Subphylum Myriapoda Myriapoda (Greek “many feet”) includes two major groups the Chilopoda (centipedes) and Diplopoda (millipedes) as well as two smaller groups the Symphyla and the Pauropoda, both of which are small soil-dwelling organisms. Myriapods have a head with simple eyes and a segmented trunk which carries paired appendages, one per segment in centipedes and in most segments two in millipedes. Subphylum Myriapoda On the head myriapods have one pair of antennae, mandibles and one or two pairs of maxillae or feeding appendages. Walking legs are found on the abdominal segments. Class Chilopoda There are about 3,000 described species and an estimated 8,000 worldwide. Centipedes (from the Latin for hundred feet) differ from millipedes in that each segment bears only one pair of appendages. They always have an odd number of pairs of legs. Distribution is worldwide ranging from Tropics to the Arctic Circle. Require moist microhabitats because the cuticle is not waxy and so is not watertight. Respiration is primarily via a tracheal system, but a few species exchange gases across the body surface. Class Chilopoda Centipedes range in size from a few millimeters long to 30cm (the Amazonian Giant Centipede), which makes them some of the largest terrestrial invertebrate predators and they are a significant element of the predatory biomass in many invertebrate communities. http://www.google.com/imgres?imgurl=http://www.cutehomepets.com/wpcontent/uploads/2011/01/giant-centipede.jpg Class Chilopoda The first pair of appendages have been modified into poison fangs (forcipules), which they use to kill prey. Centipedes are active predators that hunt in leaf litter, under logs and in similar damp, dark places for prey. The body is dorsoventrally flattened, which probably allows the centipede to squeeze into small spaces in pursuit of prey. Figure 20.01 12.32B Class Diplopoda Diplopods are commonly called millipedes (Latin “thousand feet”). Most have 36-400 legs with 750 being the maximum. They have a distinctively cylindrical body and this coupled with the fact that most segments have two pairs of legs (all but the first four thoracic segments, which have one pair each) makes millipedes hard to misidentify. Class Diplopoda There are about 10,000 described species and they occur worldwide. Millipedes range from a few millimeters in length up to 38 cm (the African giant Millipede). The largest terrestrial invertebrate ever was a 2.6 meter long millipede Anopleura from the Carboniferous (340-280 mya). Figure 20.02 12.33 Class Diplopoda Millipedes because of their large number of legs move in a distinctive flowing manner, but are generally less active than centipedes. Slow moving. Most millipedes are scavengers that feed on decaying detritus and plant material which they consume using large chewing mandibles. They prefer moist, dark places. Class Diplopoda The primary means of defense is to curl up into a tight ball protecting their vulnerable underside. However, by toxins many species are also protected Millipede mating clip http://www.youtube.com/watch?v=emvTpB BfUwk Hunting centipede clip http://www.youtube.com/watch?v=UROVf mY3NTA Subphylum Crustacea The crustaceans are another very large group of arthropods with more then 67,000 described species. The group includes crabs, lobsters, shrimp, copepods, pill bugs, barnacles, krill, and crayfish. The crustaceans are the only arthropod group whose members are almost all aquatic, and mostly marine. Crustaceans The crustaceans are the only arthropods with two pairs of antennae. They also have a diverse array or other appendages specialized for different tasks. Lobster appendages For example, in lobsters on the head there are two pairs of antennae, a pair of mandibles for eating and two pairs of other feeding appendages called maxillae. These are followed by three pairs of maxillipeds, which are food handling appendages, mounted on the thorax. Figure 19.02b 12.16B Lobster appendages After the maxillipeds, come the five pairs of walking legs, the first of which are modified in many cases into large clawed chelipeds. Abdominal appendages are modified for swimming. These swimmerets are used for locomotion, but in males the first pair are modified for copulation. The last pair of appendages (uropods) are wide and assist the animal in moving backwards quickly. Figure 19.02a 12.16A Lobster clip http://www.youtube.com/watch?v=h6nhOC hpMck Crustaceans Crustaceans, being aquatic, breathe using gills. In most cases the gills are attached to appendages and movement of the appendages aerates them. Crustaceans Most crustaceans