MARIAX H. PETT1B0NE ReVlSWTl OJ

Some Species

Referred to

Leanira Kinberg

(Polychaeta: Sigalionidae)

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • 1970 NUMBER 53
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SMITHSONIAN CONTRIBUTIONS TO

ZOOLOGY

NUMBER 53

Marian H. Pettibone Revision o f

Some Species

Referred to

Leanira Kinberg

(Polychaeta: Sigalionidae)

SMITHSONIAN INSTITUTION PRESS

CITY OF WASHINGTON

I97O
ABSTRACT

Pettibone, Marian H. Revision of some species referred to Leanira Kinberg (Polychaeta: Sigalionidae). Smithsonian Contributions to Zoology 53: 1-25. 1970.—

The numerous sigalionid species described originally as members of the genus Leanira

Kinberg, or subsequently referred to it, form a heterogeneous group. Among them, two

closely related groups are recognized and assigned to two genera: Leanira Kinberg,

limited, consisting of seven species, one of which is new, and one synonym, and

Ehlersileanira, new genus, comprising a single species and three synonyms. Definitions

of the two genera, with a key to the species of Leanira, are followed by redescription

of the species, based in large part on examination of the type-specimens.

Official publication date is handstamped in a limited number of initial copies and is recorded

in the Institution's annual report, Smithsonian Year.

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Marian H. Pettibone Revision of

Some Species

Referred to

Leanira Kinberg

(Polychaeta:

Sigalionidae)

Among the scaled polychaetous annelids of the superfamily Aphroditoidea, some of the sigalionid species

that have been referred to the genus Leanira Kinberg

are reviewed and revised, based on an examination of

available type-specimens and other specimens for

which there are published records. In addition to the

sigalionid collections in the Smithsonian Institution

(USNM), material was obtained from the following

Museums: British Museum (Natural History), London (BMNH), through J. D. George; Museum of

Comparative Zoology, Harvard (MGZ), through

H. W. Levi; Naturhistoriska Riksmuseet, Stockholm

(NRS), through R. Olerod; Queensland Museum,

Brisbane (QMB), throughB. Campbell; Rijksmuseum

van Natuurlijke Historie, Leiden (RNHL), through

J. van der Land; Zoologische Museum Universiteit

van Amsterdam (ZMA), through S. van der Spoel;

Zoologisches Museum, Berlin (ZMB), through G.

Hartwich; and Zoologisches Staats-museum, Hamburg

(ZMH), through G. Hartmann-Schroder.

I take this opportunity to thank the above-mentioned individuals for their cooperation and help in

arranging for the loan of the sigalionid material on

which this study is based. The manuscript benefited

from the suggestions of Horton H. Hobbs, Jr., and

Nancy M. Foster, both of the Smithsonian Institution.

Marian H. Pettibone, Curator, Department of Invertebrate

Zoology, National Museum of Natural History, Smithsonian

Institution, Washington, D.C. 20560.

Some thirty-eight sigalionid species have been described as members of the genus Leanira Kinberg,

1855, or subsequently referred to it (Hartman, 1959,

1965a—Catalogue). They represent a heterogeneous

group of species. Generally those sigalionid polychaetes

having spinigerous or distally pointed, camerated or

canaliculate, compound neurosetae have been assigned

to the genus Leanira. Willey (1905: 259) proposed

the genus Sthenolepis to receive those species in which

the median antenna has a distinct ceratophore

equipped with lateral auricles or ctenidia, separating

them from species of Leanira in which the ceratophore

is indistinct and lacks lateral auricles. The separation

of the two groups has not been generally accepted.

Both Horst (1917: 115) and Monro (1936: 106)

indicated that the distinction was unjustified. Hartman

(1965b) emended the two genera by following Willey

(1905) in assigning to Leanira those sigalionid species

in which the prostomium lacks antennal auricles and

the neurosetae are compound, distally pointed, and

camerated and to Sthenolepis those species with similar

neurosetae but with auricles on the ceratophore of

the median antenna. Of the species referred to the

latter by Hartman (1965a,b), some of them show closer

relationships to the type-species of Leanira—L. quatrefagesi Kinberg, 1855—than they do to the type-species

of Sthenolepis Willey—Leanira japonica Mclntosh,

1885. For this group of species, the new genus Ehlersileanira is proposed. The species of Leanira and Ehlersileanira considered here are redescribed. The other
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

species will be treated in subsequent studies of the

Aphroditoidea.

All the species, except Eupholoe cirrata Treadwell,

which are referred to Leanira in this revision were

described originally as members of that genus. The

genus Leanithalessa Hartmann-Schroder, 1965, with

type-species L. antennata Hartmann-Schroder, is

referred herein to Leanira. The following species are

included in Leanira (synonyms in parentheses):

(1) L. quatrefagesi Kinberg, 1855. Southwestern Atlantic,

off Argentina. (Leanithalessa antennata HartmanSchroder, 1965. Southeastern Pacific, off southern

Chile).

(2) L. hystricis Ehlers, 1874. Northeastern Atlantic, off

Ireland. (Leanira laevis Mclntosh, 1874. Nomen

nudum. Northeastern Atlantic.)

(3) L. robusta Verrill, 1885a,b. Northwestern Atlantic, off

New England.

(4) L. alba Moore. 1910. Northeastern Pacific, off southern

California.

(5) L. coeca Horst, 1917. Indo-Pacific, Malay Archipelago.

(6) L. cirrata (Treadwell).

(Eupholoe cirrata Treadwell, 1934. Northwestern Atlantic, off Puerto Rico.)

(7) L. adenensis, new species.

(L. vulturis.—Monro, 1937. [Not Horst, 1917.] Gulf

of Aden.)

The species referred herein to Ehlersileanira were

described originally under Sthenelais Kinberg by

Grube (1877), and Ehlers (1887), and under Leanira

Kinberg by Horst (1917), and Takahashi (1938).

They were transferred to Sthenolepis Willey by Hartman (1965a, b). The following species is referred to

Ehlersileanira, new genus (synonyms in parentheses):

E. incisa (Grube), new combination.

(Sthenelais incisa Grube, 1877. Southeastern Atlantic,

off West Africa).

(Sthenelais simplex Ehlers, 1887. Northwestern Atlantic, off Florida).

(Leanira vulturis Horst, 1917. Indo-Pacific, Malay

Archipelago.)

(Leanira izuensis Takahashi, 1938. Northwestern Pacific, Izu Peninsula, Japan.)

Abbreviations Used in the Figures

ac, aciculum

acL, acicular lobe

au, auricle

a-vB, anterior-ventral bract

br, branchia

buC, buccal cirrus

ci, cirrophore

ct, ctenidia

dTc, dorsal tentacular cirrus

dTu, dorsal tubercle

fTu, facial tubercle

IpaS, inner palpal sheath

laL, labial lobe

lAn, lateral antenna

mAn, median antenna

ne, neuropodium

no, notopodium

nuO, nuchal organ

OpaS, outer palpal sheath

pa, palp

p-lB, posterior lower bract

p-uB, posterior-upper bract

sP, segmental papilla

st, stylode

vC, ventral cirrus

vTc, ventral tentacular cirrus

General Characteristics of Species of Leanira Kinberg

and Ehlersileanira, New Genus

BODY SHAPE.—The body is elongate, with numerous

segments, tetragonal, flattened dorsoventrally, and

gently tapered posteriorly.

ELYTRA, DORSAL TUBERCLES, BRANCHIAE, and PARAPODIAL CTENIDIA.—The elytra are located on segments

2, 4, 5, 7, on alternate segments to 25 or 27, and on

all succeeding segments. The elytra are small and oval

on the first few segments, becoming progressively larger,

subrectangular to subcordiform in shape, and nearly

covering the middorsum; papillae and tubercles are

lacking except for occasional minute sensory papillae;

some have lateral indentations (Figures 2a,e-g; 6n—q;

llg—i). Dorsal cirri are absent. Conical dorsal tubercles occur on segments 6, 8, and on alternate segments

to 24 or 26 (Figures 3d; lib). Branchiae and parapodial ctenidia are absent from the anterior region.

Cirriform branchiae begin about segment 30 but there

may be smaller rudimentary ones on the elytriphores or

dorsal tubercles of more anterior segments. The parapodial ctenidia begin at about the same region as the

branchiae; they may be inconspicuous, the three

ctenidia per parapodium being nearly contiguous (Figures 1/; 3h,i; 11/).

