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