DEEP-SEA AMPHIPODA

FROM THE WESTERN NORTH

ATLANTIC
OCEAN. THE FAMILY AMPELISCIDAE1t2
Eric L. Mills"
Institute

of Oceanography,

Dalhousie

University,

IIalifax,

Nova

Scotia

ABSTRACT

Eleven species of Ampeliscidae

(Amphipoda)
occur on a deep-sea benthic transect
between Massachusetts and Bermuda,
Two species of Byblis and one of Byblisoides
are
newly described; Ampelisca uncinata Chevreux, Byb1i.s gaimardi (Kr@yer), B. sewata Smith,
Haploops setosa Boeck, and H. &n&s Stephensen are redescribed
or refigured;
and new
distributional
or ecological
information
is given for Ampelkca
aequicornis
Bruzelius,
A.
There is a rough taxonomic
and ecological
agassixi (Judd),
and A. declivitatis
Mills.
zonation of Ampcliscidac,
with species of Ampelisca in shallow water, blind species of
Byblis on the continental
slope, species of Haploops on the slope extending into the abyss,
and an endemic species of Byblisoides at fully abyssal depths which is adapted for feeding
on largo quantities of abyssal sediments by having strongly-modified
second antennae and
species of Byblis and Haploops
also appear to bc
massive antenna1 muscles. Bathyal
sediment or suspension feeders but are less strongly modified.
Zoogeographic
affinities
of the species arc varied, especially on the continental
slope.
Shallow-water
species are Virginian
and Carolinian,
outer shelf species may range to north
or south (but not both), and the Byblisoides
is endemic to the North American Basin.
The bathyal species are widespread around the North Atlantic or closely related to species
from the northeast Atlantic.

INTRODUCTION

Ampeliscid
amphipod
crustaceans are
characteristic inhabitants of shallow marine waters all over the world and are
most common on fine sand and mud scdi1nmts.
In general, they are tube dwellers
which collect suspended and settling particulatc matter or rake the bottom with
their antennae for organic-rich
material
from the sediment surface (for detailed
accounts of ecology and feeding see Eacquist 1950 and Mills 1967a).
l Contribution
No. 2087 from Woods
Hole
Oceanographic
Institution.
2 This research was supported by grants from
the National
Research Council
of Canada and
the Faculty
of Graduate
Studies,
Dalhousie
University.
3 For information
and for the loan of collections,
I am indebted to J. L. Barnard
(U.S. National
Museum ), Ebbe Kanneworff
( Marine Biological
Laboratory,
Helsinggr,
Denmark),
A. D. Michael
(Dalhousie
University
and Systematics-Ecology
Program, Woods Hole, Mass. ), Elizabeth
Schroeder
(Woods
Hole
Oceanographic
Institution,
Woods Hole, Mass. ), Wayne K. Storey (Department of Zoology, University
of Maine, Orono ),
and Torben Wolff
(Universitets
Zoologiske
Museum, Copcnhagcn ) .
LIMNOLOGY

AND

OCEANOGRAPIIY

The animals discussed here have been

collcctcd from benthic stations along a
transect between Gay Head, Massachusetts, and Bermuda. The nature of the
transect, its general oceanography, sedimcnts, and faunal diversity have been reported elsewhere (Sanders et al. 1965;
Hessler and Sanders 1967; Sanders and
is
Hcssler 1969). Thus far information
available for some Amphipoda (Mills 1966,
1967b), Bryozoa ( Schopf 1965)) Isopoda
( Hcssler 1967)) Ophiuroidea
( Schoener
1967, 1968, 1969)) Polychac,ta ( Hartman
1965)) Priapulida
( Sanders and Hcssler
1962)) and Tunicata( Monniot and Monniot 1968). Studies on the transect have
shown that bathyal and abyssal animals
are both more abundant and more diverse
than has been suspected previously for the
deep sea. Because ,of its location close to
shallow-water
zoogeographic
boundaries,
the Gay Head-Bermuda
transect is wellplaced for studies relating deep-sea faunas
to shallow-water
faunas on the adjacent
shelf and to faunas of high latitudes in the
North Atlantic.
I will discuss here 11
spccics of Ampcliscidae
from the Gay
Head-Bermuda
transect and, briefly, their
357

MARC11

1971,

V.

16(2)

358

ERIC

L.

relation to amphipods from adjacent shallow waters and other deep-sea areas.
In the text following,
WIIB refers to
benthic stations of Woods Hole Occanographic Institution.
The thoracic limbs of
amphipods (seven in all) arc designated
gnathopods 1 and 2, followed by percopods 1 to 5, in accordance with tcrminology used by Barnard ( 1969).
This monograph is dedicated to Profcssor G. E. IIutchinson, who encouraged my
interest in systematics and who has provided mc and many others with the most
exciting and exacting intellectual standards
and challenges conceivable.
SYSTEMATIC

Ampclisca

acquioornis

SECTION

Bruxelius

(Fig. 1A)

Ampeliscn
nequicowzk
Bruzelius.
G. 0. Sars
1895, p. 177-178, Pl. 82.
Loculity
record:
WIIB
slope sta. No. 3; 39
58.4 N, 70 40.3 W; 300 m. Anchor dredge.
Atlantis cruise 283; 28 August 1962.
Range:
On the continental
shelf and slope
(depth records 92-835 m) aro,uncl the North Atlantic
from Britain
and Scandinavia
to Nova
Scotia and the Middle Atlantic States.

Diagnosis: A sturdy offshore Amp&sea

with cyes an d cornea1 lenses; ant&ma 1
almost as long as antenna 2; the front
ischium,
margin of the head obliguc;
merus, and carpus of the last perelopod
unlobed and more or less quadratc; ischium of last pcreopod longer than merus;
and only a slight carina of the urosome.
Remarks: This species has been recorded previously from off northwest Europe at 92-366 m (Sars 1895)) off Iceland
and southwest #of Greenland at 169-835 m
( Stephmscn 1925)) and from cast of Nova
Scotia at 212 m (Shocmakcr 1931). The
locality rccordcd here, on the continental
slope cast of New Jersey, is the southelllmost in the western North Atlantic.
Ampelisca

agassizi (Jutlcl)

(Fig. 1B)

Ampelisca
agussixi (Judd).
Mills 1967c (for
complete synonymy).
1) Sta. S3, Sanely Cove,
Local@
racorcls:
Digby
Co., Nova Scotia; 9 July 1963. E. L.
Bousfield,
collect.
2) Shcepscott River cstllary,
Maine; summer 1955. R. W. Hanks, collect.
3)

MILLS

Various collections off Gay Head, Marthas Vineyard and off Cuttyhunk,
Massachusetts;
30-40 m.
RV A. E. VerriU; July 1967. 4) WHB sta. 170;
40 37 N, 70 50 W; 67.7 m. Epibenthic
sled.
Atlantis
II cruise 40; 2rl November
1967. 5 )
WHB sta. 171; 40 34 N, 70 51 W; 67.7-71.4
m. Anchor dredge. Atlantis ZZ cruise 40; 27 November 1967. 6) WHB sta. 172; 40 12,.3 N,
70 44.7 W to 40 10.8 N, 70 43.6 W; 119 m.
Atlantis
II cruise 40; 27 November
1967. 7)
WIIB sta. 89; 40 01.6 N, 70 40.7 W; 196 m.
Chnin cruise 50; G July 1965. 8) WHB slope sta.
No. 2; 40 01.8 N, 70 42 W; 200 m. Atlantis
cruise 283; 28 August 1962. 9) WHB sta. No. 3;
39 68.4 N, 70 40.3 W; 300 m. Atlantis cruise
283; 28 August 1962. 10)Various
stations off
Cape Lookout, North Carolina; 445-450 m. J. F.
Grasslc, collect.
Ranae:
\, Shallow inshore waters to 450 m in
muddy sediments from western Nova Scotia to
the Caribbean
Sea and in the eastern Pacific
Ocean from at least the Queen Charlotte Islands,
British Columbia, to Ecuaclor.

Diagnosis: A modcratc-sized (6-10 mm)

compressed Ampelkca
with
cx tremely
long second antennae, a short, broad basis
of ihe last percopod, and a posterodistally
lobed mcrus of the last nereoDod.
Remarks: Synonymy, Lange: and notes
on ecology of this species wcrc given elscwhcrc ( Mills 1967c). A few notes on the
ecology of A. agassixi in Block Island
Sound, Rhode Island, where the species
lives in mud containing some fine gravel,
were given by Smith (1950). Breeding
occurs from June to October; 40% of the
standing crop is consumed annually by two
main predators, winter flounder ( Pseudopleurohectes
americanus)
and whiting
( Me&c&s
bilinearis), although it is not
an important item of diet for either species. Because of its enormous range-both
geographic and bathymetric-these
obscrvations give only limited insight into the
suite of biological relationships of populations of A. agassixi.
Ampclisca

declivitatis

Mills

(Fig:. 1 C)

Ampeliscn
de&it&is
Mills 1967c (for complete synonymy).
Locality
recorcls: 1) WHB sta. C No. 1; 40
20.5 N, 70 47 W; 97 m. Anchor clreclge. Aista. 89;
lardis cruise; 25 May 1961. 2) WIIB
40 01.6 N, 70 40.7 W; 196 m. Epibenthic
sled. Chuin cruise 50; G July 1965. 3) WIIB
slope sta. No. 4; 39 56.5 N, 70 39.9 W; 400

NORTH

ATLANTIC

DVJ!Q?-SEA

AMPPIIIPOlX

359

7 mm 0.
FIG. 1. A. Ampalisca aecpicornis
Brnzeli~~s, 10 mm Q . B. Ampelisca aguxsixi (Judd),
rlec1i~~itnti.s Mills, 5 mm 9, The bar besitlc cnch species rcprcscnts 2 mm of length.
C. Ampehkx

360

ERIC L. MILLS

m. Anchor dredge.
AtZantis cruise; 30 August
1962. 4) WHB sta. D No. 1; 39 54.5 N, 70
35 W; 466.7-508.7
m. Anchor dredge, Atlantis
cruise 277; 23 May 1962. 5) WHB sta. 96; 39
55.2 N, 70 39.5 W; 498 m. Epibenthic
sled.
Chain cruise 58; 27 April 1966. 6) WHB sta.
105B; 39 56.5 N, 71 03.6 W; 530 m. Epibenthic sled. Chain cruise 58; 5 May 1966. 7)
WHB sta. E No. 3; 39 50.5 N, 70 35 W;
823.5 m. Anchor
dredge.
Atlantis
cruise; 25
May 1961. 8) WHB sta. 87; 39 48.7 N, 70
40.8 W; 1,102 m. Epibenthic
sled. Chain cruise
50; 6 July 1965.
Range: At lower continental
slope to bathyal
depths (loo-1,100
m) from west of Greenland
south to the shelf and slope of the Middle Atlantic States. Probably
also present on the continental slopes of the southeastern
United
States
and northern Europe.

