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PROCEEDINGS
OF THE =
California Academy of Sciences
Volume 47
SAN FRANCISCO PUBLISHED BY THE ACADEMY 1990-1992
SCIENTIFIC PUBLICATIONS COMMITTEE Thomas F. Daniel, Scientific Editor Ann Senuta, Managing Editor Terrence M. Gosliner Tomio Iwamoto Thomas Moritz
Wojciech J. Pulawski
(US ISSN 0068-547X)
The California Academy of Sciences Golden Gate Park San Francisco, California 94118
PRINTED IN THE UNITED STATES OF AMERICA BY ALLEN PRESS, INC., LAWRENCE, KANSAS
No. 1.
INO
No. 12.
CONTENTS OF VOLUME 47
GOSLINER, TERRENCE M. AND ALAN M. KuzirRIAN. Two new species of Fla- bellinidae (Opisthobranchia: Aeolidacea) from Baja California. Published 11 September 1990
BAUER, AARON M. AND JENS V. VinpDum. A checklist and key to the herpeto- fauna of New Caledonia, with remarks on biogeography. Published 17 October
Voss, GILBERT L. AND WILLIAM G. PEARcy. Deep-water octopods (Mollusca: Cephalopoda) of the northeastern Pacific. Published 17 October 1990...
SMITH, ALLYN G., WALTER B. MILLER, CARL C. CHRISTENSEN, AND BARRY Rotn. Land Mollusca of Baja California, Mexico. Published 17 October
LIGHTFOOT, DAvip C. AND Davip B. WEISSMAN. Review of the grasshopper genera Dracotettix and Litoscirtus (Orthoptera: Romaleidae), with a discussion of their origins and life histories. Published 7 June 1991.0
TopziA, CAROL A. AND FRANK ALMEDA. A revision of 7ibouchina section Lepidotae (Melastomataceae: Tibouchineae). Published 7 June 1991.00.
IWAMOTO, TOMIO AND YuRI N. SHCHERBACHEV. Macrourid fishes of the sub- genus Chalinura, genus Coryphaenoides, from the Indian Ocean. Published 3 OS Re) tn, ct ln i Ln, Whe rll WPA a it thd ee ek SOR 8
DANIEL, THOMAS F. A revision of Aphelandra (Acanthaceae) in Mexico. Pub- NaS OVC HOR: LD cate ea gel te By 2 ee eagle AR lh de
KoOcIOLEK, JOHN P. AND EUGENE F. STOERMER. New and interesting Gompho- nema (Bacillariophyceae) species from East Africa. Published 10 December TESS al en alee ce te cee 09k WF oe tf ety rece es BI ee, eae SE Det ch dtl
PuLAwskI, WojciEcH J. A revision of the wasp genus Kohliella (Hymenoptera: Sphecidac), Published 10 December [99 ies oy ee ee ee ee
GARTHWAITE, RONALD L. AND RoBIN LAwson. Oniscidea (Isopoda) of the San rancisco Bay Area. Published 21 February 1992". 2
RANDALL, JOHN E. AND JOHN E. MCCoskeEr. Two new damselfishes of the genus Chromis (Perciformes: Pomacentridae) from the South Pacific. Published 21 nc Deviate! 900 ae bie leew Beet} chon al awe rot 4 xox Np 30 BARD.
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235-274
275-288
289-302
303-328
329-337 339-347
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PROCEEDINGS OF THE
CALIFORNIA ACADEMY OF SCIENCES
Vol. 47, No. 1, pp. 1-15, 7 figs., 2 tables.
September 11, 1990
TWO NEW SPECIES OF FLABELLINIDAE (OPISTHOBRANCHIA: AEOLIDACEA) FROM BAJA CALIFORNIA
Terrence M. Gosliner
By
Marine Biological Laboratory LIBRARY |
California Academy of Sciences, Golden Gate Park, San Francisco, CA SEP 17 1990
and Alan M. Kuzirian Marine Biological Laboratory, Woods Hole, MA 02543
ABSTRACT:
Woods Hole, Mass.
Two new species of Flabellina are described from Baja California. Flabellina bertschi sp. nov. is
widely distributed within the Gulf of California from Puerto Penasco to the area north of La Paz. Flabellina marcusorum sp. nov. has been found on the Pacific and Gulf coasts of Baja California, as well as the coast of Brazil. This species was originally recorded from Brazil as Coryphellina rubrolineata O’Donoghue, 1929. However, when compared with Indo-Pacific material of F. rubrolineata, the presently described species is consistently distinct in its internal and external morphology. A preliminary phylogeny of the Flabellinidae supports the maintenance of Flabellina as a single, large, diverse genus.
Received February 2, 1990. Accepted March 14, 1990.
INTRODUCTION
The opisthobranch gastropod fauna of the Pa- cific and Gulf of California coasts of México has been extensively studied for much of the twen- tieth century (e.g., MacFarland 1924; Baker and Hanna 1927; Marcus and Marcus 1967; Keen 1971; Bertsch and Ferreira 1974: Bertsch 1977, 1978a, b, c; Gosliner and Behrens 1985). Al- though approximately 150 different species have been recorded from the region, many new rec- ords and taxa remain to be added to the fauna.
Seven species of the aeolidacean family Fla- bellinidae have been recorded from the Pacific coast of Baja California and the Gulf of Califor- nia. The flabellinid nudibranch Flabellina rubro-
[1]
lineata (O’Donoghue, 1929), originally described from the Red Sea, has subsequently been re- ported from Japan (Baba 1949) and Australia (Willan and Coleman 1984). Marcus and Marcus (1961, 1970) recorded specimens of this species (as Coryphellina O’Donoghue, 1929) from the coast of Brazil and from Sonora, México. Recent investigations within the Gulf of California and from the Pacific coast of Baja California have yielded specimens of Flabellina Voigt, 1834, that are identical in their external morphology to ma- terial described by the Marcuses from México and Brazil. These animals are consistently dif- ferent from Indo-Pacific specimens of F. rubro- lineata (O’Donoghue) and are here described as a new species.
iS)
Other investigations into the opisthobranch fauna of the Gulf of California have yielded spec- imens of a second undescribed species of flabel- linid, which is also described in this paper. This species has been collected from several different localities within the Gulf of California. Its anat- omy is described and compared to other closely allied species.
DESCRIPTIONS
Flabellina bertschi sp. nov. (Figs. 1A, 2, 3)
Flabellina telja Er. Marcus and Ev. Marcus, 1967:223, in part, specimen from Puerto Penasco, collected by Mary Anne Hill, 18 June 1966.
Tyre MarTerRIAL.—Holotype, California Academy of Sci- ences, San Francisco, CASIZ 066760, in tide pool, among hy- droids on Sargassum sp., Puerto Penasco, Sonora, México, 23 July 1975, Hans Bertsch and P. Cook. Five paratypes, CASIZ 066761, same date and locality as holotype. One dissected paratype, CASIZ 066762, same date and locality as holotype. One paratype, CASIZ 066763, collected from hydroids on ver- tical wall, San Diego Reef (25°12’N, 110°42’W), s.e. of Isla San Diego, Gulf of California, México, 13 m depth, 24 July 1985, T. M. Gosliner.
DistRIBUTION. — Flabellina bertschi is known within the Gulf of California from Puerto Penas- co, south to Isla San Diego.
EtyMoLoGy.—This species is named for our friend and colleague, Hans Bertsch, who first not- ed that this appears to be an undescribed species. Hans has contributed considerably to our knowl- edge of the Panamic molluscan fauna.
DESCRIPTION. — External morphology. The liv- ing animals (Fig. 1A) are thin and elongate (8- 12 mm long). The body color is translucent white with an overlaying opaque white pigment on the distal two-thirds of the rhinophores and oral ten- tacles. The opaque white pigment covers most of the dorsal surface of the animal, except for an area of translucence immediately posterior to the rhinophores. Each translucent ceras contains a central red to red-brown digestive diverticulum and terminates in a large, opaque white cnidosac.
The rhinophores are smooth, without orna- mentation, and are up to 2.5 mm in length. The oral tentacles are approximately as long as the rhinophores. The cylindrical cerata are of uni- form diameter throughout most of their length. They (Fig. 2A) are arranged in discrete clusters that are inserted on slightly pedunculate mounds of notal tissue. The anteriormost cluster on either side is most pronounced in its elevation from the
PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 47, No. 1
notum. The left and right anterior branches of the digestive tract each contain three or four rows of cerata with 1-4 cerata per row. The posterior digestive branch contains up to six clusters of cerata, each containing 1-3 rows of cerata. Each row is composed of 1-3 cerata. The anteriormost cluster contains more rows of cerata, with more cerata per row. The density of both rows per cluster and cerata per row decreases posteriorly.
The genital apertures are located on the right side of the body ventral to the first and second or second and third ceratal rows. The pleuro- proctic anus is situated within the interhepatic space, below the notum, immediately anterior to the anteriormost cerata of the right digestive branch. The nephroproct is located immediately anterodorsal to the anus. The anterior border of the foot (Fig. 2B) bears elongate foot corners that are held perpendicular to the longitudinal body axis, when the animal is actively crawling. The foot narrows posteriorly and terminates in an acutely pointed tail.
Buccal cavity. The thin, ovoid jaws (Figs. 2C, 3A) occupy much of the muscular buccal mass. They bear several rows of minute denticles on their elongate masticatory margin. From the an- terior end of the buccal mass, a pair of elongate, highly digitate oral glands extends posteriorly within the laterally elevated tissue of the notum. The posterior limit of the glands is at the ante- riormost cluster of cerata of the posterior diges- tive branches. A pair of elongate strap-like sal- ivary glands joins the posterior end of the buccal mass near its junction with the esophagus. The triseriate radula, also contained within the buccal mass, has a formula of 24-31 x 1.1.1., in five specimens examined. The rachidian teeth (Fig. 3C, D) bear 6-8 elongate denticles on either side of the elongate central cusp. The central cusp is depressed ventrally below the level of the adja- cent denticles. The teeth are broadly curved and deeply incised posteriorly. The posterior limbs of each tooth terminate in a posteriorly extended tubercle. The lateral teeth (Fig. 3B) are broadly triangular and bear an elongate primary denticle. The inner masticatory margin bears 9-11 irreg- ular denticles.
Reproductive system. The arrangement of or- gans (Fig. 2D) is androdiaulic. The narrow preampullary duct expands into the ampulla and divides into the oviduct and vas deferens dis- tally. The oviduct immediately expands into the
GOSLINER AND KUZIRIAN: BAJA CALIFORNIA FLABELLINIDAE 3
Ficure 1. Living animals. A. Flabellina bertschi sp. nov., photograph of holotype. B. Flabellina marcusorum sp. nov., photograph of a paratype specimen from San Diego Reef, Baja California Sur, México.
4 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 47, No. 1
Ficure 2. Flabellina bertschi sp. nov. A. Lateral view of holotype showing arrangement of cerata, a = anus; ga = genital aperture; n = nephroproct; pc = pericardial hump; r = rhino- phore, scale = 1 mm. B. Ventral view of holotype, scale = 1 mm. C. Jaw, scale = 0.2 mm. D. Reproductive system, alb = albumen gland; am = ampulla; me = membrane gland; mu = mucous gland; od = oviduct; pe = penis; pr = prostate; rs = receptaculum seminis, scale = 0.5 mm.
two ovoid lobes of the receptaculum seminis. The oviduct again narrows and enters the al- bumen gland, situated between the larger mem- brane and mucous glands. The vas deferens ex- pands into a simply curved prostatic portion that terminates in the indistinct, conical penial pa- pilla, which is devoid of any armature. The nida- mental and penial apertures are united into a short common genital atrium. A bursa copulatrix is absent in the three specimens examined.
Discussion. — Flabellina bertschi is sympatric with five other species of Flabellinidae: Flabel- lina telja Marcus and Marcus, 1967; F. stohleri Bertsch and Ferreira, 1974; F. iodinea (Cooper, 1863); and F. cynara (Marcus and Marcus, 1967). All possess perfoliate rather than smooth rhino- phores. Flabellina marcusorum has strongly pap- illate rhinophores. None of these species has a white body with red cerata.
Flabellina bertschi is similar to two western Atlantic species in its external morphology. Fla-
bellina dushia (Marcus and Marcus, 1963) has smooth rhinophores, a whitish body, and orange- brown cerata, but it lacks an area of translucence posterior to the rhinophores, which characterizes all specimens of F. bertschiexamined in this study. More significantly, the lateral radular teeth of F. dushia are thinner with a more acutely pointed cusp. Nothing is known about the reproductive anatomy of F. dushia. Flabellina verta (Marcus, 1970) also has smooth rhinophores, a whitish body, and brown digestive gland. However, it has opaque white lines on the oral tentacles that join and pass between the rhinophores, which are absent in F. bertschi. Its radula has only half as many rows of teeth as F. bertschi. Also, the reproductive system of F. verta lacks a receptacu- lum seminis, but has a bursa copulatrix adjacent to the genital atrium. In F. bertschi, there is a bilobed receptaculum seminis, but a bursa cop- ulatrix is absent.
The only other described species that are known to possess a bilobed receptaculum seminis but lack a bursa copulatrix are F. pedata (Montagu, 1815) (see Schmekel and Portmann 1982: fig. 7.50e) and F. albomarginata (Miller, 1971). In F. pedata the body color is purple rather than white. In F. albomarginata the rhinophores are covered with small tubercles, and an opaque white line is present around the margin of the foot.
Flabellina marcusorum sp. nov. (Figs. 1B, 4, 5)
Coryphellina rubrolineata O’ Donoghue, 1929:798; Marcus and Marcus 1961:224, figs. 1-10 (misidentification); Marcus and Marcus 1970:210, fig. 81 (misidentification).
Flabellina telja Marcus and Marcus, 1967:223; Ferreira and Bertsch 1972:414, fig. 1 (misidentification); Kerstitch 1989: 66, fig. 154 (misidentification).
Type MatTerIAL.—Holotype, California Academy of Sci- ences, San Francisco, CASIZ 066151, San Diego Reef (25°12'N, 110°42'W), s.e. of Isla San Diego, Gulf of California, México, 13 m depth, 24 July 1985, T. M. Gosliner. Paratypes, CASIZ 066152, six specimens, same date and locality as holotype. Paratypes, CASIZ 066153, two specimens, one dissected, Los Islotes (24°36'N, 110°24’W), Gulf of California, México, 10 m depth, 20 May 1985, Lynne Dunne.
OTHER MATERIAL. — Fifteen specimens, CASIZ 066154, Sa- yulita (20°52'N, 105°29’'W), Nayarit, México, intertidal, 24 Jan. 1975, Gary McDonald. Three specimens, CASIZ 066155, two dissected, Sao Sebastiao, Brazil, Eveline Marcus. One spec- imen, Arroyo San Carlos, Isla Cedros, Baja California, México, 4 m depth, 31 Dec. 1985, Hans Bertsch.
DIsTRIBUTION. — Flabellina marcusorum has been collected from the Atlantic coast of Brazil
GOSLINER AND KUZIRIAN: BAJA CALIFORNIA FLABELLINIDAE
Ficure 3. Flabellina bertschi sp. nov. Scanning electron micrographs of radula of a specimen from Puerto Pefiasco. A. Masticatory border of jaw, scale = 10 um. B. Rachidian and lateral teeth, scale = 4 um. C, D. Rachidian teeth, scale = 4 um.
6 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 47, No. 1
D
Ficure 4. Flabellina marcusorum sp. nov. A. Lateral view, a = anus; ga = genital aperture; n = nephroproct; pe = pericardial hump; r = rhinophore, scale = 2 mm. B. Ventral view, scale = | mm. C. Rhinophore, scale = 0.5 mm. D. Jaw, scale = 0.2 mm. E. Reproductive system, alb = albumen gland; am = ampulla; be = bursa copulatrix; me = membrane gland; mu = mucous gland; pe = penis; pr = prostate; rs = receptaculum seminis; v = vagina, scale = 1 mm.
GOSLINER AND KUZIRIAN: BAJA CALIFORNIA FLABELLINIDAE 7
(Marcus and Marcus 1961). On the Pacific coast of México, it has been found from Isla Cedros south to Sayulita, Nayarit (present study). With- in the Gulf of California, it is known from San Agustin, Sonora, south to Los Islotes, north of La Paz, Baja California Sur.
EtTyMoLoGy.-— This species is named for Ernst Marcus and Eveline du Bois Reymond Marcus, who first recorded this species (as Coryphellina rubrolineata) from Brazil and the Gulf of Cali- fornia.
DescriPTION. — External morphology. The liv- ing animals (Fig. 1B) are 7-30 mm in length. The general body color is translucent rose pink. The oral tentacles, foot corners, rhinophores are rose basally, with purple pigment in their middle, and opaque white present on the apical portion. The oral tentacles have a wide band of purple, while the cerata bear only a narrow ring between the pink and white pigments. The pigment on the apices of the cerata may be yellowish rather than white, in some specimens. The rhinophores bear more purple pigment on their posterior face, where it covers most of the surface of the pa- pillae. The anterior face of the rhinophores bears less purple pigment. The posterior end of the foot is also purple with an opaque white line or spot on its posterodorsal end. The digestive gland within the cerata is the same rose color as the rest of the body.
The rhinophores (Fig. 4C) are elongate (ap- proximately 4 mm in length) and conical. They each bear approximately 100 long papillae on their posterior face. The papillae are approxi- mately 0.5 mm in length and are arranged in somewhat indistinct rows, covering the middle two-thirds of the rhinophores. The oral tentacles are thin and elongate, tapering to a rounded apex. They are longer than the rhinophores.
The cerata are cylindrical and rounded apically immediately distal to the ovoid cnidosac. Within each ceras is a thin core of digestive gland that fills a variable portion of the diameter of the ceras. The cerata are arranged in distinct groups (Fig. 4A). Only the anterior cluster 1s slightly elevated from the notum. The anterior, precar- diac cluster contains 3—5 rows of cerata on either side of the animal, with 2-4 cerata per row. The postcardiac cerata are arranged in 6-8 groups per side of the body. Generally, each group contains a pair of tightly packed rows with 2-4 cerata per row. Only the posteriormost 1-2 rows are un-
paired and consist of a single row. The gonopore is situated on the right side of the body, ventral to the third and fourth ceratal rows of the pre- cardiac cluster. The pleuroproctic anus is situ- ated at the posterior end of the interhepatic space, just anterior to the first postcardiac ceratal row. The nephroproct is immediately anterodorsal to the anus.
The foot (Fig. 4B) is grooved anteriorly and possesses elongate, tentacular foot corners. Pos- teriorly, it tapers gradually to a narrow tail.
Buccal cavity. The buccal mass is short and muscular. Extending from the anteroventral por- tion of either side of the buccal mass is a large digitate oral gland, which extends into the wid- ened portion of the notum in the region of the precardiac cerata. Within the buccal mass is a pair of large chitinous jaws (Fig. 4D). They are ovoid with an elongate masticatory margin (Fig. 5A). The margin bears several rows of denticles with 20-24 denticles on the outer row. The rad- ular formula is 27-34 x 1.1.1. in six specimens studied. The rachidian teeth (Fig. 5B, C) are sim- ply arched with 5-8 triangular denticles on either side of the elongate central cusp. The central cusp is approximately the same width as the adjacent denticles but is depressed ventrally from their level. The lateral teeth (Fig. 5B, D) are triangular in shape with a broad base extending towards the outer edge. There is a single prominent, acutely pointed apex. On the inner margin of the tooth is a series of 4-12 triangular denticles.
At the posterior limit of the buccal mass, near its junction with the esophagus, is a pair of elon- gate salivary glands that extend posteriorly on the dorsolateral surface of the stomach.
Reproductive system. The arrangement of re- productive organs 1s essentially triaulic (Fig. 4E). The ovotestis 1s a diffuse aggregation of distinct follicles. The female acini are distinct. The nar- row preampullary duct widens into an ampulla consisting of two convolutions. The postampul- lary duct again narrows and passes between the lobes of the albumen and membrane glands. Af- ter a short distance it divides into the oviduct and vas deferens. After a short distance the ovi- duct joins a pair of receptacula seminis. The more proximal receptaculum is the larger of the two. The oviduct again narrows, and a short distance later enters the nidamental glands. At this branching point, a distinct, narrow vaginal duct continues towards the genital apertures. Imme-
PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 47, No. 1
Ficure 5. Flabellina marcusorum sp. nov. Scanning electron micrographs. A. Masticatory border of a specimen from San Diego Reef, Baja California Sur, México. B. Rachidian and lateral teeth of specimen from Sao Sebastiao, Brazil. C. Rachidian tooth of specimen from Sido Sebastiao, Brazil. D. Lateral tooth of specimen from San Diego Reef, Baja California Sur. Scales = 10 um.
GOSLINER AND KUZIRIAN: BAJA CALIFORNIA FLABELLINIDAE 9
diately prior to emptying into its own genital pore, it joins with a large saccate bursa copula- trix. The bursa is thin walled and short stalked. The nidamental glands comprise the bulk of the reproductive system. The mucous gland is by far the largest portion, while the albumen and mem- brane glands are smaller. The nidamental glands empty via their own gonopore, ventral to the vaginal and penial apertures. The vas deferens is initially quite narrow and expands into a smooth prostatic portion, which is slightly con- voluted. The vas deferens enters the short penial sac at its distal end. The penial papilla is tubular and thin with a slightly flared apex. There is no armature associated with the penial papilla.
Discussion. —Flabellina marcusorum was originally described from Brazil (Marcus and Marcus 1961) as Coryphellina rubrolineata O’Donoghue, 1929. Marcus and Marcus (1970) also misidentified specimens from the Gulf of California as C. rubrolineata. Flabellina mar- cusorum has also been misidentified as Flabel- lina telja Marcus and Marcus, 1967 by Ferreira and Bertsch (1972) and Kerstitch (1989). The latter species differs from F. marcusorum in hav- ing perfoliate rather than papillate rhinophores, and in several other major details of its external and internal anatomy.
Flabellina marcusorumand F. rubrolineata dif- fer in several consistent aspects of their color- ation. In Flabellina rubrolineata there is a mid- dorsal and a pair of lateral red or purple lines along the length of the animal (O’Donoghue 1929; Baba 1955, pl. 13, fig. 37; Abe 1964, pl. 30, fig. 107; Willan and Coleman 1984, fig. 133; Tan, Pai, and Hsha 1987, fig. 52). These observations are confirmed by examination of specimens of F. rubrolineata from Aldabra Atoll and Madang, Papua New Guinea, in this study. Occasionally, the red or purple lines may be interrupted as in the case of the specimen illustrated by Willan and Coleman, but are present in all material ob- served. In F. rubrolineata the rhinophores are opaque white basally with apical purple pigment. The body color is a much deeper rose in F. mar- cusorum, and longitudinal lines are never present on the body. The rhinophores are purplish ba- sally with an opaque white apex, the reverse of the colors in F. rubrolineata.
The number of radular rows and denticles on the radular teeth varies considerably in both species (Table 1). However, the shape of the ra-
chidian teeth differs consistently between the two species. In Flabellina marcusorum the indenta- tion of the posterior end of the tooth extends deeply, almost to the basal portion of the central denticles (Marcus and Marcus 1961; Fig. 5B, C). In F. rubrolineata (Fig. 6A—-C), the indentation is rounded and does not extend as far forward, leaving a triangular chitinous area between the indentation and the basal portion of the denti- cles. The lateral teeth of F. rubrolineata (O’Don- oghue 1929:fig. 219d; Fig. 6D) bear a series of striations along the middle of their outer edge, which are absent in all specimens of F. marcu- sorum that have been examined.
In their description of Flabellina marcusorum from Brazil, Marcus and Marcus (1961, as Cory- Dhellina rubrolineata) depicted the reproduc- tive anatomy in detail. Their specimens had a large spherical bursa copulatrix, but no recep- taculum seminis. Our examination of Brazilian specimens provided by Eveline Marcus and from material collected from the Pacific coast of Méx- ico indicates that all specimens do possess a bi- lobed receptaculum seminis near the branching of the oviduct. In addition, the reproductive sys- tem is entirely triaulic in all the material we ex- amined. The only aspect of the reproductive anatomy of F. rubrolineata that has been de- scribed was the presence of an unarmed penis (O’Donoghue 1929). However, Dr. Kikutaro Baba provided dissection notes of a specimen of F. rubrolineata from Japan, and we dissected specimens from Papua New Guinea. Both species possess a triaulic arrangement of organs and two proximally situated receptacula seminis; how- ever, their anatomy differs from that of F. mar- cusorum in two significant regards. The prostate of F. marcusorum is more elongate and convo- luted than in F. rubrolineata. More importantly, the bursa copulatrix is small and sessile in F. rubrolineata, while it is well developed and stalked in F. marcusorum.
GENERIC DIVISION WITHIN THE FLABELLINIDAE
The genus Coryphellina, with C. rubrolineata as the type species, was placed in synonymy with Coryphella (Miller 1971). More recently Gosli- ner and Griffiths (1981) considered Coryphella, including Coryphellina, as a junior synonym of Flabellina. This view has been generally adopted
PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 47, No. 1
10
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GOSLINER AND KUZIRIAN: BAJA CALIFORNIA FLABELLINIDAE
Ficure 6. Flabellina rubrolineata (O’Donoghue, 1929). Scanning electron micrographs of specimen from Madang, Papua New Guinea. A. Dorsal view of half-row of radula. B. Dorsal view of rachidian tooth. C. Ventral view of rachidian tooth. D. Lateral tooth. Scales = 10 um.