are predatory, although there are also many suspension feeders such as barnacles. The suspension feeders use their legs which have a thick coating of bristles to sweep food particles (such as plankton and bacteria) through the setae. Feeding barnacle Crustacean reproduction Most crustaceans have separate sexes, although in some groups males are scarce and parthenogenesis occurs. Most crustaceans brood their eggs and the offspring may be miniature versions of the adults or may pass through several larval stages. Subphylum Crustacea Subphylum Crustacea Class Branchiopoda: fairy shrimp, brine shrimp, water fleas, (e.g. Daphnia), Class Maxillopoda: copepods, barnacles. Class Malacostraca: Isopoda (pill bugs), Amphipoda (beach fleas), Euphausiacea (krill), Decapoda (lobsters, crayfish, crabs, shrimps) Subphylum Crustacea: Class Branchiopoda The Branchiopoda (Gk: “gill foot”) include a number of fairly primitive organisms including fairy shrimp and water fleas (such as Daphnia). The legs are the principal respiratory organ and as a result are flattened and leaflike. Figure 19.15 12.22 Subphylum Crustacea: Class Branchiopoda Most branchipods also use their legs for suspension feeding. The most diverse group of the branchiopods is the Cladocera, which is a major component of the zooplankton fauna in freshwater. Cladocera Close up of a cladoceran feeding http://www.youtube.com/watch?v=1KIVW8 cZQHU Subphylum Crustacea: Class Maxillopoda The maxillopoda includes several groups of crustaceans that were once considered classes themselves, but recently been grouped together based on their numbers of segments and a uniquely structured eye. Three important subclasses are the Copepoda (small, diverse planktonic), Branchiura (fish parasites) and the Cirripedia (barnacles). Copepoda Subclass Copepoda (Gk “oar foot”) are a very diverse group of small (usually only a few mm long), elongate and tapered crustaceans. They are a very important component of aquatic food webs and often are the commonest primary herbivore in aquatic communities. Figure 19.16c 12.24 Branchiura The Branchiura (Gk: “gill tail”) are a group of approximately 130 species of dorsoventrally flattened ectoparasites (usually of fish) often referred to as fish lice. The body is very flat and oval and almost entirely covered by the carapace. Argulus: a fish louse Branchiura Mouthparts and antennae have been modified into a proboscis with hooks and spines with which the parasite grips its host. Branchiurans are fast moving and often detach and leave a host before reattaching to another later. Once they grab a host they quickly move to a position behind the gill cover where they are less likely to be knocked or swept off as the fish moves. Branchiura While attached branchiurans feed on mucus and skin cells or suck blood. They can cause devastating losses to commercial aquaculture facilities where fish are confined at high densities Cirripedia The Cirripedia (L: “curl of hair” and “feet”) are the barnacles. Sessile and filter feeding as adults they are covered with a shell of calacreous plates secreted by the carapace. Cirri are extended between plates to filter feed. Figure 19.19a 12.26A Barnacle reproduction Barnacles are monoecious, but usually cross fertilize. Barnacles are cemented in place so mating is challenging. As a result, barnacles have evolved what is proportionally the longest penis in the animal kingdom. Dissected out barnacle penis Barnacles feeding and mating http://www.youtube.com/watch?v=v1SW- pl2gYs Barnacle reproduction Banacles brood their eggs until they develop into larvae, which are then released into the water to find somewhere to settle. The initial barnacle larva is a nauplius, which transforms into a non-feeding form called a cyprius. The cyprius has food reserves for about two weeks. It must find a place to attach permanently before it runs out of food. If it locates a suitable spot, it glues itself to it and transforms within 12 hours into an adult. Nauplius larva of barnacle Cyprius larva of barnacle Female barnacle brooding eggs Subphylum Crustacea: Class Malacostraca The Malacostraca (Gk: “soft” and “shell”) is the largest and most diverse group of crustaceans. Usually they have 8 thoracic and 6 abdominal segments, each of which has a pair of appendages. They include the isopods (sow bugs and pill bugs), Euphausiacea (krill) and decapods (crabs, crayfish and lobsters). Isopods Isopods include the familiar pill bugs, which live in damp places, but there are many aquatic forms and some are parasites of crustaceans and of fish. Figure 19.21a Pill bugs (Armadillidium