Palaeogeography, Palaeoclimatology, Palaeoecology, 45 ( 1984 ): 189--223 189

Elsevier Science Publishers B.V., Amsterdam -- Printed in The Netherlands

QUANTITATIVE ICHNOLOGY OF MODERN PELAGIC DEPOSITS IN

THE ABYSSAL ATLANTIC

A. A. EKDALE, L. N. MULLER 1 and M. T. NOVAK 2

Department of Geology and Geophysics, University of Utah, Salt Lake City, UT 84112
(U.S.A.)
(Received November 24, 1982)

ABSTRACT

Ekdale, A. A., Muller, L. N., and Novak, M. T., 1984. Quantitative ichnology of modern
pelagic deposits in the abyssal Atlantic. Palaeogeogr., Palaeoclimatol., Palaeoecol., 45 :
189--223.

Quantitative sedimentologic aspects of bioturbation were investigated in a series of

deep-sea box cores, which were collected over a broad latitudinal range (35N to 30S)
and bathymetric range (1.4 to 5.7 kin) in the central Atlantic Ocean. All cores were
thoroughly bioturbated, and numerous generations of burrowing could be discerned in
many instances. Preservation of individually distinct burrows ranged from poor to excel-
lent, and eight ichnogenera plus several unnamed biogenic structures were recognized.
The distribution of calcium carbonate, organic carbon, selected species of planktic
foraminifera and sediment grain size were examined in several cores. These measurements
indicated that the surficial Mixed Layer, which rhnged from 3 to 10 cm thick, was statisti-
cally more homogeneous than the underlying Transition Layer. This relationship is seen
most clearly with respect to lateral and vertical variations in the calcium carbonate con-
tent of the cores. Bioturbation of abyssal pelagic deposits appears to be a two-phase pro-
cess, consisting of (a) homogenization of the upper few centimeters (i.e., the Mixed
Layer) by shallow-burrowing meiofauna, and (b) heterogeneous mixing of the underlying
strata (i.e., the Transition Layer) by relatively deep burrowers. The heterogeneously
mixed Transition Layer becomes the preserved trace fossil record; under normal circum-
stances the homogenized Mixed Layer is not preserved.

INTRODUCTION

T h e l a r g e s t d e p o c e n t e r o f f i n e - g r a i n e d s e d i m e n t o n e a r t h is t h e a b y s s a l
o c e a n f l o o r , w h i c h is also t h e h a b i t a t o f t h e m o s t d i v e r s e m a c r o c o m m u n i t y
o f b e n t h i c life i n t h e w o r l d . P e l a g i c d e e p - s e a s e d i m e n t t y p i c a l l y c o n s i s t s o f
biogenic calcareous ooze, biogenic siliceous ooze a n d t e r r i g e n o u s " r e d " clay.
Understanding the processes of bioturbation and recognizing the charac-
t e r i s t i c p a t t e r n s o f b u r r o w i n g i n p e l a g i c d e p o s i t s are i m p o r t a n t f o r m a n y

i Present address: Marathon Off Company, P.O. Box 120, Casper, WY 82602 (U.S.A.).
2 Present address: Getty Oil Company, 345 Bearcat, Salt Lake City, UT 84115 (U.S.A.)

0031-0182/84/$03.00 1984 Elsevier Science Publishers B.V.

190

reasons. For example, mixing of the sediment column by organisms disrupts

the original stratigraphic record, so it is important for deep-sea stratigraphers
to understand the nature and magnitude of that mixing process (Berger and
Heath, 1968; Ruddiman and Glover, 1972; Guinasso and Schinck, 1975;
Peng et al., 1977, 1979; Wetzel, 1981). Moreover, textural properties of
sediment, such as grain size, sorting and porosity, as well as sediment shear
strength, may be altered significantly as a result of bioturbation (Rhoads,
1970; Howard, 1975; Ekdale and Berger, 1978; Wetzel, 1981). The chemical
and microfossil composition of the sediment may be altered also.
Because the preservation potential of most deep-sea animals as b o d y
fossils is rather low, trace fossils provide the most abundant paleontologic
information a b o u t ancient deep-sea benthic communities (Ekdale, 1977).
Organism traces reveal the activities of deep-sea creatures, including their
locomotion patterns, feeding habits and so on. Associations of various
tracks, trails and burrows in modern deep-sea deposits allow for the recogni-
tion of deep-sea ichnofacies in the geologic record.
This paper summarizes results of ichnologic studies of modern deep-sea
oozes and clays collected in the North and South Atlantic on two Scripps
Institute of Oceanography sediment-coring expeditions (INDOMED II and
XII) during 1977 and 1978.

CORE DESCRIPTIONS

Samples

Sediment samples were collected from the Atlantic Ocean b o t t o m be-

tween latitudes 35N and 30S (Fig.l) in water depths of 1.4 to 5.7 km
(Table I). The samples were recovered using a box corer, which retrieves a
relatively undisturbed cube of the sea floor measuring 50 X 50 cm in area
and up to 60 cm deep (Hessler and Jumars, 1974). The advantages of box
cores over more conventional (small-diameter) piston or gravity cores include
the virtual lack of sediment deformation along the core walls, the greater
width of sediment exposure in which to observe ichnologic and sedimento-
logic features and the relatively undisturbed sediment surface at the core
top.
The sediment cored for this investigation included aragonitic p t e r o p o d
ooze, calcitic foraminiferal and coccolith ooze and terrigenous clay contain-
ing varying amounts of siliceous plankton remains (Novak, 1980; Muller,
1982). All sediment appeared to be thoroughly bioturbated, although preser-
vation of individually distinct burrows ranged from poor to excellent.

Core stratigraphy

In most cases, especially for calcareous ooze, one could observe a general
core stratigraphy resembling that described in box cores from the abyssal
 19]

45 30 15 0 15

40" ~~ 0 ~
59 68 [0 13
T~Tz f
65 69 72

42 /
20 )4 J~,(~ ~c~ 20"

9" ic"
,T I ;04

0
IDC I(, O

, IX
20 20
23122 1"6~- 117
s z, ! 2]g ~ 1 2 0
40 _ .40 o

3,,0o J J i
4 15 0 15

Fig. 1. Map of the central Atlantic Ocean showing the locations of box-core sites investi-
gated for this study. For additional information (water depth, sediment type, etc.), see
Table I.

Pacific by Berger et al. (1979). Three major zones occur (Fig.2): a surficial
Mixed Layer, in which sediments appear to be homogenized by the burrow-
ing activities of rather tiny (meiofaunal) animals; a Transition Layer, in
which active bioturbation by the deep-burrowing infauna causes heterogene-
ous mixing of the sediment; the Historical Layer, into which no burrowers
penetrate, so no redistribution or alteration of sediment by organisms
OCCURS.
The Mixed Layer appears to be the most intensely bioturbated zone in
any core. Typically it is a uniform-colored layer capping the sediment
column. In most cores the color of this layer is darker than that of the
underlying sediment because of its higher clay content, due to increasing
dissolution of calcium carbonate occurring in sediment deposited on a sub-
siding oceanic plate.
Tiny open burrows (less than 0.5 mm in diameter), called Trichichnus,
may be abundant in the Mixed Layer; larger open burrows (3--10 mm in
diameter) are not u n c o m m o n (Fig.3). The tiny burrows, probably produced
by meiofaunal annelids and arthropods, apparently close up soon after they
are abandoned due to the low shear strength of the Mixed Layer sediment.
192

TABLE I

B o x c o r e site i n f o r m a t i o n

Core No, Depth Latitude Longitude Sediment type a %CaCO3a, b B u r r o w

(m) preservation

I n d o m e d cores (INMD), N o r t h and S o u t h A t l a n t i c

BX40 4385 28 2 1 ' N 47 29'W n a n n o - f o r a m clay 75.6 good

BX42 3775 28 3 4 ' N 46 2 2 ' W n a n n o - f o r a m clay 77.6 good
BX48 2838 29 4 9 ' N 43 1 3 ' W nanno-foram ooze 88.3 fair
BX50 3490 31 1 1 ' N 39 4 1 ' w n a n n o - f o r a m clay 82.4 good
BX59 3358 33 2 3 ' N 32 18'W n a n n o - f o r a m ooze 89.1 good
BX65 3443 34 3 7 ' N 29 5 9 ' W n a n n o - f o r a m ooze 91.6 good
BX68 2520 34 4 8 ' N 28 2 1 ' W n a n n o - f o r a m clay 84.1 good
BX69 4670 34 4 9 ' N 26 1 2 ' W n a n n o - f o r a m clay 79.5 good
BX70 5119 34 5 8 ' N 24 1 9 ' W n a n n o - f o r a m clay 74.6 optimal
BX72 5182 34 5 7 ' N 21 4 0 ' W n a n n o - f o r a m clay 68.8 optimal
BX73 4041 34 5 8 ' N 17 2 2 ' W n a n n o - f o r a m clay NM good
BX93 5744 16 5 7 ' N 54 5 2 ' W r e d clay 0.1 good
BX94 4391 16 4 3 ' N 51 3 5 ' W n a n n o - f o r a m clay 70.0 fair
BX97 3619 1639'N 4608'W n a n n o - f o r a m clay 73.3 fair
BX98 4742 15 3 2 ' N 42 5 7 ' W n a n n o - f o r a m clay 61.5 poor
BX100 5386 11 0 0 ' N 34 1 0 ' W r a d - d i a t o m clay 41.7 optimal
BX101 4093 6 57'N 26 2 7 ' W r a d - f o r a m clay 70.5 optimal
BX103 4492 5 5 3 ' N 24 2 8 ' W foram sand 64.9 optimal
BX104 3279 4 15'N 21 5 5 ' W foram sand 77.7 good
BX106 4702 2 4 4 ' N 20 5 6 ' W foram sand 69.8 optimal
BX109 3895 5 2 8 ' S 15 5 8 ' W foram ooze 91.4 fair
BXll0 1959 10 0 2 ' S 13 2 3 ' W pteropod ooze 96.0 fair
BXlll 3069 12 3 9 ' S 13 5 1 ' W p t e r o p o d ooze 94.0 fair
BXl13 3471 15 1 5 ' S 14 5 8 ' W n a n n o - f o r a m clay NM fair
BXll5 3427 17 3 8 ' S 16 1 3 ' W nanno-foram ooze 92.7 fair
BX116 4722 21 5 4 ' S 18 4 0 ' W red clay 26.0 optimal
BXll7 4398 24 0 5 ' S 19 5 0 ' W n a n n o - f o r a m clay 70.2 good
BX120 1918 28 5 0 ' S 31 1 7 ' W pteropod sand 82.5 fair
BX121 3908 28 5 9 ' S 32 0 1 ' W n a n n o - f o r a m clay 79.6 fair
BX122 3082 29 1 8 ' S 33 0 1 ' W p t e r o p o d ooze 91.3 fair
BX123 3361 39 31'S 33 1 4 ' W n a n n o - f o r a m ooze 89.3 fair
BX125 3535 29 4 4 ' S 33 02' W n a n n o - f o r a m ooze 88.5 fair
BX127 2509 29 5 3 ' S 33 5 9 ' W pteropod ooze 94.4 fair
BX128 1436 29 5 5 ' S 34 2 1 ' W pteropod ooze NM fair

Pleiades cores (PLDS), East E q u a t o r i a l Pacific

BX83 4531 0 5 7 ' N 126 ~ 3 8 ' w n a n n o - f o r a m clay 64.0 optimal

BX85 4385 0 58'N 128 23'W n a n n o - f o r a m clay 68.0 optimal
BX89 4406 0 5 8 ' N 13139'W n a n n o - f o r a m clay 70.3 optimal
BX107 4850 509'N 13817'W r a d - d i a t o m clay 34.5 optimal

aSamples at box core surface; bNM indicates no m e a s u r e m e n t .