12 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 47, No. 1
Taste 2. Morphological diversity in Flabellina.
A] oF é & 23 ees se .»,pisSefacd ¢ § Beebe Re 5 aoe g2 386s 258 Bes co ). © 8 og cece cane Se&sasqge88sas Gs Soe ao 6 fo ens eae hypothetical ancestor em Om ONO MO MORO OOM OMOMO affinis DXA TDi @Oit @ O 1 albomarginata Heese 22) ol 2) Oe Oney al alisonae POA O 1 tlk Ot @®@@ i O babai yy 3s OF | Bank lt i © Ol bertschi li@2tlese2anti @oOl eo browni OldOii@Oil tr Oil @ove capensis Oto@Oil@gri @it OO Lk O cynara Gt ZOOIOODUO?Z? L © dushia Ol O22 O2ZO Oo lt & Ow © engeli 2230 1 21 Ot OO 1 @ falklandica TOOdODi YY DY Oi © funeka 2D | Bila il @O@ it ® @ i © fusca OMI Olt @Oi®*Yeo sl o gracilis li@O@ilk@d@iligiggore iodinea Le?s3il@qr s@oil@®2 in © islandica OWOLODODOODOADO OC lineata Ol @Odil Olt Ol OO i O marcusorum Ip S252 alia?) valle Oh nobilis P9O0OOOMOrhI @GI OG i © ornata 223 2d tt @O 1 © @ 1 Oo pedata lt @Qodaaatit @ @ i pellucida t ik @Pta2 tai Ol 1 Ce poenicia 1 ewe oD 9 2 1 0 @ i i pricei i ot Me yh PAS Oh Oy i th o@ rubrolineata 1 2 i 22 Oi YO il BZ salmonacea 0OOOO2OdS OOOO Ll O telja Y 23D i 2a Oil @ GE it trilineata fritid et Ot © Od © verrucosa QHOOil@gail | Or Oo il oO verta Ol @M@A2DIGCOMDIDOI species | le ese Ds I Dee AP ay ale One 2 species 2 DD Oe ND 2 OA lO species 3 22370 til il @ ith @o it @ For all characters 9 = unknown 1. peduncles 0 = absent, | = present, 2 = well elevated 2. cerata 0 = congested, 1 = separate, 2 = single groups 3. rhinophores 0 = simple, 1 = annulate, 2 = papillate, 3 = perfoliate 4. anus 0 = posterior, | = middle of cluster, 2 = interhepatic space 5. oral glands 0 = absent, 1 = ramified, dorsal, 2 = simple, ventral 6. central cusp 0 = large, 1 = small, 2 = depressed 7. receptaculum seminis 0 = serial, 1 = semiserial, 2 = bilobed, 3 = multilobed 8. bursa copulatrix 0 = present, | = absent 9. foot corners 0 = rounded, | = tentacular 10. reproductive system 0 = diaulic, 1 = triaulic 11. lateral teeth 0 = denticulate, 1 = smooth, 2 = denticulate on outer edge 12. number of laterals 0 = 2-3, 1 =1
13. rhinophoral papillae 0 = absent, 1 = short, 2 = elongate
GOSLINER AND KUZIRIAN: BAJA CALIFORNIA FLABELLINIDAE
alisonae species 3 ornata babai engeli
telja affinis funeka albomarginata marcusorum species | rubrolineata species 2 poenicia bertschi pellucida pedata pricei trilineata gracilis cynara iodinea dushia verta browni capensis lineata fusco verrucosa falklandica nobilis solmonacea
islandica
Ficure 7. Cladogram depicting preliminary phylogeny of Flabellina.
14 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 47, No. 1
by virtually all opisthobranch systematists. The Flabellinidae is a morphologically diverse as- semblage, containing primitive and derived taxa. Subdividing the family into genera is problem- atic, as there is a continuum of gradually adding derived features, sometimes with poor correla- tion of these characters. Cladistic analysis, em- ploying Phylogenetic Analysis Using Parsimony (PAUP, by David Swofford), was conducted on all 33 species of Flabellinidae for which adequate morphological data are known, including three undescribed species. Polarity of characters was ascertained by using the Notaeolidiidae as the outgroup of the Flabellinidae. Thirteen charac- ters were considered for 33 taxa. The characters and their distributions are included in Table 2.
Subsequent cladistic analysis produced six equally parsimonious trees of similar configu- ration and a length of 37 and a consistency index of 0.568, one of which is presented here (Fig. 7). The other five trees differ only in the placement of one or two taxa. All of these cladograms dem- onstrate that genera such as Flabellina and Cory- Dhellina cannot be maintained without render- ing Coryphella paraphyletic. Maintenance of these genera as distinct requires the erection of many new monotypic genera and higher taxa for more primitive members of the family. There are no autapomorphic features to support many of these taxa. This is clearly counterproductive and pro- vides no further resolution of the phylogeny and classification of the Flabellinidae. From this analysis, it is apparent that most plesiomorphic members of the Flabellinidae are restricted to polar environments, with more derived taxa in- habiting temperate and tropical waters. Little adaptive radiation and speciation has occurred within primitive members of the family. Most speciation has occurred in the two clades that possess either perfoliate or papillate rhinophores. Accumulation of morphological data for species that are not well known, including additional undescribed taxa, will provide needed data for future re-evaluation of the generic divisions within the Flabellinidae.
ACKNOWLEDGMENTS
Several individuals assisted in the successful completion of this project. Hans Bertsch, David Behrens, Jeff Hamann, the late Eveline Marcus, and Gary McDonald kindly provided specimens of the two species described in this paper. The
collection of specimens of Flabellina rubrolinea- ta in Madang, Papua New Guinea, was made possible by a fellowship from the Christensen Research Institute. Jean DeMouthe kindly pre- pared the ink drawings.
LITERATURE CITED
Ase, T. 1964. Opisthobranchia of Toyama Bay and adjacent waters. Hokuryu-kan, Tokyo. 99 pp.
Basa, K. 1949. Opisthobranchia of Sagami Bay collected by His Majesty The Emperor of Japan. Iwanami Shoten, Tokyo. 194 pp.
1955. Opisthobranchia of Sagami Bay. Supplement. Iwanami Shoten, Tokyo. 59 pp.
Baker, F. AND G D. HANNA. 1927. Marine mollusks of the order Opisthobranchia. Proc. Calif. Acad. Sci. ser. 4, 16(5): 123-134.
Bertscu, H. 1977. The Chromodoridinae nudibranchs from the Pacific coast of America. Part I. Investigative methods and supra-specific taxonomy. Veliger 20(2):107-118.
. 1978a. The Chromodoridinae nudibranchs from the
Pacific coast of America. Part II. The genus Chromodoris.
Veliger 20(4):307-327.
. 1978b. The Chromodoridinae nudibranchs from the
Pacific coast of America. Part III. The genera Chromolaich-
ma and Mexichromis. Veliger 21(1):70-86.
. 1978c. The Chromodoridinae nudibranchs from the Pacific coast of America. Part IV. The genus Hypselodoris. Veliger 21(2):236-250.
BertscH, H. AND A. J. FERREIRA. 1974. Four new species of nudibranchs from tropical West America. Veliger 16(4):343- 353.
Cooper, J. G. 1863. Some new genera and species of Cali- fornia Mollusca. Proc. Calif. Acad. Nat. Sci. 2:202-207. FERREIRA, A. AND H. BertscH. 1972. Additional data on Flabellina telja (Gastropoda: Opisthobranchia). Veliger 14(4):
414415.
GosLinerR, T. M. AND D. W. BEHRENS. 1985. Two new species and genera of aeolid nudibranchs from the tropical eastern Pacific. Veliger 29(1):101-113.
Gos.iner, T. M. AND R. J. GriFFitus. 1981. Description and revision of some South African aeolidacean Nudibranchia (Mollusca, Gastropoda). Ann. S. Afr. Mus. 84(2):105-150.
Keen, A. M. 1971. Sea shells of tropical West America: ma- rine mollusks from Baja California to Peru, 2nd ed. Stanford Univ. Press, Stanford. 1064 pp.
KerstitcH, A. 1989. Sea of Cortez marine invertebrates. A guide for the Pacific coast, Mexico to Ecuador. Sea Chal- lengers, Monterey. 112 pp.
MacFar.tanp, F. M. 1924. Expedition of the California Academy of Sciences to the Gulf of California. Opistho- branchiate Mollusca. Proc. Calif. Acad. Sci. ser. 4, 13(25): 389-420.
Marcus, Er. AND Ev. Marcus. 1967. American opistho- branch mollusks. Stud. Trop. Oceanogr. 6:1—256.
Marcus, Ev. 1970. On some opisthobranchs from Cananeia, Brazil. Bolm. Fac. Fil. Cien. Let. Univ. Sao Paulo, Zool. 27: 207-228.
Marcus, Ev. AND Er. Marcus. 1961. On Coryphellina rubrolineata O’Donoghue, 1929 (Opisthobranchia, Eolida- cea). Proc. Malac. Soc. Lond. 34(4):224—227.
GOSLINER AND KUZIRIAN: BAJA CALIFORNIA FLABELLINIDAE 15
AND 1963. Opisthobranchs from the Lesser
Antilles. Stud. Fauna Curacao 19(79):1-76.
AND . 1970. Some gastropods from Madagascar and Mexico. Malacologia 10(1):181-223.
Mitter, M. 1971. Aeolid nudibranchs (Gastropoda: Opis- thobranchia) of the families Flabellinidae and Eubranchidae from New Zealand waters. Zool. J. Linn. Soc. 50(4):311- 3377
O’DonoGuueE, C. 1929. Report on the Opisthobranchia. Zoo- logical results of the Cambridge Expedition to the Suez Ca- nal, 1924. Trans. Zool. Soc. Lond. 22(6):713-841.
SCHMEKEL, L. AND A. PORTMANN. 1982. Opisthobranchia des Mittlesmeeres, Nudibranchia und Saccoglossa. Springer- Verlag, Berlin. 410 pp.
Tan, T., J. Pat, AND K. HsHa. 1987. An investigation on the distribution of nudibranch mollusks along the coast of Tai- wan, R.O.C. Bull. Malac. Rep. China 13:71-90.
Voict, F. S. 1834. Pp. 113-126 in G. Cuvier. Das Thier- reich 3.
WILLAN, R. C. AND N. CoLemAN. 1984. Nudibranchs of Aus- tralasia. Australian Marine Photographic Index, Sydney. 56
pp.
CALIFORNIA ACADEMY OF SCIENCES Golden Gate Park San Francisco, California 94118
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Marine Biological Laboratory LIBRARY
Nov 7 1990
PROCEEDINGS OF THE
CALIFORNIA ACADEMY OF CIENCES Hoje, Mass.
Vol. 47, No. 2, pp. 17-45, 5 figs., 1 table. October 17, 1990
A CHECKLIST AND KEY TO THE HERPETOFAUNA OF NEW CALEDONIA, WITH REMARKS ON BIOGEOGRAPHY
By Aaron M. Bauer Biology Department, Villanova University, Villanova, Pennsylvania 19085
and Jens V. Vindum
Department of Herpetology, California Academy of Sciences, Golden Gate Park, San Francisco, California 94118
Asstract: A checklist and key to all amphibian and reptile species, terrestrial and marine, from New Caledonia and the Loyalty Islands are provided. Thirty-six of 68 extant amphibian and reptile species of the New Caledonian region are represented by 1,643 specimens in the collections of the California Academy of Sciences. Specimen lists and natural history and systematic notes of these taxa are presented. The reptile fauna of the New Caledonian mainland is largely endemic. Certain taxa, especially arboreal geckos and leaf- litter-dwelling skinks, are restricted to regions of lateritic soils and/or high rainfall within the island. Other faunal components include regional endemics (New Caledonia and Loyalty Islands and Vanuatu), pan-oceanic taxa, and known human introductions in historical times. The gekkonine geckos are generally easily trans- ported and have colonized the region only after Vanuatu and the Loyalty Islands obtained their present configuration with respect to New Caledonia. The endemic diplodactyline geckos are most closely related to New Zealand and Australian forms and probably became isolated as a result of Mesozoic tectonic movements and sea-level changes. The zoogeography of most of the scincids remains obscure. Human activity in New Caledonia has likely been responsible for the extinction of additional reptile taxa representative of a once broader herpetofaunal diversity.
Received January 18, 1990. Accepted March 14, 1990.
INTRODUCTION et in their description. Recent systematic treatments
New Caledonia is unique among the islands _ of the skinks (Greer 1974; BOhme 1976; Sadlier
of the tropical Pacific in possessing a speciose herpetofauna composed largely of endemic forms (Bauer 1988a). The native terrestrial herpeto- fauna, which consists only of geckos and skinks, remains poorly known as a consequence of the isolation of New Caledonia, its mountainous ter- rain, and large size (19,103 km?). Almost every major collecting trip to the region has revealed new species or rediscovered others not seen since
[17]
1986; Ineich and Sadlier 1990) and carphodac- tyline geckos (Bauer 1986, and in press; Sadlier 1989) of New Caledonia and its satellites, the Loyalty Islands, have provided useful informa- tion regarding distribution, morphology, and phylogeny. Nonetheless, most of the taxa remain poorly known, and new forms undoubtedly re- main to be discovered.
In this paper we present a checklist and key to
18 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 47, No. 2
the herpetofauna of the New Caledonian region and summarize biological information for 36 of the 68 species inhabiting the New Caledonian area (33 of 53 terrestrial taxa). This information is derived primarily from specimens housed in the herpetological collections of the California Academy of Sciences (CAS).
The New Caledonian amphibians and reptiles in the CAS collection originate from several sources. The first eight specimens accessioned, including two paralectotypes, were obtained by exchanges from the Naturhistorisches Museum Basel, the repository of the large New Caledonian collections made by Fritz Sarasin and Jean Roux during the period 1911-1912. Until the last de- cade, Basel (NMBA) and Paris (MNHN) were the only major collections of reptile material from the island, although type specimens and smaller collections were housed in a number of other museums. Two of these collections, those of the Museu de Lisboa and the Musée de I’Ecole de Medécine Navale de Brest, are no longer extant, the former having been destroyed by fire and the latter dispersed or lost.
An important collection of 120 specimens was donated to the Academy by F. X. Williams in 1940, including material from the Ile des Pins. Unfortunately, there are no precise locality and collection data associated with some of this ma- terial, and the Academy’s archives include no field notes or records from Williams’s collecting trip. Other additions include sea snakes obtained through Stanford University (CAS-SU), a small collection from Ted Case (University of Califor- nia, San Diego), and a specimen of Leiolopisma steindachneri obtained through exchange with the Australian Museum, now a leading repository of New Caledonian reptile specimens. The remain- der of the California Academy of Sciences New Caledonian collection was built by material ob- tained by the senior author as a result of ongoing research on the systematics, morphology, and biogeography of the geckos of the southwest Pa- cific. These additions make the California Acad- emy of Sciences a repository for one of the largest collections of New Caledonian reptile material in the world, with 1,643 specimens, including eight types.
MATERIALS AND METHODS
The standard abbreviations m (meters), mm (millimeters), mybp (million years before pres-
ent), and SVL (snout—vent length) are used in the following text. Collection abbreviations follow Leviton et al. (1985). A checklist is provided listing original authors, representation in the CAS collection, and broad distributional patterns of all species in the region, including fossil taxa. An artificial key to the living taxa—using, when available, invariant characters easily determined from museum specimens—is also provided.
Species accounts are for those taxa represented in the CAS collections. Specimens examined in- clude only those in the California Academy of Sciences (CAS and CAS-SU), although New Cal- edonian specimens from 17 additional institu- tions have been examined by the senior author (see Bauer 1986 for a complete listing of gek- konid specimens examined). Synonymies are limited to first description, junior synonyms (if applied to New Caledonian populations), and first or establishing usage of the currently employed combination. More complete synonymies are available in the literature for Litoria aurea (Duellman 1977; Cogger et al. 1983), sea turtles (Wermuth and Mertens 1977), geckos (Wermuth 1965; Bauer 1986; Ineich 1987; Sadlier 1989), Ramphotyphlops braminus (Robb 1966; Mc- Dowell 1974; Ineich 1987), Candoia bibroni Stimson (1969), and sea snakes (Stejneger 1907; Smith 1926; Gail and Rageau 1958). Unfortu- nately, complete synonymies more recent than Boulenger (1887) are not available for most New Caledonian skinks, although Sadlier (1986) pro- vided partial synonymies. Remarks on natural history, diet, and reproduction are limited to CAS material, of which the authors have field knowl- edge, and are provided to supplement, confirm, or rectify information available in the literature. The basic biological data pertinent to the other large collections of New Caledonian material have already been presented (Roux 1913; Sadlier 1986, 1989). Summaries of the biology of some species may be found in Bavay (1869), Roux (1913), Meier (1979), Sadlier (1986, 1989), Bauer (1986), and Bauer and Devaney (1987). A comprehen- sive bibliography of New Caledonian herpetol- ogy was prepared by Bauer (1985b).
Localities represented by collected material are number coded and appear in Figure 1. Many of these localities were visited on several occasions during different seasons in an attempt to achieve a representative sample of the fauna. The coor- dinates, elevation, and basic vegetational char- acteristics of these sites are summarized below.
BAUER AND VINDUM: NEW CALEDONIAN HERPETOFAUNA 19
FiGure 1.
LOCALITIES REPRESENTED BY SPECIMENS IN THE CAS COLLECTIONS
Collection Dates: A = July—October 1940; B = March-April 1984; C = September—October 1984; D = May-June 1985; E = December 1985- January 1986; F = December 1986; and G = miscellaneous dates including exchanges.
Loca ity |: Boat Pass Beach (Pointe Narian), 20°05’S, 164°00’E, sea level (C). Northernmost point of mainland New Caledonia. White sandy beach bounded by mangrove vegetation.
Loca.ity 2: Cascade de Galarino, 20°30’S, 164°46’E, elevation 0-20 m (C). Base of large waterfall dropping steeply from southeast slopes of Mt. Colnett (1,505 m) to coast. Vegetation transitional from humid forest to narrow strand palm belt.
Loca.ity 3: Koumac Caves, 20°34’S, 164°17’E, elevation 20-100 m (E). Limestone outcrop in savanna/niaouli (Melaleuca quinquenervia, the
Map of New Caledonia showing collecting sites represented by specimens in the California Academy of Sciences. See text for key to numbered localities.
dominant myrtaceous tree in drier areas of New Caledonia) mosaic. Ficus and other trees are in- vested within the outcrop itself.
Loca.ity 4: Ilot de Hiénghéne, 20°38’S, 164°56’E, sea level (C). Small, flat, coralline is- land, 3.5 km off coast of Ouanpoués, near Hiénghéne. Vegetation includes palms, Ficus, and variety of other trees and shrubs, excluding truly mesic forms.
LocaLity 5: Campement Vanhalle, 20°40’S, 164°44’E, elevation ca. 800 m (C). Densely for- ested region between drainages of Hiénghéne and Ouaiéme rivers. Characterized by mature humid forest, including tree ferns and Agathis. Tra- versed by several small, rocky streams. Forest clearings bounded by palms and cut by small, weed-choked brooks.
Loca.ity 6: Hiénghéne, 20°41'S, 164°56’E, sea level (A and G). Town in region of large black limestone cliffs, crags, and caves. Vegetation in
20 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 47, No. 2
and around townsite highly disturbed, surround- ing areas—including mountains behind the town (400-450 m)—covered in dense humid forest.
Loca.ity 7: Touho, 20°47'S, 165°14’E, ele- vation 0-50 m (C and E). Grassy field on hill overlooking Touho airport and adjacent forested region inland from the town center. Vegetation in townsite largely disturbed; inland forests typ- ical humid types.
Loca.ity 8: North bank, Tiwaka River, 4 km W of Rte. 3, 20°54’S, 165°13’E, elevation 0-30 m (D). Hilly area, typified by humid forest vege- tation, broken by small riverside plantations.
Loca.ity 9: Vallée d’Amoa, 0-1.0 km W of Rte. 3, 20°55’S, 165°17’E, sea level (C, E and F). Short transect along north bank of Amoa River. Vegetation of humid forest type broken by small family dwellings and plantations.
Loca.ity 10: Vallée d’Ina, 20°56'S, 165°19’E, elevation 10-20 m (C). Grassy pasture with scat- tered rocks in forest clearing near Ina River.
Locatity 11: Poindimié, 20°56’S, 165°20’E, elevation 0-20 m (B, C, D, E and F). Disturbed roadside vegetation, house and hotel walls, and sheets of corrugated metal in town.
Loca.tity 12: Mount Koyaboa, 20°57'S, 165°20’E, elevation 20-390 m (C, D, E and F). Lower slopes (to ca. 50 m) characterized by sec- ondary-growth scrub and some primary and sec- ondary mixed non-coniferous forest associa- tions. Middle elevations (50-140 m) with fern scrub and niaouli. Above 140 m, vegetation pri- marily secondary growth with some primary humid-forest trees (see Bauer and DeVaney 1987 for detailed description of this site).
Locautity 13: St. Thérése, Vallée d’Amoa, 20°59’S, 165°13’E, elevation 0-20 m (B). Densely forested region along Amoa River, some Arau- caria (Araucariaceae, column pines) present.
LocaALity 14: Koné, 21°04’S, 164°52’E, sea level (B and C). Sclerophyll forest, residential gardens, disturbed riverbanks, and banana grove.
Loca.Lity15: Vallée de Nimbayes, 21°05’S, 165°21’E, elevation 0-30 m (C). Humid forest.
Loca.ity 16: 10 km E of Pouembout, 21°08’S, 164°59’E, elevation ca. 50 m (B). Sclerophyll for- est with thick layer of leaf litter along banks of Pouembout River in area of forest/savanna mo- saic.
Loca.ity 17: Creek crossing, | km NE of Mt. Aoupinie, 21°08'S, 165°21’'E, elevation 500 m (G). Locality described by Sadlier (1986).
LocaALity 18: Nakety, 21°22'S, 166°34’E, ele-
vation ca. 350 m (C). Open grassy area in forest/ savanna mosaic.
LocaLity 19: 16 km NW of Plage de Poé, 21°34’S, 165°24’E, elevation ca. 120 m (C and E). Open sclerophyll forest bordered by culti- vated fields. Traversed by dry creek beds, dead standing trees and thick leaf litter abundant.
LocaLiry 20: Cascade de Cin; 219345 165°58’E, elevation ca. 270 m (C). Rocky pools at top of waterfall in disturbed area of shrubs and grass.
Locauiry 21: Mt. Canala, 21°35’S, 1657356 = elevation 100—1,051 m (G). Exact site unknown. Area of high rainfall with Araucaria stands and generally humid-forest vegetation.
Locauity 22: Plage de Poé, 21°37’S, 16522478) sea level (C, D and E). Grassy site bordering white sand beach. Roadsides bordered by water- filled ditches and tall grass.
LocaLity 23: Baie de Gouaro, 21°37'S; 165°26’E, sea level (D). Beach wrack and grassy borders of white sand beach.
LocaLity 24: Baie des Tortues, 21°37’S, 165°27’E, sea level (C and D). Beach wrack and vegetation along small seaside cliffs. Area of large coastal grove of Araucaria columnaris.
Loca.ity 25: 15 km S of Bourail on Rte. 1, 21°41’S, 165°35’E, elevation ca. 50 m (D). Grassy ditch on west side of highway in hilly area of forest/savanna mosaic.
Loca.ity 26: 63 km S of Bourail on Rte. 1, 21°47’S, 165°50’E, elevation ca. 40 m (D). Grassy ditch on west side of highway in area of savanna dotted with scattered naiouli.
Loca.ity 27: Mt. Mou, 22°04’S, 166°21’E, el- evation ca. 500 m (F). Humid forest on the lower slopes of mountain. Area of very large trees and high canopy with abundant ground cover tra- versed by several small streams.
LOcALITY 28: ca. 5 km W of Yaté off Rte. 2, 22°09’S, 166°53’E, elevation ca. 500 m (F). Rocky, open area of edaphic scrub vegetation.
Loca.ity 29: Yaté (Waho), 22°09’S, 166°54’E, sea level (C). On main road through town in area of humid forest/maquis (mediterranean shrub vegetation) interface.
Locauity 30: Islet next to Yaté [Ile Potino?], 22°09'S, 166°57’E, sea level (G). Exact locality uncertain (F. X. Williams locality—no field notes). Possibly Ie Potino, a small island in the mouth of the Yaté River typified by maquis vege- tation.
Loca.ity 31: Mt. Koghis, 22°10’S, 166°32’E,
BAUER AND VINDUM: NEW CALEDONIAN HERPETOFAUNA
elevation ca. 460-800 m (D, E and F). Dense, mixed humid forest with Ficus and, more rarely, tree ferns and Araucaria. Forest floor covered by stones and moist leaf litter.
LocaLity 32: Touaourou, 22°11'S, 166°58’E, elevation 0-20 m (B, C, D, Eand F). Open grassy areas and adjacent white sand beach, maquis for- est along abandoned mining roads, and disturbed roadside habitats.
Locauitry 33: 20 km SE of St. Louis, 22°12’S, 166°42’E, elevation ca. 200 m (C). Roadside ma- quis off highway.
Locaity 34: Border of Commune de Mont Dore, on Rte. 2 at intersection of Tina-sur-Mer road, 22°13’S, 166°29’E, sea level (F). Partially flooded, low-lying grassy and marshy areas near housing development.