Diagnosis:
A small (3-5 mm) blind
Ampelisca. First antennae short, the first
two articles of the peduncle equal in
length. Basis of last percopod long, evenly
rounded below. Carina of urosome low
and sharply pointed posteriorly.
Remarks: A common bathyal species,
discussed at length
elsewhere
(Mills
1967~). The locality records listed above
extend its bathymetric range to 1,100 m
on the Gay Head-Bermuda
transect, but
do not change the conclusion that its optimum depth is about 400 m and that it
overlaps the range of A. agassixi 011 the
continental shelf and of various species of
Haploops and Byhlis (living
mainly at
greater depths) on the continental slope.
Ampclisca uncinata C hevreux
(Figs. 2A and 3)
Ampelisca
uncinuta
Chevreux
1887, p. 573574; 1900, p. 4244,
Pl. 6.
1) WHB sta. D No. 1; 39
Locality
records:
54.5 N, 70 35 W; 466.7-508.7
m. Anchor
dredge.
Atlantis
cruise 277; 23 May 1962. 2)
WHB sta. 96; 39 55.2 N, 70 39.5 W; 498 m.
Epibenthic
sled. Chain cruise 58; 27 April 1966.
3) WHB sta. 105B; 39 56.5 N, 71 03.6 W;
530 m. Epibenthic
sled. Chain cruise 58; 5 May
1966. 4) WHB sta. E No. 3; 39 50.S N, 70
cruise;
35 W; 823.5 m. Anchor dredge. Atlantti
25 May 1961. 5) WHB sta. 87; 39 48.7 N,
70 40.8 W; 1,102 m. Epibenthic
sled. Chain
cruise 50; 6 July 1965.
Ra,nge: Probably widespread
around the North
Atlantic
at bathyal depths (350-1,100
m) from
Cap Finisterre
to the Middle Atlantic States.

Diagnosis:
A medium-sized Ampelisca
lacking
cornea1 lenses and with
an
obliquely truncated head; propodus of the
last peroopod greatly expanded, mcrus
with a posterodistal sctosc lo,be partially
covering the carpus, and dactyl curving
forward into a fine point.
Adult female: Well characterized
by
Chevreux
(1887, 1900). My specimens
agree in minute detail with his descriptions and figures.
Aduk male: Not known. The juvenile
male figured by Chcvreux (1900, Pl. 6,
Fig. 3f) is certainly that. It differs very
little from the female in characters usually sexually dimorphic
(third uropods,
carina of urosome, length and setation of
antennae).
Remarks: Species recorded before only
by Chcvreux ( 1887, 1900)) who described
seven specimens taken between 363 and
510 m off Cap Finisterre in the eastern
North Atlantic, but likely to be widespread
around the North Atlantic
at bathyal
depths. Ampelisca gibba G. 0. Sars seems
the closest relative but differs in having
cornea1 lenses and a more pronounced carina of the urosome. Published records of
A. gibba from the North Atlantic refer to
collections made between 169 and 2,300 m
(Shoemaker 1931; Stophonsen 1935). The
deep collections, especially those below
300 m, may have been of A. uncinata, but
this can only be determined by examination of the original mat&al.
lacks cornea1
Although
A. uncinata
lenses, the ganglia ,of the eyes appear to
be present below the cuticle. This may
indicate that at some stage ,of the lift
history, probably breoding -time, the species ventures into lighted parts of the water column, although the greater part of
lift is undoubtedly benthic.
Byblis

brachycephala, n. sp.
(Figs. 2B and 4)

Type specimens: 3.4 mm adult 9, Type. WHB

sta. 87; 39 48.7 N, 70 40.8 W; 1,102 m.
Large collection
of paratypes, same locality.
All
in the United States National
Museum.
1) WHB sta. E No. 3; 39
Locality
records:
50.5 N, 70 35 W; 823.5 m. Anchor dredge.

NORTH

FIG. 2. A. Ampelisca uncinata

C. Byblis medialis n. sp., 8.0 mm

ATLANTIC

DEEP-SEA

AMPIIIPODS

361

Chcvreux,
6.5 mm 9. 13. Byhl& brachycephala
n. sp., 3.5 mm Q.
0. The bar beside cnch specks represents 2 mm of lengtlh.

362

ERIC L. MILLS

FIG. 3. Ampelisca uncinuta Chevreux, 8.2 mm ovigerous

0. WHB sta. E No,. 3, 39 50.5 N, 70
35 W; 823.5 m. A-Head;
B-last
somite of pleon and urosome; C, D, E-right
uropods 1, 2, 3;
N-left
mandible;
O-hypophar17, G-gnathopods
1, 2; H, I, J, K, I,-pcreopods
1-5; M- t&on;
ynx; P, Q-maxillae
1, 2; R-mnxillipecl.

NORTII

ATLANTIC

DIXP-SEA

Atlantis cruise; 25 May 1961. 2 ) WHB sta. 87;

39 48.7 N, 70 40.8 W; 1,102 m. Epibenthic
sled. Chain cruise 50; 6 July 1965. 3) WHB
sta. 73; 39 46.5 N, 70 43.3 W; 1,470-1,330 m.
Epibenthic
sled. Atlantis II cruise 12; 25 August
1964. 4) WHB sta. 131; 39 38.5 N, 70 36.5
W to 39 39 N, 70 37.1 W; 2,178 m. Epibcnthic sled. AtZa,nti I1 cruise 30; 18 December
1966.
Range: Known only from the Gay Head-Bermuda transect between
approximately
800 and
1,500 m. To be expected at equivalent
depths
around the North Atlantic Ocean.

Diagnosis:
A very small blind BybZis
with a short head (just equal in length
to the first two body segments ) and an
inflated first segment of the peduncle of
the first antenna. Lower Epont margin of
the head straight, only slightly angled.
First uropod long and narrow, extending
well beyond the end ,of the second. First
and second antennae equal in length, long.
Adult
females:
Length
3.44.0
mm.
Head shorter than or equal to first two
body segments; anterior margin blunt,
slightly oblique. No cornea1 lenses. First
and second antennae equal in length,
about % the length of the body. First
segment ,of peduncle of first antennae
much enlarged.
Mouthparts,
gnathopods and first four
pereopods as illustrated.
Fourth
co,xal
plate about as long as deep, posterior projection short, rounded at the tip. Pcrcopod
5, expansion of basis rounded distally,
sometimes slightly quadrate at the posterodistal corner; merus and carpus about
equal in length along anterior margins;
propodus relatively broad, a little more
than half the breadth of the carpus.
Second and third cpimcral plates rounded
at the posterior corner. First segment of
the urosome with a steeply-rising
bluat
carina, rounded at the posterior.
First uropod very long, narrow, rami
extending a third of their length beyond
the end of the second uropod; peduncle
with a large spine posterolaterally;
outer
ramus, outer margin with 3-5 spines, inner
margin spineless; inner ramus, inner margin with 2-3 spines, outer margin spincless. Second uropod; outer ramus slightly
shorter than inner, outer margin with 1

AMPIIIPODS

363

spine, inner margin spineless; inner ramus,

outer
margin
spineless, inner margin with
1 spine, Third uropod narrow with both
rami tapering evenly posteriorly; outer ramus longer than inner, with a small lateral seta near the tip and a small terminal
tuft of setae; inner ramus with 3 short
spines on the inner margin.
Telson narrowly rounded distally, cleft
for nearly half it-s length; 2 small spines
on upper surface at level of base of cleft.
Adult male: Length 3.7-3.9 mm. Slimmer and more streamlined than the female. Both pairs of antennae slimmer than
in the fomale; all three peduncle articles
of first antennae with close-set rows of
fine setae ventrally.
Coxal plates much
shallower than in female, but: pereopods
similar. First and second uropods slimmer than in female; rami of third uropods slightly broader at base. Urosomc
deeply depressed ahead of a steeplyrising rounded carina.
Remarks:
This little
bathyal
BybZis
shares various features with B. crassicornis
Metzgcr and B. abyssi G. 0. Sars, known
from equivalent depths in the northcast
Atlantic, Norwegian Sea, and Davis Strait
(Stephenscn 1925), and also rese.mbleas R.
guernei Chevreux, known only from a
single station off Cap Finistcrre (Chevreux
1887, 1900). It #differs from these species,
particularly
in the short, blunt head, enlarged first article of the peduncle of the
first antenna, broader propodus of percopod 5, longer uropod 1 (extending beyond
uropod 2)) more pronounced carina of the
urosome, and narrower, nondentate rami
of the third uropods.
It is peculiar #that B. brachycephaln and
the following species are known only from
the Gay Head-Bermuda
transect. They
are to be expected around the North Atlantic Ocean, as are spccics such as B.
crassicornis, B. abyssi, and B. minuticornis.
The low density of species of Byblis at
bathyal and abyssal depths may account
for the fragmentary knowledge of their
ranges in the North Atlantic. Comprchensive collections from Cape Cod north will
be necessary to fill some of the gaps.