 193

Fig.2. E x p o s e d vertical face of core INMD BX 71 with t h e t h r e e - l a y e r e d s t r a t i g r a p h y indi-

cated. Scale in c e n t i m e t e r s o n r i g h t - h a n d m a r g i n of p h o t o .

Fig. 3. Close-up views o f t h e M i x e d L a y e r in t w o box cores. A. Large, b r a n c h e d , o p e n

b u r r o w in M i x e d L a y e r c o m p o s e d o f r e d clay in c o r e INMD BX 1 1 6 ; smaller b u r r o w s are
p r e s e n t b u t n o t easily o b s e r v e d due t o t h e i r partial d e s t r u c t i o n in t h e l o w - s t r e n g t h sedi-
m e n t . Scale bar equals 2 cm. B. T i n y b u r r o w i n g a n n e l i d w o r m (a) in M i x e d L a y e r com-
p o s e d of f o r a m i n i f e r a l s a n d in core I N M D BX 106. Scale bar equals 2 cm.
194

The larger open burrows may extend down into the Transition Layer. The
thickness of the Mixed Layer in Atlantic box cores, determined visually,
ranged from about 3 cm in clay-rich deposits to as much as 10 cm in pure
calcareous ooze. These dimensions correspond favorably to Mixed Layer
thicknesses reported in Pacific deep-sea box cores by Berger et al. (1979).
Transition Layer sediments are stiffer than those of the overlying Mixed
Layer, and so burrows produced within this zone are more visually distinct,
in terms of both color contrast and sharpness of the burrow margins. Color
hues range from dark olive green to light yellowish green and from dark
reddish brown to light tan to white. Cross-cutting relationships of burrows
of different colors indicate that darker burrows are younger than the lighter
ones. Thus, burrow color fades with age.
White rinds or haloes surround many burrows in the Transition Layer and
also in the underlying Historical Layer. These bleached sites which ring the
burrows apparently form as reduced ferrous iron is removed in solution from
the immediate vicinity of the burrowed sediment, leaving the white carbon-
ate-rich sediment devoid of iron pigment. Such coloration patterns of
burrows have been referred to as "reduction burrows" in Deep Sea Drilling
Project (DSDP) cores (Ekdale, 1977).
Burrows in the Transition Layer are dominated by vertical and sub-
horizontal burrows, termed Skolithos and Planolites respectively, which
range from 3 to 10 mm in diameter and up to 20 cm long. The base of the
Transition Layer is the level below which active bioturbation cannot be dis-
cerned, and this is recognized visually as a gradational zone where burrow
definition and color contrast begin to deteriorate downwards in the core.
The lower limit of the Transition Layer is n o t as sharply defined as the horizon
at the b o t t o m of the Mixed Layer, but in the Atlantic box cores this level
varies from about 20 to 35 cm below the sediment surface.
The Historical Layer is the preserved record. It extends from the base of
the Transition Layer down to basement, and it corresponds to the trace
fossil record observed in DSDP cores and land-based outcrop sections. As
pelagic sediments accumulate, the upper boundaries of the Historical and
Transition Layers move upward more or less continuously.
At the top of the Historical Layer, biologic reworking of sediment ceases;
early diagenesis takes over as the major factor controlling the texture, com-
position and color of the sediment. Compaction of burrows begins to be
noticeable near the top of the Historical Layer, and sediment stiffness in-
creases. Coloration of the sediment fades, and burrow margins become less
distinct.

Surface features

In most cases, the tops of the box cores appeared to contain the original
microtopography and many surface features of the deep-sea floor. It was
apparent t h a t many core tops had suffered some degree of washing, however,
 195

as b o t t o m w a t e r t r a p p e d i n t h e u p p e r c o v e r e d p o r t i o n o f t h e b o x c o r e
s l o s h e d b a c k a n d f o r t h w h i l e t h e c o r e w a s b e i n g r e t r i e v e d f r o m t h e sea
b o t t o m . With very few e x c e p t i o n s , this effect was n o t c o n s i d e r e d to be
m a j o r . T h e s u r f a c e r e l i e f i n t h e A t l a n t i c b o x c o r e s v a r i e d as m u c h as s e v e r a l
c e n t i m e t e r s , due t o the p r e s e n c e of m o u n d s , l u m p s a n d trails of u n d e t e r -
m i n e d origin (Fig.4).
M o s t c o r e s u r f a c e s c o n t a i n e d o p e n h o l e s , s o m e o f w h i c h a p p e a r e d t o be
p a i r e d ( F i g . 5 A ) . T h e l o w s t r e n g t h o f t h e M i x e d L a y e r p r e v e n t e d e a s y dissec-
t i o n of a n y h o l e s t o d e t e r m i n e t h e i r s u b s u r f a c e g e o m e t r y , h o w e v e r . S i n g l e
holes ranged from 1 to 5 mm in diameter and occurred in abundances
a p p r o a c h i n g 2 0 h o l e s p e r 1 0 0 c m 2 in s o m e c o r e s . C o m m o n l y , b u t n o t i n v a r i -
a b l y , s u c h h o l e s w e r e s i t u a t e d at t h e c e n t e r o f l o w c o n i c a l m o u n d s , r a n g i n g
f r o m 5 t o 10 c m i n d i a m e t e r a n d 1 t o 2 c m h i g h . T h e m a t e r i a l c o m p o s i n g
some mounds differs from the surrounding surface sediment and more

Fig.4. Surfaces of two box cores, providing a glimpse of the microtopography of the
deep-sea floor. A. Undulating microtopography of lumpy foraminiferal sand distributed
in irregular mounds and depressions on the surface of core INMD BX 103. Scale bar
equals 5 cm. B. Lumpy radiolarian--diatom clay and several clay-lined tubes (unoccupied)
on the surface of core INMD BX 100. Scale bar equals 5 cm.

Fig. 5. Close-up views of surface features in two box cores. A. Open holes, possibly paired
openings of U-shaped burrows, in calcareous nannofossil--foraminiferal ooze at the sur-
face of core INMD BX 123. Scale bar equals 2 cm. B. Rounded conical m o u n d of tan cal-
careous nannofossil--foraminiferal ooze sitting on gray foraminiferal sand at the surface
of core INMD BX 104; the tan ooze apparently derives from the Transition Layer at this
site and was brought to the surface by the process of bioturbation. Scale bar equals 2 cm.
196

closely resembles that of the Transition Layer in color and t e x t u r e (Fig.5B).

This suggests that, in addition, to d o w n w a r d mixing o f sediment by passive
infilling of o p e n burrows, upward mixing by bioturbation also may be
c o m m o n . In one core (INMD BX 117) from 4398 m of water in the South
Atlantic, a large o p e n hole (about 6 mm in diameter) was accompanied by
numerous tiny grooves radiating o u t w a r d from the center, as if p r o d u c e d by
a worm extending tentacles or a proboscis across the sediment surface during
feeding. No animal was discovered in the burrow.
Five sets of paired open holes, a b o u t 2 mm in diameter and spaced 2--3
cm apart, o c c u r r e d on the surface of one core (INMD BX 50) from the
North Atlantic. In addition, one pair of holes, a b o u t 4 mm in diameter and
spaced 3 cm apart, occurred on the surface of a South Atlantic core (INMD
BX 123). Injection of water in one hole of the pair d e m o n s t r a t e d that the
two openings were c o n n e c t e d , but the precise g e o m e t r y of the connection
could not be ascertained. However, in X-ray radiographs of thin, vertical
slabs sampled from several box cores (INMD BS 48, 50, 121 and 127), the
concave-up outlines of curved cylindrical burrows are evident, suggesting
t h a t U-shaped burrows may be present.
Macroscopic metazoans were rare in the surface sediments of the box
cores. T h e y included only two solitary ahermatypic coral specimens, which
obviously were n o t capable of producing biogenic sedimentary structures.
Large agglutinated colonies of x e n o p h y o p h o r i d protists occasionally occurred
on core tops. Benthic micro-organisms f o u n d alive in surficial sediment of
the cores included agglutinated foraminifera and very tiny (less than 0.5 mm
in diameter) p o l y c h a e t e annelids, none of which appeared to be responsible
for any of the macroscopic trails or burrows observed in the cores, with the
possible exception of T r i c h i c h n u s .
Thin strings of sediment which p r o b a b l y represent fecal material were
c o m m o n on box core surfaces. Some of these fecal strings were r o u n d in
cross-section and others were flattened, but typically t h e y were less than
1 mm wide and up to 7 cm long.
Various t y p e s of e m p t y biogenic tubes were f o u n d on the surfaces of
several cores (Fig.6). Some tubes (in cores INMD BX 68, 72, 100, 103 and
104) were hollow and consisted of agglutinated walls of varying width,

Fig.6. Unoccupied agglutinated tubes from the surface sediment of two box cores. A.
Thick-walled tube of agglutinated clay in core INMD BX 100. Scale bar equals 2 cm. B.
Tubes preferentially lined with size-sorted foraminiferal tests, mostly Globorotal& trun-
catulinoides, in core INMD BX 69. Scale bar equals 0.5 cm.
 197

containing m u d and microfossil tests in similar proportions as the adjacent

sediment in the core. However, tubes from INMD BX 69, which measured
7--9 mm in diameter and up to 25 mm long, were composed entirely of
whole tests of large planktic foraminifera (e.g., Globorotalia truncatulin-
oides), indicating preferential selection of rather large particles by the tube-
builders. Tubes such as these resemble thickly lined burrows in the fossil
record that have been reported as Palaeophycus heberti (Pemberton and
Frey, 1982).
Simple, irregularly meandering trails c o m m o n l y were preserved in concave
epirelief on box core surfaces. They were 2--4 mm wide and extended to
lengths of several centimeters. A sinuous, branched trail occurred in one core
(INMD BX 106); it was 2 mm wide, and the longest branch was 5 cm long.
In two cores from the South Atlantic (INMD BX 113 and 115) the sur-
faces contained a total of four prominent arcuate trails preserved in convex
epirelief. The trails were smooth and unsculptured, although one possessed a
slight marginal furrow along each side; none displayed any discoloration of
the sediment. They measured 5 mm in height, 25--35 mm in width and as
much as 68 cm in length. These trails closely resemble the plowing traces of
echinoids that have been observed in numerous deep-sea b o t t o m photo-
graphs by several workers (e.g., see Heezen and Hollister, 1971; Ekdale and
Berger, 1978).
Perhaps the most exciting ichnological discoveries in the surface sediments
were those of modern graphoglyptid burrows preserved on a number of box
core tops (Figs.7 and 8). The graphoglyptids included several specimens each
of Cosrnohaphe, Paleodictyon and Spirorhaphe. Their discovery in the cores
was reported by Ekdale (1980a); their descriptions are summarized below.