vulgare) Figure 19.22 Isopod fish parasite (Anilocra) Tongue-eating louse One specialized isopod fish parasite isCymothoa exigua, or the tongue-eating louse. This parasite enters through the gills, destroys the tongue by feeding on its blood supply so it atrophies and replaces it becoming the fish’s new tongue. http://dailyorganism.blogspot.com/2011/06/ tongue-eating-louse.html Tongue eating louse http://www.youtube.com/watch?NR=1&fea ture=fvwp&v=XtwaNBsSbHQ Euphausiacea The euaphausids include only about 90 species, are only about 3-6cm long, but are very important components of the oceanic plankton. Vast swarms of them are fed on by fish and whales. Euphausiacea The total biomass of the Antarctic krill is estimated at over 500,000,000 tonnes and of that more than half is eaten annually by whales, penguins seals, squid, fish and other predators. Krill thus plays a central role in ocean food webs feeding largely on phytoplankton and converting their energy into biomass. Northern krill http://en.wikipedia.org/wiki/File:Meganyctip hanes_norvegica2.jpg Euphausids Euphausids are found world wide in the world’s oceans and live between 2 and 6 years depending on the species. The biomass losses through predation are made up through growth and reproduction. Krill undergo extensive vertical migrations in the water column descending during the day to try to avoid predators and ascending at night to feed. Euphausids Because Antarctic waters are areas where great upwellings of nutrients occur these oceans produce enormous quantities of phytoplankton in the summer and this phytoplankton is fed on by krill. The krill thus convert the phytoplankton into biomass larger animals can eat. Whale feeding Krill occur in vast, dense swarms that may include 10,000-60,000 individuals per square meter. These massive swarms are what make krill a profitable food source for whales to feed on and explains why whales migrate thousands of miles to feed in these waters. Krill and whales http://www.youtube.com/watch?v=1_BqC9 IIuKU Whale bubble netting clip http://www.youtube.com/watch?v=iREMH Ez5PdU Decapoda The Decapods (Gk: “10 legs”) have 5 pairs of walking legs the first of which usually are modified into pincers (also called chelae). The group includes the lobsters, shrimp, crabs and crayfish. Crabs differ from the others in being much broader across the carapace and having a shorter abdomen. Decapoda There are about 15,000 species of which about half are crabs and about 3,000 species of shrimp. Commercially a very important group and extensively fished for and farmed. Most decapods are scavengers and often predatory. Decapoda Decapods primarily depend on their thick exoskeleton and pincers for protection, but hermit crabs occupy abandoned mollusc shells and at least two species of crabs use sea anemones as weapons which they carry with them and thrust at anything that threatens them. Figure 19.26 12.31 Subphylum Hexapoda Subphylum Hexapoda (six legs). Members of the Hexapoda have three tagmata (head, thorax and abdomen) and three pairs of walking legs. There are two classes. Class Insecta: beetles, flies, wasps, ants, grasshoppers, bugs, caddisflies, fleas, butterflies and moths, lice, cockroaches, dragonflies, etc. Class Entognatha: collembolans, springtails and snowfleas. Subphylum Hexapoda Insecta can be distinguished from Entognatha by the fact that insects have the bases of their mouthparts visible outside the head capsule, whereas the Entognatha do not. Class Entognatha The Entognatha includes three classes the members of all of which are small (<10mm), inconspicuous organisms. The most obvious are the members of the order Collembola the springtails and snowfleas. They are very common in soil and sometimes swarm on ponds or snowbanks. They are called springtails for the springing organ they possess, which folds underneath the abdomen and can launch them high into the air if threatened. An aggregation of springtails Springtail Subphylum Hexapoda: Class Insecta The insects are spectacularly successful, there being more species of insects than the total number of species in all the other classes of animals combined. There are about one million named species of insect with probably millions more as yet undescribed. Class Insecta Insects occupy virtually all terrestrial and freshwater habitats, marine environments being the only one where they are scarce. They are a highly adaptable group that can withstand severe conditions and have evolved a suite of adaptations to minimize water loss including a waxy cuticle, the ability to close their spiracles, and the ability