class malacostraca - crabs, shrimps, lobsters class maxillopoda - ostracods, barnacles, copepods...

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s Malacostraca - crabs, shrimps, lobsters s Maxillopoda - ostracods, barnacles, copepods s Branchiopoda - brine shrimp, tadpole shrimp s Remipedia - small cave-dwellers; basal lineage s Cephalocarida - small benthic detritivores; basal SubPhylum Crustacea

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Class Malacostraca - crabs, shrimps, lobsters

Class Maxillopoda - ostracods, barnacles, copepods

Class Branchiopoda - brine shrimp, tadpole shrimp

Class Remipedia - small cave-dwellers; basal lineage

Class Cephalocarida - small benthic detritivores; basal

SubPhylum Crustacea

Class Malacostraca SuperOrder Peracarida

- carapace reduced in size

- unique endites called oostegites (brood pouch, or marsupium)

- direct development, which helped colonize land: bypass a planktonic larval stage, mother broods young & releases fully developed juveniles

Order Mysida (opossum shrimp)

Order Isopoda (isopods) made it onto land

Order Amphipoda (amphipods)

Class Malacostraca SuperOrder Peracarida

Order Mysida (opposum shrimp)

- well developed carapace covering head+thorax segments

- pelagic, intertidal, or burrowing

- often form benthic “swarms,” which are small clouds of tiny shrimp that hide among urchin spines for protection

brood pouch where young develop

Class Malacostraca SuperOrder Peracarida

Order Isopoda >10,000 species

- no carapace: you can see each segment from above- one lineage successfully colonized dry land: the pill bugs- pereopods are very structurally variable- pleopods used in gas exchange

Cirolana

rock lousekelp isopod

Class Malacostraca SuperOrder Peracarida

Order Amphipoda 8,000 species

- no carapace

- includes semi-terrestrial “beach hoppers”

- thoracic gills are epipods, extensions of pereopods

Decapods typically brood their larvae to a well developed, large-eyed zoea larva capable of fast swimming and active behaviors, which can help larvae migrate in & out of estuaries

After several molts, the final megalopa larva is produced, which looks very much like the adult - except in crabs, the abdomen is not yet tucked under the thorax

Decapod Larvae

crabzoea

stomatopod(mantis shrimp)megalopa

Class Malacostraca - crabs, shrimps, lobsters

Class Maxillopoda - ostracods, barnacles, copepods

Class Branchiopoda - brine shrimp, tadpole shrimp

Class Remipedia - small cave-dwellers; basal lineage

Class Cephalocarida - small benthic detritivores; basal

SubPhylum Crustacea

barnacles, copepods, ostracods

Class Maxillopoda

Head (5 segments)Thorax (6)Abdomen (4)

- usually biramous limbs; no abdominal appendages

- mostly small; reduced abdomen, missing some legs

Subclasses: Thecostraca (barnacles) Copepoda Ostracoda

26,000 spp.

free-living (acorn, gooseneck)

parasitic on crustaceans

Class Maxillopoda: SubClass Thecostraca

Free-living barnacles Parasitic Rhizocephalans- hermaphrodites - separate sexes- carapace produces shell - carapace lost- suspension feeders - endoparasites w/ interna

and externa

Capitulum

Cirri

Peduncle

ScutumTergum

ExternaRootlets oframifying body

- in adult barnacles, the appendages on the thorax (= thoracic legs, or cirri) are similar in appearance to tentacles of filter-feeding worms and crinoids (to come later)

- used to filter feed: scutum and tergum plates open in response to water current, cirri extended up into water, then rapidly retracted inside shell over and over again

Antennule

Antenna

Mandible

Naupliareye

Nauplius Larval Stage

antennules, antennae, mandibles; no segmentation; single eye- successive molts: remaining head, thoracic appendages

Unique feature of barnacles: non-feeding cyprid larval stage which follows the nauplius stage

- has a bivalved shell, and swims using antennae

- in free-living barnacles, cyprid glues its head onto the first suitable hard surface it encounters

Cyprid Larvae

Cyprid attaching, cementing itself head-first to substrate

become cirri of adult

- cephalic “shield”- keep single naupliar eye- primary consumer of phytoplankton- tiny; swim by flicking antennae

Class Maxillopoda: SubClass Copepoda - 8,500 spp.

Caudal rami

Ovisac(clump of embryos)

articulation point(attachment of thorax to abdomen)

Antennule

Antenna

Naupliar eye

Class Maxillopoda: SubClass Copepoda - 8,500 spp.

In males, 1st antennae and 5th thorax limb may be specially modified for “mate guarding” behaviors

- males locate, grab and ride around holding a virgin female before her terminal (final) molt

- this way, they ensure they are the male who will fertilize her eggs when she molts, and is then ready to mate

Class Maxillopoda: SubClass Ostracoda - 6,600 spp.

- bivalved carapace (how many independent origins of a bivalved shell have we seen now??..)

- reduced segmentation; body not split into thorax + abdomen

- 5 head appendages + 1-3 thoracic appendages, including male copulatory limb-- fewest limbs of any crustacean

- many are bioluminescent, using light flashes in complex mating rituals

- swim using antennae

Class Malacostraca - Crabs, shrimps, lobsters

Class Maxillopoda - Ostracods, barnacles, copepods

Class Branchiopoda - Brine shrimp, tadpole shrimp

Class Remipedia - small cave-dwellers; basal lineage

Class Cephalocarida - small benthic detritivores; basal

SubPhylum Crustacea

Class Branchiopoda

Notostracans: tadpole shrimp Anostracans: fairy shrimp

Artemia,brine shrimp

no carapace

telson

carapace

caudalramus

often inhabit freshwater vernal pools (temporary)

Sow bugs (terrestrial isopods) are most successful group

- direct development cut ties to the sea

Crustaceans on Land

Land crabs must still return to the ocean to spawn

1. 2.

