Deep--Sea Fauna,

Zonation and Biogeography

Who Lives in the Deep Sea?

Bacteria/Archaea
Protozoa
Meiofauna (42-500 m)
Macrofauna (>300 m)
Megafauna (visible, >1 cm)
Giants

BACTERIA AND ARCHAEA

in sediment
in guts
on detritus
on carcasses
as symbionts
on hard surfaces

Bacteria

aerobic
anaerobic

heterotrophic,
autotrophic

sulfate reducer
sulfide oxidizer
methane oxidizer
Fe, Mn oxidizers

Protozoa

Xenophyophores

Foraminifera
Komokiacea (superfamily)

Metazoan
Meiofauna

Nematodes

Ostracods

Gnathostomulida
Harpacticoid copepods
Kinorhynchs

Oligochaetes
Loricifera

Macrofauna
Peracarid
crustaceans

Brachiopoda

Mollusca

Amphipods, isopods
tanaids, cumaceans
Annelida:
polychaetes

Bryozoa
Priapulida

Megafauna - Echinoderms
Crinoids

Holothurians

Ophiuroids

Echinoids
Asteroids

Megafauna

Sponges

Mollusca
Cnidarians

Crustaceans
Annelida

Echiura

Hagfish

Enteropneust

Humongofauna
(Giants)
pycnogonids

giant squid

Munopsis

Colossendeis

bryozoan
Eurythenes
Kinetoskias

ascidians
Culeolus

Polychaetes often comprise half or more of the macrofauna

Site

Depth (m) Substrate

N. Carolina Margin I.
850
NC II.

NC III.

Horizon Guyot
1840
Santa Catalina Basin
1130
Central Pacific Seamounts 1480-3150
San Diego Trough
1230
Rockall Trough
2200
HEBBLE
4820
Porcupine Abyssal Plain >4500
Central North Pacific
5500
Aleutian Trench
7298

% Polychaetes

34% sand
20% sand
31% sand
Foram Sand
Mud
Calcareous muds
Mud
?
90% mud
Red Clay

43
74
66
47
77
66-67
76
59
67
35
55
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Dominant
Scalibregma
Chrysopetalid
Dorvilleid
Paraonid
Paraonid
Cirratulid

Ampharetid
Spionidae

Key Zoogeographic Features

Original view - 1 province, cosmopolitan species
(no geographic boundaries)
Ekman 1953 - 4 zoogeographic zones:
Atlantic, Indo-Pacific, Arctic, Antarctic.
Vinogradova - 1959, 62, 79. Species have limited distributions
Only 15% of species occur in > 1 ocean
Only 4% of species occur in all ocean
Genetic methods reveal cryptic species; cosmopolitanism rare
(France & Kocher 1996) Eurythenes gryllus is many species

But there is great similarity among

oceans at the generic level.
Genera are cosmopolitan
Of 143 isopod genera in the Pacific , 134 are present in the Atlantic
Desmosomatidae (Isopods):
North vs South Atlantic - same 12 genera
North Pacfic vs Atlantic - Pacific is missing 2 genera and
=77% similarity
has one not in the Atlantic
Asellote isopods
Poore et al. 94 - 67% of 98 genera
on the SE Australian slope are
in the Atlantic
Ophiomusium (brittle star) worldwide at mid to slope depths.

Vinogradova, 1979.
Zoogeographic divisions of the
abyssal and hadal zones of the world

Distinct hadal fauna (Belyaev 1959, 1989)

more amphipods, polychaetes, bivalves, echiurids, holothurians

Origins of Deep-Sea Fauna

Antarctic or shallow water origins then
submergence
Deep origins and Antarctic emergence
(Ilyarachnid isopods) in taxa with
diversity centers in the deep sea.
Isolated basins have endemics only at
species level, suggesting
invasion from shelves and recent
evolution.

A.
B.
C.

High lat
Equator
Mid lat

Red Sea and Sea of Japan have

eurybathic species from Indian
& Pacific oceans.

