Zooplankton

Planktos: “drifts” in greek Their distribution depends on currents and gyres Certain zooplankton can swim well, but distribution controlled by current patterns Zooplankton: all heterotrophic plankton except bacteria and viruses; size range from 2 µm (heterotrophic flagellates, protists) up to several meters (jellyfish)

Herbivorous zooplankton: Grazers

Nutritional modes in zooplankton Herbivores: feed primarily on phytoplankton Carnivores: feed primarily on other zooplankton (animals) Detrivores: feed primarily on dead organic matter (detritus) Omnivores: feed on mixed diet of plants and animals and detritus

Feeding modes in Zooplankton Filter feeders Predators – catch individual particles

Filter Feeder Copepod

Filter Feeder Ctenophore

Predator Chaetognath Arrow Worm

Life cycles in Zooplankton Holoplankton: spend entire life in the water column (pelagic) Meroplankton: spend only part of their life in the pelagic environment, mostly larval forms of invertebrates and fish Ichthyoplankton: fish eggs and fish larvae

Holoplankton Copepods Planktonic crustaceans

Barnacles: benthic sessile crustacean

Meroplankton Nauplius larva

Meroplankton Cypris larva

Cypris larva and metamorphosed juveniles

Barnacle population regulation

Gadidae Gadus morhua Ichthyoplankton

Gadidae Gadus morhua Ichthyoplankton

Gadidae Gadus morhua Ichthyoplankton

Gadidae Atlantic cod Gadus morhua Demersal Adult

Protists: Protozooplankton Dinoflagellates: heterotrophic relatives to the phototrophic Dinophyceae; naked and thecate forms. Noctiluca miliaris – up to 1 mm or bigger, bioluminescence, prey on fish egg & zooplankton Zooflagellates: heterotrophic nanoflagellates (HNF): taxonomically mixed group of small, naked flagellates, feed on bacteria and small phytoplankton; choanoflagellates: collar around flagella Foraminifera: relatives of amoeba with calcareous shell, which is composed of a series of chambers; contribute to ooze sediments; 30 µm to 1-2 mm, bacteriovores; most abundant 40°N – 40°S

Dinoflagellates Noctiluca miliaris

Colonial choanoflagellates Bacteriofages (Ross Sea)

Foraminifera (calcareous – all latitudes)

Radiolaria: spherical, amoeboid cells with silica capsule; 50 µm to several mm; contribute to silica ooze sediments, feed on bacteria, small phyto- and zooplankton; cold water and deep-sea Ciliates: feed on bacteria, phytoplankton, HNF; naked forms more abundant but hard to study (delicate!); tintinnids: sub-group of ciliates with vase-like external shell made of protein; herbivores Protists: Protozooplankton

Figure 3.21b Radiolarians (siliceous – low latitudes)

Live Radiolarian

Cnidaria: primitive group of metazoans; some holoplanktonic, others have benthis stages; carnivorous (crustaceans, fish); long tentacles carry nematocysts used to inject venoms into prey; box jellyfish of Australia kills humans within minutes –Medusae: single organisms, few mm to several meters –Siphonophores: colonies of animals with specialization: feeding polyps, reproductive polyps, swimming polyps; Physalia physalis (Portuguese man-of-war), common in tropical waters, Gulf of Mexico, drifted by the wind and belong to the pleuston (live on top of water surface) Invertebrate Holoplankton

Cnidaria (medusae)

Cnidaria (siphonophora)

Ctenophores: separate phylum, do not belong to Cnidaria; transparent organisms, swimm with fused cilia; no nematocysts; prey on zooplankton, fish eggs, sometimes small fish; important to fisheries due to grazing on fish eggs and competition for fish food Chaetognaths: arrow worms, carnivorous, <4 cm Polychaets: Tomopteris spp. only important planktonic genus Invertebrate Holoplankton

Ctenophora (comb jellies)

Invertebrate Holoplankton Mollusca: –Heteropods: small group of pelagic relatives of snails, snail foot developed into a single “fin”; good eyes, visual predators –Pteropods: snail foot developed into paired “wings”; suspension feeder – produce large mucous nets to capture prey; carbonate shells produce pteropod ooze on sea floor

Heteropod (Predates on Ctenophores)