364

ERIC L. MILLS

FIG. 4. Byblis brachycephala

n. sp., 4.0 mm adult 9, WHB sta. 87;
D, E-gnathopods
m. A-Head;
B-plcon
and urosome; C-t&on;
0, P,
l-5; K-right
mandible;
L, M-maxillae
1, 2; N-maxillipcd;
two somites of pleon and urosome of 3.5 mm paratype
$ , WHB sta.

39 48.7 N, 70 40.8 W; 1,102

1, 2; F, G, H, I, J-pereopods
Q-right
uropods l-3. B-Last
87.

NORTII

Byblis mcdialis

ATLANTIC

DEEP-SEA

n. sp. (Figs. 2C and 5)

Type specimens:
7.0 mm ovigerous
0, Type.
WHB sta. F No. 1; 39 47 N, 70 45 W; 1,500
m. Two paratypes, same locality.
Ail in United
States National Museum.
Locality
records:
1) WHB sta. 73; 39 46.5
N, 70 43.3 W; 1,470-1,330 m. Epibenthic
sled.
Atlantis II cruise 12.; 25 August 1964. 2) WHB
sta. F No. 1; 39 47 N, 70 45 W; 1,500 m.
Atlantis
cruise; 24 May 1961.
Anchor
dredge.
3) WHB sta. G No. 1; 39 42 N, 70 39 W;
2,000 m. Anchor dredge. Atlantis cruise; 24 May
1961. 4) WHB sta. G No. 9; 39 44.7 N, 70
38.3 W; 2,021 m. Anchor dredge. Atlantis cruise
284; 9 September 1962. 5) WHB sta. 103 ( G);
39 43.6 N, 70 37.4 W; 2,022 m. Epibenthic
sled. Chailt cruise 58; 4 May 1966. 6) WHB
sta. G No. 10; 39 44.8: N, 70 36f W; 2,030 m.
Anchor dredge. Atlantis cruise 284; 9 September
1962. 7) WHB sta. G No. 11; 39 43.2 N, 70
40 W; 2,085 m. Anchor dredge. Atlanth
cruise
284; 10 September 1962. 8) WHB sta. G No.
3; 39 41.4 N, 70 38.3 W; 2,086 m. Anchor
dredge. Atlantis cruise 277; 23 May 1962.
Range: Known only from the above localities
south and east of Gape Cod at depths of about
1,300-2,100 m.

Diagnosis: Distinguished by the combination of the following characters : article

one of the peduncle of first antenna short
and narrow; first antenna short, reaching
to the middle of article 5 od the peduncle
of second antenna; the dorsal carina of
urosome a gently-rounded
boss; car-pus of
pereopod 5 straight-margined
or slightly
convex distally; uropod 1 extending beyond uropod 2; tclson gently rounded apitally, cleft for half its length,
AduZt female: Length 5.7-7.4 mm. I-Iead
equal in length to first three body segmcnts combined,
front
margin
rather
blunt. No cornea1 lenses. First antenna
extending about a third of the way along
fourth article of second antenna. Second
antenna slightly shorter than body.
Mouthparts and pereopods l-3 as illustrated. Coxa of pcreopod 2 deep, posterior
projection short, rounded distally. Posterior margin of basis of pcrcopod 4 evenly
rounded. Pereopod 5, basis evenly rounded
distally or sometimes slightly
quadrate,
with slight angle at posterior corner; mcrus
shorter than carpus; carpus with a straight
or very slightly convex distal margin,
Second
and third
cpimcral
plates

AMPHIPODS

365

rounded at the posterior corner. First segment of the urosome with a gently-rounded
n-iddorsal boss.
First uropod long, narrow, extending bcyond the end of ,the shorter scoond uropod;
rami slightly curved laterally; #outer ramus
spineless except for 2 fine spines on thC
ventrolateral margin; inner ramus with a
series of stout spines on the fincly-scrrated inner margin, and a single spinc
proximally
on the irmcr margin. Second
uropod shorter than first, both rami slightly
curved laterally; outer ramus considerably
shorter than inner, with 2 stout spines
on the outer margin, inner margin finely
serrated; inner ramus outer margin spincless, inner margin finely serrated, with a
single stout spinc. Rami of third uropod
narrow, the facing margins slightly concave distally and finely serrated; outer
ramus inner margin with a marked tooth
proximally.
Telson cleft about half its length; a spine
on each side of the tip in a small notch;
tip gently rounded, very finely setose or
serrated.
AduZt male: Not known. A 6.8 mm subadult male from sta. G No. 3 is very similar
to adult females but has shallower coxal
plates and markod protuberances at the
end of the vasa deferentia. The boss of
the urosome should become more pronounced and complex as sexual maturity
is reached.
Remarks: Byblis medialis is so named
because its morphology places it between
B. abyssi G. 0. Sars and B. minuticornis
G. 0. Sars. It differs from B. abyssi in
having a more deeply cleft telslon, uropod
1 extending beyond uropod 2, broader and
shorter mcrus and carpus of pcreopod 5,
the merus of percopod 5 straight margin&I
(not concave) distally, and much less dcntate rami of uropod 3. Features distinguishing the species from B. minuticornis
are the longer first antenna (inchrding the
second segment of the peduncle ) , convex
carina of the urosome, more concave margins of the rami of uropod 3, and rour-rdcd
tip of the telson.
It might be possible to regard this spc-

366

ERIC L. MILLS

FIG. 5. BybZ& medialis n, sp., 7.0 mm ovigerous $?. WHB sta. F No. 1; 39 47 N, 70 45 W; 1,500
C-left
mandible; D, &--maxillae
2, I; F-maxm. A-Head;
B-h&
somite of plcon and urosome;
1, 2; 1, J, K, L, M-pereopods
1-5; N, 0, Pilliped ( palp and plates separated) ; G, Ii-gna&opc&
right urupods l-3; Q-telson.

NORTH

ATLANTIC

DEEP-SEA

ties as a subspecies of either B. abyssi or

B. minuticornis,
both found at roughly
equivalent depths in the northeast Atlantic
(Sars 1895; Stephensen 1925). The genus
ByhZis is so tight-knit in its morphological
features that such a decision would bc
hard to justify. As in other ampeliscids,
change (presumably
by
morphological
adaptive radiation)
has occurred mosaitally, so that it is difficult to draw clear
lines of morphological
relationship
bccause of the overlap of characters between
species; for example, different species may
have similar characters of head shape but
not of pcreopod 5, of pcrcopod 5 but not
of uropod 3, and so on
Older descriptions of species of Byblis
have often been almost impossible to USC
because of inadequate drawings of taxonomically
important
f eaturcs . Careful
drawings of the following features are particularly important for critical work on this
genus: head shape, shape of fourth coxal
plate, shape of basis of percopods 3 and
4, pereopod 5, all three uropods, and tclson. Uropods in particular may bc diagnostic and should be drawn with special
care.
Byblis gaimardi

(Kr$yer)

(Figs. GA and 7)

Byblis gaimardi (Krgyer),

G. 0. Sars 1895, p.
183-184, Pl. 64. Stebbing 1906, p. 113 (see this
reference
for a complete
synonymy;
see betow
for comments ) .
Mate&z2 examined:
Numerous collections
from
Labrador,
off Nova Scotia, Casco Bay (Maine),
and off Cape Cod in United
States National
Museum; also a large collection
from Point Barrow, Alaska, and an immature
from Penobscot
Bay, Maine.
Range: Circumpolar,
from Alaska and the East
Siberian Sea to the Kara Sea, Iceland, Greenland,
and the Canadian Arctic archipelago,
South in
the eastern Atlantic
to Scotland and Denmark,
and in the western Atlantic
to Cape Cod Bay.
Range in the Pacific
uncertain;
see Barnards
(19%) comments on a record from Monterey Bay,
California.
Depth
range 5-575 m (Shoemaker
1955; Stephensen 1925).