PROSTOMIUM, TENTACULAR (i), and FEW ANTERIOR

SEGMENTS (n-m).—The prostomium is oval to subrectangular in shape and partially fused to the ten-
NUMBER 5 3

tacular parapodia, which are anteriorly directed in a

position dorsal to the long palps (Figures la-c; 2a,b;

lOa-d). There are three antennae; in Leanira, the

median antenna, lacking auricles or antennal ctenidia,

emerges from a prostomial ridge (Figures la—c; 2a,b),

while in Ehlersileanira, the ceratophore of the median

antenna is distinct and has a pair of rather small lateral

auricles or cenidia (Figures 10a; 12a). The presence

or absence of lateral auricles or ctenidia is the principal diagnostic characteristic separating the two genera.

The styles of the median antennae are small, subulate

and may be 2-articled. The ceratophores of the

lateral antennae are fused to the basal parts of the

tentacular parapodia. The palps emerge ventral to the

tentacular parapodia between large inner and smaller

outer palpal sheaths; they are long, extending posteriorly to about segment 15 (10-26). Inner ciliated

tentacular lobes are lacking. Medial to the inner palpal

sheaths is a bulbous facial tubercle (Figures 5c; 10b).

A pair of small rounded nuchal organs occur posterolateral to the prostomium; they are usually visible only

when the pharynx is at least partially extended (Figures la; l0a,c). Each tentacular parapodium is supported by a single aciculum, bears two bundles of

capillary setae directed anteriorly, somtimes a few to

numerous stylodes, and a pair of tentacular cirri with

distinct cirrophores emerging close together; the dorsal

tentacular cirrus is much longer than the ventral one

(Figures 2c,d; 8a,b; lOa-d).

The parapodia of segments 2 and 3 are directed

anteriorly and contribute to the lateral and lower lips.

The lateral lips may be provided with accessory labial

lobes (Figures 26,0*; 4b). The ventral buccal cirri of

segment 2 are thicker and slightly longer than the following ventral cirri. Segment 3 lacks dorsal tubercles

and dorsal cirri. The parapodial lobes and setae are

similar to those of the following segments except the

parapodial stylodes may be more numerous (Figures

3a-c; We-g).

PARAPODIA.—The biramous parapodia have clavate

notopodia which are smaller than the neuropodia (Figures 3d-j; 8/-/; 11a-/). On the posterodorsal sides of

the notopodia there are subterminal diagonal bracts

provided with stylodes, while larger terminal stylodes

are found near the tips of the acicular lobes. The notosetae form a half circle within the subterminal bracts;

they are slender, finely to coarsely spinous, tapering to

capillary tips. The neuropodia have bilobed posterior

bracts formed of large upper lobes directed anteriorly

and provided with stylodes; the lower lobes are smaller,

directed anteriorly, and provided with stylodes. The

anterior acicular lobes are subconical, with low or inconspicuous anteroventral bracts. The neurosetae are

compound spinigers; the stems are smooth, the blades

canaliculate or camerated and taper to fine tips (Figures le,g; 10/). The neurosetae of the anterior segments are similar to those of the following segments,

none being markedly long. The neurosetae are arranged in three series: doubly curved vertical rows

anterodorsal to the posterior-upper bracts, subacicular

transversely curved rows dorsal to the posterior-lower

bracts, and diagonal anteroventrally arched rows

within the anteroventral bracts. In addition, there may

be a few (2-5) simple spinous neurosetae in the upper

parts of the neuropodia (Figures 5/; 9g; lid). The

ventral cirri are slender, tapered, and do not extend

to the distal tips of the neuropodia; small bulbous lobes

are found on the outer bases of the cirri.

PHARYNX, PYGIDIUM, and SEGMENTAL PAPILLAE.—

The muscular eversible pharynx is equipped with 11

pairs of distal papillae and two pairs of jaws. The

pygidium has not been described; presumably it is

equipped with a pair of anal cirri. Segmental papillae

are found on some species, beginning about segment 30

(23-36); they are elongate, tubular, and located anteromedial to the ventral cirri (Figure 6/,A;).

Genus Leanira Kinberg

Leanira Kinberg, 1855, Type-species: L. quatrefagesi Kinberg, 1855, by monotypy. Gender: feminine.

Leanithalessa Hartmann-Schroder, 1965. Type-species: L.

antennata Hartmann-Schroder, by original designation and

monotypy. Gender: feminine. —L. quatrefagesi Kinberg.

DIAGNOSIS.—Body elongate, with numerous segments. Prostomium oval, partially fused to tentacular

parapodia. Three small subulate antennae; ceratophore of median antenna without auricles or ctenidia;

lateral antennae fused to tentacular parapodia. Long

palps emerging ventral to tentacular parapodia between large inner and smaller outer palpal sheaths.

Bulbous facial tubercle medial to inner palpal sheaths.

Tentacular parapodia (I) directed anteriorly, with

single aciculum, pair of tentacular cirri, and 2 bundles

of capillary setae. Elytra on segments 2, 4, 5, 7, on

alternate segments to 25 or 27, and on all succeeding

segments. Elytra smooth, lacking tubercles and papillae.

Dorsal cirri and dorsal tubercles lacking on segment 3.

Cirriform branchiae and elongate platelike ctenidia,
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

3 per parapodium, from about segment 30 posteriorly.

Notopodia with well-developed bracts with stylodes.

Neuropodia with bilobed posterior bracts bearing

stylodes. Notosetae spinous capillaries. Neurosetae consisting of compound spinigers, with blades relatively

short and canaliculate. With or without additional simple neurosetae. Ventral cirri short and tapered.

Pharynx with 11 pairs of papillae and 2 pairs of jaws.

REMARKS.—The holotype and six paratypes of

Leanira quatrefagesi from off La Plata River, Argentina, deposited in the Naturhistoriska Riksmuseet,

Stockholm (NRS 431 and 223), and the holotype and

three of the paratypes of Leanithalessa antennata from

off Chile, deposited in the Zoologisches Staats-museum,

Hamburg (ZMH 438), were examined. The two

species are herein considered to be synonymous.

Key to the Species of Leanira Kinberg

1. Tentacular parapodia (I) with numerous stylodes (Figure 8a,b). Lateral antennae inserted

medially on tentacular parapodia (Figure 86). [Elytra on all segments from 27 posteriorly,

with slight lateral indentations (Figure 8n~p). Few simple spinous neurosetae (Figure 8k).

Without distinct labial lobes.] L. cirrata (Treadwell)

1'. Tentacular parapodia (I) with none to few stylodes (Figures \a,b; 2a; 4a; 5a,c,d; 6a,c,d;

7a; 9a). Lateral antennae inserted dorsally on tentacular parapodia (Figures la,b; la; 4a;

5a; 6a; 7a; 9a) 2

2. With labial lobes (Figures \c; 2b; 4b; 96) 3

2'. Without labial lobes (Figures 56; 66; 76). [Elytra on all segments from 27 posteriorly, with

distinct lateral indentations. (Figures 5m,n; 6p,q; 7m,n)] 5

3. With simple neurosetae (Figure 9g). Elytra on all segments from 27 posteriorly, with lateral

indentations (Figure 9l,m,). [Labial lobes small, globular (Figure 96).]

L. adenensis, new species

3'. Without simple neurosetae. Elytra without distinct lateral indentations (Figure 2e-g) 4

4. Neuropodial stylodes relatively few, large, clavate (Figure 4g-4). Labial lobes bulbous (Figure

46). Elytra on all segments from 25 posteriorly L. hystricis Ehlers

4'. Neuropodial stylodes more numerous, digitiform (Figures \d,f; 3e). Labial lobes auricular

(Figures \c; 26). Elytra on all segments from 27 posteriorly L. quatrefagesi Kinberg

5. Without simple neurosetae. [With tubular segmental papillae (Figure 7;)] L. coeca Horst

5'. With simple neurosetae (Figures 5/; 67) 6

6. With few stylodes on tentacular parapodia (I) (Figure ba,c,d). Without tubular segmental

papillae L. robusta Verrill

6'. Without stylodes on tentacular parapodia (I) (Figure 6a,c,d). With tubular segmental

papillae, beginning about segment 30 (Figure 6j,k) L. alba Moore

Leanira quatrefagesi Kinberg

FIGURES 1-3

Leanira quatrefagesi Kinberg, 1855, p. 388; 1858-1910, p. 30,

pi. 9: fig. 42.—Ehlers, 1901, p. 59, pi. 5: fig. 8.—Monro,

1924, p. 46; 1936, p. 105, fig. 13.—Fauvel, 1941, p. 277.—

Wesenberg-Lund, 1962, p. 27.—Hartman, 1965a, p. 51;

1967, p. 40.

Leanira hystricis.—Mclntosh, 1924, p.14; 1925, p. 38.—Day,

1963, p. 360; 1967, p. 112, fig. 1.21, a-d. [Not Ehlers,

1874.]

