin. Distance between dorsal fin and adipose fin less than dorsal-fin base
length. Interneural shield narrow, tapering to fine point anteriorly; shorter than supraoccipital spine.
Superficial ossification of dorsal-fin pterygiophores broadly united across midline, and forming a
chevron-shaped bone, pointed anteriorly (Fig. 4a).

Adipose-fin base approximately | 4 times length of dorsal-fin base; fin height about one-fourth
of its length. Adipose-fin margin straight anteriorly, slightly convex for remainder of its length;
posterior portion deeply incised.

Caudal fin deeply forked; lobes pointed, asymmetrical, lower lobe wider than upper; upper lobe
continued as filament in intact specimens. Middle rays approximately one-third as long as unbranched
principal ray of lower lobe. Principal caudal fin with rays: 1,7,8,1. Procurrent rays symmetrical and
extending only slightly anterior to fin base.

Anal-fin origin ventral to middle of adipose fin. Anal-fin margin straight; first branched ray
longest, about twice length of last ray, which lacks posterior membranous connection to body. Anal
fin with 3 or 4 unbranched, and 10 (rarely 11 or 12) branched rays.

Pelvic-fin origin at vertical through posterior end of dorsal-fin base. Fin margin straight, its first
branched ray longest; posterior-most ray about two-thirds length of first ray. Fin with one unbranched
and five branched rays. Tip of adpressed fin not reaching to anal-fin origin.

Pectoral fin with stout spine, sharply pointed at tip. Anterior spine margin smooth, posterior
margin with moderately strong serrations along entire length. Pectoral-fin margin straight anteriorly,
convex posteriorly. First branched ray longer than spine and approximately 22 times as long as last
ray. Pectoral-fin with 10 (rarely 9) branched rays.

COLOR IN PRESERVATIVE. — Body silvery grey on dorsal half of lateral surface, white or silver
ventrally. Grey region resulting from presence of fine black pigment covering side of body for most
of its length, but only dorsal to lateral line on caudal peduncle. Dorsal surface of body and head brown.
On head, brown regions restricted to exposed cranial bones and around orbit. Ventral surface of head
and abdomen without dark pigmentation.

Dorsal fin pale basally, progressively darker distally; anterior edge of spine and distal extent of
rays dusky. Dusky region of anterior rays less extensive than on more posterior rays. Adipose fin
mostly dusky; margin of fin with fine, distinct dark band. Posterior portion of fin with ovoid black
spot, smaller than orbit in diameter; spot surrounded by pale halo. Caudal peduncle with blackened
area just ventral to adipose-fin spot, appearing as extension of spot. Caudal fin dusky, ventral lobe
darker; ventral unbranched principal ray and ventral procurrent rays white. Anal fin generally pale

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FIGURE 3. Sperata acicularis, new species, 277 mm SL, holotype, CAS 209024, Myanmar, Yangon Division, South
Oak-ka-lar-pa market. Illustration by Alison Schroeer.

with only slight duskiness distally; in one specimen (CAS 209024) distal half of anal fin heavily
pigmented. Pelvic fin pale, or with broad diffuse dusky subterminal band. Pectoral fin with scattered
fine dark pigmentation on outer rays, little or no pigment on innermost rays. Pectoral spine dark
dorsally and pale ventrally.

REMARKS. — The first record of Sperata acicularis appears to be that of Day (1873), where
Myanmar (as Burma) is listed among the localities for Macrones aor and the Burmese name for this
species, Nga-joung, is noted. This species has been generally considered conspecific with Sperata
aor, but at least one author has suggested a difference between that species and the Ayeyarwaddy
River form (recognized here as S. acicularis). Kyaw Win (1973) noted that the maxillary barbel of
specimens from near Mandalay extend only to the adipose fin and not to the end of the caudal fin as
widely reported for individuals of S. aor from the Ganges River drainage. Although the maxillary
barbel of S. aor does not always extend to the tip of the caudal fin (see Remarks under that species),
it extends well past the adipose-fin origin.

Little has been reported about the biology of Sperata acicularis. Prashad and Mukerji (1929)
stated that the species was abundant both in Indawgyi Lake and elsewhere in northern Myanmar and
was a common food fish all year round.

DISTRIBUTION. — Sperata acicularis is known from the Ayeyarwaddy, Bago, and Tenasserim
river systems of Myanmar. One record from the Chao Phraya basin of Thailand (CAS 67697) is
probably incorrect since there are no literature accounts of the species or any of its congeners in the
Chao Phraya, or anywhere else in Thailand (e. g., Smith 1945; Suvatti 1950; Kottelat 1989). As this
species is quite large and distinctive, we believe it unlikely that it was overlooked until now and,
instead, suggest that the locality information associated with this specimen is erroneous.

Sperata acicularis appears to be the only species present in Myanmar, but during this study we
examined one specimen of S. seenghala (NRM 18809), which was reported from “probably Mandalay
or Yangon areas.” Whether this represents a second species in Myanmar or an erroneous locality
cannot be determined at this time.

ETYMOLOGY. — The name acicularis, Latin for needlelike (Brown 1956), refers to the long
slender supraoccipital spine that most readily distinguishes this species from its congeners.

FERRARIS AND RUNGE: SOUTH ASIAN BAGRID CATFISH GENUS SPERATA 409

Cc

Ficure 4. Anterior view of dorsal fin and associated ossifications of four species of Sperata, in dorsal view. Stippled area
indicate bony parts of interneural shield and dorsal-fin pterygiophores that are visible in intact specimens. a) Sperata acicularis
new species, NRM 31056, 214 mm; b) Sperata aor, CAS(SU) 34853, 215 mm; c) Sperata aorella, MCZ 39637, 179 mm; d)
Sperata seenghala, MCZ 8185, 259 mm. Abbreviations: in—interneural shield; dfp—exposed portion of first dorsal-fin
pterygiophore. Scale bar = 10 mm. Illustration by Kathryn Runge.

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Sperata aor (Hamilton, 1822)
Figs. 1, 2b, 4b

Pimelodus aor Hamilton, 1822:205, 379, pl. 20, fig. 68 (Type locality: Rivers of Bengal and upper parts of
Gangetic estuaries).

Bagrus aor Valenciennes, in Cuvier and Valenciennes, 1840a:405 (Bengale). Valenciennes, 1840 (in Jac-
quemont, 1835—1844): pl. 16 (India).

Bagrus aorides Jerdon, 1849:336 (Type locality: India: Cauvery River at Errode; no types known to exist).

Macrones aor Giinther, 1864:78 (India, Bangladesh). Day 1873:cclxi (in part) (plains of India); 1877:444 (in
part) (India). Lydekker, 1886:250, pl. 36, fig. 5 (India: Siwalik Hills [Tertiary deposits]). Day 1889:149 (in
part) (India); Woodward, 1901:327 (India: Siwalik Hills [Lower Pliocene]).

Mystus (Osteobagrus) aor Jayaram, 1954:548, fig. 10 (in part) (India); Srivastava, 1968:77, fig. 49 (India: Uttar
Pradesh).

Aorichthys aor Jayaram, 1973:155, fig. la, (in part, India); 1981:205, fig. 98, 99b (in part) (India, Bangladesh).
Talwar and Jhingran 1991:547, fig. 178 (in part) (India, Nepal, Bangladesh).

Mystus (Aorichthys) aor Misra, 1976:74 (in part) (India).

Mystus aor Ataur Rahman, 1989:197 (Bangladesh); Shrestha, 1994:54, fig. 83 (Nepal: Terai region).

DIAGNOSIS. — Sperata aor is distinguished from its congeners by the combination of a broadly
rounded snout margin and a large, ovoid, interneural shield that is approximately the same length as
the supraoccipital spine.

In addition, this species has a unique combination of meristic values that further distinguish it
from each of its congeners. Sperata aor has a higher modal gill-raker count than does S. acicularis
(19 or 20, vs. 14 or 15) and generally fewer branched anal-fin rays (8-10, vs. 10-12). Sperata aor
differs from S. aorella in having more preanal vertebrae (28-30, vs. 25-27), more total vertebrae
(50-52, rarely 49, vs. 47-49), and more pectoral-fin rays (10 or 11, vs. 9, rarely 10). Sperata aor has
a higher modal gill-raker count than does S. seenghala (19, vs. 14) as well as more pectoral-fin rays
(10 or 11, vs. 8 or 9) and fewer precaudal vertebrae (16—19, vs. 21—23).

DESCRIPTION. — Body long, slender; body depth at dorsal-fin origin only slightly greater than
that anterior to adipose-fin origin, more posteriorly tapering gradually. Ventral surface of head and
body flat to anal-fin base. Body slightly compressed and triangular in cross section across abdomen,
compressed and ovoid across caudal region. Caudal peduncle narrow, only slightly deeper than
diameter of eye. Anus and urogenital openings located at vertical through middle of adpressed pelvic
fin; distance from anal-fin base greater than anal-fin base length. Skin smooth. Lateral line complete,
midlateral; canal curves slightly dorsally on caudal-fin base. Lateral line in humeral region with
numerous accessory canals and pores.

Head elongate, progressively depressed anteriorly. In lateral view, profile of head acutely
triangular, with ventral surface of head nearly horizontal. Gill opening wide, extending from exposed
surface of posttemporal to beyond isthmus. Gill membranes free from, and not attached across,
isthmus. Branchiostegal rays 12 or 13. Bony elements of dorsal surface of head covered with thin
skin; bones readily visible, ornamented with fine, irregular, radial grooves. Midline of cranium with
elongate fossa extending from posterior of snout nearly to base of supraoccipital spine; posterior half
of fossa occupied by posterior fontanel, separated from slender anterior fontanel by wide epiphyseal
bar. Lateral fontanel small, shorter than orbital diameter. Supraoccipital spine elongate, with parallel
sides and blunt posterior tip. Spine of smaller specimens slender and needlelike, becoming propor-
tionally wider in larger individuals. Cephalic lateral-line canal system with extensive branching of
most canals; integument covering fontanels, cheeks, and olfactory chamber thoroughly covered with
canals and pores.

Barbels in four pairs. Maxillary barbel long, slender, without medial membrane; tip of barbel
extends at least to caudal peduncle and often past tip of caudal-fin rays. Nasal barbel slender; extending

FERRARIS AND RUNGE: SOUTH ASIAN BAGRID CATFISH GENUS SPERATA 411

past anterior margin of orbit and sometimes to its posterior margin. Inner mental-barbel origin close
to midline; barbel thicker and longer than nasal barbel, extending to level of posterior margin of orbit.
Outer mental barbel originates about one-half of eye diameter posterolateral of inner mental barbel;
barbel thicker and longer than inner mental barbel, extending past pectoral-fin origin.

Eye ovoid, horizontal axis longest; located entirely in dorsal half of head. Center of eye at middle
of head length. Upper margin of orbit reaching dorsal profile of head; frontals bowed dorsally along
lateral margin; interorbital region of neurocranium distinctly concave. Eye diameter less than, or equal
to, one-half of snout length and slightly less than interorbital width. Orbit with free margin.

Snout long and depressed; lateral margins slightly convergent anteriorly. Snout margin distinctly
rounded, with fleshy upper lip extending anteriorly beyond upper jaw. Mouth subterminal, premax-
illary tooth patch exposed when mouth closed. Oral teeth small, sharply pointed, in irregular rows on
all tooth-bearing surfaces. Premaxillary tooth band rounded, of equal width throughout. Dentary tooth
band much narrower than premaxillary band at symphysis, tapering laterally. Palatal tooth patch
unpaired, continuous across midline; smoothly arched along anterior margin, tapering laterally to
point extending posteriorly well past level of premaxillary band; band width narrower than premax-
illary band at midline, widening laterally and then tapering to sharp point posterolaterally. Gill rakers
of first arch long, slender and closely spaced. Gill rakers 12 to 15 on lower arm plus 4 or 5 (rarely 3
or 6) on upper arm. Rakers present only on outer face of first two arches, present on both faces of
next two arches.

Dorsal fin located above middle of body; middle of fin at vertical through middle of standard
length. Dorsal-fin base shorter than length of first branched ray. Dorsal-fin margin straight, first
branched ray longest, more than twice length of last ray. Last dorsal-fin ray without posterior
membranous connection to body. Dorsal fin with spinelet, spine, and 7 branched rays. Dorsal-fin
spine long, straight, and comparatively robust; slightly shorter than first branched ray. Anterior and
lateral spine margins of small specimens smooth, with fine serrations on distal half of posterior edge.
In large individuals, anterior and lateral margin of spine granular, posterior serrations less prominent.
Tip of adpressed spine reaches past adipose-fin origin. Distance between dorsal fin and adipose fin
less than one-half of dorsal-fin base length. Interneural shield elongate oval with rounded anterior
and posterior margins. Superficially first dorsal-fin pterygiophore appears narrowly united across
midline or completely separated by thick skin; anterior margin deeply concave anteriorly (Fig. 4b).

Adipose-fin base length approximately | to 2 times that of dorsal-fin base; fin height about
one-fifth of its length. Adipose-fin margin slightly convex for entire length; posterior end deeply
incised.

Caudal fin deeply forked; lobes pointed, symmetrical, except upper lobe continued as filament
in intact specimens. Middle rays approximately one-third as long as unbranched principal ray of lower
lobe. Principal caudal fin with rays: i,7,8,1. Procurrent rays symmetrical and extend only slightly
anterior to fin base.

Anal-fin base ventral to posterior half of adipose fin. Fin margin straight; first branched anal-fin
ray longest and about twice length of last ray. Last ray without posterior membranous connection to
body. Anal fin with 3 or 4 unbranched, and 8 to 10 branched rays.

Pelvic-fin origin at vertical through posterior end of dorsal-fin base. Pelvic-fin margin slightly
convex; first branched ray longest, posterior-most ray about two-thirds length of first ray. Pelvic fin
with one unbranched and five branched rays. Tip of adpressed fin not reaching to anal-fin origin.

Pectoral fin with stout spine, sharply pointed at tip. Anterior spine margin smooth, posterior
margin with moderately strong serrations along entire length. Pectoral-fin margin straight anteriorly,
convex posteriorly. First branched ray longer than spine and approximately 2/2 times as long as last
ray. Pectoral fin with 10 or 11 branched rays.

COLOR IN PRESERVATIVE. — Body silvery grey to brown dorsally and on upper half of lateral
surface, white or pale ventrally. Grey region resulting from fine black pigment that extends ventrally

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to lateral line along caudal peduncle and somewhat below lateral line more anteriorly. Dorsal surface
of head brown, most prominent on exposed cranial bones and laterally to level of ventral margin of
orbit. Ventral surface of head and abdomen pale. Humeral region with diffuse, eye-sized, dark spot
at upper extent of gill opening.

Dorsal fin dusky at base and on distal half of fin, with medial oblique pale band. Adipose fin
dusky, leading edge sometimes darker, but not forming distinct terminal band. Posterior end of fin
with ovoid black spot approximately equal in size to pupil. Adipose fin without pigmentation dorsal
and posterior to spot. Dorsal surface of caudal peduncle with tiny blackened area posterior to adipose
fin, appearing as extension of spot. Caudal fin with scattered pigmentation, primarily on branched
rays of ventral lobe and base of dorsal lobe; ventral unbranched principal ray and ventral procurrent
rays white. Anal fin pale, with little or no pigmentation. Pelvic fin with few fine, scattered pigmen-
tation, somewhat denser basally. Pectoral fin with irregularly scattered pigmentation on dorsal
surface, pale ventrally.

REMARKS. — Sperata aor has typically been distinguished from the one previously recognized
congener, S. seenghala, on the basis of three characters: a longer maxillary barbel, a rounded (vs.
truncate) snout, and fewer principal caudal-fin rays (17, vs. 19-21) (e.g., Jayaram 1981; Talwar and
Jhingran 1991). We found that the first two characters are generally accurate, although difficult to
interpret in some specimens. The maxillary barbel of Sperata aor always extends at least to the caudal
peduncle and often past the caudal-fin base while that of S. seenghala usually does not extend past
the posterior end of the dorsal-fin base. However, the smallest specimens of S. seenghala we examined
(SU 41083, 108-130 mm) had maxillary barbels that extended past the middle of the adipose fin,
nearly as far as those of some S. aor. It appears that barbel length distinguishes adults of these two
species, but not for juveniles.

The snout of a well-preserved specimen of Sperata aor is indeed broadly rounded and contrasts
markedly with the nearly straight snout margin of S. seenghala. However, specimens of S. aor
sometimes have a truncate, if somewhat asymmetrical, snout margin that is not easily distinguished
from S. seenghala. It appears to us that these specimens were first preserved with their snouts pushed
against the bottom or side of their container, making the snout shape an artifact of preservation. Thus,
snout shape may readily distinguish fresh specimens, but may lead to misidentifications of some
preserved material.

The reported difference in caudal-fin ray counts between Sperata seenghala and S. aor appeared
first in Day (1876), and has been widely repeated up to recent times (e.g., Jayaram 1981; Talwar and
Jhingran 1991). We found no species-level distinction in caudal-fin ray counts. Nearly all Sperata
specimens examined had 17 principal rays, with a few specimens having 16.

Sperata aor is the only species of the genus known from fossil material. A single specimen found
in Tertiary deposits in the Siwalik Hills (Lydekker 1886; Woodward 1901) exhibits the same shape
and size of the interneural shield as that of Recent material.

DISTRIBUTION. — Known from the Ganges river system of India, Bangladesh, and Nepal and in
peninsular India south to the Cauvery River.

Jayaram, Venkateswarlu and Ragunathan (1982) reported this species from the Cauvery River
of southern India, but Talwar and Jhingran (1991) listed only Sperata seenghala from that drainage.
We have not examined any specimens from that drainage during the course of this study and, therefore,
we cannot resolve this discrepancy. However, the original description of Bagrus aorides Jerdon, from
the Cauvery River at Errode, sheds some light on the question. The description of B. aorides states
that the maxillary barbel extends to the caudal fin, and the adipose fin originates just posterior to the
end of the dorsal-fin base. Both of these characteristics agree more closely with those of Sperata aor
than to S. seenghala. In addition, the eye was said to be “so situated that its posterior edge is more
than half the length of the head from the muzzle” (Jerdon 1849:336), which means that the eye is not
located entirely in the anterior half of the head as it is in S. seenghala. Thus, the description more

FERRARIS AND RUNGE: SOUTH ASIAN BAGRID CATFISH GENUS SPERATA 413

closely resembles that of S. aor. On that basis, we agree with Day (1877) who first placed Bagrus
aorides in the synonymy of S. aor and, thereby, provisionally include the Cauvery River in the
distribution of S. aor.

MATERIAL EXAMINED. — 26 specimens, 57-278 mm. BANGLADESH: AMNH 56289 SD, 2,
(dry skeleton, not measured). CAS 209022 (3, 60-63 mm), Shala Bazar, 36 km from Sylhet Town,
near Kuchiara River. UMMZ 208359 (2, 134-153 mm), Comilla, Meghna River, upstream from
Chandpur, just downstream from Gumti, River mouth at Kanudi, 23°19'N, 90°39’E. UMMZ 208445
(1, 179 mm), Barisal, Meghna River at Gazipur Char, 22°47'N, 90°43’E. INDIA: AMS B 7945 (1,
265 mm), Calcutta. ANSP 83981 (2, 57-109 mm), Pulta water works, on the Hughly River, about 17
miles from Calcutta, Barrackpore District. ANSP 85764 (1, 124mm), Bombay. CAS 61857 (3,
145-160 mm), Fish Market at Sonepur. CAS 62096 (8, 126-208 mm), Karnataka State, Bellary
District, Tungabahdra River and Reservoir at Hospet, Hampi, and Kampli, Krishna River basin.
CAS(SU) 14124 (1,278 mm), Pulta, Ganges Delta. CAS(SU) 34853 (1,215 mm), Pulta. MCZ 62947
(1, 63.5 mm), Khadak Vasle Dam. NRM 13414 (1, 161 mm), Delhi, fish market.

Sperata aorella (Blyth, 1858)
Figs. 2c, 4c, 5

Bagrus aorellus Blyth, 1858:283 (Type locality: India: Calcutta fish market; no types known to exist).

DIAGNOSIS. — Sperata aorella is distinguished from its congeners by having a long, slender,
supraoccipital spine that is markedly longer than the interneural shield. The supraoccipital spine is
uniquely shaped in being constricted basally (Fig. 2c). This species is also unique in the genus in
having fewer than 28 preanal vertebrae and fewer than 50 total vertebrae. It is the only species of
Sperata that has both a truncate snout and barbels that extend past the adipose fin.

In addition, this species can be further distinguished from each of its congeners by at least one
additional character. It differs from Sperata acicularis in having fewer branched anal-fin rays (9, vs.
10-12), modally fewer pectoral-fin rays (9, rarely 10, vs. 10, rarely 9), fewer ribs (13 or 14, vs. 15—17),
and more gill rakers (20-22, vs. 14-19, typically 14 or 15). Sperata aorella has fewer pectoral-fin
rays than S. aor (9, rarely 10, vs. 10 or 11), fewer preanal vertebrae (25—27, vs. 28-30), modally more
gill rakers (21, vs. 19), a truncate, rather than rounded, snout, and an interneural shield that is not
wider than the supraoccipital spine. Sperata aorella can be distinguished from S. seenghala in having
fewer precaudal vertebrae (16, vs. 21—23), fewer preanal vertebrae (25—27 vs. 29-32), fewer ribs (13
or 14 vs. 16-20), more gill rakers (20-22, vs. 13-15), the posterior margin of the orbit situated at
about the middle of the head length instead of distinctly anterior of the midline, and an interneural
shield that is not wider than the supraoccipital spine.

DESCRIPTION. — Body long, slender; deepest at dorsal-fin origin, tapering gradually both ante-
riorly and posteriorly. Ventral surface of head and body flat to anal-fin base. Body slightly compressed
and triangular in cross section across abdomen, compressed and ovoid through caudal region. Caudal
peduncle only slightly deeper than wide. Anus and urogenital openings located at vertical through
middle of adpressed pelvic fin; remote from anal-fin base. Skin smooth. Lateral line complete,
midlateral; canal curves slightly dorsally on caudal-fin base; in humeral region, canal with numerous
accessory canals and pores.

Head elongate, progressively depressed anteriorly. In lateral view, ventral surface of head nearly
horizontal. Gill opening wide, extending from exposed surface of posttemporal to anterior of isthmus.
Gill membranes free from, and not attached across, isthmus. Branchiostegal rays 12. Bony elements
of dorsal surface of head covered with only thin skin; bones readily visible, ornamented with fine,
irregular, radial grooves. Midline of cranium with elongate fossa, extending from posterior of snout
nearly to origin of supraoccipital spine; fossa occupied by slender, elongate anterior and posterior

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FiGurE 5. Sperata aorella, CAS(SU) 34852, 234 mm, Calcutta. Illustration by Alison Schroeer.

fontanels, separated by wide epiphyseal bar. Lateral fontanel shorter than diameter of orbit. Supraoc-
cipital spine elongate, lateral margins nearly parallel past basal constriction, tip rounded; surface
ornamented like that of other exposed bony elements of head. Cephalic lateral-line canal system with
extensive branching of most canals; integument covering fontanels, cheeks, and olfactory chamber
thoroughly covered with canals and pores.

Barbels in four pairs. Maxillary barbel long, slender, without medial membrane; barbel extends
at least to middle of caudal peduncle, and may reach caudal-fin margin. Nasal barbel slender, short;
extending at least to anterior margin of orbit, but not past middle of eye. Inner mental-barbel base
closer to outer mental barbel than to midline and slightly thicker and longer than nasal barbel; barbel
extending to past vertical through middle of eye. Outer mental barbel thicker and longer than inner
mental barbel, extending past pectoral-fin origin, its base about one eye diameter posterolateral of
inner mental barbel.

Eye ovoid, horizontal axis longer; located entirely in dorsal half of head. Upper margin of orbit
nearly reaching dorsal profile of head. Eye diameter about one-third of snout length and one-half
interorbital width. Orbit with free margin.

Snout long and depressed; lateral margins nearly parallel; anterior margin truncate, slightly
rounded in some individuals. Mouth subterminal, premaxillary teeth all exposed even when mouth
is closed. Oral teeth small, sharply pointed, irregularly arranged on tooth-bearing surfaces. Premax-
illary tooth band shallowly arched, width of band approximately equal throughout its length. Dentary
tooth band as broad as premaxillary band at symphysis, but tapering laterally. Palatal tooth patch
continuous across midline, but notched medially; anterior margin smoothly arched, tapering laterally
to point that extends posteriorly well beyond level of premaxillary band; band width near midline
broader than premaxillary band. Gill rakers of first arch long, slender and closely spaced; reduced to
rudiments anteriorly on lower arm. Gill rakers 15 to 17 on lower arm plus 5 or 6 on upper arm. Rakers
present only on outer face of first two arches; on both faces of next two arches.

Dorsal fin centered above middle of standard length. Dorsal-fin base shorter than length of first
dorsal-fin branched ray. Dorsal-fin margin straight, first branched ray longest, more than twice length
of last ray. Last dorsal-fin ray without posterior membranous connection to body. Dorsal fin with
spinelet, spine, and 7 branched rays. Dorsal-fin spine long, straight, slender; spine margin smooth

FERRARIS AND RUNGE: SOUTH ASIAN BAGRID CATFISH GENUS SPERATA 415

anteriorly and laterally; with fine serrations on distal half of posterior edge. Spine slightly shorter than
first branched ray. Tip of adpressed spine just reaches adipose-fin origin. Distance between bases of
dorsal adipose fins less than dorsal-fin base length. Interneural shield small, narrow; much shorter
than supraoccipital spine (Fig. 2c). Superficial ossification of dorsal-fin pterygiophores broadly united
across midline, and forming a crescent shaped bone (Fig. 4c).

Adipose-fin base length about equal to that of dorsal fin; adipose-fin height about one-third its
length. Adipose-fin margin slightly convex for most of its length, posterior end deeply incised.

Caudal fin deeply forked; lobes pointed and asymmetrical, upper lobe continued as filament when
intact. Middle rays approximately one-third as long as unbranched principal ray of lower lobe.
Principal caudal fin with rays: 1,7,8,1. Procurrent rays symmetrical and extend only slightly anterior
to fin base.

Anal fin located ventral to posterior half of adipose fin. Fin margin slightly convex; first branched
ray longest, about twice length of last ray. Last ray without posterior membranous connection to body.
Anal fin with 3 or 4 unbranched, and 9 branched rays.

Pelvic-fin origin at vertical through posterior end of dorsal-fin base. Pelvic-fin margin straight,
first branched ray longest: posterior-most ray about two-thirds length of first ray. Pelvic fin with one
unbranched and five branched rays. Tip of adpressed fin not reaching to anal-fin origin.

Pectoral fin with stout spine, sharply pointed at tip. Anterior spine margin smooth, posterior
margin with fine serrations along entire length. Pectoral-fin margin straight anteriorly, convex
posteriorly. First branched ray longer than spine and approximately 2’ times as long as last ray. Fin
with 9 (rarely 10) branched rays.

COLOR IN PRESERVATIVE. — Body silvery grey to brown dorsally and on upper half of lateral
surface, white ventrally. Demarcation between grey and white regions distinct, occurring ventral to
lateral line along most of body length, but along lateral line on caudal peduncle. Dorsal surface of
head brown; brown regions restricted to exposed cranial bones and around orbit. Posterior margin of
orbital flap white. Ventral surface of head and abdomen pale, without dark pigmentation.

Dorsal fin mostly pale; leading edge of spine and distal portion of rays dusky. Dusky region of
anterior rays broader than on more posterior rays, but less that distal one-quarter of rays darkened.
Adipose fin mostly dusky; fin margin with fine, distinct, dark terminal band. Posterior portion of fin
with ovoid black spot, smaller in diameter than orbit. Caudal peduncle with darkened area just ventral
to adipose-fin spot. Caudal fin generally dusky; white on ventral unbranched principal ray and ventral
procurrent rays. Anal fin pale basally, becoming increasingly dark distally; leading edge of fin pale.
Pelvic fin pale, with broad, diffuse dusky subterminal band. Pectoral fin pale basally, becoming
increasingly dark distally; little or no pigment on innermost rays. Dorsal surface of spine dark, pale
ventrally.

REMARKS. — Sperata aorella has been overlooked as a valid species since shortly after it was
first described. The name was placed in the synonymy of Macrones aor by Ginther (1864), without
comment. Shortly thereafter, Day (1877) placed the name in the synonymy of Macrones seenghala,
also without explanation or comment on Giinther’s action. More recently, the name Bagrus aorellus
has either been placed in synonymy of Sperata seenghala (e.g., Misra 1976), or ignored (e.g., Jayaram
1954; Jayaram 1973).

The general appearance of this species is somewhat intermediate between the two commonly
recognized Indian species, Sperata aor and S. seenghala. The maxillary barbel extends nearly to the
caudal fin, as in S. aor. However, the snout margin is clearly truncate, which is characteristic of S.
seenghala. We therefore assume that researchers who encountered this species had difficulty in
choosing which name to assign it.

Use of the name Sperata aorella for this species is of necessity somewhat tentative, inasmuch as
we have not been able to locate the types of this species. Most of the types of species described in
Blyth (1858) are not accounted for (e.g., Eschmeyer 1998) and the primary repository for Blyth

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specimens, the Zoological Survey of India, Calcutta, is not known to have specimens that may be
types of Blyth’s Bagrus aorellus (Menon and Yazdani 1968). Nevertheless, Blyth’s description,
although brief, fits the species that we are hereby associating with his name. Sperata aorella was said
to be somewhat thicker than S. aor and, most notably, to have a “more developed” occipital process
and “small bony plate, not exceeding the occipital process in breadth.” We interpret the former
statement to refer to the length of the supraoccipital spine, which is noticeably longer in S. aorella
than in either of its Gangetic congeners. The small bony plate, which we are calling the interneural
shield, is quite massive in the other Gangetic species, but is shorter, and no wider, than the
supraoccipital spine in S. aorella. Blyth also noted differences in the form of the exposed portion of
the first dorsal-fin pterygiophore, which is broadly united across the midline in S. aorella (Fig. 4c),
but divided medially on the surface by thick skin in S. aor (Fig. 4b).

