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Entomology BIO 3323 EXTERNAL ANATOMY An outer exoskeleton of hard, articulating cuticular plates is a feature shared by all arthropods. It’s a complex non-living structure formed from strengthening alpha-chitin embedded in proteins, the procuticle, and topped with a waterproof epicuticle of crosslinked proteins and waxes. The cuticle is secreted by the underlying , living epidermal layer. This outer suit of armour creates a variety of different problems for insects not the least of which, is that in order to grow, the insect must escape the old cuticle and then lay down a new one, it moults. What’s interesting about the cuticle is the different shapes and forms it has. The delicate wings of a small fly, the massive jaws of some beetles, and the feathery antennae of moths all are made of cuticle. It is the bioplastic of the living world and the plasticity of this outer body covering helped to insure the success of insects. As you look at the different external features on the animals during today's lab always remember they are all made with the same cuticular components. INSECT MORPHOLOGY The best way to appreciate the field of Insect Morphology is by approaching it from a comparative standpoint. In the lectures we have discussed the tremendous diversity that occurs within the Class Insecta and how this is reflected in the diverse and multiple numbers of environmental niches that these animals are able to occupy. Underlying all of these adaptations are certain morphological, or structural characteristics, that all insects share. In different insects These have been modified into different structures designed for a variety of different functions. The terminology associated with the external anatomy is unique to insect morphology and these terms are important as reference points for subsequent discussions in both the laboratory and lectures. The stuructures are important in insect identification, so when the keys PAGE: 1 - © JON G. HOUSEMAN BIO 3323 Entomology ask about a modification or shape of the paranotal lob of the mesotharacic segment you’re going to need to know just what that means. External Anatomy - The lubber grasshopper The Lubber Grasshopper will be used to introduce the external characteristics of insects. Traditionally the grasshopper is used for this type of laboratory exercise because it has a number of primitive characters. You should be aware that although this animal represents some ancestral characteristics it is, in its own right, a very specialised animal, it’s jumping legs for example. We’ll look at modifications of the basic body plan in the last part of the lab and with demonstration materials as the term progreses. Figure 1 Major external features of the lubber grasshopper. © BIODIDAC Head Thorax Antenna Ocelli Compound eye MesoPro- Abdomen Meta- Femur Tibia Forewing Hindwing Cercus Spiracles Identify the three major tagma of an insect: The head, thorax and abdomen. The body is covered by the exoskeleton or cuticle that is laid down by the underlying epidermis. The exoskeleton is divided into a number of plates, or sclerites, that can be separated from each other by either sutures or membranes. The result is that different PAGE: 2 EXTERNAL ANATOMY © JON G. HOUSEMAN Entomology BIO 3323 regions of hardened cuticle can bend against each other. What may appear as lines between segments are often the inflection of the cuticle to form internal skeletal elements such as apodemes. Head Vertex Coronal suture Epistomal suture Anterior tentoral pit Clypeus Labrum Mandible Ocellus Figure 2 Major external features of the head of the lubber grasshopper. © BIODIDAC Gena Frontal suture Frons Subgenal suture The insect head is composed of six segments three of which are preoral (pregnathal) and three post oral (postgnathal). (Remember though that there is another school of thought that believes the insect head had 5 segments ancestrally.) The last three segments have appendages, the mandible, maxillae and fused labium; antennae and labrum are preoral appendages. With the exception of the most posterior lines, sutures, of the head there is no visible marker for the underlying segmentation of the tagma. Most of the visible suture lines result from inflections of the cuticle acting as strengthening ridges for the head capsule. These internal inflections are called apodemes. Identify the main features of the insect head. Compound eyes are located laterally and a close examination will reveal the complex pattern of ommatidia, the fundamental repeating optic units of the eye. In addition to the compound eye, ocelli are located on the facial region and serve a light receptive structures usually monitering light levels. Insects may have up to three ocelli and in your grasshopper they may be hard to see if the specimen is still wet with the preservative. The single pair of antennae are characteristic of the Tracheata, and are composed of three basic parts; the most proximal is the scape, the pedicel is next and the most distal part is the flagellum. The latter is composed of a number of annulations and is often elaborately modified in different insects. What is the difference between an annulation and a