28.4: Superphylum Ecdysozoa

28.4.1: Superphylum Ecdysozoa

Learning Objective

Discuss the phylogenetic position of Ecdysozoa

Key Points

• The Ecdysozoans are the most diverse group of animals, containing the nematode worms and the arthropods.
• These organisms have an external covering called a cuticle that protects their soft internal organs from water loss and the outside environment.
• After they molt, or shed their cuticle, they grow in size and secrete a new shell; this is called ecdysis.
• The phylogeny of the Ecdysozoans has been the cause of much scientific debate with no definitive consensus in the scientific community.

Key Terms

coelomate

any animal possessing a fluid-filled cavity within which the digestive system is suspended.

cuticle

a noncellular protective covering outside the epidermis of many invertebrates and plants

ecdysis

the shedding of an outer layer of skin in snakes, crustaceans and insects; moulting

Superphylum Ecdysozoa

The superphylum Ecdysozoa contains an incredibly large number of species. This is because it includes two of the most diverse animal groups: Phylum Nematoda (the roundworms) and Phylum Arthropoda (the arthropods). The most distinguishing and prominent feature of Ecdysozoans is their cuticle: a tough, but flexible exoskeleton that protects these animals from water loss, predators, and other aspects of the external environment. All members of this superphylum periodically molt or shed their cuticle as they grow . After molting, they secrete a new cuticle that will last until their next growth phase. The process of molting and replacing the cuticle is called ecdysis, which is the derivation of the superphylum's name.

Molting in arthropods

This cicada is in the middle of the molting process. The old cuticle splits and the insect climbs out. At this time, the insect's body is very soft. The cicada will then eat the old shell to replace nutrients that would otherwise be lost. This encourages the new shell to harden.

Phylogenetic Hypotheses

There are two main hypotheses about the phylogeny of the Ecdysozoans. The first is called the Articulata hypothesis. This grouping scheme is widely accepted, although some zoologists still hold to the original view that Panarthropoda should be classified with Annelida in a group called the Articulata, and that Ecdysozoa are polyphyletic. Others have suggested that a possible solution is to regard Ecdysozoa as a sister-group of Annelida, though many scientists consider them unrelated. Inclusion of the roundworms within the Ecdysozoa was initially contested, but since 2003, a broad consensus has formed supporting the Ecdysozoa, placing them in a new set of groupings that include the Ecdysozoa, the Lophotrochozoa, and the Deuterostomia.

The other idea about the phylogeny of the Ecdysozoa is called the coelomate hypothesis. Before Ecdysozoa, one of the prevailing theories for the evolution of the bilateral animals was based on the morphology of their body cavities. There were three types, or grades, of organization: the Acoelomata (no coelom), the Pseudocoelomata (partial coelom), and the Eucoelomata (true coelom). With the introduction of molecular phylogenetics, the coelomate hypothesis was abandoned, although some molecular, phylogenetic support for the Coelomata continued until 2005.

28.4.2: Phylum Nematoda

Learning Objective

Describe the features of animals classified in phylum Nematoda

Key Points

• Nematodes are in the same phylogenetic grouping as the arthropods because of the presence of an external cuticle that protects the animal and keeps it from drying out.
• There are an estimated 28,000 species of nematodes, with approximately 16,000 of them being parasitic.
• Nematodes are tubular in shape and are considered pseudocoelomates because of they do not possess a true coelom.
• Nematodes do not have a well-developed excretory system, but do have a complete digestive system.
• Nematodes possess the ability to shed their exoskeleton in order to grow, a process called ecdysis.

Key Term

exoskeleton

a hard outer structure that provides both structure and protection to creatures such as insects, Crustacea, and Nematoda

Phylum Nematoda

The Nematoda, similar to most other animal phyla, are triploblastic, possessing an embryonic mesoderm that is sandwiched between the ectoderm and endoderm. They are also bilaterally symmetrical: a longitudinal section will divide them into right and left sides that are symmetrical. Furthermore, the nematodes, or roundworms, possess a pseudocoelom and have both free-living and parasitic forms.

Both the nematodes and arthropods belong to the superphylum Ecdysozoa that is believed to be a clade consisting of all evolutionary descendants from one common ancestor. The name derives from the word ecdysis, which refers to the shedding, or molting, of the exoskeleton. The phyla in this group have a hard cuticle covering their bodies, which must be periodically shed and replaced for them to increase in size.