FocArmry 35: Vallée de Yahoué, 22°14’S, 166°30’E, elevation ca. 30 m (A and F). Small, dense forest patch along Yahoué River, inter- rupted by small banana plantations and habi- tations.
LocaLity 36: Col de Mourirange, 22°14’S, 166°39’E, elevation 258 m (D). Area of short, shrubby maquis in hilly region of lateritic soils.
Loca.ity 37: Nouméa, 22°16'S, 166°27’E, sea level (A, C, D, E and F). Capital city of New Caledonia. Urban building walls, gardens, and beach front.
Loca.ity 38: Riviére Bleue, 22°18’S, 166°50’E, 50-100 m (D). Maquis vegetation.
Loca.ity 39: Prony Bay, 22°19’S, 166°49’E, sea level (A). Region of maquis vegetation. Exact collecting localities unknown.
Locatity 40: Goro, 22°19’S, 167°00’E, ele- vation 0-20 m (C, D, E and F). Area near Gite Wadiana, along coral flats and in adjacent grassy areas with scattered coconut palms; also in nar- row belt of coastal forest with rocky substrate and along the margins of roadsides, in leaf litter, grass and rocks.
Koecarizy 41: Mt. L’>Aiguillon, 22°21’S, 166°50’E, elevation ca. 200 m (D). Transitional zone between humid forest and maquis vegeta- tion. Substrate rather rocky, traversed by several small streams.
Loca.ity 42: Ile de Pins (A). Exact location unknown.
LocaLity 43: Marine environments off the coast of Nouméa (G).
Loca.ity 44: Maré, Loyalty Islands (G). Exact location unknown.
9a.
9b.
10a.
10b. lla.
Ib:
2a:
12b.
13a.
21
ARTIFICIAL KEY TO THE EXTANT REPTILES OF THE New CALEDONIAN REGION
Ls MITA Sle SCM be en 2 | Lamips absent. (Snakes) 2.2 33
. Carapace present, limbs modified as flip-
pers (turtles) . Family Cheloniidae . . . 50
. No carapace, limbs unmodified (lizards)
. Body scales generally small, granular,
juxtaposed ...... Family Gekkonidae... 4
i Body scales mmibnicate = eee ee
OE ne 2 Fass ras Family Scincidae .. . 24
. Subdigital scansors present 0. 5 . Subdigital scansors absent 0.
Beet es eater oe Nactus pelagicus
. Digit I of manus and pes clawless ...... 6 . Alidigitselawe die: 2 ent Sane Eade 8
. Digit I rudimentary, without scansors _
Hemiphyllodactylus typus
. Digit I small but well developed, with
SGATISOFS) on ek i
. Penultimate phalanx completely free
from scansorial pad Gehyra vorax
. Penultimate phalanx partially subsumed
In pad =F Rims Lepidodactylus lugubris
. Digit 1) with apical plates =. 10 . Digit I without apical plates 0.
be ne: BAY eee 7 Hemidactylus... 9 Two pairs of enlarged post-mental chin shields contact infralabials iT Bae Sin tictea tt 3, Hemidactylus frenatus Second pair of enlarged postmentals ex- cluded from infralabials by series of small
SCaleGpemn sual e tes Hemidactylus garnotii Subdigital scansors divided 2, ARENT RS LEP ER CONN Bavayia... \1 Subdigital scansors undivided 17 Claw of digit I situated medial to single termimalsscansO ne iene same ater ct 12 Claw of digit I situated between halves oo: cleft apicaliscansor = 13
Lateral surface of hindlimb with distinct, contrasting pale spots on a dark back- PrOUnGIa!. millet tte Wis Bavayia ornata Lateral surface of hindlimb without pale Spots,OF Spots Indistinci~A.2 2. J ies «mA cate Ae Bavayia sauvagii Dorsal pattern with pale, broad vertebral SUG. en et Meera ee ee 14
22 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 47, No. 2
TasLe 1. Checklist of the recent and fossil* herpetofauna of New Caledonia.
eee. Distribution specimens Loy- in CAS New alty Ma- Taxon collection Caledonia Islands _rine Class Amphibia Order Anura Family Hylidae Litoria aurea (Lesson, 1830) [introduced] 70 x XxX (including 20 tadpoles) Class Reptilia Order Chelonia Family Cheloniidae Caretta caretta (Linnaeus, 1758) — »4 Chelonia mydas (Linnaeus, 1758) — x Eretmochelys imbricata (Linnaeus, 1766) — 4 Lepidochelys olivacea (Eschscholtz, 1829) — >. 4 Family Meiolaniidae *Meiolania mackayi Anderson, 1925 _ (Walpole Is.) *Meiolania sp. (see Gaffney et al. 1984) _ x Xx Order Crocodylia Family Mekosuchidae *Mekosuchus inexpectatus Balouet and Buffetaut, 1987 - x Order Squamata Suborder Lacertilia Family Gekkonidae Bavayia crassicollis Roux, 1913 1 xX x Bavayia cyclura (Ginther, 1872) 62 Xx xX Bavayia montana Roux, 1913 1 XxX Bavayia ornata Roux, 1913 — XxX Bavayia sauvagii (Boulenger, 1883) 299 XxX x Bavayia septuiclavis Sadlier, 1989 — xX Bavayia validiclavis Sadlier, 1989 _ x Eurydactylodes symmetricus (Andersson, 1908) — 4 Eurydactylodes vieillardi (Bavay, 1869) 1 xX Gehyra vorax Girard, 1857 — x Hemidactylus frenatus Duméril and Bibron, 1836 28 4 Xx [introduced] Hemidactylus garnotii Duméril and Bibron, 1836 4 xX xX Hemiphyllodactylus typus Bleeker, 1860 2 xX X Lepidodactylus lugubris (Duméril and Bibron, 1836) 23 XxX x Nactus pelagicus (Girard, 1857) 51 XxX xX Rhacodactylus auriculatus (Bavay, 1869) 38 4 Rhacodactylus chahoua (Bavay, 1869) 4 x Rhacodactylus ciliatus (Guichenot, 1866) —- xX Rhacodactylus leachianus (Cuvier, 1829) 9 x Rhacodactylus sarasinorum Roux, 1913 1 XxX Rhacodactylus trachyrhynchus Bocage, 1873 _ Xx Family Scincidae Caledoniscincus atropunctatus (Roux, 1913) 16 xX ».¢ Caledoniscincus austrocaledonicus (Bavay, 1869) 727 > 4 x Caledoniscincus festivus (Roux, 1913) 8 > 4 Caledoniscincus orestes Sadlier, 1986 = XxX Cryptoblepharus novocaledonicus Mertens, 1928 5 x x Emoia cyanura (Lesson, 1830) — xX Emoia loyaltiensis (Roux, 1913) - Xx Geoscincus haraldmeieri (B6hme, 1979) X Graciliscincus shonae Sadlier, 1986 1 X Leiolopisma greeri BOhme, 1979 — x
BAUER AND VINDUM: NEW CALEDONIAN HERPETOFAUNA
TABLE 1.
Taxon
Leiolopisma nigrofasciolatum (Peters, 1869) Leiolopisma novaecaledoniae (Parker, 1926) Leiolopisma steindachneri (Bocage, 1873) Leiolopisma n. sp. Ineich and Sadlier Marmorosphax euryotis (Werner, 1909) Marmorosphax tricolor (Bavay, 1869) Nannoscincus gracilis (Bavay, 1869) Nannoscincus greeri Sadlier, 1986 Nannoscincus mariei (Bavay, 1869) Nannoscincus rankini Sadlier, 1986 Nannoscincus slevini (Loveridge, 1941) Phoboscincus bocourti (Brocchi, 1876) Phoboscincus garnieri (Bavay, 1869) Sigaloseps deplanchei (Bavay, 1869) Tropidoscincus aubrianus Bocage, 1873 Tropidoscincus roehssii (Andersson, 1908) Tropidoscincus variabilis (Bavay, 1869) Family Varanidae *Varanus sp. (see Gaffney et al. 1984) Suborder Serpentes Family Typhlopidae
Ramphotyphlops angusticeps (Peters, 1878)
Ramphotyphlops braminus (Daudin, 1803) [introduced]
Ramphotyphlops willeyi (Boulenger, 1900) Family Boidae Candoia bibroni (Duméril and Bibron, 1844) Family Hydrophiidae Acalyptophis peronii (Duméril, 1853) Aipysurus duboisii Bavay, 1869 Aipysurus laevis Lacépéde, 1804 Disteira major (Shaw, 1802) Emydocephalus annulatus Krefft, 1869 Hydrophis gracilis (Shaw, 1802) Hydrophis coggeri (Kharin, 1984) Hydrophis ornatus (Gray 1842) Laticauda colubrina (Schneider 1799) Laticauda laticaudata (Linnaeus 1758) Pelamis platurus (Linnaeus 1766) Total species/species represented in CAS collection (extant taxa only) Total specimens in CAS collection
23 Continued. New ey oe Coindanaa Distribution specimens Loy- in CAS New alty Ma- collection Caledonia Islands rine 21 ».4 Xx 1 Xx 1 X — XxX 1 Xx 95 xX 26 XxX 12 x 35 x _ Xx 1 x _ x. - x Xx 65 x — x 15 xX 2 x — XxX — ? fh XxX aX — x — x jt Xx = Xx _ xX = X! —_ x = Xx! i 4 ad Xx! 8 xX 3 x — Xx 68/36 48/33 20/12 15/3 1643 1629 2 12
' The occurrence of these taxa in New Caledonia has been reported in the literature, but specimens from the region were not recorded by Cogger (1975) nor have they been verified by the authors. The confusion surrounding the identification of Hydrophis- like New Caledonia sea snakes was discussed by Minton and Dunson (1985).
13b. Dorsal pattern composed of pale, trans- versely oriented blotches Preanal pores in 2 rows; supranasal gen- erally separated by a single internasal 22 Fran tt ga Bi es Bavayia validiclavis Preanal pores in a single row, internasal region fragmented .... Bavayia septuiclavis
14a.
14b.
15a. First pair of infralabials usually contact-
ing medially 2.222 ea Bavayia montana 15b. First pair of infralabials usually separat- (©. | eae et eae i a er he NT a 16
l6a. Distinct bold, dark transverse bands bordering the pale, dorsal blotches seeeceeteey stores deme beet. Bavayia cyclura
24
16b.
Wie
Vib:
18a.
18b.
19a.
. Rostral contacts nostril
. Rostral excluded from nostril
.. HOrelimbspentadactyli.2 2. es.
. Phalangeal formula for pes 2.3.4.5.4
PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 47, No. 2
Pale dorsal blotches and dark bands ob- scure and poorly defined — ES ee ee Eek Bavayia crassicollis Body scales greatly enlarged, body com- DICSSc 22ers. Eurydactylodes ... 18 Body scales small and granular or tu- bercular Rhacodactylus ... 19 Cruciform patch of raised, rounded scales on nape; continuous slit from angle of jaw to ear .... Eurydactylodes symmetricus No raised scales on nape, slit from angle of jaw to ear interrupted by skin fold .... a, ee Eurydactylodes vieillardi Body with loose folds of skin along throat and flanks, digits half-webbed _.. 20
. Body without lateral folds, digits less than
one third webbed deta deanery ZA
fv aiarnietre Rhacodactylus chahoua
Be Rhacodactylus leachianus
. Paired, converging ciliated crests on dor-
SUM ae Rhacodactylus ciliatus
. Dorsal scales generally homogeneous . 22 . Head with raised bosses or rugosities . 23 ¥ ead smooth Weer eee 6a ee
SMOUE TUE OSG weet eer eee ee ene
Rhacodactylus trachyrhynchus
. Raised orbital and aural bosses present,
snout smooth _. Rhacodactylus auriculatus
. “Ablepharine eye” (completely fused
brill)... Cryptoblepharus novocaledonicus
_ Brillinot,completelysfused 22s es DS . Ear opening minute or absent wt » Earopening not ninute 3 a = 31 . Body elongate, snout blunt and flattened;
5 supralabialiscales;. a ee
. Body elongate, snout pointed; 6 supra-
labralescalesyee es Nannoscincus ... 27
. Ear opening absent . Nannoscincus mariei | Earopening minute: = = seeei 28 . Frontoparietals distinct, lower eyelid
Sealy e.o0 ke Sas CR, Fe 29
. Frontoparietals fused, lower eyelid with
semi-translucent central area ow... 30 es ie eee ae ee Nannoscincus gracilis
i AR lsat Nannoscincus greeri
30b. 31a. Sib: 32a.
a20.
33a.
33D:
34a.
34b.
35a. 35b. 36a. 36b. 37a.
37b.
38a.
38b.
39a.
39b.
40a.
40b.
Phalangeal formula for pes 2.3.4.4.3 _.. es A a Bee ke) Nannoscincus rankini Supranasal’scales present... eee 52 Supranasal scales "absent =e 33 Lower eyelid scaly, limbs short, teeth re- CUiVed iets tt aed Phoboscincus . . . 33 Lower eyelid with transparent disk, limbs overlap when adpressed; teeth not re- Cunved seen! renin. oe Emoia ... 34 Supranasals broadly contacting; 32-34 midbody scale rows 2. = eee Pepin 8 ane Phoboscincus garnieri Supranasals narrowly separated, 66 mid- body scale rows ..... Phoboscincus bocourti Mid-vertebral stripe present; 52-64 scale rows between parietal and tail base _.. ssn BRON i AMO Se Emoia cyanura No vertebral stripe; 62-71 scale rows be- tween parietal and tail base Peseta sorted vite Medien Emoia loyaltiensis Dorsal body scales smooth _ 36 Dorsal body scales carinate 40 Frontoparictals fused _ “=e aaa 37 Frontoparietals distinct “2 aaa 38 Enlarged nuchal scales; 22-27 lamellae beneath fourth toe .. Sigaloseps deplanchei No enlarged nuchal scales; 37-47 la- mellae beneath fourth toe... eae WO Ae Leiolopisma steindachneri Lower eyelid scaly,2 2.4.2.2 2. shane Geoscincus haraldmeieri
disk 39 Paravertebral scales 74; prefrontals con- tacting .. es) aaa eee Leiolopisma greeri Paravertebral scales 82-96; prefrontals not contacting or contacting only at a POMt 22s. Leiolopisma nigrofasciolatum Ear with one much-enlarged anterior lobule == Leiolopisma novaecaledoniae Ear with small lobules or no distinct lob-
. Supralabials excluded from lower eyelid
by subocular row .. Marmorosphax . . . 42
. Supralabials not entirely excluded from
lower eyelid] LO 2 a ee 43
. Parietal scales bordered by uniform sized
scales similar to dorsals = (eee .. Marmorosphax euryotis
. Parietal scales bordered by single upper
secondary temporal and single enlarged nuchal scale ........ Marmorosphax tricolor
BAUER AND VINDUM: NEW CALEDONIAN HERPETOFAUNA
43a.
43b.
44a.
44b.
45a.
45b.
46a.
46b.
47a.
47b.
48a.
48b.
49a.
49b.
50a.
5Ob.
Sla.
S1b.
52a:
52b.
Joa.
JOU,
Prefrontal scales contacting or narrowly Separated fn Sie Bee oD gto teste NN ome Prefrontal scales well separated + aE Pee Oe ec Caledoniscincus . . . 47 Prominent postnasal suture present oat Ae ee ee Tropidoscincus ... 45 Postnasal suture absent Ble res ceo or reat Leiolopisma n. sp. Scales on posterior aspect of thighs over- lapping and evenly decreasing in size US ota ao ae A ee 46 Scales on posterior aspect of thighs small and juxtaposed, sharply demarcated from large, overlapping scales of dorsum of iret ee Tropidoscincus variabilis Midbody scale rows 32-34 0. Tropidoscincus aubrianus Midbody scale rows 36-40 oo Pa ater tert at Tropidoscincus roehssii Tail with dark, transversely aligned ENCVRONS 2. were: 24 Bese 48 Tail without chevrons, rostral scale al- ways without dark median streak Boa htcl aly Caledoniscincus atropunctatus Dorsum with either vertebral or lateral palesiriping or both. = 49 Dorsum without vertebral or lateral striping, dark median rostral streak al- ways present Caledoniscincus festivus Dorsum with pale vertebral stripe only, dark median rostral streak always pres- ji ear ER ae Caledoniscincus orsetes Dorsum with either vertebral or lateral pale striping, or both; dark median ros- tral streak variably present 0.
Sh Lele: Caledoniscincus austrocaledonicus Four costal shields on each side of car-
Five or more costal shields on each side DIEGATAP Acer CALS eee SPOOR 20S) S 32 One pair of prefrontal scales present .... sh Sass Bes ae ce Chelonia mydas Two pairs of prefrontal scales present .. int atleun ft odlt. te Eretmochelys imbricata Four enlarged inframarginals on bridge MMB Sv RE) hale nae s Lepidochelys olivacea Three enlarged inframarginals on bridge Caretta caretta Tail laterally compressed, marine _. Family Hydrophiidae . . . 57 Tail rounded in cross-section, terrestrial Peres eee Vere. eee heme BEN LI AYE 54
54a.
54b.
55a.
S50:
56a.
56b.
57a.
57:
58a. 58b.
59a.
59b.
60a.
60b.
6la.
61b.
62a.
62b. 63a. 63b. 64a.
64b.
65a.
65b.
66a.
66b.
25
Head distinct, body large, scales keeled Family Boidae . .. Candoia bibroni Head indistinct, body extremely thin, scales smooth
Family Typhlopidae ... 55 Nasal cleft contacts preocular scale AEA SM cen Fe Ramphotyphlops braminus Nasal cleft not contacting preocular scale ead Se iietaehhe trl hl tte sandy ee deel: turd 56 Nasal cleft contacts first supraorbital scale ie Ramphotyphlops angusticeps Nasal cleft contacts second supraorbital Scale gum sua ob Ramphotyphlops willeyi Ventral scales at least three times as wide aS aGjacemt SCaleS. 2.02 a 58 Ventral scales barely twice width of ad- jacemtiscalesoe d: Sectemties ti. tu tol ANAT LE 2 62 Nasals contact one another 60 Nasals separated by internasal scales ..... ed tl 9 B15 0 Laticauda ... 59 Scale rows at midbody 19 1)" ") & sd ee Laticauda laticaudata Scale rows at midbody 21 or more ........ AR cee. Serhan Oe as Laticauda colubrina Supralabral scales: 3). 27 Jes See oe eS Emydocephalus annulatus Supralabial scales 6 or more Sees ae Oo es Alpysurus ... Head shield regular, larger than scales of INGO K eA st rt ej Bist Ded Aipysurus laevis Head shield irregular, roughly equal in size to scales on neck .. Aipysurus duboisii Head scutes small, irregular; supraocu- lars without tubercles or spines (Oy OORT es Si ORY Acalyptophis peronii Head scutes enlarged, regular, supraocu- lars without tubercles of spines 63 Mental groove absent ..... Pelamis platurus Mental’eraove present = 2 64 Anterior chin shields reduced, excluded from mental groove by first infralabials eee nS eee eee ree Disteira major Anterior chin shields large, bordering mentalygroove ase. Hydrophis .. . 65
Posterior ventrals mostly undivided .. 66 Scale rows at midbody 29-35 VAR edo £008 | Hydrophis coggeri Scale rows at midbody 39-59 0.
Wena Ac WIA USES ¢ Hydrophis ornatus
26 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 47, No. 2
SPECIES ACCOUNTS
AMPHIBIA ANURA FAMILY HyLIDAE
Litoria aurea (Lesson)
Rana aurea Lesson, 1830:60. Type locality: “Macquarie River, Bathurst, New South Wales, Australia.” Lectotype locality: “Port Jackson” [New South Wales, Australia] (designated by Roux-Estéve 1979).
Litoria aurea Tyler, 1971:352.
MATERIAL EXAMINED (50 adults; 20 tadpoles).— Locality 5: 6 Oct. 1984 (CAS 157654); Locality 8: 6 June 1985 (CAS 158555); Locality 9: 15 Dec. 1986 (CAS 162169); Locality 11: 10-11 Oct. 1984 (CAS 157655-6); Locality 14: 29 Mar. 1984 (CAS 157104-5), 1 Oct. 1984 (CAS 157652-3); Locality 19: 20 Oct. 1984 (CAS 157658-73); Locality 20: 17 Oct. 1984 (CAS 157674, lot of 20 tadpoles); Locality 21: 27 May 1985 (CAS 158311-20, 159025-8); Locality 32: 17 Oct. 1984 (CAS 157657), 4 Jan. 1986 (CAS 159569), 20 Dec. 1986 (CAS 162174-5); Locality 34: 17 Dec. 1986 (CAS 162170-3); Lo- cality 35: 23 Dec. 1986 (CAS 162176); Locality 40: 5 Jan. 1986 (CAS 159570-1).
REMARKS.—The date of introduction of this Australian species into New Caledonia is un- known. Specimens referenced by Copland (1957) and Moore (1961), however, indicate that Litoria aurea has been present for over a century. By the time of Sarasin and Roux’s expedition (191 1- 1912), this frog was widely distributed across the mainland of New Caledonia (Sarasin 1925), whence it was introduced into the New Herbrides [=Vanuatu] (Tyler 1976, 1979). The few speci- mens in early collections, however, suggest that L. aurea was not abundant until quite recently. Although it is now found in almost all habitats, L. aurea is most common in disturbed lowland sites such as gardens and ditches.
Specimens from areas of native forest were typically found singly, with males calling from the ground or from low branches. The species is carnivorous (Cogger et al. 1983) and even can- nibalistic (Cogger 1986), and in Australia it has been known to take large vertebrate prey, in- cluding elapid snakes (Tyler 1976). One individ- ual collected at Locality 34 disgorged a Caledoni- scincus austrocaledonicus.
The presence of both calling males and free- swimming tadpoles in October is consistent with the reported breeding period of L. aurea in New Zealand (Bell 1982) and its native Australia (Barker and Grigg 1977). Tadpoles were collect- ed from small rock pools at the top of a waterfall, but were also observed in slow-flowing creeks
and vegetation-choked ponds from sea-level to 800 m (Campement Vanhalle—Locality 5).
REPTILIA
SQUAMATA Family GEKKONIDAE
Bavayia crassicollis Roux
Bavayia cyclura crassicollis Roux, 1913:89. Type locality: “Maré: Netché, R6, Pénélo; Lifou: Quépénée; Ouvéa: Fa- yaoué.”’ Lectotype locality: ““Maré, Loyalty Insel” (restricted by Kramer 1979).
Bavayia crassicollis Sadlier, 1989:365.
MATERIAL EXAMINED (1 specimen).— Locality 4: 13 Oct. 1984 (CAS 157695).
REMARKS. — One large specimen (70 mm SVL) was collected under loose bark of a dead standing tree on Ilot de Hiénghéne, a tiny coralline island off the coast of Hiénghéne. The species may be sympatric with other Bavayia in forest clearings (Sadlier 1989), but appears to be most common in the coastal lowland forest and mangroves on New Caledonia and the Loyalty Islands (Roux 1913).
Bavayia cyclura (Giinther)
Peripia cyclura Giinther, 1872:422. Type locality: ““New Cal- edonia.”
Lepidodactylus neocaledonicus Bocage, 1873a:206. Type lo- cality: “Nouvelle Calédonie.”
Hemidactylus (Peripia) Bavayi Sauvage, 1879:71. Type local- ity: ““Nouvelle-Calédonie.”
Bavayia cyclura Roux, 1913:88.
MATERIAL EXAMINED (62 specimens).— Locality 9: 1 Jan. 1986 (CAS 159546, 159550-1), 8 Jan. 1986 (CAS 165877), 14-15 Dec. 1986 (CAS 162203-9, 162219-21); Locality 11: 7 Apr. 1984 (CAS 165878-9), 10 Oct. 1984 (CAS 157696), 1-2 June 1985 (CAS 158549-5S0); Locality 12: 6 June 1985 (CAS 158548); Locality 19: 20-21 Oct. 1984 (CAS 157697-704, 165884-7), 13 Jan. 1986 (CAS 165861-74); Locality 35: July-Aug. 1940 (CAS 80842, 80849-50, 80853), 23 Dec. 1986 (CAS 162237- 9); Locality 37: July—Oct. 1940 (CAS 80864-71); Locality 39: July—Aug. 1940 (CAS 80863).
REMARKS.—Although widely distributed on New Caledonia and all of the Loyalty Islands, Bavaya cyclura is restricted to the vicinity of trees and rotten or hollow logs. Only in the Forét de Yahoué (Locality 35) were specimens of this species found to be syntopic with B. sauvagii. In this area, specimens of both species were col- lected from the trunks of trees at heights of 1-3 m. At Mt. Koyaboa (Locality 12) B. cyclura were found only at higher altitudes than B. sauvagii
BAUER AND VINDUM: NEW CALEDONIAN HERPETOFAUNA 27
Ficure 2. Variation in color and body proportion in Bavayia species. Specimens (bottom to top): B. crassicollis from Ilot de Hiénghéne (Locality 4); B. cyclura from Vallée de Yahoué (Locality 35); and B. montana from Mt. Koyaboa (Locality 12).
and in conjunction with logs, rather than stones, The largest individual examined (from Locality as hiding places. A few individuals were found 19) was 72 mm SVL.
in association with human habitation. Bavaya
cyclura is generally more gracile and boldly pat-
terned than are B. crassicollis and B. montana Bavayia montana Roux
(Fig. 2). A striped individual (CAS 165 885) as Bavayia cyclura montana Roux, 1913:88. Type locality: “Mount
collected with many typically patterned ones— Ignambi, altit. 700-800 m, prés Tao, altit. 400 m, Mont we thus doubt that this pattern is of significance. Canala, altit. 700 m, Coindié, altit. 250 m, Ni, altit. 250 m.”