Diagnosis:
A large Byhlis (adult fcmales 15-25 mm); the front margin of the
head concave where the first antennae insert; first peduncular segment of the first
antenna barrel- or vase-shaped. First two

AMPIIIPODS

367

coxal plates not serrated; fourth coxal plate

deeply concave behind and drawn out to
a sharp point posteriorly; percopod 5 with
a slightly quadrate (rather than evenly
rounded) cxpandcd basis; car-pus 1.3 times
as long as mcrus and 1.8 times as broad
distally as greatest width of propodus,
giving impression of narrow car-pus; carina
of the urosomc raised, slightly sinuous dorsally, ending posteriorly in a raised transvcrsc ridge; telson short, blunt-ended, cleft
for a third or less of its length.
Remarks: Although not founld on the
Gay Head-Bermuda
transect, this spccics
is included for comparison with B. serrata,
which replaces B. guimardi east and south
of Cape Cod. In general, the figures of
Sars ( 1895, Pl. 64) characterize the species
well, including the adult male, which is
differences berarely seen. Morphological
twccn adult females of B. serm-~a and B.
gaimardi arc summarized in Table 1. Judd
( 1896, p. 601) tabulated some of the differences between the sexes.
Stebbing (1906) believed that Judds
(1896) figures and description of B. serrata rcferrod properly to B. gaimarcli and
placed them in the synonymy of that species. There can be no doubt that Judds
figures and description
arc truly of B.
serrata. It is difficult to ,understand why
Stebbing, usually so critical and careful,
made such an error, even emphasizing the
difference from Smiths species with an
exclamation point. A multiple confusion
may have led to this situation, since males
of Ampelisca
were often identified
as
Byhlis. For example, Judd described A.
agassixi ( later called A. compressa) as B.
agassixi. In addition, there is in the United
States National Museum (USNM 37160)
a collection of mature malts of A. vndorum
from Vineyard Sound, Massachusetts, misidcntificd as B. gaimardi. This collection,
made by the U.S. Fish Commission in
1871, was probably identified
by S. I.
Smith, and a similar misidentification
of
specimens,
if they were called B. serrata
and sent to Stebbing by Smith, might bavc
resulted in Stcbbings empha,tic denial that
Judd was dealing with B. serrutcl.

368

ERIC L. MILLS

FIG. 6, A. ByhZk gaimatdi

( Kr@yer ) , 20 mm female.
bar beside each species represents 2 mm of length.

B.

Byblk

serrata

Smith,

10 mm

$?. The

NORTH

ATLANTIC

DEEP-SEA

AMPIIWODS

369

and Byblis serrata Smith ( lo-wer ) . Byblis gaimardi, 20

FIG. 7. Byb1i.s gaimardi ( Kr$yer ) (upper)
A-head;
B-last
two somites of pleon
mm ovigerous
9, sta. 0620; Cape Cod Bay, Massachusetts:
and urosome; C-pcreopod
5; D, E, F-uropd~
1-3; G- inner margin of outer ramus of uropod 3;
H-telson.
A-head;
B-pleon
and
Byblti serrata, 9.5 mm ovigerous
0, sta. I?; Buzzards Bay, Massachusetts:
urosome; C-pcrcopod
5; D, E, I?-uropods
1-3; C- inner margin of outer ramus of uropod 3; Htelson; I-labrum;
J-hypopharynx;
K, L-maxillae
1, 2; M-maxilliped.

370

ERIC L. MILLS

TABLE

1.

Main

features

of morphology

sepnrating Byblis gaimardi

See also Figs. 7 and 8

(Kr$yer)

and B. serrata

Smith.

13. serrata

IIead

Front

margin

Eyes

Lower
trally,

pair directed anteriorly

and slightly laterally.

Antenna

noticeably

concave.

Front margin
hind antenna

and ven-

Lower

oblique,
1.

pair directed

almost

anteriorly

straight,

bc-

and ventrally.

First segment of pcduncla with convex

upper margin, narrowing
segment distally. Second segment of peduncle 2% x
length of first.

First
segment
of peduncle
with
almost
straight upper margin,
segment not narrowed distally.
Second segment of peduncle
11/-l%.
x
length of first.

Lower margin of plates 1 and 2 not

serrated. Plate 4 deeply concave behind,
posterior corner a long, sharp point.

Lower margin on plates 1 and 2 serrated.

Plate 4 shallowly
concave behind, posterior
corner short and blunt.

Basis slightly quadrate

Carpus 1.8 x as broad
est width of propodus.

distally.
distally as great-

Basis broadly rounded distally.

Carpus
x as broad distally
as greatest width
propodus.

Urosome

Carina raised, slightly

terminating
in a slight

sinuous dorsally,
transverse ridge.

Carina a low
posteriorly.

Telson

Short, blunt-ended;
less of its length.

Coxal

plates

Pcreopod

cleft

for

a thircl

Byblis gaimardi ranges south only to

Cape Cod Bay, then to the east and south
it is replaced by B. serrata at shallow and
moderate depths (to 200 m). Records of
B. gaimardi from Vineyard Sound (Shoemaker 1931, 1955) are probably in error;
they may have been based on the collection of A. vadorum just mentioned (USNM
37160), misidentified as B. gaimardi. Collections of B. gaimardi from Cape Cod
Bay, taken in the Systematics-Ecology
Programs Biotic Census, are from depths
of 36-56 m and summeir temperatures of
4.2-6.OC (A. Michael, personal communication). By contrast, B. serruta taken at
200 m on the Gay Head-Bermuda transect
(south and east of Cape Cod) would expcricncc temperatures well above this, 7-

or

Acutely-rounded
length.

smooth

boss, slightly

at end;

cleft

for

2.2
of

raised
half

its

15C, throughout the year (Sanders et al.

1965; Schopf 1967, 1968). Summer water
temperatures in the range of 6-8C appear
critical in restricting the southward extent
of B. gaimardi and the northward
extent of B. serruta. Cape Cod, with cold
waters to the north and northeast yearround and warmer water to the south and
southeast in summer, forms a particularly
striking zoogeographic boundary for this
pair of species.
Byblis serrata S. I. Smith (Figs. 6B, 7, and 8)
Byblis sew&u S. I. Smith -in Verrill and Smith
1873, p. 561. Judd 1896, p. 59&599,
Figs. 4-8.
Holmes 1903, p. 274; 1905, p. 482, and Fig.
Kunkcl 1918, p. 67-70, Fig. 10.
Byblis serrata (in part), Stebbing 1906, p. 115116.

A-H-9.5
mm ovigerous
9, sta. P; inside Quicks Hole, Buzzards
FIG. 8. Byblis sew&z
Smith:
I-J-8.5
mm COTYPE
$, USNM 35609; Woods Hole, Massachusetts.
A, BBay, Massachusetts.
Gnathopods 1, 2; C, D, E, F-percopods
14; G- 9 right mandible; H-ventral
view of palp of right
mandible;
I-head
of adult $ ; J-last
somite of pleon and urosome of adult $ .

NQRTII

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AMPIIIPODS

371

372

ERIC L. MILLS

ByhZz.s guimardi
(Kreyer)
(in part)
Stebbing
1906, p. 113 (see remarks under B. gaimmdi).
Locality
records:
1) Station P; inside Quicks
Hole, Buzzards Bay, Massachusetts;
15 m. Sand
and silt. 5 August 1965. 2) About 4 km off
Buoy No. 29 (504);
near Gay Head; 32 m.
21 June 1967. 3) WHB slope -sta. No. 2; 40
08.8 N, 70 42 W; 200 m. Anchor dredge sampie. Atlantis cruise 283; 28 August 1962. Other
specimens
examined : several
collections
from
southern
New England
area in United
States
National
Museum, - also USNM
35609, cotypc,
Woods IIolc, Massachusetts, 3 September 1871.
Range: From the south side of Cape Cod to
at least Chesapeake
Bay and probably
to the
Gulf of Mexico, at intertidal
depths to 200 m.

Diagnosis: A medium-sized Byblis (812 mm) with a blunt front margin of the
head; first peduncular segment of the first
antenna cylindrical; ventral margins of the
first two coxal plates serrated; fourth coxal
plate short and only shallowly concave
behind, and with a short blunt projection
posteriorly; carina of the urosome a low
smooth boss (only slightly raised posteriorly ) ; telson narrowed
and slightly
pointed apically.
Adult female: Head about equal to
first three body segments in length; anterior margin usually slightly angled, very
slightly concave. Lower pair of corncal
lenses directed anteriorly
and ventrally.
First antenna slightly longer than the peduncle of second antenna; first article of
peduncle more or less cylindrical, not narrowed distally, upper margin straight or
only slightly curved; second article of peduncle 1%-l% times the length of first.
Second antenna 2/R-% the length of the
body.
Mouthparts
as illustrated.
Note especially the flattened and expanded second
article of the mandibular palp.
Pereopods as illustrated. Note especially
the indented or serrated lower margin of
the first two coxal plates, and the shape
of the fourth coxal plate (plate of pcreopod 2), which is deep, relatively narrow,
only shallowly concave behind and with
only a short blunt point posteriorly. Pcrcopod 5 with an expanded basis which is
evenly rounded and heavily sotose distally; carpus about twice as long as merus
and 2.2 times as broad distally as the

greatest width of the propodus, giving the

impression of a broad carpus and narrow
propodus.
Third epimcral plate projecting pos teriorly as a rounded-quadrate
lobe. First
segment of the uroeome with a low smooth
carina, only slightly raised posteriorly.
First uropod long, extending beyond the
second; pcdunclc with stout spines on the
inner margin, outer margin spineless; outer
ramus slightly shorter than inner, upper
margins spineless; inner ramus with spincless outer margin and 6+ stout spines on
the inner margin, Second uropod short;
pcduncle with a long spine on the inner
margin and 3 short spines on the outer
margin; outer ramus shorter than inner,
outer margin with 4+ spines, inner margin
spinclcss; inner ramus, outer margin spineless, inner with 3f long spines. Third uropod short, extending only to end of first
uropod; outer ramus with 2 spines on the
outer margin, inner margin strongly serrated in the middle, some of the teeth and
part of the distal inner ,margin with fine
denticulations;
inner ramus outer margin
strongly serrated in the middle, part of the
distal outer margin finely denticulated; inncr margin straight, with 4 stout spines.
Telson acutely-rounded
distally, with 2
small spines in notches near the apex; cleft
for half its length.
A.duZt male: Slightly smaller and narrower-bodied than the adult female. Head
similar to that of the female, but slightly
rostrate between the bases of the first
an tcnnae.
First antenna short, segments of peduncle with close-set groups of comblike setac
on the veetral surface. Second antenna
long and narrow, the third, fourth, and
fifth articles of the peduncle with closeset groups of comblike sctac on the dorsal
surf ace.
Carina of the urosome a smoothly
rounded lobe, raised well above the following segment posteriorly. First and sccond uropods similar to those of the female.
Third uropods, outer ramus inner margin
strongly scrratcd along the whole margin;
iniler
rarnus
outer margin concave near