Phylum Nematoda includes more than 28,000 species with an estimated 16,000 being parasitic in nature. Nematodes are present in all habitats.

Morphology

In contrast with cnidarians, nematodes show a tubular morphology and circular cross-section. These animals are pseudocoelomates; they have a complete digestive system with a distinct mouth and anus. This is in contrast with the cnidarians where only one opening is present (an incomplete digestive system).

The cuticle of Nematodes is rich in collagen and a carbohydrate-protein polymer called chitin. It forms an external "skeleton" outside the epidermis. The cuticle also lines many of the organs internally, including the pharynx and rectum. The epidermis can be either a single layer of cells or a syncytium, which is a multinucleated cell formed from the fusion of uninucleated cells.

The overall morphology of these worms is cylindrical, while the head is radially symmetrical . A mouth opening is present at the anterior end with three or six lips. Teeth occur in some species in the form of cuticle extensions. Some nematodes may present other external modifications such as rings, head shields, or warts. Rings, however, do not reflect true internal body segmentation. The mouth leads to a muscular pharynx and intestine, which leads to a rectum and anal opening at the posterior end. In addition, the muscles of nematodes differ from those of most animals; they have a longitudinal layer only, which accounts for the whip-like motion of their movement.

Nematode shape

Scanning electron micrograph of soybean cyst nematode and its egg. Nematodes are cylindrical in shape, often looking like thin hairs. They possess an exoskeleton that prevents them from drying out . It must be shed (a process called ecdysis) in order for them to grow.

Excretory System

In nematodes, specialized excretory systems are not well developed. Nitrogenous wastes may be lost by diffusion through the entire body or into the pseudocoelom (body cavity), where they are removed by specialized cells. Regulation of water and salt content of the body is achieved by renette glands, present under the pharynx in marine nematodes.

Nervous system

Most nematodes possess four longitudinal nerve cords that run along the length of the body in dorsal, ventral, and lateral positions. The ventral nerve cord is better developed than the dorsal or lateral cords. All nerve cords fuse at the anterior end, around the pharynx, to form head ganglia, or the "brain" of the worm (taking the form of a ring around the pharynx), as well as at the posterior end to form the tail ganglia. In C. elegans, the nervous system accounts for nearly one-third of the total number of cells in the animal.

Reproduction

Nematodes employ a variety of reproductive strategies that range from monoecious to dioecious to parthenogenic, depending upon the species under consideration. C. elegans is a monoecious species, having development of ova contained in a uterus as well as sperm contained in the spermatheca. The uterus has an external opening known as the vulva. The female genital pore is near the middle of the body, whereas the male's is at the tip. Specialized structures at the tail of the male keep him in place while he deposits sperm with copulatory spicules. Fertilization is internal with embryonic development beginning very soon after fertilization. The embryo is released from the vulva during the gastrulation stage. The embryonic development stage lasts for 14 hours; development then continues through four successive larval stages with ecdysis between each stage (L1, L2, L3, and L4) ultimately leading to the development of a young male or female adult worm. Adverse environmental conditions such as overcrowding and lack of food can result in the formation of an intermediate larval stage known as the dauer larva.

28.4.3: Phylum Arthropoda

Learning Objective

Describe the morphology of arthropoda

Key Points

• Arthropods include the Hexapoda (insects), the Crustacea (lobsters, crabs, and shrimp), the Chelicerata (the spiders and scorpions), and the Myriapoda (the centipedes and millipedes).
• Arthropods have a segmented body plan that contains fused segments divided into regions called tagma.
• Arthropods have an open circulatory system and can use book gills, book lungs, or tracheal tubes for respiration.

Key Terms

spiracle

a pore or opening used (especially by spiders and some fish) for breathing

malpighian tubule

a tubule that extends from the alimentary canal to the exterior of the organism, excreting water and wastes in the form of solid nitrogenous compounds

tagma

a specialized grouping of arthropodan segments, such as the head, the thorax, and the abdomen with a common function

Phylum Arthropoda

The name "arthropoda" means "jointed legs" (in the Greek, "arthros" means "joint" and "podos" means "leg"); it aptly describes the enormous number of invertebrates included in this phylum. Arthropods dominate the animal kingdom with an estimated 85 percent of known species included in this phylum; many arthropods are as yet undocumented. The principal characteristics of all the animals in this phylum are functional segmentation of the body and presence of jointed appendages. Arthropods also show the presence of an exoskeleton made principally of chitin, which is a waterproof, tough polysaccharide. Phylum Arthropoda is the largest phylum in the animal world; insects form the single largest class within this phylum. Arthropods are eucoelomate, protostomic organisms.

