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PLANKTON(Bio 511)
Professor Stephen T. Tettelbach
C.W. Post Campus of Long Island University
PLANKTON
Introducing the world's first exclusive formulation of nature's superfood, marine phytoplankton , micro-algae naturally grown, harvested and available in a powerful daily liquid nutritional supplement. For the first time in history, all the wonderous health benefits of Marine Phytoplankton ,
the source of ALL life in the oceans is now,
readily available to man! Phytoplankton is utilized by whales - the worlds longest living mammals .
This mammal lives for over 150 yrs, and is sexually active for life!
plankton, from Greek word for ‘drifter’ or ‘wanderer’
organisms that are free-floating or found in the water column (i.e. pelagic) but are unable to swim against typical currents
generally small
neuston = plankton living at or just below the water surface (e.g. bacterial film)
pleuston = plankton that live at the surface but protrude into the air(e.g. Portuguese Man O’War)
nekton = pelagic animals that are powerful enough swimmers to move at will in the water column, under most conditions (e.g. fish, dolphins, squid)
benthos = organisms living on or in the bottom (in the subtidal zone)
T ab le 3 . Lo co m otion and sw im m ing spee ds o f so m e co m m on n earshore zoo p lankto n w ith co m p aris on s to fish a nd hu m an s G rou p
M e a n s of lo co m o tio n
Ch ara cte ristic
m ovem ent
Sw imm ing speed (cm /se c)
B ody lengths/second
Ciliates
Cilia
0.001 -0 .06
1.5-10
Scyphom edusae
R hy thm ic c on tractions of
m uscles around the bell
Dis tinctiv e puls ing
m otion
2-5
0.5-0.7
(dia/sec)
Hydrom edus ae
As abov e
As abov e
<0.2 -1 .0
1-5
Ctenophores
8 rows of long paddle c ilia
G liding m otion
Rotifers
Beating of c il ia on the c orona
(wheel organ) a t the an terio r
end
G liding m otion
0.05 -0.15
6-14
3-8.6
Copepod nauplii
(Acartia, Oithona,
Oncaea*)
Beating of appendages from
the head region
In term ittent
sw imming or
"jum ps"
0.01-0.1
0.5 m ax
1-4
C opepods (adult
calanoids, m ean
sp ee d ) (Acartia,
femora)
Prim arily by bea ting o f 2nd
antennae and other c ephalic
appendages
Sm ooth steady
sw imming or
halting, "s ink and
sw im" motion
0.14
1 -1 .5
Adult c alanoid esca pe
resp o nses (Calanus3)
Prim arily by v igorous bea ting
of thoracic appendages
80
300
M ysids (opossum
shrim ps)
Beating of abdom inal appendages (pleopods)
Sm ooth gliding
even during direc -
tional c hanges
2-8
2 -10
Shrim p and crab
zoeae (blue crab ,
green s hore crab, —
Z1)
Beating of thoracic
appendages, occasional
abdom inal flex ions
Us ually sm ooth ,
occasional pauses
or jerks
2-3
0.067
about 1
Am eric an lobster
(postlarv a)
Beating of abdom inal
appendages
7-13
O yster (ey ed veligers)
Cilia o f the v elar lobes
Sm ooth, steady
sw imming
0.08 -0.24
0.1-0.3
about 1
C haetognaths (arrow
w orm s)
D ors ov entral flex ions o f
longitudinal m uscles
R ap id , da rting
motion
0.5-3
0.5-5
F ish larvae
U ndu la tions o f trunk and tail
R outine burs t
s w im m ing
0.5-3
1-3 2-6
B a rrac u d a
U ndula tions o f trunk and tail
Bu rsts of up to a hundred m eters
700+ (27 m ph)
5-8
H um an (O ly m pic sw im m er)
Flailing of arms and legs
"Freestyle"
180
1
aS e e P a ffe nh o fe r e t a l. 1 9 96 a n d L e n z et a l . 2 00 4 for d e scrip tio n s o f sw im m in g b e ha vio r. from: Johnson & Allen (2005)
