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Transcript
SIO 277
Deep-Sea Biology
Deep-Sea Biology
SIO-277
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Instructor – Lisa Levin
Contact: [email protected], 534-3579
Room 2236 Sv - call before visiting
Tues & Thurs, 9:30-10:50, in Vaughan 100
Website:http://cmbc.ucsd.edu/Students/Current_Stude
nts/SIO277/
• Series of 20 lectures ~ 70 min w/discussion
• Assigned readings on electronic reserve prior to each
lecture, also some book chapters
• Texts:
– Gage and Tyler - Deep-Sea Biology
– Koslow - The Silent Deep
Student Requirements:
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Read assigned papers before
each lecture
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Attend each lecture
•
Student Cruise - Oct. 31 (Saturday)
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Mid term assignment Challenger Forward – due Oct. 29th
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Research Proposal (Deep-Sea Future)
Develop research needed for stewardship of the deepsea in the face of climate change, mining, fishing, energy
extraction etc. Read about subject, identify unanswered
problem/issue, write a hypothesis-based research
proposal (5 page max).
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Abstract due Nov. 3 or earlier
Proposal due Nov. 24th
Oral presentation of proposal in
a symposium during final exam week Dec. 10th
Course Evaluation
Grading: Letter or S/U
• Take Home Mid-Term Assignment
(30%) Challenger Forward
http://19thcenturyscience.org/HMSC/HMSCINDEX/index-linked.htm
• Written Research Proposal (40%)
• Oral Presentation (25%)
• Participation in Discussion/Cruise (5%)
Reading for Sept. 29, Oct.1
Sept. 29 Physical Environment:
• Gage and Tyler 1991. Chapter 2.
Oct. 1 Faunal composition, depth zonation:
• Carney, R.S. Zonation of deep biota on
continental margins. 2005. Oceanogr. and
Mar. Biol: An Annual Review 43: 211-278.
• Gage and Tyler 1991. Skim through images
on pages 61-162.
Orientation – The global ocean
A blank slate
Heterogeneous bathymetry
Most of the Earth is Ocean
and most of that is Deep Sea
• Percent
distribution of
earth’s surface
assessed in
vertical relief.
• Hypsographic
curve
Defining the deep sea
> 200 m
(beyond the shelf break)
or
> 1000 m (hard core)
Definitions can be important much human impact is < 1000 m
Zonation Terminology
meters
200 Continental Shelf
500
Upper
Continental
Slope
Epipelagic (euphotic)
Bathyal
Mesopelagic (disphotic)
1000
3000
Lower Continental
Slope
Bathypelagic (aphotic)
Continental Rise
4000
6000
Benthopelagic
Abysss
Hadal
Continental Margins
Abyss
History: Before Exploration
Socrates (600 BC): The beginning of wisdom is to
know that one knows nothing
Aristotle (300 BC): The ocean (deep sea) is a
frontier to be explored [180 spp, recorded from the
Aegean Sea)
Pliny (50 BC): The deep sea is an inferior world.
All we know of it is all there is to be known.
Posidonius (1 BC) Mediterranean Sea is 2000 m
deep.
History of Deep-Sea Biology
• Eric Mills – Problems in Deep-Sea Biology: an
historical perspective (In: The Sea, Vo. 8 – the
deep –sea edited by G. Rowe – 1983) – book
on reserve
• Tony Koslow - Chapter 1 in The Silent Deep.
“The rise of deep-sea Exploration: Early
paradigms” pp 8-22
Chapter 2. “On the shoulders of giants: The
Challenger Expedition”pp. 23-39.
The beginning
1.
John Ross – (1818 – 1819)
Baffin Bay –deep sea ‘clamm’ - 4 samples from 850
to 2000 m [crustaceans, corals, shellfish, worms,
basket star from 1.6 km)
In searching the NW Passage
2. James Clark Ross (1841-1847)
Tasman Sea/Antarctic – fauna to 750 m
Noted similarity with high latitude fauna and
concluded uniform cold temperature at seafloor.
3. Harry Goodsir – (1845)
Davis Strait (Arctic)– fauna dredged to 550 m
Edward Forbes
(1815-1854)
• Described marine faunas of European Seas
• Described major biogeographic provinces
from Arctic to Mediterranean and Caspian
seas
• Vertical zonation of benthos - established the
science of marine benthic ecology - pattern in
species distributions
• Aegean Sea to 420 m - unfortunate focus.