NORTH

ATLANTIC

DEEP-SEA

middle, anterior edge of concavity strongly

serrated, inner margin set with long plumose setae (see Judd 1896, Fig. 8). Judd
(1896) compared the morphology of male
and female in some detail and speculated
on the functions of the setose antennae of
the male and of other parts.
Remarks: S. I. Smith described Byblis
serrata in 1873 without figuring it (in Verrill and Smith 1873), and although this
species was later included in works by
Judd ( 1896), Holmes (1903, 1905) and
Kunkel ( 1918)) complete figures have not
been prepared previously.
Smith based
his description on a female or fclmalcs, but
the type lot canrmt be located. A specimen designated cotype, an 8.5 mm adult
male, exists in the United States National
Muscum. This is apparently the specimen
figured by Kunkel ( 1918)) and I have used
it for some of the drawings in Fig. 8 and
the written description Iof the adult male
above.
Stebbing (1906) fused B. minuticornis
with B. serrata on the basis of Smiths written description,
but this is wrong, as
Stephensen (1925) has shown. The two
are actually very different in a number of
morphological
features (cf Sars 1895, Pl.
66) and also differ markedly in ecology
and distribution since B. serrata occurs at
littoral to continental
shelf depths frolm
Cape Cod south, whereas B. minuticornis
is bathyal and unknown south of the Norwegian Sea.
The distribution and ecology of B. serrata is known only in outline. Most rcco,rds
are from the southern Cape Cod region,
where the species ranges from the intcrtidal (sparsely) to 200 m. Smiths records
(Verrill and Smith 1873) were on fine
compact mud and sand in 20-29 fathoms
off Vineyard Sound and on fine sandy
mud cast of Block Island Sound, Smith
(1950), working in Block Island Sound,
outlines some aspects of the biology of B.
serrata. It breeds from June to October.
The species is a tube dweller and feeds
like other ampeliscids
(Enequist
1950;
Mills 1967a) in the tops of flattened tubes.
New tubes may be built rapidly after dis-

373

AMPHIPODS

turbancc. It is the seventh most abundant

species of invertebrate in the diet of fishes
of Block Island Sound, where it is eaten
mainly by winter flounder (5.6% of food
eaten by weight) sea robin (1.4%), skate
(1.1%)) and sculpin (0.2%). Smith calculated the ratio of annual consumption to
standing crop ,of B. serrata as 6.7 (compared to a crudely estimated 14.3 for the
abundant and ecologically-important
specks Leptocheirus pinguis) . Wass ( 1963)
recorded B. serrata in Chesapeake Bay, but
there arc apparently no well-established
records south of this.
The specimens from WHB slope sta. No.
2 are at first view unexpected, since the
cold-water species B. gaimurdi might be
expected there. Temperature records for
200 m from the Gay Head-Bermuda
transect (shown graphically in Sanders ot al.
1965, and as isotherms by Schopf 1967)
are from roughly 7-15C, not far below
those at which B. serrata breeds near Cape
Cod. Byblis gaimardi appears to be excluded by such high temperatures, as discussed above.
Genus Byblisoides

K. H. Barnard

K. H. Barnard 1931, p. 426; 1932, p. 86-87,

Fig. 41; J. L. Barnard 1969, p. 132.
With the description
of the new species belolw,
the characteristics
of the genus outlined by K. H.
Barnard (1931, 1932) and J. L. Barnard ( 1969 )
must bc changed as follows.
(Diagnosis based on
K. I-1. Barnards
definition,
with emended portions in italics. )

Diagnosis of genus: Head elongate, both

pairs of antennae arising on the anterior
margin, anteroventral corner rounded, subquadrate, or produced into an acute process. No cornea1 lenses; ocular pigment not
visible. Side plates l-4 sloping forward,
the fourth sometimes narrowed distally,
more often of uniform
width.
Telson
ovate, longer than broad, cleft nearly to
base. Antenna 1 very short, flagellum 2-3
jointod. Antenna 2 very stout, with long
stout setae. Mandibular palp slender, second joint long and curved, third joint a
third to half the length of second. Mouthparts set posteriorly on ventral surface of
head. Maxillipecl very large. Pereopo&
3

374

ERIC L. MILLS

and 4 with second joint broadly expanded,

fifth joint with marginal spines or setae.
Pcreopod 5, second joint distally
expanded, marginal setae extending round
on to margin facing third joint, sixth linear, dactyl spiniform.
Branchial lamcllae
simple. Antennae, urosome, and uropods
show marked sexual dimorphism.
Byblisoides profundi n. sp.
(Figs. 9A and 10)
specimen: 16 mm ovigerous Q, Type.

Type
WHB sta. 92; 36 20 N, 67 56 W; 4,694 m.
In United States National Museum,
Locality
records: 1) WHB sta. 70; 36 23 N,
67 58 W; 4,680 m. Epibenthic
sled. Atlantis II
cruise 12; 23 August 1964. 2) WHB sta. 92;
36 20 N, 67 56 W; 4,694 m. Epibenthic
sled.
Atlantis
II cruise 17; 13 December
1965. 3)
WHB sta. 84; 36 24.4 N, 67 56 W; 4,749 m.
Epibenthic
sled. Chain cruise 50; 4 July 1965.
4) WHB sta. 122; 35 50 N, 64 57.5 W, to
35 52 N, 64 58 W; 4,833 m. Atlantis II cruise
24; 21 August 1966. 5) WIIB sta. LL No. 1;
35 35 N, 67 25 W; 4,977 m. Atlantis cruise
273; 28 September 1961.
Range: Known only from the Gay Head-Bermuda transect at depths of 4,680-4,977
m, but
to be expected elsewhere
at abyssal depths in
the North American Basin.

Diagnosis:
Distinguished
by the combination of rounded anteroventral corner
of the head, broad coxa of pereopod 2,
narrowly elongate shape of basis of pereopod 5, and nonbilobed carina of the
urosome.
Adult female: Head large, elongated,
an teroventral corner rounded or subquadrate, not projecting.
Antenna 1 short, extending almost to end of fourth pcduncle
scgmcnt
of antenna 2; flagellum vcly short,
of 2-3 articles. Antenna 2 short, stout, last
article of peduncle and articles of flagellum with stout: double-tipped setae.
Mouthparts forming a large bundle posteriorly on the ventral surface of the head.
Labrum broad, anterior surface slightly
concave. Mandible large; second segment

of palp elongated and strongly curved, 2

to 3 times as long as third segment; right
mandible with a stout spinelike lacinia
mobilis and a row of about 4 spines betwocn incisor and molar processes; left
mandible with only a large toothed plate
and 2 or 3 sctae in the equivalent po,sition.
Other mouthparts as illustrated.
First coxal plate evenly rounded distally. Fourth coxal plate broad, distal margin not oblique.
Pereopod 5 basis long, extending to at
least midpoint
of carpus, distal margin
slightly angular, coming to a broad distal
point. Plcon no,t ridged dorsally.
First
segment of urosome with a projecting
acute carina which is a single lobe when
viewed from above.
Uropods all long and narrow; spination
and setation as illustrated.
Number of
spines reduced in immatures,
Adult male: The only complete specimen available (from WHB sta. 70) is 14.5
mm long and probably one molt: short of
the full adult male morphology.
Differs from female mostly in characters
of antenna, urosomc, and uropods. First
segment of urosome deeply concave anteriorly, extending posteriorly into a high,
rounded projection
having two lateral
lobes when viewed from above. The fused
second and third segments of the urosome
have small roundeld projections on either
side of the dorsal depression. Features of
uropods similar to adult female except as
follows. Uropod 1, outer ramus outer margin with only 6 short spines; inner ramus
outer margin with 4 small spines. Uropod
2 missing and not known.
Body slimmer and more streamlined than
in female.
A broken specimen (head and part of
thorax) of a fully adult malt from WIIB
sta. 70 has a greatly elongated and narrowed flagellum of the second antenna,

FIG. 9. A. Byblisoides
profundi
n. sp., 12 mm 9.
$. C. Haploops similis Stephensen, 7.5 mm 0. Tho
length.

B. Haploops setosa Boeck, 12 mm immature

bar beside each species represents 2 mm of

SORTH

ATLANTIC

DEEP-SEA

AMPHIPODS

B
i_
FIG. 10. Byblisoides pmfundi n. sp., 16.0 mm
m. A-Head;
B-pleon
and urosome; C-right
pods 1, 2; H, I, J, K, L-pereopods
1-5; M, N,
pleon and urosome of 14.5 mm adult $, WHB

adult 9, WHB sta. 92; 36 20 N, 67 56 W; 4,694

mandible;
D, E-right
maxillae 1, 2; F, G-gnathoO-right
uropods 1-3; P-telson.
B-last
somite of
sta. 70; 36 23 N, 67 58 W; 4,680 m.

NORTH

ATLANTIC!