Because of the scarcity of recent literature referring to S. aorel/la, it is impossible to determine
how much of the literature on Sperata seenghala or S. aor refers instead to this species, which is
reasonably abundant in the Ganges River delta.

DISTRIBUTION. — Known from the Ganges River delta of Bangladesh and India, extending
inland at least as far as Bisrampur.

MATERIAL EXAMINED. — 10 specimens, 152-277 mm. BANGLADESH: UMMZ 187866 (2,
244-245 mm), Comilla, Pond at Hajiganj (ca. 18 mi N Chandpur). UMMZ 208590-S (1, skeleton,
not measured), Chandpur. INDIA: MCZ 39637 (1, 179 mm), Ganges, Calcutta. MCZ 8190 (1, 164
mm), Calcutta. CAS(SU) 14510 (1, 277 mm), Bisrampur, Central Province [= Bihar]. CAS(SU)
34852 (4, 153-234 mm), Calcutta.

Sperata seenghala (Sykes, 1839)
higse2 de A4dver 7.8

Platystoma seenghala Sykes, 1839a:164 (Type locality: India: Deccan; no types known to exist); 1839b:61;
1841:371, pl. 65, fig. 2.

Bagrus lamarrii Valenciennes, in Cuvier and Valenciennes, 1840a:407, pl. 415 [labelled /amarii] (Type locality:
India: Ganges River; holotype: MNHN A.9343).

Bagrus aorinus Valenciennes, 1840 (in Jacquemont, 1835—1844):pl. 17, figs. 1, la (Type locality: India; no
types known to exist).

Macrones lamarrii Giinther, 1864:79 (India, ?Afghanistan).

Macrones lamarri Day 1873:cclxi (Ganges and Jumna rivers).

Macrones seenghala Day, 1877:444 (India); 1889:149 (in part) (India).

Mystus seenghala Ataur Rahman, 1989:198, fig. 120 (Bangladesh); Shrestha, 1994:54, fig. 84 (Nepal: Terai
region).

Mystus (Osteobagrus) seenghala Jayaram, 1954:555 (in part) (India, Pakistan, ? Yunnan). Srivastava, 1968:79,
fig. 50 (India: Uttar Pradesh).

Aorichthys seenghala Jayaram, 1973:155, figs. 1b, 3b (India, Pakistan, ? Yunnan). Mirza, 1980:24 (Pakistan:
Punjab, Sind, Baluchistan; Azad Kashmir). Jayaram, 1981:205, fig. 98, 99b (India, Pakistan, Bangladesh).
Talwar and Jhingran 1991:548, unnumbered plate (India, Bangladesh, ?Afghanistan, Nepal, Pakistan).

Mystus (Aorichthys) seenghala Misra, 1976:79, fig. 16 (India).

Aorichthys aor (not Hamilton) Mirza, 1980:24 (in part) (Pakistan: Punjab, Sind).

Aorichthys aor sarwari Mirza, Nawaz, and Javed, 1992:211 (Type locality: Pakistan: Ravi River at Head Balloki,
Kasur District; holotype: GCM F-19 [not seen]).

DIAGNOSIS. — Sperata seenghala differs from its congeners in that the eye is situated completely
in the anterior half of the head (Fig. 2d). It has more precaudal vertebrae (21—23) than any of its
congeners, and it is the only Sperata species in which the length of the supraoccipital spine 1s less
than that of the interneural shield. In addition, the adipose fin is relatively shorter than that of its
congeners, with its base approximately equal to, or only slightly longer than, the dorsal-fin base.

FERRARIS AND RUNGE: SOUTH ASIAN BAGRID CATFISH GENUS SPERATA 417

Additional characters help distinguish this species from each of its congeners. Sperata seenghala
differs from S. acicularis in having fewer caudal vertebrae (28-32, vs. 33-37), fewer pectoral-fin rays
(8 or 9, vs. 10, rarely 9) and modally fewer branched anal-fin rays (9, vs. 10). Sperata seenghala has
fewer pectoral-fin rays than does S. aor (8 or 9, vs. 10 or 11), and modally fewer gill rakers (14, vs.
19). Sperata seenghala differs from S. aorella in having more preanal vertebrae (29-32, vs. 25-27),
more total vertebrae (S0—53, vs. 47-49), and fewer gill rakers (13—15, vs. 20-22).

DESCRIPTION. — Body long, slender; body depth at dorsal-fin origin only slightly greater than
that anterior to adipose-fin origin, gradually tapering posteriorly. Ventral surface of head and body
flat to anal-fin base. Body slightly compressed and triangular in cross section across abdomen,
compressed and ovoid through caudal region. Caudal peduncle moderately narrow, depth approxi-
mately twice diameter of eye. Anus and urogenital openings located at vertical through middle of
adpressed pelvic fin; distance from these openings to anal-fin base greater than anal-fin base length.
Skin smooth. Lateral line midlateral and complete, curving slightly dorsally on caudal-fin base. In
humeral region, lateral line with numerous accessory canals and pores.

Head elongate, progressively depressed anteriorly. Head length one-third of standard length.
Profile of head acutely triangular in lateral view, with ventral surface of head nearly horizontal. Gill
opening wide, extending from exposed surface of posttemporal to beyond isthmus. Gill membranes
free from, and not attached across, isthmus. Branchiostegal rays 12 or 13. Bony elements of dorsal
surface of head covered with only thin skin; bones readily visible and ornamented with fine, irregular,
radial grooves. Midline of cranium with elongate fossa, extending from posterior of snout nearly to
base of supraoccipital spine; fossa occupied by anterior and posterior fontanels, separated by wide
epiphyseal bar. Lateral fontanel length less than orbital diameter. Supraoccipital spine relatively short,
parallel sided with blunt posterior tip. Spine of smaller specimens acutely pointed, with wide base.
Cephalic lateral-line canal system with extensive branching of most canals; integument covering
fossa, cheeks, and olfactory chamber thoroughly covered with canals and pores.

Barbels in four pairs. Maxillary barbel long, slender, without medial membrane; barbel extends
at least to dorsal-fin origin and may reach past dorsal-fin base in adults; in small individuals (e.g.,
CAS(SU) 41083, 108-130 mm), barbels reach to adipose-fin base. Nasal barbel slender, short; not
reaching anterior margin of orbit. Inner mental barbels originate close to midline, separated by
diameter of pupil; barbels thicker and much longer than nasal barbel; extending to, and usually past,
posterior margin of orbit. Outer mental barbel originates about one eye diameter posterolateral of
inner mental barbel; barbel thicker and slightly longer than inner mental barbel, not reaching
pectoral-fin origin.

Eye ovoid, horizontal axis longest; located entirely in dorsal half of head and anterior to middle
of head length. Upper margin of orbit nearly reaching dorsal profile of head. Eye diameter one-half
snout length and slightly less than interorbital width in small individuals. In larger specimens, eye
diameter about one-third of snout length and only about one-half interorbital width. Orbital margin
free.

Snout long and depressed, lateral margins parallel. Anterior snout margin distinctly truncate.
Mouth subterminal, anterior margin of premaxillary tooth patch exposed when mouth closed. Oral
teeth small, sharply pointed, in irregular rows on all tooth-bearing surfaces. Premaxillary tooth band
nearly straight along anterior margin. Tooth band of dentary as broad as premaxillary band at midline,
gently curved and tapering laterally. Palatal tooth patch continuous across midline; anterior margin
nearly straight, with straight, obliquely directed lateral margin. Tooth patch broader than premaxillary
band at midline, widening laterally and then tapering to sharp point posterolaterally. Gill rakers of
first arch long, slender, and closely spaced. Gill rakers 11 (rarely 12) on lower arm plus 3 (rarely 2
or 4) on upper arm. Rakers present only on outer face of first two arches, on both faces of next two
arches.

418 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 10

FIGURE 6. Sperata seenghala, illustration from Sykes (1841, pl. 65, fig. 2), presumed holotype.

FiGuRE 7. Sperata seenghala, illustration from Cuvier and Valenciennes (1840, pl. 415), holotype of Bagrus lamarrii
Valenciennes, 1840.

Dorsal fin centered above middle of standard length. Dorsal-fin base shorter than length of first
branched ray. Dorsal-fin margin straight; first branched ray longest, more than twice length of last
ray. Last dorsal-fin ray without posterior membranous connection to body. Dorsal fin with spinelet,
spine, and 7 branched rays. Dorsal-fin spine long, straight, slender; spine margin smooth anteriorly
and laterally, with fine serrations on distal half of posterior edge. Spine slightly shorter than first
branched ray; adpressed spine tip falls far short of adipose-fin origin. Distance between dorsal- and
adipose-fin bases approximately equals dorsal-fin base length. Interneural shield ovoid, longer than
wide; posterior end wider and more bluntly rounded than anterior end (Fig. 2d). Superficial ossifica-
tion of dorsal-fin pterygiophores broadly united across midline; anterior margin broadly rounded (Fig.
4d) or incised anteriorly.

Adipose-fin base equal to, or slightly longer than, that of dorsal fin; fin height about one-third of
its length. Adipose-fin margin straight anteriorly, then slightly convex following abrupt angular
transition; posterior end deeply incised.

Caudal fin deeply forked, lobes pointed and symmetrical, except for filamentous extension of
upper lobe. Middle rays approximately one-third as long as unbranched principal ray of lower lobe.

FERRARIS AND RUNGE: SOUTH ASIAN BAGRID CATFISH GENUS SPERATA 419

FIGURE 8. Sperata seenghala, illustration from Jacquemont (1835-1844, pl. 17, fig. 1) holotype of Bagrus aorinus
Valenciennes, 1840.

Principal caudal fin with rays: 1,7,8,1. Procurrent rays symmetrical and extend only slightly anterior
to fin base.

Anal-fin origin ventral to posterior half of adipose fin. Anal-fin margin straight; first branched
ray longest, about twice length of last ray. Last anal-fin ray without posterior membranous connection
to body. Anal-fin with 3 or 4 unbranched, and 8 to 10 branched rays.

Pelvic fin-origin at vertical through posterior end of dorsal-fin base. Pelvic-fin margin slightly
convex, second branched ray longest; posterior-most ray about two-thirds length of first ray. Pelvic
fin with one unbranched and five branched rays. Tip of adpressed pelvic fin not reaching to anal-fin
origin.

Pectoral fin with stout spine, sharply pointed at tip. Anterior spine margin smooth, posterior
margin with moderately strong serrations along entire length. Pectoral fin margin straight anteriorly,
convex posteriorly. First branched pectoral-fin ray longer than spine and approximately 2% times as
long as last ray. Pectoral fin with 9, rarely 8, branched rays.

COLOR IN PRESERVATIVE. — Body silvery grey to brown on upper half of lateral surface, white
or pale ventrally. Grey or brown pigmentation resulting from fine black pigment that extends to lateral
line on caudal peduncle and somewhat more ventrally for most of body. Dorsal surface of head brown;
head pigmentation most prominent on exposed cranial bones, but also extending to level of ventral
margin of orbit and on entire opercle. Ventral surface of head and abdomen pale. Humeral region with
diffuse, eye-sized, dark spot at upper extent of gill opening.

Dorsal fin dusky for distal half, pale basally. Adipose fin dusky; leading edge and margin with
distinct terminal band. In large individuals, adipose fin coloration similar to that of dorsal part of
body; smaller individuals distinctly paler. Posterior portion of fin with ovoid black spot; spot
somewhat smaller than eye diameter. Spot surrounded by pale halo, especially along dorsal and
posterior margin. Caudal peduncle with blackened area just ventral to adipose-fin spot, appearing as
extension of spot. Caudal fin with scattered pigmentation, denser on outer branched rays of ventral
lobe and base of dorsal lobe; ventral unbranched principal ray and ventral procurrent rays white. Anal
fin pale, with little or no pigmentation. Pelvic fin with fine, scattered pigment, somewhat denser
basally. Pectoral fin with scattered pigment dorsally; pale ventrally.

REMARKS. — Platystoma seenghala Sykes (1839a) was described from the Mota Mola River in
the upper reaches of the Krishna River basin. Apparently, Sykes had a single specimen in hand, as he
reported a specimen size (84 inches) while remarking that the species grows to a much larger size.
This presumed holotype is apparently lost, as it has not been reported on in subsequent publications,
and the whereabouts of it, along with virtually all other Sykes types, is unknown (see Eschmeyer
1998). The published illustration of the presumed holotype in a more thorough account of the species

420 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 10

(Sykes 1841, pl. 65, fig. 2; reproduced here as Fig. 6) shows a fish that quite closely resembles the
species which is generally associated with the name. The snout is distinctly truncated and the eye is
located in the anterior half of the head. As noted previously by Jayaram (1954), Sykes mentioned and
illustrated a puzzling condition of the pelvic fin. He reported that the “fish is remarkable for having
the first ray of the ventral fins, as well as that of the pectoral, serrated posteriorly” (Sykes 1841:372).
This condition was not found in the pelvic fin of any specimens examined by Jayaram or us. In
addition, no catfish known to us has a spine or serrated ray in its pelvic fin, which makes it unlikely
that Sykes was referring to another species.

The specimens we examined from the Krishna River basin are about the same length as, or shorter
than, the one reported on by Sykes, and therefore exhibit characteristics of the presumed juvenile
form. The supraoccipital spine is slender and the interneural shield is not as broad as in larger
specimens. The maxillary barbel extends past the base of the adipose fin and, therefore, well beyond
the dorsal fin, as is characteristic of larger specimens. We have not been able to examine larger
specimens from the Krishna, and therefore must infer that the comparatively larger interneural shield
and shorter barbel that characterizes larger specimens from other drainages also occurs in specimens
from the Krishna.

We place several nominal species in the synonymy of Sperata seenghala, despite the paucity of
relevant type material. Bagrus lamarrii Valenciennes is clearly this species and has been widely
regarded as such (e.g., Day 1877; Jayaram 1954). The holotype, a dried mounted specimen, exhibits
all of the externally visible diagnostic characters that we report for Sperata seenghala. These
characters are also clearly seen in the published illustration of the specimen (reproduced here as
Fig.a7).

The name Bagrus aorinus Valenciennes (in Jacquemont 1840) is based solely on an illustration
(reproduced as Fig. 8), for which there is no associated text, and no type specimens have been reported
(e.g., Bertin and Esteve 1950; Daget 1984). For this study we examined and failed to uncover a Sperata
specimen in the Museum national d’ Histoire naturelle in Paris that matched the figured specimen.
However, the figure is clearly that of a specimen of Sperata seenghala, as shown by the anteriorly
placed orbit, the large interneural shield, the short maxillary barbel, and the short adipose-fin base.
Contrary to current practice, we date the name Bagrus aorinus to 1840 following the study by Daget
(1984). Therein, Daget noted that Valenciennes cited the Jacquemont plates in volume 15 of Histoire
Naturelle des Poissons (Cuvier and Valenciennes 1840b), but not in volume 14 (Cuvier and
Valenciennes 1840a), even though illustrations of several fishes described in volume 14 were among
those included in the Jacquemont plates. Bailey (1951) determined the date of publication of volume
15 (Cuvier and Valenciennes 1840b) to be November, 1840, and that of volume 14 to be January
1840. This suggests that the Jacquemont plates were published some time before November, 1840,
but after volume 14 was submitted for publication (presumed to be sometime in 1839). However, for
purposes of priority, unless evidence of an earlier publication date for the Jacquemont plates comes
to light, it must be assumed that they were published sometime in 1840 but before November of that
year. Thus, the 1841 publication date for the plates, which was first reported in Bertin and Esteve
(1950) and used in Eschmeyer (1998) is incorrect, but the name Bagrus aorinus Valenciennes (in
Jacquemont, 1840) must be considered to have been published after Pimelodus seenghala Sykes,
1839, and Bagrus lamarrii Valenciennes, 1840.

Mirza (1990), and Mizra, Nawaz, and Javed (1992) demonstrated that only one species of Sperata
occurs in the Indus River drainage. Although we examined far fewer specimens than in those studies,
we independently reached this same conclusion. Based on the shape of the snout and the number of
caudal-fin rays, Mirza Nawaz, and Javed (1992) concluded that the Indus River specimens were
similar to Sperata aor, but differed sufficiently in the length of the maxillary barbel that they were
assigned to a new subspecies, Aorichthys aor sarwari. In our study, we found snout shape variation
sometimes difficult to interpret, as the snout of many specimens were deformed by preservation, and

FERRARIS AND RUNGE: SOUTH ASIAN BAGRID CATFISH GENUS SPERATA 421

that caudal-fin ray counts did not vary among the species. Thus, the decision to compare the Indus
River specimens only to Sperata aor was not appropriate. Instead, the Indus River specimens we
examined did not differ from specimens of Sperata seenghala from throughout the Indian region.
However, as mentioned above, our study was done on a relatively small sample size. If a more
thorough comparison of the various populations of Sperata seenghala indicated that the Indus River
specimens represented a distinct species, the name Sperata sarwari would be its valid name.

DISTRIBUTION. — Sperata seenghala is widely distributed in the Ganges and Indus river systems,
and also occurrs in several major rivers in peninsular India at least as far south as the Krishna River
(Jayaram 1981; Talwar and Jhingran 1991). Jayaram, Venkateswarlu and Ragunathan (1982) indi-
cated that one specimen of this species was taken from the Cauvery River, and they implied that the
species may have been introduced recently into rivers south of the Krishna River. Perhaps based on
the report of Jayaram et al. (1982), Talwar and Jhingran (1991) listed the Cauvery River in the
distribution of this species, without further comment.

Several accounts of the presence of Sperata seenghala in Yunnan, China, appear to be based on
a single report by Chaudhuri (1911). Therein, the species was said to have been collected from Lake
Tali Fu [= Er Hai], a high elevation lake that drains into the Mekong River system of western Yunnan.
The species was neither described nor illustrated, leaving some doubt about its identity. Furthermore,
no additional reports of S. seenghala, or any of its congeners, have been made for any locality within
Yunnan and the species was not included in the comprehensive inventory of Yunnan fishes (Chu and
Chen 1989). Therefore we conclude that the report in Chaudhuri (1911) is probably incorrect. Either
the specimen collected was not a Sperata, or the locality information associated with the specimen
was incorrect. A large portion of the collection reported on by Chaudhuri (1911) was from Bhamo,
Myanmar; it is possible that the specimen was taken there, rather than in Er Hai. If so, Chaudhuri
(1911) may have actually collected a specimen of Sperata acicularis.

One specimen from the Swedish Museum of Natural History (NRM 18809) is clearly a Sperata
seenghala, but it is reported to come from “probably Mandalay or Yangon areas.” If the locality
information is correct, it represents a range extension for S. seenghala, as well as the possibility of a
second species of Sperata in the Ayeyarwaddy River system.

MATERIAL EXAMINED. — 18 specimens, 108-720 mm. BANGLADESH: AMNH 56290 SD (2,
dry skeletons, not measured). UMMZ 208295 (1,301 mm), Comilla, Meghna River downstream from
Gumti River mouth, 23°19'N, 90°38’E. INDIA: CAS 62079 (1, 173 mm), Karnataka State, Tun-
gabahdra (Bellary?) District, Tungabahdra River and Reservoir at Hospet and Kampi (Krishna River
basin). MCZ 8185 (1, 259 mm), Sutle; River near Loodina [= ?Ludhiana]. CAS(SU) 41083 (2,
108-130 mm), Poona District. CAS(SU) 41084 (1, 190 mm), Lower Anicut River, Madras Pres.
INDIA/PAKISTAN: MCZ 22203, (1, 257 mm), Bengal, “North India” (possibly including Paki-
stan), sub-Himalayan region. MCZ 22208 (1, 396 mm), Bengal, “North India” (possibly including
Pakistan), sub-Himalayan region. MNHN A.9343 (1 (holotype of Bagrus lamarrii), 720 mm),
Ganges. PAKISTAN: CAS 24247 (1, 284 mm), Indus River, 5 mi N of Sukkur. MCZ 22185 (5,
228-442 mm), Punjab, Chenab River. MYANMAR: NRM 18809 (1, 221 mm), “probably Mandalay
or Yangon areas.”

ACKNOWLEDGMENTS

Most of the research for this study was undertaken while the second author was an intern at the
California Academy of Science’s Summer Systematics Institute of 1998, which was supported by the
National Science Foundation (BIR 9531307). Visits by the senior author to various museums to
examine specimens for this study was facilitated by Inhouse Research Grants from the California
Academy of Sciences, a Collection Fellowship from the Australian Museum, and a Collection Study
Grant from the American Museum of Natural History. We thank the following individuals for

422 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 10

permitting us to examine specimens in their care: Melanie Stiassny, Barbara Brown and Radford
Arrindal (AMNH); Jeff Leis, Doug Hoese, and Mark McGrouther (AMS); Dominique Didier and
William Saul (ANSP); Darrell Siebert (BMNH); William Eschmeyer and Tomio Iwamoto (CAS);
Karsten Hartel (MCZ); Jean-Claude Hureau and Patrice Pruvost (MNHN); Sven Kullander and Eric
Alhander (NRM); Douglas Nelson (UMMZ). We thank Beatrice Parisi (MNHN) for her assistance
in gaining access to rare literature in Paris. David Catania and Jon Fong (CAS) prepared radiographs
for our examination and Alison Schroeer prepared many of the illustrations published here while
serving as Academy Fellow Intern in Scientific Illustration for 1998. We thank Dave, Jon, and Alison
for their help. The paper benefitted from comments and suggestions by Mario de Pinna (MZUSP),
Sven Kullander (NRM), Alan Leviton (CAS), John McCosker (CAS), Darrell Siebert (BMNH), and
Richard Vari (USNM).

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(Siluriformes: Bagridae) in Lake Malawi, Africa. Copeia 1992(4):1084—1088.

LYDEKKER, R. 1886. Indian Tertiary and Post-Tertiary Vertebrata. Part 8. Tertiary fishes. Memoirs of the
Geological Survey of India (ser. 10) 3(8):241-258, pls. 36-38.

McKaAYE, K. R. 1986. Trophic eggs and parental foraging for young by the catfish Bagrus meridionalis of Lake
Malawi, Africa. Oecologia 69:367—369.

MENON, A. G. K. AND G. M. YAZDANI. 1968. Catalog of type-specimens in the Zoological Survey of India. Part
2: Fishes. Records of the Zoologica! Survey of India 61 (for 1963) (1-2):91-190.

MIRZA, M.R. 1980. The systematics and zoogeography of the freshwater fishes of Pakistan and Kashmir.
Proceedings of the First Pakistan Congress of Zoology 1-41.

. 1990. The freshwater fishes in Pakistan (in Urdu). Urdu Science Board, Lahore [not seen, citation
reproduced from Mirza et al. 1992].

Mirza, M. R., H. NAWAZ, AND M.N. JAVED. 1992. A note on the fishes of genus Aorichthys Wu with the
description of a new subspecies from Pakistan. Pakistan Journal of Zoology 24(3):211—213.

Misra, K. S. 1976. The fauna of India and adjacent countries. 2nd ed. Pisces, Vol. 3: Teleostomi: Cypriniformes:
Siluri. Controller of Publications, Delhi. xxi + 367 pp., 15 pls.

Mo, T. 1991. Anatomy and systematics of Bagridae (Teleostei) and siluroid phylogeny. Koeltz Scientific Books,
Koenigstein, Germany. vii + 216 pp., 63 pages of figures.

424 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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NEAVE, S. A., ed. 1939. Nomenclator Zoologicus, a list of the names of genera and subgenera in zoology from
the tenth edition of Linnaeus, 1758, to the end of 1935. Vol. 1. Zoological Society of London. 957 pp.

NEWMAN, E. 1841. Entomological notes (continued). The Entomologist 1(3):31—37.

PRASHAD, B. AND D. D. MUKERJI. 1929. The fish of the Indawgyi Lake and the streams of the Myitkyina District
(Upper Burma). Records of the Indian Museum 31(3):161—223.

SAIGAL, B. N. 1967. Studies on the fishery and biology of the commercial cat-fishes of the Ganga River system.
II. Maturity, spawning and food of Mystus (Osteobagrus) aor (Hamilton). Indian Journal of Fisheries,
Section A 11[for 1964] (1):1-44.

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188:1—622.

SRIVASTAVA, G. J. 1968. Fishes of eastern Uttar Pradesh. Vishwavidyalaya Prakashan, Varanasi, India. 163 pp.

SUNDARA RAJ, B. 1962. The extraordinary breeding habits of the catfish Mystus aor (Hamilton) and Mystus
seenghala (Sykes). Proceedings of the National Institute of Sciences of India 28B(3):193—200.

SUVATTI, C. 1950. Fauna of Thailand. [Thailand] Department of Fisheries. 1100 pp.

SYKES, W. H. 1839a. [On the fishes of the Deccan]. Proceedings of the Zoological Society of London, Part 6
(for 1838):157—165.

. 1839b. An account of the fishes of the Dukhun. Annals and Magazine of Natural History (new series)

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. 1841. On the fishes of the Dukhun. Transactions of the Zoological Society of London 2:349-378, pls.
60-67.

TALWAR, P. K. AND A. G. JHINGRAN. 1991. Inland fishes of India and adjacent countries. Oxford and IBH
Publishing Co., Pvt. Ltd., New Delhi, Bombay and Calcutta. xx + 1158 pp.

WHITE, E. I. AND J. A. Moy-THOMAS. 1940. Notes on the nomenclature of fossil fishes. Part I: Homonyms A-C.
Annals and Magazine of Natural History (ser. 11) 7(30):502—507.

WooDWARD, A. S. 1901. Catalogue of the fossil fishes in the British Museum (Natural History). Part IV,
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Civico de Storia Nationale de Genova, (ser. 2a) 9:129-362, pls. 7-11.

© CALIFORNIA ACADEMY OF SCIENCES, 1999
Golden Gate Park
San Francisco, California 94118

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES

Volume 51, No. 11, pp. 425-448, 10 figs., 2 tables. NOV 9g 1999 November 23, 1999
w

Preliminary Phylogeny of Halgerda (Nudibranchia: Halgerdidae)
from the Tropical Indo-Pacific,
with Descriptions of Three New Species

by

Shireen J. Fahey and Terrence M.Gosliner _
Department of Invertebrate Zoology and Geology, California Academy of Sciences
Golden Gate Park, San Francisco, California 94118

This paper discusses the systematics and phylogeny of the genus Halgerda from the
tropical Indo-Pacific. Morphological and anatomical data from Halgerda species were used
to construct a preliminary phylogeny. The phylogenetic analysis demonstrates the mono-
phyly of Halgerda and its relationship to its outgroup Asteronotus Ehrenberg, 1831.

Two new species of Halgerda from Thailand are named and described anatomically.
These two species, Halgerda stricklandi sp. nov. and H. bacalusia sp. nov., are highly derived
in several aspects of their morphology. They appear to be sister species and are most closely
related to Halgerda aurantiomaculata Allan, 1932. A third new species, Halgerda diaphana
sp. nov., is the sister taxon to the three taxa named above.

Due to external morphological similarities, comparison of the three new species is made
with the original descriptions of Halgerda carlsoni, Rudman 1978, H. malesso and H.
guahan Carlson and Hoff, 1993, and H. aurantiomaculata. The coloration, reproductive
system and the radular morphology of the new species differ significantly from H. carlsoni
and the other previously described Halgerda species. Although the two new species from
Thailand share some characteristics, they differ significantly in external and radular
morphology. The new species from Okinawa, Halgerda diaphana shares some external
characteristics with H. aurantiomaculata and H. guahan, but the reproductive morpholo-
gies differ significantly.

Fahey and Gosliner (1999) provided justification for placement of Halgerda, Asteronotus,
Aphelodoris, and Sclerodoris in the Halgerdidae. Valdes and Gosliner (submitted) have shown that
Sclerodoris is more closely related to other caryophyllidia-bearing dorids than to Halgerda and
Asteronotus.

All twenty-two described species placed in Halgerda are found in the tropical Indo-Pacific, from
South Africa as the western-most locality to the Hawaiian Islands in the east. Some of the species are
known only from the original description and there is one undescribed Halgerda species that has been
included in this analysis. That specimen is currently being studied.

This paper describes three new species of the genus, two from Thailand and one from Okinawa,
based on material collected from these locations and deposited in the Department of Invertebrate
Zoology of the California Academy of Sciences (CASIZ).

A preliminary analysis of the phylogeny of the genus Halgerda is presented for the first time,
using one outgroup, Asteronotus to establish the polarity of the characters.

425

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SPECIES DESCRIPTION

Family Halgerdidae Odhner, 1926
Genus Halgerda Bergh, 1880
Type species: Halgerda formosa Bergh, 1880, by monotypy.

Halgerda stricklandi sp. nov.
Pies. 1A. 2A, 3

MATERIAL EXAMINED. — HOLOTYPE: CASIZ 115298, one specimen, dissected, Ko Ha, Thailand, Andaman
Sea. 13 meters depth, November 1997. Collected and photographed by M. Strickland.

DISTRIBUTION. — This animal is known only from Thailand, Andaman Sea (this study).

ETYMOLOGY. — This animal is named for Mark Strickland who has kindly provided many slides
and specimens of nudibranchs to the authors. His thoughtfulness and generosity are greatly appreci-
ated.

NATURAL HISTORY. — The animal was found in the open on a rock wall.

EXTERNAL MORPHOLOGY. — The preserved animal studied (CASIZ 115298) measured 35 mm.
The body is firm and smooth, but rigid. The body profile is high and the dorsum has a series of
orange-tipped, conical tubercles arranged in a reticulate pattern. There are larger, orange-tipped,
conical tubercles at the junctions of the reticulate pattern. Just below the orange tips on the largest
tubercles is a faint white ring. The ground color of the dorsum and foot is whitish with a gray tinge.
There are small orange tubercles scattered thickly over the entire dorsum, extending down to the
mantle margin. There are four midline points connecting this network. The oral tentacles are short
and digitiform. The wide foot has an orange margin.

The long rhinophores are tapered towards the tips and angled posteriorly. The white translucent
rhinophores have 17-19 transverse lamellae that are outlined in black on the anterior side. The
rhinophores have yellow background coloration on the upper half. A vertical black line is present
along the posterior face of the entire length of each rhinophore.

The gill has four branchial leaves that are pinnate. These leaves have black spots on the anterior
side of the branches. Within the gill branches are numerous flattened, translucent structures that are
glandular. The anal papilla is long and tubular with black coloration on both the posterior and anterior
sides.