28 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 47, No. 2
Lectotype locality: ““Mount Ignambi, 700-800 m, Neu-Ka- ledonien”’ (restricted by Kramer 1979). Bavayia montana Sadlier, 1989:365.
MATERIAL EXAMINED (1 specimen).— Locality 12: 3 Oct. 1984 (CAS 157694).
REMARKS.—One individual (Fig. 2) was col- lected inside a moist log at about 200 m elevation on Mt. Koyaboa. In life the specimen displayed pale speckling over the basic pattern of indistinct dorsal blotches, and had a canary-yellow venter.
Bavayia sauvagii (Boulenger)
Lepidodactylus sauvagii Boulenger, 1883:122. Type locality: New Caledonia by implication (the original description seems to refer to the species now recognized as Bavayia sauvagii; however, the purported holotype is conspecific with B. cy- clura and possibly is not the specimen designated by Bou- lenger).
Bavayia sauvagei Roux, 1913:91.
MATERIAL EXAMINED (299 specimens).—Locality 5: 6 Oct. 1984 (CAS 157706, 157711-4); Locality 11: 4 Oct. 1984 (CAS 157705, 157707-10); Locality 12: 3-5 Oct. 1984 (CAS 157914— 61), 8 Oct. 1984 (CAS 165903-10), 11-12 Oct. 1984 (CAS 157962-70, 165881-3, 165888-9, 165893-4), 17 Oct. 1984 (CAS 165880), 1-9 June 1984 (CAS 158430-508, 165875-6), 29 Dec. 1985 (CAS 15951325), 1 Jan. 1986 (CAS 159528- 43), 8-9 Jan. 1986 (CAS 159555-61, 159566-7), 13-15 Dec. 1986 (CAS 162193-200, 162213-8); Locality 27: 11-12 Dec. 1986 (CAS 162186-92); Locality 31: 25 May 1985 (CAS 158378), 15-16 Jun. 1985 (CAS 158379-83), 11 Dec. 1986 (CAS 162184-S); between localities 32 and 39: 21 Oct. 1984 (CAS 157715); Locality 35: July-Aug. 1940 (CAS 80823-36, 8083841, 80843-8, 80851-2, 80854-8), 17 Dec. 1986 (CAS 162225), 23 Dec. 1986 (CAS 162240-3); Locality 39: July— Aug. 1940 (CAS 80859-62); Locality 40: 18-20 Oct. 1984 (CAS 157716-8), 19-22 May 1985 (CAS 158321-31), 15 June 1985 (CAS 158332), 4 Jan. 1986 (CAS 159552-3), 18-20 Dec. 1986 (CAS 162228-32, 162235); Locality 41: 30 May 1985 (CAS 158386-8); Locality 43: exch. from NMBA, 1914 (CAS 38826). CAS 80837 exchanged to G. Pasteur.
REMARKS. —Like Bavayia cyclura, B. sauvagii is extremely variable in pattern, and to a lesser extent, body form. In general, the specimens from the west coast of New Caledonia are larger (max. SVL 62 mm, CAS 162184) than those from the east coast. Most specimens were collected during daylight hours from under rocks in humid forest (Bauer and DeVaney 1987), although some were collected at night on bushes and érees. The species is broadly insectivorous with crickets making up the bulk of the diet on Mt. Koyaboa (Bauer and DeVaney 1987). Bavayia sauvagii is broadly sympatric and syntopic with Nactus pelagicus throughout the range of the former and is also frequently found in association with Marmoros- phax tricolor.
Eurydactylodes vieillardi (Bavay)
Platydactylus Vieillardi Bavay, 1869:10. Type locality: ‘““Houa- gape” [=Wagap, Nouvelle-Calédonie].
Eurydactylodes vieillardi Wermuth, 1965:30.
MATERIAL EXAMINED (1 specimen).—Locality 36: 10 May 1985 (CAS 158556).
REMARKS. — Little is known about this arboreal species. It occurs in the southern lateritic region of New Caledonia and in regions of edaphic vege- tation throughout the remainder of the island.
Hemidactylus frenatus Duméril and Bibron
Hemidactylus frenatus Duméril and Bibron, 1836:366. Type locality: ““l’ Afrique australe, et. . . tout l’archipel des grandes Indes” (restricted to “Java” by Loveridge 1947).
MATERIAL EXAMINED (28 specimens).—Locality 11: 10 Oct. 1984 (CAS 157738-40), 2-4 June 1985 (CAS 158551-2); Lo- cality 14: 1 Oct. 1984 (CAS 157737); Locality 37: 27 Sept. 1984 (CAS 157727-36), 16 May—16 June 1985 (CAS 158289, 158294-303); Locality 38: June 1985 (CAS 172736).
REMARKS. — These geckos are extremely com- mon in urban habitats in New Caledonia. They are also present in much lower densities in areas of native vegetation. This species was not re- corded by Roux (1913), nor was it collected by F. X. Williams in 1940. As in other parts of the Pacific, Hemidactylus frenatus has spread swiftly in recent years. It was not listed among the Hawaiian fauna by Oliver and Shaw in 1953, but it has established itself since being introduced in the 1950s (Hunsaker 1966; Hunsaker and Breese 1967). By 1978 it was the most common gecko in the islands, supplanting H. garnotii from both urban and native habitats (McKeown 1978). Likewise, H. frenatus was not recorded in Fiji by Pernetta and Watling (1978), but by 1985 it had displaced H. garnotii in edificarian habitats (Gib- bons 1985). In New Caledonia, as in Hawaii, its introduction may have been associated with troop movement during World War II (Hunsaker and Breese 1967), and has certainly been facilitated by increased post-war contact with neighboring islands. Hemidactylus frenatus is approximately the same size as H. garnotii (max. SVL 60 mm [CAS 158552] vs. 61 mm [CAS 162223] for H. garnotii) and appears to outcompete it in natural and disturbed situations (Hunsaker and Breese 1967). In the Pacific the two species are known to be sympatric only in Hawau, the Marquesas, and New Caledonia, and it appears likely that the spread of H. frenatus, especially on smaller
BAUER AND VINDUM: NEW CALEDONIAN HERPETOFAUNA 29
islands, may cause local extinctions of H. gar- notil.
Hemidactylus garnotii Duméril and Bibron
Hemidactylus Garnotii Duméril and Bibron, 1836:368. Type locality: “Vile de Taiti.”
MATERIAL EXAMINED (4 specimens).—Locality 9: 15 Dec. 1986 (CAS 162222-3); Locality 37: July—Aug. 1940 (CAS 80876, 80878).
REMARKS.— Although formerly widespread in New Caledonia (Bavay 1869; Roux 1913), Hemidactylus garnotii appears to have retreated in the face of expansion by H. frenatus. Extensive searches in Nouméa indicate that the species has become extinct in the area since F. X. Williams’s visit in 1940. The specimens collected in 1986 were active at night on forest tree trunks along with Nactus pelagicus and Lepidodactylus lugu- bris. The patchy distribution of this species in the Pacific (Ineich 1982; Blanc et al. 1983; Crom- bie and Steadman 1988) suggests that its spread has been mediated by human movements, and that its success on small islands is dependent on the absence of competitors, particularly H. fre- natus.
Hemiphyllodactylus typus Bleeker
Hemiphyllodactylus typus Bleeker, 1860:327. Type locality: “‘Agam, Sumatra” [Indonesia] (see Kluge 1968 for a discus- sion of the type locality).
Platydactylus crepuscularis Bavay, 1869:8. Type locality: Nou- velle-Calédonie by implication.
MATERIAL EXAMINED (2 specimens). — Locality 38: June 1985 (CAS 172739-40).
REMARKS. — Little is known about the biology of this species in New Caledonia. It is rarely encountered and appears to be restricted to areas of undisturbed vegetation.
Lepidodactylus lugubris (Duméril and Bibron)
Platydactylus Lugubris Duméril and Bibron, 1836:304. Type locality: “Vile d’Otaiti.”
Lepidodactylus lugubris Fitzinger, 1843:98.
Gymnodactylus Caudeloti Bavay, 1869:13. Type locality: “Nouvelle-Calédonie, surtout dans le Nord.”
MATERIAL EXAMINED (23 specimens).— Locality 9: 10-12 Oct. 1984 (157741-6), 14-15 Dec. 1986 (CAS 162210, 162224): Locality 11: 6 June 1985 (CAS 159498), 8 June 1985 (CAS 158553-4), 8 Jan. 1986 (CAS 159511), 13-15 Dec. 1986 (CAS 162201-2, 162211); Locality 37: July-Aug. 1940 (CAS 80872- 5, 80877), 17 May 1985 (CAS 158290); Locality 38: June 1985 (CAS 172737-38).
REMARKS. — Lepidodactylus lugubris is the most widely distributed gecko in the Pacific Basin. In New Caledonia it is common in and around houses and in lowland vegetation including hu- mid forest and palm stands. In situations of co- occurrence with Hemidactylus frenatus, the smaller L. lugubris (max. SVL 44 mm, CAS 162201) is relatively secretive, foraging near shelter and on the periphery of the area of Hemi- dactylus activity. This situation parallels that seen in areas of sympatry in Hawaii (Hunsaker and Breese 1967). As in most areas of its range (Cue- llar and Kluge 1972; Ineich 1982, 1988), the New Caledonian populations of L. /ugubris appear to be all female. Adults collected in October con- tained enlarged ovarian or oviductal eggs, and communal egg clutches of up to nine eggs were observed under bark in December 1986. All specimens, regardless of date of collection, pos- sessed enlarged extracranial endolymphatic sacs clearly visible through the skin.
Nactus pelagicus (Girard)
Gymnodactylus Arnouxii Duméril, 1851:44. Type locality: ““Nouvelle-Zélande” (sic) (see Kluge 1983 for a discussion of probable locality).
Heteronota pelagica Girard, 1857:197. Type locality: “Feejee and Navigator Islands” [=Fiji and Samoa].
Nactus pelagicus Bauer and DeVaney, 1987:353.
MATERIAL EXAMINED (51 specimens).—Locality 9: 11 Oct. 1984 (CAS 157769), 1 Jan. 1986 (CAS 159544-S, 159547-9), 8 Jan. 1986 (CAS 159562-5); Locality 12: 3-11 Oct. 1984 (CAS 157758-67), 2-9 June 1985 (CAS 158509-21, 158543), 1 Jan. 1986 (CAS 159526-7), 8 Jan. 1986 (CAS 159554), 15 Dec. 1986 (CAS 162212); Locality 30: 25 Oct. 1940 (CAS 80882-3, 80885-6); Locality 40: 18 Oct. 1984 (CAS 157768), 20 May 1985 (CAS 158333-4), 11 June 1985 (CAS 158335), 18-20 Dec. 1986 (CAS 162226-7, 162233-4, 162236). CAS 80884 exchanged to G. Pasteur.
REMARKS.— Kluge (1983), when erecting the genus Nactus, placed N. pelagicus into the syn- onymy of N. arnouxii. However, the use of the older but unused name arnouxii disrupts stabil- ity and causes confusion in usage (Zug 1985a, 1989). We therefore use the specific epithet pe- lagicus. Nactus pelagicus (max. SVL 69 mm, CAS 158520) is widespread in New Caledonia but rather secretive. It was collected under stones during the day, frequently in dry streambeds (Bauer and DeVaney 1987), and low (<1 m) on the trunks of trees at night. The species has a broad insectivorous diet, similar in most respects to that of sympatric Bavayia sauvagii. All adult
30 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 47, No. 2
specimens in the collection are female, and Mo- ritz (1987) has demonstrated that the same is true of populations from southern Vanuatu and the eastern Pacific, suggesting a parthenogenetic species (2N = 35) of hybrid origin.
Rhacodactylus auriculatus (Bavay)
Platydactylus auriculatus Bavay, 1869:6. Type locality: ‘““Mont d’Or” [=Mont Dore, Nouvelle Calédonie].
Ceratolophus hexaceros Bocage, 1873a:205. Type locality: “Nouvellé Calédonie.”
Rhacodactylus auriculatus Boulenger, 1883:127.
MATERIAL EXAMINED (38 specimens).— Locality 29: 19 Oct. 1984 (CAS 157684); Locality 32: 18-19 Oct. 1984 (CAS 157679, 157681-2), 20 May 1985 (CAS 158923), 11 June 1985 (CAS 158925); on road between localities 32 and 39: 18-21 Oct. 1984 (CAS 157676-8, 157680, 157683, 165858-9, 165892, 165901), 18-20 May 1985 (CAS 158919-20, 158922), 4 Jan. 1986 (CAS 159512), 20-22 Dec. 1986 (CAS 162179-83); Lo- cality 33: 19 Oct. 1984 (CAS 165891); Locality 40: 20 May 1985 (CAS 158921, 158924), 12-14 June 1985 (CAS 165860, 165902), 2-5 Jan. 1986 (CAS 165895-900), 19 Dec. 1986 (CAS 162178); Locality 41: 30 May 1985 (CAS 158389 -90).
REMARKS. — Rhacodactylus auriculatus is en- demic to southern New Caledonia. Within this region it has been collected widely. By night the animals are active in trees (generally at heights of <3 m) and on the ground. Most CAS speci- mens were collected along dirt roads from dusk until about 2100 hr. The diet of this species in- cludes flowers (Bavay 1869), lizards (Bauer and DeVaney 1987), and probably insects and fruits as well. Females with shelled eggs were collected in October, December—January, and May-June; in captivity R. auriculatus breeds year round. This is the smallest species of Rhacodactylus; the largest specimen examined, CAS 162181, was a female of SVL 118 mm.
Rhacodactylus chahoua (Bavay)
Platydactylus chahoua Bavay, 1869:3. Type locality: ““Kanala, Lifou” (sic, see Bauer 1985a for a discussion of this locality). Neotype locality: “Vallée d’ Amoa, near St. Thérése, approx. 15 km NW of Poindimié, New Caledonia”’ (neotype desig- nated by Bauer 1985a).
Rhacodactylus chahoua Boulenger, 1883:125.
MATERIAL EXAMINED (4 specimens). — Locality 9: 1 Jan. 1986 (CAS 167764), 13 Dec. 1986 (CAS 162177); Locality 13: 6 Apr. 1984 (CAS 156691, 156692 [NEOTYPE)).
REMARKS.— Bauer (1985a) designated a neo- type to replace the lost type of Bavay (1869), which bore the locality ““Kanala, Lifou’’ (prob- ably a lapsus for Canala, New Caledonia). All specimens were collected at heights of 2-5 m in trees along the Amoa River in eastern New Cal- edonia. Rhacodactylus chahoua appears to be ac-
tive after dusk on the trunks and branches of trees of moderate size (maximum trunk diameter ca. 2 m).
Rhacodactylus leachianus (Cuvier)
Ascalabotes Leachianus Cuvier, 1829:54. Type locality: not given.
Rhacodactylus Leachianus Bocage, 1873a:201.
Rhacodactylus Aubrianus Bocage, 1873a:202. Type locality: “Nouvelle Calédonie.”
MATERIAL EXAMINED (9 specimens).— Locality 6: July-Aug. 1940 (CAS 80881); Locality 7: Nov. 1985 (CAS 159510); Lo- cality 13: 6 Apr. 1984 (CAS 156690); Locality 15: 28 Aug. 1984 (CAS 165890); Locality 27: 11 Dec. 1986 (CAS 172734); Locality 35: July-Aug. 1940 (CAS 80879-80), 12 Dec. 1986 (CAS 165857, 172735).
REMARKS. — This is the largest living species of gecko, attaining 245 mm SVL (CAS 165890). Animals were collected from the trunks and low- er branches of trees in humid forests throughout the wetter areas of New Caledonia, most fre- quently those immediately bordering streams or rivers. Males are much more commonly cap- tured and observed at lower heights on trees, whereas females generally remain in the canopy. In addition to specimens captured, many more R. leachianus were observed at heights of 7-30 m in trees where they could not be collected. In the wild this species eats birds, insects, and fruit, and in captivity will also eat mice and other small mammals (Bauer and DeVaney 1987).
Rhacodactylus sarasinorum Roux
Rhacodactylus sarasinorum Roux, 1913:99. Type locality: ““Forét de Prony (env. 100 m. d’altitude)” [New Caledonia].
MATERIAL EXAMINED (1 specimen).—Locality 32: 18 Oct. 1984 (CAS 157675).
REMARKS. — The single specimen was collected at ca. 2000 hr on the branches of a small tree in an area where both Rhacodactylus auriculatus and R. leachianus are common. The specimen is large (125 mm SVL) and extremely dark, lack- ing the spots and nuchal collar common in ju- veniles and smaller adults (Henkel 1987).
Family ScINCIDAE
Caledoniscincus atropunctatus (Roux)
Lygosoma austro-caledonicum atropunctatum Roux, 1913:117. Type locality: ‘“‘en Nouvelle-Calédonie et aux trois iles Loy- alty.”’ Lectotype locality: ““Outbatche, Neu-Kaledonien” (re- stricted by Kramer 1979).
Caledoniscincus atropunctatus Sadlier, 1986:42.
BAUER AND VINDUM: NEW CALEDONIAN HERPETOFAUNA 31
MATERIAL EXAMINED (16 specimens).— New Caledonia: ex- change from NHMB, 1914 (CAS 38828 [PARALECTO- TYPE]); Locality 12: 3 Oct. 1984 (CAS 158073), 4 Oct. 1984 (CAS 158080), 1 June 1985 (CAS 158542), 1-2 Jan. 1986 (CAS 159639-40), 15 Dec. 1986 (CAS 162311); Locality 31: 28 Dec. 1985 (CAS 159616), 11 Dec. 1986 (CAS 162273-4); Locality 40: 4 Jan. 1986 (CAS 159643-5), 19-20 Dec. 1986 (CAS 162332, 162347, 162363).
REMARKS.— Museum specimens of species of this genus are often difficult to distinguish from one another, as the primary diagnostic features relate to coloration, which is extremely variable. The specimens listed above include all those that are unambiguously assignable to Caledoniscin- cus atropunctatus. Although this small species (max. SVL 51 mm, CAS 162274) is broadly sym- patric and syntopic with C. austrocaledonicus, and to a lesser extent C. festivus, it is more often associated with forests than is C. austrocaledoni- cus (Sadlier 1986). All CAS specimens come from dense forest or forest-edge habitats. In Vanuatu this species is primarily insectivorous and sim- ilar to C. austrocaledonicus in the breadth of prey categories taken (Medway and Marshall 1975).
Caledoniscincus austrocaledonicus (Bavay)
Lygosoma Austro-Caledonica Bavay, 1869:21. Type locality: “partout en Calédonie et dans les iles Loyalty.” Neotype locality: “4 km from summit of Mt. Aoupinie by road, New Caledonia, 21°11'S, 165°16’E” (designated by Sadlier 1986).
Euprepes haplorhinus Giinther, 1872:419. Type locality: ““New Caledonia.”
Lygosoma austro-caledonicum dorsovittatum Roux, 1913:118. Type locality: ““du nord au sud de la Calédonie et sur les 3 iles Loyalty.” Lectotype locality: ““Hienghiene (sic) Neu-Ka- ledonien” (restricted by Kramer 1979).
Leiolopisma dorsovittatum bodei Borner, 1980:8. Type locality: “unter einerm Baumstamm auf einer trockenen Wiese bei Kutio auf der Ile des Pins siidlich von Neu Kaledonien.”
Caledoniscincus austrocaledonicus Sadlier, 1986:37.
MATERIAL EXAMINED (727 specimens). — New Caledonia: ex- change from NHMB, 1914 (CAS 38829 [PARALECTOTY PE of Lygosoma austrocaledonicum dorsovittatum Roux, 1913], 38830); Locality 1: 2 Oct. 1984 (CAS 158072); Locality 2: 9 Oct. 1984 (CAS 158081); Locality 3: 10-11 Jan. 1986 (CAS 159648-55); Locality 4: 13 Oct. 1984 (CAS 158048-59, 158084); Locality 5: 6-7 Oct. 1984 (CAS 157984-8013); Locality 6: July—Aug. 1940 (CAS 80793-5); Loclaity 7: 10 Oct. 1984 (CAS 158014~34); Locality 10: 12 Oct. 1984 (CAS 158045-7, 158082- 3); Locality 11: 12 Oct. 1984 (CAS 158035-44), 2 Jun. 1985 (CAS 158875); Locality 12: 3 Oct. 1984 (CAS 158079), 6 June 1985 (CAS 158546-7), 8 Jan. 1986 (CAS 159646); Locality 14: 1 Oct. 1984 (CAS 157983); Locality 16: 29 Mar. 1984 (CAS 169524); Locality 18: 17 Oct. 1984 (CAS 158064-8, 158096-7); Locality 19: 21 Oct. 1984 (CAS 158086-7); Lo- cality 22: 13-14 Jan. 1986 (CAS 159656-85); Locality 23: 26 May 1985 (CAS 158895-918); Locality 24: 20 Oct. 1984 (CAS 158060); Locality 25: 10 June 1985 (CAS 158867-70); Locality 26: 10 June 1985 (CAS 158866); Locality 28: 18 Dec. 1986
(CAS 162316); Locality 31: 25 May 1985 (CAS 158384-5), 28-29 Dec. 1985 (CAS 159611-5, 159618-37), 10-12 Dec. 1986 (CAS 162244-5, 162271-2, 162275-90); Locality 32: 30 Sept. 1984 (CAS 157974-82, 158069-71, 158077-8), 3 Oct. 1984 (CAS 158074-6), 18 Oct. 1984 (CAS 158061-3, 158088- 95); Locality 35: July—Aug. 1940 (CAS 80770, 807724, 80776, 80778-82, 80784, 80786-92); Locality 37: 18-20 May 1985 (CAS 158291-3), 28 Dec. 1985 (CAS 159610), 10 Dec. 1986 (CAS 162262); Locality 38: June 1985 (CAS 172743-46); Lo- cality 40: 18-19 Oct. 1984 (CAS 158085, 158098), 18-23 May 1985 (CAS 158557-673), 11-14 June 1985 (CAS 158674— 863), 4 Jan. 1986 (CAS 159641-2), 18-22 Dec. 1986 (CAS 162318-21, 1620323-31, 162333-51, 16235462, 162364— 461); Locality 41: 30 May 1985 (CAS 158393-8); Locality 42: 24 Oct. 1940 (CAS 80887-8). CAS 80785, 80796-7 missing from collection.
REMARKS.— As mentioned above, the current means of distinguishing species of Caledoniscin- cus from one another is not satisfactory. Some specimens of C. austrocaledonicus approach the coloration patterns of each of its congeners, as well as exhibiting a wide array of intermediate patterns. The intermediate size of this species (max. SVL 55 mm, CAS 158561) places indi- viduals well within the range of at least some individuals of each of the other Caledoniscincus species. The ventral coloration in life of a series of C. austrocaledonicus collected on 12 June 1985 at Locality 40 exemplifies the intrapopulational variability (salmon, orange, yellow, bronze, cream, grey, white, and brown venters were re- corded). In other populations, such as that of Plage de Poé, dorsal and ventral coloration are more or less uniform. The chevron marks on the tail employed in the accompanying key are oc- casionally absent from specimens with original tails, and frequently lacking on regenerated tails. An electrophoretic analysis of variation within and between putative C. austrocaledonicus pop- ulations may clarify systematic relationships in this group. At least two species are presently in- cluded under this name (R. A. Sadlier, pers. comm.).
Caledoniscincus austrocaledonicus is the most common lizard in New Caledonia. It appears to be excluded from dense forest but is common in clearings, forest margins, grassy fields, road- banks, and nearshore and beach habitats from Boat Pass to the Ile des Pins as well as on the Loyalty Islands.
Bauer and DeVaney (1987) reported a wide range of prey items in the diet of C. austroca- ledonicus (as Leiolopisma austrocaledonicum), including large numbers of crickets and cater- pillars. They found some evidence for the pref-
32 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 47, No. 2
erential feeding of juveniles on collembolans and other very small prey. Beach-wrack-dwelling specimens feed to a great extent on amphipods living in the sand in the splash zone. The species, in turn, is prey to a wide variety of saurian and avian predators, as well as Litoria aurea and probably predaceous insects.
Caledoniscincus festivus (Roux)
Lygosoma austrocaledonicum intermedium Roux, 1913:119. Type locality: ‘‘Nouvelle-Calédonie: Canala et environs (al- tit. 300 m), Coindé, Vallée de la Ngoi.”’ Lectotype locality: ““Ciu oberhalb Canala, Neu-Kaledonien (restricted by Kra- mer 1979).
Lygosoma austro-caledonicum festivum Roux, 1913:120. Type locality: ‘“‘“Nouvelle-Calédonie: Coné, environs de Canala, La Foa, Vallée de la Ngoi, Yaté.”” Lectotype locality: ““Cana- la, Neu-Kanedonien” (restricted by Kramer 1979).
Caledoniscincus festivus Sadlier, 1986:45.
MATERIAL EXAMINED (8 specimens). — Locality 5: 6 Oct. 1984 (CAS 157971-3); Locality 12: 9 Jan. 1986 (CAS 159647); Lo- cality 26: 10 June 1985 (CAS 158864—5); Locality 31: 28-29 Dec. 1985 (CAS 159617, 159638).
