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Smaller Ecdysozoans

Chapter 18

Ecdysozoans
Many protostomes

possess a cuticle.
Non-living outer
layer secreted by
epidermis.

Cuticle restricts

growth and must be


molted via ecdysis.
Regulation of
molting achieved
by the hormone
ecdysone.
Members of Ecdysozoa molt cuticle as they grow.

Ecdysozoan Phyla
Phylum Nematoda
Phylum Nematomorpha
Phylum Kinorhyncha
Phylum Loricifera
Phylum Priapulida

All have a nonliving cuticle that is shed as the organism


grows.

Diversity
Ecdysozoans do not share the same body
plan.
Members of Nematoda, Nematomorpha, and
Kinorhyncha are pseudocoelomate.
Members of Priapulida are presumed to be
pseudocoelomate, but have not been carefully
studied yet.
Loricifera species may be pseudocoelomate or
acoelomate.

Phylum Nematoda: Roundworms


About 25,000 species of nematodes are
described, but as many as half a million may
exist.
Many prefer the name Nemata for this phylum.

Found in virtually all habitats in all biomes.


Topsoil may contain billions per acre.

Phylum Nematoda: Roundworms


Free-living nematodes

feed on bacteria, yeasts,


fungal hyphae, and
algae.

Predatory nematodes

eat rotifers, tardigrades,


small annelids, and
other nematodes.

Important as food for

mites, insects, larvae,


and fungi.

Phylum Nematoda: Roundworms


Caenorhabditis
elegans is an
important model for
studies of genomics
and cell
development and
differentiation.

Phylum Nematoda
Some species
of nematodes
are important
parasites of
plants and
animals.

Nematode

parasites exist
in nearly all
animal and
plant species.

Phylum Nematoda
Triploblastic three embryonic germ layers.
Pseudocoel used as hydrostatic skeleton.
Cylindrical shape
Lack of motile cilia or flagella
Nonsegmented
Nonliving cuticle

Phylum Nematoda
The cylindrical bodies of nematodes are
covered by a tough coat called a cuticle.

Phylum Nematoda
Complete digestive system.
Circulatory and respiratory systems are
lacking.

Ring of nerve tissue and ganglia around the


pharynx lead to dorsal and ventral nerve cords.

Phylum Nematoda
Nematodes have separate sexes.
Dioecious, females are larger.
Fertilization is internal.
Eggs are stored in uterus until deposited.
Cuticle is shed between each of four juvenile stages.

Phylum Nematoda: Roundworms


Sensory papillae at
head and tail.

Amphids, pair of
sensory organs on
head, lead into a
deep cuticular pit
with modified cilia.

Phylum Nematoda
Some larvae are free-

living, some require and


intermediate host.

Parasitic nematodes of

humans:
Intestinal roundworm
(Ascaris)
Hookworm
Pinworm
Trichina worm
Whipworm

Phylum Nematomorpha
Phylum

Nematomorpha, the
horsehair worms, are free
living as adults, and
parasites in arthropods
as juveniles.
Currently placed as the
sister taxon to
nematodes
About 320 species are
known.
Occur worldwide
Pseudocoelomate
Triploblastic

Phylum Nematomorpha
Horsehair worms leaving the
body of a katydid found dead
near a river in Illinois.
Photos by Bryan Suson

Phylum Nematomorpha
Adults utilize stored nutrients.
Recent studies reveal that they can absorb

organic molecules through vestigial gut and


body wall.

Circulatory, respiratory and excretory systems


are absent.

Nematomorphs are dioecious.


Females discharge eggs into water in long
strings.

Phylum Nematomorpha
Juveniles may encyst on vegetation and are

eaten by an arthropod such as a grasshopper.

Larval stages have hooks that may be used to


bore into a host.

May also cause infection via drinking water.


Larvae encyst within host.
Marine nematomorphs infect hermit crabs and
other crabs.

Phylum Nematomorpha
After months in an arthropod host, mature
worm emerges into nearby water or during
rainfall.

Parasite stimulates terrestrial insects to seek


water.

Phylum Kinorhyncha
Kinorhynchs are usually under 1 mm long.
About 179 species are known.
Found worldwide, from intertidal areas to 6000
m deep.

Most live in mud, but some have been found in


algae, sponges, and other invertebrates.

Phylum Kinorhyncha
Spines (scalids) function
in locomotion,
chemoreception, and
mechanoreception.

Retractile head has a


circle of spines and a
retractile proboscis
(introvert).

Body wall made of a


cuticle and syncytial
epidermis.

Phylum Priapulida
The phylum Priapulida
contains 16 species of
marine worms that
occur in colder waters.

Found from intertidal


zones to deep ocean
floors, several thousand
meters deep.

Some are tube dwellers


and feed on detritus.

Phylum Priapulida
Cylindrical bodies under 15 cm
long.

Burrow by body contractions and


orient mouth at the surface.

Retractable introvert has papillae


and rows of curved spines to
sample and capture prey.

Chitinous cuticle covers body and


is molted regularly.

Sexes are separate.

Phylum Loricifera
Nine circlets of scalids on

the introvert, similar to those


of kinorhynchs.

Entire forepart can be

retracted into the circular


lorica.

Diet is unknown, possibly


feed on bacteria.

Brain fills the head and

nerves innervate scalids.