Size classes of the plankton
(from Johnson & Allen (2005)
see handout
Other definitions sometimes used:
ultraplankton: < 2 µm = femtoplankton + picoplankton
holoplankton = organisms that spend their entire life in the plankton
meroplankton = organisms that spend only part of their life in the plankton, usually as larvae
Major groups of organisms that comprise the plankton:
• phytoplankton
• zooplankton
• protozooplankton (= protozoans)
• mycoplankton (= fungi)
• bacterioplankton (= bacteria)
• viruses (sometimes called virioplankton)
For some amazing photographs of plankton – see: Wim van Egmond’s Micropolitan Museum: http://www.microscopy-uk.org.uk/micropolitan/index.html
viruses
– obligate intracellular parasites
- when in the plankton: ‘between hosts’
-most abundant members of the plankton, up to 108 -109 viruses/ml mostly bacteriophages; also: viruses of eukaryotic algae, others
#’s vary spatially, seasonally
bacteria
• tremendous diversity• exist virtually everywhere • heterotrophic forms: most get nutrition via dissolved solutes, plus, a few predaceous bacteria• autotrophic forms: chemosynthetic photosynthetic: cyanobacteria + others • 2 Kingdoms of bacteria: Eubacteria and Archaeobacteria
some important cyanobacteria: Synechococcus,Trichodesmium (marine)
Nostoc, Anabaena (fw)
bacteria
prochlorophytes: Kingdom Eubacteria Division Cyanobacteria Class Prochlorophyta most: genus Prochlorococcus • picoplankton (<2 µ)• unique divinyl derivatives of both chl a and chl b• tropical and subtropical oceans; as deep as 150-200 m • may account for 30-80% of 1° productivity in oligotrophic regions of the ocean
http://biology.kenyon.edu/Microbial_Biorealm/bacteria/prochlorococcus/prochlorococcus.htm
protozooplankton (= protozoa)
•all heterotrophic• In 6 Kingdom scheme of Woese, Kingdom Protista Subkingdom Protozoa In plankton, most important are: Phylum Ciliata (ciliates) Phylum Sarcomastigophora Subphylum Sarcodina (amoebas, shelled amoebas) Subphylum Mastigophora (zoo)flagellates many authors group these with photosynthetic flagellates - we will not.
Phylum Ciliata (>7000 spp.) move, feed with cilia very important in the food chains some filter feed on phytoplankton, some ingest phytoflagellates, some ingest bacteria. a few get nutrition from endosymbiotic microalgae.
Paramecium
protozooplankton
Phylum Sarcomastigophora Subphylum Sarcodina
as a group, feed on protozoa, very small metazoans (mostly microcrustaceans), bacteria, phytoplankton;
many get nutrition from endosymbiotic microalgae
amoebas (=amebas) pseudopodia used in locomotion and feeding
Amoeba proteusextending pseudopodia to feed on a desmid(phytoplankter)
protozooplankton
Phylum Sarcomastigophora Subphylum Sarcodina
shelled amoebas (non-motile) extend actinopods - fine, stiff cytoplasmic projections, through openings in shell, for feeding and locomotion.
Actinosphaerium, showing extended actinopods
protozooplankton
Phylum Sarcomastigophora Subphylum Sarcodina Foraminifera (bearing many small openings) = forams (~ 4000 spp.)• made 1° of CaCO3• exclusively marine and brackish; most common in coastal areas of the ocean.
Protozooplankton - Foraminifera
protozooplankton
Phylum Sarcomastigophora Subphylum Sarcodina
The radiolarians - exclusively marine and brackish; most common in open ocean.
vast areas of deep ocean have sediment referred to as radiolarian ooze.