Phylum Arthropoda includes animals that have been successful in colonizing terrestrial, aquatic, and aerial habitats. This phylum is further classified into five subphyla: Trilobitomorpha (trilobites, all extinct), Hexapoda (insects and relatives), Myriapoda (millipedes, centipedes, and relatives), Crustaceans (crabs, lobsters, crayfish, isopods, barnacles, and some zooplankton), and Chelicerata (horseshoe crabs, arachnids, scorpions, and daddy longlegs). Trilobites are an extinct group of arthropods found chiefly in the pre-Cambrian Era that are probably most closely related to the Chelicerata. These are identified based on fossil records .

Trilobite fossil

Acadoparadoxides, possibly A. briareus, a large trilobite from about 500 million years ago from Morocco, North Africa (Middle Cambrian)

Morphology

A unique feature of animals in the arthropod phylum is the presence of a segmented body and fusion of sets of segments that give rise to functional body regions called tagma. Tagma may be in the form of a head, thorax, and abdomen, or a cephalothorax and abdomen, or a head and trunk. A central cavity, called the hemocoel (or blood cavity), is present; the open circulatory system is regulated by a tubular, or single-chambered, heart. Respiratory systems vary depending on the group of arthropod. Insects and myriapods use a series of tubes (tracheae) that branch through the body, open to the outside through openings called spiracles, and perform gas exchange directly between the cells and air in the tracheae. Other organisms use variants of gills and lungs. Aquatic crustaceans utilize gills, terrestrial chelicerates employ book lungs, and aquatic chelicerates use book gills . The book lungs of arachnids (scorpions, spiders, ticks, and mites) contain a vertical stack of hemocoel wall tissue that somewhat resembles the pages of a book. Between each of the "pages" of tissue is an air space. This allows both sides of the tissue to be in contact with the air at all times, greatly increasing the efficiency of gas exchange. The gills of crustaceans are filamentous structures that exchange gases with the surrounding water.

Book gills

The ventral side of a horseshoe crab showing the book gills located near the telson (tail). These gills flap back and forth bringing oxygen to the blood.

Groups of arthropods also differ in the organs used for excretion. Crustaceans possess green glands while insects use Malpighian tubules, which work in conjunction with the hindgut to reabsorb water while ridding the body of nitrogenous waste. The cuticle is the covering of an arthropod. It is made up of two layers: the epicuticle, which is a thin, waxy, water-resistant outer layer containing no chitin; and the chitinous procuticle, which is beneath the epicuticle. Chitin is a tough, flexible polysaccharide. In order to grow, the arthropod must shed the exoskeleton during a process called ecdysis ("to strip off"); this is a cumbersome method of growth. During this time, the animal is vulnerable to predation.

28.4.4: Subphyla of Arthropoda

Learning Objective

Differentiate among the subphylums hexapoda, myriapoda, crustacea, and chelicerata

Key Points

• The Hexapoda include insects; the Crustacea include lobster, crabs, and shrimp; the Myriapoda include centipedes and millipedes; and the Chelicerata include spiders, scorpions.
• The Hexapoda are the largest grouping of Arthropods, containing the more than one million species of insects, having representatives with six legs and one pair of antennae.
• The Myriapoda are terrestrial, prefering humid environments; they have between 10 and 750 legs.
• The Crustacea are primarily aquatic arthropods, but also include terrestrial forms, which have a cephalothorax covered by a carapace.
• The Chelicerata, which includes the spiders, horseshoe crabs, and scorpions, have mouth parts that are fang-like and used for capturing prey.

Key Terms

forcipule

a modified pincer-like foreleg in centipedes, capable of injecting venom

cephalothorax

the fused head and thorax of spiders and crustaceans

Representatives of Phylum Arthropoda

Subphylum Hexapoda

The name Hexapoda denotes the presence of six legs (three pairs) in these animals, which differentiates them from the number of pairs present in other arthropods. Hexapods are characterized by the presence of a head, thorax, and abdomen, constituting three tagma. The thorax bears the wings as well as six legs in three pairs. Many of the common insects we encounter on a daily basis, including ants, cockroaches, butterflies, and flies, are examples of Hexapoda.