to extract almost all water from food and fecal material. Class Insecta Insects have three pairs of legs and usually two pairs of wings on the thorax. Head has a pair of large compound eyes, each eye being made up of up to 30,000 ommatidia, which are individual tube-like structures. Insect eyes are especially good at detecting motion and have a very wide field of view. Figure 20.22 12.20 Class Insecta The head also has a pair of antennae and complex, multipart mouthparts. The mouthparts equip different species of insect to feed on a wide variety of foods from nectar, to blood, to other insects, to plants. Figure 20.19 12.44 Class Insecta: flight Insects possess wings that were evolved independently (not surprisingly) of the birds and bats. The wings are outgrowths of the cuticle of the thorax. In most insects there are two pairs, but in flies only one. The second pair in flies has been modified into halteres (balancing organs) Figure 20.04a Class Insecta: flight Flight offers numerous advantages to insects enhancing the ability to search for food, mates and habitat, escape from predators, and migrate. It also is expensive in terms of the musculature required to fly and in some species (e.g. crickets) different flying and non-flying morphs occur depending on whether flying is ecologically beneficial or not. Class Insecta: flight Wings differ in size and shape from the long narrow wings of dragonflies to the broader shorter wings of many butterflies and moths. Wing beat rate varies greatly from a low of about 4 per second for butterflies to 100/s for bees to 1,000/s for some midges. Class Insecta: flight The fastest flying insects can reach about 30 miles per hour (horseflies and sphinx moths) with dragonflies reaching about 25mph. The kings of long distance migration are the monarch butterflies which travel hundreds of miles on migration at a speed of about 6mph. Class Insecta: Gas Exchange Gas exchange in insects is achieved by a series of tubes called tracheae. These tubes open to the outside via spiracles that can be opened to allow air in and closed to reduce water loss. The tracheae branch into a very fine network of fluidfilled tracheoles so that no cell is located far from a tracheole. In aquatic insects tracheal gills have evolved which are extensions of the body wall penetrated by many tracheoles. Figure 20.20a Figure 20.20b Class Insecta: Metamorphosis Insects go through a series of developmental stages before adulthood. Most insects (88%) go through complete metamorphosis: egg, larva, pupa, adult. In complete metamorphosis individuals of the same species at different stages of the life cycle do not compete with each other. Class Insecta The larval stage is the primary feeding stage where growth occurs. Indeed in some insects the adults are non-feeding and often short-lived e.g. mayflies, some butterflies. The pupal stage is a non-feeding stage in which the larva transforms into an adult. Often this stage is an overwintering stage. The adult stage is the reproductive stage in which eggs are laid and the cycle repeats. Figure 20.24 12.46 Class Insecta Insects such as grasshoppers, mayflies, lice, and bugs do not undergo complete metamorphosis but instead undergo incomplete metamorphosis. In incomplete metamorphosis the young are called juveniles (or nymphs) and look like the adult but smaller, gradually increasing in size via a series of molts. Class Insecta: important orders Coleoptera: beetles; Forewings modified into hardened covers or elytra. Biting, chewing mouthparts. waynesword.palomar.edu Hymenoptera: ants, bees, wasps; narrow-waisted, includes a wide variety of social species. Lepidoptera: butterflies and moths; membranous wings covered in fine scales. Mouthparts are a sucking tubes that curls up when not in use. en.wikipedia.org Diptera: true flies; single pair of wings and a pair of modified wings (halteres) which act as gyroscopes. en.wikipedia.org Hemiptera: true bugs; Piercing sucking mouthparts. www.oocities.org www.earthlife.net Orthoptera: grasshoppers, locusts, mantids; chewing mouthparts. en.wikipedia.org Siphonaptera: fleas; Wingless with laterally compressed bodies. Specialized for blood sucking. hardinmd.lib.uiowa.edu Trichoptera: caddisflies; small softbodied. As larvae inhabit cases they construct of sand, gravel and plants material in streams. Isoptera: termites; highly social wood eating insects. www.evrimteorisi.info Social behavior In many groups of insects there is a considerable degree of social behavior. This ranges from temporary associations that are quite uncoordinated such as groups of hibernating