Fig. 7. S e l e c t e d i c h n o g e n e r a . A. T h e h o r i z o n t a l , m e a n d e r i n g , g r a p h o g l y p t i d trail Cosmor-

haphe in s e l e c t e d n a n n o f o s s i l - - f o r a m i n i f e r a l clay at the surface of core INMD B X 98.
Scale bar e q u a l s 1 cm. B. Large (left) a n d small (right) s p e c i m e n s of t h e mesh-like,
g r a p h o g l y p t i d b u r r o w Paleodictyon in p t e r o p o d - r i c h calcareous o o z e at t h e surface o f
core INMD B X 128. Scale bar e q u a l s 0.5 cm.
198

Fig. 8. Selected ichnogenera. A. The double-spiralling, graphoglyptid trail Spirorhaphe in

calcareous nannofossil--foraminiferal ooze at the surface of core INMD BX 94. Scale bar
equals 1 cm. B. Vertical face of foraminifera-rich sediment in the Transition Layer of core
INMD BX 106, containing tiny Chondrites (e), Planolites (19)and Skolithos (s). Scale bar
equals 2 cm.

ICHNOLOGY

Systema tic Ichnology

Note: Although it is usual practice in ichnology to identify modern trails and burrows in
terms of their creators instead of assigning them to ichnotaxa, the producers of the traces
reported here are unknown. Because depositional rates are very low in the abyss, it is diffi-
cult to determine precisely how " m o d e r n " these deep-sea traces actually are. Thus, in this
paper we employ ichnogenus names for the following biogenic sedimentary structures which
conform to established ichnotax& Abbreviated diagnoses are summarized from H~intzschel
(1975); partial synonymies include other reported occurrences in deep~sea cores. The
distribution of ichnogenera and other biogenic structures is reported in Table II.

Chondrites S t e r n b e r g 1 8 3 3

Chondrites Sternberg 1833. Van der Lingen, 1973, figs.15--18; Warme et al., 1973, pls.l-A
to K; Chamberlain, 1975, text-figs.2, 3, pls.l-4 to 7, 3-1; Kennedy, 1975, fig.17.6;
Kauffman, 1976, pls.2--10; Ekdaie, 1977, p l . l - A to C; Chamberlain, 1978, f i g s . l l - - 1 3 ,
84, 116; Ekdale, 1978, pl.l-1, 2, 4, 5; Ekdale and Berger, 1978, fig.9; Harrington,
1978, pp. 933--934; Berger et al., 1979, fig.13-A; Ekdale, 1980b, plsl-4 to 6 (photo-
graphs 4, 5, and 6 mislabelled by printer as 6, 4 and 3, respectively); Wetzel, 1981,
figs.2, 3, 24, pl.9.4.F; McMillen and Lundberg, 1982, pp. 654--655.

Diagnosis: H i g h l y b r a n c h e d s y s t e m o f c y l i n d r i c a l t u n n e l s o f u n i f o r m d i a m -
e t e r , w h i c h t y p i c a l l y d o n o t c r o s s e a c h o t h e r o r a n a s t o m o s e (see H ~ i n t z s c h e l ,
1975, pp. W49--52). ?Cambrian; Ordovician--Holocene.

Remarks: A l t h o u g h v e r y c o m m o n in w o r l d w i d e D S D P c o r e s ( C h a m b e r l a i n ,
1 9 7 5 ; E k d a l e , 1 9 7 7 ) , Chondrites is n o t o f t e n o b s e r v e d in d e e p - s e a b o x c o r e s .
T h i s m a y b e b e c a u s e Chondrites-producing a n i m a l s a r e n o l o n g e r a b u n d a n t
 199

TABLE II

Biogenic structures in the 34 Atlantic box cores

Structure No. of cores Depth range Latitude range %CaCO~ range

(m)

Ch ondrites 1 4702 3 N 70
Cosmorhaphe 2 (+4?) 3427--4742 16 N--18 S 62--93
Paleodic tyon 2 1436--3895 5S--30S 89--100
Planolites 32 (+2?) 1436--5744 35 N--30 S 0--96
Skolithos 15 (+4?) 3069--5182 35 N--30 S 26--94
Spirorhaphe 7 3358--4702 33 N--30 S 65--91
Trichichnus (occurrences not recorded)
Zoophycos 2(+1?) 4093--4492 7 N--6 N 65-~71
Foram-walled tubes 4 2520--4670 35 N--4 N 70--84
Mu d-walled tubes 7 2520--5386 35N--4 N 42-~92
Open holes 22 2520--5744 35 N--30S 0--93
Surface mounds 15 2520--5182 35N--30 S 65--93
Surface trails 8 3427--5744 28 N--18 S 0--93
? U-shaped burrows 5 2509--3908 31 N--34 S 80--94

or, m o r e l i k e l y , b e c a u s e t h e b u r r o w s are p o o r l y visible d u e t o t h e i r small size

a n d c o m m o n l y f a i n t c o l o r a t i o n . C h o n d r i t e s was seen in o n l y o n e o f t h e
A t l a n t i c b o x c o r e s ( I N M D BX 1 0 6 ) , w h i c h c o n t a i n e d several small s p e c i m e n s
(Fig.8B).

Cosmorhaphe Fuchs 1895

Cosmorhaphe Fuchs 1895.

Cosmoraphe sinuosa (lapsus calami). Ekdale, 1980a, fig. l-C.

Diagnosis: R e g u l a r l y m e a n d e r i n g h o r i z o n t a l trail c o n s i s t i n g of f i r s t - o r d e r a n d
s e c o n d - o r d e r l o o s e m e a n d e r s (see H E n t z s c h e l , 1 9 7 5 , p. W53). ? O r d o v i c i a n ;
Silurian--Holocene.

Remarks: C o s m o r h a p h e is a " g r a p h o g l y p t i d " b u r r o w ( S e i l a c h e r , 1 9 7 7 ) ,

which includes m a n y of the g eo m et ri cal l y organized feeding a n d / o r dwelling
structures that typify Seilacher's (1967) Nereites Ichnofacies. Portions of
C o s m o r h a p h e , r e p r e s e n t e d as d i s t i n c t l y i n c i s e d s i n u s o i d a l g r o o v e s , are f a i r l y
c o m m o n on b o x c o r e surfaces. O n e A t l a n t i c c o r e ( I N M D BX 9 8 ) c o n t a i n e d a
nearly complete specimen consisting of both first-order and second-order
m e a n d e r s ( F i g . 7 A ) . T h e t r a i l s a r e t h o u g h t to be c o n s t r u c t e d a n d m a i n t a i n e d
as o p e n b u r r o w s j u s t a f e w m i l l i m e t e r s b e n e a t h t h e w a t e r - - s e d i m e n t i n t e r f a c e
(Ekdale, 1980a).
200

Paleodictyon Meneghini 1850

Paleodictyon Meneghini 1850. Ekdale, 1980a, fig.l-D.

Diagnosis: Mesh-like system of horizontal tunnels forming regular hexagonal

polygons (see H~ntzschel, 1975, pp. W89--91). Ordovician--Holocene.

Remarks: Paleodictyon is an easily recognized graphoglyptid burrow because

of its distinctive hexagonal net pattern, but it is u n c o m m o n in box cores of
recent sediment. Two Atlantic box cores (INMD BX 109 and BX 128)con-
rained Paleodictyon specimens of various sizes, with polygons ranging in
width from 2 to 5 mm (Fig.7B). As with Cosmorhaphe, Paleodictyon was
seen only on slightly washed core tops, and the trails forming the net were
sharply incised 1 or 2 mm into the sediment (Ekdale, 1980a).
Worthy of n o t e here is the recent interpretation of branched forms of
m o d e r n xenophyophores, which are agglutinated benthic protists f o u n d only
in the deep sea, as possible producers of Paleodictyon-like structures in the
sediment (Swinbanks, 1982). Although some forms do branch to form poly-
gons within the size range of Paleodictyon observed in the two box cores,
neither regular hexagons nor a complete net have yet been described among
the xenophyophores.

Planolites Nicholson 1873

Planolites Nicholson 1873. Warme et al., 1973, pls.2-A to F; Kennedy, 1975, fig.17.6;
Kauffman, 1976, pp. 512--513; Ekdale, 1977, pls.2-A, B; Chamberlain, 1978, figs.40,
41; Ekdale, 1978, pls.l--3, 6, 7; Ekdale and Berger, 1978, figs.6--8; Berger et al., 1979,
fig.13-B; Ekdale, 1980b, figs.4--8 (photographs 4, 5, 6, 7 and 8 mislabelled by printer
as 6, 3, 4, 8 and 7, respectively); Wetzel, 1981, figs.8, 24, pls.9.4B--F; McMillen and
Lundberg, 1982, pp. 654--655.

Diagnosis: Unlined, cylindrical or subcylindrical, infilled burrows, straight to

gently curved, usually non-branching, usually more or less horizontal or
slightly oblique to bedding planes, may cross one another (H~intzschel, 1975,
p. W95, see also pp. W95--97; Pemberton and Frey, 1982, p. 865). Precam-
brian--Holocene.

Remarks: Simple horizontal and sub-horizontal burrows, ranging from a few

millimeters to more than a centimeter in diameter, occur in almost every box
core (Fig.8B). These burrows are Planolites, because they are unlined and
typically are unbranched, u n o r n a m e n t e d and unoriented. Due to their
straight or gently curved geometry and their lack of transverse annuli, they
should be referred to Planolites cf. P. beverlyensis (Pemberton and Frey,
1982). Planolites is c o m m o n throughout the entire bathymetric and litho-
logic range represented by the Atlantic box cores, and in fact it is the single
most abundant type observed in the cores.
 201

Skolithos H a l d e m a n 1 8 4 0

Sholithos Haldeman 1840. Ekdale and Berger, 1978, fig.6; Berger et al., 1979, fig.13-B;
Ekdale, 1980b, p. 602.

Diagnosis: " S t r a i g h t t u b e s or pipes p e r p e n d i c u l a r t o b e d d i n g and parallel to

each o t h e r , subcylindrical, u n b r a n c h e d " (H~intzschel, 1975, pp. W 1 0 6 - - 1 0 7 ,
see also pp. W 1 0 6 - - 1 0 8 ) . P r e c a m b r i a n - - H o l o c e n e .

Remarks: Straight, u n b r a n c h e d , vertical b u r r o w s are c o m m o n in c a r b o n a t e -

rich deep-sea sediments, and such b u r r o w s are r e f e r r e d to Skolithos (Fig.8B).
The p r o b a b i l i t y of intersecting a vertical b u r r o w in a core t a k e n parallel to it
o b v i o u s l y is n o t as high as t h a t o f intersecting a h o r i z o n t a l b u r r o w , so visible
Skolithos rarely o u t n u m b e r Planolites in e x p o s e d vertical faces o f b o x cores,
even t h o u g h t h e y m a y be p r e s e n t in similar a b u n d a n c e s .
It appears t h a t Skolithos originates as o p e n shafts which fill in with sedi-
merit after being vacated. Thus, d o w n w a r d m i x i n g o f s e d i m e n t is a c o m m o n
o c c u r r e n c e in the deep-sea e n v i r o n m e n t . The a b u n d a n c e o f Skolithos seems
to be a f u n c t i o n o f the shear strength of the s e d i m e n t , which is a f u n c t i o n o f
its calcium c a r b o n a t e c o n t e n t , which in t u r n is a f u n c t i o n o f b a t h y m e t r y
relative t o the calcite c o m p e n s a t i o n d e p t h (Ekdale and Berger, 1978). There-
fore, the n u m b e r s o f Skolithos are highest in cores c o m p o s e d o f m o r e t h a n
70% CaCO3 (i.e., in less t h a n 4 7 0 0 m of w a t e r ) and a p p r o a c h zero as the car-
b o n a t e c o n t e n t decreases with w a t e r d e p t h ; v e r y few Skolithos specimens
have been observed in clay-rich deposits cQllected f r o m m o r e t h a n 5 0 0 0 m of
w a t e r in the Atlantic.