DEEP-SEA

and the pcdunclc of antenna 1 is also

somewhat elongatod.
Remarks: The discovery of B. profundi
has made necessary a revision of the charactors of the genus as given by K. H.
Barnard (1931, 1932) and J. L. Barnard
(1969), since the post antenna1 angle
(anteroventral
corner of the head) is not
produced in B. profundi.
Byblisoides profundi
differs from the
four other species of the genus in the
combination of characters given in the diagnosis above. Its only unique characteristic is the rounded (not produced)
anteroventral corner of the head, just below
the second antenna.
J. L. Barnard (1961, 1964) discussed
the rather small taxonomic differences bctwccn the four species known previously
(B. juxtacornis K. H. Barnard, B. arcillis
J. L. Barnard, B. esferis J. L. Barnard, and
B. bhensis J. L. Barnard) and concluded
that specific, not subspecific distinction is
warranted.
I fully agree with him, espccially now that a distinct new species is
known from abyssal depths in the northern hemisphcrc and there are also indications of an undescribed species from the
Peru-Chile
Trench at depths (of 3,0004,000 m.
It appears likely that B. profundi is a
characteristic
species of abyssal depths
below about 4,000 m in the western North
Atlantic Ocean. B yblisoides juxtacornis is
known from the area of the southern
Antarctic Peninsula (Palmer Archipelago)
at 160335 m (K. H. Barnard 1932), B.
arcillis has been taken in the Tasman Sea
at 610 m, and B. esferis in Makassar Strait,
Indonesia, at 1,560 and 2,000 m (J. L.
Barnard 1961). The first species of BybZisoides found in the northern hemisphere,
B. blasensis, was taken in the Caribbean
Sea north of Panama at 1,715 m (J. L.
Barnard 1964). The fact that all arc cycless and adapted for feeding in soft ooze
(see remarks following)
suggests that the
bathyal species of Byblisoides may also be
taken at depths greater than Ithose recorded. Barnards ( 1964) suggestion that
Byblisoides is a bathyal genus displaced

AMl?IIJJ?ODS

377

from the abyss will probably prove wrong

as more abyssal species arc discovered.
Several morphological
features are of
great interest in this unusual genus. The
short brushlike first antennae and stout
swecplike second antennae are found in no
other ampeliscid genus. The second antennae are probably
used in powerful
sweeping movements across the surface of
deep-sea scdimcnts. The extrinsic muscles
of the scoond antenna are greatly enlarged
in length and volume compared to those
in the genus Ampelisca. Ampelisca, which
uses the second antonnac in feeding (Mills
1967a; Enequist 1950), in turn has relativcly larger extrinsic muscles of the sccond antenna than do members of the
genus Gammarus, where the second antennae are not actively used in feeding mothen, B yblisoides has
tions. Apparently,
carried the USC of the second antennae in
feeding further than most other amphipods. The actual mechanism of feeding
will have to await functional
studies.
Other morphological features which I bclieve will prove to be related to an unusual
mode of feeding are the forw,at-d-slanted
coxal plates l-4 and the highly setose
gna thopods. Gnathopod 1, in particular,
must be closely concerned with feeding.
In the female from WHB sta. 92 (the
largest and best-prescrvcd specimen available), gnathopod 1 is tucked up under the
head, with sharp bends at the basi-ischial
and ischio-meral joints, making it U-shaped,
and bringing merus, car-pus, and propodus
into contact with the maxilliped.
There is
also a sharp bend at the carpo-propodal
joint in which much fine yellowish material, similar to that in the mouthparts and
foregut, has accumulated
,on the long
lower setac of carpus and propodus. The
first gnathopod is positioned like a second
maxillipod
and may well act like one,
being closely integrated with maxillipod
and maxillae in processing food.
The fine material removed from mouthparts and gnathopods is made up almost
entirely of fine mineral grains (most of
them clear and apparently siliceous), small
Foraminifera and fragments of larger ones,

378

ERIC L. MILLS

about 50% amorphous material (fine organic matter?), and a few sponge spicules.
Clearly, B. profundi like its shallow-water
cousins is a selective deposit feeder, but
the actual means of collecting and processing food is not known.
Haploops

sctosa Boeck (Figs. 9B und 11)

Haploops
setosa Boeck.
G. 0. Sars 1895, p,
194-195, Pl. 68. IIO~ES 1905, p. 525, St&bing
1906, p. 117-118.
Stephensen 1925, p. 152-154;
1926, p. 52; 1928, p, 117; 1929, p. 78-79, Fig,
83; 1933, p. 26; 1935, p, 138-139; 1940, p. 18.
Schellenberg
1925, p. 201. Shoemaker 1931, p.
12-13.
Gurjanova
1951, p. 323-324,
Figs, 190
and 191. Dunbar 1954, p. 721-723,. Ziegelmeyer
1959, p, 99-102.
J. L. Barnard 1961, p. 66-67.
Kanneworff
1966, p. 198-200.
Haploops
robusta.
G. 0. Sars 1895, p. 195196, Pl. 68. Holmes 1905, p. 525. Stebbing 1906,
p. 118. Stephensen 1925; p. 154; 1935, p, 139;
1940, p. 18. Gurjanova
1951, p, 324-326, Fig.
192. J. L. Barnard 1961, p. 66-67.
Locality
records:
1) WHB sta. 73; 39 46.5
N, 70 43.3 W; 1,470-1,330 m. Epibenthic
sled.
Atlantis II cruise 12; 25 August 1964. 2) WIIB
sta. F No. 1; 39 47 N, 70 45 W; 1,500 m.
Anchor dredge.
Atlantis
cruise; 24 May 1961.
3) WHB sta. G No. 1; 39 42 N, 70 39 W;
2,000 m. Anchor dredge. Atlantis cruise; 24 May
1961. 4) WHB sta. G No. 9; 39 44.7 N, 70
38.3 W; 2,021 m. Anchor
dredge.
Atlantis
cruise 284; 9 September 1962.. 5) WHB sta. 103;
39 43.6 N, 70 37.4 W; 2,022 m. Epibenthic
sled. Chain cruise 58; 4 May 1966. 6) WHB
sta. G No. 10; 39 44.8 N, 70 36 W; 2,030 m.
Anchor
dredge.
Atlantis
cruise;
9 September
1962. 7) WHB sta. G No. 5; 39 41.5 N, 70
37.8 W; 2,034 m. Anchor dredge. Atluntis cruise;
17 January 1962. 8) WHB sta. G No. 11; 39
43.2 N, 70 40 W; 2,085 m. Anchor dredge.
Atlantis cruise 284; 10 September 1962. 9) WHB
sta. 66; 38 46.7 N, 70 08.8 W; 2,802 m. Epibenthic sled. Atlantis
II cruise 12; 21 August
19fX
10) WHB sta. 64; 38 46 N, 70 06 W;
2,886 m. Epibenthic
sled. Atlantis II cruise 12;
21 August 1964.
Other material
examined:
Collections
in the
United
States National
Museum
(USNM ) from
Ungava
Bay, Labrador,
St. Lawrence
estuary,
Scotian Shelf, Bay of Fundy, off Georges Bank,
and off Delaware
Bay.
Range: Widely
distributed
around the North
Atlantic
Ocean and in contiguous
seas from the
Kara Sea, Barents Sea (271-400
m) and Spitzbergen (69-330 m) south to the western British
Isles ( 190-1,375 m) and west to Iceland (2601,469 m), Greenland
( 19-2,702 m), and Ungava
Bay (92-230 m); thence south to Newfoundland,
St. Lawrence
estuary, Nova Scotia region (loo-

350 m), and Georges Bank (90 m), apparently

reaching a southern limit at about 39 N (depths
about 1,300-3,000
m). Range and depths summarized from Dunbar (1954), Shoemaker (1931),
and Stephensen (1925) as well as above locality
records.

Diagnosis: A large blind Haploops, probable size at breeding 15-30 mm, First
antenna almost as long as second; secon#d
segment of peduncle slightly more than
twice as long as the first, but becoming
relatively
shorter with increasing
size.
Long setae on the dorsum of last two
thoracic and first three abdominal segments. Basis of last pereopod broad, extending at least to end of ischium, usually
about halEway along merus; posterodistal
lobe of basis rounded or slightly angled.
Subadults: Most individuals in the collections from the transect are ,subadult and
characterized by Fig. 11. Sars (1895) gave
useful drawings of northern specimens.
Compared with arctic and boreal specimens, those from the transect are generally slimmer-bodied
(probably a function
of smaller size), have a slightly narrower
basis and carpus of pereopod 5, more spinose carpus ,of pereopod 5, the head is
blunter, with its front margin less angled,
and the last urosomal segment lacks or
has only very small tubercles on each side.
In northern specimens )the urosomal tubercles arc conspicuously present, even in the
smallest immatures.
Adult male: Adult males are very rare;
only the head of a mature male is present
in the transect collections (WHB sta. G
No. 11, 2,085 m), but a complete adult
male in somewhat battered condition was
collected from the stomach of a cod at
Port Burwell, Ungava, in 1927 (USNM
112832). Main differences from the figured specimens follow.
First antenna long and slim, ventral surfaces of peduncle segments with regular
rows of brushlike setae. Second antenna
very long and slim, equaling the body in
length; peduncle segments 3, 4, and 5 with
regular rows of brushlike setae dorsally.
Gnathopods and pereopods l-2 heavily set
with plumosc setae. Gills long and heavily
pleated. Carpus of! pereopod 5 elongate,

NORTH

ATLANTIC

DEEP-SEA

AMPIILPODS

379

FIG. 11. IIupEoops sdosu Boeck, 13.4 mm immature

8, WIIB sta. G No. 1. 39 42 N 70 39 W*
2,000 m. A-Head;
B-pleon
and urosome; C--telson;
D, E-maxillae
1 2; FLinner
and outer plate;
of maxillipcd;
G, H-gnathopods
1, 2; I, J, K, L, M-pereopods
1-5; i-right
mandible. 0 P Q3 3 ,
right uropods 1, 2, 3.