Unproductive waters, little in deep water.
• Ignored work of earlier scientists
Azoic Hypothesis (1859)
life absent > 550 m (300 fathoms).
• Deep sea is
– Dark
– Cold
– High Pressure
– Stagnant and anoxic
How could life survive?
Azoic Hypothesis
• Attacked by G.C. Wallich
• 1860 - 13 starfish recovered from a sounding
line at 1260 fathoms (2300 m) in the N.
Atlantic off Greenland
• 1861 - Allman and Milne Edwards –
15 species recovered from a broken
telegraph cable in the Mediterranean
between Sardinia and N. Africa at 2300 m
including stony coral.
Yet the azoic paradigm persisted
John Jeffreys (1861 – 1868)
Shetland Island dredging to 311m
204 species
Osprey
• 51-ft sailing
vessel
• No auxillary
power
• Hemp lines for
dredging
Azoic theory in question
Otto Torell (1861-1865)
a. Benthic fauna at 2560m off Spitzbergen
Michael Sars and G.O. Sars (1864-1868)
a. Dredging to 550 m in Norwegian fjords
b. 427 species of invertebrates including
asteroids and crinoids (connections to fossil
record)
Led to the idea of deep sea a refuge for extinct faunas.
Azoic theory in question
L.F. de Portales & L. Agassiz (1867-1868)
dredged fauna off Grand Bahama Bank
to 1555 m depth.
W.B. Carpenter & C. Wyville Thomson
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H.M.S. Lightning (1868)
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Dredged fauna to 1189 m N.E. Atlantic
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Deep water temperatures low (0 – 8.5oC)
Beginning of Big Science
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Government funding
Networking
Politicking
Manipulating the public’s imagination
Wyville Thompson and William Carpenter
HMS Lightening 1868
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North of Scotland between Shetlands and Faroes
Dredged 10 d of 6 wk at sea! To 1180 m
Abundant life everywhere, relict species
Brisingid starfishes, Hexactinellid sponges
Arctic outflows and NA Deep Water (0oC)
separated by a ridge from warmer (6.4oC) Gulfstream-influenced Atlantic waters. Fauna varied
with water temperature
H.M.S. Porcupine (1869-70)
First ship specifically equipped for
oceanographic studies in deep water (first fully
organized oceanographic expedition)
Carpenter and Thomson - chief scientists
West of Ireland - dredged to 2700 m
South of England/France - 4450 m
Mediterranean Sea w of Spain
Recovered all major groups (mollusks, crustaceans,
echinoderms, sponges, stalked crinoids, banks of
Lophelia pertusa, primitive urchins - Living fossils
1st use of protected thermometer
Led to hypothesis of density-driven, deep-water
circulation.
W. Thompson
H.M.S. Challenger (1872 – 1876)
Beginning of modern oceanography
(Institutional, collaborative, multidisciplinary)
C. Wyville Thomson – Expedition leader
+ 4 naturalists (Murray, Buchanan, von WillimoesSuhm, Moseley) + 1 artist (Wild)
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3.5 years
68,890 nautical miles
362 stations
40% time spent in ports
19th century equivalent of the US space program in 20th century
Challenger Expedition
Evolved from a one page proposal!
Objectives - To map:
1. Global patterns of deep-water circulation
2. Chemistry of world’s oceans
3. Geology of the deep-sea floor
4. Distribution/abundance/origin of deep-sea
organisms
Determine: chemical composition of seawater, physical conditions
of the deep sea, characteristics of sediment deposits,
distribution of organic life
H.M.S. Challenger
(226 ft corvette)
• Dredging equipment
– Winch
– Dredging platform
– Accumulators (safety
springs)
– Beam trawl
H.M.S.