REMARKS.—This large species of Caledoni- scincus (max. SVL 68 mm, CAS 157972) was encountered in open grassy habitat (Locality 26) and in clearings and grassy patches at medium (Locality 12) to high elevations (Locality 5). Campement Vanhalie is a northern record for the species and is very close to Mt. Panié, the only known locality of the very similar C. orestes. These skinks appear to be generalist carnivores; CAS 158864 (recorded by Bauer and DeVaney 1987 as Leiolopisma austrocaledonicum) had a 2.0 g Caledoniscincus austrocaledonicus in its stomach.
Cryptoblepharus novocaledonicus (Mertens) Cryptoblepharus boutonii novo-caledonicus Mertens, 1928:88. Type locality: ““Hienghiéne [=Hiénghéne] Neukaledonien.” Cryptoblepharus novocaledonicus Sadlier, 1986:53. MATERIAL EXAMINED (5 specimens).— Locality 6: 1911 (ex- changed from NMBA, 1921) (CAS 53991); Locality 24: 20
Oct. 1984 (CAS 157725-6); Locality 32: 29 Sept. 1984 (CAS 157724); Locality 35: July-Aug. 1940 (CAS 80783).
REMARKS.—Despite the high densities and widespread occurrence in nearshore habitats of its congener, Cryptoblepharus poecilopleurus, on the islands to the north and east of New Cale- donia (Loveridge 1945), C. novocaledonicus ap- pears to be greatly outnumbered by members of the genus Caledoniscincus over most of its range. CAS 157725-6 from the Baie des Tortues on the central west coast confirm Sadlier’s (1986) as-
sumption that the absence of C. novacaledonicus from the west of the island was a collecting ar- tifact. The largest CAS specimen (157725), 42 mm SVL, is an adult female carrying one ovi- ductal egg.
Graciliscincus shonae Sadlier
Graciliscincus shonae Sadlier, 1986:12. Type locality: “4 km along the Mt Gouemba road from turnoff 3 km south of La Fause Yaté Bridge, New Caledonia, 22°09’S, 166°54’E.”
MATERIAL EXAMINED (1 specimen).—Locality 31: 28 Dec. 1985 (CAS 159576 [PARATYPE]).
REMARKS. —CAS 159576 is one of two known in this monotypic genus. It was collected under a large stone in warm rainy weather.
Leiolopisma nigrofasciolatum (Peters)
Lygosoma (Mocoa) nigrofasciolatum Peters, 1870:435. Type locality: ““Neu Kaledonien.”
Lygosoma arborum Bavay, 1869:19. Type locality: Nouvelle- Calédonie by implication. Neotype locality: ““Mouac Island off Poum, New Caledonia, 20°13’S, 164°00’E” (designated by Sadlier 1986).
Lygosoma Deplanchei Bocage, 1873b:229 (non Lygosoma De- planchei Bavay, 1869). Type locality: ““Nouvelle Calédonie.” Neotype locality: ““Outbache, New Caledonia, 20°26’S, 164°38’E” (designated by Sadlier 1986).
Leiolopisma nigrofasciolatum Greer, 1974:17.
MATERIAL EXAMINED (21 specimens).— New Caledonia, July— Aug. 1940 (CAS 80890); Locality 6: July-Aug. 1940 (CAS 80801); Locality 14: 29 Mar. 1984 (CAS 156757); Locality 32: 29-30 Sept. 1984 (CAS 157719-21); Locality 35: July—Aug. 1940 (CAS 80809-1 1); Locality 40: 18 Oct. 1984 (CAS 157722- 3), 23 May 1985 (CAS 158344), 11 June 1985 (CAS 158345), 14 June 1985 (CAS 158346), 4-5 Jan. 1986 (CAS 159573-5), 21 Dec. 1986 (CAS 162463-5); Locality 44: exchanged from NMBH, 1914 (CAS 38825).
REMARKS.—Bavay’s (1869) name Lygosoma arborum has priority over L. nigrofasciolatum Peters, 1870. However, the former name has not been used by subsequent authors and we regard it as a nomen oblitum.
Although generally regarded as arboreal (Meier 1979; Sadlier 1986), Leiolopisma nigrofasciola- tum has been collected in a wide variety of open habitats. Many of the specimens from Goro were collected in vertical crevices in boulders, includ- ing several in the splash zone. One L. nigrofas- ciolatum (CAS 158344) was found sharing the same crevice with a Laticauda colubrina (CAS 158309). The Goro and Touaourou records are the first published for the southeastern coast of the New Caledonian mainland. The species preys upon cockroaches and other large insects (Bauer and DeVaney 1987), as well as smaller skinks
BAUER AND VINDUM: NEW CALEDONIAN HERPETOFAUNA 33
CM
STS ise po | vl tiettt attire
tae Tan
FicureE 3. Marmorosphax euryotis (CAS 80889) from the Ile des Pins showing dorsal color pattern.
(Roux 1913). The largest specimen is CAS 38825 (112 mm SVL), containing 10 oviductal eggs, presumably collected in December, 1911 (Roux 1913). This is the only CAS specimen lacking a solid dark brown median stripe on the nape and shoulders.
Leiolopisma novaecaledoniae (Parker)
Lygosoma (Leiolopisma) novae-caledoniae Parker, 1926:493. Type locality: “Upper Houailou River, New Caledonia.” Leiolopisma novaecaledoniae Sadlier, 1986:21.
MATERIAL EXAMINED (1 specimen).—Locality 6: July—Aug. 1940 (CAS 80799).
REMARKS.—Leiolopisma novaecaledoniae is known from only a few specimens and localities in New Caledonia. CAS 80799 (SVL 44 mm) fills in part of the gap between the eastern coastal records of Oue Camme and Houailou (Sadlier 1986). It closely matches the squamation and coloration of specimens in previously published accounts.
Leiolopisma steindachnerii (Bocage)
Lioscincus Steindachnerii Bocage, 1873b:228. Type locality: “Nouvelle Calédonie.”’ Neotype locality: ““Mt Panie (500 m), New Caledonia, 20°36’S, 164°46’E” (designated by Sadlier 1986).
Lygosoma (Liolepisma) steindachneri Roux, 1913:129.
Leiolopisma steindachneri Greer, 1974:17.
MATERIAL EXAMINED (1 specimen).—Locality 17: 13 Dec. 1978 (CAS 162121).
REMARKS.—Received in exchange from Aus- tralian Museum, original number AMS 77723.
Marmorosphax euryotis (Werner)
Lygosoma euryotis Werner, 1909:271. Type locality: “Ile des Pins, Nouvelle-Calédonie.” Marmorosphax euryotis Sadlier, 1986:36.
MATERIAL EXAMINED (1 specimen).—Locality 41: 24 Oct. 1940 (CAS 80889).
REMARKS. — This species 1s evidently quite rare, and Sadlier (1986) examined only two speci- mens. CAS 80889 comes from the type locality and matches the holotype in squamation and approximate size (SVL 36 mm). The coloration of this specimen differs in some respects from that described by Sadlier (1986) and exhibits some features not reported for either of the two known specimens, particularly a distinct, dark nuchal band (see Fig. 3).
Marmorosphax tricolor Bavay
Lygosoma tricolor Bavay, 1869:17. Type locality: Nouvelle- Calédonie by implication. Neotype locality: ‘“‘summit of Mt Aoupinie (1,086 m), New Caledonia, 21°11'S, 165°16’E” (designated by Sadlier 1986).
Marmorosphax tricolor Sadlier, 1986:32.
34 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 47, No. 2
MATERIAL EXAMINED (95 specimens). — Locality 6: July—Aug. 1940 (CAS 80800); Locality 9: 11 Oct. 1984 (CAS 157756); Locality 12: 3-10 Oct. 1984 (CAS 157747-55), 12 Oct. 1984 (CAS 157757), 1-9 June 1985 (CAS 158399-429, 158544-5), 29 Dec. 1985-1 Jan. 1986 (CAS 159590-8), 8 Jan. 1986 (CAS 159600-5), 15 Dec. 1986 (CAS 162306-10); Locality 21: 1913 (exchanged from NHMB, 1921) (CAS 53988); Locality 31: 25 May 1985 (CAS 158367-71), 15-16 June 1985 (CAS 158372- 7), 28 Dec. 1985 (CAS 159586, 159588), 10-11 Dec. 1986 (CAS 162258-61, 162263-4); Locality 35: July-Aug. 1940 (CAS 80802-8); Locality 40: 5 Jan. 1986 (CAS 159599), 18 Dec. 1986 (CAS 162317), 19 Dec. 1986 (CAS 162322).
REMARKS.—This is the most common skink in heavily forested areas of the wetter regions of New Caledonia. These moderately large skinks (max. SVL 66 mm, CAS 162264) were often found under large stones, where they feed on ants, scorpions, and terrestrial arthropods (Bauer and DeVaney 1987). Juveniles of this viviparous species were collected only during December and January 1985/86 and measured 22-25 mm SVL, confirming Sadlier’s (1986) observations.
Nannoscincus gracilis (Bavay)
Lygosoma gracilis Bavay, 1869:24. Type locality: Nouvelle- Calédonie by implication.
Mocoa micropus Giinther, 1872:420. Type locality: “‘Feejee Islands” (amended to New Caledonia in British Museum [Natural History] catalogue).
Nannoscinicus gracilis Sadlier, 1986:58.
MATERIAL EXAMINED (26 specimens).— Locality 12: 4—5 Oct. 1984 (CAS 157685-9), 1-9 June 1985 (CAS 158522-3, 158527- 41), 1 Jan. 1986 (CAS 159608-9), 13 Dec. 1986 (CAS 162305) 15 Dec. 1986 (CAS 162313).
REMARKS.— This is the largest (max. SVL 41 m, CAS 162305) and most widespead member of the genus. All material was obtained in very moist conditions under logs and stones (Bauer and DeVaney 1987). The species appears partic- ularly prone to dehydration. Like all members of the genus, Nannoscincus gracilis appears to be at least partially fossorial, and it frequently at- tempts to escape by burrowing rapidly into the substrate. The largest individual, collected in mid- December, contained two very large eggs filling most of the body cavity. This confirms Sadlier’s (1986) report and suggests that the rainy Decem- ber—January period and the periods immediately bracketing it are the periods of breeding and/or birth/hatching for many New Caledonian lizard species.
Nannoscincus greeri Sadlier
Nannoscincus greeri Sadlier, 1986:63. Type locality: “lower east slope of Mt Koyaboa, Poindimié, ca. 20°56'S, 165°20’E” [New Caledonia].
MATERIAL EXAMINED (12 specimens).—Locality 12: 12 Oct. 1984 (CAS 157692-3), 2 June 1985 (CAS 15825-6 [PARA- TYPES]), 29 Dec. 1985 (CAS 159606 [HOLOTYPE]), 1 Jan. 1986 (CAS 159607 [PARATYPE)), 15 Dec. 1986 (CAS 162312, 162314—5); Locality 35: July-Aug. 1940 (CAS 80819-21), CAS 158524 [PARATYPE] exchanged to Australian Museum, reg- istered as AMS R-123000.
REMARKS.—This smallest of Nannoscincus species (max. SVL 34 mm, CAS 162312) has been reported only from central eastern and southwestern New Caledonia. Its microhabitat is identical to that of the sympatric N. gracilis. The smallest individual (CAS 162315, 22 mm SVL) was collected in December and differs from larger individuals (as reported by Sadlier 1986) in possessing a prominent light spot on the nape and a broken, pale midvertebral stripe.
The date of collection of the holotype was in- correctly reported as January 1, 1986, by Sadlier (1986).
Nannoscincus mariei (Bavay)
Anotis Mariei Bavay, 1869:29. Type locality: Nouvelle-Calé- donie by implication.
Nannoscincus fuscus Giinther, 1872:421. Type locality: ““Fee- jee Islands” (amended to New Caledonia in British Museum [Natural History] catalogue).
Nannoscincus mariei Sadlier, 1986:56.
MATERIAL EXAMINED (35 specimens).—Locality 27: 11-12 Dec. 1986 (CAS 162297-304); Locality 31: 25 May 1985 (CAS 158358-9), 15-16 June 1985 (CAS 158360-66), 28-29 Dec. 1985 (CAS 159583-5), 10 Dec. 1986 (CAS 162254-7), 11-12 Dec. 1986 (CAS 162291 -S); Locality 35: July-Aug. 1940 (CAS 80813-8). CAS 80812 exchanged to Museum of Comparative Zoology.
REMARKS. — This is another southern New Ca- ledonian endemic and was collected only in ex- tremely moist microhabitats, under rocks and logs, and among the roots of trees. All females of this small skink (max. SVL 39 mm, CAS 158363) collected in December 1986 contained one or two large, elongate eggs.
Nannoscincus slevini (Loveridge)
Lygosoma slevini Loveridge, 1941:193. Type locality: “Mont Canala, New Caledonia.” Nanoscincus slevini Sadlier, 1986:61.
MATERIAL EXAMINED (1 specimen).— Locality 35: July—Aug. 1940 (CAS 80822).
REMARKS.—This species had not been previ- ously recorded from southwestern New Cale- donia. Other specimens are from the region of Canala. The Yahoué locality is unique in sup- porting three species of Nannoscincus—N. greeri, N. mariei, and N. slevini.
BAUER AND VINDUM: NEW CALEDONIAN HERPETOFAUNA 35
Phoboscincus bocourti (Brocchi)
Eumeces Bocourti Brocchi, 1876:95. Type locality ““Nouvelle- Calédonie.” Phoboscincus bocourti Greer, 1974:15.
MATERIAL EXAMINED: No CAS specimens.
REMARKS.—This species is known only from the holotype (SVL 275 mm). The senior author observed what was apparently an individual of this species at ca. 2030 hr on 21 December 1986 just south of the Kuébini River. The sighting may have been of a Phoboscincus garnieri or an un- known species, but the estimated size of the an- imal (ca. 400 mm total length) is consistent with P. bocourti.
According to Greer’s (1974) hypothesis of ly- gosomine relationships, the separation of Pho- boscincus and Tachygia from Eugongylis leaves the last genus paraphyletic. Bohme (1976) con- sidered the generic distinctions proposed by Greer (1974) to be continuously variable in a morpho- cline of constituent taxa and relegated Phobo- scincus spp. as well as a new species, Eugongylus haraldmeieri (Geoscincus haraldmeieri of Sadlier 1986), to the single genus Eugongylus. Present evidence tends to favor BOhme’s treatment of the group, but in the absence of strictly cladistic analysis, we retain the genera Phoboscincus and Geoscincus as lineages putatively independent of Eugongylus.
Sigaloseps deplanchei (Bavay)
Lygosoma Deplanchei Bavay, 1869:23 (non Lygosoma De- planchei Bocage, 1873). Type locality: Nouvelle-Calédonie by implication.
Hinulia tetragonurus Ginther, 1872:420. Type locality: ““Fee- jee Islands” (amended to New Caledonia in British Museum [Natural History] catalogue).
Sigaloseps deplanchei Sadlier, 1986:50.
MATERIAL EXAMINED (65 specimens).—Locality 31: 25 May 1985 (CAS 158871-4, 158876), 15-16 June 1985 (CAS 158877- 94), 28 Dec. 1985 (CAS 159577-82, 159587, 159589), 10-12 Dec. 1986 (CAS 162246-53, 162265-70, 162296); Locality 35: July-Aug. 1940 (CAS 80768-69, 80771, 80775, 80777); Locality 40: 19 Oct. 1984 (CAS 157690-1), 19 May 1985 (CAS 158336-8), 14 June 1985 (CAS 158339-43), 20 Dec. 1986 (CAS 162352-3); Locality 41: 30 May 1985 (CAS 158391-2).
REMARKS. — This small (max. SVL 46 mm, CAS 162246) skink is a southern New Caledonian en- demic. Specimens were collected from under- neath rocks and debris in shaded areas of forest floor in humid montane forest (Localities 31 and 41) and in coastal forest belt (Locality 40). It is intermediate in habitat between Marmorosphax tricolor and Nannoscincus spp. and eats a variety
of small terrestrial prey (Bauer and DeVaney 1987).
Tropidoscincus roehssii (Andersson)
Lygosoma rohssii Andersson, 1908:4. Type locality: ‘““New Cal- edonia.” Tropidoscincus rohssii Sadlier, 1986:30.
MATERIAL EXAMINED (15 specimens).— Locality 38: June 1985 (CAS 172741-42); Locality 40: 22 May 1985 (158349-50), 11-14 June 1985 (CAS 158347-8, 158351-7), 4 Jan. 1986 (CAS 159572), 21 Dec. 1986 (CAS 162462).
REMARKS. — Like all members of this genus, juveniles of Tropidoscincus roehssii have dis- tinctive yellowish dorsolateral and midlateral stripes on the forebody, and a generally reddish- brown tail. Adults are greyish to olive-brown and lack stripes. The venter of adult males is orange- red, fading somewhat on the limbs and tail; the chin and anterior part of the throat are white. Juveniles were most frequently encountered dur- ing May-June 1985 (nine of 11 specimens col- lected during this period). The largest adult spec- imen measured (CAS 158348) was 75 mm SVL. Comments about the ecology of Leiolopisma var- iabile in Bauer and DeVaney (1987) refer to this taxon.
Tropidoscincus variabilis (Bavay)
Tropidolopisma variabilis Bavay, 1869:26. Type locality: Nouvelle-Calédonie by implication. Neotype locality: “Mt. Panie (750 m), New Caledonia, 20°33’S, 164°45’E” (desig- nated by Sadlier 1986).
Tropidoscincus variabilis Sadlier, 1986:24.
MATERIAL EXAMINED (2 specimens).—New Caledonia: ex- changed from NMBA, 1914 (CAS 38827); Locality 6: July- Aug. 1940 (CAS 80798).
REMARKS.—The larger of the two specimens (CAS 38827) measures 67 mm SVL.
Family TYPHLOPIDAE
Ramphotyphlops braminus (Daudin)
Eryx braminus Daudin, 1803:279. Type locality: “Vizagata- pam, Bengale, Inde.” Ramphotyphlops braminus Robb, 1966:676.
MATERIAL EXAMINED (7 specimens).— Locality 37: Nov. 1986 (CAS 162092-4), 12 Dec. 1986 (CAS 162095-6), 18 Dec. (CAS 162097), 22 Dec. 1986 (CAS 162098).
REMARKS. — These are the first collected spec- imens (Bauer 1987) of this widespread, parthe- nogenetic species and range in size from 66 to 161 mm SVL. The only other typhlopid reported
36 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 47, No. 2
Ficure 4. Hydrophis coggeri (CAS-SU 12430). First record from New Caledonian waters.
from the New Caledonian mainland is Rampho- typhlops angusticeps, and only the type of this species, otherwise distributed in the Solomons, bears this locality. It is probable, however, that Peters’s (1877) holotype locality of ““Neu Kale- donien” is an error (Roux 1913; McDowell 1974), most likely a lapsus for Neu Pommeranien or Neu Brittanien, as this taxon is otherwise known only from the Solomon Islands. Discussion with natives in New Caledonia suggests that typhlop- ids probably became established within the last decade and that they are now present in most disturbed areas of both the east and west coasts of the island.
Family HyDROPHIIDAE
Hydrophis coggeri (Kharin)
Leioselasma coggeri Kharin, 1984:1538. Type locality: “Fiji, Port Suva” [translated from original Russian]. Hydrophis coggeri Cogger, 1986:658.
MATERIAL EXAMINED (1 specimen).— Locality 43: Apr. 1945 (CAS-SU 12430).
REMARKS.— This specimen (SVL 1,024 mm) (Fig. 4) represents the first confirmed New Cale- donian record for the species. It is similar to sea snakes from the Coral Sea and the vicinity of Fiji first reported by McDowell (1972) and Cogger (1975) as Hydrophis melanocephalus and later
BAUER AND VINDUM: NEW CALEDONIAN HERPETOFAUNA 37
used by Kharin (1984) in his revision of Leio- selasma. Minton and Dunson (1985) collected specimens probably also referable to this species at the Chesterfield Reefs. We here retain the more familiar generic designation Hydrophis for this species.
Laticauda colubrina (Schneider)
Hydrus colubrinus Schneider, 1799:238 (Linnaeus 1758 based his Coluber laticaudatus partly on this species and partly on the following species, fide Andersson 1899). Type locality: unknown.
Laticauda colubrinus Stejneger, 1907:406.
MATERIAL EXAMINED (8 specimens). — Locality 39: 19-23 May
1985 (CAS 158304—-10); Locality 43: Apr. 1945 (CAS-SU
12438).
REMARKS.—This species is by far the most common nearshore sea snake in New Caledonia. Hundreds of individuals may come ashore, par- ticularly during the period of egglaying (Saint Girons 1964). Most individuals collected were resting in rock crevices at water’s edge during daylight hours. The largest specimen examined is CAS 158306, SVL 860 mm.
Laticauda laticaudata (Linnaeus)
Coluber laticaudatus (part) Linnaeus, 1758:222. Type locality: “Tndiis.”
Laticauda laticaudata Stejneger, 1907:402.
MATERIAL EXAMINED (3 specimens).— Locality 40: 2 Jan. 1986 (CAS 159568); Locality 43: Apr. 1945 (CAS-SU 12441-2).
REMARKS. — This species occurs with Laticau- da colubrina in most New Caledonia nearshore habitats. The specimen collected in 1986 was obtained in 0.5 m of water at ca. 2100 hr and was one of several specimens that were swim- ming towards shore. CAS-SU 12441 (SVL 1,088 mm) is the largest specimen.
BIOGEOGRAPHY OF THE NEw CALEDONIAN HERPETOFAUNA
Roux (1913) first analyzed New Caledonian herpetofaunal characteristics and identified four faunal elements: strictly New Caledonian en- demics, regional endemics (New Caledonia and the Loyalty Islands), Melanesian-Polynesian ele- ments, and cosmopolitan elements. Sternfeld (1920), Sarasin (1925), Burt and Burt (1932), Mertens (1934), Brown (1956), Virot (1956), Gibbons (1985), and Bauer (1986, 1988a), among others have since discussed the biogeographic
affinities of the herpetofauna of the region. Most of these authors stressed the high levels of ende- mism and the contrast provided by New Cale- donia to the trend of Papuan diminution that typifies many of the islands of the central and eastern Pacific, and have focused on the potential faunal ties of New Caledonia to Australia and New Zealand. With an increased knowledge of the historical geology of the region and the phy- logeny of its herpetofauna, it is now possible to begin to redefine the faunal elements of the re- gion.
The marine taxa of New Caledonian waters show little tendency towards endemism, al- though Hydrophis coggeri appears to be restrict- ed to a relatively small area between the Coral Sea and Fiji (Kharin 1984). Most of the sea snakes are primarily inhabitants of the waters of the Australian continental shelf and reach their east- ernmost limits in New Caledonia, or like Hy- drophis ornatus and Laticauda spp., are distrib- uted broadly from western Asia to the western Pacific (Cogger 1975). These taxa favor shallow waters, and their occurrence around New Cale- donia is not surprising given that the Chesterfield Reefs and the northern tip of the Lord Howe Rise provide a shallow-water corridor from the Queensland coast, with only two narrow deeper basins intervening (O.R.S.T.O.M. 1981). The re- maining New Caledonian sea snakes, Emydo- cephalus annulatus and Pelamis platurus, are both more widely distributed in the Pacific, the latter to the coast of the Americas. All of the New Caledonian sea turtles range widely in the trop- ical and subtropical Pacific (Iverson 1986). Al- though turtle populations are conspicuous and easily monitored during nesting periods, the same is not true of the hydrophiids. Some of the sea snake records for New Caledonia are doubtful (see checklist), and more sampling is needed to complete the species list accurately for the region.
Among terrestrial species, Litoria aurea and Hemidactylus frenatus, and almost certainly Ramphotyphlops braminus, have been intro- duced by man into New Caledonia during his- torical times. Their arrival in New Caledonia is easily explained by human transport from ad- jacent, or even distant, regions. Litoria aurea presumably arrived from Australia, although the species was already established in parts of New Zealand by the late 1860s (Bell 1982), and the North Island could conceivably have been the
38 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 47, No. 2
source of at least some of the New Caledonian populations. The advent of rapid and frequent shipping contact in the South Pacific during and after World War II makes the determination of the origin of the other two species more prob- lematic. Although this European-era faunal ele- ment is of little interest to historical biogeogra- phy, it is of prime ecological concern since, at least in the case of H. frenatus, native forms ap- pear to be adversely affected by recent migrants (see Hemidactylus species accounts).
A second element includes Lepidodactylus lu- gubris, Nactus pelagicus, Hemidactylus garnotii, and Hemiphyllodactylus typus—species that are widespread in Oceania or the Pacific and South- East Asia. These are “weedy” species (Bauer 1988a), which by virtue of small size, the cal- careous gekkonine eggshell (Werner 1972; Dun- son 1982), and parthenogenesis (at least in the first three species and possibly the fourth [Kluge and Eckardt 1969; Cuellar and Kluge 1972; Mo- ritz 1987; Ineich 1988]) have been able to col- onize successfully over water.
McCann (1953), Brown (1956), Kluge (1969) and Ineich (1987) outlined suites of biological characteristics conducive to over-water dispersal and colonization. All of the taxa in the first and second faunal-element categories possess most of these features. However, the additional fea- tures of human-aided dispersers as outlined by Kluge (1969) are: (1) undifferentiated from prob- able parent stock; (2) distributed primarily in areas of human settlement; (3) primarily coastal (associated with ports); and (4) dates of intro- duction established do not necessarily hold true for the second faunal element.