Leanithalessa antennata Hartmann-Schroder, 1965, p. 88,

figs. 36-40.

MATERIAL EXAMINED.—South Atlantic, off mouth of

La Plata River, Argentina, 91 meters, Eugenie Expedition—holotype (NRS 431) and 6 paratypes (NRS 223)

of Leanira quatrefagesi. Puerto Angusto, Magellan

area, South America, 18 meters, 25 March 1896, Nordenskjold, collector, Ehlers, determined—3 specimens

(NRS 1612). Strait of Magellan, 4-58 meters, H. M. S.

Alert—3 specimens (BMNH 1925: 1: 28: 41^3).

Cape of Good Hope, South Africa, Mclntosh collection (as L. hystricis)—1 specimen (BMNH 1924: 7:

21: 23). South Pacific, off Chile, Station 93, 15 March

1960, mud with fine sand and detritus, 84 meters—

holotype (ZMH 438); 1 paratype (ZMH) and 2 paratypes (USNM 40996) of Leanithalessa antennata.

DESCRIPTION.—The maximum length and width, including setae, are 185 and 7 mm, respectively, and the

maximum number of segments 140. The elytra are thin,

transparent, and found on all segments from 27 posteriorly; they are small and oval on the first few segments, become progressively larger and subrectangular

to subcordiform in shape posteriorly, nearly covering

the middorsum; distinct lateral indentations are lacking (Figures \a,b; 2e-g).
NUMBER 53

FIGURE 1.—Leanira quatrefagesi (a, NRS 1612; b-c, paratype of L. quatrefagesi, NRS 223; d-g,

holotype of L. quatrefagesi, NRS 431): a, Anterior end, dorsal view, pharynx partially extended;

right elytron removed; b, anterior end, dorsal view; left palp missing; right parapodia of II and

III not shown; c, anterior end, ventral view; d, parapodium from anterior region, posterior view;

e, upper, middle, and lower neurosetae from same; /, parapodium from middle region, anterior

view; g, upper and lower neurosetae from same.

The oval prostomium and tentacular segment (I)

are partially fused (Figures \a-c; la-d). The short

subulate median antenna may be 2- or 3-segmented,

attached to an indistinct ridge on the anterior half

of the prostomium; similar lateral antennae are attached to the inner dorsal sides of the tentacular parapodia, connected by indistinct ceratophores to the

prostomium. Eyes may be lacking (on types of L.

quatrefagesi; Figure \a,b; two eyes lateral to the

median antenna, according to Kinberg) or there may

be two pairs of eyes: a pair of larger rounded spots

anteriorly on the prostomium and a pair of small spots

in the middle of the prostomium; in addition, there

may be darker pigmented areas along the lateral bases

of the prostomial ridge (types of L. antennata; Figure

2a). Small rounded nuchal organs may or may not

be visible. The palps extend to about segment 14

(9-26). The inner palpal sheaths are large and subtriangular, the outer ones, short and rounded. Medial

to the inner palpal sheaths is a bulbous facial tubercle.

The tentacular cirri have distinct cirrophores; the dorsal tentacular cirri are much longer than the ventral

ones. There are usually 1-2 short stylodes medial to

the cirrophores of the dorsal tentacular cirri or the

stylodes may be lacking. There are two bundles of

capillary setae directed anteriorly. Auricular labial

lobes are found on the lateral lips; they may appear

as lateral extensions when the lips are somewhat ex-
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 2.—Leanira quatrefagesi (paratype of Leanithalenessa antennata, USNM 40996): a,

Anterior end, dorsal view, right elytra removed; palps not shown; b, anterior end, ventral view;

c, right tentacular parapodium (I), lateral or outer view; d, left tentacular parapodium, inner

view; e, right first elytron; /, right anterior elytron (seventh); g, right middle elytron.

tended (Figure Ic) or as medial extensions when the

lips are withdrawn (Figure 2b). The notopodia of

segments 2 and 3 have a circlet of stylodes; the neuropodia have upper and lower groups of stylodes; the

buccal cirri of segment 2 and the ventral cirri of segment 3 are short, thick and subconical; the neurosetae are similar to those of the following segments;

both dorsal cirri and dorsal tubercles are lacking on

segment 3 (Figures \a-c; 1a,b; 3a-c).

Branchiae are present from about segment 25 posteriorly, reaching full size about segment 30; occasionally smaller rudimentary branchiae occur more

anteriorly. The parapodial ctenidia begin at about the

same region as the branchiae; they may be inconspicuous, the 3 ctenidia per parapodium being nearly contiguous (Figures 1/; 3h,i).

The notopodia and neuropodia are subequal in

length (Figures ld-g; 3d-j). The small clavate notopodia have circlets of stylodes on subterminal posterodorsal bracts and a larger terminal stylode near

the tips of the acicular lobes. The notosetae are slender, coarsely to finely spinous, and taper to capillary

tips. The large neuropodia have large posterior-upper

and small posterior-lower bracts, both equipped with

stylodes, the latter being more numerous in the anterior parapodia. The acicular lobes are subconical

with low anteroventral bracts. The neurosetae are all

compound spinigers; the blades are canaliculate and

taper rather abruptly to fine tips. The neurosetae are

arranged in three series: doubly curved vertical rows

anterodorsal to the posterior-upper bracts; subacicular

curved rows dorsal to the posterior-lower bracts, and

ventral arched rows within the anteroventral bracts.

The neurosetae of the latter series are more slender

than the others. The ventral cirri are slender, tapered,

and do not extend to the distal tips of the neuropodia;

small bulbous lobes are found on the outer bases of

the cirri.
NUMBER 53

FIGURE 3.—Leanira quatrefagesi (paratype of LeanithaUnessa antennata, USNM 40996): a,

Parapodium from segment 2, posterior view; b, upper, middle, and lower neurosetate from same;

c, parapodium from segment 3, anterior view; d, parapodium from anterior region, anterior view;

e, same, posterior view; /, diagrammatic end-view of same, showing position of bracts and setae; g,

middle and lower neurosetae from same; h, parapodium from middle region, posterior view; i,

same, anterior view; j , upper, middle, and lower neurosetae from same.

875-841 O—70
8

DISTRIBUTION.—South Pacific off Chile, South

Atlantic off Argentina, Magellan area, Falkland Islands, South Africa, Antarctic. Intertidal to 4758

meters.

Leanira hystricis Ehlers

FIGURE 4

Leanira hystricis Ehlers, 1874, p. 292; 1875, p. 35, pi. 2:

figs. 5-11; 1908, p. 55.—Mclntosh, 1876, p. 408, pi. 73:

figs. 6-8; 1885, p. 155; 1900, p. 434, pi. 28: fig. 17, pi.

31: figs. 12, 13, pi. 42: figs. 20-22.—Fauvel, 1914, p. 84;

1923, p. 118, fig. 43, h-m.—Ditlevsen, 1917, p. 48.

Leanira laevis Mclntosh, 1874, p. 268 [Nomen nudum.]

MATERIAL EXAMINED.—Off southwest coast of Ireland, 1478 meters, Porcupine Expedition, station 2,

1869, Mclntosh collection—3 specimens (BMNH

1921:5: 1:653).

REMARKS.—The type-specimens of Leanira hystricis

Ehlers (1874, 1875), collected by the Porcupine Expedition in 1869 in the northeastern Atlantic, in 1214

to 2640 meters, apparently no longer exist. At least

they were not found in the Zoological Museum, Berlin;

Zoological Institute, Gottingen; or the Zoological

Museum, Hamburg (G. Hartwich, P. Kuenzer, and G.

Hartmann-Schroder, in litteris). Among the collections of Mclntosh, now deposited in the British Museum, are three specimens of the species from the same

Expedition, Porcupine station 2, and reported by

Mclntosh in 1876. The specimens are comparatively

small, 18 mm in length for 51 segments (incomplete

posteriorly) and 2.5 mm in width, including parapodia; all the elytra are now missing and most of the

setae are broken. Mclntosh (1876: 408) referred

Leanira laevis Mclntosh (1874: 268), nomen nudum,

to L. hystricis Ehlers, 1874.

The specimens from South Africa, reported by Mclntosh (1924: 14; 1925: 38) and by Day (1963: 360;

1967: 112) as L. hystricis, are referred herein to L.

quatrefagesi. Whereas Hartman (1942: 105) synonymized Eupholoe cirrata Treadwell with L. hystricis,

the species are here considered to be distinct. Pettibone

(1963: 53) considered L. robusta Verrill and L. alba

to be synonyms of L. hystricis, but they, too, are now

believed to be valid.

DESCRIPTION.—The elytra are thin, transparent, and

found on all segments from 25 posteriorly; they are

small, rounded to elongate-oval, and do not cover the

middorsum; distinct lateral indentations are lacking.