Challenger
• Natural
history
laboratory
Primordial ooze
• Origin of life from inanimate
matter
• Bathybius haeckelii
primordial ooze
• Missing evolutionary links
between fossil and modern
organisms
Bathybius haeckelii, 1868-76. Viewed under the
microscope the small discoids are the exoskeletons
of tiny sea creatures, while the jelly within which these
are suspended is the gelatinous gypsum precipitate.
www.creationism.org/ books/TaylorInMindsMen
In 1858 Huxley had found gelatinous
matter - identified as Monera and
called Bathybius haeckelii formed living sheet over much of
the ocean and provided food
source for higher organisms
Alcohol + water yielded
(Gypsum - Ca Sulfate)
H.M.S. Challenger Expedition
Departed Portsmouth – 21 Dec. 1872
South to Madeira, St. Thomas, Bermuda,
Halifax, Azores, Cape Town, Kerguelen
to Antarctic pack ice, Melbourne, Fiji,
Hong Kong. Philippines, Japan, Hawaii,
Tahiti, Valparaiso, Falkland Islands.
Arrived Portsmouth – 24 May 1876
Challenger Expedition Results
1.
2.
Animals collected throughout the ocean to 5500 m depth
Many deep-sea species of many taxa, high proportion of
rare species, many with direct development
3. Decreasing abundance and diversity with depth generated paradigm of the Depauperate Deep.
4. Different taxa in deep than shallow water (zonation)
5. Stability of deep-sea environment (temperature, chemical
composition, lack of seasonal changes)
6. Constant seawater constituent ratios.
7. Deep-sea sediments (calcareous, siliceous oozes) of
pelagic origin (foraminifera, radiolarians, coccolithophores,
pteropods, diatoms, etc)
8. Red clay of terrestrial origin in central oceans, manganese
nodules rich in metals
9. Description of water masses based on T and S
10. Description of shelf , cont. slope upper and base
Theories Laid to Rest
1.
Azoic Theory disproved. Animals present throughout the
deep sea to 5500 m one sample at 7000 m Japan Trench.
2.
Huxley’s Bathybius -(artifact of preservation) addition of
alcohol to seawater caused precipitation of gypsum.
3.
No living fossils - trilobites or Belemnites [extinct
cephalopods] Rather deep fauna evolved from continental
shelf and slope forms … relatively recent - with onset of
Glaciation in Cenozoic.
4.
No large, cosmopolitan deep-sea species, but genera
widely distributed (5-7% at high and low latitudes, 3-4%
had bipolar distributions). 1 species in common between
Pacific and Atlantic at mid equatorial latitudes.
Challenger Results
Scientific results published in 50 volumes with final
summary by John Murray (1895- 13 years after
Thomson’s death at 53).
29,500 pages 3,000 plates
Samples distributed to experts globally
a. high species diversity in deep-sea
b. common taxa in high latitudes
c. greater depth ranges for deeper species, sharper
zonation in shallow water
d. endemism common in deep water
e. first proof of deepwater plankton (between 915
and 1830 m depth)
f. Discovery of mid Atlantic Ridge
Post-Challenger Expeditions
Alexander Agassiz (1877- 1888) USA
a. Steamer Blake in Gulf of Mexico and
Caribbean.
b. replacement of hemp with wire rope (less
deck volume, more efficient handling)
c. abundant fauna dredged to 3567 m depth
d. existence of midwater plankton
e. reiterated question of food supply to deep
sea – sinking of plankton, role of terrestrial
debris.
f. caught nothing below 200 m (100 fathoms)
with Sigsbee gravitating plankton net
Italian circumnavigation by the Vetter Pisani
(1882-1885)
– Chierchia and Palumbo using a crude openingclosing net discovered deep-water plankton (i.e.
siphonophores) to 2300 m depth.
– Chun, inspired by these studies, sampled off
Naples finding a rich pelagic fauna exists to
1400 m depth – hypothesized a “ladder of
vertical migrations” (Chun, 1887).
Valdivia expedition (1898-1899)
– Carl Chun expedition leader
– European and African coasts to Antarctica,
Indian Ocean, Red Sea, Suez Canal,
Mediterranean Sea to Hamburg
– Opening/closing nets revealed extensive
deepwater pelagic fauna to 2000 m depth
– below which fauna became sparse.
French Oceanographic Expeditions
1.
2.
Travailleur – (1880) – Bay of Biscay
Talisman – (1888-1927) – E. Atlantic, Mediterranean.
– Distinctive deep-water ichthyofauna, with
vertical migrators
– Distinctive abyssal molluscan fauna originating at
high latitudes
– Homogeneity of deepwater fauna across the
Atlantic
– Ancient annelids are widespread, modern ones
are not.
Prince Albert of Monaco
• Series of cruises on Hirondelle, Princess Alice,
Princess Alice II, between 1885 and 1914.