What is Zonation?
Pattern of uniform change in species
Step-like boundaries between regions of
homogeneous composition.

Current usage
Non-repeating, sequential pattern of species replacement
measurable as changes in the overall rate of change
in faunal composition.

Why should we care about zonation?

Resource exploitation is increasing (fisheries, petroleum)
Sound management demands understanding of distributions

Zonation Terminology
meters
200 Shelf
500

Epipelagic (euphotic)
Mesopelagic (disphotic)

Upper
Slope

Bathyal
Bathypelagic (aphotic)

1000
Lower Slope
3000
Continental Rise
4000
6000

Abysssal

Benthopelagic
Hadal

Zonation in the Past

Since Challenger Expedition:
Scientists have recognized that species change much more rapidly
with depth down the continental margin than horizontally.
Depth-related changes in the ocean are considered one of the
greatest environmental gradients on this planet (Gage and Tyler ).

What can generate zonation in the Ocean?

What can generate zonation in the Ocean?

Temperature
Salinity
Bottom type
Food levels

Challenger/Sars expeditions
Murray and Hjort (1912)
3 ZONES:
Shelf (to 200/300 m)
Archibenthic transition (600/800 - 2000/3000) (bathyal)
(based on megafauna, topography, temperature)
Abyss
(upper boundary set at 4o C isotherm - Bruun 1957)
(lower boundary at 6000 m - topography)

shelf
transition

There is little area

between 1000 & 2500 m
Yet, this is the zone of
highest diversity

abyss

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Controlling Gradients: Light

Light: present to 1000 m
Childress 1995: the use of vision by fishes, crustaceans, squid
above 1000 m elevates metabolic rate and affects forms present

Controlling Gradients: Hydrostatic pressure

only variable directly related to depth (10 m = 1 atm)

pressure accelerates reactions in which the molar volume

of the products are less than the molar volumes of the
reactants.
Pressure retards reactions in which there is a volume
expansion (Le Chatelier effect)

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Effects of pressure in the deep sea

Increased dissolution of calcium carbonate
Increasing metabolic cost associated with maintaining
carbonate structures at depth (molluscs,
foraminiferans, echinoderms show effects)
Influence on chemosynthesis based on H2S? Stability
of gas hydrates (>450 m)
Protein structure, membrane fluidity, lipid content
Different propreties of gellatinous/membranous
megafauna?
A piezo barrier to downward colonization: 500-1000
m, 2000-3000m

Controlling Gradients: Temperature

55oN to 55oS - Warm, low density upper layer
Thermocline may form a barrier. Much of the deep sea is 2-4oC
Increasing temperature increases chemical reaction rates.
2 to 3 x rate increase with 10o C temperature increase.
Adaptations to cold:

increasing concentrations of enzymes

adopting enzymes effective at low temperatures

incorporating modulator compounds that help maintain

enzyme reactions over a range of temperatures
Are water mass effects the result of temperature (?)

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Controlling Gradients: Organic Flux

Since Forbes (1859) - importance of food recognized.
Decreasing food input with depth
Different rates of mixing within sediment with depth
Food effects on zonation may be modified by competitors.
Effects of OM flux include changes in sediment geochemistry.
High flux depletes oxygen. .

TROX model foraminifera distributions

based on oxygen and food

TROX model
-applied to
foraminifera

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Controlling Gradients:
Oxygen
Oxygen minima at 100-1000 m
(<0.5 ml/l = 22 M)
Sharp faunal changes at upper and lower boundaries
Maximum densities, low diversity at boundaries
Low density, diversity in OMZ core
Composition changes within OMZ associated with depth
OMZ faunas