shells made 1° of amorphous silica with organic inclusions
Protozooplankton - radiolarians
protozooplankton
Phylum Sarcomastigophora Subphylum Sarcodina
Acantharia - shells of strontium sulfate
Lychnaspis miranda
protozooplankton
Phylum Sarcomastigophora Subphylum Sarcodina
Heliozoans – marine, brackish, but 1° fw. Some have silica
Actinosphaerium
protozooplankton
Phylum Sarcomastigophora Subphylum Mastigophora (flagellates)
Phytoplankton
•photosynthetic organisms, usually unicellular, but may form colonies• Miller (2004) estimate: spp. ~ 5000 spp., some authors: total # of phytoplankton spp. may be >100,000 much debate about algal taxonomy:
for our purposes, cyanobacteria and other photosynthetic bacteria will all
be treated as bacteria (procaryotes)
eucaryotic algae are all placed in the Kingdom Protista
[see handout: Table 26.2 from Prescott, Harley and Klein (2005): Comparative Summary of Some Algal Characteristics]
Phytoplankton - Diatoms
Kingdom Protista Division (=Phylum) Chrysophyta (golden plant) Class Bacillariophyceae
Diatoms (Gr. dia: across, temnein: to cut) • 2 parts of cell wall fit together like parts of a Petri dish
• unicellular (may form chains), or sometimes filamentous• cell wall: hard mineral shell (= frustule) composed of hydrated, polymerized silicic acid Si(OH4) -same as opal
• golden-brown (yellow-brown) when healthy; greenish when not
• marine, brackish, fw• planktonic and benthic; also in air and ice.• found from poles to tropics; most abundant in polar to temperate zones.• usually 5-200µm, but may be up to 4 mm.
Phytoplankton – Diatoms
2 major forms: centric – rounded; essentially radially symmetric
Coscinodiscus
Dityum brightwelli
Phytoplankton – Diatoms
Chains of centric diatoms
Thalassionema nitzschioides
Eucampia zodiacus
Phytoplankton – Diatoms
pennate (L. pinnatus: feathered) – elongate; bilaterally symmetric pinnate refers to markings on walls of some of these forms
Cymbella affinis
Pseudo-nitzschia
cleanedfrustule of pennatediatom
Phytoplankton – Diatoms
Chains of centric diatoms
Chaetoceros affinis
Actinoptychis
Phytoplankton – chain of Antarctic centric diatoms
Phytoplankton - Dinoflagellates
Kingdom Protista Division Dinophyta (Gr. dinos: whirling; L. flagellum: whip) (=Pyrrophyta) (Gr. pyros: fire, and phyton: plant) – refers to bioluminescence of many forms
• most have 2 flagella• may be armored (thecate)
or
unarmored (naked)
Noctiluca
Dinophysis
Peridinium
Ceratium
Phytoplankton – Dinoflagellates
• marine, brackish, and fw• usually ~2-200µ , but may be up to 2 mm in size• very interesting ecologically: some forms do not have chloroplasts and are therefore completely heterotrophic - these may prey on diatoms, protozoans or even copepod nauplii
• ~ 60 species (Horner, 2002) have been identified to secrete powerful neurotoxins (cause paralytic shellfish poisoning, ciguatera,etc.)
• very important as endosymbionts in coral reef spp. (e.g. Symbiodinium)
Phytoplankton – Dinoflagellates
2 main taxonomic groups, based on where flagella insert
desmokonts: 2 flagella arise from anterior part of cell
Phytoplankton – Dinoflagellates
dinokonts: – one flagellum in transverse groove (cingulum): whirls cell around,
2nd simpler flagellum in longitudinal groove (sulcus) helps pull cell thru water
Important representatives:
Alexandrium, Gonyaulax,Gambierodiscus, Pfiesteria
Phytoplankton – Green Algae
Kingdom Protista Division Chlorophyta (Gr. Chloros: green)
• many multicellular examples, but also some unicellular and planktonic forms; these include spherical, filamentous and colonial forms; some w/ flagella, some without• marine, brackish, but mostly in freshwater
fw examples:
Volvox
Closterium (floater)
Chlamydomonas
Phytoplankton – Green Algae
Kingdom Protista Division Chlorophyta (Gr. Chloros: green)
desmids (Gr. desma: bond) – cells are arranged in mirror image halves, joined by isthmus, where spherical nucleus is located • all fw , usually in more oligotrophic waters
© Wim van Egmond: http://www.microscopy-uk.org.uk/micropolitan/index.html
Phytoplankton – Pelagophytes
Kingdom Protista Division Heterokontophyta Class Pelagophyceae
• very small (<2 µ) = picoplankton• most in oceanic waters, recently discovered • most in genus Chlorococcus
• now thought to be the principal eukaryotic picoplankton: extremely important in terms of the amount of C they fix
Chlorococcus
Phytoplankton - Silicoflagellates and relatives
Kingdom Protista Division Heterokontophyta Class Dictyophyceae
• external silica skeleton, 1 flagellum.
