Crustaceans

Zoology 250 Lecture 25 ARTHROPODA (IV): Crustacea

(Crustacea study images)

SubPh. CRUSTACEA (crabs, shrimps & relatives; >50,000 marine, freshwater & terrestrial spp.); share 3 traits: i) 2-pair of antennae, ii) terminal moveable spine(s) on telson, & iii) a nauplius larval stage 2) The body plan is surprisingly stereotyped for such a diverse group:

a) head has 5 segments with limbs: 2 antennae, 1 mandible, 2 maxillae b) head is typically covered by a carapace; the carapace may extend over a few or all thoracic segments (-> cephalothorax) c) some thoracic segments may fuse with the head and their limbs modified as maxillipeds for feeding; walking legs= periopods d) abdominal segments may or may not have limbs (pleopods)

3) Crustacean diversity

a) 5 classes: 3 major (Branchiopoda, Maxillopoda, Malacostraca) & 2 small, phylogenetically important (Remipedia, Cephalocarida) b) most are marine, but one class (Branchiopoda) is primarily freshwater and three others have freshwater members c) most are free living; 2 classes have a number of parasitic species, and one subclass (Cirripidea) is entirely sessile as adults d) body size is generally small (<20 mm) in most classes/subclasses, but large size (>100 mm) occurs in three subclasses e) classes are distinguished by: number of tagmata, number of segments per tagmatum, number of anterior segments covered by a carapace, number of segments fused with the carapace, and presence/absence of movable spines (rami) on the telson

f) most diverse class (Malacostraca) share 3 characters: i) all have 21 somites, ii) stalked eyes in adults, iii) limbs on abdominal segments

4) Primitive Crustacea had an elongate body with many similar biramous limbs; most have a carapace that covers the head & part/all of thorax 5) Particle feeding in Crustacea is accomplished in many ways

a) primitively, crustaceans were probably particle feeders b) filter feeding at small size is difficult because of high viscosity: water has no momentum, 'wall effects' are large, flow is reversible c) in fairy shrimp a filter box generates high pressure that forces water through a sieve of setae; other small species use similar methods

1) SubPh.CHELICERATA (sea spiders, horseshoe crabs, spiders, mites) a) body generally has 2 tagmata; lack antennae!; have uniramous legs b) Cl. PYCNOGONIDA (sea spiders) ~600 marine spp - typically small bodied (<5 mm) but deep sea & Antarctic species have leg spans > 25 cm!; predators mainly on hydroids - very unusual body form: legs make up 50 - 80% of body mass! - typically have 1 pair of chelicerae, 1 pair of ovigers, 4 pair of walking legs, a reduced abdomen, and simple eyes - have an unusual reproductive biology: males brood embryos c) Cl. MEROSTOMATA (horseshoe crabs) ~50 marine spp - considered 'living fossils' because 400 million-year old fossils resemble living forms so closely; only 4 living species - large-bodied predators (to 40 cm) on small, infaunal invertebrates (food shredded or crushed by gnathobases of walking legs) - have 1 pair of chelicerae, 5 pairs of walking legs (first 4 with claws), abdominal book gills, and a prominent tail spine

- the only chelicerate with compound eyes (independently evolved) d) Cl. ARACHNIDA (spiders, mites & ticks, scorpions, etc.) ~70,000 mostly terrestrial spp; a few marine & freshwater spp - generally inspire more fear in humans than any other animal group - have 1 pair of chelicerae; modified as small pincers (most groups) or hollow fangs for injecting poison (spiders) - have 1 pair of pedipalps; may be leg-like & sensory (spiders) or form large pincers (scorpions) - 4 pair walking legs (abdomen may have other limbs), simple eyes - Or Araneae (spiders); nearly half of all arachnid species - liquify prey with digestive enzymes & imbibe fluids with a sucking pharynx & stomach - have the impressive ability to spin silk from the abdomen - some have quite elaborate social & courtship behaviors - Or Acari (mites & ticks) >500,000 spp? are everywhere! (marine, terrestrial, aquatic,aerial, subterranean, deep sea, hot springs) - most (except ticks) are very small (< 1mm); lack tagamata - all ingest liquids (from host, or dissolved by saliva)

2) SubPh. TRILOBITOMORPHA (extinct trilobites) a) were once the most numerous arthropods on earth; some reached very large size (>40cm); all thought to be particle feeders b) 3 tagmata; head with 1 pair antennae & 4 pair of thoracic-like legs c) variable number of thoracic segments with similar biramous limbs d) variable number of abdominal segments fused as pygidium