BUCCAL ARMATURE. — The buccal mass is not pigmented. The labial cuticle is smooth and
devoid of any jaw rodlets. The radular sac is elongate and extends well behind the posterior end of
the buccal mass. The radula (Fig. 3) of the holotype has a formula of 58 x 53.0.53 (CASIZ 115298).
The middle lateral teeth are hamate (Fig. 3C). The five or so inner lateral teeth are smaller and have
shorter hooks than the middle lateral teeth (Fig. 3B). The three outer lateral teeth are much smaller
than the inner and middle lateral teeth and have blunt denticles (Fig. 3A).

REPRODUCTIVE SYSTEM. — The reproductive system is triaulic (Fig. 2A). The wide ampulla is
elongate with a single curve. The ampulla narrows into the postampullary duct, which bifurcates into
the vas deferens and oviduct. The short oviduct enters the female gland mass. The short vas deferens
separates from the ampulla and widens into the large, greatly expanded portion of the glandular

_>

FiGuRE 1. Living animals. A. Halgerda stricklandi sp. nov. (CASIZ 115298). Specimen, 35 mm, from Ko Ha, Thailand,
Andaman Sea; 13 m depth, November 1997. Photo by M. Strickland. B. Halgerda bacalusia sp. nov. (CASIZ 115299).
Specimen, 40 mm, from Richelieu Rock, Thailand, Andaman Sea; 28 m depth, November 1997. Photo by M. Strickland. C.
Halgerda diaphana sp. nov. (CASIZ 070118). Specimen, 30 mm, from Okinawa, Ryukyu Islands, 2 km ENE southern tip;
23 m depth, March 1987. Photo by R. Bolland.

FAHEY AND GOSLINER: PRELIMINARY PHYLOGENY OF HALGERDA

427

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FIGURE 2. Reproductive systems. A. Halgerda stricklandi sp. nov. (CASIZ 115298; B. Halgerda bacalusia sp. nov. (CASIZ
115299): C. Halgerda diaphana sp. nov. (CASIZ 070118). Abbreviations: am = ampulla, be = bursa copulatrix, dp = dark
pigment, ej = ejaculatory duct, fgm = female gland mass, ga = genital atrium, p = penis, pr = prostate, rs = receptaculum
seminis, V = vagina, vg = vaginal gland. Scale bar = 1 mm.

prostate. The prostate consists of two distinct glandular types that are well differentiated, as is the
prostate of most other members of Halgerda. The muscular portion of the vas deferens exits the
prostate in a long, single convoluted duct, then widens sharply into the wide penial bulb. The female
gland mass is about the same size as the prostate gland. The short uterine duct emerges from the
female gland mass and joins the spherical receptaculum seminis near its base. The duct connecting
the receptaculum and the bursa is long and convoluted. The receptaculum seminis is smaller than the
spherical bursa copulatrix. The large, thin-walled bursa is covered by the prostate. The vaginal duct
that emerges from the base of the bursa copulatrix is long and thin. Near its exit adjacent to the base
of the male aperture, is an enlarged, and obviously glandular portion of the vagina. The genital aperture
is wide and large and the interior has dark pigmentation.

DISCUSSION. — Halgerda stricklandi has very similar external morphology to H. malesso
Carlson and Hoff, 1993 and H. carlsoni Rudman, 1978. Halgerda malesso has the same smooth but
firm, high body profile with angled tubercles tipped with orange, with orange spots scatted over the
dorsum. In H. malesso, there is a network of orange lines between the ridges. At the juncture of the
ridges are large, rounded tubercles with orange tips. Smaller orange-tipped tubercles are found along
the ridge crests. The foot margin is orange.

FAHEY AND GOSLINER: PRELIMINARY PHYLOGENY OF HALGERDA 429

Ficure 3. Halgerda stricklandi sp. nov. (CASIZ 115298). Scanning electron micrographs of radula. A. Outer lateral teeth,
scale bar = 25 uum; B. Inner lateral teeth, scale bar = 43 tum; C. Middle lateral teeth, scale bar = 75 um.

Halgerda carlsoni also has a high body profile with rounded tubercles tipped with orange or red
as in the type specimen, H. stricklandi. There is an opaque white ring below each orange or red tip.
Between the largest tubercles on the ridges are smaller orange or red-tipped tubercles. Scattered over
the dorsum are numerous small orange or red spots. The foot margin has a band of orange or red.
Halgerda stricklandi has the same orange-tipped tubercles as H. malesso and H. carlsoni, but they
are more numerous. All three species have the same grayish white background color. Halgerda
stricklandi differs from H. malesso and H. carlsoni in several other distinct ways:

430 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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1) The rhinophores of all three species are long and tapering, with the clavus angled posteriorly.
The rhinophores of H. stricklandi have a yellowish club and black-edged lamellae with a black line
on the posterior face. There is also a black line extending the length on the posterior side. Halgerda
malesso and H. carlsoni have brown spots with brown lamellae and lack a line on the posterior side.

2) The four gill branchia of H. stricklandi are outlined with black on the posterior side, and there
are black spots on the anterior side only. Halgerda malesso and H. carlsoni have brown spots around
the branchia and they are not outlined in black.

3) All three species have a glandular portion of the vagina, but the vaginal duct of H. malesso is
extremely wide compared to H. stricklandi and H. carlsoni. The vaginal ducts of H. stricklandi and
H. carlsoni are similar and both are narrower than that of H. malesso.

4) The penis and the vagina are not joined basally into a common atrium in H. malesso, whereas
in H. stricklandi and H. carlsoni they share a common opening. The penis in H. malesso is more
bulbous than that of H. stricklandi and H. carlsoni, both of which have a more tubular penis that is
larger than the vagina.

5) Both H. malesso and H. stricklandi have dark pigmentation on the genital atrium, however,
the location of the pigmentation differs. In H. malesso, the pigmentation is at the top of the atrium,
equidistant from both the vagina and penial openings. In H. stricklandi, the pigmentation lies in the
penis, at the junction with the vagina. Halgerda carlsoni has no pigmentation.

6) The three outer lateral teeth of both species are smaller than the middle and inner lateral teeth.
But all three outer teeth of H. stricklandi are denticulate whereas in H. malesso, only two are
denticulate and the third is simple. Rudman (1978) reported the three outer teeth of H. car/soni from
Fiji as degenerate and not denticulate. However, specimens examined from Papua New Guinea had
three denticulate outer teeth. In addition, H. stricklandi has very long, hamate, middle lateral teeth
with much thinner hooks than those of H. malesso and H. carlsoni. Both H. carlsoni and H. malesso
have very similarly-shaped inner and middle lateral teeth.

Halgerda bacalusia sp. nov.
Figs. 1B, 2B, 4

MATERIAL EXAMINED. — HOLOTYPE: CASIZ 115299, one specimen, dissected, Richelieu Rock, Thailand,
Andaman Sea. 28 meters depth, November 1997. Collected and photographed by M. Strickland.

DISTRIBUTION. — This animal is known only from Thailand, the Andaman Sea (this study) and
from a photograph of a specimen from Mindanao, Philippines (M. Miller, pers. comm.).

ETYMOLOGY. — The trivial name bacalusia is Latin for a kind of confection or sweet, which this
animal resembles with its bright, candylike colors.

NATURAL HISTORY. — This animal was found in the open on coral rubble substrate.

EXTERNAL MORPHOLOGY. — The preserved animal studied (CASIZ 115299) measured 40 mm.
The body is firm and smooth, but rigid. The body profile is high and the dorsum has a series of orange
ridges arranged in a reticulate pattern. There are four, orange-tipped, conical tubercles along the
midline at the junctions of the ridges. Just below the orange tips on the largest tubercles is a faint
yellow ring. The ground color of the dorsum and foot is whitish with a gray tinge. There are small
orange dots scattered thickly over the entire dorsum, extending near, but not to the mantle margin.
The small orange dots also terminate just below the orange coloration on the ridge crests. The white
ground color is visible on the sides of the ridges. The mantle is lined with a bright yellow marginal
band. The oral tentacles are short and digitiform. The wide foot has the same bright yellow margin
as the mantle.

The long rhinophores are tapered towards the tips and angled posteriorly. They are translucent
and have 17—19 transverse lamellae, which begin halfway up the stalk, which is ornamented with

FAHEY AND GOSLINER: PRELIMINARY PHYLOGENY OF HALGERDA

Ficure 4. Halgerda bacalusia sp. nov. (CASIZ 115299). Scanning electron micrographs of radula. A. Outer lateral teeth,
scale bar = 30 um; B. Outer lateral teeth, scale bar = 4.3 xm; C. Inner lateral teeth, scale bar = 75 jum; D. Middle lateral teeth,
scale bar = 100 um.

black on the anterior side. The rhinophores have yellow background coloration on the upper half. A
black longitudinal line is present on the posterior face and extends from the base to the club.

The gill has four branchial leaves that are highly pinnate. These leaves have black pigment on
the margins and posterior sides of the branches. The tips of the branches are white. Within the gill
branches are numerous flattened, translucent structures that are glandular.

432 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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BUCCAL ARMATURE. — The buccal mass is not pigmented. The radular sac is elongate and
extends well behind the posterior end of the buccal mass. The radula (Fig. 4) of the holotype has a
formula of 59 x 57.0.57 (CASIZ 115299). The middle lateral teeth are hamate, with long, thin hooks
(Fig 4D). The inner lateral teeth are hamate, shorter than, and gradually increasing in size towards
the middle of the half-row (Fig. 4C). The three outer lateral teeth are much smaller than the middle
teeth and are not hooked but are denticulate (Fig. 4A, B). The denticles are blunt and numerous.

REPRODUCTIVE SYSTEM. — The reproductive system 1s triaulic (Fig. 2B). The wide ampulla is
elongate and flattened. The ampulla narrows into the postampullary duct, which bifurcates into the
vas deferens and oviduct. The short oviduct enters the female gland mass. The female gland mass is
longer than the prostate gland. The short vas deferens separates from the ampulla and widens into the
large, widened portion of the glandular prostate. The prostate consists of two distinct glandular types
and they are well differentiated as in most other members of Ha/gerda. The muscular portion of the
vas deferens leaves the prostate in a long, single duct, then widens slightly into the penial bulb. The
short uterine duct emerges from the female gland mass and joins the spherical receptaculum seminis
near its base. The duct connecting the receptaculum and the bursa is long and coiled. The spherical
receptaculum seminis is much smaller than the thin-walled spherical bursa copulatrix. The bursa is
covered by the larger prostate. The vaginal duct that emerges from the base of the bursa copulatrix is
long and thin. Near its exit adjacent to the base of the male aperture, is an enlarged and obviously
glandular portion of the vagina. The genital atrium is wide and large and the interior has dark
pigmentation.

DISCUSSION. — Halgerda bacalusia has very similar external and internal morphologies to H.
carlsoni Rudman, H. aurantiomaculata Allan, and H. guahan Carlson and Hoff, but differs in several
ways. Rudman (1978) described H. carlsoni from Fiji. The type specimen was described as having a
smooth, ridged body, with a series of angled ridges. At the junctions of these ridges are red-tipped
tubercles, which have a ring of white below the red tip. Between the largest tubercles on the ridges
are smaller red-tipped tubercles. Scattered over the dorsum are numerous small red spots. The mantle
margin is white with a border of orange or red spots. The foot has an orange-red border band.

Willan and Brodie (1989) recently redescribed H. aurantiomaculata (Allan, 1932) from Fiji.
They described their two specimens as having a low body profile, with prominent pustules and a
mid-dorsal ridge connecting the pustules. The pustules were capped with bright orange tips and the
ridges had orange streaks along the summits. The mantle margin was orange and there were 3-8
orange spots of varying sizes in the depressions between the ridges. The foot had an orange margin.

Halgerda guahan Carlson and Hoff has the same firm, low body profile as H. aurantiomaculata.
However, while it does have the characteristic ridges, it does not have pustules. The ridges are edged
in yellow and the mantle margin and foot are edged in a thin, opaque white line.

Halgerda bacalusia also has a smooth, firm, high body profile with a series of angled orange
ridges. There is also yellow coloration alongside the orange lines. At the junctions of these ridges are
orange-tipped tubercles, which have a ring of yellow below the orange tip. Between the largest
tubercles on the ridges are smaller, orange-tipped tubercles. Scattered over the dorsum are numerous
small orange spots. The mantle has a bright yellow border, as does the foot.

In addition to the external coloration differences, Halgerda bacalusia differs from H. carlsoni,
H. aurantiomaculata and H. guahan in the following ways:

1) All four species have long, tapering rhinophores. The rhinophores of both H.car/soni and H.
bacalusia have a yellowish club, and the edges of the lamellae have black pigment. The rhinophores
of H. aurantiomaculata and H. carlsoni are covered with small, regularly-spaced, brown spots, which
are darker on the stalk than on the club. The brown spots on the rhinophores are larger on H. guahan
and the posterior side of the base has a thick brown streak on some specimens. Halgerda bacalusia
has a black posterior line. The rhinophoral sheath of H. guahan also has a yellow margin. Halgerda
carlsoni has 18-20 lamellae, all located near the tip, whereas H. bacalusia has 17-18 lamellae,

FAHEY AND GOSLINER: PRELIMINARY PHYLOGENY OF HALGERDA 433

covering half the stalk and H. aurantiomaculata has 25 lamellae. The clavus of the rhinophores is
angled posteriorly in all species.

2) The gill branches of H. aurantiomaculata, H. guahan and H. carlsoni have brown spots
whereas H. bacalusia has dark pigment and no distinct spots.

3) There is a glandular portion of the vagina in H. malesso, H. carlson, H. aurantiomaculata and
H. bacalusia. Halgerda guahan does not have this structure. The vaginal ducts of H. bacalusia, H.
guahan and H. carl/soni are thinner than either H. aurantiomaculata or H. malesso. Halgerda malesso
has a long, extremely wide vagina.

4) There is dark pigmentation on the genital atrium of H. aurantiomaculata, H. malesso and H.
bacalusia, whereas H. carlsoni does not have this pigmentation. There is a common genital atrium
in three species while in H. guahan the penis and vagina are separate to their distal ends.

5) The radular sac of H. carlsoni, H. aurantiomaculata and H. bacalusia is elongate. No
information is available on the radular sac of H. guahan or H. malesso.

6) The three outer lateral teeth of H. carlsoni are much smaller than the middle lateral teeth.
Although Rudman (1978) reported the three outer teeth as not denticulate, specimens examined from
Papua New Guinea did have three denticulate outer teeth. The outer lateral teeth of H. auwrantiomacu-
lata are also smaller than the middle lateral teeth and sometimes weakly bifid (Willan and Brodie,
1989). Both H. guahan and H. malesso have three outer lateral teeth which are smaller than the middle
teeth; two are denticulate and the third only slightly so. In all four species, the middle laterals are
hamate.

Although H. bacalusia shares similar characters with H. aurantiomaculata, H. carlsoni and H.
guahan, its unique combination of characters confirm it as a new species. The parsimony based
analysis of the Halgerda genus also confirms that H. bacalusia is more closely related to H. stricklandi
than to other members of the genus.

Halgerda bacalusia differs from H. stricklandi in the following ways:

1) External morphology. Halgerda stricklandi has orange-tipped conical tubercles, not connected
by ridges. Halgerda bacalusia has orange and yellow-tipped tubercles connected by orange crested
ridges. Halgerda bacalusia has a yellow-orange mantle margin and H. stricklandi has no coloration
along the margin. The orange spots on the dorsum of H. bacalusia are tiny, whereas those of H.
stricklandi are much larger and fewer.

2) Reproductive system. Halgerda stricklandi has a female gland mass approximately the same
size as the prostate. Halgerda bacalusia has a female gland mass that is larger than the prostate. The
ampulla of H. stricklandi is elongate with a single curve, and in H. bacalusia, the ampulla is elongate
and flat. The dark pigmentation on the genital atrium of H. stricklandi is near the opening of the genital
atrium, whereas the pigmentation on the genital atrium of H. bacalusia is near the juncture of the
vagina and the penis.

3) Buccal armature. Halgerda bacalusia has blunt denticles on the three outer lateral teeth, and
the denticles on H. stricklandi are sharper. The inner teeth of H. stricklandi are hamate with short
hooks and those of H. bacalusia are hamate with long hooks.

Halgerda diaphana sp. nov.
Rigs mliGs2CrS. 647

MATERIAL EXAMINED. — HOLOTYPE: CASIZ 079369, one specimen, 34 mm. 1 km WNW Onna Village,
Horseshoe Cliffs, Ryukyu Islands, Okinawa. 50 meters depth, September, 1991. Collected by R. Bolland.
PARATYPES: CASIZ 070118, one specimen, 30 mm, dissected. 2 km ENE southern tip, Ryukyu Islands, Okinawa.
23 meters depth, March, 1987. Collected and photographed by R. Bolland. CASIZ 074688, one specimen, 25
mm. | km WNW Onna Village, Horseshoe Cliffs, Ryukyu Islands, Okinawa. 60 meters depth, March, 1984.
Collected by R. Bolland. CASIZ 079368, one specimen, dissected, 39 mm. 1 km WNW Onna Village, Horseshoe
Cliffs, Ryukyu Islands, Okinawa. 27 meters depth, August, 1991. Collected by R. Bolland. CASIZ 084877, one

434 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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specimen, 49 mm. Seragaki Beach, 1.3 km ENE of Maeki-zaki, Ryukyu Islands, Okinawa. 47 meters depth,
October, 1991. Collected by R. Bolland. CASIZ 086598, one specimen, 37 mm. Seragaki Beach, 1.3 km ENE
of Maeki-zaki, Ryukyu Islands, Okinawa. 3 meters depth, May, 1992. Collected by R. Bolland. CASIZ 087900,
one specimen, 69 mm. Seragaki Beach, 1.3 km ENE of Maeki-zaki, Ryukyu Islands, Okinawa. 43 meters depth.
August, 1992. Collected by R. Bolland. CASIZ 089002, one specimen, 49 mm. 1.3 km ENE of Maeki-zaki,
Seragaki Beach, Ryukyu Islands, Okinawa. 3 meters depth. September, 1992. Collected by R. Bolland. CASIZ
089012, one specimen, dissected, 50 mm. | km WNW Onna Village, Horseshoe Cliffs, Ryukyu Islands,
Okinawa. 55 meters depth, December, 1992. Collected by R. Bolland. CASIZ 104698, one specimen, 29 mm.
1 km WNW Onna Village, Horseshoe Cliffs, Ryukyu Islands, Okinawa. 30 meters depth. November, 1994.
Collected by R. Bolland. CASIZ 105305, one specimen, dissected, 53 mm. Seragaki Tombs, Ryukyu Islands,
Okinawa. 25 meters depth. November, 1994. Collected by R. Bolland.

DISTRIBUTION. — This animal is known only from the Ryukyu Islands, Okinawa (this study).

ETYMOLOGY. — The trivial name diaphana is from the Greek word diaphanes meaning a gauzy
texture that is translucent or transparent, like the dorsum of this new species.

NATURAL HIsToRY. — This animal is found from 3-60 meters depth, on various substrates from
rock and coral rubble to silty sand. It has also been found on vertical rock walls and on the reef forefront
on coral rubble and in crevices at the reef edge.

EXTERNAL MORPHOLOGY. — The preserved animals dissected (CASIZ 070118, 079368 and
105305) are 30 mm, 25 mm, and 30 mm respectively, in length. The body is firm and smooth, but
rigid. Although the body profile is slightly arched, this species does not have the high body form
common in most other Halgerda. The dorsum has orange crested ridges arranged in a distinct
triangular pattern of three points per side of the dorsum. There are four pustules at the centerline of
the dorsum that join the orange crests. There are secondary orange lines in the concavities of the
ridges. The ground color of the dorsum and foot is translucent white, with the viscera clearly visible
through the dorsum. The mantle submargin is lined in bright orange, and some animals have yellow
coloration on the outside of the orange margin. The mantle edge is bright white. The oral tentacles
are short and digitiform. The wide foot has the same bright orange submargin as the mantle.

The long rhinophores are tapered towards the tips and the bulb is angled posteriorly. The white
translucent rhinophores have 18 transverse lamellae, which begin halfway up the stalk. The rhino-
phores have black speckles sprinkled liberally along their length. The extreme tip of the rhinophore
is white.

There are two main branchial leaves that are moderately pinnate. Each of the two main leaves
divides into three branches, with the posterior two being more pinnate than the anterior pair. The gills
have the same black speckles that completely cover the rhinophores. The extreme tip of each branchia
is white. Within the gill rachis are numerous flattened, translucent structures that are glandular. The
anal papilla is long and tubular with black speckles on both the posterior and anterior sides.

BUCCAL ARMATURE. — The buccal mass is not pigmented. The radular sac is elongate and
protrudes well behind the posterior end of the buccal mass. The radular formulae of the dissected
specimens are 53 x 43.0.43 (CASIZ 070118), 49 x 50.0.50 (CASIZ 079368) and 55 x 51.0.51
(CASIZ 105305). The outer and middle lateral teeth are hamate (Figs. 5, 6, 7). The 10 innermost
lateral teeth are hamate and small, then significantly larger towards the middle of the half-row (Figs.
5, 6, 7). The three outer lateral teeth are much smaller than the middle teeth and are not hooked but
some are denticulate (Figs. 5, 6, 7). The denticles are blunt and numerous.

REPRODUCTIVE SYSTEM. — The reproductive system is triaulic (Fig. 2C). The wide tubular
ampulla is elongate and lies tightly against the female gland mass. The ampulla narrows into the
postampullary duct, which bifurcates into the vas deferens and oviduct. The short oviduct enters the
female gland mass. The female gland mass is about the same size as the prostate gland. The short vas
deferens separates from the ampulla and widens into the large glandular prostate. The prostate consists
of two distinct glandular types, which are well differentiated as in most other members of Halgerda.

FAHEY AND GOSLINER: PRELIMINARY PHYLOGENY OF HALGERDA 435

FiGure 5. Halgerda diaphana sp. nov. (CASIZ 070118). Scanning electron micrographs of radula. A. Outer lateral teeth,
scale bar = 25 um; B. Inner lateral teeth, scale bar = 25 tum; C. Middle lateral teeth, scale bar = 60 um.

The vas deferens leaves the prostate in a long, single duct, then widens substantially into the long,
muscular portion of the penial bulb. The short uterine duct emerges from the female gland mass and
joins the pyriform receptaculum seminis at its base. The duct connecting the receptaculum and the
bursa is moderately long and convoluted. The pyriform receptaculum seminis is much smaller than
the thin-walled spherical bursa copulatrix. The bursa is covered by the larger prostate. The vaginal

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 11

FIGURE 6. Halgerda diaphana sp. noy. (CASIZ 079368). Scanning electron micrographs of radula. A. Outer lateral teeth,
scale bar = 15 um. B. Outer lateral teeth, scale bar = 43 um. C. Inner lateral teeth, scale bar = 43 tum. D. Middle lateral teeth,
scale bar = 60 yum.

FAHEY AND GOSLINER: PRELIMINARY PHYLOGENY OF HALGERDA

FiGureE 7. Halgerda diaphana sp. nov. (CASIZ 105305). Scanning electron micrographs of radula. A. Outer lateral teeth,
scale bar = 15 um; B. Outer lateral teeth, scale bar = 60 um; C. Inner lateral teeth, scale bar = 43 tum: D. Middle lateral teeth,
scale bar = 100 uum.

438 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 11

duct that emerges from the base of the bursa copulatrix is short and immediately widens into the very
large and glandular vagina. The openings of the vagina and the penis share the common genital atrium,
which has dark pigmentation half-way down its length on the interior of the duct.

DISCUSSION. — Halgerda diaphana shares similar external characteristics with H. auran-
tiomaculata and H. guahan. Carlson and Hoff (1993) described H. guahan from Guam and Willan
and Brodie (1989) redescribed H. aurantiomaculata from Fiji. Halgerda guahan and H. auran-
tiomaculata have the same firm, low body profile as 1. diaphana. However, while H. guahan has the
characteristic ridges, it does not have pustules. Halgerda aurantiomaculata has prominent pustules,
with four along the midline of the dorsum like H. diaphana. The ridges of Halgerda guahan are edged
in yellow and the mantle margin and foot are edged in a thin, opaque white line. Halgerda diaphana
and H. aurantiomaculata both have orange ridge crests and orange coloration submarginally along
the mantle margin. The orange submarginal band on the mantle of H. diaphana is edged in bright
white whereas the mantle margin of H. aurantiomaculata is orange. Halgerda aurantiomaculata has
orange spots of various sizes on the mantle in the ridge concavities, while H. diaphana has secondary
orange lines in the concavities.

There are several other distinct differences between H. diaphana, H. aurantiomaculata and H.
guahan:

1) The rhinophores of both 1. diaphana and H. aurantiomaculata have black speckles along the
entire length. Halgerda guahan has brown spots on the rhinophores which are larger and the posterior
side of the base has a thick brown streak on some specimens. The rhinophoral sheath of H. guahan
also has a yellow margin. The club of the rhinophores is tilted backwards in all three species.

2) There is a glandular portion of the vagina in H. aurantiomaculata, whereas the entire vagina
of H. diaphana is glandular. Halgerda guahan does not have this glandular structure. The vaginal
duct of H. guahan is longer and thinner than that of either H. aurantiomaculata or H. diaphana.

3) There is dark pigmentation on the genital atrium of both H. aurantiomaculata and H. diaphana
whereas H. guahan does not have this pigmentation (C. Carlson, pers. comm.). There is a common
genital atrium in H. aurantiomaculata and H. diaphana while in H. guahan the penis and vagina are
separate.

4) The outer lateral teeth of H. diaphana are much smaller than the middle lateral teeth, and are
denticulate. The outer lateral teeth of H. aurantiomaculata are also smaller than the middle lateral
teeth and sometimes weakly bifid (Willan and Brodie 1989). Halgerda guahan has three outer lateral
teeth which are smaller than the middle teeth; two are denticulate and the third only slightly so. In all
three species, the middle laterals are hamate.

Although H. diaphana shares similar characters with H. aurantiomaculata and H. guahan, its
unique combination of characters establishes it as a new species. The parsimony based analysis of
the Halgerda genus confirms that H. diaphana is a sister taxon to H. aurantiomaculata, H. bacalusia,
and H. stricklandi and is more closely related to them than to other members of the genus.

PHYLOGENETIC RELATIONSHIPS

Phylogeny of Halgerda

Species of Halgerda have been recently described by several authors (Rudman 1978; Bertsch
and Johnson 1982; Willan and Brodie 1989; Carlson and Hoff 1993; Yonow 1993; Gosliner and
Fahey 1998; and Fahey and Gosliner, 1999). Additional data were taken from the original publications
on Halgerda (Bergh 1880; Eliot 1904; Farran 1905). Valdés and Gosliner’s data (submitted) were
used for the character description of Asteronotus. In addition, the following species were examined
directly: Halgerda formosa Bergh, 1880; H. elegans Bergh, 1905; H. albocristata Gosliner and Fahey,
1998; H. willeyi Eliot, 1904; H. dalanghita Fahey and Gosliner, 1999; H. bacalusia Fahey and
Gosliner, 1999; H. carlsoni Rudman, 1978; H. tessellata Bergh, 1880; H. stricklandi Fahey and

FAHEY AND GOSLINER: PRELIMINARY PHYLOGENY OF HALGERDA 439

Gosliner, 1999; H. dichromis Fahey and Gosliner, 1999; H. toliara Fahey and Gosliner, 1999; H.
terramtuentis Bertsch and Johnson, 1982; H. sp. 2, undescribed species.

Thus, previous literature and direct observation and dissection of 13 species of Halgerda and
members of the outgroup have provided the information on the characters for the present study of the
phylogenetic relationships of Halgerda species. To establish the polarity of the morphological
characters (Table |) used in this study, one outgroup was selected based on a review of anatomical
characters. We used Asteronotus as the outgroup rather than Sclerodoris, based on the phylogenetic
study of Valdés and Gosliner (submitted).

The following characters were considered in this analysis:

[Note: Characters in parentheses were deleted from the first analysis, because they are phyloge-
netically uninformative. Those character descriptions preceded by an asterisk were deleted from a
second analysis that tested how color characters (and phylogenetically uninformative characters)
affect the tree topology. |

1. *Rhinophores——All species of Halgerda have elongate rhinophores. The outgroup taxon,
Asteronotus has short rhinophores and this character is considered plesiomorphic.

2. *Ridges——All Halgerda have dorsal ridges. These ridges can be low-lying and almost level
with the rest of the notum or they can be quite high and distinct. Asteronotus does not have ridges.
The absence of ridges is considered the pleisiomorphic state.

3. Tubercles——Rounded dorsal tubercles are absent in seven Halgerda species. Distinct rounded
tubercles are present on nine Halgerda species, while pointed tubercles are present on another seven
species. Asteronotus has prominent rounded tubercles, which are connected in concentric circles. The
absence of tubercles is considered pleisiomorphic.

4. Small marginal tubercles —Small tubercles are found along the mantle edge of four species
of Halgerda: H. brunneomaculata, H. formosa, H. tessellata, and H. toliara. The outgroup taxon,
Asteronotus does not have these marginal tubercles, and their presence is considered apomorphic.

5. *(Caryophyllidia——No species of Halgerda or Asteronotus have caryophyllidia. This charac-
ter is not applicable to the taxa being studied since caryophyllidia are absent from the outgroup and
ingroup taxa studied).

6. *Body color.—tThe plesiomorphic state of an orange-yellow or reddish body color is shared
by the outgroup taxon and four Halgerda species: H. brunneomaculata, H. xishaensis, H. dalanghita,
and H. toliara. All other Halgerda have a white to gray-white background color.

7. Ridge color—tThree Halgerda have white dorsal ridges: H. albocristata, H. xishaensis, H.
dalanghita. All others have orange or yellow ridges, while the outgroup taxon has the same coloration
over its entire dorsum. Ridges that are the same color as the dorsum are considered the plesiomorphic
state.

8. *Dark marginal lines—Black marginal lines are unique to four Ha/gerda: H. albocristata, H.
elegans, H. iota, and H. willeyi. The marginal lines lie perpendicular to the mantle edge. These
marginal lines are not present on the outgroup taxon. Lack of marginal lines is the plesiomorphic
state.