380

ERlC L. MILLS

strongly concave below; propodus very

narrow, with 4 spines on the posterior
margin. Carina of the urosome a strongly
raised lobe, projecting
posteriorly
as a
squarish process. Large spine at outer corner of pedunclc ,of uropod 1 absent. The
whole inner margin of inner ramus of uropod 1 set with spines. Margins of rami
of uropod 2 with many close-set spines.
Rami of uropod 3 narrower than in subadult, all margins with long plumose setae.
Length 25 mm.
Adult female: There are no adult females in the collections from the transect,
but a specimen from a cod stomach, Port
Burwell, Ungava (USNM 112832)) allows
the following
differences from subadults
to be noted.
Peduncles of first and second antennae
set with many long slim setae. Pereopods
l-4 heavily set with long plumose sctac.
Bases of pcroopods 3-5 with many long
plumose setae. Carpus of pcrcopod 5 with
5 spines set in notches on anterior margin,
posterior margin with spines and plumose
setae. Well-markeld tubercles on either
side of urosome behind carina. Length
21 mm.
Remudcs: This large and widespread
species has been refigured to clarify some
of the drawings of Sars ( 1895). The taxonomy of the genus Haploops is particularly difficult,
so I have figured some
pereopods
important
parts, particularly
3-5, uropods l-3, and the telson in considcrable detail to allow comparison with
other species.
The relationship
between H. setosu
Bocck and H. robustu G. 0. Sars has been
a matter of difficulty, now apparently rcsolved (at least as a nomenclatural problcm) by H. robustu being placed in the
synonymy of I-I. setosu by Dunbar (1954)
and Kanneworff (1966). However, the nature of the forms originally described as
robustu and setosu remains a problem
Dunbar found a combination of the characters distinguishing
I-I. setosa and F1. rohustn in his specimens from Ungava Bay,
and the same is true of specimens from
the Gay Head-Bermuda
transect. Most

specimens of this latter material have a

head shape intermediate between E-I. setosu and H. rohustu, antenna 1 pcdunclc
like H. setosu (with exceptions in large
animals nearing adulthood),
basis of pereopod 5 like 11. setosu (a few like I-I.
robusta), toothing and shape of the third
epimeral plate like I-I. setosu ( except for
large subadult females) and telson like H.
robustu (a few like H. setosa).
Kanneworff
(1966) suggests that H.
robustu is the young male of H. setosu,
but examination of the specimens from
the Gay Head-Bermuda
transect leads to
different conclusions. In most #of lthe small
specimens, the preponderance of characters is like H. setosu (although the telson
i.s usually more deeply cleft). The largest
specimens ( 13-17 mm), independent
of
apparent sex, show a shortened second article of the poduncle of antenna 1, while
large females (2 examined) have deeper
side-platos, more curved posterior margin
of the third epimeral plate ( with little
toothing at the oorncr) and straighter inner margin of the inner ramus of uropod
3 (the last 2 characters found in H. robustu) than a similar-sized subadult male.
The characters of H. robusta appear to
arise as adulthood is neared and some are
associated with sexual dimorphism.
A
more detailed ,analysis on greater numbers
of specimens is desirable.
The depth range occupied by II. setosu
( lOO-2,800 m) is constant from 78 N to
about 44 N. South oE 44 N, the minimum depth appareatly increases, reaching
about 500 m at 39 48 30 N. From
published records, the temperature range
occupied
throughout
the geographical
range is essentially constant, extremes bcing -l.lC and 6.2C. There is an apparent
downward dip of distribution south of 40
N which can be accounted for by high
water temperatures at upper slope depths
south of Cape Cod (cf figures in Schopf
1967). The distribution
of IT. setosu is
apparently governed by the 7C isotherm.
North of Cape Cod, animals at 100-200 m
will not encounter water much warmer
than 7C, whereas south of Cape Cod tem-

NORTII

ATLANTIC

peratures at these depths ,range from lo16C. At the latitudes of the transect: (39
48 and south), 7C water is reached at
about 500 m (Fig. 4 of Sanders, et al. 1965).
This corresponds closely with #the occurrence of II. setosu at 461 m at 39 48 30
N, 70 54 W (Fish IIawk Sta. 880, USNM
33785). Thus, II. setosu appears to follow
the 7C isotherm to greater depths at the
southern end of its range aad can bc regarded as showing a form of low latitude
submergence.
Haploops similis Stephensen
(Figs. 9C, 12, und 13)
ZZupZoops sZm%.s Stephensen 1925, p. 156, Fig.
46; Shoemaker 1931, p. 18.
Locality
records:
1) WHB sta. E No. 3; 39
50.5 N, 70 35 W; 823.5 m. Anchor dredge.
Atlantis cruise; 25 May 1961. 2) WHB sta. 87;
39 48.7 N, 70 40.8 W; 1,102 m. Epibenthic
sled. Chain cruise 50; 6 July 1965. 3 > WHB
sta. G No. 9; 39 44.7 N, 70 38.3 W; 2,021 m.
Anchor dredge. Atlantis cruise 284; 9 September
1962. 4) WHB sta. 103; 39, 43.6 N, 70 37.4
W; 2,022 m. Epibenthic
sled. Chain cruise 58;
4 May 1966. 5) WHB sta. G No. 11; 39 43.2
N, 70 40 W; 2,085 m. Anchor dredge. AtZunth
cruise 284; 10 September
1962. 6) WHB sta.
131; 39 38.5 N, 70 36.5: W to 39 39 N,
70 37.1 W; 2,178 m. Epibenthic
sled. Atlantis
II cruise 30; 18 December
1966. 7) WHB sta.
GH No. 3; 39 27.5 N, 70 33.5 W; 2,478 m.
Anchor dredge.
Atluntis cruise 273; 3 October,
1961. 8) WHB sta. 62; 39 26r N, 70 33 W;
2,496 m. Epibenthic
sled. Atlantis ZZ cruise 12;
21 August 1964. 9) WHB sta. 76; 39 38.3 N,
67 57.8 W; 2,862 m. Epibenthic
sled. Chain
cruise 50; 29 June 1965. 10) WHB
sta. 64;
38 46 N, 70 06 W; 2,886 m. Epibenthic
sled.
Atlantis ZZ cruise 12; 21 August 1964.
Range:
Probably
widely-distributed
around
the North Atlantic
Ocean, but known at present
only from the mouth of Davis Strait, 2,702 m
(Stephenscn
1925), three localities
southeast of
Nova Scotia on the Scotian Shelf, 104-238 m
(Shocmakcr
1931) and along the Gay HeadBermuda transect between 39 50 N, 70 35 W
and 38 46 N, 70 06 W at depths from about
800-2,900 m.

Diagnosis. A blind Haploops of relatively small size (size at breeding estimated as 8-9 mm) lacking setae on dorsal
terga. Second article of peduncle of first
antenna long (almost twice length of first
article).
Coxal plate 4 with a straight
anterior margin, sharp anterior corner and

DEEP-SEA

AMl?IIIPODS

381

jlmost straight lower margin. Carpus of

pereopod 5 with a long anterior margin,
so that anterior lobe is longer than postcrior one. Inner ramus of uropod 1 short,
% the length of the outer ramus. Urosome
carina gently rounded, not keeled, with
l-2 small setac posteriorly.
Remarks: Figure 12 supplements Stephensens (1925) figure and illustrates
a
much larger specimen than he had available. In many respects, II. similis rescmbles H. tubicolu. It differs, however, in its
relatively small size, lack of cornea1 lenses,
somcwhat longer second article of the
pcdunclc of the first antenna, shortened
inner ramus of uropod 1 (H. tubicolu =
II. spinosu is variable in this feature-see
Shoemakor 1931, p. 17) and particularly
in the shape of the fourth ooxal plate.
Haploops similis appears to be a scarce
but regular inhabitant Iof the continental
shelf and slope from the Arctic to areas
off the cast coast of the United States.
No doubt it will also be found off northdistriwestern Europe. Its bathymetric
bution, like that of I-I. setosu, is strongly
affected by water temperature, since approximately
the 5C isotherm is followed
beneath warmer surface waters in the
southern part of the range.
Changes in the shape and armature of
the distal part of the ramus of pereopold
5 with increasing size are shown in Fig.
13. Stephenscn (1925) and Kanncworff
(1966) illustrated the same 4.5 mm type
specimen from off West, Greenland at
2,702 m. It appears to be roughly cquivalent in morphology to a 4.0 mm animal
from the Gay Head-Bermuda transect (cf
Kanneworffs Figs. 7B and D with Figs.
13D, E, and E here),
The difference
could rcflcct a reduced frequency of molting at the northern station, but the causative factors are unknown, since tcmpclraturc at 2,000-3,000 m is likely to be about
the same off West Greenland and on the
transect.
The marked changes in shape and armature of percopod 5 shown in Figs. I3C,
D, I& and F and E and F illustrate the
difficulty of working with immature mate-

382

ERIC L. MILLS

9, WHB sta. 62; 39 26 N, 70 33 W;

FIG. 12. Hnploops similis Stephensen, 7.5 mm immature
G, H-gnatho-t&on;
2,496 m. A-IIead;
B-pleon
and urosome; C, D, E-right
.uropods 1-3; F
O-anterior
view of
of
maxilla
2;
pods l-2;
I, J, K, L, M-pereopods
1-5; N-posterior
view
maxilliped.