Phytoplankton – Raphidophytes
Kingdom Protista Divison Chromophyta Class Raphidophyceae (according to Horner (2002)• flagellates – all photosynthetic• marine, estuarine • important fish killers (produce toxins) especially important in aquaculture
Dictyocha
Heterosigma
Phytoplankton – Euglenophytes
Kingdom Protista Division Euglenophyta
•unicellular flagellates, with 1 very long and 1 very short flagellum
• 1° freshwater
Assorted euglenoids
Phytoplankton – Prymnesiophytes
Kingdom Protista Division Prymnesiophyta
• golden-brown flagellates, usually have a thread-like, variable length appendage called a haptonema (Gr. haptein: to fasten) located between the 2 flagella of the cell.
• end of haptonema is sticky and used to anchor cell to a substrate or to capture food• cells are covered by 1 or more layers of unmineralized organic scales and/or by CaCO3 scales called coccoliths
• some harmful and produce toxins that harm finfish, benthic inverts and even macroalgae
Chrysochromulina
Phytoplankton – Prymnesiophytes
Kingdom Protista Division Prymnesiophyta coccolithophorids
(have CaCO3 scales called coccoliths)
Scales can reflect light so water appears whitish during blooms
Emiliana huxleyi
Coccolithophorid blooms in the Celtic Sea and Gulf of Biscay
www.co2.ulg.ac.be/objects/intro_cocco.htm
Zooplankton
• many of the roughly 35 animal phyla have representatives in the plankton: marine and/or fresh waters
• some groups are exclusively holoplankton, • some are exclusively meroplankton,
• many phyla have members in both categories
Zooplankton
Phylum Cnidaria (= Coelenterata)
• simple body plan: with central mouth, usually surrounded by tentacles• radial symmetry• all members possess stinging cells (cnidoblasts = nematoblast)• 2 major body forms: polyp and medusa• 1st larval stage: planula
• ~9,000 spp.
4 classes: Hydrozoa (hydroids, Hydra, some jellyfish) Scyphozoa (true jellyfish) Anthozoa (corals, sea anemones) – only larvae are planktonic Cubozoa (box shaped jellyfish, including sea wasp of Australia)
some authors include within Scyphozoa
Zooplankton
Phylum Cnidaria Class Hydrozoa
actinula larva
hydromedusa: Maeotias
hydromedusa: Benthocodon (deepwater)
Hydromedusa Colobonema sericeum
Phylum Cnidaria Class Hydrozoa
Zooplankton
Phylum Cnidaria Class Hydrozoa
Physalia (Portuguese Man O’War) Vellela (by-the-wind sailor)
Phylum Cnidaria Class Hydrozoa
Physalia
(Portuguese Man O’War)
Zooplankton
Phylum Cnidaria Class Scyphozoa
Stomolophus
www.marlin.ac.uk
Lion’s Mane Jellyfish – Cyanea capillata
Sea Nettle – Chrysaora
quinquecirrha
http://www.bbep.org/images/seanettle.jpg
Phylum Cnidaria Class Scyphozoa
Zooplankton
Phylum Cnidaria Class Cubozoa
Chironex flexneri (sea wasp)
Zooplankton
Phylum Cnidaria Class Anthozoa
tube anemone larva
Zooplankton
Phylum Ctenophora (comb jellies)
• closely related to Cnidaria• simple body plan, with central mouth, w/ or w/o tentacles• radial symmetry• have 8 cteni (rows of cilia) used in locomotion and feeding• often bioluminescent• only marine• ~150 spp.
Beroe
animaldiversity.ummz.umich.edu/.../view.html
www.huriisgaard.biology.sdu.dk/Mnemiopsis%209...