Among the hexapods, the insects are the largest class in terms of species diversity as well as biomass in terrestrial habitats ). Typically, the head bears one pair of sensory antennae, mandibles as mouthparts, a pair of compound eyes, and some ocelli (simple eyes), along with numerous sensory hairs. The thorax bears three pairs of legs (one pair per segment) and two pairs of wings, with one pair each on the second and third thoracic segments. The abdomen usually has eleven segments and bears reproductive apertures. Hexapoda includes insects that are winged (like fruit flies) and wingless (like fleas).

Insect showing wings and body segments

Protaetia fieberi in flight posture. Hexapods are characterized by having three distinct tagma, or body segments. This beetle is just one of over one million different species of insects that inhabit the Earth.

Subphylum Myriapoda

Subphylum Myriapoda includes arthropods with numerous legs. Although the name is hyperbolic in suggesting that myriad legs are present in these invertebrates, the number of legs may vary from 10 to 750. This subphylum includes 13,000 species; the most commonly-found examples are millipedes and centipedes. All myriapods are terrestrial animals, prefering a humid environment.

Myriapods are typically found in moist soils, decaying biological material, and leaf litter. Centipedes, such as Scutigera coleoptrata, are classified as chilopods . These animals bear one pair of legs per segment, mandibles as mouthparts, and are somewhat dorsoventrally flattened. The legs in the first segment are modified to form forcipules (poison claws) that deliver venom to prey such as spiders and cockroaches, as centipedes are predatory. Millipedes bear two pairs of legs per diplosegment, a feature that results from embryonic fusion of adjacent pairs of body segments, are usually rounder in cross-section, and are herbivores or detritivores. Millipedes have visibly more numbers of legs as compared to centipedes, although they do not bear a thousand legs.

House centipede

The house centipede (Scutigera coleoptrata) is one of the 13,000 species of Myriapoda. They bear one pair of legs per segment and can inject venom. The Myriapods contain the millipedes and centipedes.

Subphylum Crustacea

Crustaceans are the most dominant aquatic arthropods since the total number of marine crustacean species stands at 67,000. However, there are also freshwater and terrestrial crustacean species. Krill, shrimp, lobsters, crabs, and crayfish are all examples of crustaceans . Terrestrial species like the wood lice (Armadillidium spp.) (also called pill bugs, rolly pollies, potato bugs, or isopods) are also crustaceans, although the number of non-aquatic species in this subphylum is relatively low.

Mediterranean green Crab

This crab (Carcinus aestuarii) is one of the 67,000 species of crustaceans inhabiting the world's oceans. Most crustaceans are decapods, having ten legs.

Crustaceans possess two pairs of antennae, mandibles as mouthparts, and biramous ("two branched") appendages: their legs are formed in two parts, as distinct from the uniramous ("one branched") myriapods and hexapods.

Unlike that of the Hexapoda, the head and thorax of most crustaceans is fused to form a cephalothorax, which is covered by a plate called the carapace, thus producing a body structure of two tagma. Crustaceans have a chitinous exoskeleton that is shed by molting whenever the animal increases in size. The exoskeletons of many species are also infused with calcium carbonate, which makes them even stronger than in other arthropods. Crustaceans have an open circulatory system where blood is pumped into the hemocoel by the dorsally-located heart. Hemocyanin and hemoglobin are the respiratory pigments present in these animals.

Subphylum Chelicerata

This subphylum includes animals such as spiders, scorpions, horseshoe crabs, and sea spiders and is predominantly terrestrial, although some marine species also exist. An estimated 77,000 species, found in almost all habitats, are included in subphylum Chelicerata.

The body of chelicerates may be divided into two parts: prosoma and opisthosoma, which are basically the equivalents of cephalothorax (usually smaller) and abdomen (usually larger). A "head" tagmum is not usually discernible. The phylum derives its name from the first pair of appendages, the chelicerae, which are specialized claw-like or fang-like mouthparts. These animals do not possess antennae. The second pair of appendages is known as pedipalps. In some species, such as sea spiders, an additional pair of appendages, called ovigers, is present between the chelicerae and pedipalps.

Chelicerae are used primarily for feeding, but in spiders, these are often modified into fangs that inject venom into their prey before feeding . Members of this subphylum have an open circulatory system with a heart that pumps blood into the hemocoel. Aquatic species have gills, whereas terrestrial species have either trachea or book lungs for gaseous exchange.