9. *Dark spots—Black dorsal spots are present on four Halgerda: H. brunneomaculata, H.
formosa, H. punctata, and H. dichromis. These spots can range from deep purple-black to dark brown.
The outgroup taxon does not have these distinguishing spots, and this state is plesiomorphic.

10. Gill branching —Only two Halgerda species share the tripinnate gill branching pattern with
the outgroup taxon. Those two are H. xishaensis and H. dalanghita. The tripinnate gill pattern is
considered plesiomorphic.

11. *Gill spots—Dark spots on the gill branches are considered the apomorphic state. Nine
species have these symmetrical round spots. Fourteen Halgerda species lack spots and share the
plesiomorphic state with the outgroup taxon.

440

TABLE |. Characters and states considered for the phylogeny of Halgerda.

Yr CDOOMAANDMNAHRWN —

we

— —
O Nn

NON
=) \OniCo]

>)
>}

Nhe
We

ys)

46

Character

Rhinophores

Ridges

Tubercles

Small marginal tubercles
Caryophyllidia

Body color

Ridge color

Dark marginal lines
Dark spots

Gill branching

Gill spots

Gill stripes

Gill tip color

Gill shape

Gill pinnation

Rhinophore base color

Rhinophore bulb color
Gill glands

Oral tentacles shape
Foot border color

Foot lines

Foot spots

Body shape

Secondary orange lines
Orange dots on dorsum
Radula sac shape
Buccal mass

Radula shape

Middle lateral teeth
Outer lateral teeth shape
Outer lateral teeth size
Fimbriate tooth length
Ampulla length

Vaginal duct length
Vaginal duct shape
Vagina duct termination
Vaginal base shape
Vaginal sphincter
Prostate

Differentiation of prostate
Ejaculatory duct
Receptaculum seminis
Tubercular vaginal glands
Penis shape

Uterine duct insertion
Prostate size

Atrium pigment
Mantle margin

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES

Volume 51, No. 11

_____ Plesiomorphic
0 = short
0) = absent
0 = absent
0 = absent
0 = present
0 = reddish or orange
0 = body color
0 = absent
() = absent
0 = tripinnate
() = absent
0 = absent
0 = white
0 = erect

0 = highly pinnate
0 = same as body color
0 = white

0 = same as gill color
0 = short and rounded

() = absent

() = absent

0 = absent

0 = broad

0 = absent

0 = absent

0 = short

0 = unpigmented

(0) = square

0 = hamate

0 = hamate

( = same size as middle laterals
0 = short

0 = short

0 = short

0 = narrow

0 = enters common atrium
0 = narrow

0 = absent

0 = one part

0 = poorly differentiated

0 = short

0 = same size as bursa copulatrix
0 = absent

0 = tubular

0 = base of bursa

0 = smaller than female gland

mass
0) = absent
0 = absent

ae

1 = elongate

| = present

1 = rounded, 2 = pointed

| = present

| = absent

| = whitish

1 = orange/yellow, 2 = white

| ='present

| = present

| = bipinnate

| = present

| = present

1 = black

| = flat

| = moderately, 2 = sparsely

pinnate

| = dark blotches, 2 = dark spots
3 = dark stripes

1 = black

| = opaque white

| = long and tapered

| = present

| = present

= present
= elongate
= pigmented
= rectangular
= elongate

= fimbriate

= wide

= separate duct

= wide

= present

= two parts

= well differentiated

l
l
l
l
l
l
l
l
l
l
l
1 = elongate
l
l
l
l
|
|
l
l
l
l
1 = more distal
1 = equal to female gland mass
| = present
= orange/yellow, 2 = white

FAHEY AND GOSLINER: PRELIMINARY PHYLOGENY OF HALGERDA 44]

12. Gill stripes—tLack of dark gill stripes is the plesiomorphic state, and twelve species share
this state with the outgroup. Ten species have dark stripes on the gill branches. The stripes run from
bottom to top, along the posterior of the gill rachis.

13. *Gill tips—AII but five Ha/lgerda species share the plesiomorphic character with the
outgroup taxon with white gill tips that are the same color as the rest of the gill. The five Halgerda
species that have black gill tips are: H. albocristata, H. elegans, H. formosa, H. iota, and H. toliara.

14. Gill shape——Only two Halgerda species have gill branchia which lie flat on the dorsum: H.
punctata and H. dichromis. All other species including the outgroup taxon have erect gill branchia.
This is considered the plesiomorphic state.

15. Gill pinnation—The plesiomorphic character state is a highly pinnate gill. This state is shared
by the outgroup taxon and H. dalanghita. All other Halgerda have a moderately pinnate gill except
for H. brunneomaculata and H. toliara which have a sparsely pinnate gill.

16. *Rhinophore base—The rhinophoral base is the same as the body color in Asteronotus. Three
Halgerda species share this plesiomorphic character state: H. albocristata, H. elegans, and H. toliara.
The three other character states: dark blotches, dark spots and dark stripes are distributed among the
other Halgerda species. This character is treated as unordered.

17. Rhinophore bulb.—A rhinophoral bulb that is the same color as the body 1s the plesiomorphic
state, and only Asteronotus has this character state. All Halgerda species have dark coloration on the
bulb.

18. (Gill glands——All Halgerda share the apomorphic character state of opaque white glands
internal to the gills. This character state could not be determined for H. xishaensis. The outgroup taxon
has gill glands which are not opaque.)

19. (Oral tentacles—Short rounded oral tentacles are plesiomorphic. Half of the Halgerda
species share this character state with the outgroup taxon. The character state could not be determined
from the literature for H. iota).

20. Foot border.— Asteronotus does not have a colored margin around the foot. Nine Halgerda
species have a yellow or orange foot margin. This character state could not be determined for H.
dichromis. Lack of a foot margin is considered plesiomorphic.

21. *Foot lines —Dark-colored foot lines are absent in the outgroup taxa, and in most Halgerda
species. However, H. iota, H. tessellata, and H. willeyi have dark lines along the edge of the foot. The
presence of dark lines on the foot is considered apomorphic.

22. *Foot spots—Over half of the Halgerda species have dark spots on the foot. Asteronotus
and the other half of the Halgerda species do not have dark foot spots. This is considered the
plesiomorphic state.

23. *Body shape.— Asteronotus has a broad body shape, the plesiomorphic state. This shape ts
shared by half the Halgerda species. The other half have a more elongate body shape.

24. *Secondary orange lines.—Ten Halgerda species have secondary orange lines on the notum
between the ridges. Asteronotus does not have these secondary lines, and this is considered the
plesiomorphic state.

25. *Orange dots.—Orange dots on the notum is the apomorphic character state of five Halgerda
species: H. aurantiomaculata, H. carlsoni, H. punctata, H. stricklandi, and H. bacalusia. The
outgroup taxon and the remaining Ha/gerda do not have orange dots on the notum.

26. Radular sac—The radular sac of most Halgerda is elongate. Only H. dalanghita has a short
radular sac, which is the plesiomorphic character state of the outgroup taxon Asteronotus. This
character state could not be determined for five taxa.

27. *Buccal mass——Most Halgerda species have an unpigmented buccal mass as does Asterono-
tus. However, H. albocristata, H. formosa, H. wasinensis, and H. willevi all have dark pigmentation
on the buccal mass. An unpigmented buccal mass is the plesiomorphic state. This character state could
not be determined for H. graphica, H. xishaensis or H. dichromis.

442 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 11

28. Radula shape——The shape of the radular ribbon is nearly square in Asteronotus and H.
dalanghita. This radula shape is the plesiomorphic state. All other Ha/lgerda species have a rectangular
radular ribbon.

29. Middle lateral teeth—AlII Halgerda species except H. dalanghita share the plesiomorphic
character state of hamate middle lateral teeth. Halgerda dalanghita has elongate, middle lateral teeth.

30. Outer lateral teeth shape—Pointed, undivided outer lateral teeth is the plesiomorphic
character state which is shared by Asteronotus, H. graphica, H. iota, H. willeyi, and H. dalanghita.
The remaining Halgerda species have fimbriate outer lateral teeth.

31. Outer lateral teeth size-——Only H. dalanghita has outer lateral teeth that are the same size as
the middle lateral teeth. 4steronotus and all other Halgerda species have smaller outer lateral teeth
than the middle teeth. The presence of smaller outer lateral teeth is the plesiomorphic state.

32. Fimbriate teeth Of the species which have fimbriate outer teeth, only H. alhocristata, H.
brunneomaculata, and H. elegans have elongate fimbriate teeth. All other species have short fimbriate
teeth. The presence of short, fimbriate, outer lateral teeth is considered the plesiomorphic character
state. This character state is not relevant for five species, including the outgroup (see character 30
above).

33. Ampulla length.—A short ampulla is considered plesiomorphic. This character state is shared
by both the outgroup taxon and five Halgerda species. All other Halgerda have a long ampulla. Data
were unavailable for four species: H. graphica, H. iota, H. punctata, and H. xishaensis.

34. Vagina length—Six Halgerda species share the plesiomorphic character state of a short
vagina with Asteronotus. All other Halgerda have an elongate vagina. Data were unavailable for the
same four species as 1n character 33.

35. Vagina shape——The outgroup taxon and nine Halgerda species have a narrow vagina. The
other Halgerda, for which data were available, have a wide vagina. A narrow vagina Is the
plesiomorphic character state.

36. Vagina duct—The vaginal duct enters the common atrium proximally in the outgroup taxon
and in all Halgerda except H. albocristata, H. aurantiomaculata, H. elegans, and H. guahan in which
they are separate to the base. Data were unavailable for the same four species noted above. A common
atrium is the plesiomorphic state.

37. Vaginal base—A narrow vaginal base is considered the pleisiomorphic character state. All
Halgerda share this state with the outgroup taxon except H. formosa, H. tessellata, H. wasinensis, H.
willevi, and H. diaphana. No data are available for the four Halgerda species mentioned previously.

38. Vaginal sphincter—A sphincter at the top of the vagina is present in five Ha/gerda species:
H. formosa, H. tesselata, H. wasinensis, H. willeyi, and H. dichromis. No sphincter is present in the
outgroup taxon, and this is considered the pleisiomorphic state. No data are available for the same
four Halgerda species as above.

39. (Prostate— All Halgerda and Asteronotus have a two-part prostate. Data were unavailable
for the same four species as noted above).

40. (Differentiation of prostate —All taxa studied except H. dalanghita have a well-differentiated
prostate. A well-differentiated prostate is the apomorphic state. Data were missing for the same four
species noted above).

4]. Ejaculatory duct—A short ejaculatory duct is the plesiomorphic character state, and the
outgroup taxon shares this state with eight Halgerda species. The ejaculatory duct is elongate in all
other Halgerda for which data were available.

42. Receptaculum seminis.—The receptaculum seminis is about the same size as the bursa
copulatrix in the outgroup taxon, in H. brunneomaculata and H. carlsoni. \n all other Halgerda for
which data were available, the receptaculum is smaller than the bursa.

FAHEY AND GOSLINER: PRELIMINARY PHYLOGENY OF HALGERDA 443

43. Tubercular vaginal glands —Tubercular vaginal glands are present in eight Halgerda species.
This is considered apomorphic. Lack of tubercular vaginal glands is the character state shared by the
outgroup taxon and the remaining Ha/gerda species for which data were available.

44. Penis Four Halgerda species have a tubular penis. All other Ha/gerda for which data were
available have a bulbous penis like Asteronotus. A bulbous penis is the plesiomorphic state.

45. Uterine duct insertion —The uterine duct inserts at the base of the bursa in the outgroup taxon,
and in half the Ha/gerda species. In the other Ha/gerda species, the duct inserts at a point more distal
to the bursa, closer to the receptaculum seminis. This is considered apomorphic. No data were
available for the same four Halgerda species.

46. (Prostate —The prostate is smaller than the female glana mass in the outgroup taxon and in
half the Halgerda species. In the other half, the prostate is the same size as the female gland mass.
The smaller prostate is considered plesiomorphic. No data were available for the same four species
as above).

47. *Atrium pigment—Dark pigmentation is present on the genital atrium of six Halgerda
species. All other species, including the outgroup taxon have no pigmentation on the genital atrium.
This 1s the plesiomorphic state.

48. Mantle margin——A colored margin around the perimeter of the mantle is a character state
shared by six Halgerda species. The outgroup taxon does not have a colored mantle margin. Lack of
a colored mantle margin is considered pleisiomorphic.

Methods

In order to develop phylogenetic hypotheses regarding Halgerda, the above described characters
were placed into a data matrix (Table 2) from MacClade version 3.04 (Maddison and Maddison 1998).
All characters used have equal weight and are unordered. Six characters were deleted from the first
analysis due to being parsimoniously uninformative. The characters deleted are indicated in paren-
theses in the character descriptions (previous section). The data were analyzed by Phylogenetic
Analysis Using Parsimony (PAUP) version 4.0b2 by David Swofford (1999). A heuristic search was
performed with the optimality criterion of maximum parsimony. The stepwise addition option of
Random Trees was used, with 100 repetitions, starting from random start trees. Four most parsimo-
nious trees were produced. The trees required 128 steps and had a consistency index of 0.375. The
strict consensus tree is shown in Figure 8. Figure 9 shows the majority rule tree with the character
numbers and character reversals. The underlined numbers indicate reversals.

An evaluation was performed of one million trees sampled randomly from the set of all possible
trees. The mean of that evaluation 1s 205.2, the standard deviation is 7.72, the gl statistic is —0.34 and
the g2 statistic is 0.08.

A decay analysis was performed using a heuristic search by PAUP for all trees <129 steps. The
stepwise addition option of Random Trees was used, with 100 repetitions, starting from random start
trees. A 50% majority-rule consensus of 390 trees was computed. The Consistency Index (CI) is 0.372
for this tree.

A second analysis was performed deleting color-based characters to determine the impact of these
characters on the tree topology. An asterisk preceding the character description (previous section)
indicates the characters deleted for this analysis. The data were analyzed by the same method using
PAUP as described for the first analysis. Figure 10 shows the strict consensus of 72 most parsimonious
trees obtained by deletion of the color characters. The tree required 77 steps and had a CI of 0.42.

Discussion

Character analysis indicated that nearly half of the characters were subject to at least one instance
of homoplasy, either parallelism or reversal. Figure 9 shows those characters by number. Some of
these characters are: small marginal tubercles; elongate, fimbriate outer teeth; a long ampulla; a wide

444 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES

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446 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 11

Strict
albocristata

elegans
brunneomaculata
guahan
aurantiomaculata
carlsoni
malesso
terramtuensis
H. sp. 2
stricklandi
bacalusia
diaphana
formosa
tessellata
wasinensis
graphica

fota

willeyi
punctata
dichromis
toliara
xishaensis
dalanghita

Asteronotus

FiGure 10. Strict consensus of 72 most parsimonious trees, derived from color characters deleted from data matrix
(Table 2).

vagina; a wide vaginal base; and the uterine duct insertion at a point more distal to the bursa copulatrix.
The other characters all relate to color, and it was for this reason that the second analysis, which
deleted these characters, was performed. Figure 10 shows that the same tree topology 1s maintained
after deletion of the color characters, although there is less resolution at the terminal branches.

Several apomorphies exhibit no instance of parallelism or reversal. They include: an elongate
vagina, a vaginal sphincter, a tubercular vaginal gland, a bipinnate gill, flat gill branchia, an elongate
ejaculatory duct, a tubular penis, and a receptaculum seminis that is smaller than the bursa copulatrix.

Further character analysis and testing of the data are needed to strengthen the phylogenetic
hypothesis of the genus Halgerda.

FAHEY AND GOSLINER: PRELIMINARY PHYLOGENY OF HALGERDA 447

Cladogenesis

The pattern of species cladogenesis of Halgerda is similar to that of other opisthobranch lineages,
such as Thuridilla (Gosliner 1995). In short, there 1s little adaptive radiation in the lower branches of
the tree and the most derived clades tend to undergo more speciation than lower clades.

Previous studies (Gosliner 1995) have shown that the opisthobranch clades which undergo more
adaptive radiation are strictly tropical. Halgerda fits this model and they are found only in tropical
waters.

ACKNOWLEDGMENTS

The authors are grateful to Mark Strickland who collected and photographed the specimens of
Halgerda stricklandi and H. bacalusia. Robert Bolland provided the photos and specimens of H.
diaphana. Dong Lin of the California Academy of Sciences provided the color plates and developed
the SEM micrographs. Our thanks to Clay Carlson for constructive comments and to Michael Miller
for the phographs of H. bacalusia. Angel Valdés and an anonymous reviewer provided excellent
suggestions for the improvement of the manuscript.

LITERATURE CITED

ALLAN, J. K. 1932. Australian nudibranchs. Australian Zoology 7:87—105.

BERGH, R. 1880. Beitrage zur Kenntniss der japanischen nudibranchien. I. Verhandlungen der koniglich-kais-
erlichen Zoologisch-botanischen Gesellschaft in Wien 30:155—200.

BERTSCH, H. AND S. JOHNSON. 1982. Three new species of dorid nudibranchs (Gastropoda: Opisthobranchia)
from the Hawaiian Islands. The Veliger 24(3):214-218.

CARLSON, C. H. AND P. J. Horr. 1993. Three new Halgerda species. (Doridoidea: Nudibranchia: Opisthobran-
chia) from Guam. The Veliger 36(1):16—26.

EHRENBERG, C. G. 1831. Symbolae physicae seu icones et descritiones animalium evertebratorum sepositis
insectis quae ex itinere per Africam Borealem et Asiam Occidentalem — novae aut illustratae redierunt.
Decas | Mollusca.

ELIOT, C. 1904. On some nudibranchs from East Africa and Zanzibar. Part III. Proceedings of the Zoological
Society of London. 2:371—374.

FAHEY, S. J. AND T. M. GOSLINER. 1999. Description of three new species of Halgerda from the western Indian
Ocean with a redescription of Halgerda formosa Bergh, 1880. Proceedings of the California Academy of
Sciences 51(8):365—383.

FARRAN, G. P. 1905. Report to the government of Ceylon on the pearl oyster fisheries of the gulf of Manaar
Supplementary Reports, Ceylon Pearl Oyster Fisheries. XX1:339-340.

GOSLINER, T. M. 1995. The genus Thuridilla (Opisthobranchia: Elysiidae) from the tropical Indo-Pacific, with
a revision of the phylogeny and systematics of the Elysiidae. Proceedings of the California Academy of
Sciences 49(1):1—54.

GOSLINER, T. M. AND S. J. FAHEY. 1998. Description of a new species of Halgerda, with a redescription of
Halgerda elegans Bergh 1905. Proceedings of the California Academy of Sciences 50(15):347-359.
MADDISON, W. P. AND D. R. MADDISON. 1998. MacClade 3.04. Distributed by the authors. Cambridge,

Massachusetts.

ODHNER, N. H. J. 1926. Die Opisthobranchien. Jn Further Zoological Results of the Swedish Antarctic Expedi-
tion 1901-1903 under direction of Dr. Otto Nordenskjold 2(1):1—100, pls. 1-3.

RUDMAN, W.B. 1978. The dorid opisthobranch genera Halgerda Bergh and Sclerodoris Eliot from the
Indo-West Pacific. Zoological Journal of the Linnean Society, London 68:59-87.

SWOFFORD, D. L. 1999. Phylogenetic analysis using parsimony (PAUP), version 4.0b2. Illinois Natural History
Survey, Champaign.

VALDES, A. AND T. M. GOSLINER. Submitted. Systematics and phylogeny of the caryophyllidia-bearing dorids
(Mollusca, Nudibranchia) with a description of a new genus and four new species from Indo-Pacific deep
waters. Zoological Journal of the Linnean Society, London.

448 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 11

WILLAN, R. C. ANDG. A. BRODIE. 1989. The nudibranch Halgerda aurantiomaculata (Allan, 1932) (Doridoidea:
Dorididae) in Fijian Waters. The Veliger 32(1):69—80.

Yonow, N. 1993. Opisthobranchs from the Maldive Islands, including descriptions of seven new species
(Mollusca: Gastropoda). Revue Frangaise d’ Aquariologie 20(4):97—130.

© CALIFORNIA ACADEMY OF SCIENCES, 1999
Golden Gate Park
San Francisco, California 94118

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES

Volume 51, No. 12, pp. 449-460, 5 figs., 3 tables. | November 23, 1999

| NOV 29 1999

Four New Species of the Genus Platymantis (Amphibia: Ranidae)
from Luzon Island, Philippines

by
Walter C. Brown
Department of Herpetology, California Academy of Sciences
San Francisco, California 94118

Angel C. Alcala
SUAKCREM, Silliman University,
Dumaguete City, 6200 Philippines

and

Arvin C. Diesmos
College of Arts and Sciences, University of the Philippines,
Los Banos, Laguna, Philippines

Four new species of Platymantis (P. cagayanensis, P. indeprensus, P. pseudodorsalis, and P.
taylori) are described. Two are from Mt. Banahao, one from the Central Cordillera, and
one from the Sierra Madre, Luzon Island, Philippines. These species are in the dorsalis
Group as diagnosed at present. The distinguishing characters and distribution of the
Philippine species of the dorsalis Group are discussed.

Since 1994, our field work on Mt. Maquiling and Mt. Banahao (two mountain peaks in southern
Luzon), the Central Cordillera, and Sierra Madre (northern Luzon, Philippines), has resulted in the
recognition of 11 species of Platymantis, dorsalis Group as currently defined (Brown et al. 1997a).
Seven of these have been described: P. corrugatus, P. dorsalis, P. spelaeus, P. levigatus, P. mimulus,
P. naomii, and P. pygmaeus.

These species are clearly distinguished by their advertisement calls, although they share a number
of morphological features. These similarities, particularly of the digits, are the main reason for the
failure of earlier herpetologists to recognize them as separate species.

MATERIALS AND METHODS

Materials examined include: (1) holotypes of Platymantis: P. corrugatus, P. dorsalis, P. mimulus,
P. levigatus, P. spelaeus, P. naomii, and P. pygmaeus; (2) all specimens from recent collections on
Mt. Banahao, Mt. Maquiling, Central Cordillera, and Sierra Madre that are referable to the dorsalis
Group of species. These materials are in the custody of the following institutions: Museum National
d’Histoire Naturelle (MNHN), the California Academy of Sciences (CAS or CAS-SU), Philippine
National Museum (NMPH), Silliman University Biology Museum (SUBM).

Morphometric characters: snout-vent length (SVL), head length (HL), head breadth (HW), snout
length (SnL), diameter of eye (ED), diameter of tympanum (TD), tibia length (TiL), and third finger
length from proximal edge of basal tubercle (3FL) were measured to the nearest 0.1 mm, using a

449

450 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 12

Helios dial caliper. Other morphological characters include webbing of toes, tubercles of hands and
feet, skin ornamentation, head shape, vomerine teeth, and color pattern.

Wide band spectrograms of the advertisement calls were prepared using a Kay Electrics
Sonagraph (Model #550) and SIGNAL Sound Analysis System software. No temperature data were
recorded.

SYSTEMATIC SECTION

Platymantis cagayanensis new species
Figs |

Holotype. — NMPH 6691, an adult female collected in disturbed Dipterocarp forest in Taggat
Forest Reserve in Santa Praxedes Town, Central Cordillena, Cagayan Province, Luzon, at 50-100 m
elevation, by A. C. Alcala and party, September 4, 1998.

Paratypes. — CAS 207447-207451, NMPH 6692-6693, and SUBM 2321-2322, collected at
the same locality as the holotype on September 4, 1998.

Description of holotype. — An adult female (measurements in mm): SVL 34.7, HL 13.7, HW
13.8, SnL 6.2, ED 4.9, TD 2.1, TiL 19.5, 3FL 5.2; head relatively smooth; dorsum shagreened with
many small tubercles and low, short, irregular ridges; venter and thighs smooth. Color of dorsum dark
brown, with a dark X-mark at shoulder region; a dark interorbital bar; dark cross bars on fore and
hind limbs. Color of venter cream; dark mottling on throat and breast.

DIAGNOSIS. — Advertisement call distinct (Fig. SA); large size, based on SVL of small sample
(Table 1); TD/SVL 6-9% (Table 2); a few tubercles and numerous short ridges on dorsum; dorsal
ground color reddish brown to brown with darker patches associated with ridges.

DESCRIPTION. — SVL 26.4-30.8 mm for seven males and 34.7—37.4 mm for two females; HL
39-43% of SVL and HW 40-46% of SVL; snout rounded, upper jaw moderately protruding, SnL
17-19% of SVL; eye moderate, ED 14-16% of SVL; tympanum distinct, TD 6—-9% of SVL; canthus
rounded; lores slightly oblique and concave; vomerine teeth prominent, patches widely separated;
fingers without webs; disks small and grooved; first, second, and fourth finger about equal in length;
3FL 15-16% of SVL; subarticular tubercles strongly protruding, blunt to pointed; one row of
supernumerary tubercles; palmar area otherwise smooth; hind limbs long, TiL 55-62% of SVL; toes
webbed at base, reaching distal edge of basal tubercle or beyond on fifth toe; disks of toes about equal
to or slightly larger than those of fingers; inner metatarsal tubercle elongate; outer small and rounded;
plantar area smooth; dorsum with a few tubercles and numerous short ridges; venter relatively smooth
or with granules under head and throat; posterior thighs granular.

COLOR. — In life, ground color of dorsal and upper lateral surfaces reddish brown (Fig. |). There
are a few small, dark spots on dorsum, a dark bar behind the eye, and weak, dark bands on the hind
limbs.

In preservative, dorsal ground color brown with small darker areas associated with pattern of
ridges to predominately darker, almost blackish; upper lateral surfaces darker; a dark interorbital bar;
venter creamy with brownish mottling anteriorly; limbs with dark, transverse bars.

REPRODUCTION. — Non-pigmented ovarian eggs, measuring about 1.75 mm in diameter, are
present in one of the females. No other information is available.

ADVERTISEMENT CALL. — The advertisement call (Fig. 5A) is distinct. It sounds like “kree-eek-
kree-eek.” The first, second, fourth and sixth parts (pulses) of each note are between about 1500 and
2000 Hz. The third and fifth parts are between 2700 and 3300 or 3400 Hz. The duration of each note
ranges from about 0.25 to 0.27 seconds. The interval between notes ranges from about 0.52 to 0.55
seconds.

BROWN, ALCALA, AND DIESMOS: FOUR PHILIPPINE PLATYMANTIS 45]

—

a

FiGure |. Platymantis cagavanensis (CAS 207448) showing characteristic color and ornamentation.

ETYMOLOGY. — The name is derived from that of the province, Cagayan, in the Central
Cordillera, Luzon where the population was discovered.

COMPARISONS. — This species, in terms of size (SVL 26.0-35.0 mm), is one of the six Philippine
species of the dorsalis Group that are termed large (Table 1). In general appearance this species is
most similar to Platymantis dorsalis. However, the advertisement call is very different (Fig. 5A). For
the call of P. dorsalis, see Brown et al. 1997b, fig. 1B.

HABITAT. — Forest floor in disturbed primary forest is the only known habitat for this species.

Platymantis taylori new species
Figs 2

Holotype. — NMPH 6684, an adult female collected by Arvin C. Diesmos in disturbed lowland
rain forest in eastern Sierra Madre Mountains in Sitio Natapdukan, Barangay Didian, municipality of
Palanan, Province of Isabela, Luzon Island on March 21, 1997.

Paratypes. — NMPH 6512-6513, 6524, 6526-6529, SUBM 2326, and CAS 207440207442
from the same site as the holotype on March 19-23, 1997; and NMPH 6671, 6674-6675, 6687-6688,
SUBM 2327, and CAS 207443-207446 Sitio Blos, Barangay Reina Mercedes municipality of
Maconacon, Isabela Province about 50 km north of type locality.

Description of holotype. — A mature female (measurements in mm): SVL 35.5, HL 14.7, HW
13.9, SnL 6.5, ED 4.9, TD 3.3, TiL 22.0, 3FL 5.9; dorsum with several low, short ridges anteriorly;
dorsum with dark interorbital bar and a few dark spots on body; limbs with dark crossbars; venter
creamy with some dark spots anteriorly.

DIAGNOSIS. — Advertisement call distinct (Fig. 5B); large size, based on SVL of males (Table
1); TD/SVL 7-9% (Table 2); a few low ridges and tubercles on back; dorsal ground color light brown
to grayish brown with scattered dark spots.

452 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 12

TABLE 1. Size categories (SVL in mm) for Philippine species of the dorsalis Group (N =
number in sample).

SVL
Species N Range Mean
Very large
P. spelaeus
Females 11 52.8-60.5 =
Males 6 41.5-46.9 =
Large
P. cagayanensis
Females 2 34.7-37.4 =
Males if 26.4-30.8 28.9
P. corrugatus
Females 2 35.2-50.8 45.7
Males 14 28.2-33.0 30.4
P. dorsalis
Females 10 30.8-42.9 34.5
Males 16 26.7-35.0 29.4
P. levigatus
Females 2 34.3-38.5 —
Males 3 29.5-30.3 -
P. taylori
Females 10 30.7-39.8 36.8
Males 10 26.6-33.0 29.6
P. pseudodorsalis
Females - -- _
Males 10 22.4-31.5 28.4
Intermediate
P. indeprensus
Females - - _
Males 12 23.6-27.2 DSi
Small
P. mimulus
Females 18 21.0-28.1 wal)
Males 3) 18.4-25.8 21.9
P. naomii
Females 14 21.6-28.0 24.3
Males 18 20.3-23.9 21.6
P. pygmaeus
Females 15.0 -
Males 10 14.1—15.7 ISI

DESCRIPTION. — SVL 27.6—-28.3 mm for three mature males and 37.3 mm for one mature
female; HL 40-43% of SVL; HW 39-43% of SVL; SnL 18-19% of SVL; ED 14-15% of SVL; TD
7-9% of SVL; tympanum exposed; canthus rostralis rounded; lores deeply oblique, concave;
vomerine teeth strongly protruding, patches widely separated, fingers without webs; fingers with
grooved disks; first, second and fourth fingers nearly equal in length; third finger longest, 3FL 14-17%
of SVL; subarticular tubercles large, strongly protruding but not pointed; one row of large supernu-
merary tubercles; metacarpal tubercles oval, inner and middle large and outer small; hind limbs long,
TiL 57-63% of SVL; toes webbed at base: first, second and third to base of subarticular tubercle, to

BROWN, ALCALA, AND DIESMOS: FOUR PHILIPPINE PLATYMANTIS 453

_ TABLE 2. Proportional measurements as a percent of SVL for the species described.