• Technological advances using wire rope, steam
winches, large closing trawls, baited traps to abyssal
depths
• Areas of study – N.E. Atlantic, Mediterranean
• Circulation of N. Atlantic studied with drifters
• Sardine fishery off N. Spain, marine mammals
• Successful trawling to 6035 m* off Cape Verde
• Confirmed vertical migrations by deep pelagic fauna.
• Used baited traps - confirmed existence of scavengers
at depth (lysiannasid amphipod 14 cm long)
• Once worked a 120 h continuous station at 5940 m off
Portugal
*deepest until 1947
Prince Albert - cont.
• Significantly advanced technology to study
deep-sea communities
• Migrating bathypelagic fauna exists
supporting Chun’s “ladder of migrations”
hypothesis concerning transport-food supply
to deep ocean
• Unique combination of creativity and financial
independence.
Pre WWI
Michael Sars cruise (1910)
• a European collaboration
• John Murray (Great Britain) and Johan Hjort
(Norway) – N.E. Atlantic
a. Extensive pelagic trawling, minimal benthic
trawling (5160 m)
b. Further speculation on sources of nutrition
for deep sea- phytoplankton – zooplankton –
detritus-dissolved organic matter
c. Influential text on oceanography (Murray
and Hjort –1912- The Depths of the Ocean.
WWI through WWII
• From 1910 until after WWII – deemphasis of deep-sea research
– More emphasis on coastal fisheries
problems and plankton dynamics
Post WWII
1. Swedish Deep-Sea Expedition(1947-1948)
– Circumnavigation in Albatross –
– Seismic studies, piston coring, bottom
water sampling
– only last 3 months devoted to deep-sea
benthic trawling
– Benthic trawling in Puerto Rican Trench –
to 7900 m – deepest trawl in N. Atlantic
Post WWII
Danish Deep-Sea Expedition – Galathea (1950 – 1952)
around the world.
- Anton Bruun – expedition leader
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1st thorough study of abyss & hadal zones
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1st microbiology in the deep sea
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1st quantitative abyssal samples (grab)
• Trawled to 10,190 m in Philippine Trench (sea anemones,
amphipods, isopods, bivalves, holothurians).
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- Trawled in five trenches recovering 115 species > 6000m depth. Found
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a distinct hadal fauna.
- Isolation of barophilic bacteria from deep-sea (Zobell and Morita, 1959)
- First use of 14C to estimate primary productivity (Steeman Nielsen)
Reports published even in 1980’s
Cold war era
Russian deep-sea expeditions (1950s)
- Vitiaz – extensive grab sampling to
determine benthic biomass in deep basins
and trenches of Atlantic, Pacific and Indian
Oceans (Zenkevitch, 1963; Belyaev, 1972).
Emphasis on feeding
ecology
Modern era
1949: there were less than 100 oceanographers in
the USA
1959: Oceanography budget 21 million
1969: Oceanography budget 221 million.
In those 10 years: 20 new vessels and 8 new
laboratories
American non-expedition studies (1960’s)
- Strong financial support from U.S. government
- Establishment of Gay Head Bermuda transect (55 to
5000 m depth) – Howard Sanders and Bob Hessler in
1965
- Anchor dredge, epibenthic sled, finer sieves
- Higher abundance, greater species diversity
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Antifreeze protein systems in Antarctic fish
Avian diversification
Biodiversity in protista
Biological interaction on islands
Collecting poisonous sea snakes
Deep-sea fish at the Antarcti c
Dissolved Organic Matter
Enzymes from the Ikaite columns in Greenland
Fluorescent proteins
Gingers
Oceanic oxygen deficiency zones
Parasites in zooplankton
Plankton dynamics in the Andaman Sea
Plant communities in the Galapagos Islands
Roseobacter bacteria – the ocean’s stars
Sea turtels in the major sea current systems
Sound in the Oceans
The Benthic Fauna of the Solomon Sea
The DNA of the Polar Seas
The European Eel
The Horseshoe Crab
The Marine Carbon Cycle
The origin of the vertebrate immune system
The physiology of antarctic fish
The significance of the climate the degree of isolation on biological interplay and
biodiversity in lakes
Galathea 3 - Danish
Vaedderen “the ram” ( 2006-2007)
Ram’) 3
Observatories
(e.g., Neptune Canada)