Spirorhaphe F u c h s 1895

Spirorhaphe Fuchs 1895.

Spiroraphe (lapsus calami). Ekdale and Berger, 1978, fig.10; Berger et al., 1979, fig.13-D.
Spiroraphe involuta (lapsus ealami). Ekdale, 1980a, figs.l-A, B.

Diagnosis: Tightly spiralled h o r i z o n t a l trail which coils inward, t h e n loops

back and coils o u t w a r d alongside the inward-coiling trail (H~intzschel, 1975,
pp. 1 0 8 - - 1 0 9 ) . C r e t a e e o u s - - H o l o e e n e .

Remarks: T h e distinctive d o u b l e spiral of Spirorhaphe is fairly c o m m o n on

box core surfaces (Fig.8A). It is the m o s t a b u n d a n t g r a p h o g l y p t i d b u r r o w in
deep-sea box cores, and it o c c u r r e d in several Atlantic cores t a k e n f r o m 3 3 0 0
to 4 7 0 0 m of water. As with the o t h e r graphoglyptids, the coiled trail of
Spirorhaphe o c c u r s in distinct concave epirelief o n slightly w a s h e d core tops,
incised a few millimeters i n t o the sediment. It is t h o u g h t to have been an
o p e n t u n n e l s y s t e m situated just b e n e a t h the s e d i m e n t surface (Ekdale,
1 980a).
202

Trichichnus F r e y 1 9 7 0

Trichichnus Frey 1970. Frey, 1975, fig.2.8c; Bellaiche and Blanpied, 1979, pl.40; Wetzel,
1981, fig.13, pl.9.4A.

Diagnosis: Tiny, thread-like, cylindrical b u r r o w , straight or slightly curved,

b r a n c h e d or u n b r a n c h e d , vertical or subvertical (H~intzschel, 1975, p. W118).
Upper Cretaceous--Holocene.

Remarks: The M i x e d L a y e r in m a n y box cores, including b o t h c a r b o n a t e

o o z e and red clay, o f t e n contains n u m e r o u s tiny o p e n b u r r o w s which are
r e f e r r e d t o Trichichnus. T h e burrows generally are m u c h less t h a n 1 m m in
diameter, b u t t h e y m a y be several c e n t i m e t e r s long. T h e y are usually slightly
curved; branches o c c u r b u t are rare. M e i o f a u n a l annelids ( a n d / o r o t h e r meio-
faunal w o r m s and a r t h r o p o d s ) have been c o l l e c t e d f r o m M i x e d L a y e r sedi-
m e n t s by sieving and m a y be responsible for c o n s t r u c t i n g these tiny burrows.
Such w o r m s t y p i c a l l y are 0.5 m m or less in d i a m e t e r (Fig.3B).
Trichichnus has n o t been o b s e r v e d in T r a n s i t i o n L a y e r or Historical L a y e r
deposits, p r o b a b l y because the low shear strength o f the M i x e d L a y e r causes
it t o collapse as soon as b u r r o w s are a b a n d o n e d . Due t o their p o o r preserva-
tion and low visibility, the specific o c c u r r e n c e s of Trichichnus in the box
cores were n o t r e c o r d e d with c o n f i d e n c e and so are n o t r e p o r t e d in Table II.

Zoophycos Massalongo 1855

Zoophycos Massalongo 1855. Seilacher, 1967, pt.l-E; Piper and Schrader, 1973, pl.l-1 to
4; Reineck, 1973, figs.7--9; Van der Lingen, 1973, figs.3, 5--10; Warme et al., 1973,
pls.4-A to C, 5-A to G; Chamberlain, 1975, text-figs.2, 7, pls.l-2, 2-2, 5, 3-1, 3, 5, 6;
Kennedy, 1975, figs.17.6, 17.7; Ekdale, 1977, pls.3-A to E; Chamberlain, 1978,
figs.67--69, 116; Ekdale, 1978, pl.2-1 to 7; Ekdale a~d Berger, 1978, fig.1 ; Harrington,
1978, p. 934; Ekdale, 1980b, pl.l-4 to 6 (photographs 4, 5 and 6 mislabelled by printer
as 6, 3 and 4, respectively); Wetzel, 1981, figs.14, 15, 17, 19, 24, pl.9.4. E--G; Wetzel
and Werner, 1981, figs.l, 2, 5--11, 13; McMillen and Lundberg, 1982, pp. 654--655,
pl.l-1.

Diagnosis: B u r r o w s y s t e m consisting of flat or helically coiled spreiten

f o r m e d by tightly spaced, arcuate tunnels radiating f r o m a central axis which
is p e r p e n d i c u l a r t o bedding (see H~intzschel, 1975, pp. W 1 2 0 - - 1 2 2 ) . Ordo-
vician--Holocene.

Remarks: Zoophycos is easily r e c o g n i z e d in deep-sea cores, because its

spreite in vertical cross-section appears as a string o f crescentic backfill
structures. In m a n y cases a stack o f parallel spreiten can be seen, and some-
t i m e s the central axis o f the b u r r o w system itself is dissected (e.g., see
Warme et al., 1973; Ekdale, 1977). Zoophycos o c c u r r e d in several of the
Atlantic b o x cores, b u t no stack or coil of multiple spreiten was observed;
 203

each spreite was strictly flat and solitary. All the Zoophycos s p e c i m e n s in
the A t l a n t i c cores were pelleted, suggesting t h a t t h e y were s t u f f e d with fecal
material. T h e b u r r o w s were o b s e r v e d in b o t h cross-section a n d plan view,
and t h e m o r p h o l o g y of the spreiten i n d i c a t e d the p r e s e n c e of p r i m a r y , or
major, lamellae (i.e., radiating a r c u a t e tunnels) b u t no s e c o n d a r y , or m i n o r ,
lamellae joining the p r i m a r y lamellae, as have b e e n o b s e r v e d in m a n y Zoo-
phycos s p e c i m e n s in l a n d - b a s e d sections ( S i m p s o n , 1970).

Burrow preservation

T h e p r e s e r v a t i o n a l s t a t e o f b u r r o w s as t h e y a p p e a r e d in vertical cross-
section in the b o x cores was c o d e d as follows: " p o o r " if a l m o s t no b i o t u r b a -
tion was evident; " f a i r " if b u r r o w s a n d / o r trails were evident, b u t their
outlines were difficult to trace and their d e n s i t y was v e r y low; " g o o d " if
b u r r o w s were e v i d e n t a n d m o s t of the outlines c o u l d be traced; " o p t i m a l "
if b u r r o w s c o n t r a s t e d vividly with the m a t r i x s e d i m e n t , if outlines c o u l d be
t r a c e d c o m p l e t e l y , and if c r o s s - c u t t i n g of b u r r o w s c o u l d be observed. T h e
r a n k i n g of each b o x c o r e with r e s p e c t to b u r r o w p r e s e r v a t i o n is s h o w n in
T a b l e I.
T h e b o x cores with o p t i m a l b u r r o w p r e s e r v a t i o n include I N M D BX 100,
101, 103, 106 and 116. All these cores are either relatively low in c a r b o n a t e ,
or else t h e y have m u c h greater c a r b o n a t e values at core surfaces in H o l o c e n e
s e d i m e n t s t h a n in o l d e r glacial s e d i m e n t s . Core I N M D BX 100 is f r o m the
eastern slope o f the M i d - A t l a n t i c Ridge in the G a m b i a A b y s s a l Plain; c o r e s
I N M D BX 101, 103 and 106 are f r o m ' t h e Sierra L e o n e Rise; c o r e I N M D
BX 116 is f r o m the w e s t e r n slope o f the M i d - A t l a n t i c Ridge in the Brazil
Basin.
Cores with g o o d b u r r o w p r e s e r v a t i o n include I N M D BX 93, 104 and 117.
Core I N M D B X l 1 7 has a r e l a t i v e l y l o w c a r b o n a t e p e r c e n t a g e (70%), a n d
I N M D BX 93 has a l m o s t no c a r b o n a t e (0.08%). B o t h are f r o m d e e p w a t e r
( 4 3 9 8 a n d 5 7 0 8 m , r e s p e c t i v e l y ) , b u t core I N M D BX 1 0 4 was r e c o v e r e d
f r o m a b o v e the c a r b o n a t e c o m p e n s a t i o n d e p t h (CCD) ( 3 2 9 6 m ) a n d has a
higher c a r b o n a t e p e r c e n t a g e t h a n all o t h e r Sierra L e o n e Rise cores. T h e
c o n t r a s t b e t w e e n t h e coarse f r a c t i o n of H o l o c e n e and P l e i s t o c e n e s e d i m e n t s
in I N M D BX 104 is n o t as great as in the o t h e r Sierra L e o n e Rise cores.
All the cores f r o m the Sierra L e o n e Rise have either g o o d or o p t i m a l pre-
servation of b u r r o w s , t t o b a r t et al. ( 1 9 7 5 ) s h o w e d t h a t b o t t o m c u r r e n t s flow
a l o n g the K a n e G a p on the Sierra L e o n e Rise, and this d e e p - w a t e r c i r c u l a t i o n
p r o b a b l y reflects the e f f e c t s o n e a s t e r n basin c i r c u l a t i o n o f the crossing of
A n t a r c t i c B o t t o m Water t h r o u g h t h e M i d - A t l a n t i c Ridge at either the R e -
m a n c h e or V e m a F r a c t u r e Z o n e . A l t h o u g h the surfaces o f Sierra L e o n e Rise
b o x cores are rich in f o r a m i n i f e r a l tests, t h e y are b a r r e n in c o c c o l i t h s (S. A.
Walker, oral c o m m u n . , 1980), w h i c h m a y i n d i c a t e t h a t the s e d i m e n t has
b e e n w i n n o w e d . In a n y case, the core surfaces are radically d i f f e r e n t f r o m
c o r e b o t t o m s in t e r m s of s e d i m e n t t y p e . T h e t o p 20 c m or so consists o f
204

coarse-grained foraminiferal sand; the b o t t o m portion of the cores consists

of fine-grained red clay. The upper boundary of this clay may represent the
end of a glacial dissolution phase in the Atlantic. In these cores, burrows are
most visible below the carbonate-rich surface layer, i.e., where burrows have
been excavated in carbonate-poor, finer-grained sediment and then filled in
from above by foraminifera-rich, coarse-grained sediment.
The Sierra Leone Rise cores exemplify the "historical m o d e l " of box core
stratigraphy proposed by Berger et al. (1979). Because of either decreased
dissolution or winnowing out of the fine fraction, these cores consist of red
clay overlain by foraminifera-rich sand. The transition between the two sedi-
m e n t types corresponds to a time during which sedimentation and fertility
were changing in the Atlantic. Increased fertility during glacial time, due to
increased circulation and upwelling, possibly caused a decrease in quality of
burrow preservation, because a greater intensity of bioturbation may have
allowed homogenization of the sediment. Burrows are preserved in this inter-
mediate zone, not because of increased bioturbation, but rather in spite of it.
The preserved burrows are mostly those which were made in carbonate-poor
clay and, when abandoned, were passively filled with foraminiferal sand.
Shallow-water cores containing deposits which have always been above the
CCD, as well as deep-water cores which have always been below it, can best
be considered using the "steady state m o d e l " of burrow preservation (Berger
et al., 1979). In this model it is important t h a t the burrower makes the
burrow below the Mixed Layer, so that it will n o t be destroyed by subse-
quent bioturbation. Since sediments in and below the Mixed Layer in these
cores have nearly the same color and texture, the organisms that alter the
sediment in some way, either by ingesting the sediment or by stuffing
burrows with organic-rich fecal pellets have a better chance of leaving a
record in the homogeneous sediments than do organisms whose burrow will
be passively infilled by overlying sediment.
Shallow water cores usually represent a high sedimentation rate which is
not balanced by dissolution; therefore, the sediment accumulation rate is
high, and the carbonate percentage also is high. Pigment-producing metal
oxides are diluted by white carbonate, and color differences between burrow
and matrix are slight or non-existent. Burrow preservation is minimal in
these cores.
Deeper-water cores appear to have better burrow preservation, because the
removal of carbonate by dissolution concentrates the metal oxides and
sulfides which give the sediment its most vivid colors. Thus, slight differences
in burrow fill and matrix sediment are highlighted. Burrows preserved in low-
carbonate cores probably are actively filled burrows in sediment where the
original organic carbon was low, so that reduction of metallic cations was
lessened, and the pigment-producing oxides stayed in the sediment where
t h e y were deposited and did n o t migrate.
 205