P. cagayanensis P. indeprensis P. pseudodorsalis P. taylori

HL/SVL 39-43 37-45 39-43 40-43
HW/SVL 40-46 40-48 39-44 39-43
SnL/SVL Ia) 14-19 16-18 18-19
ED/SVL 14-16 INS 14-16 14-15
TD/SVL 6-9 6-9 8-1] 7-9

3FL/SVL 15-16 15-18 =I 7 14-17
TiL/SVL 55-62 35=05 54-60 57-63

much below base on fourth, and to distal edge of basal tubercle on fifth; disks of toes with grooves,
about equal to or slightly larger than those of fingers; subarticular tubercles, one on first and second
toes, two on third and fifth, and three on fourth; inner metatarsal tubercle elongate and outer round,
both strongly protruding; plantar area smooth; tubercle at heel absent; dorsum, head, snout, eyelids
and limbs generally smooth, with few low tubercles and short, low ridges on back. Throat, chin and
belly smooth; posterior thighs with small granules.

COLOR. — In life, ground color of dorsal and upper lateral surfaces light brown to grayish brown
(Fig. 2). There are a few dark spots on the dorsum and dark bars on the lips. The venter has dark
blemishes under the head and throat.

In preservative, dorsal background color light brown. A middorsal, light irregular band extends
from snout to anal region. In this band and in areas bordering it are dark spots. A vague, dark W-shaped
mark on the back at the shoulder level and a dark, interorbital band are present. Lips have dark bars
and limbs dark crossbands. Chin, throat, breast and sides of body have dark spots. Posterior belly and
ventral parts of thighs are whitish in color.

FiGure 2. Platymantis taylori (NMPH 6512) showing characteristic color and ornamentation.

454 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 12

REPRODUCTION. — Reproduction has not been observed. However, the large, unpigmented eggs
in the holotype are indicative of direct development, characteristic of all known species of the genus.

ADVERTISEMENT CALL.— The call (Fig. 5B) sounds like “kraak-kraak-kraak.” Each note
consists of several pulses between 1200-3900 Hz, with the terminal group very closely spaced. The
duration of each note is about 0.3—0.4 seconds. The interval between notes 1s short, 0.13—0.14 seconds.

ETYMOLOGY. — This species is named for Edward H. Taylor who devoted the period 1912 to
1923 to the study of the Philippine herpetofauna.

COMPARISONS. — This species, like P. cagayanensis, is one of the large species of the Philippine
dorsalis Group. The advertisement call (Fig. 5B) 1s very distinct, however.

HABITAT. — The specimens of this species were taken from the floor of disturbed primary forest.

Platymantis pseudodorsalis new species
Fig. 3

Holotype. — NMPH 6689, an adult male, collected in upper Dipterocarp forest on Mt. Banahao,
Luzon Island, Philippines by Roy Quiber and Ronnie Rizo on May 29, 1998.

Paratypes. — CAS 201536, 207455—207459, NMPH 6690, and SUBM 2323, collected between
upper Dipterocarp and lower montane forest (600—1,200 m) on Mt Banahao, Luzon Island, Philip-
pines.

Description of holotype. — A mature male (measurements in mm): SVL 30.7, HL 12.8, HW
13.5, SnL 5.3, ED 4.4, TD 3.0, TiL 18.0, 3FL 4.7; snout, head and dorsum smooth; upper eyelid with
few low tubercles; a light vertebral line; upper surface of hind limbs with many tubercles; venter
smooth; four small dark spots in inter-orbital space; skin on back uniformly brownish tan; dorso-lat-
eral area with interrupted, dark stripe; sides of body dark brown; upper surfaces of hind limbs with
dark mottling.

DIAGNOsIS. — Advertisement call distinct (Fig. 5C); large size, based on SVL of males (Table
1); TD/SVL 8—-11% (Table 2); dorsum nearly smooth or with a few, low ridges and tubercles; dorsal
ground color light to medium brown, lighter than upper lateral surface.

DESCRIPTION. — SVL 22.4-31.5 mm for 10 males; HW 39-44% of SVL; snout rounded to
somewhat round pointed; upper jaw moderately protruding; SnL is 16-18% of SVL; eye large, ED
14-16% of SVL; tympanum distinct, TD 8—11% of SVL; canthus rounded; lores slightly oblique and
concave; fingers without webs, with small disks and narrow dermal flanges, first finger about as long
or slightly longer than second, both somewhat shorter than fourth, third finger longest, 3FL 15-17%
of SVL; subarticular tubercles large, strongly protruding; inner and middle metacarpal tubercles large,
oval; outer small and elongate: one row of supernumerary tubercles; hind limbs long, TiL 54-60%
of SVL; toes with minute webs: first and second toes about to base of tubercle, third about to middle
of basal tubercle, fourth much below basal tubercle, and fifth to mid-point or distal edge of basal
tubercle; disks of toes slightly larger than those of fingers; subarticular tubercles oval, strongly
protruding, inner metatarsal tubercles high, elongate, outer small, round, strongly protruding; plantar
area smooth; snout and head generally smooth; upper eyelids with few to many low tubercles; back
smooth or with a few low, short ridges; upper surfaces of limbs usually with many low tubercles;
throat and belly smooth.

COLOR. — In life, ground color of dorsum light brown to medium brown, somewhat darker on
the upper lateral surfaces (Fig. 3). Some specimens have a pale vertebral stripe. The hind limbs have
dark transverse bands. There is a pale line on the anterior forearm. The venter has dark blemishes
anteriorly, sometimes forming a network.

In preservative, general color above light brown to brownish tan; head and back lighter than sides
of body; most specimens with a light vertebral line; two or more irregular dark spots in inter-orbital
space, dark spots on head between eyes and nostrils in most specimens; throat, breast and anterior

BROWN, ALCALA, AND DIESMOS: FOUR PHILIPPINE PLATYMANTIS 455

FIGURE 3. Platvmantis pseudodorsalis (NMPH 6689) showing characteristic color and ornamentation.

belly with dark concentrations of pigments, sometimes forming a network; posterior belly generally
immaculate; a light line in all specimens either continuous or broken on the antero-ventral portion of
forearm.

REPRODUCTION. — Nothing is known about this species.

ADVERTISEMENT CALL. — The call of this species sounds like “tsolek-tsolek” (Fig. 5C). Each
note is divided into three parts (pulses). The first and third parts are between 2500 and 3100 to 3200
Hz. The middle part is between 1400 and 2100 Hz. The duration of each note is very uniform between
(0.12 and 0.13 seconds. The intervals between notes range from about 0.8 to 1.2 seconds.

ETYMOLOGY. — Species name is based on the similarity of specimens with P. dorsalis.

ECOLOGICAL NOTE. — This species inhabits the forest floor at elevations of 700 to 1,200 m from
the upper Dipterocarp forest to lower montane forest. It has been observed from January to September,
but calls are generally heard from May to July.

COMPARISONS. — This species, like P. indeprensus, has been confused with P. dorsalis. It
differs from both P. dorsalis and P. indeprensus in the very different advertisement call (Fig. SC). In
size it is most similar to P. indeprensus (Table 1).

RANGE. — Known only from Mt. Banahao, Quezon Province on Luzon Island.

Platymantis indeprensus new species
Fig. 4

Holotype. — CAS 201196, an adult male collected in submontane forest on Mt. Banahao, Luzon
Island at about 1,080 m by A. C. Diesmos, April 20, 1996.

Paratypes. — CAS 201013-201014, 201178-201183, 201185, 207452—207454, SUBN 2325,
NMPH 6685-6686 Mt. Banahao, Luzon Island, Philippines (850—1,300 m in upper Dipterocarp and
montane forest).

456 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 12

Ficure 4. Platymantis indeprensus (CAS 201183) showing characteristic color and ornamentation.

Description of holotype. — An adult male (measurements in mm): SVL 27.0, HL 10.1, HW
11.4, TiL 14.4, SnL 4.1, ED 3.8, TD 1.9, 3FL 4.1; snout, upper eyelids, upper lateral surfaces and
limbs with tubercles; dorsum with tubercles and a few short ridges anteriorly; venter relatively
smooth.

DIAGNOSIS. — Advertisement call distinct (Fig. 5D); intermediate size, based on SVL of males
(Table 1); TD/SVL 6-9% (Table 2); dorsum with scattered tubercles and low ridges; dorsal ground
color mottled tannish and blackish brown.

DESCRIPTION. — SVL 21.5—27.3 mm for 14 males (females not identified); HW 40-48% and
HL 37-45% of SVL; snout rounded to round pointed; upper jaw moderately protruding; SnL 14-19%
of SVL, eye large, ED 12-16% of SVL; tympanum distinct TD 6—9% of SVL; canthus rounded; lores
only slightly oblique and concave; fingers without webs; disks small with narrow, dermal flanges;
first finger about as long as or slightly longer than second, both somewhat shorter than fourth; 3FL
15-18% of SVL; subarticular tubercles large, strongly protruding, rounded; inner and middle
metacarpal tubercles large, oval; outer moderate, elongate; one row of supernumerary tubercles; hind
limbs long; TiL 55-63% of SVL toes with minute webs: first, second, and third toes about to base of
tubercle on inside, fifth toe to midpoint or beyond basal tubercle; disks of toes slightly larger than
those of fingers; subarticular tubercles oval, strongly protruding; inner metatarsal tubercles high,
elongate, outer small, round, strongly protruding; plantar area smooth; snout, head, and upper eyelids
with numerous, flattish tubercles; dorsum posterior to eyes with scattered tubercles, sometimes
tubercles fused to form a jumbled pattern of short ridges; throat and belly smooth; proximal half of
posterior thighs granular.

COLOR. — In life, ground color of dorsal and upper lateral surfaces is mottled blackish brown
(Fig. 4). Some specimens have a pale vertebral stripe. The limbs have dark transverse bands. There
are dark blemishes under the head and throat and sometimes a few on the belly.

In preservative, part of interorbital space lighter than the rest of head; dorsum mottled blackish
brown and tannish brown or occasionally with a narrow, pale vertebral stripe; lateral surfaces with

BROWN, ALCALA, AND DIESMOS: FOUR PHILIPPINE PLATYMANTIS 457

blotches of grayish brown to grayish tan, mottled brown under head and throat, more cream with or
without scattered brown spots on belly; distinct crossbars on limbs.

REPRODUCTION. — Nothing is known for this species. Females have thus far eluded capture or
identification.

ADVERTISEMENT CALL. — The calls sound like “eyak-eyak.” The main part of each note is
between 2500 and 3500 Hz, closing with a series of pulses ranging from 2000 to 4000 Hz. The
duration of each note is between 0.2] and 0.31 seconds. Each series is usually two or three notes at
intervals from about 0.18 to 0.31 seconds (Fig. 5D). The males tend to call while perching on more
exposed spots on the forest floor.

ETYMOLOGY. — The species name is Latin for “unobserved” or “undiscovered” and refers to the
fact that this species was confused with P. dorsalis.

ECOLOGICAL NOTE. — The species inhabits the forest floor from upper Dipterocarp to lower
montane forest on Mt. Banahao, Quezon Province, Luzon Island. Males have been observed during
the rainy months from April through August.

COMPARISONS. — This species from Mt. Banahao has been confused with P. dorsalis. The
distinct advertisement call (Fig. 5D) and intermediate size (Table 1) distinguish it from P. dorsalis.

RANGE. — Known only on Mt. Banahao and Mt. San Cristobal.

DISCUSSION

The dorsalis Group as currently diagnosed includes eleven species from the Philippines and
sixteen from Melanesia, New Guinea, and Palau Islands (Table 3). These species are characterized
by: (1) blunt digits or small, nearly undilated disks, (2) subtending parts of digits narrow, (3)
subarticular tubercles strongly protruding and usually pointed, (4) terminal phlanges pointed not
T-shaped. Pending completion of phylogenetic studies of platymantine frogs, the probable relation-
ships among the species of the dorsalis Group is nothing more than a guess.

The Philippine species (P. corrugatus, P. dorsalis, P. mimulus, P. pygmaeus, P. spelaeus, P.
levigatus, P. naomii, P. indeprensus, P. pseudodorsalis, and P. taylori) are distributed in the Greater
Mindanao, Greater Negros-Panay, Greater Luzon, and Mindoro Island groups. They are differentiated
by their advertisement calls and some morphological characters. They also are separated into

TABLE 3. Species currently referred to the dorsalis Group of Platymantis.

Species known to occur only on islands outside Species known to occur only in the Philippines

P. acrochordus (Solomon Is.) P. cagayanensis (Luzon)

P. aculeodactylus (Solomon Is.) P. corrugatus (most islands)
P. akarithymus (Bismarck Is.) P. dorsalis (most islands)

P. boulengeri (Bismarck Is.) P. indeprensus (Luzon)

P. gilliardi( Bismarck Is.) P. levigatus (Tablas and Sibuyan Is.)
P. macrops (Solomon Is.) P. mimulus (Luzon)

P. mimulus (Bismarck Is.) P. naomii (Luzon)

P. myersi (Solomon Is.) P. pseudodorsalis (Luzon)
P. papuensis (New Guinea and satellites) P. pygmaeus (Luzon)

P. parkeri (Solomon Is.) P. spaeleus (Negros)

P. pelewensis (Palau Is.) P. taylori (Luzon)

P. rhipiphaleus (Bismarck Is.)

P. schmidti (Bismarck Is.)

P. solomonis (Solomon Is.)

P. vitianus (Fiji Is.)

P. weberi (Solomon Is.)

458 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 12

Sy Ge BUM Ie eat eS ence eT Cs ee co ae a ce

by sstetatee: ®t bx

TIME (sec)

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459

BROWN, ALCALA, AND DIESMOS: FOUR PHILIPPINE PLATYMANTIS

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460 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 12

relatively distinct subgroups, based on SVL at maturity (Table 1). Very large includes only one
species, Platymantis spelacus. Large includes six species, P. cagavanensis, P. corrugatus, P. dorsalis,
P. levigatus, P. pseudodorsalis, and P. taylori. Small includes three species, P. mimulus, P. naomii,
and P. pygmaeus.

Platymantis spelaeus does not overlap any of the other species in SVL. There is extensive overlap
in SVL for the six species classified as large, and both range and means of SVL are given for both
males and females for those species represented by sufficiently large samples (Table |). Of the large
species, P. cagayanensis, P. dorsalis, P. pseudodorsalis, and P. taylori are most similar both in size
and appearance and were first recognized on the basis of distinct advertisement calls. Platymantis
corrugatus, however, is readily recognized because of the blunt digital tips and distinctive color
pattern; and P. /evigatus from its very smooth skin and, to some extent, color pattern. Platymantis
indeprensus, although long confused with P. dorsalis, is intermediate in SVL (Table |). The three
species (P. mimulus, P. naomii, and P. pygmaeus) categorized as small do not overlap with other
species in SVL.

Populations of P. dorsalis on Luzon and Negros have similar calls and are considered the same
species. However, recent observations on advertisement calls of some populations in Leyte, Samar,
Mindoro, and western Mindanao indicate that populations in these areas currently referred to P.
dorsalis may belong to separate species. We are conducting field work on these populations and
analyzing calls at the present time.

ACKNOWLEDGMENTS

We are indebted to Arne Jensen, Nordic Agency for Development and Ecology (NORDECO)
and W. Pollisco, Director of the Protected Areas Bureau of the Department of Environment and
Natural Resources, Philippines for making available specimens of P. tayvlori. We also wish to thank
Robert Drewes, Jens Vindum, and Andrea Jesse (CAS) for their assistance.

LITERATURE CITED

ALCALA, A.C., W.C. BROWN, AND A.C. DIESMOS. 1998. Fwo new species of the genus
Platymantis (Amphibia: Ranidae) from Luzon Island, Philippines. Proceedings of the California
Academy of Sciences 50(17):381-388.

BROWN, W.C., R. M. BROWN, AND A.C. ALCALA. 1997a. Species of the hazelae Group of
Platymantis (Amphibia: Ranidae) from the Philippines, with descriptions of two new species.
Proceedings of the California Academy of Sciences 49(11):405—421.

BROWN, W.C., A.C. ALCALA, AND A.C. DIESMOS. 1997b. A new species of the genus
Platvmantis (Amphibia: Ranidae) from Luzon Island, Philippines. Proceedings of the Biological
Society of Washington 110(1):18-23.

DiesMos. A.C. 1998. The amphibian faunas of Mount Banahao, Mount San Cristobal, and
Mount Maquiling, Luzon Island, Philippines. Thesis (M.Sc.) in Wildlife Studies, University of the
Philippines, Los Banos, Laguna. Unpublished manuscript.

© CALIFORNIA ACADEMY OF SCIENCES, 1999
Golden Gate Park
San Francisco, California 94118

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES

Volume 51, No. 13, pp. 461-472, 7 figs. | November 23, 1999

Two New Species of Chromodoris (Mollusca, Nudibranchia,
Chromodorididae) from Southern India, with a Redescription of
Chromodoris trimarginata (Winckworth, 1946)

by
Angel Valdés
Department of Invertebrate Zoology and Geology, California Academy of Sciences
Golden Gate Park, San Francisco, California 94118

Ernesto Mollo
Istituto per la Chimica di Molecole di Interesse Biologico, CNR.
Via Toiano 6, 80072 Arco Felice, Napoli, Italy

and

Jesus Ortea
Departamento de Biologia de Organismos y Sistemas, Laboratorio de Zoologia,
Universidad de Oviedo, c/ Catedratico Rodrigo Uria s/n, 33071 Oviedo, Spain

Chromodoris trimarginata (Winckworth, 1946) is redescribed on the basis of specimens
from southern India. The reproductive system of this species is illustrated for the first time.
New characters, such as the rhinophores and gill coloration, the width of the white marginal
band, and the radular morphology, are provided to distinguish this species from the similar
Chromodoris preciosa (Kelaart, 1858). In addition, two new species of Chromodoris from
southern India are described. Chromodoris mandapamensis sp. nov. is characterized by a
dorsum covered with brown spots, a broken line of bright orange around the mantle
margin, very large branchial leaves, and radular teeth with large denticles. Chromodoris
naiki sp. nov. is a pale gray species with bright orange and purple spots, and a number of
white dots on the dorsum. Internally it lacks rachidian radular teeth, has long cusps in the
mid lateral teeth, and lacks a vestibular gland. The external morphology and anatomy of
these two species is described, and compared to other similar Indo-Pacific species.

The Chromodorididae of India and Sri Lanka (Ceylon) have received little attention compared
to well studied areas in the Indo-Pacific. Few species were described in the classic papers by Alder
and Hancock (1864) and Kelaart (1958; 1859a; 1859b). In the first half of the 20th century, Farran
(1905), Eliot (1906a; 1906b; 1910), O’Donoghue (1932), Winckworth (1945) and White (1948)
described more new species or redescribed those introduced by Kelaart or other authors. More
recently, Narayanan (1969) and Rudman (1973) studied several specimens from India, providing
range extensions and the description of a new species.

The present paper studies three species of the genus Chromodoris Alder and Hancock, 1864,
collected by the second author from Mandapam (Tamil Nadu) and Muttom (Kerala), southern India.
The specimens studied are deposited at the Department of Invertebrate Zoology, California Academy
of Sciences, designated by the abbreviation CASIZ.

461

462 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 13

SPECIES DESCRIPTIONS

Chromodoris trimarginata (Winckworth, 1946)
Figs. LA, 2, 3

Glossodoris trimarginata Winckworth, 1946:156—157.
Chromodoris trimarginata (Winckworth): Rudman, 1985:267—268, fig. 16.

MATERIAL EXAMINED. — Mandapam, Tamil Nadu, India, May 1998, two specimens 30 mm long, one of
them dissected, collected by E. Mollo (CASIZ 115219); March 1999, one specimen 20 mm long, collected by
E. Mollo (CASIZ 115228).

EXTERNAL MORPHOLOGY. — The body is oval to elongate (Fig. 1A). The posterior end of the
foot is not covered by the notum. The living animal is pale gray, with a number of reddish spots
irregularly scattered over the central part of the dorsum. At the extreme edge of the mantle there is a
white, thin band, then a band of deep red and on the inside a slightly wider band of yellow. The mantle
glands are situated on the inside of the yellow band and form a wide, opaque white band around the
whole mantle margin. The rhinophore stalks are translucent white; the clubs are cream, and the apexes
opaque white. The perfoliate rhinophores are composed of 20 lamellae. The gill consists of 11
unipinnate branchial leaves. The leaves are translucent white with opaque white lamellae.

ANATOMY. — The buccal mass is divided evenly into an anterior glandular portion and a
posterior muscular one. At the posterior end of the mass there are a pair of short, elongate salivary
glands. The jaws are composed of a number of elongate, bifid rodlets (Fig. 2D) about 20 um in length.
The radular formula is 57 x 53.1.53 in one specimen (CASIZ 115219) examined. The rachidian teeth
are small, triangular plates. The innermost lateral teeth (Fig. 2A) have three to four denticles on the
inner side of the cusp and four to five denticles on the outer side. The remaining lateral teeth (Fig.
2B) are hook-shaped, lack denticles on the inner side of the cusp and have a series of four to six
denticles along the outer edge. The outer laterals (Fig. 2C) are elongate, having five to six denticles
situated at the tip of the teeth.

The reproductive system (Fig. 3) has a short, tubular ampulla that divides into the oviduct and
the prostate. The oviduct is short and enters the female glands near the center of the mass. The prostate
is long, tightly coiled with several loops. It narrows and then expands into a long, muscular deferent
duct. The deferent duct is also highly coiled, and opens into a common atrium with the vagina. The
penis is unarmed. Near the exit of the female glands there is a small, saccate vestibular gland. The
vagina is very long, slightly coiled. Near the end of the vagina the uterine duct emerges. It is very
long and convoluted and opens into the female glands. The curved, club-shaped seminal receptacle
and the elongate, thin-walled bursa copulatrix open proximally in the vaginal duct.

DISTRIBUTION. — Chromodoris trimarginata is only know from India; it was originally de-
scribed from Bombay (Winckworth 1946) and the present record from Mandapam constitutes the
second collection of this species.

REMARKS. — Chromodoris trimarginata was described by Winckworth (1946) in the binomen
Glossodoris trimarginata, based on 88 specimens collected from Bombay (India). The living animals
were described as being cream or very pale green with irregular reddish spots scattered over the central
area. At the extreme edge of the mantle there was a white, thin band, then a band of red and on the

—>

FiGuRE |. Living animals. A. Chromodoris trimarginata (CASIZ 115219); B. Chromodoris mandapamensis sp. nov. (CASIZ
115220); C. Chromodoris naiki sp. nov. (CASIZ 115221).

VALDES, MOLLO, AND ORTEA: NEW CHROMODORIDIDAE FROM SOUTHERN INDIA

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 13

FIGURE 2. Chromodoris trimarginata (CASIZ 115219), scanning electron micrographs. A. Inner lateral teeth, scale bar =
25 ym; B. Lateral teeth from the central portion of the half-row, scale bar = 30 jum; C. Outer lateral teeth, scale bar = 25 um;
D. Jaw rodlets, scale bar = 15 um.

VALDES, MOLLO, AND ORTEA: NEW CHROMODORIDIDAE FROM SOUTHERN INDIA 465

ud

FIGURE 3. Chromodoris trimarginata (CASIZ 115219), reproductive system, scale bar = 1 mm. A. General view; B. Detail
of several dissected organs. Abbreviations: am = ampulla, be = bursa copulatrix, dd = deferent duct, fg = female glands, pr =
prostate, sr = seminal receptacle, ud = uterine duct, v = vagina, vg = vestibular gland.

inside a slightly wider band of yellow. The rhinophores were described as very pale green with white
tips, and the eleven, simply pinnate branchial leaves as white, with the lamellae and rachis opaque
white. Rudman (1985) redescribed this species based on the preserved syntypes deposited at the
Museum of Natural History of London (BMNH 1960725). He transferred this species to the genus
Chromodoris and distinguished it from the similar Chromodoris preciosa (Kelaart, 1858) and
Chromodoris sinensis Rudman, 1985, due to differences in color and radular morphology.

The coloration of our specimen from Mandapam resembles the original description of C.
trimarginata by Winckworth (1946), and the radula is identical to the photographs published by
Rudman (1985) for this species.

The present study confirms the differences in color between C. trimarginata and C. preciosa.
The latter has reddish rhinophores and branchial leaves (Kelaart 1858; Rudman 1985), whereas in C.
trimarginata the rhinophores are cream and white and the branchial leaves are white. Also, the
marginal white band of C. preciosa is much wider than that of C. trimarginata. Another important
difference between these two species is in the radular morphology. The radula of C. preciosa, figured
by Rudman (1985, fig. 17), has much shorter innermost lateral teeth than C. t7imarginata. Also, in
C. preciosa the innermost teeth have two or three large, acute denticles on the inner side of the main
cusp, whereas in C. trimarginata there are three or four smaller, rounded denticles.

Chromodoris sinensis differs from C. trimarginata in color and radular morphology. The radula
of C. sinensis figured by Rudman (1985, fig. 18) has large rachidian teeth, and two or three acute
denticles on the inner side of the innermost lateral teeth. In C. trimarginata, the rachidian teeth are
much smaller and the innermost lateral teeth have three or four rounded denticles on the inner side.
According to Rudman (1985) the external differences between these two species are: (1) the

466 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 13

translucent white border is much narrower in C. sinensis than in C. trimarginata; (2) the branchial
leaves and rhinophores of C. sinensis are edged with red, whereas they are uniformly white in C.
trimarginata;, and (3) the dorsal spots of C. sinensis are regular minute orange-brown spots, instead
of the irregular reddish spots of C. trimarginata. The study of our living specimens of C. trimarginata
confirmed all these differences, except for the white band, that is also very narrow in C. trimarginata.
The reproductive system of C. trimarginata, here studied for the first time, has a vestibular gland that
is absent in C. sinensis (see Rudman 1985). In addition, the uterine duct and the seminal receptacle
of C. trimarginata connect to the vaginal duct near the bursa copulatrix, whereas in C. sinensis they
connect in the middle of the vagina (Rudman 1985, fig. 15C).

Chromodoris mandapamensis sp. nov.
Figs. 1B, 4, 5

MATERIAL EXAMINED. — HOLOTYPE: Mandapam, Tamil Nadu, India, March 1999, 50 mm long, collected
by E. Mollo (CASIZ 115229). PARATYPES: Mandapam, Tamil Nadu, India, May 1998, two specimens 30 mm
long, one of them dissected, collected by E. Mollo (CASIZ 115220).

Twenty six additional specimens collected in March 1999 from Mandapam (Tamil Nadu, India) and one
specimen collected in May 1998 from Muttom (Kerala, India), were used for chemical studies.

EXTERNAL MORPHOLOGY. — The body is elongate (Fig. 1B). The posterior end of the notum is
very elongate and covers the foot. The living animal is cream to pale gray. The entire dorsum is densely
covered with numerous, rounded, dark brown spots. These spots appear to be diffuse in the mantle
edge. A broken line of bright orange spots edges the mantle margin. There are several, ramified,
subepidermical mantle glands around the mantle margin (Fig. 5A). The rhinophore stalks are cream
with several dark brown spots. The clubs are brown, with white edged lamellae. The apexes are reddish
with white tips. The perfoliate rhinophores are composed of 17 lamellae. The gill consists of seven
large, bipinnate branchial leaves. The leaves are pale brown with numerous, small opaque white dots
scattered all over the surface, and large brown spots near the base of the leaves.

ANATOMY. —The buccal mass is divided evenly into an anterior glandular portion and a
posterior muscular one. At the posterior end of the mass there are a pair of long salivary glands. The
jaws are composed of a number of elongate, bifid rodlets (Fig. 4D) about 15 um in length. The radular
formula is 67 x 53.0.53 in one specimen (CASIZ 115220) examined. Rachidian teeth are absent. The
innermost lateral teeth (Fig. 4A) have two denticles on the inner side of the cusp and three denticles
on the outer side. The remaining lateral teeth (Fig. 4B) are hook-shaped, lack denticles on the inner
side of the cusp, and have a series of four to six denticles along the outer edge. The outer laterals (Fig.
4C) are elongate with four to seven denticles situated on the tips of the teeth.

The reproductive system (Fig. 5B, C) has a long, tubular ampulla that divides into the oviduct
and the prostate. The oviduct is short and enters the female glands near the center of the mass. The
prostate is long, tightly coiled with several loops. It narrows and then expands into the muscular
deferent duct, which is bulbous distally. The deferent duct is very short and wide, and opens into a
common atrium with the vagina. The penis is unarmed. The vagina is short and slightly coiled. The
uterine duct emerges near the end of the vagina. The uterine duct is long and opens into the female
glands. More proximally there are the curved, club-shaped seminal receptacle and the rounded,
thin-walled bursa copulatrix.

ETYMOLOGY. — The name is derived from Mandapam, the type locality of this species.

DISTRIBUTION. — So far, this species is only known from Mandapam (Tamil Nadu) and Muttom
(Kerala), southern India.

REMARKS. — Chromodoris mandapamensis is externally different from other species of Chro-
modoris previously described.

VALDES, MOLLO, AND ORTEA: NEW CHROMODORIDIDAE FROM SOUTHERN INDIA

FiGuRE 4. Chromodoris mandapamensis sp. nov. (CASIZ 115220), scanning electron micrographs. A. Inner lateral teeth,
scale bar = 25 um; B. Lateral teeth from the central portion of the half-row, scale bar = 30 jum; C. Outer lateral teeth, scale bar
= 30 um; D. Jaw rodlets, scale bar = 20 tum.

468 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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FiGuRE 5. Chromodoris mandapamensis sp. nov. (CASIZ 115220). A. Disposition of the mantle glands, scale bar = | mm;
B. General view of the reproductive system, scale bar = 1 mm; C. Detail of several dissected organs, scale bar = | mm.
Abbreviations: am = ampulla, be = bursa copulatrix, dd = deferent duct, fg = female glands, pr = prostate, sr = seminal recep-
tacle, ud = uterine duct, v = vagina.