3. Insects – molecular evidence indicates the most successful animal lineage evolved from a crustacean ancestor

Regier et al. 2010, Nature

Analysis of 62 genes redefined arthropod relationships

(1) Insects are nested within the“Crustacea”, which is therefore a paraphyletic group!

New name, “Pancrustacea”, refers to crustaceans + insects

Regier et al. 2010, Nature

(1) Insects are nested within the“Crustacea”, which is therefore a paraphyletic group!

2) Maxillopoda not a true group

- barnacles, copepods & ostracods do not form a clade

- despite sharing the same “formula” for body segments, these are all unrelated groups

Regier et al. 2010, Nature

Insects are nested within the“crustacea”, which is therefore a paraphyletic group!

2) Maxillopoda not a true group

2a – barnacles are sister to Malacostraca (crabs, etc)

2b – copepods are sister to (Malacostraca + barnacles)

2c – ostracods are basal to rest of Crustacea

Tardigrades

Onychophorans

Arthropods

Ecdysozoa

Priapulids

Kinorhynchs

Loriciferans

Nematodes

Nematomorphs

- clade Ecdysozoa was named in 1997 from an analysis of 18S ribosomal RNA gene

- members all molt to grow, controlled by hormone ecdysone

- microscopic 8-legged “water bears” from semi-aquatic habitats prone to drying out (e.g. moss)

- display extreme cryptobiosis when environments dry out: the ability to enter a state of suspended animation, with no metabolism or apparent ageing, until conditions improve

- 4 pairs of unjointed legs

- uncalcified cuticle lines parts of gut

- main cavity = hemocoel; no gills, respire across wet body wall

Phylum Tardigrada - “water bears”3 Orders, 10 Families ~ 800 species

Cryptobiosis in Tardigrades

During environmental stress, legs are pulled and body coated in a double-walled cuticular envelope, forming a cyst

Then, a single-walled tun state develops, where metabolism is undetectable and individuals can survive long periods of dessication (= dryness)

Tuns have survived… - up to 10 years without water - complete vacuum, long periods with no oxygen - temperatures near absolute zero: 8 hr at -272°C !! - toxic liquids (ether, 100% alcohol)

Allows tardigrades to persist in an unstable ecological niche

styletused topunchholesin plantsor algae;then tardigrade sucks out cytoplsm

mostly feed on plants, algae

convergent evolution: very similar to the single tooth and method of feeding in sacoglossan sea slugs (my group, the herbivores)

Tardigrades

Onychophorans

Arthropods

Ecdysozoa

Priapulids

Kinorhynchs

Loriciferans

Nematodes

Nematomorphs

clade Cycloneuralia– brain forms a loop around esophagus– anterior end = introvert, which can be withdrawn into body

Cycloneuralia

Tardigrades

Onychophorans

Arthropods

Ecdysozoa

Priapulids

Kinorhynchs

Loriciferans

Nematodes

Nematomorphs

clade Cephalorhyncha

– spiny collar of scalids around the introvert

Cephalorhyncha

Phylum Nematoda ~25,000species

- body round in cross-section, but no circular muscles; longitudinal muscles let them thrash around to move

- covered in cuticle, shed in 4 molts during juvenile growth allows survival in hostile environments

- extensions of muscles called muscle arms branch out to contact neurons, instead of other way ‘round like most animals

- unique excretory system featuring renette cells; no circulatory structures

Includes model organism for developmental biology, C. elegans

Phylum Nematoda

Ecological importance:

- tremendously abundant decomposers - 200 per ml of coastal mud - 90,000 per rotting apple - 9 billion per acre of soil

Medical importance: - cuticle allows them to resist immune system, function as endoparasites (endo = living inside the host’s body)

- parasitic forms cause diseases including river blindness, elephantiasis, trichinosis

Nematomorphs – “horsehair worms”

- long thin body encased in well-developed cuticle

- over 1 meter long, less than 1 mm wide

- no apparent segmentation

- all growth accomplished by larvae, which are parasites in

arthropod hosts

- gut, but no mouth; nutrients are

absorbed from insect host across

body wall, taken up by gut tissue

~320species

Priapulids - benthic marine “worms”

- introvert with hooked spines

- complete gut, protonephridia for excretion

- no circulatory system

- thin cuticle surrounds body

- loricate larva has special cuticle that’s

shed at metamorphosis

18 species

Introvert

Trunkrings

Caudal appendages

Phylum Kinorhyncha- all marine; interstitial = live between grains of mud or sand

- body with 13 segments: head, neck, and 11-segment trunk

- no cilia; crawl by extending head, locking spines in place

- head = introvert; can retract into neck

- ring of backward-facing spines (scalids) used to lock body in

place when crawling

convergent evolution with how annelids use chetae to dig

- well-developed cuticle

- no larval stage!

~ 180 species

Phylum Kinorhyncha

Phylum Loricifera- all marine; only discovered in 1983!

- body tiny (< 0.5 mm) but complex (>10,000 cells)

- head with piercing stylets around mouth

- head, neck + thorax retract into abdomen

- 9 rings of spines called scalids

- complete gut, but no circulatory system

or nephridia

~ 100 species

Phylum Loricifera

Head

Thorax

Abdomen