QUANTVI'ATIVE SEDIMENTOLOGIC ASPECTS

Effects of bioturbation on sediment composition and texture

General. Biologic mixing of deep-sea sediment not only obscures its original
stratigraphy and primary stratification, but burrowing also alters such sedi-
ment properties as composition and texture. In order to determine these
effects of bioturbation, we examined the distributions of calcium carbonate,
organic carbon, selected species of planktic foraminifera and sediment grain
size in several box cores. Samples from various levels in the cores were ana-
lyzed to determine the extent of compositional and textural variation, both
vertically and laterally, that might be caused by bioturbation. In addition,
samples from selected burrows were analyzed to observe the compositional
and textural differences between burrow-fill material and the surrounding
sediment.

Calcium carbonate. The CaCO3 content of deep-marine sediment has been

studied by many workers as an aid in paleoecologic interpretation and pale-
oceanographic reconstruction of ancient oceans. The major factors influen-
cing the a m o u n t of CaCO3 in oceanic deposits include (1) productivity/
fertility of calcareous plankton (coccoliths, foraminifera and pteropods) in
surface waters, (2) dilution by non-carbonate material, such as pelagic and
hemipelagic clay, and (3) dissolution of CaCO~ at the sea floor, which
increases with bathymetry.
For this study, the CaCO3 distribution in selected box cores was analyzed
by means of a LECO WR-12 carbon determinator. Sediment samples were
dried, pulverized and split evenly into two duplicate subsamples weighing
approximately 0.2 grams each. One subsample was baked at 500C to burn
off organic carbon, and both subsamples were analyzed in the LECO deter-
minator to obtain their total carbon contents. The CaCO3 percentage was
calculated by multiplying the carbon c o n t e n t of the baked sample by 8.33
(i.e., the weight percent of carbon in a CaCO3 molecule). Although it is
possible that other carbonates may be present in the study samples, it is not
likely t h a t their concentrations are significant; thus, for this investigation
all carbonate carbon was considered to reside in CaCO3. The percentage of
organic carbon in the samples was obtained by subtracting the total carbon
of the baked subsample from that of the unbaked subsample.
The CaCO3 distributions within several box cores are indicated in Figs.9
and 10. Superimposed on these plots are two dashed horizontal lines, one
(M. L. ) representing the b o t t o m of the Mixed Layer, as determined by visual
inspection, and the other line (11K B.P.) representing the beginning of the
Holocene interglacial at 11,000 years B.P., as determined by the appearance
of tests of Globorotalia menardii menardii. In most cores, CaCO3 decreases
markedly below the 11,000 year horizon. This trend probably is due to
decreased calcite dissolution in bottom waters resulting from major climatic
changes beginning at the Pleistocene/Holocene boundary.
206

DEPTH ,c OEPIH
IN IN
C3RE CORE
cm, (era)
2O
_ /lic ,/7 IJ

:I
.~)C lO0 (53#6 ~ , l l I v )
C, - - - ,'0 A ~<0 4'0 50 40 50 60 TO 80 R~O
A ,cooo3 B %0o003

Fig.9. Profiles o f calcium c a r b o n a t e in t w o box cores. M . L . = base o f the M i x e d Layer;

1 1 K B . P . = P l e i s t o c e n e / H o l o c e n e b o u n d a r y at a p p r o x i m a t e l y 1 1 , 0 0 0 years B . P . A . Core
INMD BX 100. B. Core I N M D B X 101.

5 . . . . . . . . . . .

M ~
l0 L ~

DEPTH
IN 15
\ DEPTH
iN 20 . . . . . . . . . . . . . . /JP
CORE CORE
(cm) /3P (cm)

20 25

25 30

30 ~5

/2X i o , (3.279 ,~, ,A~ ) dX IO9 (3#95~, 5g )

40 510 610 i'lo 80 90 40 ~ 86--~- 9'0 ?5
A % CaCO$ B % CaD03
Fig.10. Profiles o f calcium c a r b o n a t e in t w o box cores. M . L . = base o f the M i x e d Layer;
1 1 K B . P . = P l e i s t o c e n e / H o l o c e n e b o u n d a r y at a p p r o x i m a t e l y 1 1 , 0 0 0 years B . P . A . Core
I N M D BX 104. B. Core I N M D B X 109.

As mentioned earlier, visual inspection of box cores indicates that the

Transition Layer is heterogeneously mixed and that the overlying Mixed
Layer is relatively homogenized. In order to test this hypothesis quantita-
tively, several samples from the same horizons in the cores were analyzed for
 207

CaCO3. Standard deviations were calculated for CaCO3 in the Mixed Layer
and also in the Hol ocene (i.e., p o s t - l l , 0 0 0 years B.P.) p o r t i o n of the Transi-
tion Layer (Table III). Data from the lower, Pleistocene p o r t i o n of the
Transition Layer were not considered, so t ha t carbonate differences due to
climatic changes would n o t confuse the results.
Although the n u m b e r of measurements was small, the standard deviation
o f CaCO3 percentages below the Mixed Layer in Holocene sediment is
double that of Mixed Layer sediment in cores INMD BX 100, 101 and 104
and is four times that of the Mixed Layer in core INMD BX 109 (Table III).
This suggests that the Transition Layer is two to four times as heterogeneous
as the surficial Mixed Layer with respect to CaCO3. Thus, the Mixed Layer
indeed seems to be hom oge ne ous , n o t only visually but also with regard to
its composition relative to the underlying sediment.
In order t o assess the effects of burrowing (including possible ingestion
and ex cr etio n ) o n sediment composition, we measured CaCO3 in specific
burrows and c o m p a r e d it to that in the surrounding matrix sediment. Three
box cores were analyzed, including INMD BX 72 from the N ort h Atlantic
and two cores collected on a previous expedition in the eastern equatorial
Pacific, PLDS BX 89 and PLDS BX 107 (Bryant, 1977).
For PLDS BX 89 two types of burrows were analyzed: those filled with
green sediment and those filled with sediment of similar color to ot her
matrix sediment in the core. Carbonate analyses show t hat some burrows of
b o t h types fall within the range of variation of matrix sediment from the
same depths in the core ( F i g . l l ) . Other burrows deviate significantly from
the matrix sediment of their own level, but m ost fall within the range of vari-
ation of o th er matrix sediment in the core. Four of the five burrow samples
in this latter group are green, and one has a lower carbonate c o n t e n t than
any o th er sample from the same core.
Carbonate data alone suggest t ha t the burrows which differ from the
surrounding sediment may be r e w o r k e d from above. The green color of the
burrows suggests biological alteration of the burrow-fill sediment, which may
result in the destruction or sorting out of carbonate particles, thus enriching
the sediment in n o n- c a r bonat e c o m p o n e n t s .
Samples of matrix sediment taken t h r o u g h o u t INMD BX 72 vary greatly

TABLE III

Calcium carbonate in selected box cores

Core No. Mixed layer Transition layer (Holocene)

2 s 2 s

BX100 41.7% 7.11% 34.9% 14.60%

BX101 70.4 1.30 72.9 2.34
BX104 78.1 2.23 80.3 4.23
BX109 90.9 0.36 90.8 1.30
208

IO
DEPTHIN 8 I OEPIH
IN
CORE 12
CORE 15
(cm) ! (cry)
16 ~

I
20b
25

3E'
i i i i . ~

20 30 40 5C 60 ?0 3O 40 50 60 lO 80 90
A % CoCO3 B % COCO]

8
DEPTH
IN 12
CORE
(cm) 16

20

24

28 IH ~)'I07 f#850~, 5/~)

19 20 30 40 50
C % C~C03

F i g . l l . Profiles o f calcium c a r b o n a t e in three b o x cores. The lettered aste~ ..... c o r r e s p o n d

to samples o f material filling particular burrows. A. Core INMD BX 72: A = b r o w n P l a n o -
l i t e s ; B = gray " P l a n o l i t e s " ; C = light b r o w n S k o l i t h o s ; D = dark b r o w n P l a n o l i t e s ; E =
b r o w n P l a n o l i t e s ; F = light b r o w n P l a n o l i t e s ; G = light gray P l a n o l i t e s . B. Core P L D S
Bx 89: A = g r e e n P l a n o l i t e s ; B = greenish b r o w n P l a n o l i t e s ; C, D = g r e e n P l a n o l i t e s ; E , F =
light brown P l a n o l i t e s ; G = w h i t e r e d u c t i o n halo; H = green S k o l i t h o s ; I = light brown
" P l a n o l i t e s " ; J = u n i d e n t i f i e d burrow. C. Core P L D S B X 107: A = dark b r o w n " P l a n o -
l i t e s " ; B = light brown P l a n o l i t e s ; C = dark b r o w n P l a n o l i t e s ; D = dark b r o w n P l a n o l i t e s ;
E = grayish brown P l a n o l i t e s ; F = light brown P l a n o l i t e s re-burrowed by smaller dark
b r o w n P l a n o l i t e s ; G -- dark b r o w n P l a n o l i t e s ; H = dark b r o w n " P l a n o l i t e s " .