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

The oval prostomium and tentacular segment (I)

are partially fused (Figure 4a,b). The short, subulate,

median antenna is attached to an indistinct ridge on

the anterior third of the prostomium; similar lateral

antennae are attached to the inner dorsal sides of the

tentacular parapodia, connected by indistinct ceratophores to the prostomium. Eyes are lacking (slightly

dusky areas are present on the anterior borders of

prostomium, according to Mclntosh). Nuchal organs

were not observed. The palps extend to about setiger

18 (15-19). The inner palpal sheaths are large and

rounded, the outer ones, short. Medial to the inner

palpal sheaths is a bulbous facial tubercle. The tentacular cirri have distinct cirrophores; the dorsal tentacular cirri are much longer than the ventral ones;

stylodes are lacking; setae are few in number or lacking

(no setae on specimens examined; none, according to

Mclntosh; few setae indicated by Ehlers). The lateral

lips bear a pair of small bulbous labial lobes (Figure

4b). The notopodia of segments 2 and 3 have a circlet

of stylodes; the neuropodia have upper and lower

groups of few stylodes; the buccal cirri of segment 2

and the ventral cirri of segment 3 are short and subconical; the neurosetae are similar to those of the following segments; both dorsal cirri and dorsal tubercles

are lacking on segment 3 (Figure Aa-d).

Branchiae are present from about segment 30, with

smaller rudimentary ones more anteriorly. The parapodial ctenidia begin at about the same region as the

branchiae; they are rather inconspicuous, the 3 ctenidia per parapodium appearing nearly contiguous

(Figure 4t-/).

The notopodia and neuropodia are subequal in

length in the anterior region, but the latter surpass the

former more posteriorly (Figure Ae-m). The small

clavate notopodia have a few stylodes on subterminal

posterodorsal bracts and a large terminal stylode near

the tips of the acicular lobes. The notosetae are slender,

coarsely to finely spinous, and taper to capillary tips.

The large neuropodia have large posterior-upper and

small posterior-lower bracts, both equipped with stylodes; the latter are relatively few in number, large

and clavate: 1 or 2 stylodes on the posterior-upper

bracts and one or no stylodes on the posterior-lower

bracts. The acicular lobes are subconical with inconspicuous anteroventral bracts. The neurosetae are all

compound spinigers; the blades are canaliculate, tapering to fine tips; the neurosetae of the lower series are

more slender than the others. The ventral cirri are
NUMBER 5 3

m

FIGURE 4.—Leanira hystricis (BMNH 1921: 5: 1: 653): a, Anterior end, dorsal view, styles of

dorsal tentacular cirri missing; b, anterior end, ventral view; c, parapodium from segment 2,

posterior view; d, parapodium from segment 3, anterior view; notosetae and notopodial stylodes

all missing; e, parapodium from segment 4, posterior view; /, upper, middle, and lower neurosetae

from same; g, parapodium from anterior region, posterior view; setae mostly broken; h, same,

anterior view; i, parapodium from middle region, posterior view; ;, same, anterior view; k,

parapodium from middle fragment, posterior view; I, same, anterior view; m, upper, middle, and

lower neurosetae from same.
10 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

slender, tapered, and do not extend to the distal tips

of the neuropodia; small bulbous lobes are found on

the outer bases of the cirri.

DISTRIBUTION.—Northeastern Atlantic: south of

Iceland, off Great Britain, off Azores. In 957 to 2640

meters.

Leanira robusta Verrill

FIGURE 5

Leanira robusta Verrill, 1885a, p. 682, pi. 40: fig. 175;

1885b, p. 426.—Hartman, 1942, p. 104, fig. 8a; 1965a, p.

51.

Leanira hystricis.—Pettibone, 1963, p. 53 (part). [Not

Ehlers, 1874.]

MATERIAL EXAMINED.—Off Martha's Vineyard,

Massachusets, 220 meters, Fish Hawk station 876, 13

September 1880—2 syntypes (USNM 10320), 5 small

syntypes (USNM 10323).

DESCRIPTION.—The length is more than 100 mm,

the maximum width 10 mm, including setae, with segments more than 100. The elytra are thin, transparent,

and found on all segments from 27 posteriorly; they

are small and oval on the first few segments, become

progressively larger, and subrectangular to subcordiform in shape, with lateral indentations (Figure

5m,n).

The oval prostomium and tentacular segment (I)

are partially fused (Figure 5a-d). The short subulate

median antenna is attached to an indistinct ridge on

the anterior third of the prostomium; similar lateral

antennae are attached to the inner dorsal sides of the

tentacular parapodia, connected by indistinct ceratophores to the prostomium. Eyes and nuchal organs

were not observed. The palps extend to about segment

14. The inner palpal sheaths are large and subtriangular, the outer ones, short and rounded. Medial to

the inner palpal sheaths is a bulbous facial tubercle.

The tentacular cirri have distinct cirrophores; the dorsal tentacular cirri are about twice as long as the

ventral ones. There are usually 2 or 3 short stylodes

medial to the cirrophores of the dorsal tentacular cirri;

two bundles of capillary setae are directed anteriorly.

The lateral lips lack distinct labial lobes (Figure 5b).

The notopodia of segments 2 and 3 have a circlet of

stylodes; the neuropodia have upper and lower groups

of stylodes; the buccal cirri of segment 2 and the ventral cirri of segment 3 are short, thick and subconical;

the neurosetae are similar to those of the following

segments; both dorsal cirri and dorsal tubercles are

lacking on segment 3 (a minute papilla may or may

not be present; Figure 5e,f).

Branchiae are present from about segment 30, with

smaller rudimentary ones more anteriorly. The parapodial ctenidia begin at about the same region as the

branchiae; they are low and wide, the 3 ctenidia per

parapodium appearing nearly contiguous (Figure

5/Jfc).

The notopodia and neuropodia are subequal in

length (Figure 5g-k). The small clavate notopodia

have circlets of stylodes on subterminal posterodorsal

bracts and a larger terminal stylode near the tips of the

acicular lobes. The notosetae are slender, coarsely to

finely spinous, and taper to capillary tips. The large

neuropodia have large posterior-upper and small posterior-lower bracts, both equipped with stylodes; the

latter are filiform and numerous all along the body;

in the neuropodia of the middle region, the posteriorupper bracts are extended anteriorly such that a few

presetal stylodes are found (Figure 5k). The acicular

lobes are subconical with inconspicuous anteroventral

bracts. Most of the neurosetae are compound; the

blades are canaliculate, and taper to fine tips; those of

the lower series are more slender (Figure 5i). A few

slender, simple spinous neurosetae occur in the upper

parts of the neuropodia, beginning in the middle parapodia or more anteriorly (Figure 5/). The ventral cirri

are slender, tapered, and do not extend to the distal

tips of the neuropodia; small bulbous lobes are found

on the outer bases of the cirri.

DISTRIBUTION.—Northwestern Atlantic, off Massachusetts. In 183 to 230 meters.

Leanira alba Moore

FIGURE 6

Leanira alba Moore, 1910, p. 387, pi. 33: figs. 99-104.—

Hartman, 1960, p. 82; 1965a, p. 51.

MATERIAL EXAMINED.—Vicinity of San Diego, California, 1181 meters, Albatross station 4354, 14 March

1904—holotype (USNM 16883). Off southern California, 32°40' N., H7°3 r W, 1503 meters, green mud,

Albatross station 2923, 19 January 1889—1 specimen

(USNM 40998).