or roosting carpenter bees to true societies of the Hymenoptera (bees and ants) and termites (Isoptera) Evolution of Eusociality In the complex eusocial (truly social) societies of Hymenoptera and Isoptera communities are permanent and all stages of the life cycle occur within the nest. There is division of labor and highly coordinated behavior. Many individuals do not reproduce. Instead they act as helpers at their parents’ nests for their entire life. This is an extreme type of altruism. Evolution of Eusociality in insects Eusociality describes social systems with three characteristics: Overlap in generations between parents and offspring. Cooperative brood care. Specialist castes of non-reproductive individuals. Honey Bee nests Honey bees have a very complex social system that lasts potentially indefinitely as the group survives from season to season. Up to 70,000 bees may occupy a hive and there are 3 castes: a single reproductive queen, a few hundred males (drones), and many thousands of female workers who are non breeders. Honey Bee nests The workers tend to the hive’s young, gather nectar and pollen and make honey, and guard the hive. What caste individuals belong to is determined by fertilization (males develop from unfertilized eggs) and what the developing larva is fed. Honey Bee nests Larvae that will become queens are fed royal jelly, which is secreted by the salivary glands of nurse workers. Workers only produce royal jelly when the level of “queen substance” in the hive declines (because the queen has died or is too old). Queen substance is a pheromone produced by the queen that suppresses sexual maturation of workers. Honey bee foraging Honeybees are remarkable for their ability to collaborate in their search for food. An individual who has found food dances to convey information about the food source’s location to other foragers. Karl von Frisch pioneered the work on dancing bees. Honey bee foraging A honey bee that has found food dances to indicate the location of the food to other foragers. If the food is close (<50m) the bee performs a round dance. This tells the other bees that food is near the hive. Round dance Round dance Honey bee foraging If food is further away bee performs a “waggle” dance. Bee performs dance on a path that is roughly figure 8 shaped. Bee travels in straight line while waggling her body. Then turns left or right to circle back to the beginning. If bee outside hive, direction of waggle dance points directly at source of food. Honey bee foraging Inside in the hive it is dark and the bee performs the dance on a vertical surface. Vertical indicates the direction of the sun and the direction of the dance relative to the vertical indicates the angle of the food relative to the sun. The length of the waggle portion indicates the approximate distance to the food. The fewer circuits the bee performs in a period of time the further away the food is located. Vertical orientation in hive Waggle dance. Waggle dance http://www.youtube.com/watch?v=aUCoLe I5Qxg Ant sociality The ants who are close relatives of the bees (both are members of the Hymenoptera) also have highly organized colonies. As in bees a queen is responsible for egg laying and the workers are females. There may be several different castes of worker females: soldiers specialized for defense; and different sized workers specialized for food gathering and tending the young. Males are produced only for reproduction and die after mating. Ant sociality Ants have evolved many remarkable behaviors including: Slave making: some species raid the nests of other species of ants and steal their larvae, which grow up to become the workers in their new nest. Specialist slave makers produce no workers of their own. Ant sociality As well as: Farming: Numerous species of ants farm various different species of fungus gathering leaves to turn into compost and destroying pests that attack the fungus. Herding: ants protect “herds” of aphids from parasitoids and predators and eat the honeydew (partially digested plant juices) that the aphids secrete. Termite colonies Termites belong to a different order of insects than bees and ants (the Isoptera) and their colonies have a similar but not identical organization. As in the Hymenoptera there are both sterile and fertile individuals in a termite colony. Termite colonies Nonreproductive females become either workers or soldiers (which have large heads and mandibles with which they protect the colony). Termites specialize in eating wood and depend on bacteria to digest it as they do not produce enzymes capable of digesting