in carbonate content. This variation roughly corresponds to visible stratifica-

tion in the core ( F i g . l l ) . Samples from the Mixed Layer have a higher car-
bonate c o n t e n t than the rest of the core and show relatively little horizontal
variation. Transition Layer samples from the 16-cm level, however, show
much horizontal variation.
Burrow samples from INMD BX 72 fall within the wide range of variation
of the matrix sediment in the core. Some burrow samples plot close to sedi-
 209

m e n t from their own level, while others resemble overlying or underlying

sediment more than their immediate surroundings. The best explanation for
the differences between the burrows and the surrounding sediment is vertical
reworking by burrowing organisms. Any biological alteration of burrow-fill
sediment in this core is difficult t o d o c u m e n t because of the great variability
of the matrix sediment. Some burrows, however, are gray, indicating possible
reducing conditions.
The matrix sediment of PLDS BX 107 shows a steady decrease in carbon-
ate with d ep th (Fig.11). Below approximately 20 cm there is less than 1%
carbonate, and the sediment is a clay-rich siliceous ooze. The greatest varia-
tion in the matrix sediment is near the surface; however, n o t all of the Mixed
L a y er in this core was sampled at the time of coring because of its soupy
consistency (Bryant, 1977).
Some burrow samples fall within the range of variation of matrix sediment
from their level in core PLDS BX 107. Two burrows in the lower, non-car-
b o n a te part of the core also contain little carbonate. These burrows probably
were made before carbonate sedimentation began at that site. A n o t h e r group
of burrows contained different carbonate percentages than the surrounding
sediment. The carbonate contents of these burrows fall between those of the
upper and lower parts of the core, and t h e y m ost closely resemble the car-
bonate c o n t e n t s in the 12--16-cm zone. These burrows range from 4 to 20
cm in d ep th in the core.
The burrows which are depleted in c a r bonat e relative to their surround-
ings may represent biological destruction of carbonate material, but the
presence of burrows with a similar carbonate c o n t e n t in the lower, non-
c a r b on ate layers of the core suggests upward and dow nw ard reworking of
sediment from depths of 12--16 cm.
In general, the carbonate analyses show the Mixed Layer to be distinct
from the rest o f the sediment in the cores. In the two Pacific cores, PLDS
BX 89 and 107, there is m uc h horizontal variation among samples taken
from the same depths in the Mixed Layer. Below the Mixed Layer is a hori-
zon with some of the lowest horizontal variation in the core. This relation-
ship opposes that seen in the Atlantic box cores. In INMD BX 72 the Mixed
Layer has a higher car bona t e c o n t e n t than the rest of the core, and it shows
some vertical variation of carbonate but relatively little horizontal variation.
Carbonate data suggest t ha t burrowing organisms m a y cause upward and
d o w n w a r d reworking of sediment, or reworking with no vertical displace-
ment. Biological destruction or size-sorting of carbonat e material also may
have taken place in PLDS BX 89.

Organic carbon. The a m o u n t of organic (i.e., non-carbonate) carbon material

in deep-sea sediment is largely a f unct i on of the oxidation potential of inter-
stitial waters. Generally this remains high enough to oxidize m ost of the resi-
dual carbon, unless circulation of the b o t t o m water becomes restricted, as in
a silted basin {e.g., the Black Sea). All samples in this study were collected
210

from unrestricted basins, so b o t t o m waters and sediments were well-oxygen-

ated, and residual organic carbon in the sediment was n o t expected to be
high. Indeed, analyses using the LECO carbon determinator showed that the
percentage of organic carbon in the box-cored sediment ranged from zero to
about 1.3%, with a mean of only 0.4% organic carbon in the surface layers of
the 23 Atlantic box cores in which organic carbon was measured.
The distributions of organic carbon within selected box cores are shown in
Figs.12 and 13. As with the CaCO3 plots, two dashed horizontal lines indi-
cate the bottom of the Mixed Layer (M.L.) and the beginning of the Holo-
cene interglacial (11K B.P. ). Absent from the carbon plots is any significant
trend, such as the pre-Holocene drop in CaCO3 observed in the carbonate
plots. Organic carbon, at least in the small quantities measured in our
samples, thus does n o t seem to be as directly influenced by climatic changes
as CaCO3.
Comparisons of the relative homogeneity of Mixed Layer and Transition
Layer sediments with respect to organic carbon were accomplished in the
same manner as for CaCO3 (Table IV). Only in core INMD BX 104 is there a
significant difference between standard deviations of organic carbon in the
Mixed Layer and Transition Layer; in this core the Transition Layer is twice
as heterogeneous as the Mixed Layer with respect to its organic carbon con-
tent.

Grain size. At the Pleistocene/Holocene boundary, the coarse (sand-size)

fraction of some Atlantic sediment changes considerably. The decrease in
number of larger foraminiferal species during glacial periods could result in

//12

I0

/;P
DEPIH 15 DEIPJ.H
IN
CORE CORE
(cm) (cm)
20

~5

30

~X ioo ( ~3,~6 ,,,, ~ioiv ) ~X ioi ( ~<ogm ,,,, 7 h )

I I ! i I I
0 I0 20 0 I0 20
A o~ ORGANICCARBON B '/o ORGANIC CARBON

Fig.12. Profiles of organic c a r b o n in t w o b o x cores. M . L . = base of t h e M i x e d L a y e r ; 1 1 I i

B.P. = P l e i s t o c e n e / H o l o c e n e b o u n d a r y a t a p p r o x i m a t e l y 1 1 , 0 0 0 years B.P. A, Core INMD
B X 100. B. Core INMD B X 101.
 211

tiff ff
Ol

10 DEPTH
,N
CORE
(cml
DEPTH 15
IN
CORE
( m)
.... _ ~ .......... 2 / ~ ,x ,0
20

i
25 3o~

]0

/2X / o , (3279 ~, ,A; ) /iX Io (3,~95,., 5 S ',

I I I I 210
0 I0 0 I0
A % ORGANIC CARBON L~ % ORGANIC CARBON

Fig.13. Profiles o f organic carbon in t w o b o x cores. M . L . = base o f the M i x e d Layer; 1 1 K

B.P. = Pleistocene/Holocene b o u n d a r y at a p p r o x i m a t e l y 1 1 , 0 0 0 years B . P . A . Core INMD
B X 104. B. Core I N M D BX 109.

a decrease in mean grain size of glacial sediments (Diester-Haass, 1975). Also,

dissolution contributes significantly to a decrease in mean grain size at depth
by causing fragmentation of larger foram tests and by corrosion of finer car-
bonate particles.
Grain size analyses of Atlantic box cores were accomplished by weighing
two duplicate samples, and then drying one and weighing it again to deter-
mine the percentage of water in the sediment. The other sample was sieved
wet on a 62-pm sieve, and the coarse fraction was dried and weighed. Weight
percent of the coarse (sand-size) fraction of sediment then was calculated.
The coarse fraction mainly consisted of remains of planktic foraminifera,
pteropods, radiolaria and diatoms, and some rare volcanic rock fragments or
micrometeorite debris. The fine-grained component consisted of clay par-

T A B L E IV

Organic carbon in s e l e c t e d b o x cores

Core No. M i x e d layer Transition layer ( H o l o c e n e )

)C s X s

BX100 1.25% 0.33% 1.08% 0.30%

BX101 0.97 0.27 0.61 0.32
BX104 0.67 0.15 0.44 0.35
BX109 0.36 0.12 0.15 0.15
212

ticles and remains of calcareous nannoplankton, small radiolaria, small

diatoms and fragments of larger plankton.
Samples of surface sediment from all South Atlantic and most North
Atlantic box cores were sieved. Additional samples of both Holocene and
Pleistocene sediment were taken in cores where it was suspected that grain
size differences may have been significantly different, i.e., in cores with low
sedimentation rates or good burrow preservation. Where the difference in
grain size between tops and bottoms of cores was greater than 35%, samples
were taken at 4--8 cm vertical intervals to determine patterns of change.
Grain size differences were most dramatic between tops and bottoms of
cores collected along the Sierra Leone Rise (INMD BX 101, 103, 104 and
106), indicating a possible increase in b o t t o m current velocity since the
Pleistocene. The grain size of Holocene sediments generally is larger than
t h a t of the Pleistocene (Fig.14). Other cores show little change from the
Pleistocene to the Holocene, either because they are so far above the lyso-
cline that changes in bottom-water temperature and circulation would have
very little effect on them (e.g., cores from the Rio Grande Rise and Mid-
Atlantic Ridge), or because t h e y are so far below the lysocline that the
coarse fraction has been continually dissolved out (e.g., cores from abyssal
plains ).
In order to examine possible textural changes in the sediment caused by
bioturbation, the grain size of material filling specific burrows was deter-
mined for two box cores collected previously from the equatorial Pacific
(PLDS BX 83 and 107). In both cores, the ratio of coarse- to fine-grained
material in the burrows fell within the range of those valves for the surround-
ing matrix sediment in the same cores (Fig.15). Thus, at the present time we
have no compelling evidence for large-scale textural alteration of the sedi-
ment by deep-sea burrowers, although size-sorting of grains for tube con-
struction has been observed (Fig.6).

Micropaleontology. The deep-sea oozes sampled for this study contain high
number of planktic foraminiferal tests, up to 2500 individuals per gram of
sediment. Different species show differing susceptibilities to dissolution in
sea water {Parker and Berger, 1971; Berger, 1973) and perhaps to mechanical
breakage as well. For this reason, relative abundances of different foramini-
feral species might be a sensitive indicator of changes in sediment composition
due to bioturbation as well as a means to trace the m o v e m e n t of sediment
between different layers in the core due to burrowing.
Samples of sediment filling obvious burrows and sediment adjacent to the
same burrows were dried, weighed and wet-sieved through a 62-t~m sieve. All
individuals of each planktic foraminiferal species in the sample were counted.
Tests showing preservation of 75% or more were counted as whole tests;
more poorly preserved tests were counted as fragments. Counts were stan-
dardized for comparison by calculating (1) the number of individuals of each
species per gram of sediment and (2) the percentage of total planktic fora-
minifera represented by each species.
 213

5 .... y/AL
I0 IO
DEPTH DEPTH
IN 15 IN
CORE CORE
(cm)
15
/ ';!f//. #.
(ore) 20 20

25 25

30 10
ILl ~ /01 (#0.~3~, 7'/11) ~_J( / o 6 (#z.?, 5%~

9 4 06 08 1.0 04 0 0.8 ~0
/I Send Silt & Cloy B Sand Sil~ ~, Cloy

DEPfH
IN
CORE
(ore) 20
IO
15

25

50
_ #-/

/1"~.#.
-
DEPTH IO
IN
CORE
(cm)
i5
0
5 . . . .

20

25

3.0
#.J_.

i/ 8.#.

/2Y/O# ~ # 7 0 2 ~ , 3"ID
I O 1.2 14 04 06 0'8 1.0 ILl
C Sand Silt & Cley D Sand Silt & Clay

Fig. 14. Profiles of s e d i m e n t texture, expressed as the ratio of the w e i g h t percent o f sand-
size material to that of silt-size and finer material, in four box cores from the South
Atlantic. M . L . = base of the Mixed Layer; 1 1 K B . P . = P l e i s t o c e n e / H o l o c e n e boundary at
a p p r o x i m a t e l y 1 1 , 0 0 0 years B.P. A, B> C, D. Cores INMD BX 101, 1 0 3 , 1 0 4 , 1 0 6 .