Southwest mouth Columbia River, Oregon, MS

Commando, M. S. Alton, collector; 45°56' N,

124°46.4' W, 596 meters, 26 May 1962—1 specimen

(USNM 40999). 45°58.7' N, 124°45.5' W, 457 meters,

9 May 1963—1 specimen (USNM 41000). 45°57.3' N,
NUMBER 5 3 11

OpoS

FIGURE 5.—Leanira robusta (syntype, USNM 10320): a, Anterior end, dorsal view; b, anterior

end, ventral view; c, prostomium and left tentacular parapodium, inner or medial view; d, right

tentacular parapodium, outer or lateral view; e, parapodium from segment 2, anterior view; /,

parapodium from segment 3, posterior view; g, parapodium from anterior region, posterior view;

h, same, anterior view; i, upper, middle,and lower neurosetae from same; ;, parapodium from

middle region, posterior view; k, same, anterior view; /, upper simple neuroseta from same; m,

right seventh elytron; n, right middle elytron.
12

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 6.—Leanira alba (USNM 41001): a, Anterior end, dorsal view; b, anterior end, ventral

view; c, tip of prostomium and right tentacular parapodium, outer or lateral view; d, left tentacular parapodium, inner or medial view; e, parapodium* from segment 2, anterior view; /,

parapodium from segment 3, posterior view; g, parapodium from anterior region, anterior view;

h, same, posterior view; i, middle and lower neurosetae from same; ;, parapodium from middle

region, anterior view; k, same, posterior view; /, upper simple neuroseta from same; m, middle and

lower neurosetae from same; n, right second elytron; o, right third elytron; p, right elytron from

anterior region; q, right elytron from middle region.
NUMBER 53

124°48.7' W, 596 meters, 10 May 1963—1 specimen

(USNM 30914). 45°58.6' N, 124°47' W, 553 meters,

23 May 1964—1 specimen (USNM 41001).

DESCRIPTION.—The length is more than 70 mm,

the maximum width 7.5 mm, including setae, with

segments more than 100. The elytra are soft, flexible,

smooth, and found on all segments from 27 posteriorly;

they are small and oval on the first few segments, become progressively larger and subreniform to subcordiform in shape, with lateral indentations (Figure

6n-q). None of the elytra examined show the bleblike

elevations along the lateral margins as noted by Moore

for some of them; evidently they were abnormal, as

he surmised.

The oval prostomium and tentacular segment (I)

are partially fused (Figure 6a-d). The short subulate

median antenna is attached to the short, free portion

of an indistinct ridge on the anterior third of the

prostomium; similar lateral antennae are attached to

the dorsal sides of the tentacular parapodia, connected

by indistinct ceratophores to the prostomium. Eyes

and nuchal organs were not observed. The palps extend to about segment 14. The inner palpal sheaths

are large and subtriangular, the outer ones, short and

rounded. Medial to the inner palpal sheaths is a bulbous facial tubercle or ridge. The tentacular cirri have

distinct cirrophores; the dorsal tentacular cirri are

about three times longer than the ventral ones. The

setae of the two bundles are relatively few and delicate,

directed anteriorly; stylodes are lacking. The lateral

lips lack distinct labial lobes (Figure 6b). The notopodia of segments 2 and 3 bear a circlet of stylodes;

the neuropodia have upper and lower groups of stylodes; the buccal cirri of segment 2 and the ventral

cirri of segment 3 are short, thick and subconical; the

neurosetae are similar to those of the following segments; both dorsal cirri and dorsal tubercles are lacking on segment 3 (Figure 6e,f).

Branchiae are present from about segment 30 posteriorly, with rudimentary ones more anteriorly. Fully

developed branchiae are large, digitiform, and inflated.

The parapodial ctenidia begin at about the same region as the branchiae; they are low and wide, the 3

ctenidia per parapodium being nearly contiguous

(Figure 6;^:).

The notopodia and neuropodia are subequal in

length in the anterior parapodia (Figure 6g,h); more

posteriorly, the neuropodia surpass the notopodia (Figure 6j,k). The small clavate notopodia have circlets

13

of stylodes on subterminal posterodorsal bracts and a

larger terminal stylode near the tips of the acicular

lobes. The notosetae are slender, coarsely to finely

spinous, and taper to capillary tips. The large neuropodia have large posterior-upper and small posteriorlower bracts, both equipped with stylodes; the latter

are filiform to digitiform and are numerous all along

the body; in the neuropodia of the middle region, the

posterior-upper bracts are extended anteriorly, such

that a few presetal stylodes are found (Figure 6;). The

acicular lobes are subconical, with inconspicuous anteroventral bracts. Most of the neurosetae are compound; the blades are canaliculate and taper rather

abruptly to fine tips; those of the lower series are more

slender (Figure 6i,m). There are as many as 5 slender,

simple, spinous neurosetae in the upper parts of the

neuropodia, first appearing in the middle parapodia

or more anteriorly (about setiger 25, according to

Moore; Figure 6/). The ventral cirri are slender,

tapered, and do not extend to the tips of the neuropodia; small bulbous lobes are found on the outer

bases of the cirri. Slender, tubular, segmental papillae,

situated anteromedial to the ventral cirri, are present

from about segment 28 (26-36) posteriorly (Figure

6;,*).

DISTRIBUTION.—Northeastern Pacific, from southern

California to Oregon. In 553 to 1503 meters.

REMARKS.—The specimen from Australia (QMB

G3972), identified by Rullier (1965) as L. alba, is a

member of the genus Sthenolepis, for on examination

the ceratophore of the median antenna was found to

have two large ctenidia or auricles.

Leanira coeca Horst

FIGURE 7

Leanira coeca Horst, 1917, p. 120, pi. 26: figs. 5-7 (part;

not Siboga station 211).—Hartman, 1965a, p. 51.

MATERIAL EXAMINED.—Malay Archipelago, Siboga

Expedition: station 5, 10 March 1899, 07°46' S, 114°-

30.5' E, 330 meters, mud—syntype (ZMA 524.5);

station 178, 2 September 1899, 02°40/ S, 128°37.5' E,

835 meters, blue mud—syntype (ZMA 524.4); station

271, 21 December 1899, 05°46.7' S, 134° E, 1788

meters, bluish green mud—14 syntypes (ZMA 524.2),

5 syntypes (RMNH 1170); station 314, 17 February

1900, 07°36' S, 117°3O.8' E, 694 meters, fine sandy

mud—3 syntypes (ZMA 524.1, RMNH 1169, USNM

40995).
14

DESCRIPTION.—The length is more than 42 mm, the

maximum width 6 mm, including setae, with more

than 75 segments. The elytra are soft, flexible, smooth,

and found on all segments from 27 posteriorly; they

are small and oval on the first few segments, become

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

progressively larger and subreniform to subpyriform in

shape, with lateral indentations (Figure 7a,l-n).

The oval prostomium and tentacular segment (I)

are partially fused (Figure 7a,b). The short, subulate,

median antenna is attached to the short free portion

FIGURE 7.—Leanira coeca (syntype, ZMA 178): a, Anterior end, dorsal view, right palp missing;

b, anterior end, ventral view; c, parapodium from segment 2, posterior view; d, parapodium from

segment 3, anterior view; e, upper, middle, and lower neurosetae from same; /, parapodium from

anterior region, posterior view; g, same, anterior view; h, neurosetae from same; i, parapodium

from middle region, posterior view; j , parapodium from posterior region, anterior view; k, upper,

middle, and lower neurosetae from same; /, left second elytron; m, left sixth elytron; n, left

middle elytron.
NUMBER 5 3 15

of an indistinct ridge on the anterior third of the prostomium; similar lateral antennae are attached to the

inner dorsal sides of the tentacular parapodia, connected by indistinct ceratophores to the prostomium.

Eyes are lacking. A pair of small rounded nuchal

organs, posterolateral to the prostomium, were observed when the pharynx was fully extended. The palps

extend to about segment 16. The inner palpal sheaths

are large and subtriangular, the outer ones, short and

rounded. Medial to the inner palpal sheaths is a bulbous facial tubercle or ridge. The tentacular cirri have

distinct cirrophores; the dorsal tentacular cirri are

about three times longer than the ventral ones. The

setae of the two bundles are relatively few, delicate, and

directed anteriorly; 1 to 3 stylodes are usually present

near the bases of the dorsal tentacular cirri. The lateral

lips lack distinct labial lobes (Figure 7 b). The notopodia of segments 2 and 3 have a circlet of stylodes;

the neuropodia have upper and lower groups of stylodes; the buccal cirri of segment 2 and the ventral

cirri of segment 3 are short, thick and subconical; the

neurosetae are similar to those of the following setigers; both dorsal cirri and dorsal tubercles are lacking

on segment 3 (Figure 7 c—e).

Branchiae are present from segment 30 posteriorly,

with smaller rudimentary ones more anteriorly. Fully

developed branchiae are large and digitiform. The

parapodial ctenidia begin at about the same region as

the branchiae; they are low and wide, the 3 ctenidia

per parapodium being nearly contiguous (Figure 7t).

The notopodia and neuropodia are subequal in

length in the anterior and middle regions (Figure

7f,g,i) ; more posteriorly the neuropodia surpass the

notopodia in length (Figure 7j). The small clavate

notopodia have circlets of stylodes on subterminal posterodorsal bracts and a larger terminal stylode near

the tips of the acicular lobes. The notosetae are slender, coarsely to finely spinous, and taper to capillary

tips. The large neuropodia have large posterior-upper

and small posterior-lower bracts, both equipped with

stylodes; the latter are filiform to digitiform and are

fairly numerous all along the body; in the neuropodia

of the posterior region, the posterior-upper bracts are

extended anteriorly, such that a few presetal stylodes

are present (Figure 7;'). The acicular lobes are subconical with inconspicuous anteroventral bracts. All

the neurosetae are compound spinigers; the blades arc

canaliculate and taper to short fine tips; those of the

lower series are more slender (Figure 7h,k). The ventral cirri are slender, tapered, and do not extend to the

tips of the neuropodia; small bulbous lobes are found

on the outer bases of the cirri. Slender tubular segmental papillae, anteromedial to the ventral cirri, are

found from about segment 30 posteriorly (Figure 7;).

