Plankton.Lecture.1
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PLANKTON(Bio 511)
Professor Stephen T. Tettelbach
C.W. Post Campus of Long Island University
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PLANKTON
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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!
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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)
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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)
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Size classes of the plankton
(from Johnson & Allen (2005)
see handout
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Other definitions sometimes used:
ultraplankton: < 2 µm = femtoplankton + picoplankton
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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
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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
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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
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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)
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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
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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
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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)
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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
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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.
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Protozooplankton - Foraminifera
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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
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Protozooplankton - radiolarians
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protozooplankton
Phylum Sarcomastigophora Subphylum Sarcodina
Acantharia - shells of strontium sulfate
Lychnaspis miranda
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protozooplankton
Phylum Sarcomastigophora Subphylum Sarcodina
Heliozoans – marine, brackish, but 1° fw. Some have silica
Actinosphaerium
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protozooplankton
Phylum Sarcomastigophora Subphylum Mastigophora (flagellates)
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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]
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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.
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Phytoplankton – Diatoms
2 major forms: centric – rounded; essentially radially symmetric
Coscinodiscus
Dityum brightwelli
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Phytoplankton – Diatoms
Chains of centric diatoms
Thalassionema nitzschioides
Eucampia zodiacus
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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
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Phytoplankton – Diatoms
Chains of centric diatoms
Chaetoceros affinis
Actinoptychis
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Phytoplankton – chain of Antarctic centric diatoms
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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
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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)
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Phytoplankton – Dinoflagellates
2 main taxonomic groups, based on where flagella insert
desmokonts: 2 flagella arise from anterior part of cell
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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
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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
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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
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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
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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
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Phytoplankton – Euglenophytes
Kingdom Protista Division Euglenophyta
•unicellular flagellates, with 1 very long and 1 very short flagellum
• 1° freshwater
Assorted euglenoids
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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
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Phytoplankton – Prymnesiophytes
Kingdom Protista Division Prymnesiophyta coccolithophorids
(have CaCO3 scales called coccoliths)
Scales can reflect light so water appears whitish during blooms
Emiliana huxleyi
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Coccolithophorid blooms in the Celtic Sea and Gulf of Biscay
www.co2.ulg.ac.be/objects/intro_cocco.htm
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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
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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
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Zooplankton
Phylum Cnidaria Class Hydrozoa
actinula larva
hydromedusa: Maeotias
hydromedusa: Benthocodon (deepwater)
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Hydromedusa Colobonema sericeum
Phylum Cnidaria Class Hydrozoa
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Zooplankton
Phylum Cnidaria Class Hydrozoa
Physalia (Portuguese Man O’War) Vellela (by-the-wind sailor)
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Phylum Cnidaria Class Hydrozoa
Physalia
(Portuguese Man O’War)
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Zooplankton
Phylum Cnidaria Class Scyphozoa
Stomolophus
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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
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Zooplankton
Phylum Cnidaria Class Cubozoa
Chironex flexneri (sea wasp)
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Zooplankton
Phylum Cnidaria Class Anthozoa
tube anemone larva
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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
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animaldiversity.ummz.umich.edu/.../view.html
www.huriisgaard.biology.sdu.dk/Mnemiopsis%209...
Phylum Ctenophora: Mnemiopsis leidyi (sea walnut)
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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
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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
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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.
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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
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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
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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
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Swarming polychaete epitokes at night, Glover's Reef, Belize
www.ryanphotographic.com/epitoke.htm
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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
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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
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Zooplankton – adult holoplanktonic gastropods
pteropod (sea butterfly)
Glaucus glaucus
hydrozoan Porpita porpita
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Zooplankton – adult holoplanktonic gastropods
heteropod
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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
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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.
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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
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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
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Zooplankton
Phylum Arthropoda Subphylum Crustacea
Ostracods
• bivalved shells (mostly opaque)• small (<2 mm)• many benthic, some holoplanktonic
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Zooplankton
Phylum Arthropoda Subphylum Crustacea
Cladocerans (4 orders)
• most with calcareous shell (usually clear)• mostly fw• holoplankton
Polyphemus
Daphnia pulex
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Zooplankton
Phylum Arthropoda Subphylum Crustacea
barnacles (Cirripedia)
• meroplankton only• larvae: nauplius, then cypris (cyprid)
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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
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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
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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
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Euphausids (krill) comprise the greatest biomass of any animal in the ocean
photography.nationalgeographic.com/photograph...
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Zooplankton
Phylum Arthropoda Subphylum Crustacea
Decapods
Crab zoea 2
Spiny lobster phyllosoma
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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
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Phylum Echinodermata
swimming sea cucumber,Enypniastes eximia
http://www.whoi.edu/cms/images/media2-2002-214-Enypniastes_eximia_49415.jpg
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Zooplankton
Phylum Hemichordata (acorn worms)
• wormlike, with 3 body parts, with collar being middle section• pharyngeal gill slits• unsegmented• larva: tornaria
tornaria larva
adult
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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
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Sea squirts, Molgula manhattensis, on lantern nets used for growing bay scallops
S. Tettelbach
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Zooplankton
Phylum Urochordata Class Thaliacea (salps)
• holoplankton• no larval form• gelatinous• may form colonies >2 m long
solitary adult
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A string of salps in the Red Sea. www.itsnature.org/sea/other/salps/
Phylum Urochordata Class Thaliacea (salps)
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Zooplankton
Phylum Urochordata Class Larvacea = (Appendicularia)
• gelatinous house with filter feeding animal inside
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Zooplankton
Phylum Chordata Class Vertebrata
fish larvae
eel leptocephalus larva
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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
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Zooplankton
Phylum Chordata Class Vertebrata
adult Mola
king mackerel larvae