3) SubPh. UNIRAMIA (myriapods & insects)

a) 2nd antennae lost, limbs exclusively uniramous, possess a spiracle/trachea system for respiration b) SuperCl. MYRIAPODA: - body has 2 tagmata (head, trunk); the trunk has many similar legs - compound eyes are lost - Cl. CHILOPODA (centipedes) 15 - 177 pairs of legs; 1st pair bears poison claws; mostly predatory - Cl. DIPLOPODA (millipedes): 100 - 700 pairs of legs; adjacent trunk segments fuse yielding 2 pairs of legs per body unit (diplosegment); mostly herbivores
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c) SuperCl. HEXAPODA (=Insecta): - body has 3 tagmata (head, thorax, abdomen); thorax bears 3 pairs of walking legs; abdomen lacks limbs - 2nd maxillae fuse as labium - typically have 1 pr compound eyes & 3 medial ocelli (simple eyes) - Cl. APTERYGOTA: insects lacking wings (ametabolous devel.) - Cl. PTERYGOTA: winged insects - SubCl. EXOPTERYGOTA: wing buds are external; mostly hemimetabolous devel. (nymphs resemble adults but lack wings) - SubCl. ENDOPTERYGOTA: wing buds arise from internal imaginal discs; holometabolous development (larvae may bear no resemblence to adult; pass through a pupal stage where a radical metamorphosis takes place; eclose as fully formed adult) - an extraordinarily successful arthropod group, but very curiously never successfully invaded the sea

Approximately 30,000 species make up this Subphylum. Most are aquatic; of these, the majority are marine but some are found in fresh water. Members of the Subphylum include lobsters, crabs, crayfish, shrimp, copepods, barnacles, and several other groups of organisms. All have two pairs of antennae, a pair of mandibles, a pair of compound eyes (usually on stalks), and two pair of maxillae on their heads, followed by a pair of appendages on each body segment (crustacean bodies usually are made up of head, thorax, and abdomen, although the segments composing these tagmata differ among different Classes). The appendages are primitively branched (biramous), and although this condition is modified in many species, adults always have at least some biramous appendages. Crustaceans respire via gills. Like other arthropods, all have a hard but flexible exoskeleton. Most crustaceans are free-living, but some are sessile and a few are even parasitic. Most use their maxillae and mandibles to take in food. The walking legs, including specialized chelipeds, may be used to help capture prey. Some crustaceans filter tiny plankton or even bacteria from the water; others are active predators; while still others scavenge nutrients from detritus. Most crustaceans are dioecious. The actual mechanisms by which fertilization is achieved vary greatly. Some crustaceans hatch young that are like miniature adults; others go through a larval stage called a nauplius. Many species, including lobsters, crayfish, barnacles, and crabs are important to human economies, some very much so. Others, such as krill, are at the base of extremely important marine food chains. Still others are crucial in recycling nutrients trapped in the bodies of dead organisms.

Brachiopoda Approximately 800 species of branchiopods are found worldwide in freshwater ponds, lakes, and inland saline waters such as the Great Salt Lake in Utah. Their fossil record includes the extinct order Lipostraca and dates back to the Devonian period (approximately 400 - 360 million years ago). Some references recognize four extant orders: Anostraca (fairy shrimps), Notostraca (tadpole shrimps), Cladocera (water fleas), and Conchostraca (clam shrimps). Members of these orders are commonly used for aquarium fish food, scientific research, and once were marketed as pets called sea monkeys. These small crustaceans are a very important source of food for fish and birds in nature. Distinguishing features of a branchiopod include a small body (0.25 mm - 10 cm long), paired compound eyes, single simple eye, simple mouth parts, leaflike or phyllopodous appendages, and minimal body tagmosis. The nervous system and sensory system are simple, although some species vibrate their compound eyes to gather more visual information. Branchiopods use their leaf-like appendages for feeding, locomotion, and respiration. Gathered food particles are pushed into a groove that leads to the mandibles , then the ventral food gut .