Reductions in large organisms, lots of small ones

Dominance by nematodes, annelids

Shift in nematode: harpacticoid ratios

Drop in species richness and evenness

Features Affecting Vertical Zonation

Physiological Parameters - Mainly affect top 1000 m
Light - declines with depth
Temperature - cooler with depth
Salinity - polar/surface freshening
Oxygen - midwater minima
Pressure - increases 1 atm with 10 m depth
(larvae have specific sensitivities as do adults)
Depth-Related Features - usually act as gradients
Substrate Type - grain size, grain composition
Organic matter source (terrestrial vs marine)
Flow Regime - accelerated near topography
Water Masses- carry larvae
Changing Resources with Depth
Organic Matter Availability- reflects surface production &
horizontal advection
Habitat area/space
(least from 1000-3000 m, greatest from 4000-5000 m)

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Pelagic Zonation
Governed largely by light but in some places temperature.
Evidence of the importance of competition
comes from non-overlapping distributions of related species
Ontogenetic zonation is common
Often larvae are found shallowest and gravid females deepest
(Tradeoffs between food and mortality from predation)

Zonation may be forced by physiography

of the ocean floor
SHELF BREAK - Boundary between shelf and deep-sea fauna.
Referred to as the mud line by John Murray (1895)
= upper limit for muddy bottom, deep-sea conditions
may occur at 200 m (W. Europe), 500 m ( Antarctic)
or few m (fjords)
HEMIPELAGIC VS PELAGIC SEDIMENTS Ekman (1953)
Break between hemipelagic (with considerable terrigenous input)
and pelagic sediments is a key zoogeographic boundary.
ANTARCTIC - Little difference between shelf and deep-sea faunas

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Bathyal (200-3000 m) vs
Abyssal (4000 - 6000 m)
Bathyal has shallow and deep faunas
Abyssal with true deep faunas.
But new data by Rex for molluscs suggests that most abyssal species
also occur in the bathyal realm and few are endemic.
(Rex et al. 2005; Am. Nat. 165).
Menzies:
Greater continuity of shelf and bathyal faunas at high latitudes
and of bathyal and abyssal faunas at low latitudes. Why?

Means of Evaluating Zonation

Multivariate Classification Procedures rely on similarity matrices
similarity coefficients: Percent Similarity Coefficient,
Bray Curtis coefficient, Normalized Expected Species Shared [NESS]
Cluster Analysis -highly dependent on parameters chosen
(can achieve multiple forms of clustering with same data set)

Ordination - employs linear models, best when only a small

part of the depth gradient is being examine.

Vertical ranges of species zonal breaks are those

where greater number of upper and lower depth limits occur
(requires large data set)
Cumulative species curves

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Porcupine Seabight
zonal
breaks
are
those
where
greater
number
of
upper
and
lower
depth
limits
occur

Cascadia Basin

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What Changes with Depth?

Taxonomic composition
Diversity
Density, Biomass
Body Size
Trophic structure

Megafauna - New England Margin

(Hecker 1990- Photographic survey)
(284,692 indiv. 94,380 m2 of seafloor)

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Hecker, 1990
Upper slope: to 700 m Dasmosmilia lymani, Flabellum alabastrum
Upper-middle slope: (1100-1200 m)
Geryon quinqueidens, Synaphobranchus
Nezumia, Phycis chesteri, Glyptocephalus
Transitional Lower Middle Slope: 1500-1700
Distichoptilum gracile + Anemones
Lower Slope: Ophiomusium lymani, Cerianthid anemone,
Distocptilum gracile, Echinus affinus

Hecker, 1990
Species replacement with depth was gradual in flat areas and
abrupt in steep areas.
Geographic variation was observed related to
glacial inputs/hard substrate.
Lowest densities on middle slope
Depth ranges are narrow on upper slope and broad on lower slope

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HECKER, 1990

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Examples
Along Gayhead Bermuda Transect
Gastropods and cumaceans show narrower depth ranges
than polychaetes and brittle stars.
In N. Atlantic, sharpest changes are at shelf,
and a zone of 400-1000 m is reflected in megafaunal changes
(Sanders, Grassle, Haedrich, Rex).
N. Atlantic studies support a step model, with biggest changes
at 200 m, 400-600 m, 1000 m, 1400 to 1600 m, 2000 m.
Echinoderms 800-1200 m and 1800 m - Rockall Trough.
Agree with Bay of Biscay (Sibuet).