DISTRIBUTION.—Malay Archipelago. In 330 to 1788

meters.

REMARKS.—The syntype from Siboga station 211

(ZMA 524.3) differs from the other syntypes and does

not agree with the description or figures of L. coeca

by Horst. There are short, conical, dorsal tubercles on

segment 3; the median antenna has a large bulbous

ceratophore, with the style broken. The specimen is in

such a poor condition that its specific identity cannot

be determined.

Leanira cirrata (Treadwell)

FIGURE 8

Eupholoe cirrata Treadwell, 1934, p. 5, pi. 1: figs. 9-12, pi.

2: figs. 13-16; 1939, p. 198, fig. 28.

Leanira hystricis.—Hartman, 1942, p. 105. [Not Ehlers,

1874.]

Leanira cirrata.—Hartman, 1965a, p. 51.

MATERIAL EXAMINED.—North of Puerto Rico,

Johnson-Smithsonian Deep-sea Expedition: station

14, 2 February 1933, 18°3O/ N, 66°03' W, 366-439

meters—holotype (USNM 20033); station 23, 4 February 1933, 18°32' N, 66°21' W, 475 meters—2 paratypes (USNM 20068).

DESCRIPTION.—The length is more than 90 mm, the

maximum width 7 mm, including setae, with more

than 110 segments. The elytra are delicate, transparent, with small, opaque areas lateral to their

places of attachment to the elytriphores (referred to as

brown spots by Treadwell). The elytra are located on

all segments from 27 posteriorly; they are small and

oval on the first few segments, become progressively

larger, and subpyriform to subreniform in shape, with

slight lateral indentations (Figure 8m-p).

The oval prostomium and tentacular segment (I)

are partially fused (Figure 8a,b). The short subulate

median antenna is attached to the short free portion

of an indistinct ridge on the anterior third of the prostomium; slightly more slender, lateral antennae are

attached medially on the tentacular parapodia (overlooked by Treadwell, 1934; observed by Hartman,

1942). Eyes and nuchal organs were not observed. The

palps extend to about segment 15. The inner palpal
16

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

OpaS

FIGURE 8.—Leanira cirrata (holotype of Eupholoe cirrata, USNM 20033): a, Prostomium and

left tentacular parapodium, outer or lateral view; b, same, inner or medial view; c, parapodium

from segment 2, anterior view; d, neurosetae from same; e, parapodium from segment 3, anterior

view; /, parapodium from anterior region, posterior view; g, same, anterior view; h, neurosetae

from same; i, diagrammatic end view of parapodium from anterior region showing arrangement of

setae; ;, parapodium from middle region, anterior view; k, upper simple neuroseta from same; /,

neuroseta from same; m, left second elytron; n, left fourth elytron; o, left tenth elytron; p, left

middle elytron.
NUMBER 5 3 17

sheaths are large and subtriangular, the outer ones,

low. Medial to the inner palpal sheaths is a bulbous

facial tubercle. The tentacular cirri (overlooked by

Treadwell) have distinct cirrophores; the dorsal tentacular cirri are about three times longer than the ventral ones. The setae of the two bundles are relatively

few, delicate, and directed anteriorly; the tip of the

single aciculum is reddish; numerous filiform stylodes

are medial to the cirrophores of the dorsal tentacular

cirri. The lateral lips lack distinct labial lobes. The

notopodia of segments 2 and 3 have a circlet of stylodes; the neuropodia have upper and lower groups of

stylodes; the buccal cirri of segment 2 and the ventral

cirri of segment 3 are short, thick and subconical; the

compound neurosetae are similar to those of the following segments; in some of the neurosetae the articulations appear rather indistinct; both dorsal cirri and

dorsal tubercles are lacking on segment 3 (Figure

8c-e).

Branchiae are present from segment 30, with smaller

rudimentary ones more anteriorly. Fully developed

branchiae are large and digitiform. The parapodial

ctenidia begin at about the same region as the

branchiae; they are low and wide, the 3 ctenidia per

parapodium being nearly contiguous (Figure 8;).

The notopodia and neuropodia are subequal in

length (Figure 8f,g,j). The small clavate notopodia

have circlets of stylodes on subterminal posterodorsal

bracts and 1 to 3 larger terminal stylodes near the

tips of the acicular lobes. The notosetae are slender,

coarsely to finely spinous, and taper to capillary tips.

The large neuropodia have large posterior-upper and

small posterior-lower bracts, both equipped with numerous stylodes; the latter are filiform and numerous

all along the body; on the neuropodia of the middle

region, the posterior-upper bracts are extended anteriorly, such that some presetal stylodes are found (Figure 8;'). The acicular lobes are subconical with inconspicuous anteroventral bracts. Most of the neurosetae

are compound; the blades are canaliculate and taper

rather abruptly to fine tips; the shafts are reddish

amber-colored; the neurosetae of the lower series are

more slender than the others (Figure 8h,l). There are

a few slender, simple spinous neurosetae in the upper

parts of the neuropodia, beginning about segment 30

(Figure 8k; overlooked by Treadwell and Hartman).

The ventral cirri are slender, tapered, and do not extend to the distal tips of the neuropodia; small bulbous

lobes are present on the outer bases of the cirri. Slender

tubular segmental papillae, anteromedial to the ventral cirri, are found in more posterior segments

(Figure 8;).

DISTRIBUTION.—Northwestern Atlantic, north of

Puerto Rico. In 366 to 475 meters.

REMARKS.—Hartman (1942: 105) examined the

holotype of Eupholoe cirrata and referred the species

to Leanira hystricis. Later, Hartman (1965a: 51) recognized L. cirrata as a distinct species.

Leanira adenensis, new species

FIGURE 9

Leanira vulturis.—Monro, 1937, p. 263. [Not Horst, 1917.]

TYPE MATERIAL.—Gulf of Aden, 274 meters, the

John Murray Expedition 1933-34, station 191-holotype and 2 paratypes. [The holotype (BMNH 1937:

9: 2: 61) consists of an anterior and two middle fragments, with a total length of 65 mm, width of 4 mm,

including setae, and more than 117 segments. A

smaller paratype (USNM 40997) consists of an anterior fragment of 28 mm, width of 3 mm, including

setae, and 53 segments. The smallest paratype (BMNH

1937: 9: 2: 62) consists of an anterior fragment of

10 mm, width of 2 mm, including setae, and 33

segments.]

DESCRIPTION.—The elytra are delicate, transparent,

smooth, and found on all segments from 27 posteriorly; they are small and oval on the first few segments,

become progressively larger and subrectangular to

subcordiform in shape, with lateral indentations (Figure 9k-m).

The oval prostomium and tentacular segment (I)

are partially fused (Figure 9a,b). The ceratophore of

the median antenna is attached to the anterior fourth

of the prostomium; the style is short, subulate, 2-

jointed; similar lateral antennae are attached to the

dorsal sides of the tentacular parapodia, connected by

indistinct ceratophores to the prostomium. Two pairs

of eyes are faintly visible on the holotype: a large pair

on the anterior border and a minute pair in the middle

of the prostomium; a pair of semilunar nuchal organs,

posterolateral to the prostomium, are visible on the

holotype (the pharynx is partially extended, causing the

anterior parapodia to be spread apart). The palps

extend to about segment 15. The inner palpal sheaths

are large and subtriangular, the outer ones, rounded

and about half as long. Medial to the inner palpal

sheaths is a bulbous facial tubercle. The tentacular
18

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

dTc

FIGURE 9.—Leanira adenensis, new species (a, c-m, holotype, BMNH 1937: 9: 2: 61-62; b,

paratype, USNM 40997): a, Anterior end, dorsal view, pharynx partially extended; b, same,

ventral view; c, parapodium from segment 2, posterior view; d, parapodium from segment 3,

anterior view; e, parapodium from anterior region, posterior view; /, same, anterior view; g,

upper simple neuroseta from same; h, middle and lower compound neurosetae from same; i,

parapodium from middle region, posterior view; ;, same, anterior view; k, right first elytron;

/, right elytron from anterior region; m, right elytron from middle region.
NUMBER 5 3

cirri have distinct cirrophores; the dorsal tentacular

cirri are about three times longer than the ventral ones.