Chelicera of spiders

This photo shows the chelicera of a spider being held open with a stick. Some chelicerae, such as those found in spiders, are hollow and contain (or are connected to) venom glands which are used to inject venom into prey or a (perceived) threat.

The nervous system in chelicerates consists of a brain and two ventral nerve cords. These animals use external as well as internal fertilization strategies for reproduction, depending upon the species and its habitat. Parental care for the young ranges from absolutely none to relatively-prolonged care.

Attributions

• Superphylum Ecdysozoa
  • "Boundless." http://www.boundless.com/. Boundless Learning CC BY-SA 3.0.
  • "Ecdysozoa." http://en.wikipedia.org/wiki/Ecdysozoa. Wikipedia CC BY-SA 3.0.
  • "coelomate." http://en.wiktionary.org/wiki/coelomate. Wiktionary CC BY-SA 3.0.
  • "cuticle." http://en.wiktionary.org/wiki/cuticle. Wiktionary CC BY-SA 3.0.
  • "OpenStax College, Biology. October 17, 2013." http://cnx.org/content/m44667/latest/?collection=col11448/latest. OpenStax CNX CC BY 3.0.
  • "ecdysis." http://en.wiktionary.org/wiki/ecdysis. Wiktionary CC BY-SA 3.0.
  • "Cicada Molting." http://commons.wikimedia.org/wiki/File:Cicada_Molting.jpg. Wikimedia CC BY-SA.
• Phylum Nematoda
  • "Boundless." http://www.boundless.com/. Boundless Learning CC BY-SA 3.0.
  • "OpenStax College, Biology. October 17, 2013." http://cnx.org/content/m44667/latest/?collection=col11448/latest. OpenStax CNX CC BY 3.0.
  • "exoskeleton." http://en.wiktionary.org/wiki/exoskeleton. Wiktionary CC BY-SA 3.0.
  • "Soybean cyst nematode and egg SEM." http://commons.wikimedia.org/wiki/File:Soybean_cyst_nematode_and_egg_SEM.jpg. Wikimedia CC BY.
• Phylum Arthropoda
  • "Boundless." http://www.boundless.com/. Boundless Learning CC BY-SA 3.0.
  • "malpighian tubule." http://en.wiktionary.org/wiki/malpighian+tubule. Wiktionary CC BY-SA 3.0.
  • "tagma." http://en.wiktionary.org/wiki/tagma. Wiktionary CC BY-SA 3.0.
  • "spiracle." http://en.wiktionary.org/wiki/spiracle. Wiktionary CC BY-SA 3.0.
  • "OpenStax College, Biology. October 17, 2013." http://cnx.org/content/m44667/latest/?collection=col11448/latest. OpenStax CNX CC BY 3.0.
  • "BLW Trilobite (Paradoxides sp.)." http://commons.wikimedia.org/wiki/File:BLW_Trilobite_(Paradoxides_sp.).jpg. Wikimedia CC BY-SA.
  • "Horseshoecrab2." http://commons.wikimedia.org/wiki/File:Horseshoecrab2.jpg. Wikimedia CC BY.
• Subphyla of Arthropoda
  • "Boundless." http://www.boundless.com/. Boundless Learning CC BY-SA 3.0.
  • "forcipule." http://en.wiktionary.org/wiki/forcipule. Wiktionary CC BY-SA 3.0.
  • "cephalothorax." http://en.wiktionary.org/wiki/cephalothorax. Wiktionary CC BY-SA 3.0.
  • "OpenStax College, Biology. October 17, 2013." http://cnx.org/content/m44667/latest/?collection=col11448/latest. OpenStax CNX CC BY 3.0.
  • "Solifugae Chelicera lateral aspect 2012 01 24 0999s." http://commons.wikimedia.org/wiki/File:Solifugae_Chelicera_lateral_aspect_2012_01_24_0999s.JPG. Wikimedia CC BY.
  • "Carcinus aestuarii 2009 G1." http://commons.wikimedia.org/wiki/File:Carcinus_aestuarii_2009_G1.jpg. Wikimedia CC BY-SA.
  • "Proteatia vol." http://commons.wikimedia.org/wiki/File:Proteatia_vol.jpg. Wikimedia CC BY-SA.
  • "House centipede." http://commons.wikimedia.org/wiki/File:House_centipede.jpg. Wikimedia CC BY.