Kelaart (1859b) described Doris humberti as a white species with purple and brown spots on the
dorsum; white mantle margin with a row of bright red spots, red rhinophores and gill with white spots,
nine small branchial leaves irregularly pinnated. This species is clearly distinguishable from C.
mandapamensis, which has seven very large, brown branchial leaves and dark brown rhinophores.
O’ Donoghue (1932) redescribed D. humberti based on newly collected material from southern India
and transferred it to the genus Glossodoris Ehrenberg, 1831. O’Donoghue’s paper includes a
description of the radula that is elongate, with a formula of 74 x 46.0.46, and has innermost lateral
teeth with a single, large denticle on the inner side, and two on the outer side. This is very different
from C. mandapamensis, which has a formula of 67 x 53.0.53 and the innermost lateral teeth have
two denticles on the inner side and three on the outer side.

Two other Indo-Pacific species of Chromodoris with a similar coloration to C. mandapamensis
are Chromodoris lineolata (van Hasselt, 1824) and Chromodoris striatella Bergh, 1876, redescribed
by Rudman (1982) and Rudman and Darvell (1990). These latter two species have a white background
color with numerous brown or black lines, and an orange mantle margin. They are clearly distinguish-
able from C. mandapamensis, which has dorsal spots instead of lines.

Chromodoris naiki sp. nov.
Figsy Gy Gs 7

MATERIAL EXAMINED. — HOLOTYPE: Mandapam, Tamil Nadu, India, May 1998, one specimen 13 mm
preserved length, dissected, collected by E. Mollo (CASIZ 115221).

EXTERNAL MORPHOLOGY. — The body is oval (Fig. 1C). The notum covers the posterior end of
the foot. The living animal is pale gray-brown, with a pale blue band at the edge of the notum. Along
the margin and in the central part of the dorsum, there are scattered large spots of dark purple.
Immediately inside the area of outer purple spots there is an area of reflective yellow or orange-yellow
mantle glands. There are also yellow or orange-yellow mantle glands (Fig. 6A) associated with the
dark purple spots in the center of the dorsum. Numerous, small opaque white spots are scattered all

VALDES, MOLLO, AND ORTEA: NEW CHROMODORIDIDAE FROM SOUTHERN INDIA 469

FIGURE 6. Chromodoris naiki sp. nov. (CASIZ 115221). A. Disposition of the mantle glands, scale bar = | mm; B. General
view of the reproductive system, scale bar = 1 mm. Abbreviations: am = ampulla, be = bursa copulatrix, dd = deferent duct,
fg = female glands, pr = prostate, sr = seminal receptacle, ud = uterine duct, v = vagina.

over the central part of the dorsum. The rhinophore clubs and gill are gray-brown. The bases of the
rhinophores are translucent gray. There are numerous small opaque white spots arranged in distinct
rows along the edges of the rhinophoral lamellae and gill pinnae. The gill consists of six unipinnate
branchial leaves. The perfoliate rhinophores are composed of 20 lamellae.

ANATOMY. — The buccal mass is divided evenly into an anterior glandular portion and a
posterior muscular one. At the posterior end of the mass there are a pair of large, elongate salivary
glands. The jaws are composed of a number of elongate, bifid rodlets (Fig. 7E) about 15 tm in length.
The radular formula is 38 x 41.0.41 in the holotype (CASIZ 115221). Rachidian teeth are absent. The
innermost lateral teeth (Fig. 7A, B) have one large denticle on the inner side of the cusp and three to
four denticles on the outer side. The remaining lateral teeth (Fig. 7C) are hook-shaped, lack denticles
on the inner side of the cusp and have a series of six to seven denticles along the outer edge. The outer
laterals (Fig. 7D) are elongate with six to seven denticles situated on the tips of the teeth.

The reproductive system (Fig. 6B) has an elongate and tubular ampulla that divides into the
oviduct and the prostate. The oviduct is very short and enters the female glands near the center of the
mass. The prostate is long, tightly coiled with several loops. It narrows into the muscular deferent
duct. The deferent duct is also very long and coiled, and opens into a common atrium with the vagina.
The penis is unarmed. The vagina is short and wide, slightly coiled. Near the end of the vagina the
uterine duct emerges. It is short and convoluted and opens into the female glands. More proximally
are the tightly coiled, digitiform seminal receptacle and the rounded, thin-walled bursa copulatrix.

ETYMOLOGY. — The species is named after Dr. Chandrakant G. Naik, National Institute of
Oceanography, Goa, India, for his support during the field work.

DISTRIBUTION. — So far, this species is only known from Mandapam, southern India.

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 13

FIGURE 7. Chromodoris naiki sp. nov. (CASIZ 115221), scanning electron micrographs. A. Inner lateral teeth, scale bar =
20 um; B. Inner lateral teeth, scale bar = 15 jm; C. Lateral teeth from the central portion of the half-row, scale bar = 25 um;
D. Outer lateral teeth, scale bar = 25 jum; E. Jaw rodlets, scale bar = 20 um.

REMARKS. — Chromodoris naiki is externally very similar to Chromodoris kitae Gosliner, 1994,
described from Madagascar. However, there are several external and internal differences that clearly
separate these two species. In C. kitae the yellow glands are only present in the mantle margin (see

Gosliner 1994), whereas in C. naiki they are also in the center of the dorsum, associated with the
purple spots. In addition, C. naiki has a number of small white spots in the center of the dorsum that

VALDES, MOLLO, AND ORTEA: NEW CHROMODORIDIDAE FROM SOUTHERN INDIA 47]

are absent in C. kitae. The number of branchial leaves is also different between them: C. kitae has
eight whereas C. naiki has six.

Internally these two species differ in their radular morphology. Whereas C. kitae has rachidian
teeth, they are absent in C. naiki. Also, the innermost lateral teeth of C. Aitae have one or two small
denticles on the inner side, clearly separated from the main cusp, whereas in C. naiki there is only
one, large denticle situated next to the main cusp. On the outer side of the cusp, C. kitae has two to
three denticles instead of three to four in C. naiki. The mid lateral teeth of C. naiki have a much longer
cusp than those of C. kitae. The reproductive system of C. kitae differs from that of C. naiki in having
a vestibular gland and a short, stalked seminal receptacle.

ACKNOWLEDGMENTS

We are very grateful to Dr. Chandrakant G. Naik and all the staff of the National Institute of
Oceanography, Goa, India, for their support during the field work. Terry Gosliner, Rebecca Johnson
and one anonymous reviewer made constructive comments on the manuscript.

Financial support for field work was made possible by an agreement between the Istituto per la
Chimica di Molecole di Interesse Biologico (CNR), Naples, Italy and the National Institute of
Oceanography, Goa, India. This paper has also been supported by the Ministerio de Educacion y
Cultura of Spain (SEUI), through its postdoctoral fellowships program.

LITERATURE CITED

ALDER, J. AND A. HANCOCK. 1964. Notice of a collection of nudibranchiate Mollusca made in India by Walter
Elliot, Esq., with descriptions of several new genera and species. Transactions of the Zoological Society of
London 5:113—147, pls. 28-33.

ELIOT, C. 1906a. On the nudibranchs of southern India and Ceylon, with special reference to the drawings by
Kelaart and the collections belonging to Alder and Hancock preserved in the Hancock Museum at
Newcastle-on-Tyne. Proceedings of the Zoological Society of London 1906:636—-691, pls. 42-47.

. 1906b. On the nudibranchs of southern India and Ceylon, with special reference to the drawings by

Kelaart and the collections belonging to Alder and Hancock preserved in the Hancock Museum at

Newcastle-on-Tyne. No. II. Proceedings of the Zoological Society of London 1906:999—1008.

. 1910. Notes on nudibranchs from the Indian Museum. Records of the Indian Museum 5:247—252,
jolly Wee

FARRAN, G. P. 1905. Report on the opisthobranchiate Mollusca collected by Professor Herdman, at Ceylon, in
1902. Ceylon Pearl Oyster Fisheries, Supplementary Reports, 21:329-364, pls. 1-6.

GOSLINER, T. M. 1994. New species of Chromodoris and Noumea (Nudibranchia: Chromodorididae) from the
western Indian Ocean and southern Africa. Proceedings of the California Academy of Sciences
48(12):239-252.

KELAART, E. F. 1858. New and little known species of Ceylon nudibranchiate molluscs, and zoophytes. Journal
of the Royal Asiatic Society, Ceylon Branch 3:84—139, 2 pls.

. 1859a. Descriptions of new and little-known species of Ceylonese nudibranchiate mollusks. Annals and

Magazine of Natural History (3)3:291—304.

. 1859b. On some additional species of nudibranchiate mollusks from Ceylon. Annals and Magazine of
Natural History (3)4:267-270.

NARAYANAN, K. R. 1969. On the opisthobranchiate fauna of the Gulf of Kutch. Pp. 188-213 in Proceedings of
the Symposium on Mollusca held at Cochin, I. Marine Biological Association of India, Mandapam, India.

O°’ DONOGHUE, C. H. 1932. Notes on nudibranchiata from southern India. Proceedings of the Malacological
Society of London 20:141—166.

RUDMAN, W. B. 1973. Chromodorid opisthobranch Mollusca from the Indo-West Pacific. Zoological Journal
of the Linnean Society 52:175—199, pls. 1-2.

472 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 13

. 1982. The Chromodorididae (Opisthobranchia: Mollusca) of the Indo-West Pacific: Chromodoris
quadricolor, C. lineolata and Hypselodoris nigrolineata colour groups. Zoological Journal of the Linnean
Society 76:105—173.

. 1985. The Chromodorididae (Opisthobranchia: Mollusca) of the Indo-West Pacific: Chromodoris
aureomarginata, C. verrieri and C. fidelis colour groups. Zoological Journal of the Linnean Society
83:241-299.

RUDMAN, W. B. AND B. W. DARVELL. 1990. Opisthobranch molluscs of Hong Kong. Part 1: Goniodorididae,
Onchidorididae, Triophidae, Gymnodorididae, Chromodorididae (Nudibranchia). Asian Marine Biology
7:31-79.

WHITE, K. M. 1948. Ona collection of marine molluscs from Ceylon. Proceedings of the Malacological Society
of London 27:199-205.

WINCKWorTH, H. C. 1945. Glossodoris from Bombay. Proceedings of the Malacological Society of London
26:155—160.

© CALIFORNIA ACADEMY OF SCIENCES, 1999
Golden Gate Park
San Francisco, California 94118

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES

Volume 51, No. 14, pp. 473-481, 6 figs., 1 table. December 9, 1999
DEC 20 1999

Acanthurus reversus, a New Species of Surgeonfish
(Perciformes: Acanthuridae) from the Marquesas Islands

by
John E. Randall and John L. Earle

Bishop Museum, 1525 Bernice St.,
Honolulu, Hawaii 96817-2704

Acanthurus reversus is described as a new acanthurid fish from five specimens from the
Marquesas Islands. It is most similar to Acanthurus olivaceus of the central and western
Pacific, differing principally in color. The caudal fin is pale yellow instead of gray-brown
with small dark spots, and it has a black crescent posteriorly instead of a dark-edged white
crescent; there is a broad, deep blue, longitudinal band in the humeral region with an
elongate orange spot centered anteriorly, not reaching more than halfway back within the
deep blue band (in A. olivaceus orange band is broader and extends nearly to the end of
the blue band). Like A. olivaceus the young are bright yellow. Acanthurus reversus also
differs in having longer dorsal spines (longest spine 15.2-16.3% SL, compared to
12.0-14.5% in A. olivaceus and the caudal spine is shorter on the average (7.7—10.8% SL,
compared to 9.8—11.5% in A. olivaceus).

While snorkeling in Takaroa Atoll in the northern Tuamotu Islands in November 1956 one of us
(J. E.R.) speared a surgeonfish, 175 mm standard length (measured fresh). It was much like the
wide-ranging Acanthurus olivaceus, but there was only a small elongate orange spot anteriorly within
the broad, deep blue band of the shoulder region, and the color of the caudal fin was very different,
pale yellow without dark spots, and with a black crescent posteriorly in the fin instead of a white one.
Typical A. olivaceus was common at the atoll, and the odd-colored fish was swimming with several
individuals of this species. Four of the 4. olivaceus specimens were speared from the area and
compared with the atypical specimen. No morphological differences were found, but two of the four
specimens of A. olivaceus had a narrower humeral orange band that was diffuse posteriorly; their
caudal coloration, however, was typical of 4. olivaceus. Due to limited formalin, only the odd-colored
specimen was preserved. It was deposited in the U. S. National Museum of Natural History as USNM
16981; the standard length is now 168 mm.

In July, 1957 J. E.R. sailed his 10-m ketch Nani from Tahiti through the Marquesas Islands.
There he found the odd-colored surgeonfish to be abundant, but no typical A. olivaceus were observed.
One specimen of the Marquesas Is. form, 196 mm SL, was speared at Anaho Bay, Nuku Hiva and
deposited in the Stanford Natural History Museum. Unfortunately, this specimen could not be found
in the collection of the California Academy of Sciences where the Stanford University fish collection
is now housed.

It was concluded that the one fish at Takaroa was a Marquesan form that was transported as a
larva from the Marquesas Is. in the prevailing current. Randall (1960) discussed the Marquesan
variant and indicated that it should be regarded as a subspecies of 4. olivaceus. He declined to name
it and indicated that additional specimens should be procured from both the Marquesas Is. and
Tuamotu Archipelago.

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In 1971 J. E. R. and colleagues visited the Marquesas Is. on the 30-m schooner Westward and
collected fishes for a month at all the major islands of the archipelago. An adult and a juvenile of the
Marquesan form of A. olivaceus were collected. An additional adult specimen was later obtained from
the Honolulu Laboratory of the National Marine Fisheries Service. When these were compared with
typical 4. olivaceus, no meristic differences and no obvious morphological differences were noted.
In a checklist of the fishes of French Polynesia, Randall (1985a) listed the Marquesan specimens as
A. olivaceus.

In October, 1998, the authors spent eight days on a dive cruise at the northern Marquesan islands
of Nuku Hiva and Eiao, mainly for underwater photography. Seeing the Marquesan color form of 4.
olivaceus again, and knowing how consistently colored the typical A. o/ivaceus is throughout the rest
of the Pacific (Fig. 1), led us to make a more careful comparison of Marquesan specimens with typical
A. olivaceus of similar size. We now find that the dorsal spines of the Marquesan form are longer,
and the caudal spine shorter on the average. Coupled with the marked color differences, we conclude
that the Marquesan form should be regarded as a distinct species. It seems likely that an occasionai
individual of this Marquesan surgeonfish drifts to the northern Tuamotu Archipelago as a larva and
ultimately interbreeds there with 4. o/ivaceus. Hybrids of closely related species of fishes may occur
when one is rare and the other abundant. Such is the case with 4. achilles and A. nigricans [=A.
elaucopareius| at the island of Hawaii where A. achilles is common and A. nigricans is rarely seen
(Randall 1985b).

MATERIALS AND METHODS

Type specimens of the new species are deposited in the Bernice P. Bishop Museum, Honolulu
(BPBM); California Academy of Sciences, San Francisco (CAS); Museum National d’ Histoire
Naturelle, Paris (MNHN); and the U.S. National Museum of Natural History, Washington, D. C.
(USNM).

Lengths of specimens are given as standard length (SL), which is measured from the median
anterior end of the upper lip to the caudal-fin base (posterior end of hypural plate); head length is
taken from the same anterior point to the posterior end of the opercular flap; body depth 1s the greatest
depth from the base of the dorsal spines to the ventral margin of the abdomen (correcting for any
obvious malformation of preservation); body width is measured just posterior to the gill opening;
orbit diameter is the greatest fleshy diameter, and interorbital width the least bony width; upper-jaw
length is taken from the front of the upper lip to the posterior end of the maxilla; caudal-peduncle
depth is the least depth, and caudal-peduncle length the horizontal distance between verticals at the
rear base of the anal fin and the caudal-fin base; lengths of fin spines and rays of the median fins are
measured from their extreme bases; caudal concavity is the horizontal distance between verticals at
the tips of the longest and shortest caudal-fin rays; pectoral-fin length is the length of the longest ray;
pelvic-fin length is measured from the base of the pelvic spine to the tip of the longest soft ray.
Pectoral-ray counts include the upper rudimentary ray. Counts of gill-rakers were made on both the
anterolateral and posteromedial side of the first gill arch (here termed anterior and posterior); the
counts include all rudiments.

Data in parentheses in the description refer to paratypes. Table | presents 26 measurements as
percentages of the standard length. Ratios of proportional measurements in the text are rounded to
the nearest 0.05.

RANDALL AND EARLE: NEW SPECIES OF SURGEONFISH 475

Figure |. Adult of Acanthurus olivaceus, about 240 mm TL, Kona, Hawaii.

SPECIES DESCRIPTION

Acanthurus reversus n. sp.
Figs. 2-6

MATERIAL EXAMINED. — HOLOTYPE: BPBM 12474, 209 mm, female, Marquesas Islands, Nuku
Hiva, off point at S end of Marquisienne Bay, 15 m, spear, J. E. Randall, 10 May 1971. PARATYPES:
USNM 169871, 168 mm male, Tuamotu Archipelago, Takaroa Atoll, outer reef slope, E side of
entrance to pass, spear, J. E. Randall, 16 November 1956; CAS 204429, 210 mm, female, Marquesas
Islands, Tahuata, Hana Hevane Bay, rotenone, Charles H. Gilbert Cruise 54, J. Manguson, 16 October
1961; MNHN 1999-60, 216 mm, male, Marquesas Islands, Ua Huka, small bay 0.4 miles NE of Motu
Takatai, 6 m, spear, J. E. Randall, 7 May 1971; BPBM 12441, 36 mm, same locality as preceding,
4.5—9 m, rotenone, J. E. Randall, J. R. Haywood, and R. M. McNair, 7 May 1971.

DIAGNOSIS. — Dorsal rays [X,24—25; anal rays III,23—24; pectoral rays 16-17; anterior gill
rakers 22—28: posterior gill rakers 21-27; body depth 2.1—2.2 in SL; head length 3.5—3.65 in SL; ninth
dorsal spine 1.7(1.8—1.85) in head length; caudal spine 2.3—2.65 in head length; caudal fin strongly
lunate in adults, the caudal concavity 3.35—4.7 in SL; color in life brown, the posterior half of body
sometimes abruptly paler, with an elongate orange spot extending posteriorly from upper end of gill
opening, the posterior half enclosed by a broad, deep blue band that extends beyond tip of pectoral
fin; dorsal and anal fins with a faint orange line at base, the soft portion of the dorsal with three dark
longitudinal bands on outer one-half of fin, the anal fin with a narrow blue margin; caudal fin pale
yellow with a broad black posterior border that narrows as it extends onto elongate lobes of fin; outer
one-third of pectoral fins abruptly white; juveniles bright yellow with blue margins on dorsal and anal
fins.

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FIGURE 2. Holotype of Acanthurus reversus 209 mm SL, Nuku Hiva, Marquesas Is.

DESCRIPTION. — Dorsal rays IX,25(24), the first 3 soft rays unbranched; anal rays 23(23—24),
the first 2 soft rays unbranched; pectoral rays 16(16—17), the upper 2 and lowermost unbranched;
pelvic rays I,5; principal caudal rays 16, the outermost unbranched; upper and lower procurrent caudal
rays 7; scales small, 157 in longitudinal series from upper end of gill opening to caudal-fin base of
holotype; anterior gill rakers 22(22—28); posterior gill rakers 24(21—27).

Body deep, the depth 2.15(2.1—2.2) times into SL, and compressed, the width 3.0(2.55—3.15) in
depth; head length 3.65(3.5-3.6) in SL; dorsal profile of head strongly convex; snout length
4.9(4.75-4.8) in SL; orbit diameter 4.65(3.75—4.3) in head length; interorbital strongly convex, the
least width 3.25(3.0-3.2) in head length; caudal-peduncle depth 2.6(2.6—2.7) in head length.

Mouth small, the gape slightly oblique, the upper jaw length 4.1(3.64.1) in head length; teeth
uniserial in jaws, incisiform, spatulate with denticulate edges, and close-set; holotype with 20 upper
and 18 lower teeth; (36-mm juvenile paratype with 12 upper and 14 lower teeth; 169-mm paratype
with 16 upper and lower teeth; 216-mm paratype with 19 upper and 20 lower teeth).

Scales on body coarsely ctenoid, the cteni on posterior edge of scales in middle of body 10 to 18;
scales on head embedded; lateral line obscure, approximately following dorsal contour of body and
ending at front of caudal-spine socket; a finely-scaled fleshy sheath at base of dorsal and anal fins;
very small scales extending about one-third distance out on membranes of posterior one-half of dorsal
and anal fins; extremely small scales extending out on caudal fin more than three-fourths distance to
outer margin: pectoral fins with extremely small, embedded scales extending about three-fourths
distance to outer margin; lateral surface of pelvic fins with a narrow column of tiny scales on first
soft ray and membrane about one-half distance to tip of fin, the medial edge with a narrow band of
extremely small scales on each membrane extending about one-half distance to posterior edge of fin.

A deep groove extending anteriorly and slightly ventrally from lower edge of orbit, its length
about three-fourths orbit diameter; anterior nostril small, centered above groove anterior to eye, with
a membranous rim and rounded posterior flap which completely covers nostril aperture when folded
forward, and just reaches slit of posterior nostril when folded back.

Dorsal-fin origin above upper end of gill opening, the predorsal length 2.55(2.6—2.7) times into
SL; first dorsal spine about one-half length of second spine; second dorsal spine 2.5(2.45—2.5) in head
length; ninth dorsal spine longest, 1.7(1.8-1.85) in head length; first dorsal soft ray longest,

RANDALL AND EARLE: NEW SPECIES OF SURGEONFISH

>

FIGURE 3. Adult of Acanthurus reversus about 240 mm TL, Eiao, Marquesas Is.

FiGuRE 4. Group of adult Acanthurus reversus feeding, Eiao, Marquesas Is.

478 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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1.6(1.65—1.7) in head length; anal-fin origin below base of seventh dorsal spine, the preanal length
2.0(2.0—2.05) in SL; first anal spine very short; second anal spine 2.85(2.55—2.8) in head length; third
anal spine 2.4(2.2—2.45) in head length; middle anal soft rays longest, 1.75(1.7—1.75) in head length;
caudal fin of adults long, the fin length 2.65(2.05—2.4) in SL, and extremely lunate, the caudal
concavity 4.35(3.35—4.7) in SL; third and fourth pectoral rays longest, 3.9(3.6—3.7) in SL; pelvic fins
reaching anal-fin origin, the first ray longest, 4.8(4.45—-4.65) in SL; caudal spine 2.35(2.3—2.65) in
head length.

Color of holotype in alcohol dark brown with a horizontally elongate, ellipsoidal, pale orangish
spot extending posteriorly from upper end of gill opening, its length 1.5 times orbit diameter, this
spot contained within a dark brown band a little broader than orbit diameter, extending slightly beyond
tip of pectoral fin; caudal fin lighter brown than body, with a black posterior margin that is one-half
orbit diameter in width centrally, but narrows to a black line on the prolonged caudal lobes; upper
and lower edges of caudal fin narrowly black; dorsal, anal, and pelvic fins dark brown, the outer part
of soft portion of dorsal with three faint longitudinal darker brown bands; pectoral fins dark brown,
the outer one-third abruptly whitish except second ray brown to tip.

The color of the holotype when fresh is shown in Figure 2. Note that the dark brown humeral
band as seen on preserved specimens is deep blue with a black edge in life, and only the black edge
continues around the front of the orange spot. In life (Figs. 3, 4), the caudal fin of adults is pale yellow,
contrasting sharply with the black posterior crescent of the fin, and the blue margin of the anal fin is
more evident.

The bright yellow juvenile of Figure 4 was estimated to be 45 mm total length; note the truncate
caudal fin. Figure 5 shows two subadults estimated to have been 90 mm total length; their caudal fins
are more yellow than that of adults, but not as yellow as the color of the juvenile.

ETYMOLOGY. — This species 1s named Acanthurus reversus from the Latin with the same
meaning as English, in reference to the reversal of the color pattern of the caudal fin compared to that
of its close relative, 4. olivaceus.

REMARKS. — Acanthurus reversus is endemic to the Marquesas Islands, and the one specimen
collected at Takaroa in the northern Tuamotu Archipelago 1s regarded as a stray from the Marquesas.
The species is common inshore at all the islands of the Marquesas, but it has a different native
Marquesan name at each of the major islands (Lavondes and Randall 1978).

Acanthurus olivaceus Forster in Bloch and Schneider, 1801, type locality, Tahiti, is clearly its
closest relative. This surgeonfish remains unknown from the Marquesas, but it is common throughout
the rest of the central and western Pacific and ranges into the eastern Indian Ocean to western
Australia, Christmas Island, and Cocos (Keeling) Islands; in the western Pacific it extends from
southern Japan to New South Wales. It appears to be replaced in the western Indian Ocean by
Acanthurus tennenti Giinther (Randall, 1956).

As noted above, the Marquesan surgeonfish was first considered to be only a color variant of A.
olivaceus and subspecific status was suggested. The most obvious color differences are the reduced
size of the orange part of the humeral band in the Marquesan fish and its having a pale yellow caudal
fin with a black posterior crescent instead of a gray-brown fin with small dark brown spots and a
black-edged white posterior crescent. When four adults of 4. reversus were recently compared with
11 adults of A. olivaceus of about the same size from the collections of the Bishop Museum and
California Academy of Sciences (specimens listed below), no meristic differences were detected, but
the dorsal spines of 4. reversus were noted to be longer, and the caudal spine shorter. No overlap was
found in the measurements of the longest (ninth) dorsal spine, 15.2-16.3% SL for 4. reversus
compared to 12.0-14.5% for A. olivaceus specimen. Two of the 11 specimens of A. olivaceus have
shorter caudal spines than the A. reversus specimen with the longest spine. The caudal spine of the
four A. reversus specimens are 7.7—10.8% SL (X = 9.4 ), compared to 9.8-11.5% (X = 11.0) for the
eleven A. olivaceus specimens.

RANDALL AND EARLE: NEW SPECIES OF SURGEONFISH

Ficure 5. Juvenile of Acanthurus reversus about 45 mm TL, Nuku Hiva, Marquesas Is.

FiGuRE 6. Subadults of Acanthurus reversus about 90 mm TL, Eiao, Marquesas Is.

480 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 14

TABLE |. Proportional measurements of type specimens of Acanthurus reversus expressed as percent of
standard length (% SL).

Holotype

Paratypes
BPBM 12474 USNM 169871 MNHN 1999-60

CAS 204429

Sex Female Male Female Male
Standard length (mm) 209 168 210 216

Body depth 46.9 47.0 45.6 45.8
Body width a7) 14.9 17.9 ga
Head length DAES 28.7 DES 28.4
Snout length 203 2 20.9 22
Orbit diameter 5.8) Te 6.6 6.6
Interorbital width 8.4 OF 2) 8.9
Upper-jaw length 6.7 7.8 7.8 6.9
Caudal-peduncle depth 10.5 HO 10.6 11.0
Caudal-peduncle length 11.0 eS 11.4 ned
Predorsal length 39.0 37.7 38.6 38.0
Preanal length 50.5 50.2 50.3 49.0
Prepelvic length 32.8 34.4 33.9 33.4
Dorsal-fin base O12 69.5 67.0 6955
Second dorsal spine 11.0 broken lies ale?
Last dorsal spine fos 16.0 15.7 sea
Longest dorsal soft ray ives 16.7 16.8 16.8
Anal-fin base 45.0 44.4 45.6 44.5
Second anal spine oN 10.3 10.9 10.2
Third anal spine eS) 12.5 27 Lhe
Longest anal soft ray 5) 16.9 16.6 16.2
Caudal-spine length deh 10.8 8.4 10.7
Caudal-fin length SHAS broken 41.2 49.]
Caudal concavity 23.0 —~ 2 30.0
Pectoral-fin length 26.3 PLT Dial 275
Pelvic-spine length 14.3 14.6 13.8 14.1
Pelvic-fin length 20.8 22.4 INNS) 21.6

An updated checklist of the shore fishes of the Marquesas Islands has been completed (Randall
and Earle, in press). The total number of shore fishes is 401, a relatively impoverished fish fauna
compared to the Society Islands with 593 species (Randall 1985a). But the level of endemism of the
fishes in the Marquesas Is. is high, 11.2%. This is third highest in the Pacific, after the Hawaiian
Islands and Easter Island, with 23.1 and 22.2%, respectively (Randall 1998). The Marquesas Is. are
not as isolated as Hawaii and Easter Island, the nearest shoal water being the northern atolls of the
Tuamotu Archipelago some 500 km to the south. However, the prevailing South Equatorial Current
sweeps to the west, and the nearest shoal water to the east that could be the source of larvae of inshore
marine animals is the Galapagos Islands, 3700 km away. Only a rare change in current pattern could
result in recruiting larvae from the Tuamotu Archipelago or Caroline Atoll to the west.

MATERIAL OF ACANTHURUS OLIVACEUS EXAMINED. — CAS 53992, 214 mm, Hawaiian Islands,
Oahu; CAS 31961, 2:180—-185 mm, Hawaiian Islands, Oahu; BPBM 4284, 208 mm, Johnson Island;
BPBM 4287, 227 mm, Johnson Island; BPBM 6222, 2:200—-219 mm, Line Islands, Baker Island; CAS
96922, 196 mm, Caroline Islands, Kapingamarangi Atoll; CAS 96926, 186 mm, Caroline Islands,
Kapingamarangi Atoll.

RANDALL AND EARLE: NEW SPECIES OF SURGEONFISH 481

LITERATURE CITED

BLocH, M. E. AND J. G. SCHNEIDER. 1801. Systema ichthyologiae iconibus cx illustratum. Post obitum auctoris
opus inchoatum absoluit, correxit, interpolavit. Sanderiano Commissum, Berlin. |x + 584 pp.

LAVONDFS, H. AND J.E. RANDALL. 1978. Les noms de poissons marquisiens. Jour. Soc. Oceéanistes
34(60):79-1 12.

RANDALL, J. E. 1956. A revision of the surgeon fish genus Acanthurus Pacif. Sci. 10(2):159-235.