The setae of the two bundles are relatively few and

delicate, directed anteriorly; sty lodes are usually lacking (single one on right tentacular parapodium of

holotype). The lateral lips have small globular labial

lobes (Figure 9b). The notopodia of segments 2 and 3

have a few stylodes; the neuropodia have upper and

lower groups of stylodes; the buccal cirri of segment 2

and the ventral cirri of segment 3 are short, thick and

subconical; the neurosetae are similar to those of the

following segments except that the blades are somewhat longer; both dorsal cirri and dorsal tubercles are

lacking on segment 3 (Figure 9c,d).

Branchiae are present from about segment 25 posteriorly, with smaller rudimentary ones more anteriorly. Fully developed branchiae are large and digitiform.

The parapodial ctenidia begin at about the same region

as the branchiae; they are low and wide, the 3 ctenidia

per parapodium being nearly contiguous (Figure

The notopodia and neuropodia are subequal in

length (Figure 9e,f,i,j). The small clavate notopodia

have circlets of stylodes on subterminal posterodorsal

bracts and a larger terminal stylode near the tips of

the acicular lobes. The notosetae are slender, coarsely

to finely spinous, and taper to capillary tips. The large

neuropodia have large posterior-upper and small posterior-lower bracts, both equipped with stylodes; the

latter are digitiform and fairly numerous all along the

body; on the neuropodia of the middle region, the posterior-upper bracts are extended anteriorly, such that

a few presetal stylodes are found (Figure 9j). The

acicular lobes are subconical, with inconspicuous

anteroventral bracts. Most of the neurosetae are compound spinigers; the blades are canaliculate and taper

to fine tips; those of the lower series are more slender

(Figure 9h). There are a few (2-5) slender, simple

spinous neurosetae in the upper parts of the neuropodia, beginning in the anterior region (about segment 16; Figure 9g). The ventral cirri are slender,

tapered, and do not extend to the tips of the neuropodia; small bulbous lobes are found on the outer

bases of the cirri. Slender tubular segmental papillae,

anteromedial to the ventral cirri, are present from

about segment 23 posteriorly (Figure 9/).

DISTRIBUTION.—Gulf of Aden. In 274 meters.

19

Ehlersileanira, new genus

TYPE-SPECIES: Sthenelais incisa Grube, 1877. Gender: feminine.

DIAGNOSIS.—Body elongate, with numerous segments. Prostomium oval, partially fused to tentacular

parapodia. Three antennae; ceratophore of median

antenna with lateral auricles or ctenidia; style short;

lateral antennae short, fused to tentacular parapodia.

Long palps emerging ventral to tentacular parapodia

between large inner and smaller outer palpal sheaths.

Bulbous facial tubercle medial to inner palpal sheaths.

Tentacular parapodia (I) directed anteriorly, with

single aciculum, pair of tentacular cirri, and 2 bundles

of capillary setae. Elytra on segments 2, 4, 5, 7, on

alternate segments to 27, and on all succeeding segments. Elytra smooth, lacking tubercles and papillae.

Dorsal cirri and dorsal tubercles lacking on segment 3.

Cirriform branchiae and elongate platelike ctenidia, 3

per parapodium, from about segment 25-30 posteriorly. Notopodia with well-developed bracts with

stylodes. Neuropodia possessing bilobed posterior bracts

bearing stylodes. Notosetae spinous capillaries. Neurosetae consisting of compound spinigers, with blades

relatively short and canaliculate and additional simple

ones. Ventral cirri short and tapered. Pharynx with 11

pairs of papillae and 2 pairs of jaws.

ETYMOLOGY.—The genus is named for the late

Professor Ehlers, whose figures of Sthenelais simplex are

the most detailed and accurate of all those existing for

the species referred to the genus.

REMARKS.—Sthenelais luxuriosa Grube (1875: 78;

1878: 54), from the Philippine Islands, should perhaps be referred to Ehlersileanira. No figures accompanied the original descriptions and type-specimens are

no longer extant (G. Hartwich, in litteris). Because of

the inadequate description, this combination must be

considered a species inquirendum.

Ehlersileanira incisa (Grube), new combination

FIGURES 10-12

Sthenelais incisa Grube, 1877, p. 519.

Sthenelais simplex Ehlers, 1887, p. 60, pi. 13: figs. 2, 3, pi. 14:

figs. 1-5 (part; not pi. 14: fig. 6).—Treadwell, 1901, p.

187; 1939, p. 193, fig. 23. [Lectotype (MCZ 722) here

designated.]

Leanira vulturis Horst, 1917, p. 118, pi. 25: figs. 5-7 (part of

Siboga station 47).

Leanira incisa.—Augener, 1918, p. 107.—Monro, 1930, p.

70.—Hartman, 1944, p. 13, pi. 1: figs. 1-4, pi. 2: figs.
20

9-11; 1959, p. 120.—Fauvel, 1953, p. 9.—Rullier, 1964,

p. 1081.

Sthenelais luxuriosa.—Tre&dvrttt, 1920, p. 592 (part; not

Albatross station D5257). [?Not Grube, 1875.]

Leanira izuensis Takahashi, 1938, p. 199, pi. 20: fig. C, textfigs. IV, V.—Hartman, 1959, p. 116.

Sthenolepis izuensis.—Imajima and Hartman, 1964, p. 43.—

Hartman, 1965a, p. 14.

Sthenolepis vulturis.—Hartman, 1965a, p. 14.

Sthenolepis incisa.—Hartman, 1965a, p. 14.

MATERIAL EXAMINED.—West Africa, mouth of

Congo, Gazelle Expedition—holotype of Sthenelais

incisa (ZMB 912). Off St. Paul de Loanda, Angola,

64-65 meters, grey mud, Discovery station 274, 4 August 1927—1 specimen (BMNH 1930: 10: 8: 1175).

Off Cape Lopez, French Congo, 58-67 meters, mud

and fine sand, Discovery station 279—1 specimen

(BMNH 1930: 10: 8: 1172).

Northwest Atlantic, 28°42' N, 88°40' W, 587

meters, Blake station 47, 1877-78—lectotype of

Sthenelais simplex (MCZ 722). South of Tortugas,

Florida, W. L. Schmitt, collector: 201 meters, 15 July

1930—5 specimens (USNM 41062); 152-165 meters,

23 July 1932—1 specimen (USNM 41058); 110

meters, 5 August 1932—1 specimen (USNM 41060).

Gulf of Mexico, 29°02' N, 88°40/ W, 309 meters,

Pelican station 9, 4 February 1938—1 specimen

(USNM 41057). 28°43' N, 91°49' W, 29 meters,

Pelican station 85-4, 12 July 1938—1 specimen

(USNM 41061).

Caribbean, Point del Algarrabo, Puerto Rico, 177—

220 meters, Fish Hawk station 138 (6066), 20 January 1899—2 specimens (USNM 15684). 18°32' N,

66°21' W, 475 meters, Johnson-Smithsonian Deep-sea

Expedition, station 23, 4 February 1933—1 specimen

(USNM 20051). 18°3rN,66°12'W,256-366meters,

station 21, 4 February 1933—1 specimen (USNM

20053). 18°37' N, 65°05' W, 640 meters, station 93,

2 March 1933—1 specimen (USNM 20052).

Southwest Atlantic, east coast South America off

Brazil, 23°O8' S, 41°34' W, 108 meters, blue mud,

Albatross station 2762, 30 December 1887—1 specimen

(USNM 41059).

Malay Archipelago, Siboga Expedition: Madura

Strait, 07°27.5' S, 113°08.5' E, 37 meters, station 1,

7 March 1899—syntype of Leanira vulturis (ZMA

529.1). Madura Strait, 07°25' S, 113° 16' E, 56 meters,

station 2, 8 March 1899—2 syntypes (ZMA 529.2).

Bay of Bima, 55 meters, station 47, 8/12 April 1899—

syntype (ZMA 529.3); 2 syntypes (RMNH 1164).

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Strait of Makassar, 03°30' S, 116°33' E, 30.6 meters,

M. P. N. van Kampen, collector—syntype (RMNH

1163).

Vicinity Philippine Islands, Albatross stations:

Lusaran Light, between Panay and Negros, 10°32' N,

122° 26' E, 176 meters, soft green mud, station D5183,

30 March 1908—1 specimen (USNM 17503). Tayabas Light, 13°42'N, 121 °50' E, 196 meters, green mud,

2 March 1909, station D5375—1 specimen (USNM

17516). Tubig Point, Destacado Island, 247 meters,

green mud and sand, station D5392, 1 March 1909—1

specimen (USNM 17513). Panalangan Point, Talajit

Island, 11°57' N, 124° 10' E, 245 meters, gray mud,

station D5397, 15 March 1909—2 specimens (USNM

17517). Cagayan Island, Jolo Sea, 09°38' N, 121°11'

E, 929 meters, gray mud, coral sand, station D5423,

31 March 1909—1 specimen (USNM 17514).

REMARKS.—The syntypic series of three specimens

of Sthenelais simplex Ehlers (MCZ 722) contains three

species:

(1) Anterior fragment of about 100 segments, with

the pharynx withdrawn. It agrees for the most part

with the figures by Ehlers (1887), except for plate 14:

figure 6. This specimen is herein selected as the

lectotype.