Series of s e d i m e n t s a m p l e s f r o m inside and o u t s i d e b u r r o w s w e r e a n a l y z e d

in cores t a k e n f r o m b o t h the A t l a n t i c and the Pacific. B o t h d i f f e r e n c e s and
similarities appeared b e t w e e n the m e m b e r s o f each pair o f samples, b u t the
results are s o m e w h a t difficult t o interpret. For e x a m p l e , a d a r k - c o l o r e d
Planolites f r o m c o r e P L D S BX 1 0 7 s h o w e d a m a r k e d d e p l e t i o n in w h o l e
tests and f r a g m e n t e d tests o f f o r a m i n i f e r a relative to t h e s u r r o u n d i n g sedi-
merit. T w o green Planolites f r o m core P L D S BX 89 w e r e e n r i c h e d in non-
resistant f o r a m i n i f e r a l tests relative to the s u r r o u n d i n g s e d i m e n t . Other
b u r r o w s , such as a Planolites f r o m c o r e I N M D BX 72 w h i c h w a s slightly
lighter than t h e s u r r o u n d i n g s e d i m e n t in c o l o r , w e r e fairly similar to the
o u t s i d e s e d i m e n t in f o r a m i n i f e r a l c o n t e n t .
In order t o gain a clearer u n d e r s t a n d i n g o f the variation o f s e d i m e n t w i t h -
214

0 O

5 5

I0 I0
DEPTH DEPTH
IN IN ~B
15 15
CORE CORE
(cm) (cm)
20 20

25 25

30 30
/3X3 (~53/~, /~] ~.~'107 (#gSO~, 61~]
I I I I I l I I I I I I
0105 0 I0 0.15 0.20 0.25 0.30 0105 010 0ll5 020 0.25 0,30
Sand~Silf $ Clay Sond: Silt & Cloy

Fig.15. Profiles of s e d i m e n t t e x t u r e , e x p r e s s e d as t h e r a t i o of t h e w e i g h t p e r c e n t of sand-

size m a t e r i a l t o t h a t o f silt-size a n d f i n e r m a t e r i a l , in t w o b o x cores f r o m the e q u a t o r i a l
Pacific. T h e l e t t e r e d asterisks c o r r e s p o n d t o samples of m a t e r i a l filling p a r t i c u l a r b u r r o w s .
A. Core PLDS B X 83: A = green P l a n o l i t e s ; B = light b r o w n Planolites. B. Core PLDS
B X 107; A = w h i t e Planolites; B, C, D = dark b r o w n Planolites.

in the core, two cores (PLDS BX 89 and INMD BX 72) were selected for
more detailed study. A vertical series of samples of the matrix sediment (i.e.,
that lacking clearly defined burrows) were taken at 4-cm intervals, and the
foraminifera, counts were standardized for comparison.
In PLDS BX 89 the burrows which were sampled differ from the rest of
the sediment in several ways. The most striking difference is in foraminiferal
species composition. Fig.16A shows the c o m m o n non-resistant foraminifers
( Globigerinoides ruber and G. sacculifer ) versus the c o m m o n resistant species
(Globorotalia tumida, G. menardii, Neogloboquadrina dutertrei and Pullenia-
tina obliquiloculata), with respect to the number of individuals per gram of
sediment. It is evident t h a t the burrow samples have higher numbers of non-
resistant species and lower numbers of resistant species than most of the
other matrix samples in the core. Only a matrix sample from the 24-cm hori-
zon has as many non-resistant individuals as do the burrow samples. This
graph also shows t h a t samples from the surface, as well as from the 2-, 4-, 8-
and 12-cm levels, plot close to each other, demonstrating the similarity be-
tween Mixed Layer and upper Transition Layer sediments in this core.
Similar relationships are seen in the graph of the percentage of total indivi-
duals represented by each species (Fig.16B).
A plot of the ratio of c o m m o n non-resistant species to c o m m o n resistant
species in core PLDS BX 89 (Fig. 17A) shows that the burrow samples are
enriched in non-resistant species relative to the surrounding matrix sediment.
A plot of the total number of foraminiferal tests per gram of sediment
against depth in core (Fig.17B) shows abrupt changes in the matrix samples
which correspond to color changes between the different strata in the core.
 215

300

25O

200 ~20

150 15

100 ~0
o
5O z5

0 7 9 ~ 0
500 1000 1500 2000 70 80 90 100
A Resisfonf sp. (per gm) e % Resisfonf sp

Fig.16. Comparisons of solution-resistant planktic foraminiferal species with relatively

non-resistant species in core PLDS BX 89. A. Data based on the number of whole fora-
miniferal tests per gram of sediment. B. Data based on the percentage of individuals of
resistant and non-resistant species in the sediment. In both graphs A and B, the numbered
solid circles correspond to matrix sediment samples from various levels in the cores, the
numbers referring to the number of centimeters below the core surface where the samples
were taken. The lettered asterisks correspond to samples of material filling particular
burrows: A = green Planolites, 26 cm below the surface; B = green Planolites, 28 cm be-
low the surface; C = light tan Planolites, 21 cm below the surface; D = green Planolites, 17
to 19 cm below the surface; E = green Skolithos, 23 to 26 cm below the surface.

T h e b u r r o w s a m p l e s g e n e r a l l y h a v e f e w e r t e s t s t h a n t h e s u r r o u n d i n g sedi-
ment.
The foraminiferal data from core PLDS BX 89 indicate that sediment
inside burrows have a microfossil species composition different from most
other sediment in the core. The burrows could have obtained their high
n u m b e r s o f n o n - r e s i s t a n t f o r a m i n i f e r s o n l y if t h e y h a d b e e n i n f i l l e d w i t h
sediment of a composition similar to that of the 24-cm level sample, unless
s o m e u n k n o w n p r o c e s s a c t e d t o e n r i c h t h e i r c o n t e n t in n o n - r e s i s t a n t t e s t s .
0 0
4 4 -Q
/
8
12 O[PTH ~2
DEPTH
IN
IN 16 ~o CORE 16
CORE i
{m) 20 . .c (cm) 28 ~c
24 4t.A
/-----~.11. '~E 24
88 28
32 / /2Y #'9
~X#; -~- ~--~-o
e" I i I ._ 32
05 I0 15 20 25 500 I000 1500 2000
A Nonresiston f sp :Resistont sp. e Whole rests (per gin)
Fig. 17. Profiles of planktic foraminifers in box core PLDS BX 89. A. Data based on the
ratio of number of individuals of non-resistant species to that of solution-resistant species
in the sediment. B. Data based on the total number of whole foraminiferal tests (resistant
and non-resistant) per gram of sediment. In both graphs A and B, the lettered asterisks
correspond to samples of material filling the same burrows referred to in Fig. 16.
216

T h e r e are o t h e r i m p o r t a n t differences b e t w e e n the b u r r o w samples and

the m a t r i x samples in P L D S BX 89. The b u r r o w s are d e p l e t e d in w h o l e fora-
miniferal tests a n d large f r a g m e n t s . The s e d i m e n t filling these b u r r o w s prob-
ably was altered biologically in such a w a y as t o cause d e s t r u c t i o n or sorting
o u t of the larger c a r b o n a t e particles. This process a p p a r e n t l y affects the
c h e m i c a l l y resistant tests t o a greater degree t h a n the c h e m i c a l l y non-resis-
t a n t tests.
A p r o c e s s w h i c h m a y a c c o u n t for these differences and also the green
c o l o r of these b u r r o w s m a y be ingestion o f s e d i m e n t b y animals w h i c h back-
fill t h e i r b u r r o w s w i t h feces. The digestive process m a y cause m e c h a n i c a l
a n d / o r c h e m i c a l d e s t r u c t i o n o f s o m e larger c a r b o n a t e particles, a n d it is
possible t h a t this p r o c e s s does n o t favor t h e preservation of large, n o r m a l l y
solution-resistant foraminiferal tests.
F o r core INMD BX 72, p l o t s o f n o n - r e s i s t a n t f o r a m i n i f e r a l species (Orbu-
lina sp., Globigerinella siphonifera, G. ruber, Globigerinoides conglobatus)
against resistant species (Globorotalia truncatulinoides and G. inflata) s h o w
t h a t the b u r r o w samples generally fall within the range of variation o f m a t r i x
samples f r o m the core (Fig.18). Variations in species c o n t e n t in the burrows,

800

~ 600 o~

-%~. z/~ ~ 30

~ ,,, 20

20o / /I *%~ =~ 15
0 1 2 - / o , ,

.o
/])C7.2 /]% 7.2
0 - ~ I I 5 L J i L ~ J
500 1000 1500 40 50 60 l0 80 90
A Resistant sp. (per gm) B % Resistant sp.
Fig.18. Comparisons of solution-resistant planktic foraminiferal species with relatively
non-resistant species in core INMD BX 72. A. Data based on the number of whole fora-
miniferal tests per gram of sediment. B. Data based on the percentage of individuals of
resistant and non-resistant species in the sediment. In both graphs A and B, the numbered
solid circles correspond to matrix sediment samples from various levels in the cores, the
numbers referring to the number of centimeters below the core surface where the samples
were taken. The lettered asterisks correspond to samples of material filling particular
burrows; A = light brown Planolites, 26 cm below the surface;B = grayish tan "Plano-
lites", 18 cm below the surface; C = matrix sediment immediately adjacent to B; D =
dark brown Planolites, 23 cm below the surface; E = l i g h t t a n Skolithos, 12 to 16 cm
below the surface; F = light tan Planolites, 28 cm below the surface; G = dark brown
Planolites, 17 cm below the surface ; H = matrix sediment immediately adjacent to A.
 217

therefore, could easily result from animals filling their burrows with sedi-
ment from other layers, or else simply leaving their burrows open to the sur-
face to be passively infilled with surface sediment.
When the total number of foraminifers per gram of sediment is plotted
against depth in core, some aspects of the nature of the mixed layer may be
seen (Fig.19). Below the 8-cm horizon the matrix samples plot in fairly
smooth trends. Two replicate samples from 24 cm (taken horizontally 8 and
16 cm away from the original sample) allow for a possible variation of 400
individuals per gram in this layer. Above 8 cm, however, the original matrix
samples show great variation; two replicate samples from the 4-cm level show
that the possible variation in this layer is at least 1300 individuals per gram.
Foraminiferal particles larger than 2 are unevenly distributed in the core,
especially in the Mixed Layer. The most likely explanation for this variation
may be that burrowing organisms fractionate the sediment by particle size,
and this process appears to be most intense in the Mixed Layer. Such size
fractionation does not seem to affect the species composition of larger fora-
miniferal tests. Percentage data of each species of the total planktonic fora-
minifers may be used to test this hypothesis statistically. Because of the
gross similarities in the proportions of the different species, however, most
of the samples from INMD BX 72 would be judged homogeneous by a ~2
distribution test at most c o m m o n l y used levels of significance (Novak,
1980).
The species composition of sand-size foraminiferal tests in the Mixed
Layer of INMD BX 72 is constant, while the distribution of that size fraction
throughout the Mixed Layer is n6t. Sand-size particles at the 24-cm level,
and probably other levels below the Mixed Layer as well, are homogenous in
terms of species composition. However, vertical changes in the sand-size frac-
tion of the sediment may be quite abrupt below the Mixed Layer. Sediment

DEPTH 12 ~ ~E
IN j '

(cm) 20
24 ~ . - ~

28 ,,!~Z 7.?