. 1960. A new species of Acanthurus from the Caroline Islands, with notes on the systematics of other

Indo-Pacific surgeonfishes. Pacif. Sci. 14(3):267-279.

. 1985a. Fishes. Pp. 462-481 in French Polynesian Coral Reefs. B. F. Delesalle, R. Galzin and B. Salvat,

eds., Vol. 1, Fifth International Coral Reef Congress, Tahiti, 27 May 1|-June 1985.

. 1985b. Guide to Hawaiian reef fishes. Harrowood Books, Newtown Square, Pennsylvania. 74 pp.

. 1998. Zoogeography of shore fishes of the Indo-Pacific region. Zool. Stud. 37(4):227-268.

RANDALL, J. E. AND J. L. EARLE. In press. Annotated checklist of the shore fishes of the Marquesas Islands, with
40 new records. Occ. Papers, B. P. Bishop Mus., Hawaii.

© CALIFORNIA ACADEMY OF SCIENCES, 1999
Golden Gate Park
San Francisco, California 94118

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PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES

Volume 51, No. 15, pp. 483-492, 3 figs. December 9, 1999
DEC 2 0 1999

Taxonomic and Distributional Notes on Neotropical
Justicia (Acanthaceae)

by
Thomas F. Daniel

Department of Botany, California Academy of Sciences
Golden Gate Park, San Francisco, California 94118

Three species of Justicia are newly described as J. chimalapensis, J. novogaliciana, and J.
wendtii. All are endemic to Mexico. Macromorphological and palynological aspects of these
species are considered in an attempt to elucidate the subgeneric affinities of each. The new
name, J. isthmensis, is proposed for the Panamanian species originally described as
Chaetochlamys panamensis. The geographic distribution of J. candelariae is extended into
Panama.

Justicia L. is the largest genus of Acanthaceae with estimates of up to 600 species worldwide.
Numerous genera in which Neotropical species have been treated are now included within an
expanded circumscription of Justicia (Graham 1988; Daniel 1990, 1995a); these include Beloperone
Nees, Chaetothylax Nees, Chaetochlamys Lindau, /xtlania M. E. Jones, Jacobinia Nees, Neohallia
Hemsl., Siphonoglossa Oerst., and Tabascina Baill. Additional Neotropical genera that are currently
recognized, but insufficiently studied, will likely be added to this growing list.

Graham (1988) presented the most recent and comprehensive infrageneric classification of
Justicia to date. Although her treatment, based on and encompassing 295 species, provides a useful
starting point for additional work on the genus, numerous Neotropical species cannot be classified in
any of the subgeneric taxa recognized (e.g., Daniel 1990, 1993b, 1995a, 1998; Durkee and McDade
1996). Additional systematic studies of Justicia and its allies are greatly needed.

Seventy-eight species of Justicia are known from Mexico (Daniel 1993a) and approximately 100
species are known from the Mesoamerican region (Durkee and Daniel, unpublished). Numerous
collections from Mexico and Central America do not correspond to any of these species and many of
them undoubtedly represent undescribed taxa. Several of these are described below and, where
possible, their subgeneric affiliations are noted. In addition, a new name is proposed for a species
previously treated in Chaetochlamys and the distributional range of J. candelariae is expanded to
Panama.

NEW SPECIES DESCRIPTIONS

Justicia wendtii T. F. Daniel, sp. nov.
Fig. |

TyPE. — MEXICO. Veracruz: Mpio. Minatitlan, 13.7 km E de La Laguna, sobre terraceria a
Uxpanapa, luego 7.2 km N sobre camino nuevo a Belisario Dominguez, 130 m, lat. 17°20.5’N, long.
94°23'W, 21 March 1981, 7. Wendt, A. Villalobos C., and I. Navarrete 3031 (holotype: CAS!;
isotypes: CAS!, CHAPA, MEXU, TEX).

483

484 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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Herbae perennes usque ad 7 dm altae. Folia petiolata, laminae ovato-ellipticae vel ellipticae,
60-120 mm longae, 24-51 mm latae, 2—3-plo longiores quam latiores. Inflorescentia floribus in
spicas (vel paniculas spicarum) terminales vel axillares; dichasia subopposita vel alterna, sessilia,
uniflora. Bracteae aliquando + heteromorphae, obovatae vel oblanceolatae vel lineari-ellipticae,
3.3-8.5 mm longae, 1—2.5 mm latae, pagina abaxialis pubescens trichomatibus eglandulosis et
glandulosis. Flores sessiles. Calyx 5-lobus, 34.5 mm longus, lobis subaequalibus vel inaequalibus.
Corolla viridi-alba labio inferno purpureo-notato, 7-8.5 mm longa, extus pubescens trichomatibus
glandulosis (aliquando sparsis) et eglandulosis. Stamina thecis 0.8—1.2 mm longis, impariter insertis,
theca inferna basi calcarata. Capsula 7-8 mm longa, pubescens trichomatibus eglandulosis et glan-
dulosis.

Erect to decumbent perennial herbs to 7 dm tall. Young stems subterete to subquadrate, pubescent
with retrorsely appressed eglandular trichomes 0.2—0.4 mm long, trichomes + evenly disposed or
concentrated in 2 lines. Leaves petiolate, petioles to 35 mm long, blades ovate-elliptic to elliptic,
60—120 mm long, 24-51 mm wide, 2.0—3.0 times longer than wide, acute to acuminate at apex, acute
to subattenuate at base, adaxial surface glabrous, abaxial surface pubescent along major veins with
antrorse to antrorsely appressed eglandular trichomes to 0.3 mm long, margin entire to subsinuate.
Inflorescence of axillary and terminal pedunculate dichasiate spikes (or panicles of spikes) to 85 mm
long (including peduncles and excluding flowers), spikes 6-9 mm in diameter (excluding flowers)
near midpoint of fertile portion, spikes and/or panicles near shoot apex sometimes subtended by
greatly reduced leaves and collectively forming an open terminal compound panicle, axillary spikes
(or panicles of spikes) alternate to opposite, | per axil, peduncles to 40 mm long, pubescent with
flexuose to retrorse eglandular trichomes to 0.4 mm long, rachis of panicles (if present) pubescent
like peduncles, rachis of spikes evenly pubescent with erect to flexuose (and sometimes with some
appressed trichomes near base of rachis) eglandular and glandular trichomes 0.05—0.2 mm long;
dichasia subopposite to alternate, 1-flowered, | per axil, sessile. Bracts subopposite to alternate,
sometimes somewhat heteromorphic (i.e., sterile bracts sometimes somewhat smaller than fertile
bracts), obovate to oblanceolate, 3.3-8.5 mm long, I|—2.5 mm wide, acute to rounded at apex, abaxial
surface pubescent like rachis of spike, margin ciliate with trichomes like those of abaxial surface and
with scattered flexuose eglandular trichomes to 0.3 mm long as well. Bracteoles obovate to oblanceo-
late to linear-elliptic, 2.7-7.3 mm long, 0.7—1.2 mm wide, pubescent like bracts. Flowers sessile.
Calyx 5-lobed, 34.5 mm long, lobes lance-subulate, subequal to unequal, 2.64 mm long, 0.4—0.6
mm wide, abaxially pubescent like bracts. Corolla greenish cream with purplish markings on lower
lip, 7-8.5 mm long, externally pubescent with eglandular and glandular (sometimes sparse) trichomes
0.05—0.1 mm long, tube cylindric, 44.8 mm long, 1.5—1.7 mm in diameter near midpoint, upper lip
2.5-4 mm long, emarginate, lobes 0.2-0.4 mm long, lower lip 3.5-4.5 mm long, lobes 1—1.7 mm
long, 0.7-2 mm wide. Stamens inserted near apex of corolla tube, 3.3-3.4 mm long, filaments
glabrous, thecae 0.8—1.2 mm long (including basal appendage), unequal, parallel, unequally inserted
(overlapping by up to 0.2 mm), dorsally pubescent (especially upper theca) with eglandular trichomes,
lower theca with a prominent basal appendage 0.3—0.4 mm long; pollen 3-aperturate, 6-pseudocolpate
(i.e., apertures flanked on each side by both a continuous band of exine and a pseudocolpus), exine
reticulate. Style 4.5-5.5 mm long, sparsely pubescent with antrorse eglandular trichomes, stigma
0.1-0.2 mm long, only one lobe evident. Capsule 7-8 mm long, externally pubescent with erect
glandular and erect to flexuose to retrorse eglandular trichomes 0.05—0.1 mm long, stipe 2.6—3.5 mm
long, head subspheric to subellipsoid, 44.8 mm long. Seeds 4, lenticular, 1.92.3 mm long, 1.7—2.2
mm wide, surfaces minutely rugose and sparsely pubescent with apically branched eglandular
trichomes 0.05—0.1 mm long, margin entire, conspicuously ciliate with apically branched trichomes.

DANIEL: NOTES ON NEOTROPICAL JUSTICIA A485

FIGURE |. Justicia wendtii. a. habit (Wendt et al. 3031); b. inflorescence nodes (Wendt et al. 3031); c. corolla opened
lengthwise showing stamens (Wendt and Hernandez 6727), d. apex of stamen with bithecous anther (Wendt and Hernandez
6727); e. apex of style and stigma (Wendt and Hernandez 6727), f. capsule (Vazquez T. et al. V-2413); g. seed (Vazquez T. et
al. V-2413): h. seminal surface (Vazquez T. et al. V-2413). Scale: a, bar = 15 mm; b, bar = 2.1 mm: c, bar = 1.6 mm; d, bar =
0.57 mm; e, bar = 0.5 mm; f, bar = 2 mm; g, bar = 0.7 mm; h, bar = 0.37 mm. Drawn by Jenny Speckels.

486 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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PHENOLOGY. — Flowering and fruiting: March—April.

DISTRIBUTION AND HABITAT. — Southeastern Mexico (Veracruz, Fig. 2); plants occur on highly
karsted limestone in lowland rain forests at elevations from 120—130 m.

PARATYPES. — MEXICO. Veracruz: Mpio. Jesus Carranza, 2 km N del Poblado 2, Ejido F. J.
Mina, 17°16’N, 94°40'W, M. Vazquez T. et al. V-2413 (CAS, LL); Mpio. Minatitlan, 13.7 km E of
La Laguna toward Uxpanapa, then 6.5—7.2 km N toward Belisario Dominguez, 17°19’N, 94°24'W,
T. Wendt and H. Hernandez G. 6727 (CAS), 5606 (CAS, TEX).

Based on the infrageneric classification of Graham (1988), this species “keys” closest to sections
Sarotheca and Plagiacanthus. Although Justicia wendtii shares some characteristics (e.g., glandular
inflorescence) with the former and other characteristics (e.g., calyx lobes more or less equal and
strongly compressed seeds) with the latter, it differs from both of these sections in several features,
including its 3-aperturate pollen (i.e., Graham’s “Type 1”; Fig 3). Graham (1988) found “Type 1”
pollen only in sections Betonica and Rhaphidospora, both of which are restricted to the Old World,
and in one Neotropical species (J/. alainii Stearn) of section Drejerella. Thus, the subgeneric affinities
of J. wendtii are ambiguous.

A collection from cloud forest habitat on granitic substrate at an elevation between 1150 and
1250 meters in this same general region of Mexico (Wendt et al. 6803 from the Sierra de Tres Picos
in adjacent Oaxaca) resembles J. wendtii in most characteristics (e.g., habit, inflorescence, stamens,
and pollen). In addition to the contrasting ecological parameters noted, this collection differs from J.
wendtii most notably by its somewhat longer calyx (4.5—-6 mm long) that lacks glandular trichomes
on the lobes, longer corolla (9-12 mm long), capsular pubescence (with the glandular trichomes
inconspicuous and sessile vs. conspicuous and stipitate in J. wendtii), and seed surface ornamentation
(minutely pitted, the margin with a few inconspicuous trichomes less than 0.05 mm long that lack
apical branching). Only a few capsules and seeds of this collection were available for study. Study of
additional plants from this region might help to clarify whether Wendt et al. 6803 should be recognized
as a distinct taxon or included within J. wendtii.

The species is named in honor of Tom Wendt, botanical explorer in southeastern Mexico and
collector of this and other unusual Acanthaceae.

Justicia chimalapensis T. F. Daniel, sp. nov.

TyPE. — MEXICO. Oaxaca: Mpio. Santa Maria Chimalapa, camino hacia Arroyo San Vicente,
990 m, 23 March 1995, E. Torres B. 536 (holotype: CAS!).

Herbae perennes vel frutices usque ad 1.5 m alti. Folia petiolata, laminae ovatae vel ellipticae,
105-210 mm longae, 33-93 mm latae, 2.3—3.2-plo longiores quam latiores. Inflorescentia floribus in
spicas vel paniculas spicarum axilares; dichasia alterna, sessilia, uniflora, et unilateralia. Bracteae
homomorphae, subulatae, 1-2 mm longae, 0.3—0.4 mm latae, pagina abaxialis glabra vel sparse
pubescens. Flores sessiles vel subsessiles. Calyx 5-lobus, 4.5-6 mm longus, lobis homomorphis.
Corolla viridi-alba labiis purpureo-notatis, 16—20.3 mm longa, extus pubescens trichomatibus glan-
dulosis. Stamina thecis 1.8—2.2 mm longis, subpariter vel impariter insertis, theca inferna basi
calcarata. Capsula pubescens trichomatibus glandulosis.

Erect perennial herbs or shrubs to 1.5m tall. Young stems multiridged with shallow and
somewhat undulating ridges, glabrous, + conspicuously constricted just above nodes. Leaves petio-
late, petioles to 33 mm long, blades ovate to elliptic, 105-210 mm long, 33—93 mm wide, 2.3—3.2
times longer than wide, + abruptly acuminate to acuminate-caudate at apex, acute to + abruptly
attenuate at base, surfaces glabrous or with antrorse eglandular trichomes on midvein adaxially (at

DANIEL: NOTES ON NEOTROPICAL JUSTICIA 487

@ J. chimalapensis

O

J. novogaliciana

J. wendtii

hy
Sa

FiGuRE 2. Map of Mexico showing the distributions of Justicia chimalapensis, J. novogaliciana, and J. wendtii.

least near base of blade), margin entire to sinuate. Inflorescences of axillary subsessile to short-pedun-
culate dichasiate spikes or basally branched panicles of spicate axes to 50 mm long (including
peduncles and excluding flowers), spikes or panicles opposite, | per axil, peduncles to 3 mm long,
glabrous or pubescent with eglandular trichomes, bracts subtending panicle branches (= inflorescence
bracts) subulate, 1.5 mm long, 0.3—0.5 mm wide, rachis of spikes or spicate axes nearly glabrous or
evenly pubescent with erect to flexuose glandular trichomes to 0.2 mm long (glandular pubescent)
and antrorse eglandular (at least proximally) trichomes 0.05—0.1 mm long; dichasia alternate, |-flow-
ered, | per axil, sessile, unilateral. Bracts opposite, homomorphic, subulate, |—2 mm long, 0.3—0.4
mm wide, abaxial surface glabrous or sparsely pubescent with trichomes like those of rachis, eciliate.
Bracteoles lance-subulate, 1.8—2 mm long, 0.3—0.7 mm wide, abaxial surface glabrous or glandular
pubescent. Flowers sessile to subsessile (1.e., pedicels to 0.5 mm long). Calyx 5-lobed, 4.5-6 mm
long, lobes homomorphic, lance-subulate, 3.34.5 mm long, 0.5—0.7 mm wide, abaxially glandular
pubescent. Corolla with long axis + vertically oriented, greenish white with purplish markings,
16—20.3 mm long, externally glandular pubescent, tube + cylindric, subsaccate below midpoint,
10-12 mm long, 1.7—2.5 mm in diameter near midpoint, upper lip 6—9.3 mm long, apically 2-lobed,
lobes 0.1—0.5 mm long, lower lip 6-9 mm long, lobes 1.3—2 mm long, 1.6—2.5 mm wide. Stamens
inserted below midpoint of corolla tube, 12-16 mm long, filaments glabrous distally, pubescent with
eglandular trichomes near base, thecae 1.8—2.2 mm long (including basal appendage), subequal,
subparallel, subequally to unequally inserted (overlapping by up to 1.8 mm), upper theca dorsally
pubescent with eglandular trichomes to 0.05 mm long, lower theca glabrous and with a prominent
basally papillose basal appendage 0.3—0.5 mm long; pollen 2-aperturate, apertures flanked on each
side by | row of insulae (peninsulae sometimes present as well), exine reticulate; 2 staminode-like

488 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
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thickenings or pouchlike protrusions of corolla present at point of insertion of filaments, these
pubescent with eglandular trichomes. Style 15—15.5 mm long, pubescent with eglandular trichomes,
stigma 0.2—0.3 mm long, only | lobe evident. Capsule (immature) 5.5 mm long, externally sparsely
pubescent with glandular trichomes to 0.05 mm long.

PHENOLOGY. — Flowering: March—April. Immature fruits present in March.

DISTRIBUTION AND HABITAT. — Mexico (Oaxaca, Fig. 2), in the Isthmus of Tehuantepec; plants
occur in cloud forest (bosque mesofilo de montafia) at elevations from 990 to 1200 m.

PARATYPE. — MEXICO. Oaxaca: Mpio. Santa Maria Chimalapa, Sierra de Tres Picos, ca. 1.5
km W of Cerro Piicotzuc, ca. 17.5 km SSE of La Laguna, Veracruz, 17°07'30"N, 94°27'55"W, T.
Wendt.et al. 6830 (TEX).

This species cannot be readily affiliated with any one of the subgeneric taxa recognized by
Graham (1988). Its pollen (“Type 5” of Graham 1988; Fig. 3) is characteristic of several of her
sections, as well as numerous species of uncertain affinities. Like several other species of Justicia
(i.e., J. chol T. F. Daniel, J. costaricana Leonard, and J. nevlingii Wassh. and T. F. Daniel; see
Wasshausen and Daniel 1995), J. chimalapensis has two pubescent, staminode-like thickenings
adjacent to the point of insertion of the filaments in the corolla tube. These thickenings form two
conspicuous, pouchlike protrusions of the corolla into the tube. Justicia chimalapensis differs from
all of these species by having homomorphic bracts. It appears unlikely that these staminode-like
thickenings represent a synapomorphy uniting these taxa because they differ in numerous macromor-
phological and micromorphological attributes. Wasshausen and Daniel (1995) summarized the
differences among the species with heteromorphic bracts.

Wendt et al. 6530 lacks the conspicuous glands on the inflorescence rachises and bracteoles
(although a few glands are present on these structures), but otherwise resembles the holotype.

The name of this species is derived from the municipality in which it was collected.

Justicia novogaliciana T. F. Daniel, sp. nov.

Type. — MEXICO. Jalisco: Sierra de Manantlan Occidental, slopes of La Calera, just NW of
KM 188 on Autlan-Manzanillo hwy (Mex. 80), 9 km NNE of La Resolano (Casimiro Castillo) and
ca. 16 km SE of Autlan, 19°40'N, 104°24’—25'’W, 11 March 1992, H. /Itis, L. Guzman H., and B. Benz
31028 (holotype: WIS!; isotype: CAS!).

Frutices usque ad 2 m alti. Folia petiolata, laminae ovatae vel ellipticae (vel obovato-ellipticae),
18-82 mm longae, 10-32 mm latae, |.3—3-plo longiores quam latiores. Inflorescentia floribus in
dichasia axillaria; dichasia alterna vel opposita, sessilia vel pedunculata, (I—) S—multiflora. Flores
subsessiles vel pedicellati. Calyx pariter 4-lobus vel impariter 5-lobus, 45.5 mm longus. Corolla
aurantiaca vel rubens, 23-33 mm longa, extus pubescens trichomatibus glandulosis et eglandulosis.
Stamina thecis 2—2.5 mm longis, subpariter insertis, basi ecalcaratis. Capsula 20 mm longa, pubescens
trichomatibus glandulosis et eglandulosis.

Shrub to 2 m tall. Young stems striate with alternating light green and darker green-brown lines,
subquadrate to subhexagonal, (+ evenly to) bifariously pubescent with (flexuose to) retrorse eglan-
dular trichomes 0.3—0.8 mm long (trichomes sparse, + unifarious, or merely concentrated in 2 lines
on some internodes). Leaves petiolate, petioles to 9 mm long, blades ovate to elliptic (to obovate-el-
liptic), 18-82 mm long, 10-32 mm wide, 1.3-3 times longer than wide, (rounded to) acute to
acuminate (to apiculate) at apex, (rounded to) acute to subattenuate at base, surfaces pubescent with
flexuose eglandular trichomes to 0.5 mm long, trichomes concentrated along major veins, margin
entire to subsinuate. Inflorescence of axillary sessile to pedunculate dichasia to 50 mm long (including

DANIEL: NOTES ON NEOTROPICAL JUSTICIA

FiGuRE 3. Scanning electron micrographs of pollen. a. Justicia wendtii (Wendt et al. 3031), interapertural view. b. J.
chimalapensis (Torres B. 536), apertural view. c. J. novogaliciana (IItis et al. 31028), apertural view. d. J. novogaliciana (IItis
etal. 31028), interapertural view. Scale for a = 5 ym: scale for b-d = 10 um.

490 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 15

peduncle and excluding flowers), dichasia alternate or opposite, (1—) 5—many-flowered, | per axil,
sessile or pedunculate, peduncles to 2.5 (-22) mm long, pubescent with flexuose to retrorse to antrorse
eglandular trichomes. Bracteoles lance-subulate to triangular-subulate, 1.72.5 mm long, 0.4-0.8 mm
wide, abaxial surface pubescent with erect to flexuose to antrorse eglandular (and sometimes
glandular) trichomes 0.1—0.3 mm long. Flowers subsessile to pedicellate, pedicels to 3 mm long.
Calyx equally 4-lobed or unequally 5-lobed, 4-5.5 mm long, 4 lobes similar, subulate, equal, 34.5
mm long, 0.5—0.9 mm wide, abaxially pubescent with erect to antrorse eglandular (and sometimes
glandular) trichomes 0.1—0.2 mm long, 5th lobe (if present) 2.5 mm long, 0.1—0.2 mm wide. Corolla
orange to red, 23—33 mm long, externally pubescent with erect to flexuose glandular and eglandular
trichomes 0.1—0.4 mm long, tube gradually expanded distally, 12—18 mm long, 2.54 mm in diameter
near midpoint, upper lip 11-16 mm long, apically entire, lower lip 1 1—14 mm long, lobes |.2-3 mm
long, 1.5—3 mm wide. Stamens inserted near apex of corolla tube, 8-20 mm long, filaments glabrous,
thecae 22.5 mm long, subequal, sagittate, subequally inserted, dorsally glabrous, lacking basal
appendages; pollen 2-aperturate, apertures flanked on each side by 2 rows of insulae (peninsulae
sometimes present as well), exine reticulate. Style 23—31 mm long, sparsely pubescent near base with
antrorse eglandular trichomes, stigma 0.1 mm long, only | lobe evident. Capsule 20 mm long,
externally pubescent with erect (to retrorse) glandular and eglandular trichomes 0.05—0.1 mm long,
stipe 8 mm long, head subellipsoid, 12 mm long. Seeds 4, sublenticular, 3.5 mm long, 3.5 mm wide,
surfaces tuberculate.

PHENOLOGY. — Flowering: January—March and July; fruiting: March.

DISTRIBUTION AND HABITAT. — Western Mexico (Jalisco and Guerrero, Fig. 2); plants occur in
tropical deciduous forests and tropical subdeciduous forests at elevations from 780—1650 m.

PARATYPES. — MEXICO. Jalisco: Sierra de Manantlan Occidental, Arroyo La Calera, ca. 9 km
N de Casimiro Castillo en el camino entre Autlan y la costa, 19°41'N, 104°25'W, E. Judziewicz et al.
5117 (WIS); Mpio. Autlan, Rancho Chiquihuitlan, 6 km O de Autlan, E. Lott and A. Solis M. 443
(CAS). Guerrero: 24 km N de Valle de Zaragoza, E. Martinez S. and J. Soto N. 3710 (CAS).

This species is not readily assignable to any one of the subgeneric taxa proposed by Graham
(1988). Its pollen (Fig. 3) conforms to “Type 7” (Graham 1988), which occurs in numerous sections.
Justicia novogaliciana differs from all of these sections in the structure of its inflorescence and in
various other characters.

The few known collections of Justicia novogaliciana show variation in several features. Martinez
S. and Soto N. 3710 from Guerrero differs from the collections from Jalisco by having both glandular
and eglandular trichomes on the secondary peduncles, pedicels, bracteoles, and calyx. In the speci-
mens from Jalisco these structures have only eglandular trichomes. Lott and Solis M. 443 differs from
the other collections by its more or less evenly pubescent internodes and long-pedunculate (i.e.,
peduncles 5—22 mm long) dichasia that comprise a single flower. In the other collections, the
internodes are generally bifariously pubescent and the dichasia are sessile or short-pedunculate (1.e.,
peduncles to 2.5 mm long) with five or more flowers.

This species is named for the old Spanish province of Nueva Galicia, from which the type and
most other collections were taken.

A NEW NAME
Justicia isthmensis T. F. Daniel, nom. nov.

Justicia panamensis (Lindau) V. A. W. Graham, Kew Bull. 43:605. 1988; non J. panamensis Durkee (as
“panamense’’) (1978).

DANIEL: NOTES ON NEOTROPICAL JUSTICIA 49]

Chaetochlamys panamensis Lindau, Repert. Spec. Nov. Regni Veg. 11:124. 1912. Type. — PANAMA. “Ad
solum araneosum ad rivos et ad umbrosas vias prope Alhaguela et Las Cruces 30—100 m” (from protologue),
Jan—Feb 1911, Pittier 2338 (syntype: B?, destroyed), Pittier 2614 (syntype: B?, destroyed). [Graham (1988)
noted that Pittier s.n. from the Chagres Valley at BM was an isosyntype; additional isosyntypes are likely
at US].

Chaetochlamys Lindau was included within Justicia by Graham (1988). The combination J.
panamensis (Lindau) V. A. W. Graham (Graham 1988) is a later homonym of J. panamensis Durkee
(Durkee 1978). Thus, a new name is needed in Justicia for the former species. The epithet selected
refers to the Isthmus of Panama.

A RANGE EXTENSION
Justicia candelariae (Oerst.) Leonard

PANAMA. Veraguas: region de la Yeguada, Cerro Cristal, 1360 m, 19 February 1987, M.
Correa et al. 4842 (PMA).

This species is reported to occur in Mexico, Guatemala, Belize, Honduras, and Costa Rica (Daniel
1995b). The collection noted above represents the first known occurrence of the species in Panama.
Twenty-five species of Justicia are now known from Panama (Daniel and Wasshausen 1990; Daniel
and McDade 1995), making it the largest genus of Acanthaceae in that country. Justicia candelariae
can be distinguished from other species of the genus in Panama by the following combination of
characters: petioles glandular abaxially, inflorescence of terminal and subterminal pedunculate spikes,
bracts ovate to elliptic to obovate and 34 mm wide, calyx with 4 similar lobes and a reduced posterior
lobe, corollas whitish marked with maroon and 7—10.5 mm long, thecae dorsally pubescent and
lacking basal appendages, and capsules S—6 mm long and externally pubescent. Intraspecific variation
and interspecific relationships of this species were discussed by Daniel (1995b). Leonard’s (1958)
description of J. chlorostachva from Colombia does not seem to differ significantly from that of /.
candelariae from Chiapas, Mexico (Daniel 1995b). However, I have not studied specimens attributed
to the former species.

ACKNOWLEDGMENTS

I thank Tom Wendt, Emily Lott, and Ted Cochrane for sending me collections of Acanthaceae;
Mireya Correa and William Trauba for logistical assistance in the field; Darrell Ubick for assisting
with the scanning electron microscope; and Jenny Speckels for illustrating one of the species.

LITERATURE CITED

DANIEL, T.F. 1990. New and reconsidered Mexican Acanthaceae. IV. Proc. California Acad. Sci.
46(12):279-287.

. 1993a. Mexican Acanthaceae: diversity and distribution. Pp. 541-558 in Biological Diversity of

Mexico: Origins and Distribution, T. P. Ramamoorthy et al., eds. Oxford University Press, New York.

. 1993b. New and reconsidered Mexican Acanthaceae. V. Contr. Univ. Michigan Herb. 19:271—291.

. 1995a. New and reconsidered Mexican Acanthaceae. VI. Chiapas. Proc. California Acad. Sci.

48(13):253-284.

. 1995b. Acanthaceae. Pp. 1—158 in Flora of Chiapas, Pt. 4, D. E. Breedlove. ed. California Academy of

Sciences, San Francisco.

492 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES
Volume 51, No. 15

. 1998. Pollen morphology of Mexican Acanthaceae: diversity and systematic significance. Proc.
California Acad. Sci. 50(8):217—256.

DANIEL, T. F. AND L. A. MCDADE. 1995. Additions to the Acanthaceae of Panama. Ann. Missouri Bot. Gard.
82:542-548.

DANIEL, T. F. AND D. C. WASSHAUSEN. 1990. Three new species of Justicia (Acanthaceae) from Panama. Proc.
California Acad. Sci. 46(13):289-297.

DuRKEE, L. H. 1978. Family 177. Acanthaceae. Pp. 155—284 in Flora of Panama, R. E. Woodson et al., eds.
Ann. Missouri Bot. Gard. 65.

DurKEE, L. H. AND L. A. MCDADE. 1996. Three new species of Justicia (Acanthaceae) from Costa Rica. Novon
6:13-21.

GRAHAM, V. A. W. 1988. Delimitation and infra-generic classification of Justicia (Acanthaceae). Kew Bull.
43:551-624.

LEONARD, E. C. 1958. The Acanthaceae of Colombia, III. Contr. U. S. Natl. Herb. 31:323—781.

WASSHAUSEN, D.C. AND T. F. DANIEL. 1995. Justicia nevlingii (Acanthaceae), a new species from Mexico.
Novon 5:114—117.