mouth and, in turn, a complete

Undigested particles exit through the

anus and nitrogenous wastes are eliminated through

maxillary glands. These glands, located near the maxillae , also function in osmoregulation. Locomotion is generally achieved by metachronal beating of the appendages. The leaflike portions of the appendages have a large surface area that functions in gas exchange. Since most branchiopods are small with a thin cuticle, gas exchange can occur across the body wall as well. The circulatory system includes a heart that pumps blood into an open body cavity or hemocoel. The pigment in the blood of some species is hemoglobin. Brine shrimp, of the order Anostraca, are most noted for their desiccation-resistant eggs that will hatch in salt water. Most species of branchiopods are gonochoric and some are parthenogenetic. Those species that have indirect development produce nauplius larvae. Maxillopoda (ostracods, copepods, barnacles) Maxillopods include barnacles, copepods, mystacocarids, tantulocarids, branchiurans, ostracods, and related groups. Most species are small. Most feed by means of their maxillae (rather than filter feeding using thoracic appendages to move water); barnacles, however, are an exception. Barnacles feed with thoracic appendages, but in a way that is unique among crustaceans. Other characteristics of maxillopods including a basic plan of 5 head and 10 trunk segments (6 thoracic and usually 4 abdominal), followed by a terminal telson. The abdominal segments typically lack appendages; appendages elsewhere on the body are usually biramous. Barnacles grow to encrust solid structures placed in marine environments, including docks and pilings and also the bottoms of ships. Malacostracans Malacostracans are distributed worldwide in marine, freshwater, and terrestrial environments. There are roughly 25,000 species in as many as fifteen orders. Decapoda (crabs, lobsters, and shrimps) is the most speciose group within the Malacostraca. Malacostracans exhibit the hard, calcified exoskeleton typical of crustaceans. The body is divided into three tagmata, cephalon, thorax, and abdomen. The head and thorax are fused into a cephalothorax and may be difficult to distinguish. All malacostracans possess five segments in the head, eight in the thorax, and six in the abdomen, excepting the 20-odd species in the Phyllocarida, which have seven abdominal segments. As a general rule, each segment bears a pair of appendages, but in some organisms appendages are lacking on several abdominal appendages. The anterior one to three thoracic appendages are modified into maxillipeds, which are used in feeding. The anteriormost five abdominal appendages are, almost without exception, biramous. In most malacostracans, the posteriormost abdominal appendages, if present, are flattened and form a tail fin with the telson. Orders are often categorized by the specialization of specific limbs and body segments. There is a great deal of morphological diversity within the class, which is the most familiar of all the crustacean taxa.

Malacostracans have the typical crustacean body plan. Internal gills are protected by the carapace. The circulatory system in large organisms may be highly developed and extensively venous, though it is still considered open (as opposed to closed). The nervous system is highly centralized. A large brain near the eye is connected to a number of ganglia via a paired ventral nerve cord, which runs the length of the body. Green glands in the second antennae serve an osmoregulatory and excretory function. Disposal of nitrogenous wastes probably also occurs across the gills or body wall itself. The mouth of malacostracans leads to a two-chambered stomach, which possesses a grinding structure called the gastric mill. Digestion occurs throughout the gut, and waste matter is expelled through a posterior anus on the telson. Malacostracans are dioecious, and sex is genetically determined. The gonads are located in the sixth thoracic segment in females, and the eighth in males. Copulation is the rule as the unflagellated sperm are nonmotile. The anterior one or two abdominal appendages in males are modified into reproductive structures designed to aid in sperm delivery. Development ranges from direct to metamorphic among members of class Malacostraca. In peracardians, eggs are brooded behind the thorax. In other malacostracans, eggs are laid. Most metamorphosing malacostracans have a nauplius larva, but in many species eggs hatch into zoea larvae. Virtually every imaginable feeding strategy is demonstrated by at least one member of the class. Many malacostracans are strictly carnivorous, and are active hunters. Organisms representing many orders possess thoracic appendages modified for spearing or catching and crushing prey. Several malacostracan taxa are parasitic. Still others are scavengers. Herbivorous malacostracans, as well as filter-feeders, also exist. Malacostracans are generally active. Among benthic taxa, however, some burrowing species remain fairly inactive. Many pelagic forms are active hunters. Decapods are known for elaborate courtship displays, such as those demonstrated by the fiddler crab. Malacostracans have a powerful role in the economy. Humans consume large amounts of decapods, and huge industries have developed around the capture or farming and sale of shrimp, lobster, and crabs. There is also a large aquarium trade, supplying animals both as pets and as food for fish and amphibians. Most malacostracan parasites invade fishes and crustaceans. For this reason, parasitic malacostracans have a negative impact on fish, shrimp, lobster, and crab industries. Malacostracans play such an important role in aquatic ecosystems that their conservation is an important issue. Commercial overfishing may eventually put populations in danger. Ironically, it is the important role that malacostracans play in the human economy that is endangering them. Both the fishing and farming of malacostracans can be environmentally damaging. The taxonomy of malacostracans centers around the specialization and arrangement of appendages and body segments. Unfortunately, many researchers suspect that a high degree of convergence is obscuring the phylogeny of malacostracan orders. For this reason, the taxonomic divisions between

many groups of malacostracans should be viewed as a good general guidelines rather than strict phylogenetic relationships.