Little work has been done on the variability of the wide-ranging taxa of oceanic lizards. Mor- phological differentiation among Polynesian populations of Lepidodactylus lugubris is low (Blanc and Ineich 1985), and there is high his- tocompatibility between Hawaiian and French Polynesian populations (Cuellar 1984), but ge- netic studies have demonstrated that there are at least two bisexual populations and five clonal races in the central South Pacific (Ineich 1987, 1988). In contrast, only a single genetic clone was found by Moritz (1987) among several widely distributed populations of Nactus pelagicus from the western Pacific (including New Caledonia). Genetic homogeneity may be an indication of recent establishment or fragmentation of popu- lations, as suggested for the skink Lipinia noctua
(Zweifel 1979), or it may be a reflection of in- trinsically limited genetic variability, as in clonal lineages such as the parthenogenetic New Cale- donian taxa Lepidodactylus lugubris, Nactus pe- lagicus, Hemidactylus garnotii, and probably Hemiphyllodactylus typus (Cuellar and Kluge 1972; Cuellar 1984; Pasteur et al. 1987). Thus, for parthenogens, Kluge’s (1969) first criterion does not distinguish between human-aided and naturally occurring dispersal.
The distribution of Hemiphyllodactylus, Nac- tus, Lepidodactylus, and Hemidactylus garnotii is not limited to human habitations or to coastal localities in New Caledonia, and at least the for- mer two taxa are restricted to more undisturbed habitats there. Finally, all of these taxa were re- corded by Bavay in 1869, during the earliest stages of French colonization of the island. Their pres- ence in undisturbed areas at this early date argues strongly against dispersal by Europeans, al- though the transport of some taxa by ship is so easy that it cannot be ruled out entirely (Schnee 1901).
Transportation to New Caledonia via human aid in pre-European times remains a strong pos- sibility, and this means of dispersal has been argued for many lizard species in the central and eastern Pacific (Garman 1908; Zweifel 1979; Gibbons 1985; Crombie and Steadman 1988). Crombie and Steadman (1988) dismissed natural rafting as statistically unlikely between the tiny islands of the central Pacific and suggested that Lepidodactylus lugubris arrived in the Cook Is- lands with the Polynesians, while Hemidactylus garnotii, which is spottily distributed and fre- quents edificarian habitats in Polynesia, may have been an import of the post-Polynesia era. Ineich (1982), however, believed that L. /ugubris, at least, may have antedated man in Oceania. While low- ered sea levels in the Pleistocene (and earlier) (Gibbons 1985) may have had only a minor effect on the small, isolated islands of the central and eastern Pacific, their effect in the New Caledo- nian region was marked. More “stepping stones” would have been emergent on routes from Aus- tralia and New Guinea (Gibbons 1985:135, fig. B))
Although the navigational prowess and fre- quent travels of the Polynesians are well estab- lished (see Crombie and Steadman 1988), the pre-European movements of Melanesians to and from New Caledonia are less well understood. Nonetheless, data obtained from Lapita pottery
BAUER AND VINDUM: NEW CALEDONIAN HERPETOFAUNA 39
(White et al. 1988) and largely untapped eth- nobotanical sources (see O.R.S.T.O.M. 1981) clearly indicate a strong cultural link at least 6,000 years old to northern Melanesia, in particular the Solomons and Bismarks, as well as more recent connections via Fiji into Polynesia. Clearly the possibility exists for the human transport of at least some of the second-element taxa. By the same token, however, other seemingly easily transported taxa such as Gehyra mutilata and Emoia cyanura, which occur both southeast and northwest of New Caledonia along the probable routes of prehistoric contact, are absent from New Caledonia. For the present, the origin of the four species of second-element lizards remains prob- lematic, although a reasonably recent (post-Plio- cene) but pre-European introduction, whether human-aided or not, appears likely.
All species in the first two categories are pres- ent in the Loyalty Group as well as on the New Caledonian mainland. Only one species, Cale- doniscincus atropunctatus, occurs in New Cale- donia, the Loyalty Islands, and parts of neigh- boring Vanuatu, and it is part ofa primarily New Caledonian radiation of skinks (Sadlier 1986). It is likely that this species has only recently estab- lished itself in Vanuatu (Bauer 1988b), as the paleoposition of Vanuatu would have precluded easy dispersal prior to 6-8 mybp (Chase 1971).
Native regional endemics (occurring both in New Caledonia and the Loyalty Islands) include the geckos Bavayia crassicollis, B. cyclura, B. sauvagii, and the skinks Caledoniscincus austro- caledonicus, Cryptoblepharis novocaledonicus, Leiolopisma nigrofasciolatum, and Phoboscincus garnieri. All but C. novocaledonicus and P. gar- nieri are representatives of New Caledonian ra- diations, probably of great age (Sadlier 1986; Bauer 1986, in press). Colonization of the Loy- alty Islands from New Caledonia probably took place recently, perhaps during the Pleistocene pe- riod of low sea levels. Most of the Loyalty Islands have probably only been emergent since this time (Bauer 1988a), and being low-lying, have only been able to support those species occurring in coastal forest or marginal strand habitats. Inter- estingly, Bavayia sauvagii is restricted to Maré, the only island with substantial non-coralline areas. This is consistent with this gecko’s pref- erence for rocky substrates (Bauer and DeVaney 1987), otherwise lacking in the Loyalties.
Cryptoblepharis novocaledonicus is a member ofa pan-Pacific genus of 36 morphologically sim-
ilar skinks formerly considered as subspecies of a single variable species (Mertens 1934; Greer 1974). Distinct morphological differentiation within the genus supports the pre-human occur- rence of Cryptoblepharis in the region (Mertens 1931), but its age in the region is uncertain. Like Bavayia sauvagii, C. novocaledonicus favors rocky habitats and has been recorded only from Maré in the Loyalties. Phoboscincus garnieri (with P. bocourti restricted to the New Caledonian main- land) is an offshoot of the genus Eugongylus (Greer 1974), a Papuan lineage. Its ancestors may well have arrived via over-water dispersal, and like the previous taxon, it represents an unam- biguous pre-human arrival. This is supported by the known dispersal abilities of Eugongylus sen- su stricto, but too little is known of the biology of the endemic Caledonian taxa to shed light on the issue.
A few species occur only in the Loyalty Islands or in the Loyalties and Oceania at large, exclud- ing New Caledonia (Emoia cyanura), or in the Loyalties and the Solomon Islands and other Melanesian Outer Arc (Holloway 1979) islands (Candoia bibroni, Gehyra vorax). It appears like- ly that these taxa reached the Loyalties quite re- cently via over-water dispersal from northern and/or eastern source areas, and that there has been insufficient time for speciation to occur in the region. The presence of Candoia in the Loy- alty Islands remains puzzling. The genus shows the typical diminution of species expected for a group of Papuan origin (Gibbons 1985; Bauer 1988b), but if they are correctly identified as boas, their occurrence in the Pacific at all is difficult to understand. Phylogenetic analysis being un- dertaken by Kluge (1988) may shed light on the problem. Only Emoia loyaltiensis, which shares great affinity with the Emoia samoensis group, has apparently speciated in the Loyalties (Brown in press). Rhamphotyphlops willeyi, described from Lifou in the Loyalties, has also been re- corded from the Solomon Islands (McCoy 1980). However, snakes in these two areas may be spe- cifically distinct from one another (McDowell 1974), although they are certainly closely related.
The absence of these Loyalty taxa from New Caledonia has long been recognized (Roux 1913), but its basis is by no means well established. The recent movement of Vanuatu to its present po- sition (Chase 1971) and the emergence of the Loyalty Islands may reduce the problem to one that is analyzable under the equilibrium model
40 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 47, No. 2
of island biogeography (MacArthur and Wilson 1967). The prevailing southern tropical counter current (O.R.S.T.O.M. 1981) would tend to bring propagules to the Loyalties from New Caledonia, while other currents lead from Fiji and Vanuatu toward the Loyalties (Gibbons 1985). Appar- ently the preference of Gehyra vorax and Emoia spp. for vegetated over edificarian habitats has prevented accidental transportation and estab- lishment of these taxa on the mainland, in spite of daily boat and air crossings of cargo and pas- sengers.
Among the remaining taxa, 29 species belong- ing to 8 genera occur only on the New Caledonian mainland or adjacent islands, such as the Ile de Pins, Ile Ouen, and the Iles Belep, which were connected to the mainland during the Pleistocene (Gibbons 1985, fig. 3). These species are limited in vagility by saltwater-permeable eggs (all), spe- cific habitat requirements (such as rainforest; e.g., some Rhacodactylus), and low resistance to des- iccation (e.g., Nannoscincus).
The carphodactyline geckos Rhacodactylus, Eurydactylodes, and Bavayia represent a natural group that has invaded Australia (Bauer 1986, in press). King (1987) has proposed a Gondwa- nan origin for geckos as a whole and subsequent phylogenesis associated with the breakup of the supercontinent. We believe that evidence sup- ports the origin of the subfamily Diplodactylinae as a result of the isolation (both geographical and climatical) of eastern Gondwanaland from its neighbors. Immunological investigations in the context of molecular clock dating suggest that the New Caledonian carphodactylines last shared a common ancestor with Australian members of the group at least 66 mybp (King 1987; Bauer and Rainey in prep.). This is compatible with the accepted age of the opening of the Tasman and Coral seas some 80 mypb (see Bauer 1986 for complete discussion).
The origin of the endemic skinks of New Cal- edonia is less easily explained. Phoboscincus bo- courti and Geoscincus haraldmeieri, following the reasoning outlined above, probably arrived as a result of over-water rafting of a Eugongylus-like ancestor. Most of the remaining taxa fall into Greer’s (1974) Leiolopisma group and were ar- ranged by Sadlier (1986) into the genera Leiolo- pisma sensu stricto, Caledoniscincus, Tropidos- cincus, Marmorosphax, and Sigaloseps. The first genus is apparently paraphyletic, but Sadlier (1986) grouped its New Caledonian members and
the following three genera as each other’s closest relatives, with Sigaloseps, Graciliscincus, and Nannoscincus (including the Australian N. mac- coyi) as separate lineages. Even after the reshuf- fling of Leiolopisma, the biogeography of the group makes no more sense than it did following Greer’s generic concepts (see Zug 1985b), and neither vicariance nor dispersal hypotheses alone adequately account for the observed distribution patterns.
The minor radiation of some taxa such as Tro- pidoscincus and Nannoscincus in New Caledonia demonstrates that the New Caledonian land mass is large enough and diverse enough ecologically to have supported “‘continental” speciation (sen- su Diamond 1984) in reptiles. The occurrence of the purportedly monophyletic Leiolopisma group skinks certainly suggests a long period of evo- lution in isolation in New Caledonia. Unfortu- nately, the absence of a broader, explicit phy- logenetic hypothesis for the Lygosominae does not permit an application of narrative historical biogeographical methods (sensu Humphries and Parenti 1986), nor are the phylogenies of other biotic components of the region well established enough to effectively utilize the analytical ap- proach of cladistic biogeography. The scincids thus represent the most glaring gap in our general understanding of the zoogeography of the New Caledonian herpetofauna.
Within New Caledonia, some ecogeographic trends are seen. Most noticeably there is a faunal element which, via the vegetation, is tied to the regions of lateritic soils that cover about one third of the land area of the mainland (O.R.S.T.O.M. 1981; Morat et al. 1986; Jaffré et al. 1987). This group includes Graciliscincus shonae, Tropido- scincus roehssii, Sigaloseps deplanchei, Nanno- scincus mariei, Rhacodactylus auriculatus, and R. sarasinorum. All of these taxa occur only in the southern third of the island, the largest single region of lateritic soils. Unfortunately, the exact nature of the relationship between the soil type and these lizards is unclear, although the effect of soil on the peculiar ““New Caledonian growth form” of much of the flora is well known (Jaffré 1980).
Other components of the endemic fauna ap- pear to have ranges associated with pluviometric features of the island. The greatest diversity of species occurs on the more humid east coast, where the winter wet season is most pronounced. This is especially true of geckos (Bauer 1986) and
BAUER AND VINDUM: NEW CALEDONIAN HERPETOFAUNA 41
ths
FiGure 5.
Soil types of New Caledonia; 1, alluvial soils; 2, feralitic soils on ultramafic rocks; 3, brown soils over basic rocks;
4, other non-carbonate soils; 5, carbonate soils. Many patterns of intra-island endemism are associated with the lateritic soil types (2 in legend), especially in the large continuous region in southern New Caledonia. (Map modified from Atlas de la Nouvelle-Calédonie et Depéndences, 1981, O.R.S.T.O.M.., Paris.)
the sub-fossorial skink species (Sadlier 1986). El- evation appears to play less of a role in New Caledonia than on other high islands, and there are no known high-elevation endemics, although herpetological investigations of the many peaks over 1,000 m has been limited and should have high priority for future research.
Perhaps one of the most intriguing features of the zoogeography of New Caledonia concerns the taxa now extinct, including meiolaniid turtles (Gaffney et al. 1984), mekosuchian crocodiles (Buffetaut 1983; Balouet and Buffetaut 1987), and varanids (Gaffney et al. 1984). All of these ani- mals were probably contemporaneous with man. The occurrence of these fossil forms is somewhat unsettling, as they disrupt the established picture of reptile distribution in the Pacific and may weaken existing hypotheses if applied in the con- text of cladistic biogeography. Their presence
again raises the question of the role of man in shaping the present herpetofauna, not only by means of introductions, but by extinction as well (Cassels 1984; Steadman 1985). For example, Varanus may have been a native form that was exterminated by early settlers, or, asin Palau and the Caroline islands, it may have been brought to New Caledonia as food and subsequently died out. Fossil geckos and skinks (perhaps including new taxa) have also recently been discovered, but probably predate the arrival of man in New Cal- edonia (Balouet and Olson 1989).
The addition of both phylogenetic and geologic hypotheses that are well corroborated has added much to the progress in interpretation of the her- petofauna of New Caledonia. Perhaps the most significant insights since the work of Roux (1913) are that (1) some faunal elements, such as the carphodactyline geckos and some of the skinks,
42 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 47, No. 2
are probably present as the result of vicariant events associated with the breakup of Gon- dwanaland (Bauer 1986, in press), and (2) the recognition that the fauna of the region, like that of other Pacific high islands (Gibbons 1985; Bauer 1988a, b), is a mosaic of elements derived from a variety of sources.
Among the priorities for establishing a firmer understanding of the biogeography of the new Caledonian herpetofauna are: (1) the erection of an explicit hypothesis of lygosomine relation- ships; (2) the genetic analysis of Pacific popula- tions of widespread taxa in order to trace poten- tial migration or transport routes (this work has already begun, e.g., Ineich 1988; Moritz 1987); and (3) the study of additional fossil material that may indicate a once broader faunal diversity. Perhaps most basically, more locality data and careful field work in neglected regions of New Caledonia, particularly fossorial and montane habitats, are needed in order to establish the pat- terns of ecogeographic variation and complete the herpetofaunal list for the region.
ACKNOWLEDGMENTS
The senior author thanks the New Caledonian authorities for permission to collect the speci- mens cited herein. Logistical support in New Caledonia was provided by M. Rémy LeGoff (S.L.N.), O.R.S.T.O.M. (Drs. Jean Chazeau and Luc Brun), and M. Alain Renevier and family. Field work was carried out with the assistance of Griff Blackmon, Kathy DeVaney, Larry Wish- meyer, Debra Wadford, and Katie Muir. Finan- cial support for ongoing research in New Cale- donia has been provided by the Department of Herpetology and the G. Lindsay fund, California Academy of Sciences. The manuscript was typed by Nancy Stevenson and Figures 2—4 were pho- tographed by Mare Charnow (CAS). We thank Margaret Berson and Cecily Bloomfield for ed- itorial comments. The manuscript has benefited from the constructive comments of Ron Crom- bie and Kevin de Queiroz.
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STEADMAN, D. W. 1985. Fossil birds from Mangaia, southern Cook Islands. Bull. Brit. Ornithol. Club 105:58-66.
STEJNEGER, L. 1907. Herpetology of Japan and adjacent ter- ritory. U.S. Natl. Mus. Bull. 58:xx + 1-577.
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1985b. A new skink (Reptilia: Sauria: Leiolopisma)
from Fiji. Proc. Biol. Soc. Wash. 98:221-231.
1989. Heteronota pelagica Girard, 1857 (currently Gymnodactylus, Cyrtodactylus or Nactus pelagicus; Reptilia, Sauria): proposed conservation of the specific name. Bull. Zool. Nomencl. 46:38-40.
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COLLIS A: C7 PHALOPODA) OF 7 PASTERN PACIFIC 5
PROCEEDINGS OF THE
CALIFORNIA ACADEMY OF S
Vol. 47, No. 3, pp. 47-94, 20 figs., 19 tables.
Marine Biological Laboratory LIBRARY
Nov 7 1990
IF Wes Hole, Mass.
October 17, 1990
DEEP-WATER OCTOPODS (MOLLUSCA; CEPHALOPODA) OF THE NORTHEASTERN PACIFIC
By
Gilbert L. Voss!
Rosenstiel School of Marine and Atmospheric Science, University of Miami, 4600 Rickenbacker Causeway, Miami, Florida 33149
and William G. Pearcy
College of Oceanography, Oregon State University, Corvallis, Oregon 97331
Asstracr: Descriptions and illustrations are given of nine species of Octopoda from depths of 1,000 m to 4,000 m off the coast of Oregon. Eight are new: Grimpoteuthis bathynectes, G. tuftsi, Benthoctopus robustus, B. canthylus, B. oregonensis. B. yaquinae, B. macrophallus, and Graneledone pacifica. Four genera are treated: Cirroteuthis and Grimpoteuthis of the Cirrata and Benthoctopus and Graneledone of the Incirrata. Cirroteuthis muelleri is first recorded from the Pacific Ocean and is redescribed. The genus Benthoctopus in the North Pacific is discussed and a provisional key for the nine recognized species is provided. The variability and form of the radula of Graneledone are considered to be due to degeneracy and not primitiveness, and to lack
of selective feeding pressure.
Received August 21, 1989. Accepted March 14, 1990.
INTRODUCTION
The deep-water octopod fauna of the north- eastern Pacific below 2,000 m is poorly known. Little research on the systematics of deep-sea octopods has been completed since Robson’s (1932) monograph, with the exception of Voss’s (1976) partial revision of Graneledone. Two re- cent papers, however, provide important reviews of the evolution, phylogeny, and biogeography of deep-sea octopods and relevant background for this paper (see Voss 1988a, b).
' Deceased January 23, 1989.
[47]
The present paper is based on a unique col- lection of about 90 specimens of deep-water oc- topods, all from the continental slope and two abyssal plains off the Oregon coast. The speci- mens were collected by the ships CAYUSE, ACONA and YAQUINA of Oregon State Uni- versity (OSU) in depths from 1,000 to nearly 4.000 m, with most of the material from depths in excess of 2,500 m. Most of the specimens belong to new species in the genera Grimpoteu- this and Benthoctopus, with the largest series of specimens in the genus Graneledone. All three of these poorly known genera consist of a large number of species, most of which were inade-
Ps a
48 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 47, No. 3
quately described and illustrated; thus, reference to type material was necessary.
All of the type specimens of deep-water oc- topods from the northern Pacific in the collec- tions of the United States National Museum of Natural History (USNM) were borrowed, but their poor condition prevented direct compari- son of many features. There was little non-type material at the USNM, although some fine spec- imens of Graneledone verrucosa were borrowed for comparison with the Pacific specimens.
In 1975 Voss traveled to Europe to examine the extensive collections at the British Museum (Natural History), the Muséum National d’His- toire Naturelle in Paris, and the Institut Océano- graphique in Monaco. Much of the type material was in poor condition and its examination failed to assist in the comparisons except in a few cases which bore indirectly upon the problems of the Oregon material. Specimens of Benthoctopus were borrowed from Takashi Okutani, Tokai Region- al Fisheries Research Laboratory (currently To- kyo University of Fisheries). Examination of these specimens confirmed the separate identity of B. canthylus, but left the others in doubt.
Characters previously used to identify deep- water octopods were not sufficient for species recognition. Therefore the specimens were care- fully dissected, and when possible, detailed ob- servations were made for each species of the beaks, radulae, digestive tracts, genitalia, sper- matophores, and stellate ganglia, as well as the more traditional counts and measurements. All these features were illustrated.
We hope that the details given here will permit positive identification of new material from the northeastern Pacific without necessitating the onerous, frustrating, and detailed examination of every specimen, as encountered in this study. The deep-sea benthic octopods are, however, in need of monographic revision.
CLASSIFICATION OF NORTHEASTERN PACIFIC DEEP-WATER OCTOPODS
Suborder Cirrata Grimpe, 1916 Family Cirroteuthidae Keferstein, 1866 Genus Cirroteuthis Eschricht, 1836 Cirroteuthis muelleri Eschricht, 1836 Family Opisthoteuthidae Verrill, 1896 Genus Grimpoteuthis Robson, 1932 Grimpoteuthis bathynectes new species Grimpoteuthis tuftsi new species
Suborder Incirrata Grimpe, 1916 Family Octopodidae Orbigny, 1845 Subfamily Bathypolypodinae Robson, 1928 Genus Benthoctopus Grimpe, 1921 Benthoctopus robustus new species Benthoctopus canthylus new species Benthoctopus oregonensis new species Benthoctopus yaquinae new species Benthoctopus macrophallus new species Subfamily Graneledoninae Voss, 1988a Genus Graneledone Joubin, 1918 Graneledone pacifica new species
MATERIALS AND METHODS
Collections were made with 2.7 to 3.0 m wide beam trawls with 13 mm (stretch) mesh and with otter trawls having a 7 m foot rope and 38 mm mesh with a 13 mm mesh liner in the cod end. Both nets were towed at two knots (3.7 km hr~') (see Carey and Heyamoto 1972 for descriptions of nets and sampling methods). Over 600 tows were taken on the continental slope and abyssal plains in the northeastern Pacific from depths of 400 to 5,180 m, 32 to 2,225 km offshore, between 1961 and 1974 (Fig. 1, see Pearcy et al. 1982 for details). Of these, 164 tows were from 2,500 to 3,045 m on the Cascadia Abyssal Plain, about 400 tows from 400 to 2,780 m on the continental slope off Oregon, and 35 tows from 2,780 to 5,182 m on the Tufts Abyssal Plain. Octopods from this study were captured at depths from 1,000 m on the continental slope to 3,932 m on Tufts Abyssal Plain from the research vessels ACONA, CAYUSE and YAQUINA. One spec- imen was caught in a trap at 3,660 m on the Aleutian Abyssal Plain.
Collections, including octopods, were pre- served in 10% formalin and seawater at sea. Oc- topods were moribund or dead after capture, so they were not relaxed or killed prior to preser- vation.
After the cruises, the muscular, firm-bodied specimens were transferred to 70% ethanol, while the soft-bodied forms were stored in 40% iso- propyl alcohol. The delicate cirroteuthids were stored in 10% formalin. Most of the material was transferred to 70% ethanol in Miami. Material stored in formalin, especially cirroteuthids, showed signs of deterioration. Storage in ethanol results in some shrinkage but is preferrable to loss of the specimens.
All descriptions of color are based upon spec-
VOSS AND PEARCY: DEEP-WATER OCTOPODS
49
sis)
Choo tte)
25 (e)
TUFTS ee PLAIN
155° 150° 145° 140°
Ficure 1.
V t 50°
CALIF. v
Location of trawl collection areas on the continental slope, Cascadia Abyssal Plain, and Tufts Abyssal Plain.
Obliquely hatched areas denote general sampling areas on Cascadia Plain, solid dots specific sampling stations on Tufts Plain and the slope. The station on the Alaskan Abyssal Plain shows the location of capture for one Benthoctopus robustus in a trap.
Vertical hatched areas indicate ridges and hilly topography.
imens preserved in ethanol. No color notes were made of living specimens.
All specimens were carefully examined, and measurements, body proportions, and counts were obtained from the whole animals as de- scribed by Voss (1963). The mantle of all spec- imens was opened to permit examination of the internal anatomy and to count gill lamellae. The digestive tract and genitalia were dissected to permit detailed descriptions and illustrations for each species. Because of the paucity of good spe- cific characters in these octopods, the buccal mass was removed from nearly all specimens and the beaks and radulae extracted, cleaned, and illus- trated. The buccal mass was typically immersed in tap water with one or two pellets of potassium hydroxide and gently heated. Beaks and radular ribbons were cleaned in household bleach solu- tion (Chlorox). The beaks were illustrated lying free in a Petri dish, while the radulae were mounted in either Euparol or CMC 10 and drawn with the aid of a camera lucida. The radular rib- bon of Grimpoteuthis tuftsi was destroyed if the buccal mass was treated as described above; even
quick cleaning in bleach caused some radular ribbon deterioration.
Spermatophores were obtained only from Benthoctopus. They were so large that mounting was impractical; they were described and illus- trated lying free in a Petri dish.
Special attention has been given to the stellate ganglion, particularly in the cirroteuthids, as this structure appears to be of systematic value. The ganglion and its associated nerve fibers were dis- sected from the mantle wall, mounted on a slide, and drawn with the aid of a camera lucida.
The indices used are those defined by Voss (1963) with the exception of those for fin length (FLI) and width (FWI) of cirroteuthids and the hectocotylized arm index (HcAI), which is ex- pressed as a percentage of the length of its fellow arm on the opposite side. The range and mean of ratios and measurements is given as follows: 60-80-100, in which the italicized number is the mean. All measurements are in mm unless oth- erwise stated.