Dioecious with dimorphic


males and females.

Body cavity is a pseudocoel.

Clade Panarthropoda
Clade Panarthropoda contains Arthropoda
and two allied phyla, Onychophora and
Tardigrada.

In onychophorans and arthropods coelom develops


by schizocoely, but enterocoelic in tardigrades.
A new cavity, hemocoel, forms from fusion of the
main coelomic cavity with the blastocoel.
Blood from open circulatory system enters the
hemocoel and surrounds the internal organs.
Contains a muscular heart but limited muscular
blood vessels.

Phylum Onychophora
About 70 living species of velvet worms in the
phylum Onychophora.

1.4 to 15 cm in length.
Limited to moist, leafy rain forest habitat in tropical
and subtropical regions.

Changed little over 500 million years.


Fossil Aysheaia similar to modern forms.

Share traits with annelids and arthropods and were


considered a missing link.

Most are predaceous, some live in termite nests.

Phylum Onychophora
No external segmentation
except for paired appendages.
Skin is soft - cuticle contains
protein and chitin but does not
harden as in arthropods.
Body covered with tiny
tubercles bearing sensory
bristles.
Minute scales on tubercles
impart an iridescent and
velvety appearance.
Head has one pair of large
antennae and an annelid-like
eye at base.

Phylum Onychophora
14 to 43 pairs of unjointed legs.
Legs move by waves of body
contractions.

Slime glands on each side of body


cavity open on oral papillae.

Mouth leads to a straight digestive


tract.

Each segment contains a pair of

nephridia, each containing a


vesicle, ciliated funnel, and duct.

Phylum Onychophora
Tracheal system provides respiration to all
parts of body.

Cannot close spiracles, so they are restricted


to moist habitats.

Important differences suggest the tracheal


system evolved independently from that of
arthropods.

Phylum Onychophora
Open circulatory system.
Nervous system organized much like a ladder.
Sense organs include pigment cup ocelli, taste spines,
tactile papillae, and hygroscopic receptors that orient
the animal toward water vapor.

Phylum Onychophora
With exception of one parthenogenetic species, all are
dioecious with paired reproductive organs.
Males deposit spermatophores on back of female.
White blood cells dissolve the skin and sperm
migrate to ovaries.

May be oviparous, ovoviviparous, or viviparous.

Phylum Tardigrada
Tardigrades, or

water bears, are


less than a
millimeter in
length.
Freshwater or
marine
Live in spaces
between sand
grains.

http://www.youtube.com/watch?v=W9WWysGBAlU&feature=related

http://www.youtube.com/watch?v=blX7Ef1gxcM&feature=related

Phylum Tardigrada
Trunk bears four pairs of short, stubby,

unjointed legs, each with four to eight claws.

Body covered by non-chitinous cuticle that is


molted four or more times during lifetime.

Mouth leads to a muscular pharynx adapted for


sucking.

Two stylets protrude to pierce integument of


nematodes or walls of plant cells and allow
them to suck juices.

Phylum Tardigrada
They share many characteristics with
arthropods.
But legs are unjointed.
Non-chitinous cuticle that is molted.

Phylum Tardigrada
No circulatory or respiratory systems.
Gas exchange occurs at body surface.
Muscles are all longitudinal.
Uses hydrostatic pressure as a skeleton.
Brain is relatively large.

Phylum Tardigrada
Sexes are separate.
In parthenogenetic
freshwater and
moss-dwelling
species, males are
unknown.

Egg-laying, like

defecation, occurs at
molting.
Eggs may be highly
ornate.

Phylum Tardigrada
Tardigrades can enter a state called
cryptobiosis, where metabolism is
imperceptible.

Tardigrades can dehydrate from 85% water to only

3% water.
In this state they can resist extreme temperatures,
ionizing radiation, oxygen deficiency, etc. for years.
When water is available, they become metabolically
active again.

Phylogeny
Evolutionary relationships among ecdysozoans
are not well-understood.

Members of this clade do not share a common


cleavage pattern:
Nematodes and nematomorphs - cleavage is unique,
not spiral or radial.
Cleavage in kinorhynchs, lorificiferans, and
tardigrades has yet to be studied.
Priapulid cleavage is nearly radial.

Phylogeny
Recent phylogenies place Nematoda and

Nematomorpha as sister taxa since they share a


collagenous cuticle.

Phylum Kinorhyncha and Priapulida are considered

sister taxa because they share a two-layered pharynx.

Velvet worms, phylum Onychophora, are the sister


taxon to tardigrades.

Phylogeny
Onychophorans share a number of
characteristics with annelids:

Metamerically arranged nephridia, muscular body


wall, pigment cup ocelli, and ciliated reproductive
ducts.

Onychophorans also share features with the


arthropods:

Including a cuticle, tubular heart, and hemocoel with


open circulatory system, presence of tracheae
(possibly not homologous), and large brain.

Phylogeny
Sequence analysis supports placement of
Onychophora in clade Panarthropoda.

Tardigrades and arthropods also share


arthropod-type setae and muscles inserted on
the cuticle.

Adaptive Diversification
Nematodes show impressive adaptation they
are found in almost every habitat available to
animals.

Body structure is plastic enough to allow


adaptation.

Life cycle ranges from simple to complex.


Have been known to survive in suboptimal
conditions.

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