Phylum Ctenophora: Mnemiopsis leidyi (sea walnut)
Zooplankton
Phylum Rotifera (Rotatoria) – wheel animals• very small, most < 1 mm• have circular corona of cilia that looks like a rotating wheel – used in locomotion and feeding• radial symmetry• no planktonic larvae• mostly fw• ~2,000 spp.
Collotheca with egg
Lecane
Zooplankton
Phylum Platyhelminthes – (flatworms)
• flat, bilateral symmetry• many spp. are parasitic• of the free-living members of the Class Turbellaria - usually benthic, but many spp. swim in water column for brief periods• Muller’s larva• ~12,000 spp.
Dugesia
Zooplankton
Phylum Nematoda (roundworms)
• small, unsegmented, smooth; bilateral symmetry• mostly benthic• move via ‘lashing’ movement rather than ‘peristaltic’ motion• no planktonic larvae• ~12,000 spp.
Zooplankton
Phylum Nemertea (ribbon worms)
• large, flat, unsegmented; bilateral symmetry• mostly benthic but enter plankton as larvae (pilidium) or when swarm during mass spawning• > 800 spp.
Cerebratulus
pilidium larva
Zooplankton
Phylum Annelid (segmented worms)• bilateral symmetry• initial larval stage is trochophore • ~12,000 spp.
3 classes: Polychaeta (~9,000 spp.)- pronounced head- each segment with paired appendages called parapodia, plus several bristles (setae) on each segment- marine only- several spp. holoplanktonic- 3 families produce epitokes = reproductive adults that swarm en masse
into water column
Zooplankton
Phylum Annelida Class Polychaeta
holoplanktonic adult polychaete
Class Oligochaeta (incl. earthworm)• few bristles • usually reduced head• no parapodia• fw and marine; usually benthic
Class Hirudinea (leeches) • no setae • reduced head• have suckers for attachment • often parasitic
Swarming polychaete epitokes at night, Glover's Reef, Belize
www.ryanphotographic.com/epitoke.htm
Zooplankton
Phylum Mollusca (mollusks)
• most important groups have calcareous shells• initial larval stage is trochophore• some classes have holoplanktonic adults, but mostly meroplanktonic larvae• ~100,000 spp.
7 classes in all:
Class Cephalopoda• includes squids, octopus, Nautilus • considered nekton as adults• marine only
larvaloctopus
trochophorelarva
Zooplankton
Phylum Mollusca Class Gastropoda (snails and slugs) • usually 1 shell (or none)• mostly benthic • have radula for feeding • some holoplanktonic; many with planktonic larvae: trochophore, veliger (shelled) • terrestrial, fw, marine Conus spp.
Conus larva
Cypraea guttata
Zooplankton – adult holoplanktonic gastropods
pteropod (sea butterfly)
Glaucus glaucus
hydrozoan Porpita porpita
Zooplankton – adult holoplanktonic gastropods
heteropod
Zooplankton
Phylum Mollusca Class Pelecypoda (Bivalvia) – clams, scallops, etc.
• 2 shells• benthic • most filter feeders• most with planktonic larvae: trocophore, veliger (shelled)• fw and marine 8 mm bay scallop on eelgrass
bivalve veliger larvae
S. Tettelbach
Zooplankton
Phylum Chaetognatha (‘bristle jaws’) (arrow worms)
• small (<2 mm)• torpedo or arrow shaped• bilateral symmetry• rapid swimmers (many can avoid most plankton nets)• all marine – very specific temperature/salinity preferences• holoplanktonic• ~100 spp.
Zooplankton
Phylum Arthropoda (jointed feet)
• includes insects, spiders, crustaceans• > 1 million spp.• mostly marine; some fw; some terrestrial• many holoplankton as well as meroplankton
Subphylum Chelicerata Class Merostomata
Horseshoe crabs (Limulus) trilobite larva
© Joan Krispyn, 2007
Zooplankton
Phylum Arthropoda Subphylum Crustacea
• mostly marine, also fw, terrestrial•~50,000 spp.