The cirroteuthids present certain problems not encountered in other octopods, e.g., the method
50 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 47, No. 3
of taking measurements of the cirri, the fins, and the mantle length. Some writers have considered that the cirri can contract differentially depend- ing on the type and strength of the preservative. However, cirri lengths do not vary appreciably within a species and average cirri lengths for con- specific animals are the same whether preserved in formalin, isopropyl alcohol, or ethanol. The cirrus length was taken from the longest cirrus on the arms and measured intact from the base to the tip. In some very flaccid specimens the cirrus was removed, stretched out on a glass slide, and measured.
Fin measurements entailed a reconsideration of terms applied both to squid and cirroteuthids. In squid, fin width is the greatest width across the fins measured from outer angle to outer angle, while length is measured from the anterior lobe to the posterior insertion or the tip of the mantle in fins united posteriorly. This measurement is misleading, however, when applied to the pad- dle-like fins of cirroteuthids, where fin length was measured from the midpoint of a line drawn across the fin base to the outer angle of the fin. There is considerable difference in measure- ments made along the posterior and anterior margins, particularly if the fins are directed somewhat anteriorly. Fin width is measured across the individual fin at the widest point when the fin is flattened to include the delicate thin anterior border. Fin-width index is expressed as a percentage of fin length. Fin-length index is expressed as a percentage of the interocular width for reasons given below.
Standard length in octopods has convention- ally been considered to be the dorsal mantle length (ML) or the length of the body measured from mantle apex to a line drawn across the head from pupil to pupil. In cirroteuthids this measurement is fraught with difficulties. In some species of Grimpoteuthis the posterior end of the mantle coincides with the posterior surface of the shell vestige. In others, particularly Cirroteuthis and Stauroteuthis but also in some Grimpoteuthis, in life or in freshly preserved material, the mantle extends well beyond the shell vestige as a gelat- inous, often fluid-filled sac, projecting posteriorly equal to the length of the mantle from the inter- ocular line to the shell vestige. In preservation, this mantle extension is often completely con- tracted against the shell vestige. As a result, dis- cretion must be used, and as Robson (1932) and others have often given fin length as a ratio of
the interocular width, the fin-length index used here is expressed similarly.
Holotypes of the new species described are de- posited in the United States National Museum of Natural History (USNM). Other material ex- amined, including the paratypes, are in the col- lections of the California Academy of Science (CAS), the Santa Barbara Museum of Natural History (SBMNH), the invertebrate collections of the Rosenstiel School of Marine and Atmo- spheric Science, University of Miami (UMML), and USNM.
THE StuDy AREA
The area of sampling has been described by Pearcy et al. (1982). Briefly, the steep and topo- graphically complex continental slope off Oregon adjoins Cascadia Abyssal Plain, a basin bordered by submarine ridges on the west and south (Fig. 1). Its floor slopes gradually to the south and west. The Astoria and Nitinat fans, two large Pleistocene fans associated with the Columbia and Fraser rivers, are dominant features of the basin, producing a gradual slope from depths of 2,100 m on the fans adjacent to the slope to over 3,000 m in the southern portion of the basin.
Tufts Abyssal Plain is farther offshore and much less influenced by terrigenous sedimenta- tion than is Cascadia Plain. It is bordered by the East Pacific Rise on the east and hilly seamounts on the south; it is connected with the Alaskan and Aleutian Abyssal plains on the north and west. Depth of the Tufts Plain increases gradually to the west. Cascadia Channel, which originates in Cascadia Plain and penetrates the East Pacific Rise, is the only deepwater connection between Tufts and Cascadia plains.
The sediments of these plains are mainly fine muds and clays. In general the rate of sedimen- tation and organic carbon content of the sedi- ments decrease from east to west across both plains (Griggs et al. 1969). The biomass of ben- thic megafauna, both invertebrates and fishes, also decreases from east to west (Carey 1965; Pereyra and Alton 1972; Pearcy et al. 1982).
SYSTEMATICS
Order OcTOPODA Suborder CrrrATA Grimpe, 1916
Deep-sea octopods that live on or just above bottom. Body gelatinous to subgelatinous, with
VOSS AND PEARCY: DEEP-WATER OCTOPODS
lateral paddle-shaped fins, uniserial suckers, and cirri alternating with suckers; shell vestige car- tilaginous U-, V-, or saddle-shaped, well devel- oped, serving mainly as fin support; radula may be present. Representatives have been reported from throughout the world ocean.
Family CirROTEUTHIDAE Keferstein, 1866
As currently recognized, this family contains two monotypic genera— Cirroteuthis and Cirro- thauma (see Voss 1988b:296). Body elongate, gelatinous, fragile, colorless to only lightly pig- mented except on oral surface of arms and web; cirri long, conspicuous; fins large and wide; sim- ple crop present; radula absent; gills ““sepioid”’; median pallial adductor vestigial; ““secondary” web present; shell vestige saddle-shaped.
Cirroteuthis Eschricht, 1836 With the characters of the family.
Tyre Species: Cirroteuthis muelleri Eschricht, 1836, by monotypy.
Cirroteuthis muelleri Eschricht, 1836 (Fig. 2, 3)
Cirroteuthis muelleri Eschricht, 1836:627, pls. 46-48. Morch 1857:440. Hoyle 1886:60. Lonnberg 1891:6. Appelléf 1893: 1. Posselt 1898:269. Appelléf 1899:7-8, pl. 1, figs. 6-7. Friele and Grieg 1901:123. Pfeffer 1908:23, fig. 13. Robson 1926: Passim; 1932:130, fig. 18. Grieg, 1933:9.
Sciadephorus muelleri Reinhardt and Prosch, 1846:187, pls. 1-5.
MATERIAL EXAMINED (12 specimens all collected by R/V YAQUINA).—2 males, ML 74-79 mm, Cr. Y7102B haul 262, 45°38.3'N, 126°43.8’W in 2,721 m, 17 Feb. 1971, USNM 817580.—1 female, ML 66 mm, Cr. Y7105B haul 276, 45°56.7'N, 127°38.6’W in 2,761 m, 17 May 1971, SBMNH 35142.—1 female, ML 67 mm, Cr. Y7102B haul 263, 45°36.4’N, 126°44.9'W in 2,730 m, 17 Feb. 1971, UMML 31.1935.—1 male ML 72 mm, | female ML 21 mm, Cr. Y7105B haul 278, 45°24.0'N, 127°39.0’W in 2,811 m, 18 May 1971, UMML 31.1936.—1 female, ML 64 mm, Cr. Y7102B haul 265, 45°18.6'N, 126°31.5'W in 2,750 m, 18 Feb. 1971, CAS 067786.—1 female, ML 63 mm, Cr. Y7102B haul 270, 44°38.4'N, 126°42.0’W in 2,830 m, 20 Feb. 1971, SBMNH 35143.—2 females, ML 47-63 mm, Cr. Y7102B haul 268, 44°58.8'N, 126°37.4’W in 2,770 m, 19 Feb. 1971, USNM 817581.—1 male, ML 53 mm, Cr. Y7005C haul 230, 44°27.0'N, 132°14.0'W in 3,655 m, 1 June 1970, CAS 067787.—1 female, ML 40 mm, Cr. Y7301F haul 317, 44°44.5'N, 127°29.0'W in 2,810 m, 3 Feb. 1973, CAS 067788.
DESCRIPTION. — Moderately large species with soft, almost gelatinous consistency; body cov- ered by thin, tough, outer layer except on arms
51
and web where easily torn; all of animal very fragile except for parts of internal anatomy.
Mantle elongate, somewhat tubular but gently rounded posteriorly (Fig. 2a), about half as wide as long (Tables 1-4); head width slightly greater than mantle width with no discernible constric- tion between head and mantle; mantle aperture narrow, mantle fits snugly around funnel; eyes small, fully formed, with large lenses; eyelid ab- sent but skin transparent over orbit.
Funnel long, conical, narrow, free for about half of its length; funnel organ thin, flat, A -shaped, with sharp median point and truncate lower limbs; small cup-shaped olfactory organ located about midway on either side of funnel on an edge of shallow funnel groove.
Fins approximately median, located slightly nearer apex of mantle than eyes (Fig. 2a); each fin paddle-shaped, about twice as long as wide (Tables 1-4); posterior edge nearly straight, curves anteriorly just past midpoint; anterior margin strongly rounded and proximally turns posteri- orly to form narrower base; heavy muscular por- tion of fins well developed, usually occupies most of fin width near base, extends at least halfway to fin end; outer lobe of fin nearly forms semi- circle; fins do not project at 90° angle from body axis but are directed slightly anteriorly.
Arms translucent, not muscular, appear filled with fluids in life but usually collapsed, wrinkled when preserved. Arm lengths difficult to obtain due to contractions of arms and distortion by contraction of web; measurements therefore ap- proximate. Arms (Fig. 3a) subequal with perhaps lateral arms slightly longest; single row of arm suckers as far as distal attachment of primary web, beyond which arms slender, smooth, de- void of suckers, whip-like; inner oral surface of arms broad and bear suckers, aborally arms nar- rower, triangular in cross-section, outer edge of arm located inside secondary or intermediate web extends as lamella to outer or primary web.
Primary web with thin tough outer layer that keeps web intact in preserved specimens; pri- mary web usually torn from arms during capture, attached only to arm near tip on dorsal side at point where suckers terminate; arms along rest of their length attached to primary web by dorsal and ventral intermediate webs (Fig. 3a); interior spaces appear to be filled with fluids; dorsal in- termediate web begins at base of arms, widest at about midpoint, and tapers to terminate near primary web-attachment point; ventral inter-
PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 47, No. 3
2a wefie io oe \ ) \ \. a ee _ eo aes Bis { / Ne Sea a | Nee ; S ae of / | | 7 fb ee : \ ¢ Ga yee Su Ee Se aE a a is oa 7 \ \ va / \ ] 5 — + ( zx L : AS | e d Ns e — = —_-, ( \ f
| Sa mm ML.
j NS Ficure 2. Cirroteuthis muelleri Eschricht. (a) Dorsal view of male; ML 79 mm. (b-c) Upper and lower beaks. (d-f) Ventral,
dorsal, and lateral views of shell vestige of male, 74 mm ML. (g-h) Dorsal and ventral views of shell vestige of male, 53
VOSS AND PEARCY: DEEP-WATER OCTOPODS
53
za d sem. ves.
vas. def.
Ficure 3. Cirroteuthis muelleri Eschricht. (a) Oral view of arms and web of male (dorsal side up), 79 mm ML. (b) Genitalia and egg of female, 67 mm ML. (c-d) Genitalia of male, 72 mm ML. (e) Left stellate ganglion.
mediate web originates same as dorsal, is wider, and terminates as moderately wide membrane along inner edge of large, fleshy, thinly tapered process on ventral side of arm a few millimeters proximal to attachment of primary web. This whip-like process (nodule) also the fusion point of primary web, which unites on aboral surface,
intermediate web on its oral surface; no indica- tion of unions of webs when viewing primary web aborally.
Suckers small (Tables 1-4); first suckers sur- rounding mouth small, 2nd and 3rd succeeding suckers usually largest, then size gradually de- creases to about 7th or 8th sucker (Fig. 3a). Seven
54 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 47, No. 3
TaBLE 1. Measurements and counts of four males of Cir- roteuthis muelleri Eschricht, 1836.
Specimen
USNM USNM UMML_ CAS
817580 817580 31.1936 067787 Mantle length 79.0 74.0 72.0 53.0 Mantle width 47.0 26.0 40.0 27.0 Head width 52.0 34.0 37.0 32.0 Arm length I 170.0 — 145.0 = Arm length II 178.0 140.0 = _ Arm length III 170.0 - - a Arm length IV 170.0 = = = Total length 277.0 210.0 222.0 178.0 Fin length 50.0 47.0 47.0 53.0 Fin width 30.0 21.0 30.0 23.0 Sucker diameter — iN) Del 1.0 Cirrus length 19.2 16.0 18.4 11.0 Gill lamellae 8 9 8 7 Arm formula 21=3=4 _ = —
or eight basal suckers cup-shaped, usually raised on broad, heavy pads; suckers smaller distal to 8th sucker, often less than half size of basal suck- ers, delicate, thin-lipped, often with no apparent suction cup, and formed of two, almost parallel, elements. Sucker bases in undamaged specimens widely spaced, formed of large, delicate, mam- milar, fluid-filled structures, surmounted by minute, apparently non-functional sucker cups. When fluid-filled bases collapse, suckers lie flat on arm or appear to be mounted on thin delicate pedicels, according to condition of collapsed bas- es. Distal to small suckers, at about 28th sucker, normal appearance regained. Ventral web nodule joins at about 3rd or 4th enlarged sucker; suckers gradually become smaller, terminate at union of primary web with arm on dorsal surface; largest of terminal suckers about same diameter as basal suckers.
Cirri small, occur in pairs between either suck- ers | and 2, or 2 and 3, one on each side of sucker slightly proximal to base, appear to alternate with suckers, extend distally to terminal sucker; larg- est in midportion of arm, attain length of over 19 mm, or 9 to 13 times diameter of largest sucker. (Cirri difficult to measure unless removed from arm and placed on glass slide.)
Gills small, little modified, with 7-8-9 lamel- lae per outer demibranch, including large, par- tially divided terminal lamella. Thin, transpar- ent, distinct median pallial adductor muscle, 10- 16% ML, unites mantle to viscera; inner attach- ment on membrane overlaying male or female genitalia in median position.
TABLE 2. Indices of bodily proportions and counts of four males of Cirroteuthis muelleri Eschricht, 1836.
Specimen
USNM USNM UMML_ CAS
817580 817580 31.1936 067787 25 ML 79.0 74.0 72.0 53.0 — MWI 58.7 B5n1 55:5 50.4 49.9 HWI 65.0 45.9 SES 60.3 55.6 MAI 47.0 52.8 49.6 46.9 49.1 ALI 64.2 66.6 65.3 63.4 64.9 SIn — 17; 2.9 1.8 Zal Cil 24.0 21.6 25.6 20.8 23.0 FLI 96.2 138.2 127.0 165.0 131.6 FWI 60.0 44.7 63.8 43.4 52.9
Posterior third of mantle cavity devoid of structures but filled with tough, viscous, gel-like material that gives body its shape posterior to skeletal structure; may be reservoir for ammo- nium ions for buoyancy. Area is easily damaged, collapses with too much handling, difficult to measure.
Shell vestige saddle-shaped; originates as flat sheet that curves dorsally on anterior and pos- terior borders, curls in toward itself, thickens to form supports for large basal fin muscle (Fig. 2d— h).
Beaks thin, weak, and indistinctive (Fig. 2b, c); radula absent, surface normally covered by radula thickened, slightly ridged transversely, bears series of longitudinal folds that resemble lines of various sizes of teeth; cuticular lining not apparent.
Esophagus slender at connection with buccal mass, passes posteriorly between two anterior salivary glands (posterior salivary glands absent); widens into croplike area without diverticulum, but expanded to two to three times diameter of anterior section; posterior to “crop’’ esophagus narrows, enters distinctly two-parted stomach; spiral caecum nearly as large as both elements of stomach; two large ducts lead into digestive gland through digestive duct appendage; diges- tive gland round but slightly flattened; intestine large, nearly straight, not divided into parts. Lon- gitudinal folds run full length of intestine, seen clearly through its transparent wall. Ink sac ab- sent; anus without flaps.
Male genitalia unusual (Fig. 3c, d), difficult to compare with those of other octopods: duct from testis large, leads into folded mass with thin- walled out-folding that may correspond to Need-
VOSS AND PEARCY: DEEP-WATER OCTOPODS 55 TABLE 3. Measurements (in mm) of seven females of Cirroteuthis muelleri Eschricht, 1836. Specimen CAS USNM USNM SBMNH CAS SBMNH UMML
067788 817581 817581 35143 067786 35142 31.1935 Mantle length 40 47 63 63 64 66 67 Mantle width 19 22 32 - 46 43 40 Head width 21 23 35 — 42 42 47 Arm length I - - _ — 114 96 _ Arm length II — = _ - — 96 — Arm length III _ = - - — 94 — Arm length IV 67 95 108 _ _ 91 124 Total length 113 157 190 — 180 172 203 Fin length 41 52 61 52 50.5 46.5 49.5 Fin width 18 24 25 38.0 35.0 36.0 30.0 Sucker diameter 0.6 1.0 1.0 Pegi 1.6 1.4 2.4 Cirrus length 8.0 9.0 13.0 16.0 19.2 19.2 13.0 Gill lamellae 7 7 8 8 8 8 7 Arm formula - _ — — — 1=234 —
ham’s sac; distal portion of folded mass smooth with small slender duct (penis?) that protrudes from its center; Needham’s sac contains about 10 small ovoid structures, apparently spermato- phores (similar to structures found in Grimpo- teuthis); they do not resemble typical octopodon spermatophore and probably consist only of sperm reservoir.
Female genitalia (Fig. 3b) unpaired, nearly round ovary contains about 12 large, apparently ripe, ovoid, flesh-colored eggs with light wavy linear markings; 10.4 x 9.3 mm; numerous small to minute eggs also present; proximal oviduct single, about 4—5 times length of distal oviduct, rather inflated, swollen at distal end at junction with oviducal gland; oviducal gland large, dark colored, forms ring around oviduct; distal ovi- duct short, strongly flared at distal opening. Ma- ture egg at end of oviduct, with another lodged in oviducal gland (Fig. 3b) exactly as figured by Reinhardt and Prosch (1846, pl. 4).
Optic ganglia distinct, globular, darkly pig- mented, almost same diameter as orbit of eye. The stellate ganglion shown in Figure 3e.
Color in ethanol varies from pale white to pur- ple; inner surfaces of arms and web invariably darkest, purple or purplish-brown.
MEASUREMENTS AND Counts. — Males: mantle length 53-69.5-79; mantle width 26-35.0-47; head width 32-38.8—52; arm length I 145-170, II 140-178, III 170, IV 170; total length 178- 221.8-277, fin length 47-49. 3-53; fin width 21- 26-30; sucker diameter 1—/.5—2.1; cirrus length 11.0-16.2—19.2; number of gill lamellae 7-S-9.
Females: mantle length 40-58.6-67; mantle width 19-33.3-46; head width 21-—35.0-47; arm length I 96-114, II 96, Ill 94, IV 67-97-124; total length 113-/69.2—203; fin length 41-50.4— 61; fin width 18-29.4-38; sucker diameter 0.6- 1.4-2.4; cirrus length 8.0-/3.9-19.2; number of gill lamellae 7-7.6-8.
Type.— Zoological Museum, Copenhagen. Robson (1932: 130) erroneously stated that it was in Naturhistorischen Verein of Breslau and Bonn. In a letter to Robson from Sparck dated 8 February 1932, Sparck stated that Robson had made a mistranslation and that the holotype is in the Zoological Museum of Copenhagen (see Kristensen and Knudsen 1983).
TYPE-LOCALITY.—Jakobshavn, West Green- land (Eschricht 1836:627).
Discussion. —A careful examination of the 12 specimens available has revealed no important differences between our material and the descrip- tions of Cirroteuthis muelleri given in the liter- ature (Eschricht 1836; Reinhardt and Prosch 1846; Robson 1932). Some minor differences oc- cur.
Robson (1932:131) drew attention to a pecu- liar modification of the small suckers on the arms of the British Museum specimen—they were flat. If he referred to the sucker only and not its total base, this condition is also found in our speci- mens. Eschricht’s fig. 1 indicates that the median suckers were smaller in the type, and the color- ation seems to indicate that they were on large mammilar bases, just as in our specimens.
The differences in the origin of the cirri (often
UMML 31.1935 67 59.7 70.2 54.0
66
65.2 63.6 68.8
SBMNH 35142
CAS 067786 64 T9 65.6 56.1
Specimen 63 50.8 55.6 58.3
USNM 817581
SBMNH 35143 63
47 46.8 48.9 49.5
Indices of bodily proportions and counts of seven females of Cirroteuthis muelleri Eschricht, 1836. USNM 817581
TABLE 4.
CAS 067788 40 47.5 59.7
MWI HWI MAI ALI SIn
ML
PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 47, No. 3
between the Ist and 2nd suckers in our speci- mens, only between the 2nd and 3rd in the oth- ers) seems to be a matter of individual variation.
S “ BS u S In our specimens cirri may first appear between a suckers | and 2 in some arms and between 2 and 3 on other arms. Often only one cirrus is present between | and 2, both cirri between 2 and 3. Robson (1929:26) pointed out the possible sys- tematic value of the size of the mantle aperture, eaans and in C. muelleri indicated that while Esch- WIADRD Oh . ee . . . Oe richt’s original description showed a wide aper- ture (Robson’s ‘C’), the British Museum speci- men has a small aperture (at least “B’). We believe the original illustration is of a very flaccid and perhaps stretched specimen. Our specimens all show at least the B condition and often A. - & S So The shape of the fins also is different. Both =] Eschricht (1836) and Reinhardt and Prosch (1846) show broad flabellate fins with a very un- natural appearance. We believe that this is ar- tistic treatment; our specimens show a remark- able homogeneity of appearance and closely Mie Goh Lt correspond to the more detailed structure shown STRESS in pl. 5, fig. | of Reinhardt and Prosch (1846).
A more basic difference is the presence or ab- sence of a median pallial adductor muscle. This is discussed by Robson (1932:119-120), who stated that there was no median adductor in the British Museum specimens. A thin but distinct median adductor is present in our specimens; it is easily overlooked because it is small and trans- parent. Robson was not noted for the delicacy of his dissections and it is possible that, being familiar with the thick, muscular adductors of the Incirrata, he overlooked the pallial adductor in his specimen. Voss examined two fine USNM specimens of C. muelleri obtained by the R/V WALTHER HERWIG in the North Atlantic. Both had the narrow, transparent, easily over- looked median pallial adductor muscle.
DIsTRIBUTION.—Cirroteuthis muelleri was pre- viously known only from the northern part of the North Atlantic (see Robson 1932:130; Voss 1988b). Its presence in the northeastern Pacific indicates either a very broad distribution or a discontinuous circumboreal pattern (Ekman 1953:161). While the shallow waters of the Arctic Ocean may be a barrier, vertical distribution there is not limited by temperature, as shown by the capture of Cirrothauma murrayi in an ice hole off Point Barrow (Voss 1967:527). Furthermore, cirromorphs resembling Cirroteuthis have been photographed in the Arctic Ocean in 2,360 to
33
60.5
NX
19.2 226.0 46.2
1.5
59.3 20.0 195.2 44.0
Cil FLI FWI
VOSS AND PEARCY: DEEP-WATER OCTOPODS
3,786 m (Pearcy and Beal 1973), suggesting a continuous distribution through the Arctic Ocean.
Family OpISTHOTEUTHIDAE Verrill, 1896
Body gelatinous to semigelatinous; bell-shaped or disc-like; fins long to small, and lateral to su- perior (dorsal); simple crop present or absent; radula present or absent; gills “half orange”’ type; median pallial adductor present; web single; shell vestige U- or V-shaped. This family contains two genera: Opisthoteuthis and Grimpoteuthis (Voss 1988b).
Grimpoteuthis Robson, 1932
Tyre Species: Cirroteuthis umbellata Fischer, 1883:404. By original designation, Robson (1932: 136).
Grimpoteuthis bathynectes new species (Figs. 4, 5)
MATERIAL EXAMINED (15 specimens all collected by R/V YAQUINA).—Holotype: male, ML 50 mm, Cr. Y7210A haul 308, 45°01.1'N, 135°12.0’W in 3,932 m, USNM 730715. Para- types: 2 males ML 46.5-48 mm, | female ML 31 mm, Cr. Y7210A haul 300, 44°58.1'N, 132°14.7'W in 3,585 m, 10 June 1972, CAS 067789.—1 male, ML 42 mm, Cr. Y7105B haul 281, 44°38.5'N, 127°39.5’W in 2,816 m, 19 May 1971, SBMNH 35144.—1 male ML 30 mm, | female ML 85 mm, Cr. Y7210A haul 303, 45°05.1'N, 133°10.9’W in 3,700 m, 10 July 1972, UMML 31.1938.—1 female, ML 57 mm, Cr. Y7005C haul 232, 44°40.2'N, 133°35.7'W in 3,742 m, 3 June 1970, SBMNH 35145.—2 males, ML 17.5-53 mm, Cr. Y7210A haul 299, 44°56.8'N, 132°11.5'W in 3,580 m, 10 June 1972, UMML 31.1937.—1 male, ML 17 mm, Cr. Y7210A haul 307, 45°03.5'N, 134°45.0’W in 3,900 m, 10 Oct. 1972, CAS 067790.—2 males ML 30-36 mm, | female ML 29 mm, Cr. Y7210A haul 305, 45°05.2'N, 134°43.4’W in 3,900 m, 9 Oct. 1972, USNM 817582.—1 male, ML 28 mm, Cr. Y7206B haul 288, 44°06.2'’N, 125°22.7'W in 2,940 m, 14 June 1972, CAS 067791.
Diacnosis.— A Grimpotheuthis with long fins; short cirri; suckers largest proximally, suckers of males larger than those of females; gills compact with 7-9 lamellae; radula absent.
DEscRIPTION. — Mantle saccular (Fig. 4a), little more than half as wide as long, rounded poste- riorly; aperture narrow, of type B; head wider than mantle, not set off by neck region; eyes large, somewhat protruding.
Funnel large, projects far beyond mantle mar- gin, distal quarter or less free; funnel organ broad, thick-limbed, /A-shaped pad with all angles rounded.