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Pacific Examples
In North Pacific Fish (Oregon/Washington) (Pearcy et al. 1982)
New species appear at:
400-700 correspond to OMZ boundary.
1900 to 2000m.
Species disappear at
500-900 m
2800-3100 m (floor of Cascadia Plain)
Holothurians show only gradual replacement (Carney & Carey 1977)

Mediterranean - (Emig 1997)

Expression of upper bathyal zone is directly related to slope
physiography, thus slope profile should be given in studies.
Upper bathyal - belts of suspension feeders related to flow structure
Maximal density in upper bathyal (higher than shelf)
Zones: 100-600 m, 600-1300 m, 1400-1600 m, 1600-2000, > 2000

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Carneys Overview
Carney 2005
Upper Boundary
Biota Shelf to <2000 m
Intermediate
Boundary Biota encountered by 1500 m
(dont extend to upper
or lower boundaries
Lower Boundary
Biota
extend up from abyssal
plain (fewer in
isotermal waters (MED,
Arctic)

Zonation below the Abyss?

Vertical and trophic partitioning by amphipods
A

Blankenship
et al. 2006
Deep-Sea Res.

Blankenship &
Levin 2007
L and O

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Compositional Zonation
MACROBENTHOS
Shallower Taxa Amphipods, molluscs, polychaetes
Increasing in deep water -tanaid and isopod (peracarid) crustaceans
-aplacophorans (mollusca)
-sipunculans, priapulids

MEGABENTHOS
Sponges more common
on the slope,
echinoderms
(holothurians,
asteroids,
brittle stars)
below

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Zonation of Body Size

WITHIN TAXA
Larger size species found at deep end of taxon range
Asellote isopods - smallest genera < 250 m, largest > 1000 m
Tanaids,
Gastropods
Foraminifera.
ACROSS TAXA
Decrease in average body size with depth
- replacement of large by small species
(except fish in NW Atlantic)

Lampitt et al. 1986 -Porcupine Seabight

At shallow depths large species dominate biomass
At deeper depths small species dominate biomass
*****

Small species - shallow:deep

11% vs 34% of biomass
93% vs 98% of numbers

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Zonation of Feeding Mode

Reduction in suspension feeding &
increase in deposit feeding with depth
Replacement of sedentary deposit feeders
with mobile deposit feeders at deeper depths.
(Fauchald and Jumars 1979)
50:50 surface and subsurface-deposit feeders at depth.

Taxon Rates of Species Change with Depth

Do big and little animals show similar zonation?
Is zonation pattern taxon specific?
Rex 1977
Megafauna changed most rapidly with depth
but holothurians changed slowly (Carney and Carey 1982)
Predatory gastropods changed at an intermediate rate
Infaunal polychaetes changed slowest
Higher trophic levels have narrower bathymetric zones

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Testing Causes of Zonation

Barnacle Manipulative Experiments
Dayton et al. 1982
-transplanted 250 Bathylasma coroliforme
from 400 m to 25-40 m.
Normally live > 100 m
Individuals survived for 2 y and produced larvae
Authors argued water movement restricted distribution.

Faunal Age Change with Depth

Original belief :
the deeper the zone the more ancient the fauna
(primitive protobranch bivalves, brotulid fishes)
primitive generalists do best
For holothurians and starfish,
bathyal forms are most primitive,
abyssal and hadal forms are specialized
Radiation of diversity in the deep sea occurs for few taxa
Ilyarachnidae (isopods)
Early belief - diversification in Antarctic then submergence to deep

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Horizontal/Lateral Extent
Do vertical zonation patterns persist over wide horizontal ranges?
Wide lateral ranges off S. Australia (bathed in homogeneous
Antarctic intermediate water).
Atlantic margin - more restricted lateral ranges
Associated with eastern, western boundary water masses
And strong gradients in productivity
Gayhead-Bermuda transect bivalves (Allen and Sanders 1996)
Fauna more similar between adjacent basins than depths
Increasingly cosmopolitan than depth.

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