Pteropod

Protochordate Holoplankton Appendicularia: group of Chordata, live in gelatinous balloons (house) that are periodically abandoned; empty houses provide valuable carbon source for bacteria and help to form marine snow; filter feeders of nanoplankton Salps or Tunicates: group of Chordata, mostly warm water; typically barrel-form, filter feeders; occur in swarms, which can wipe the water clean of nanoplankton; large fecal bands, transport of nano- and picoplankton to deep-sea; single or colonies

Appendicularia

Pelagic Salps

Arthropoda: crustacean zooplankton Cladocera (water fleas): six marine species (Podon spp., Evadne spp.), one brackish water species in the Baltic Sea; fast reproduction by parthenogenesis (without males and egg fertilization) and pedogenesis (young embryos initiate parthenogenetic reproduction before hatching) Amphipoda: less abundant in pelagic environment, common genus Themisto; frequently found on siphonophores, medusae, ctenophores, salps Euphausiida: krill; mm, pronounced vertical migration; not plankton sensu strictu; visual predators, fast swimmers, often undersampled because they escape plankton nets; important as prey for commercial fish (herring, mackerel, salmon, tuna) and whales (Antarctica)

Amphipoda

Amphipoda (parasites of gelatinous plankton)

Euphasids (krill)

Arthropoda: crustacean zooplankton Copepoda: most abundant zooplankton in the oceans, “insects of the sea“; herbivorous, carnivorous and omnivorous species –Calanoida: most of marine planktonic species –Cyclopoida: most of freshwater planktonic species –Harpacticoida: mostly benthic/near-bottom species Copepod development: first six larval stages = nauplius (pl. nauplii), followed by six copepodit stages (CI to CVI) Tropical species distinct by their long antennae and setae on antennae and legs (podi)

Copepods

Mollusca: clams and snails produce shelled veliger larvae; ciliated velum serves for locomotion and food collection Cirripedia: barnacles produce nauplii, which turn to cypris Echinodermata: sea urchins, starfish and sea cucumber produce pluteus larvae of different shapes, which turn into brachiolaria larvae (starfish); metamorphosis to adult is very complex Polychaeta: brittle worms and other worms produce trochophora larvae, mostly barrel- shaped with several bands of cilia Common Meroplankton

Decapoda: shrimps and crabs produce zoëa larvae; they turn into megalopa larvae in crabs before settling to the sea floor Pisces: fish eggs and larvae referred to as ichthyoplankton; fish larvae retain part of the egg yolk in a sack below their body until mouth and stomach are fully developed Common Meroplankton

Meroplankton

Meroplanktonic Larvae Planktotrophic –Feeding larvae –Longer Planktonic Duration Times –High dispersal potential Lecithotrophic (non-feeding) –Non-feeding larvae –Shorter planktonic Duration Times –Low dispersal potential

Molluscs: Meroplankonic Veliger larvae PLANKTOTROPHIC

Vertical Distribution Epipelagic: upper m water column; high diversity, mostly small and transparent organisms; many herbivores Mesopelagic = 300 – 1000 m; larger than epipelagic relatives; large forms of gelatinous zooplankton (jellyfish, appendicularians) due to lack of wave action; some larger species (krill) partly herbivorous with nightly migration into epipelagic regimes Oxygen Minimum Zone: 400 – 800 m depth, accumulation of fecal material due to density gradient, attract high bacterial growth, which in turn attracts many bacterial and larger grazers; strong respiration reduces O2 content from 4-6 mg l-1 to < 2 mg l-1 Bathypelagic: 1000 – 3000 m depth, many dark red colored, smaller eyes Abyssopelagic: > 3000 m depth, low diversity and low abundance Demersal or epibenthic: live near or temporarily on the seafloor; mostly crustaceans (shrimp and mysids) and fish

Diel Vertical Migration DAILY (diel) vertical migrations over distances of 800 m –Nocturnal: single daily ascent beginning at sunset, and single daily descent beginning at sunrise –Twilight: two ascents and descents per day (one each assoc. with each twilight period) –Reversed: single ascent to surface during day, and descent to max. depth during night

Scattering Layer

Horizontal distribution: patchiness

Exotic Planktonic species New England Ctenophore  Black Sea

Water Tank Ballast Holoplankton Meroplankton

Black Sea Ballast Invasions Mnemiopsis

Black Sea Ballast Invasions Mnemiopsis Beroe ovata

European Green Crab – Carcinus maenas