Fins long, moderately wide, situated about halfway between apex of mantle and eyes; pos- terior margin nearly straight, tips rounded with
57
slight point, anterior margin convex, turning slightly posteriorly at line of mantle to form mi- nor constriction; fins with narrow muscular base, muscular area extends about three-quarters of length of fins.
Arms gelatinous, moderately long, subequal, stout, deeply set into thick web. Arm formula generally L.II.IILIV.
Web single (secondary web absent), thick, fleshy, soft; web formula variable, usually ABCDE; in most large males sector C on left side noticeably larger than C on nght; web depth about half arm length, extensive along arms but un- equally distributed; large fleshy nodule or finger- like process located on ventral side of arms (Fig. 4b), about one-half to two-thirds of arm length from base, unites web to ventral side of arms, beyond which web may extend only very short distance; nodules on dorsal side of arms absent, web extends to, or nearly to, tip of each arm. Consequently, sector A lacks processes, so web extends to tip of arm on both sides to make this sector deepest; conversely, web sector E is the shallowest with nodules on each side of arms.
Suckers in single row extend from mouth to tips of arms. Dorsal arms with 47 to 58 suckers set into oral surface, their apertures projecting freely; first four to five suckers small to minute, followed by abruptly larger suckers that gradually increase in size for about one-third arm length, after which they gradually decrease in size to arm tip. Suckers over proximal half of arms largest, abruptly enlarged suckers absent indicating lack of hectocotylization; considerable sexual dimor- phism in size and shape of suckers (Tables 5-8); in males, suckers (Fig. 4d) larger (SIn 4.3-6.7), more globular, with short, narrow apertural rims that rise from free globular bases where web usu- ally joins suckers; suckers in females and small males (Fig. 4c) smaller (SIn 2.6-3.5), globular base missing or nearly so, and whole sucker smaller, tubular, more erect than those of large males, without webbing between suckers.
Cirri short, about two-thirds to 1.5 sucker di- ameter in length; located on sides of arms be- tween bases of suckers (Fig. 4c, d), they first ap- pear between suckers 3 and 4 and extend to tips of arms.
Median pallial adductor thin, narrow, length 2.8-9.2% ML. Gills compact, spherical, of “half orange” or Opisthoteuthis type (Robson 1932: 124), 7-9 lamellae per gill, closely packed, slight- ly rugose in appearance.
58 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 47, No. 3
d | —
Ficure 4. Grimpoteuthis bathynectes new species. (a) Dorsal view of holotype, USNM 730715, male, 50 mm ML. (b) Oral view of left dorsal arm of male, 50 mm ML. (c) Suckers and cirri of female, 58 mm ML. (d) Suckers and cirri of male, 42
mm ML.
Digestive tract of male shown in Figure 5a; beaks shown in Figure 5d, e. Radula absent; an- terior salivary gland apparently absent, two small oval white bodies immediately adjacent and ven- tral to buccal mass may represent posterior sal- ivary glands; esophagus widens gently posterior
to buccal mass but does not form crop; stomach slightly two-parted; buccal mass, esophagus, and stomach densely covered with deep reddish-pur- ple gelatinous tissue, obscuring details until re- moved; caecum large with strong, almost com- plete turn; digestive ducts stout, lead from
VOSS AND PEARCY: DEEP-WATER OCTOPODS 59
pied sem.ves IIJ
=i) sem. ves. IT
sem. ves. I
stomach
testis
vas. def.
Ficure 5. Grimpoteuthis bathynectes new species. (a) Digestive tract of male, 53 mm ML. (b) Genitalia of female, 85 mm ML. (c) Genitalia of male, 53 mm ML. (d-e) Upper and lower beaks of male, 53 mm ML. (f) Left stellate ganglion of male, 53 mm ML. (g) Dorsal view of shell vestige and lateral view of end, male, 53 mm ML.
caecum-stomach to digestive gland, pass through Male genitalia complicated (Fig. 5c). Sper-
small but distinct digestive duct appendage area; matophores present, similar to those in Cirro- digestive gland small, globular; intestine small, — teuthis.
stout, straight; anal flaps and ink sac absent. Female genitalia (Fig. 5b) simple, unpaired;
PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 47, No. 3
60
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VOSS AND PEARCY: DEEP-WATER OCTOPODS
TasLe 7. Measurements (in mm) of four females of Grim- poteuthis bathynectes new species.
Specimen
USNM CAS SBMNH UMML
817582 067789 35145 31.1938 Mantle length 29 31 aiff 85 Mantle width 16 18 32 67 Head width 24 19 39 59 Arm length I V2 65 124 143 Arm length II 78 YP 104 150 Arm length III 73 60 101 143 Arm length IV 67 56 99 141 Total length 95 110 185 240 Fin length 26 28 33 39 Fin width 10 14 21 24 Sucker diameter 1.0 1.0 IES 3.0 Cirrus length 165 IES 1S 3.0 Gill lamellae 7 7 8 7 Suckers on I 51 51 58 58 Arm formula 2314 2134 1234 2134
ovary of largest female small with no mature eggs: proximal oviduct long, slender; oviducal gland two-parted, proximal third white, distal two-thirds very dark brown to almost black, both parts strongly and finely plicate; distal oviduct about one-third length of proximal oviduct, slen- der proximally but broadens distally, appears bulbous.
Shell vestige in male (Fig. 5g) deep, U-shaped, formed of rounded-edged flat strip that tapers to thin flat point distally; fin insertions long and form slight angle near base with rounded apex.
Stellate ganglion of male illustrated in Figure S16
Color of mantle and head grayish, tinged with purplish brown; posterior edges of fins lined with dark reddish brown; aboral surface of arms and web dark purple or reddish brown; oral surface of arms and web deep purplish brown or choc- olate, often with light brown or flesh-colored suckers.
MEASUREMENTS AND Counts.— Holotype: mantle length 50; mantle width 36, head width 41; arm length I 132, II 124, HI 115, IV 106; total length 170; fin length 42; fin width 21; suck- er diameter 3.0; cirrus length 2.2; number of gill lamellae 9; number of suckers on arm I 57.
Males: mantle length 17—36.2-53; mantle width 11—25.8-38; head width 17-—33.0-51; arm length I 45-99. 5-141, If 43—95.8-147, III 43-91.5-146, IV 39-86.5—-137; total length 70-—/35.8—200; fin length 19-34.2—43; fin width 6—/5.8—22; sucker
61
TABLE 8. Indices of bodily proportions and counts of four females of Grimpoteuthis bathynectes new species.
Specimen
USNM CAS SBMNH UMML
817582 067789 35145 31.1938 Mantle length 29 31 5)i/ 85 MwWI 55 58 56 79 HWI 83 62 69 70 MAI 36 42 46 56 ALI 84 66 67 63 SIn 3).5) 3 2.6 31.5 Cil SI) 4.8 2.6 Bh8) FLI 108.3 147.4 84.6 66.1 FWI 38.5 50.0 63.6 61.5 Arm formula 2314 2134 1234 2134
diameter 0.6—/.8—3.0; cirrus length 0.7—/.7-3.0; number of lamellae 7—7.7—9; number of suckers on arm I 47-50.5-57.
Females: mantle length 29-50.5-85; mantle width 16-33.3-67; head width 19-35.3-59; arm length I 72—/01.0-143, II 78—10/.0-150, II 60- 94.3-143, IV 56-90.8-141; total length 95- 157.5-240; fin length 26-3 7.5-39; fin width 10- 17.3-24; sucker diameter |—/.6-3; cirrus length 1.5-/.9-3.0; number of gill lamellae 7—7.3-8; number of suckers on arm I 51—54.5-58.
Type.— Holotype, male, 50 mm ML, USNM 730715 (fixed in formalin and preserved in 50% isopropyl alcohol).
TyYPE-LOCALITY.—Off Oregon, Tufts Abyssal Plain, 45°01 N, 135212:0' W, 3.932 am:
Discussion. — Only two species of Grimpoteu- this, G. hippocrepium (Hoyle, 1904) and G. al- batrossi (Sasaki, 1920) have been reported from the North Pacific Ocean (Voss 1988b). Grim- poteuthis hippocrepium was poorly described from a mutilated specimen and is unrecognizable as far as many important characters are concerned. The type (USNM 382467) is in very poor con- dition and little can be learned from it. Grim- poteuthis albatrossi is well described (Sasaki 1929: 7) but the type (USNM 332949) dried up, and although it has been realcoholed, it is nearly worthless for comparative studies. As Robson remarked (1932:150), G. albatrossi resembles the Atlantic G. grimaldi (Joubin, 1903) in having small fins and large suckers near the tips of arms. Neither of these species resembles G. bathy- nectes. It appears closely related to G. umbellata (Fischer, 1883), but differs from that species in the shape of the shell vestige, particularly in the muscle insertion, the digestive tract, the male
Pie Se LAI RA
Sates
FIGURE 6. (c-d) Upper and lower beaks of male, 72 mm ML.
genitalia, and the shape of the stellate ganglion. Examination of the syntypes in Museum Na- tional d’Histoire Naturelle, Paris, did not aid in clarifying the picture; the larger syntype had a different sucker arrangement but the shell vestige was missing; the syntype specimen may be a dif- ferent species.
While many problems exist in the genus, pres-
PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 47, No. 3
x oN
vi ee
self 4 5 00'0'626 Ortoee ¥ id ) >
Weovaryrdh
Grimpoteuthis tuftsi new species. (a) Dorsal view of male, 93 mm ML. (b) Oral view of left dorsal arm of same.
ent material is sufficiently distinct to warrant de- scription as a new species. The specific name bathynectes is from the Greek and means deep swimmer.
Thirteen specimens of G. bathynectes were col- lected from Tufts Abyssal Plain and two were from Cascadia Plain. Sampling effort was greater on the Cascadia Plain, hence this species inhabits
VOSS AND PEARCY: DEEP-WATER OCTOPODS
63
FIGURE 7.
deeper offshore waters and may be found in other abyssal waters of the North Pacific.
Grimpoteuthis tuftsi new species (Figs. 6-8)
MATERIAL EXAMINED (7 specimens, all collected by R/V YA- QUINA):—Holotype: male, ML 101 mm, Cr. Y7210A haul 305, 45°05.2'N, 134°43.4’W in 3,900 m, 9 Oct. 1972, USNM 730714. Paratypes: 1 male, ML 72 mm, Cr. Y7210A haul 300, 44°58.1'N, 132°14.7’W in 3,585 m, 6 Oct. 1972, UMML 31.1939.—1 male, ML 93 mm, Cr. Y7210A haul 302, 44°58.0'N, 133°14.5’W in 3,700 m, 7 Oct. 1972, CAS 067792.— 1 male, ML 100 mm, Cr. Y7210A haul 306, 45°02.0'N, 134°42.2'W in 3,900 m, 9 Oct. 1972, SBMNH 35146.—1 fe- male, ML 65 mm, Cr. Y7210A haul 307, 45°03.5'N, 134°45.0’'W in 3,900 m, 10 Oct. 1972, UMML 31.1940.—2 females, ML 74-102 mm, Cr. Y7210A haul 305, 45°05.2'N, 134°43.4'W in 3,900 m, 9 Oct. 1972, USNM 817583.
Diacnosis.—A Grimpoteuthis with long fins; long cirri; suckers largest distally at web border, suckers of males and females of equal size; gills loosely arranged, with 7-8 lamellae; radula pres- ent.
DESCRIPTION. — Mantle (Fig. 6a) saccular, flac- cid, width nearly three-quarters length, rounded posteriorly; aperture moderately narrow (27-39%
Grimpoteuthis tuftsi new species. View of web, arms and suckers.
ML); head wider than mantle, not set off by neck region; eyes large, protrude slightly.
Funnel large, broad, projects well beyond mantle margin, distal third or quarter free; funnel organ thick; A-shaped pad covers most of fun- nel; olfactory organ distinct round tubercle on each side of funnel at corners of aperture.
Fins long, moderately wide, united to mantle about halfway from apex; posterior margin near- ly straight for about three-quarters of its length, then turns forward; anterior margin rounded, tip somewhat pointed, base narrow, stout; muscular portion of fins extend for more than half fin length.
Arms somewhat gelatinous, moderately long, stout, deeply set in web. Arm formula generally LIL.IIL.1V. but varies.
Web (Fig. 7) simple, thick, fleshy; secondary web absent; web formula variable, generally ABCDE; web depth about half arm length; web unequal on two sides of arms (Fig. 6b): on dorsal side web extends nearly to tip of arm, on ventral side nodule or lappet present at about midpoint of arm, apparently strengthener for web attach- ment; web ends slightly distal to nodule.
Suckers (Fig. 8a) extend in single row from
64 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 47, No. 3
Re NS i < & NS
\\ aN f | \ f \ 1 \ / aN y: \ ae nee Sf \ * 2 j k FIGuRE 8.
| DAQ 5 Gang
Q ~
Grimpoteuthis tuftsi new species. (a) Lateral view of suckers and cirri of male, 93 mm ML. (b) Digestive tract of
male, 72 mm ML. (c) Genitalia of male, 72 mm ML. (d) Genitalia of female, 65 mm ML. (e) Radula of male, 72 mm ML. (f) Radula of female, 65 mm ML. (g-h) Radula of male, 93 mm ML; g is uncleaned, h cleaned with ribbon transparent. (i) Stellate ganglion from right side of male, 100 mm ML. (j-k) Ventral and dorsal view of shell vestige of male, 72 mm ML.
mouth to tip of arms; dorsal arms with about 63 to 75 small suckers (SIn 2.5—5.3) set into oral surface of arms with raised apertures; suckers cylindrical or tubular with small apertures, set about two to three sucker diameters apart in some sections but crowded together in others, united by thin transparent web or membrane along oral surface of arms (Fig. 8a). First 6-7 suckers small, then a gradual increase in size to their maximum
diameter at or just beyond level of ventral nod- ule. No sexual dimorphism (Tables 9, 10).
Cirri moderately long, longest 1.5 to 3.5 times longer than diameter of largest suckers; first ap- pear between suckers 5 and 6, or 6 and 7 and extend to arm tip; difficult to distinguish near tip.
Median pallial adductor thin, narrow, length 5.4-11.8% mantle length. Gills nearly spherical
VOSS AND PEARCY: DEEP-WATER OCTOPODS 65 TABLE 9. Measurements (in mm) of four males and three females of Grimpoteuthis tuftsi new species. Specimen
UMML CAS SBMNH USNM UMML USNM USNM
31.1939 067792 35146 730714 31.1940 817583 817583 Sex M M M M FE FE F Mantle length 72 93 100 101 65 74 102 Mantle width 49 75 69 63 50 52 64 Head width 60 101 98 73 60 67 78 Arm length I 289 309 322 312 207 224 285 Arm length II 275 288 320 295 186 222 238 Arm length III 260 270 283 258 170 210 250 Arm length IV 217 261 261 239 _ 208 253 Total length 346 457 475 460 285 322 460 Fin length 65 90 95 85 59 60 81 Fin width 30 38 40 31 27 29 35 Sucker diameter 3.8 4.0 4.5 DS DES DES 3.0 Cirrus length 6.0 9.0 8.0 9.0 9.0 6.0 5.0 Gill lamellae 8 8 8 7 8 8 7 Suckers on I 71 67 73 63+ 69 74 Ws, Arm formula 1234 1234 1234 1234 1234 1234 1432
(“half orange” type), 7 to 8 lamellae per gill, not tightly compacted as in G. bathynectes, lamellae rather loose and somewhat elongate in overall appearance; filaments branch off each lamella distinctly separated, to form overlapping leaflike layers.
Digestive tract of 72 mm ML male (Fig. 8b). Beaks (Fig. 6c, d) lack specifically distinct char- acters. When beaks removed from buccal mass, dark brown radular ribbon discovered. Radulae removed from three animals, cleaned in KOH, and mounted in CMC10. Ribbon delicate, all of dark brown pigment lost and ribbon partly eaten away in four or five seconds. Radulae show great variation (Fig. 8e—-h). Female of 65 mm with five homodont teeth, rachidian tall, slender (Fig. 8f);
second and third laterals broad to slender tri- angular, both narrow on one side of band, wide on other; traces of small first laterals in some rows on one side but not on other; no trace of marginal teeth or plates. Radula from 72 mm male (Fig. 8e) with five teeth in a row, three median ones (rachidian and second laterals?) elongate triangular; third laterals broad, flat- topped teeth, those on one side sharply truncate. Teeth from 101 mm male (Fig. 8g, h) all broadly triangular.
Posterior salivary glands absent; esophagus widens in mid-part to form simple crop, entire length covered by dark purple tissue. Esophagus narrows abruptly before entering small round stomach; spiral caecum equal in size to stomach,
Taste 10. Indices of bodily proportions of four males and three females of Grimpoteuthis tuftsi new species. Specimen UMML CAS SBMNH USNM UMML USNM USNM 31.1939 067792 35146 730714 31.1940 817583 817583 Sex M M M M F In FE Mantle length 72. 93 100 101 65 74 102 MwI 68.1 80.7 69.0 62.4 76.9 70.3 62.8 HWI 83.3 108.0 98.0 723 92.3 90.5 76.5 MAI 24.2 30.0 29.4 31.9 30.7 Biles 34.0 ALI 85.8 67.8 71.6 69.6 64.4 73.0 65.2 SIn 553 4.3 4.5 DS 3.9 3.4 2.9 cil 8.3 9.7 8.0 8.9 13.9 8.1 4.9 FLI 108 89 97 116 98 90 104 FWI 46 42 42 37 46 48 43 Arm formula 1234 1234 1234 1234 1234 1234 1432
66 PROCEEDINGS OF THE CALIFORNIA ACADEMY OF SCIENCES, Vol. 47, No. 3
two ducts lead to large round digestive gland; digestive duct appendage prominent; intestine short, inflated; anal flaps and ink sac absent.
Male genitalia complex (Fig. 8c), differ from those of G. bathynectes; various glands and their numbering not yet determined; both genitalia similar, spermatophores absent.
Female genitalia (Fig. 8d) simple (immature); large, deflated, thin-walled ovary, eggs absent; proximal oviduct thick, comparatively short; oviducal gland large, two-parted, finely plicate; distal oviduct long, comparatively slender, ap- erture with flared, incised, or plicate collar.
Shell vestige (Fig. 8j, k) U- or rounded V-shape; stout, strong, somewhat constricted at posterior end; limbs angle slightly at outer posterior in- ception of fin insertion area, taper to thin, flat, straplike, truncated tips.
Stellate ganglion of male shown in Figure 81.
Mantle and head pigmented light brownish purple, posterior borders of fins dark purple; web, arms darkish purple aborally, dark purple orally with lighter colored suckers and cirri.
MEASUREMENTS AND Counts.— Holotype: mantle length 101; mantle width 63; head width 73xarmylencth 132. e295. Tl 258 -3nV 239: total length 460; fin length 85; fin width 31; suck- er diameter 2.5; cirrus length 9.0; number of gill lamellae 7; number of suckers on arm 63+.
Males: mantle length 72-9/.5-101; mantle width 49-64-75; head width 60-83-101; arm length I 289-308-322, If 275-294. 5-320, II 258- 267.8-283, IV 217-244.5-261; total length 346- 434.5475: fin length 65—83.8—95; fin width 30- 34.8-40; sucker diameter 2.5-3.7-4.5; cirrus length 6—8.0-9; number of gill lamellae 7-7.8- 8; number of suckers on arm I 67—70.3-73.
Females: mantle length 65-80.3-102; mantle width 50—55.3-64; head width 60-68.3-78; arm length I 207—238.7-285; II 186-—2/5.3-238; Ill 170-210-250, IV 208-253; total length 285-—355- 460; fin length 59-66.7-81; fin width 27-30.3- 35; sucker diameter 2.5—2.7—3.0; cirrus length 5— 6.7-9; number of gill lamellae 7—7.7—8; number of suckers on arm I 69-72.7-75.
Type. — Holotype, male, 101 mm ML, USNM 730714 (fixed in formalin and preserved in 50 percent isopropyl alcohol).
TYPE-LOCALITY.—Off Oregon on the Tufts Abyssal Plain, 45°05.2'N, 134°43.4’W, 3,900 m.
Discussion.—Grimpoteuthis tuftsi is some- what similar to G. bathynectes but many char- acters differentiate them. Grimpoteuthis tuftsi has:
lack of sexual dimorphism in sucker shape, long- er cirri, different genitalia, more elongate gills with separate lamellae, enlarged suckers near nodules, larger funnel organ, and a radula.
The discovery of a radula in this species, un- known elsewhere in the cirrate octopods, re- quires a careful reexamination of all species, as its absence has been taken for granted by some earlier students of the group and may have been overlooked. It is obviously degenerate and in its simplicity, variability, and loss of teeth resem- bles the radula of Graneledone pacifica (de- scribed in this work) and G. antarctica (Voss, 1976). Similarly, the radulae of Thaumeledone and Vosseledone show degeneration in the near loss of all teeth except the rachidian. We hy- pothesize that in these genera the radula is de- generate and in the process of being lost because of diet and loss of selective pressure.
All specimens of G. tuftsi were caught in bot- tom trawls on Tufts Abyssal Plain, none in Cas- cadia Basin nearer the North American conti- nent.
Suborder INcIRRATA Grimpe, 1916
Deep-sea, shallow-water, and pelagic octopods with gelatinous to muscular bodies; fins and cirri absent; shell, when present, reduced to pair of slender cartilaginous or calcareous stylets; radula present in all known species; suckers uniserial or biserial; representatives known from throughout world oceans.
Family OcTopoDIDAE Orbigny, 1845
Benthic octopods with semigelatinous to mus- cular bodies.
Subfamily BATHYPOLYPODINAE Robson, 1928
Restricted to benthic octopods with biserial suckers; hectocotylus well developed; crop large with diverticulum reduced or absent; ink sac ab- sent; radula Octopus-like to degenerate.
Benthoctopus Grimpe, 1921
Deepwater octopods of normal Octopus-like appearance with short to long arms; suckers bise- rial; hectocotylus Octopus-like, ligula slightly to moderately excavated with indistinct midrib, smooth or bearing low, often indistinct rugae,
VOSS AND PEARCY: DEEP-WATER OCTOPODS
never laminate; crop present, usually with di- verticulum; ink sac absent; radula usually with strongly, seldom weakly, multicuspid rachidian; body entirely smooth, papillae or ocular cirri ab- sent.
Tyee Species: Octopus piscatorum Verrill, 1879:470. By orig- inal designation, Grimpe 1921:299.
DiscussIon.— Examination of Verrill’s type specimen (USNM 574641) has shown that it probably is not a Benthoctopus but more likely belongs to Bathypolypus (personal observation by Voss). If this proves correct, the name Ben- thoctopus should be preserved to maintain no- menclatural stability in this group.
Benthoctopus robustus new species (Figs. 9, 10)
MATERIAL EXAMINED (2 specimens).— Holotype: male, ML 137 mm, R/V ACONA Sta. NAD22, 44°38.5'N, 126°03.8’W in 2,800 m, 1 June 1963, USNM 729994. Paratype: 1 male, ML 114 mm, 140 nautical miles off Tasu Sound, Queen Char- lotte Island, B.C., Canada, 52°27'42’N, 135°34'36’W in 3,660 m, 16 Oct. 1978, in sablefish trap, E. Houde, USNM 730895.
Diacnosis. —A Benthoctopus with large, strong, transversely ridged lingula; long, sharp calamus; radula with simple rachidian with cusps.
DESCRIPTION. — Body (Fig. 9a) compact, ro- bust, muscular; mantle nearly round, about as wide as long (MWI 80.3), no distinct neck region; head small, narrow (HWI 51.8); eyes and eyelids small; mantle aperture wide.
Funnel large, free for about half of its length, aperture small; funnel organ (Fig. 9f) large, W-shaped, with long, pointed lateral limbs that project slightly anterior to large, round median limb.
Arms (Table 11) moderately long (MAI 64.3; ALI 56.7), stout at base (AWI 12.4), taper to stout, sharp tips, arm formula IV.I.II.III, sub- equal; web heavy, deep (WDI 34.7), extending along ventral side of arms for about two-thirds their length; web formula DBCAE; suckers bise- rial, erect on stout bases, well separated, rather small (SIn 5.1) for size of animal; enlarged suck- ers absent; dorsal arms with about 38-40 suck- ers.
Third right arm hectocotylized, shorter than its fellow (HcAI 85.1), stout, bordered ventrally by heavy, thickened, in-rolled web that forms incomplete spermatophoral groove, ligula (Fig. 9b) large (LLI 12.4), with deep groove and thick- ened margins crossed by 14-16 grooves and
67
membranous ridges or folds; calamus large, long, slender, sharply pointed (CLI 45.4).
Gills large with 11 lamellae per outer demi- branch.
Upper beak with very small, sharp, curved ros- trum (Fig. 9g); rostrum of lower beak forms right angle at jaw angle (Fig. 9h). Radulae with rachid- ian somewhat different from other Pacific Ben- thoctopus (Fig. 10d, e), rachidian simple, narrow, with smooth-sided, slender darkened portion bordered by narrow clear portion that bears small, irregular cusps, one on each side; admedians with tall sharp ectocone; inner edges of second laterals strongly, smoothly rounded in holotype, nearly straight in paratype; third laterals slender, sabre- like, curved.
Esophagus slender, leads into distinct crop with anterior diverticulum (Fig. 9d, e); crop of holo- type unexpanded, without food; posterior esoph-