Copepods – (> 10,000 spp.) most abundant animals in oceans mostly holoplankton (some parasitic) swim with antennae, thoracic legs, or mouthparts
Candacia
Cyclops
Zooplankton
Phylum Arthropoda Subphylum Crustacea
Ostracods
• bivalved shells (mostly opaque)• small (<2 mm)• many benthic, some holoplanktonic
Zooplankton
Phylum Arthropoda Subphylum Crustacea
Cladocerans (4 orders)
• most with calcareous shell (usually clear)• mostly fw• holoplankton
Polyphemus
Daphnia pulex
Zooplankton
Phylum Arthropoda Subphylum Crustacea
barnacles (Cirripedia)
• meroplankton only• larvae: nauplius, then cypris (cyprid)
Zooplankton
Phylum Arthropoda Subphylum Crustacea
Pericaridans• no carapace• no free larval stages• young brooded in ventral pouch
Mysids (opposum shrimp) usually < 15 mm prominent thoracic legs, large eyes ▼
Cumaceans ▲ small quasi-shrimplike usually <5 mm big heads, little tails eyeless
Zooplankton
Phylum Arthropoda Subphylum Crustacea
Pericaridans
Amphipods (‘beach fleas’)
- laterally compressed - mostly benthic, some planktonic ▼
▲ Isopods (‘pill bugs’) - dorso-ventrally compressed- mostly benthic, some planktonic
gammarid amphipod
Zooplankton
Phylum Arthropoda Subphylum Crustacea
Decapods – shrimps, crabs, lobsters, euphausids (=krill)
• most benthic; some holoplanktonic (e.g. krill)• complex life cycles with many larval stages (some shrimp with ~16);• larvae: nauplius, protozoea, zoea• planktonic postlarvae: crabs (megalops) lobsters (puerulus or phyllosoma)
euphausid Meganyctiphanes
Euphausids (krill) comprise the greatest biomass of any animal in the ocean
photography.nationalgeographic.com/photograph...
Zooplankton
Phylum Arthropoda Subphylum Crustacea
Decapods
Crab zoea 2
Spiny lobster phyllosoma
Zooplankton
Phylum Echinodermata (sea stars, urchins, cucumbers, etc.)
• hard calcified endoskeleton• pentaradial symmetry• marine only• in plankton usually only as larvae
brittle star pluteus
coral planula larva
sea star bipinnaria, branchiolaria
Phylum Echinodermata
swimming sea cucumber,Enypniastes eximia
http://www.whoi.edu/cms/images/media2-2002-214-Enypniastes_eximia_49415.jpg
Zooplankton
Phylum Hemichordata (acorn worms)
• wormlike, with 3 body parts, with collar being middle section• pharyngeal gill slits• unsegmented• larva: tornaria
tornaria larva
adult
Zooplankton
Phylum Urochordata
• primitive chordates• notochord and dorsal hollow neural tube
Class Ascidacea (=Tunicata) (sea squirts) • adults benthic• (tadpole larva) – has notochord and dorsal hollow neural tube
tadpole larva
Molgula adults
Sea squirts, Molgula manhattensis, on lantern nets used for growing bay scallops
S. Tettelbach
Zooplankton
Phylum Urochordata Class Thaliacea (salps)
• holoplankton• no larval form• gelatinous• may form colonies >2 m long
solitary adult
A string of salps in the Red Sea. www.itsnature.org/sea/other/salps/
Phylum Urochordata Class Thaliacea (salps)
Zooplankton
Phylum Urochordata Class Larvacea = (Appendicularia)
• gelatinous house with filter feeding animal inside
Zooplankton
Phylum Chordata Class Vertebrata
fish larvae
eel leptocephalus larva
THE PHOTOGRAPHIC WEB GUIDE TO THE LARVAE OF
CORAL REEF FISHES:THE 21st CENTURY
Benjamin C. Victor
www.coralreeffish.com/larvae.html
Late Stage Coral Reef Fish Larvae Collected at Night
Zooplankton
Phylum Chordata Class Vertebrata
adult Mola
king mackerel larvae