(2) Middle fragment of a different sigalionid. Considered indeterminable.

(3) Anterior fragment, with pharynx extended.

The jaws correspond to plate 14: figure 6, in Ehlers

(1887) and are of the polyodontid type. The parapodia

have spinning glands and the setae are similar to those

found in Eupanthalis. Obviously, this specimen is a

member of the family Polyodontidae.

It is doubtful that the specimen recorded by Augener

(1906: 106) as Leanira simplex, from Blake station

344 (MCZ 2238), was correctly identified. The figure

of the prostomium by Augener (plate 1: figure 16)

shows no lateral auricles on the ceratophore of the

median antenna, such as were illustrated by Ehlers

(1887, plate 13: figure 2) for S. simplex. The specimen, which was examined in this study, is in such poor

condition that it cannot be identified. Sthenelais simplex Ehlers, 1887, was declared a synonym of L. incisa

(Grube, 1877) by Augener (1918: 107).

Of the four syntypes of Leanira vulturis from Siboga

station 47, one proved to be Pareulepis malayana

(Horst) (see Pettibone, 1969: 12). The specimens

from the Gulf of Aden, referred to L. vulturis by

Monro (1937: 263) were examined and are referred

herein to L. adenensis, new species.
NUMBER 5 3 21

DESCRIPTION.—The length is more than 330 mm,

the maximum width 12 mm, including setae, with segments more than 200. The elytra are rather thick,

opaque, and found on all segments from 27 posteriorly;

they are small and oval on the first few segments,

become progressively larger and subpyriform in shape,

with anterior indentations and lateral notches; more

opaque spots are found lateral to their places of attachment (Figure llg-i).

The oval prostomium and tentacular segment (I)

are partially fused (Figures lOa-d; \2a-d). The

ceratophore of the median antenna, located on the

anterior fourth to third of the prostomium, has a pair

of lateral auricles, which are joined medially; the style

is short, subulate, and 2-articled. Similar lateral antennae are attached to the dorsal sides of the tentacular parapodia, connected by indistinct ceratophores to

the prostomium. Eyes may be lacking (Figure 10a)

or four eyes present—anterior larger pair and posterior

pair lateral to the ceratophores of the median antenna

(Figure \2a,c). Small semicircular nuchal organs may

or may not be visible. The palps extend to about segment 16 (10-23). The inner palpal sheaths are large

and subtriangular, the outer ones, shorter and

rounded. Medial to the inner palpal sheaths is a bulbous facial tubercle. The tentacular cirri have distinct

buC

FIGURE 10.—Ehlersileanira incisa (holotype of Sthenelais incisa, ZMB 912): a, Anterior end,

dorsal view; right parapodia of segments 2 and 3 not shown; b, same, ventral view; only bases of

dorsal tentacular cirri and palps shown; c, prostomium and right tentacular parapodium (I),

outer view; d, left tentacular parapodium, inner view; e, parapodium from segment 2, anterior

view; /, middle and lower neurosetae from same; g, parapodium from segment 3, posterior view.
22 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 11.—Ehlersileanim incisa (holotype of Sthenelais incisa, ZMB 912): a, Parapodium

from anterior region, posterior view; b, same, anterior view; c, diagrammatic end view of same,

showing arrangement of setae and bracts; d, simple neuroseta from same; e, upper and lower

neurosetae from same; /, middle parapodium, posterior view; g, right second elytron; h, left

fourth elytron; i, left middle elytron.

cirrophores; the dorsal tentacular cirri are much

longer than the ventral opes. Few (1-5) short stylodes

are usually present medial to the cirrophores of the

dorsal tentacular cirri. Two bundles of relatively few

capillary setae are directed anteriorly. The lateral lips

lack auricular labial lobes (Figures 106; 126). The

notopodia of segments 2 and 3 have a circlet of stylodes; the neuropodia have upper and lower groups

of stylodes; the buccal cirri of segment 2 and the ventral cirri of segment 3 are short, thick and subconical;

the neurosetae are similar to those of the following

segments, except that the articulations of the compound spinigers are rather indistinct; both dorsal cirri

and dorsal tubercles are lacking on segment 3 (Figures

l0a,e-g;l2a,e-g).

Branchiae are present from about segment 30 posteriorly, with smaller rudimentary ones more anteriorly. The parapodial ctenidia begin at about the same

region as the branchiae: they are low and elongated,

the 3 ctenidia per parapodium being nearly contiguous

(Figures 11/; 12*,/).

The small clavate notopodia extend beyond the

large neuropodia (Figures lla-/; \2h-l). The notopodia have circlets of stylodes on subterminal posterodorsal bracts and a larger terminal stylode near

the tips of the acicular lobes. The notosetae are slender, coarsely to finely spinous, and taper to capillary

tips. The large neuropodia have large posterior-upper

and small posterior-lower bracts, both equipped with

stylodes. The acicular lobes are subconical with low

anteroventral bracts. The neurosetae are mostly compound spinigers; the blades are rather long, canaliculate, and taper to fine, flexible tips; the articulations

of the blades to the stems are rather indistinct (Figures We; \2g,j). The neurosetae are arranged in 3

series (Figure We): curved vertical rows anterodorsal

to the posterior-upper bracts; subacicular curved rows
NUMBER 53 23

FIGURE 12.—Ehlersileanira incisa (syntype of Leanira vulturis, ZMA 529.2): a, Anterior end,

dorsal view; b, same, ventral view; c, prostomium and right tentacular parapodium (I), outer

view; d, right tentacular parapodium, inner view; e, parapodium from segment 2, anterior view;

f, parapodium from segment 3, posterior view; g, neuroseta from same; h, parapodium from

anterior region, posterior view; i, same, anterior view; ;, middle and lower neurosetae from same;

k, parapodium from middle region, posterior view; I, same, anterior view.

dorsal to the posterior-lower bracts; and ventral

arched rows within the anteroventral bracts. The

neurosetae of the latter series are more slender than

the others. A few simple spinous neurosetae occur in

the upper posterior parts of the neuropodia (Figure

1 Id); they may be lacking in the more anterior neuropodia. The ventral cirri are slender, tapered, and do not

extend to the distal tips of the neuropodia; small bulbous lobes are found on the outer bases of the cirri.

Tubular segmental papillae, anteromedial to the ventral cirri, begin about segment 26 or more anteriorly

(Figures 11/; 12/).

DISTRIBUTION.—North and South Atlantic: off

West Africa, South and Central America, Gulf of Mexico, Florida, West Indies; Malay Archipelago; Philippine Islands; Izu peninsula, Japan. In 15 to930 meters.
24 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

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ICE: 1970 O—375-441





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should not contain rules. Formal tables must be submitted as pages separate from the text, and

each table, no matter how large, should be pasted up as a single sheet of copy.

For measurements and weights, use the metric system instead of (or in addition to) the

English system.

Illustrations (line drawings, maps, photographs, shaded drawings) can be intermixed

throughout the printed text. They will be termed Figures and should be numbered consecutively;

however, if a group of figures is treated as a single figure, the individual components should be

indicated by lowercase italic letters on the illustration, in the legend, and in text references:

"Figure 96." If illustrations (usually tone photographs) are printed separately from the text as

full pages on a different stock of paper, they will be termed Plates, and individual components

should be lettered (Plate 9b) but may be numbered (Plate 9: figure 2). Never combine the

numbering system of text illustrations with that of plate illustrations. Submit all legends on pages

separate from the text and not attached to the artwork.

In the bibliography (usually called "Literature Cited"), spell out book, journal, and article

titles, using initial caps with all words except minor terms such as "and, of, the." (For capitalization of titles in foreign languages, follow the national practice of each language.) Underscore

(for italics) book and journal titles. Use the colon-parentheses system for volume, number, and

page citations: "10(2) :5-9." Spell out such words as "figures" and "plates" (or "pages" when

used alone).

For free copies of his own paper, a Smithsonian author should indicate his requirements on

"Form 36" (submitted to the Press with the manuscript). A non-Smithsonian author will receive

50 free copies; order forms for quantities above this amount with instructions for payment will be

supplied when page proof is forwarded.