500 I000 1500 2000 2500

Whole Tests (per gin)
Fig. 19. Profile of planktic foraminifers in box INMD BX 72. Data based on the total
number of whole foraminiferal tests (resistant and non-resistant) per gram of sediment.
The lettered asterisks correspond to samples of material filling the same burrows referred
to in Fig. 18.
218

from cylindrical, gray burrows 18 cm deep in the core was compared with an
averaged Mixed-Layer species composition and with sediment samples from
16- and 20-cm depths in the core. The burrow sample was most similar to
the Mixed Layer, closely followed by the 20-cm sample, but it is quite dis-
tinct from the 16-cm sample (Novak, 1980). It is probable, then, that the
burrow filling represents downward reworking of surficial sediment.
In summary, foraminiferal data demonstrate three different effects of
burrowing. In PLDS BX 89, most of the burrows that were sampled are
filled with sediment which is different from all other samples in the core,
and these differences are probably biological in origin. In INMD BX 72, the
burrows generally fall within the range of variation of other sediment
samples from the core, and differences between the burrows and surrounding
matrix sediment are mainly due to vertical displacement of sediment by
burrowing organisms. Finally, burrowers in the Mixed Layer act to homo-
genize the species composition of the sand-size foraminiferal tests in that
horizon, b u t they also cause this size fraction to be distributed inhomogene-
ously throughout the Mixed Layer.

Quantitative m i x i n g m o d e l s

Because the omnipresent burrowers disrupt the original stratigraphy of

deep-sea sediment, many workers have sought to develop quantitative
models of sediment-mixing by organisms. Berger and Heath (1968) proposed
a simple model which assumes homogeneous mixing in order to explain the
redistribution of sedimentary particles when the rate of mixing is much
greater than the rate of sedimentation. In such a situation, which is the usual
case for deep-sea sediments, burrowing organisms would completely homo-
genize the upper few centimeters of a deposit. As sedimentary particles fall
o n t o the surface, they are dispersed throughout the upper "mixed layer" of
the sediment. Because this "mixed layer" has a constant thickness, sediment
is incorporated into "historical layers" below the "mixed layer" at a rate
approximately equal to the sedimentation rate. The "mixed layer" moves
continuously upward to keep pace with sedimentation. This model was used
to predict the distribution of microfossils in the sediment column following
the sudden appearance or extinction of a species.
Guinasso and Schink (1975) expanded on this idea by allowing for differ-
ing mixing and sedimentation rates. In their model, the various types of sedi-
ment mixing can be characterized by a dimensionless parameter G, defined
by three variables,
G = D/Lv
where: D = eddy diffusivity or mixing rate (cm 2 per 1000 years); L = depth
of " m i x e d layer" (cm); and v = sedimentation rate (cm per 1000 years).
When G is greater than 10, the sediment is totally homogenized before burial,
and the model approximates that of Berger and Heath (1968). When G is
 219

less than 0.1, there is little mixing of the sediment before final burial because
of the relatively high sedimentation rate.
Peng et al. (1977) modified Berger and Heath's (1968) model to predict
the distribution of a non-conservative tracer with continuous input, such as
x4C, in deep-sea cores. They found that biologic mixing causes 530- to 1800-
year deviations in the '4C ages of sediment from the true ages. They also
found relatively constant 14C ages in the upper 10 cm of sediment, which can
best be explained by the thorough mixing of old and new sediment by organ-
isms. Below this " m i x e d layer", the 14C ages they observed were predicted by
their model with moderate accuracy.
By employing modifications of the model of Guinasso and Schink (1975),
several workers have c o m p u t e d coefficients of sediment mixing for the
Atlantic and Pacific. Nozaki et al. (1977) calculated a mixing coefficient of
190 cm 2 per 1000 years for deposits on the Mid-Atlantic Ridge, based on the
profile of 21pb in the sediment; Peng et al. (1979) calculated a mixing coef-
ficient of 120 cm 2 per 1000 years for calcareous ooze in the western equato-
rial Pacific, based on both 14C and 2~Pb profiles.
Williams et al. (1978) examined the distribution of 14C produced by
atomic testing in box cores of abyssal red clay. They f o u n d relatively high
14C activities in the upper " m i x e d layer" of the cores, reflecting the rapid
fixation and sinking of organic carbon in the sea and its slow oxidation on
the sea floor. They also found unexpected penetration of organic carbon
from the " m i x e d layer" in the upper 4 cm of the sediment down to at least
12 cm in the core. Plutonium isotopes also were present in the upper 12--14
cm of the sediment. These observations indicate bioturbation well below the
"mixed layer", although bioturbation alone may not explain all the features
of isotope distribution that they observed.
The presence of a homogeneously mixed surface stratum (i.e., Mixed
Layer) in the Atlantic box cores which we studied was demonstrated earlier
in this paper (see sections on "Core stratigraphy" and "Effects of bioturba-
tion on sediment composition and texture"). The 3--10-cm range of thick-
ness of the Mixed Layer in these cores is not inconsistent with that predicted
by the quantitative models just discussed. In fact, Berger and Johnson
(1978) suggested that ~4C stratigraphy in cores from all three major oceans
indicates a Mixed Layer averaging roughly 8.5 cm thick.
However, Berger et al. (1978) examined stable oxygen and carbon isotopes
in one deep-sea core and concluded that bioturbation also can introduce
" l u m p y " noise into the signals of isotope variation with time. This type of
disturbance would not be expected if there were only homogenous sediment
mixing. Indeed, all the box cores investigated for this study do contain an
easily observed, heterogeneously mixed Transition Layer beneath the Mixed
Layer. The stratigraphic errors caused by heterogeneous mixing cannot be
eliminated with certainty by any mathematical means known at present. We
simply require much more data on such aspects of the deep-burrowing
infauna as their average body size, population density, burrowing speed and
specific style of bioturbation.
220

CONCLUSIONS

The major conclusions of this research may be summarized as follows:

(1) Regardless of latitude, water depth or sediment type, bioturbation of
pelagic abyssal deposits in the Atlantic Ocean is intense and total. The origi-
nal fabric of the box-cored sediment had been obscured and overprinted by
the effects of bioturbation in all cases; neither primary lamination nor
cttrrent structures could be discerned in any of the cores.
(2) Optimal preservation of burrows occurred in cores which possessed
either very little carbonate throughout or else a much greater carbonate
content of the Holocene sediment than of the underlying glacial Pleistocene
sediment, thus allowing visible differences between burrow fill and matrix
sediment to be highlighted. Cores from many of the deeper sites tended to
exhibit greater visibility of burrows, due to the dissolution of carbonate and
the resultant concentration of metal oxides and sulfides which give the sedi-
ment its most vivid colors.
(3) Several well-known ichnogenera could be recognized in the box-cored
sediment. Planolites was ubiquitous; Skolithos was abundant only in carbon-
ate-rich deposits; Trichichnus was cosmopolitan but occurred only in the
uppermost few centimeters of any core; Chondrites and Zoophycos, which
are very abundant in DSDP cores, occurred in several box cores but were
uncommon. Slightly washed core tops contained the partially e x h u m e d
passageways of Spirorhaphe and, less c o m m o n l y , Cosrnorhaphe and Paleo-
dictyon. There was evidence of U-shaped burrows in some cores, but their
geometries could n o t be determined with sufficient accuracy to a t t e m p t
ichnogeneric identifications.
(4) With the exception of several uniform-colored, shallow-water (i.e.,
pure white carbonate) and deep-water (i.e., pure red-brown clay) cores, the
visible stratigraphy within the box cores consisted of three zones: a surficial
Mixed Layer, extending 3--10 cm below the water/sediment interface; an
intermediate Transition Layer, extending 20--35 cm below the water/sedi-
ment interface; and the Historical Layer, extending down to the b o t t o m of
each core.
(5) The Mixed Layer can be recognized as a visibly uniform and typically
dark-colored stratum capping the box core. It c o m m o n l y contains numerous
open burrows, although its shear strength may be very low. This is a zone of
intense bioturbation, especially by meiofaunal organisms, which results in
the total destruction of original stratification and the homogenization of
sediment composition and texture.
(6) The Transition Layer is a zone of heterogeneous mixing, and it ex-
hibits m a x i m u m color contrasts of burrows. Calcium carbonate and, to a
lesser extent, organic carbon and grain size distributions within selected cores
indicate that the Transition Layer may be two to four times as heterogeneous
as the overlying Mixed Layer.
(7) The Historical Layer is that in which no active bioturbation occurs,
 221

and it c o r r e s p o n d s t o the t r a c e fossil r e c o r d . N e a r t h e t o p of this z o n e ,

b u r r o w c o m p a c t i o n and s e d i m e n t stiffness increase, a n d the brightness o f
s e d i m e n t colors and the distinctness of b u r r o w margins decrease.
(8) T e x t u r a l a n a l y s e s y i e l d e d n o e v i d e n c e f o r or against large-scale altera-
t i o n o f s e d i m e n t grain size by deep-sea b u r r o w e r s , a l t h o u g h sorting of grains
b y size, and p o s s i b l y also b y shape, was e v i d e n t in the walls o f c e r t a i n dwel-
ling t u b e s c o n s t r u c t e d b y u n k n o w n organisms.
(9) T h e d i s t r i b u t i o n s of p l a n k t i c f o r a m i n i f e r s w i t h i n several b o x cores,
including inside and o u t s i d e p a r t i c u l a r b u r r o w s , i n d i c a t e d t h a t b o t h h o r i z o n -
tal and vertical m i x i n g o f m i c r o f a u n a s can result f r o m b i o t u r b a t i o n . In o n e
core, the m i c r o f o s s i l c o m p o s i t i o n of b u r r o w fill m a t e r i a l was significantly
richer in foran~iniferal species t h a t are relatively susceptible t o calcite disso-
lution t h a n the s u r r o u n d i n g m a t r i x s e d i m e n t . H o w e v e r , t h o s e same b u r r o w s
c o n t a i n e d far f e w e r u n f r a g m e n t e d f o r a m i n i f e r a l tests t h a n the s u r r o u n d i n g
s e d i m e n t . This suggests a sorting process, p e r h a p s i n g e s t i o n / d i g e s t i o n / e x c r e -
t i o n , w h i c h m e c h a n i c a l l y b r e a k s d o w n tests t h a t o r d i n a r i l y w o u l d be resis-
t a n t to calcite dissolution.
(10) O b s e r v e d t h i c k n e s s e s o f t h e M i x e d L a y e r in the A t l a n t i c b o x c o r e s
are c o n s i s t e n t w i t h a p p r o x i m a t i o n s p r e d i c t e d b y p u b l i s h e d q u a n t i t a t i v e
m o d e l s of s e d i m e n t - m i x i n g . H o w e v e r , o b s e r v e d t h i c k n e s s e s o f t h e T r a n s i t i o n
L a y e r , w h i c h are quite variable, c a n n o t be easily q u a n t i f i e d and p r e d i c t e d b y
existing m i x i n g m o d e l s .

ACKNOWLE DGEMENTS

This research was s u p p o r t e d b y the N a t i o n a l Science F o u n d a t i o n ( G r a n t s

O C E 7 3 - 2 1 6 0 1 a n d O C E 7 8 - 2 5 8 4 4 to E k d a l e ) a n d the U n i v e r s i t y of U t a h
( D e p a r t m e n t of G e o l o g y and G e o p h y s i c s Special F u n d s grants to Muller and
N o v a k ) . We are i n d e b t e d to W. H. Berger for his e n c o u r a g e m e n t , assistance
a n d ideas, and we t h a n k P. H. R o t h a n d S. A. Walker for their help w i t h the
i d e n t i f i c a t i o n o f f o r a m i n i f e r s . F o r t h e i r h e l p at sea, we also t h a n k P. F.
B r y a n t , N. B. B r y a n t , D. Ripley, S. A. Walker, T. Walsh, o t h e r m e m b e r s of
t h e I N D O M E D II and X I I scientific parties, and t h e c r e w of the R/VMelville.

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