cellulose. Haplodiploidy and eusocial Hymenoptera One idea advanced to explain eusociality is the unusual genetic system (haplodiploidy) of the Hymenoptera (ants, wasps, bees, etc.). Males are haploid and females diploid because males develop from unfertilized eggs and females from fertilized eggs. Haplodiploidy and eusocial Hymenoptera Daughters receive all of their fathers genes and half of their mothers genes. Thus, daughters share ¾ of their genes. This suggests females would be better off if they favored the production of reproductive sisters rather than their own offspring. Haplodiploidy and eusocial Hymenoptera Queens are equally related to all offspring and so should prefer a 1:1 ratio of sons to daughters among reproductives. Females workers however should prefer a 1:3 ratio of brothers to sisters among reproductives. Haplodiploidy and eusocial Hymenoptera It has been shown in wood ants that queens produce equal numbers of male and female eggs, but the hatching ratio is heavily female biased. Workers apparently selectively destroy male eggs. Haplodiploidy and eusocial Hymenoptera Haplodiploidy appears to influence worker behavior, but consensus among scientists today is that it does not explain the evolution of eusocial behavior in Hymenoptera. There are several reasons why. Haplodiploidy and eusociality First, haplodiploid explanation assumes all workers have the same father. However, honeybee queens mate with more than 17 males on average. As a result relatedness between worker honeybees often below 1/3. Haplodiploidy and eusociality Second, in many species, more than one female founds a nest. In this case workers may be completely unrelated. Haplodiploidy and eusociality Third, many eusocial species are not haploid (e.g. termites) and many haplodiploid species are not eusocial. Haplodiploidy and eusociality Phylogenetic analysis of Hymenoptera by Hunt (1999) emphasizes that eusociality is relatively rare even though haplodiploidy occurs in all groups. Eusociality occurs in only a few families which are scattered around the tree, which suggests eusociality has evolved independently multiple times. Haplodiploidy and eusociality Hunt also points out that eusociality has only evolved in groups that build complex nests, and care for young for a long time. Association between nest building, long term care and eusociality suggests main driving force for eusociality is ecological not genetic. Haplodiploidy and eusociality Nest building and need to supply offspring with a steady stream of food make it impossible or very difficult for a female to breed alone. Also, if predation rates are high, solitary breeding individuals may not live long enough to raise their young. Insects and humans: benefits Insects and humans obviously interact in many ways and insects are both harmful and beneficial. Benefits include the fact that insects produce a number of commercially valuable products including honey and silk and historically were used to make certain dyes. Insects and humans:benefits Another very important role insects play is in the pollination of food crops. Today in the U.S., however, the spread of bee mites, which wipe out hives of honey bees, currently threatens this. Historically the introduction of European honey bees greatly reduced the numbers of native pollinating insects, such as bumblebees, and these species now are too scarce to make up the pollination deficit. Insects and humans: costs There are a huge number of insects that consume crops, trees and ornamental plants and we fight an ongoing battle with these pests. The success of some of these pests is because they were introduced here and escaped their native predators and parasites (e.g. Japanese beetles, gypsy moths, various bark beetles, scale insects) or simply because they thrive in monocultures of crops. Insects and humans: costs The costs both economic and ecological of pesticide based control of insect pests are enormous. Attempts to reduce dependence on pesticides have led to increased use of biological control agents and the use of integrated pest management approaches in which strategies such as rotating crops, planting multiple different crops, planting pest resistant varieties, and employing biological controls are used together to limit pesticide use. Insects and humans: costs Insects also are major vectors of disease and transmit many serious disease agents. Vectors include: Mosquitoes: malaria, yellow fever, filariasis, West Nile virus. Blackflies: river blindness Fleas: bubonic plague Tsetse flies: sleeping sickness Rhodnius bugs: Chaga’s disease Lice: typhus