© CALIFORNIA ACADEMY OF SCIENCES, 1999
Golden Gate Park
San Francisco, California 94118

INDEX TO VOLUME 51

(Compiled by Katie Martin and Hillary Culhane)

Acanthurus reversus
Caelorinchus amydrozosterus
Caelorinchus charius
Caelorinchus gaesorhynchus
Caelorinchus goobala
Caelorinchus lasti
Caelorinchus mayiae
Caelorinchus pardus
Caelorinchus thurla
Ceratoglanis pachynema
Chromodoris mandapamensis
Chromodoris naiki
Dibranchus accinctus
Dibranchus tremendus
Dibranchus velutinus
Gadomus pepperi
Gargamella bovina
Gargamella gravastella
Halgerda bacalusia
Halgerda dalanghita
Halgerda diaphana
Halgerda dichromis
Halgerda stricklandi

A
Acanthaceae 483
Acanthicini 3
Acanthicus 3
Acanthoptilon 29
Acanthoptilum 25, 29, 33
agassizi 33
album 33
annulatum 33
gracile 33
oligacis 33
pourtalesii 33
scalpelifolium 33
Acanthurus
achilles 474
glaucopareius 474
nigricans 474

New Taxa

Halgerda toliara

Justicia wendtii

Justicia chimalapensis
Justicia novogaliciana
Nezumia kapala

Nezumia leucoura
Nezumia merretti
Nezumia soela

Nezumia wularnia
Paraminabea
Paraminabea arborea
Platymantis cagayanensis
Platymantis indeprensus
Platymantis pseudodorsalis
Platymantis taylori
Solocisquama
Solocisquama carinata
Sperata acicularis
Thamnophis atratus zaxanthus
Trachonurus yiwardaus
Ventrifossa gomoni
Ventrifossa paxtoni
Ventrifossa sazonovi

General Index to Taxa

olivaceus 473, 474, 478
reversus 475, 478
tennenti 478
Acestra 3
Acestridiinae 3
Acestridium 3
Acestrini 3
Acrochordonichthyini 3
Acrochordonichthys 3
Acrophytum 337, 362
Actinoptilon 29
Actinoptilum 25, 29, 33
molle 33
Actinoptinum 29
Ageneiosi 3
Ageneiosus 3
Ailia 3

[493]

494

Ailichthyoidei 3
Akyses 3
Akysidae 3, 8
Akysis 3

Alcyontidae 337, 338, 362
Alcyonium indicum 338, 357, 361

Amblycepinae 4
Amblyceps 4
Amblycipitidae 4
Ameiurus 4
Amiurina 4
Amiurus 4
Amphiacme 25, 29, 33
abysorrum 33
Amphianthus 29
Amphiliidae 4, 6
Amphiliinae 4
Amphilius 4
Anchariidae 4
Ancharius 4
Ancistri 4
Ancistrus 4
Andinichthyidae 4
Andinichthys 4
Aneides lugubris 327
Anesipoma 4
Anodontes 4
Anthomastus 337, 362
agilis 344
phalloides 339, 362
Anthoptilum 25, 29, 33
grandiflorum 33
murrayi 33
Aoria 400, 401
Aorichthys 398, 400, 401
aor sarwari 420
Aphelodoris 366, 425
Argeini 4
Argentella 29
Arges 4
Ari 4
Ariidae 4-7, 10
Ariobagri 4
Ariodontes 4
Arius 4
Aspidoradidi 4
Aspidoras 4
Aspredinae 4
Aspredinidae 4, 6, 8
Aspredo 4
Astephinae 4
Astephus 4

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 51

Asteronotidae 366
Asteronotus 366, 425, 438, 439, 441-443
Asterophysus 4
Asthenomacrurus victoris 114
Astroblepidae 4, 5
Astroblepiformes 4
Astroblepus 4

Astrophysi 4
Auchenipteridae 3—6, 8, 10
Auchenipterinae 4
Auchenipterini 4
Auchenipterus 4
Auchenoglanidinae 4
Auchenoglanis 4
Austroglanididae 5
Austroglanis 5

B

Bagarina 5
Bagarius 5
Bagre 5
Bagreidae 5
Bagrichthyes 5
Bagrichthyoidei 3, 5
Bagrichthys 3, 5
Bagridae 5, 7-9
Bagroides 5
Bagroidinae 5
Bagrus 3,5
aorellus 397, 415, 416
aorides 397, 412, 413
aorinus 397, 420
lamarrii 397, 398, 400, 401, 420, 421
meridionalis 402
Balticina 29
Baptodoris 253-255, 257
cinnabarina 253-255
fongosa 257
mimetica 253-255
Batasinae 5
Batasio 5
Bathyalcyon 362
Bathygadidae 107, 115
Bathygadus 115
cottoides 116, 117
spongiceps 115, 116
Bathypenna 29
Bathyptilum 29
Batrachocephalinae 5
Batrachocephalus 5
Batrachoseps attenuatus 327
Bellonella indica 338, 345, 361, 362

INDEX

Beloperone 483
Bensonularia 33
spatulata 33
Benthoptillum 29
Benthoptilum 29

Betonica 486
Bunocephalini 5
Bunocephalus 5

Cc
Caelorinchus 108, 110, 113, 122, 127, 144,

147, 148, 150, 160, 166

acanthiger 123, 125, 145, 166, 167

acutirostris 122,125,127

amydrozosterus 122, 127, 129, 152

argentatus 123, 129, 130, 132, 137, 154,
1552 161

argus 166

asteroides 140

bollonsi 129

charius 123, 132, 136, 164

commutabilis 136

cookianus 129

denticulatus 132, 155

fasciatus 128, 129, 152, 157

formosanus 132

gaesorhynchus 123, 136, 137, 161

goobala 123, 138, 140

innotabilis 123, 140, 141

kaivomaru 141

kamoharai 132, 137, 154

kamohari 130

kermadecus 159

lasti 123, 125, 141, 144, 145

longissimus 132

macrorhynchus 123, 145-147, 164

maculatus 123, 147, 148

matamua 122, 148

maurofasciatus 122, 128, 129, 150, 152

mayiae 123, 130, 132, 137, 152, 154, 155,
161

mirus 122,128. 155, 157

multispinulosus 132, 155, 161

mycterismus 112, 125, 145, 157—159

mystax 129

pardus. 123,130; 132, 137, 154.1554,159:
161

parvifasciatus 122, 128, 129, 152

radcliffei 140

smithi 123, 136, 147, 161, 164

spinifer 147

thurla 123, 164, 166

trachycarus 123, 125, 166, 167
Calibelemnon 31, 33

hertwigi 35

indicum 35

symmetricum 35
Callichthini 5
Callichthyidae 4—6
Callichthys 5
Callophysinae 5
Callophysus 5
Calophysinae 5
Calophysus 5
Carotalcyon 362
Cascadura 5
Cascaduridi 5
Cavernularia 25,31, 33, 35

capitata 35

chuni 35

clavata 35

dayi 35

dedeckeri 35

elegans 35

glans 35

habereri 35

lutkeni 35

luetkeni 35

lutkeni 35

malabarica 35

mirifica 35

obesa 35

pusilla 35
Cavernulina 25, 35

cylindrica 35

darwini 35

grandiflora 35

orientalis 35
Centrochir 5
Centrochirinae 5
Centromochli 5
Centromochlus 5
Ceratiidae 263
Ceratioidei 306
Ceratoglanis 385-388, 390

pachynema 385, 387, 389, 390, 393

scleronema 385, 387, 389, 390, 393
Cetonurichthys 167

subinflatus 114, 167
Cetonurus 167, 169

crassiceps 169, 170

globiceps 114, 169, 170
Cetopsidae 5, 6
Cetopsini 5

495

496 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 51

Cetopsis 5

Chaca 5

Chacidae 5

Chacini 5

Chaetochlamys 483, 491

Chaetostomidi 5

Chaetostomus 5

Chaetothylax 483

Chalinura 170
yaquinae 170

Chaunacidae 306

Chiloglanidinae 5

Chiloglanis 5

Chromodorididae 461

Chromodoris 461, 465, 466, 468
kitae 470, 471
lineolata 468
mandapamensis 466, 468
naiki 468, 470, 471
preciosa 465
sinensis 465, 466
striatella 468
trimarginata 462, 465, 466

Chrysichthyinae 5

Chrysichthys 5

Chunella 25, 35
gracillima 35

Cladiella 362

Cladiscus 29

Clarias 5

Clariidae 5, 6

Clariini 5

Claroteidae 4—6

Claroteini 5

Clarotes 5

Clavella 31

Clypeati 5

Coelophrys 260, 297, 305, 306
brevicaudata 305

Coluber 321

Conta 5

Continae 5

Copionodon 5

Copionodontinae 5

Corydoradinae 5

Corydoras 5

Coryphaenoides 108, 113, 114, 170
armatus 170
filicaudus 170
macrocephalus 171
memillani 170, 173
murrayi 170

paradoxus 171
rudis 170-171
serrulatus 171, 173
striaturus 173, 174
sp. 174
Cranoglanidae 5
Cranoglanididae 5
Cranoglanis 5
Crassophyllum 35
cristatum 35
thessalonicae 35
Crinillum 31
Crispella 31
Cyclopidae 5
Cyclopium 5

D
Deutocaulon 31
Diadophis amabilis 327
Dianema 5
Dianemidi 5
Diaulula 254—255
sandiegensis 254, 255
Dibranchichthys 260
nudivomer 260
Dibranchopsis 260
spongiosa 260
Dibranchus 259-264, 266, 267, 270, 275,
279-281, 284, 293-295, 297, 298,
305-307
accinctus 263, 270, 279, 295, 307
alberti 260
asper 281
atlanticus 259, 262—264, 266, 267, 270,
NEDA SOAS PZ CUT)
cracens 263, 284
discors 263, 266, 292, 294, 307
erinacea 260
erinaceus 263, 290, 307
erythrinus 259
hystrix 262, 263, 270, 280, 281, 307
infranudus 261
japonicus 259, 263, 270, 294, 307
micropus 260
nasutus 260
nudiventer 261
nudivomer 263, 266, 292
obscurus 261
scaber 262, 281
simulus 260
sparsa 260, 307
sparsus 263, 287, 307

INDEX

spinosa 260

spinosus 260, 262-263, 266, 289, 307

spinulosa 260, 289

spongiosus 262, 263, 266, 280, 281
stelliferus 260

stellulatus 259, 297

tremendus 259, 263, 266, 270, 275, 307

velutinus 262, 263, 285, 287
Dicamptodon 327
Diplomystes 6
Diplomystidae 6
Diptéronotes 6
Distichoptilum 25, 31, 35

gracile 35
Doiichthyidae 6
Doiichthys 6
Dorididae 5—7, 366
Doradini 6
Doras 6
Doris humberti 468
Doumea 6
Doumeinae 6
Drejerella 486
Dibenia 31

E
Echinoptilum 25, 35
asperum 35
echinatum 35
elongatum 35
macintoshi 35
minimum 35
roseum 35
Eleutherobia 352
somaliensis 352
Encrinus 31
Ensatina 327, 328
eschscholtzii 327
Eremophilini 6
Eremophilus 6
Erethistes 6
Erethistidae 5—7
Erethistides 6
Euanemini 6
Euanemus 4,6
Eutainia
atrata 333
infernalis vidua 333
Exostoma 6
Exostomatina 6

F

Farlowella 6

Farlowellidi 6

Funiculina 25, 31, 33, 35, 38
armata 38
parkeri 38
quadrangularis 38

Fusticularia 31

G
Gadomus 108, 113, 117, 119, 121
aoteanus 117, 119, 121
capensis 119, 121
denticulatus 119
colletti 119
introniger 119
magnifilis 119
melanopterus 119, 121, 122
multifilis 119, 121
pepperi 117, 119, 121, 122
sp: ch. collettr N17; AV95 ID?
Gargamella 245, 252-257
bovina 249, 250, 252-256
gravastella 246, 248-250, 252-256
immaculata 245, 248-250, 252-256
latior 245, 252
novozealandica 245
Gephyroglanidini 6
Gephyroglanis 6
Glanapteryginae 6
Glanapteryx 6
Glani 6
Glossodoris 468
trimarginata 462
Glyptosterni 6
Glyptosternum 6
Glyptothoracini 6
Glyptothorax 6
Godeffroyia 31
Godefroyia 31
Gondul 31
Goniodontes 6
Graphularia 22, 25, 38
ambigua 38
badenia 38
belgica 38
crecelii 38
desertorum 38
groenwalli 38
incerta 38
irregularis 38
kalimnae 38

497

498 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 51

longissima 38
meijeri 38

nigra 38
quadrata 38
salisburgensis 38
sulcata 38
wetherelli 38

africana 38
californica 38
christii 38
finmarchica 38
heptazooidea 38
willemoesi 38
Halisceptrum 31

Haplomacrourus nudirostris 114
Harttia 6

Harttiinae 6

Helicoptilum 31

Helogenes 6

Helogenidae 6

H Hemidoradinae 6

Halgerda 365, 366, 371-374, 376, 378, 379, Hemidoras 6

382, 425, 426, 428, 432-434, 438, 439, Hemiodontichthyina 6
441-443, 446, 447 Hemiodontichthys 6

yamakawai 38
Gunneria 31
Gyrophyllum 25, 29, 38

hirondellei 38

sibogae 38

albocristata 382, 438, 439, 441, 442 Hemipimelodinae 6
Hemipimelodus 6

Hemisilurus 387

aurantiomaculata 432, 433, 438, 441, 442

bacalusia 430, 432, 433, 438, 441
brunneomaculata 439, 441, 442

carlsoni 428-430, 432, 433, 438, 441, 442

dalanghita 373, 374, 381, 382, 438, 439,
441, 442

diaphana 433, 438, 442

dichromis 374, 376, 378, 379, 381, 382,
439, 441, 442

elegans 374, 438, 439, 441, 442

formosa 365, 366, 369, 371-373, 378, 379,

382, 426, 438, 439, 441, 442
graphica 441, 442
guahan 432, 433, 438, 442
iota 439, 441, 442
malesso 428-430, 433
punctata 365, 371-373, 439, 441, 442
stricklandi 426, 428-430, 433, 438, 441
terramtuentis 439
tessellata 438, 439, 441, 442
toliara 380-382, 439, 441
wasinensis 382, 441, 442
willeyi 438, 439, 441, 442
xishaensis 439, 441, 442
sp.2 439
Halgerdidae 366, 425, 426
Halicmetus 297, 298, 305—307
Halieutaea 260, 305

mekongensis 387
moolenburghi 387

scleronema 386
Heptapterinae 6
Heptapterus 6
Herklotsia 31
Heterobranchia 6

Heterobranchoidei 6

Heterobranchus 6
Heteropneustes 6

Heteropneustidae 6, 9

Heteropolypus 362
Hopliancistrini 6 -
Hopliancistrus 6
Hoplomizoninae 6
Hoplomyzon 6
Hoplosterninae 6
Hoplosternum 6

Hoplostethus atlanticus 224

Hymenocephalus 108, 112, 114, 175, 179, 219

adelscotti 176
kuronumai 176
lethonemus 178
longibarbis 177
longiceps 177
megalops 180

Halieutichthys 260, 305 nascens 177, 180

Halieutopsis 260-262, 264, 297, 305, 306 striatissimus aeger 178-180
micropa 306 striatulus 178, 180

simulus 261 Spay S0 Say
vermicularis 261 Hymenosomatidae 390

Halipteris 25, 29, 31, 33, 38 Hyomacrurus 189

INDEX

Hypophthalmini 6
Hypophthalmus 4, 6
Hypoptopoma 6
Hypoptopominae 6
Hypostomiden 3, 7
Hypostomus 7
Hypsidoridae 7
Hypsidoris 7

I

Ichthaelurus 7

Ichthyaelurinae 7

Ichthyaelurus 7

Ictalurni 7

Ictaluridae 4, 7

Ictalurus 7

Idiolophorhynchus 234, 235
andriashevi 113, 235, 236

Inermes 7

Ixtlania 483

J

Jacobinia 483

Jorunna pantherina 245

Juncoptilum 31

Justicia 483, 488, 491
alainii 486
candelariae 483, 491
chimalapensis 486, 488
chlorostachya 491
chol 488
costaricana 488
isthmensis 490
nevlingii 488
novogaliciana 488, 490
panamensis 491
wendtii 483, 486

K

Kophobelemnon 25, 29, 31, 38
affine 38
heterospinosum 38
hispidum 38
irregulatus 38
leucharti 38
macrospinosum 38
molanderi 38
pauciflorum 39
stelliferum 39

Kryptopterini 7

Kryptopterus 7

499

Kumba 192

Kuronezumia 108, 114, 180, 192
leonis 181, 182
pallida 181, 182

L
Laguvia 7
Laguviini 7
Lampropeltis zonata 328
Leioptillum 31
Leioptilum 31
Leioptilus 31
Lepidorhynchus 182
denticulatus 114, 182, 184
Leptoptilum 31
Lictores 7
Lionurus 170
Lioptilum 31
Lithodoradinae 7
Lithodoras 7
Lithogeneinae 7
Lithogenes 7
Lithoxina 7
Lithoxus 7
Lituaria 25, 31, 39
amoyensis 39
australasiae 39
breve 39
habereri 39
hicksoni 39
kuekenthali 39
molle 39
phalloides 39
philippinesis 39
valenciennesi 39
Lobophytum 362
Lophiidae 263
Lophiodes 263
Lophiomus 264
Lophius 263
Loricaria 2,7
Loricariichthyina 7
Loricariichthys 7
Loricariidae 3—9
Lucigadus 184, 210
acrolophus 185
microlepis 185
nigromaculatus 184—186
nigromarginatus 184, 185
ori 114, 184, 185
Luciopimelodinae 7
Luciopimelodus 7

500 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 51

Lygomorpha 31
Lygus 31

M
Macrones 397, 400, 401
aor 408, 415
seenghala 415
Macronichthys 400, 401
Macropterotere 6
Macrouridae 107
Macrourinae 122
Macrouroides 234
inflaticeps 113
Macrouroidinae 234
Macruronus novaezelandiae 184
Malacacanthus 337, 362
Malacobelemnon 39
stephensoni 39
Malacocephalus 186
boretzi 187
hawaiiensis 187
laevis 114, 186, 187
luzonensis 187
nipponensis 187
occidentalis 187
okamurai 187
Malapteruridae 7
Malapterurini 7
Malapterurus 7
Malthe 260
Malthopsis 260, 305, 306
spinulosa 290
Masticophis 321
lataeralis euryxanthus 323

Mataeocephalus 108, 113, 187—189

acipenserinus 114, 188
sp. 189
Mesobelemnon 31
Mesobius 108, 114, 189, 190, 192
antipodum 189-192
berryi 190, 191
Metaloricaria 7
Metaloricariina 7
Microptilum 31

Minabea 337-339, 344, 358, 361, 362

acronocephala 338
aldersladei 338
ozakii 337, 344, 362
phalloides 337-339, 344, 362
Mochokidae 5, 7, 8-10
Mochokus 7
Mystidae 7

Mystus 7, 398, 401
aor 401
seenghala 401

N
Nangra 7
Nangrina 7
Nematogenyidae 7
Nematogenyini 7
Nematogenys 7
Nematognathi 7, 9
Neoceratiidae 263
Neohallia 483
Neoplecostominae 7
Neoplecostomus 7
Neorhynchoplax prima 390
Nezumia 108, 114, 192, 197, 208
burragei 199
coheni 192, 195, 202
condylura 201
evides 197, 201
holocentra 199
kapala 193-195, 202, 208
leucoura 195, 197
merretti 197, 199, 204
namatahi 192, 195, 202
propinqua 197, 199, 201
soela 195, 201, 202
spinosa 199, 202, 204
wularnia 205, 208
Nidalia indica 357
Norticina 31

O

Octobasis 39
Odontomacrurus murrayi 114
Ogcocephalidae 305, 306
Ogcocephalus 260, 261, 305
Olyra 7

Olyrinae 7

Ombellula 31
Ombellulaires 31
Ombellularia 31

Ompok 387

Oplophoria 7

Osteobagrus 398, 401
Osteocella 31
Osteogeneiosus 7
Osteogeniosinae 7
Otocinclini 7

Otocinclus 7

Otothyrini 7

INDEX

Otothyris 7

P
Pangasianodon 7
Pangasianodonidi 7
Pangasiidae 7
Pangasini 7
Pangasius 7
Parabelemnon 31
Paracetonurus 208
Parakysidae 8
Parakysis 8
Paraminabea 338, 344, 352, 358, 361, 362
acronocephala 345, 358, 361, 362
aldersladei 361, 362
arborea 352, 357, 362
cosmarioides 361
goslineri 361, 362
indica 352, 357, 361
kosiensis 361
robusta 362
sp. 361
Pareiodon 8
Pareiodontinae 8
Pavonaires 31
Pavonaria 31, 39
portisi 39
Pelteobagrini 8
Pelteobagrus 8
Penna 31
Pennatula 25,31, 39
aculeata 39
argentina 39
delicata 39
fimbriata 39
grandis 39
indica 39
inflata 39
mollis 39
phosphorea 39
prolifera 39
rubra 39
Pennatulites 39
manzonil 39
Phalacronotini 8
Phalacronotus 8
Phosphorella 31
Phreatobinae 8
Phreatobius 8
Pimelodidae 4—9
Pimelodinae 8
Pimelodini &

Pimelodus 8
aor 397, 398, 400
seenghala 420
Pinirampidae 8
Pinirampus 8
Pinus sabiniana 328
Plagiacanthus 486
Planiloricaria 8
Planiloricariina 8
Platydoris 253 —257
angustipes 254-256
argo 253-256
formosa 253
maculata 254-256
stomascuta 254-256
Platymantis 449
cagayanensis 450, 454, 460
corrugatus 449, 457, 460
dorsalis 449, 451, 455, 457, 460
indeprensus 455, 457, 460
levigatus 449, 457, 460
mimulus 449, 457, 460
naomi 449, 457, 460
pseudodorsalis 454, 457, 460
pygmaeus 449, 457, 460
spelaeus 449, 457, 460
taylori 451, 457, 460
Platystacinae 8
Platystacus 8
Platystoma seenghala 397, 419
Plecostomiformes 8
Plecostomus 8
Plotose 6
Plotosichthyoidei 3, 8
Plotosidae 8
Plotosus 3,8
Policella 31
Porcinae 8
Porcus 8
Prochunella 31
Prographularia 39
tradica 39
Protocaulon 31
Protoptilum 25, 39
carpenteri 39
celebense 39
cyaneum 39
denticulatum 39
smitti 39
thomsoni 40
Pseudacanthicina 8
Pseudacanthicini 8

502

Pseudacanthicus 8
Pseudauchenipterini 8
Pseudauchenipterus 8
Pseudecheneidina 8
Pseudecheneis 8
Pseudonezumia 208

cetonuropsis 208, 209

flagellicauda 208
japonicus 208
parvipes 208, 209

pusilla 114, 208, 209

Pseudopimelodidae 8
Pseudopimelodinae 8
Pseudopimelodus 8
Pterocryptis 387

Pteroeides 25,29, 31, 33, 40

acutum 40
argenteum 40
bankanense 40
bestae 40
breviradiatum 40
caledonicum 40
carnosum 40
crossieri 40
densum 40
dofleini 40
duebeni 40
durum 40
esperi 40
flexuosum 40
humesi 40
hymenocaulon 40
isosceles 40
Jungerseni 40
laboutei 40
latissimum 40
lusitanicum 40
malayense 40
morbusus 40
oblongum 40
sagamiense 40
sparmanni 40
spicatum 40
spinosum 40
tenerum 40
timorense 40
triangulum 40
Pteromorpha 31
Pteronotidae 8
Pteronotus 8
Ptilella 31
Ptilosarcus 25,31, 40

PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 51

gurneyi 40
Pygidiidae 8
Pygidium 8

R
Rana aurora 327
Reganella 8
Reganellina 8
Renila 31
Renilla 25,31, 40
kollikeri 40
koellikeri 40
kollikeri 40
mulleri 40
muelleri 40
mulleri 40
musaica 40
octodentata 40
reniformis 40
Renillina 31
Rhamdia 8
Rhamdiae 8&
Rhaphidospora 486
Rhinoglanina 8
Rhinoglanis 8
Ricola 9
Ricolina 9
Rineloricaria 9
Rineloricariina 9
Rita 9
Ritae 9

S
Saccobranchini 9
Saccobranchus 9
Sarcobelemnon 31
Sarcoglanidinae 9
Sarcoglanis 9
Sarcophyllum 33
Sarcophyton 31, 362
Sarcophytum 362
Sarcoptilus 25, 33, 41
bollonsi 41
grandis 41
nullispiculatus 41
rigidus 4]
shaneparkerii 41
tentaculata 41
Sarotheca 486
Sceptonidium 33
Schilbe 9
Schilbeini 9

INDEX

Schilbidae 3, 9
Scleracanthi 9
Sclerobelemnon 25, 31, 41
burgeri 41
elongatum 4]
gracile 41
gravieri 41
koellikeri 41
magniflorum 41
schmeltzi 41
theseus 41
Sclerodoris 366, 374, 425, 439
tuberculata 374
Scleroptilum 25, 41
grandiflorum 4]
Scoloplacidae 9
Scoloplacinae 9
Scoloplax 9
Scytaliopsis 25, 41
djiboutiensis 41
ghardagensis 41
ghardagana 41
Scytalium 25, 41
balssi 41
martensi 41
sarsii 41
tentaculatum 41
Siluranodon 9
Siluranodontinae 9
Silurichthyoidei 3, 9
Silurichthys 3,9, 387
Siluridae 7— 9, 385
Anomalopterae 9
Branchicolae 9
Heteropterae 9
Homalopterae 9
Opisthopterae 9
Proteropodes 9
Proteropterae 9
Stenobranchiae 9
Siluroides 3, 9
Silurus 3,9, 387
Simuldentinae 9
Sinularia 344, 362
Siphonoglossa 483
SISOP S550 O
Sisorichthyoidei 3,9
Sisoridae 5—8

Solocisquama 259, 297, 298, 307

carinata 297, 298, 301, 304
erythrina 297, 299, 301, 304
stellulata 297, 298, 301, 304

Sorubim 9
Sorubinae 9
Sorubium 9
Spectracanthicina 9
Spectracanthicus 9

Sperata 398, 400-403, 413, 416, 420, 421

503

acicularis 400, 403, 408, 410, 413, 417, 421

aor 400, 401, 403, 408, 410, 412, 413,

415-417, 420, 421

aorella 400, 403, 410, 413, 415-417

sarwari 421

seenghala 403, 408, 410, 412, 413,

415-417, 420, 421

vittata 401
Sphagemacrurus 210, 212

grenadae 212

hirundo 212

pumiliceps 114, 210-212

richardi 210, 212
Spicomacrurus 175, 176
Squalogadus 234

modificatus 113, 234
Stachyptilum 25, 41

dofleini 41

macleari 41

superbum 41
Stegophilina 10
Stegophilus 10
Stephanoptilum 33
Stichoptilum 33
Struthiopteron 33
Stylatula 22, 25, 31, 41

antillarum 41

brasiliensis 41

darwini 41

diadema 41

elegans 41

elongata 46

gracilis 46

kinbergi 46

lacazi 46

polyzoidea 46
Stylobelemnoides 33
Stylobelemnon 33
Svava 33
Svavopsis 33
Synodontini 10
Synodontis 10

T
Tabascina 483
Tachysurinae 10

504 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 51

Tachysurus 10 lindahli 46
Thamnophis 311, 325 monocephalus 46
agiatms S01, 312, 314, 320, 321, 324, 325; pellucida 46
327, 328, 332, 334 spicata 46
atratus aquaticus 311, 312, 314, 321, 323, thomsoni 46
324, 332-334 Umbellularia 33
atratus atratus 311, 312,314, 315,320,321, | Umbellulidae 21
323-325, 329, 330, 332-334 Uroctonus mordax 328

atratus hydrophilus 311, 312, 314, 315, 320,

321, 323-325, 330, 332, 334 V
atratus zaxanthus 328, 330, 332-334 Vandellia 10
couch 311, 320, 321, 327 Vandelliini 10
elegans 311, 312, 315, 320, 321, 327 Ventrifossa 108, 112-114, 219, 224
elegans aquaticus 333 atherodon 233
elegans nigrescens 321 ctenomelas 228
elegans terrestris 315, 320, 327, 334 fusca 223
gigas 311 gomoni 220, 221, 230
hammondii 311, 327, 328, johnboborum 219, 221, 223
ordinoides 311 longibarbata 221, 228
sirtalis 330 macrodon 233
sirtalis fitchi 323, 328 macropogon 112, 219, 221, 223, 224, 230
sirtalis pickeringii 321 macroptera 233
Thesioides 33 misakia 223
Thuridilla 447 nigrodorsalis 219, 221, 224, 228
Torpedininae 10 paxtoni 221, 228, 230
Torpedo 10, petersoni 228
Trachelyopterini 10 rhipidodorsalis 221, 228
Trachelyopterus 10 saikaiensis 221, 230
Trachonurus 108, 112, 114, 212, 218 sazonovi 219, 221, 229, 231, 233
gagates 212,213,218, 219 Veretillum 25, 31, 46
robinst 212 australis 46
sentipellis 112, 212, 213, 216, 218 cynomorium 46
villosus 212, 213 leloupi 46

yiwardaus 217-219

malayense 46
Trachyrincinae 234

manillensis 46

Trachyrincus 113, 234 tenuis 46
longirostris 234 vanderbilti 46
Trichogenes 10 Verrilia 33
Trichogeninae 10 Verrillia 33
Trichomycteridae 5, 6, 8-10 Verseveldtia 337, 362
Trichomycterini 10 Virgularia 22, 25, 29, 31, 33, 46
Trichomycterus 10 abies 46
Trichoptilum 33 brochi 46
Tridens 10 bromleyi 46
Tridentinae 10 densa 46
galapagensis 46
U glacialis 46
Umbellaria 33 gracillima 46
Umbellula 25, 31, 33, 46 gustaviana 46
durissima 46 halisceptrum 46
encrinus 46 jJuncea 46

huxleyi 46 kophameli 46

INDEX 505

loveni 46 tuberculata 47
mirabilis 46 Vorhisia 10
patachonica 46 Vorhisiidae 10
patagonica 46 Vorticella 33
presbytes 47

reinwardti 47 Zo

rumphi 47 Zalieutes 305

schultzei 47

© CALIFORNIA ACADEMY OF SCIENCES, 2000
Golden Gate Park
San Francisco, California 94118

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