NORTII

ATLANTIC

DEEP-SEA

AMPHIPODS

383

FIG. 13. Huploops similis Stephensen. A-Right

mandible;
B-maxilla
1. Both 7.5 mm immature
9, WHB sta. 62. C-F-Merus
to dactyl of pereopod 5, showing changes with increasing size and
C-2.2
mm immature, WHB sta. 87; D-3.6
mm immature, WHB sta. G No. 9;
sexual dimorphism.
E-4.9
mm immature
8, WHB sta. 131; E4.6
mm immature
9, WHB sta. 103 (posterior plumose
sctae of merus omitted); F-7.0
mm immature
$, WHB sta. 193; F-7.2
mm immature
9, WHB
sta. 62.

rial of this species. Full breeding adults

can be expected to show even heavier
spination and broader limb articles than
F and F (7.0 mm immature 8 and 7.2
mm immature 5!) and although the sexual
dimorphism in the specimens illustrated is
slight, fully mature males will be markedly different in number of spines and
setation from females.
DISCUSSION

Table 2 lists ampcliscid species known

or cxpccted from the Gay Head-Bermuda
transect and adjacent areas to the north by
their depth preferences, as they are now
known. Examination of this list shows that
species of Ampelisca, in general, occur in
shallow waters, while #on the continental
slope species of Byblis and Huploops prc-

dominate.
Only B. profundi, apparently
an endemic species, occurs solely at abyssal depths on the transeot. Because of
the arrangement of species by their shallowest depth record in Table 2, the actual
relationship between occurrences of Haploops and Byblis is somewhat obscured.
Two species of Byblis, B. brachycephala
and B. medialis, are best considered midslope species, judging by their abundance
in collections, while, although they are
abundant nowhere, both H. setosu and H.
simih extend to abyssal depths. There is
thus a rough taxonomic (and probably
ecological) zonation of Ampcliscidae along
this transect, with Ampelisca in shallow
water, blind Byblis at midslope deplths,
Haploops at slope and abyssal depths, and
Bybhoides
abyssal only. This is also a
rough gradient of morphological
adapta-

384

ERIC

TABLE 2. Ampelticidae
recorded from the Northwest Atlantic
north of Cape Cod and the Gay
Head-Bermuda
transect

MILLS

mens have been collected from the carefully-worked

Gay Head-Bermuda
transect), and it is quite possible that members
of this genus will be found more widely
Expected
or recorded
depth
range
Literature
as rare but widespread specialists on fine
Species
citation
(m)
sediments
at both bathyal and abyssal
Amp&sea
ubdita*
Intertidal-30
Mills
1967a
depths.
Ampelisca vemilli*
Intertidal-50
Mills
19G7c
Ampclisca vndorum*
In gcncral, the abundance of AmpelisIntertidal-70
Mills
1963
Jntertidal-200
Byblis serrata
This
paper
cidae declines with depth and members
Ampelisca agassizi*
5(?)-450
Mills
1967c
and
of this family are ,far from significant
this paper
numerically
at lower slope and abyssal
Byblis gaimardi
5-575
This
paper
Ampelisca macrocephnla
Mills
1967c
lo-280
depths. This decline in abundance and
Ampelisca eschvichti
20->256
Stephenscn
itnportance parallels a similar decline of
1925,
Shoeother amphipod families and a relative inmaker
1931,
Dunbar
1954,
crease in the Isopoda and Tanaidacea in
Brunel
1956
the abyss. Whether the Amphipoda are
IInploops tubicoln
22-238(?)
Shoemaker
1931
(as H. spinosa)
just beginning an invasion of the deep sea
? (37-110
Shoemaker
1931
Ampelisca typica
$or are competitively excluded from it canin Ewo~x )
(Sars
1895)
not be decided conclusively.
The great
This
paper
and
Ampelisca nequico~ni.9
92-835
Shoemaker
1931
success
and
adaptive
radiation
of
the order
Ampeliscu dsclivitatis*
loo-1,100
Mills
19670
and
in shallow marine waters, along with the
this paper
This
paper
IIa~~loops similis*
fact that deep-sea Amphipoda are relaloo-2,900
This
paper
IIaploops setosa*
lOO-3,000
tively unspecialized morphologically
(J. L.
Shoemaker
1931
Ampelisca amblyops
295
Barnard
1961,
1964),
indicates
either
a
Shoemaker
1931
Ampelisca
gibba
295
recent colonization of the deep sea (reaThis
paper
Ampeliscn
uncinatn
350-1,100
sons unknown)
or actual exclusion of all
This
paper
Byblis hmchycephaln*
800-1,500
This
paper
Byblis medinli.9
1,300-2,000
but a relatively few generalized and pcrThis
paper
Byblisoides profun&*
4,680-4,977
haps eurytopic species.
* Species
found
on the Gay Head-Bermuda
or cxpccted
Ampeliscidae on the transect show an
transect.
interesting mosaic of zoogeographic relations. The abyssal B. profundi is unique
and probably endemic to the North Amertion to food available, since the bathyal
and abyssal Byblis and Haploops have ican Basin; both species of Haploops, at
long, setose antennae, which previous wlork bathyal and upper abyssal depths, range
(Enequist
1950) suggests may be an around the North Atlantic; and the two
adaptation for collection of suspended or species of bathyal BybZk share features
The acme of with species found farther north and in
settling organic matter.
adaptation to deep-sea life, as far as am- European waters. AmpeZisca aequicornis
and A. uncinata range at lower slope and
peliscids are concerned, is shown in Bybbathyal depths to Europe, while many
Zisoides, which has the second antennae
markedly shortened and provided with a species of Ampelisca known from closely
adjacent waters to the north and considremarkably heavy musculature for swceping in volumes of food-poor abyssal scdi- erable depths (e.g., A. eschrichti, A. macrocephala, H. tubicola, B. gaimurdi),
do
ments to the mouthparts for processing,
It is puzzling that elsewhore species not occur on the transect (Table 2). Two
spccics found from the shelE to slope-edge
of Byblisoides are known primarily from
bathyal depths. J, L. Barnard (1964) has or part way down the slope, A. agassixi
speculated that the genus is an abyssal and B. serrata, are unknown or very rare
north of the transect. In total, the transect
one that has been forced into shallower
may be envisioned as beginning in shallow
water by ecological #or physical changes.
waters
where the fauna has mostly VirByblisoicles is rare (only about 9 spcci-

NOIWII

ATLANTIC

ginian and Carolinian elements (e.g., A.

abdita, A. verrilli, A. vadorum),
passing
into a relatively warm-water area on the
shelf and at its edge where a mosaic of
warmer water southern and colder water
northern species co-occur (e.g., A. egassixi,
B. serrata, A. aequicornis, A. uncinuta),
and moving to slope and upper abyssal
depths where widely ranging stenothermal
species of the north Atlantic (H. setosa,
H. similis) or species (B. brachycephala,
B. medialis) closely related to those of
the northern and eas(tern Atlantic occur.
Although B. profundi is ;the only species
found solely at abyssal depths, this is
in accord with the conclusions of J. L,
Barnard (1961, 1966, 1967) that abyssal
amphipods show a high degree of endemism in basins and trenches-a
conclusion
strongly supported for abyssal faunas in
general by Vinogradova ( 1962).
There is certainly also justification
for
considering the continental slope a sort of
mixing ground for shallow water, abyssal,
and cndcmic slope species of varied geographic origin. So,me elements of the slope
fauna may show low latitude submergence.
For example, H. setosa has a minimum
depth of about 500 m on the transect,
whereas to the north it occurs to 100 m
an d even shallower depths occasiunally.
This is an isolated example, and there is
certainly no other justification
for considering deep-sea (including bathyal) faunas
to have strong affinities with the shallowwater faunas of polar latitudes, when the
distribution
of Ampeliscidae
around the
North Atlantic, in the Arctic, and on the
Gay Head-Bermuda
transect is used as
evidence. Rather, a picture emerges of
isolated deep-sea basins, with their endcmic stenotopic sp&es, ringed by cornplex bathyal zones, where a mosaic of
faunal elements exists, derived from other
latitudes, shallow water, and in situ spcciation. The bathyal depths, are,as of great
ecological and evolutionary
tension, dcscrvc incrcascd and critical attention.

385

AMFIIIPODS

Pacific Ocean by the Velero III nnrl Velcro

Allan
Hancock
Pac. Exped.
18( 1) :
IV.
1-137.
1961. Gammaridean
Amphipoda
from
-.
depths of 400 to 6000 meters.
Galathea
Rep. 5: 23-128, 83 Figs.
Deep-sea Amphipoda
(Crusta1964,
-.
tea) coI.lected by the R/V Vema
in the
eastern Pacific Ocean and the Caribbean and
Mediterranean
Seas. Bull. Amer. Mus. Natur. Hist. 127( 1) : 46 p.
1966. Submarine canyons of Southern
-.
California.
Part 5. Allan Hancock Pac. Exped. 27( 5) : 166 p.
-.
1967. Bathyal
and abyssal gammaridcan Amphipoda
of Cedros Trench, Baja CaliU.S. Nat. Mus. Bull. 260. 205 p.
fornia.
-.
1969. The families and genera of marine gammaridean
Amphipoda.
U.S. Nat.
Mus. Bull. 271. 535 p.
I~ARNARD, K. H.
1931. Diagnosis of new genera and species of amphipod Crustacea collected during the Discovery
Investigations,
1925-1927.
Armu. Mag. Natur. Hist. Ser.
10 7: 425-430.
-.
1932. Amphipoda.
Discovery
Rep. 5:
l-326.
BRUNEL, P. 1956. The bathymetric
distribution
of the benthic Amphipoda
(Crustacea:
MaIacostraca)
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