segment? Examine the antennae closely using the dissecting microscope and observe the sensory or setal hairs. These have a primarily chemosensory role and may not be fully apparent until the preservative has dried off the surface. The head is divided into a number of regions by grooves or sutures. The epistomal suture, for example, lies between the frons and the clypeus and reinforces the head against the forces generated by the mandibular musculature. While we refer to insects as having an PAGE: 3 - EXTERNAL ANATOMY © JON G. HOUSEMAN BIO 3323 Entomology external skeleton that are points were the cuticle expands internally, for example apodemes, to increase the surface area available for muscle attachment. The small pits in this suture are the inflections of cuticle that form the endoskelelatal tentorium of the head. The other apodemes of the tentorium is located at the back of the head. Here the occiput and postocciput are all the reamins of the heads original segmentation and near the base of the plates is the second apodeme of the tentorium. This very specialised piece of internal skeleton is required for the muscles that move the mouthparts. A grasshopper head that has bean cleared (the tissues are dissolved), by boiling it in alkali shows this internal element. The inverted Y located on the head is an ecdysial line, or suture, which splits open when the insect moults. Familiarise yourself with the main plates of the head which include the vertex, frons, gena, occiput, postocciput, and the various sutures.. Figure 3 Major external features of the head of the lubber grasshopper. © BIODIDAC Ocular suture Gena Ocellus Antenna Occipital suture Postoccipital suture Subocular suture Occiput Frons Postocciput Cervex Posterior tentorial pit Post gena Labium Clypeus Subgenal suture Labrum Mandible Maxilla The opening to the digestive system lies at the base of a buccal cavity consisting of the upper labrum; the labial bottom; and the mandibular and maxillary sides. The grasshopper mouthparts are similar to the generalised, or primitive form. Carefully remove the mouthparts by pulling the head forward from the thorax and cut thorough the membranous neck region. The labrum is suspended from the clypeus and forms the upper lip, or roof of the buccal cavity. The mandibles are highly sclerotized and hardened with both a cutting , incisor, and grinding , molar, region, locate both. The two points where the mandible articulates with the head should also be PAGE: 4 EXTERNAL ANATOMY © JON G. HOUSEMAN Entomology BIO 3323 obvious. The maxillae are also paired and composed of a tooth-like lacinea, a galea and a sensory leg-like maxillary palp which is used to taste the food. Figure 4 Mouthparts of the lubber grasshopper. © Clypeus BIODIDAC Labrum Grinding Mandible Cutting Mentum Hypopharynx Cardo Stipes Prementum Paraglossa Labial suture Glossa Palpifer Palpus Lacinia Ligula Labium Galea Palpus Maxilla This sensory structure is also covered with sensory hair similar to those on the antennae. The labium represents two fused mouthpart appendages and forms the bottom to floor to the buccal cavity. Again sensory labial palps are present. The hypopharynx is not an appendage but lies in the buccal cavity as a tongue like structure with oral opening above and salivary opening below. As we will see later in the lab this plan can be extensively modified and has allowed the insects to feed on a tremendous variety of different foods. Thorax The thorax is locomotory in function and is composed of three segments, the prothorax, mesothorax and metathorax. Each of these thoracic segments bears a pair of appendages and, if present, wings are on the mesothoracic and metathoracic segments. The thoracic segments can be divided into three major sclerites which include the dorsal notum (tergite), lateral pleura and the ventrally located sternum. The movements of these three plates relative to each other are important in understanding the flight mechanism. In most insects the lateral pleura have fused with the ventral sternites to form the bottom three sides to the thoracic box. The notum is connected by a membranous part of the pleura to the box, forming the top. As the notum rises the wings are lowered, as it falls, the wings are raised. Look closely at the pleural plates of those with wings. A central pleural suture divides the plural sclerite into an anterior episternum and posterior epimeron. The inflextion of the suture braces the thoracic box against the muscular forces of the flight muscles. Locate PAGE: 5 - EXTERNAL ANATOMY © JON G. HOUSEMAN BIO 3323 Entomology the basalare and subalare sclerites in the pleural membrane under the wing . These two scerites and the pleural wing process, the extension of the pleura sclerite between them, are important in wing movement. Two pairs of spiracular openings are located on the thorax and these regulate air intake into the internal tracheal system which is also an unusual feature of insects that oxygenate their tissues by the direct supply of air rather then by using a circulating respiratory pigment. Locate the spiracles and the auditory tympanum. The pleural sclerite also articulates with the leg which is attached in a membranous area below the pleural suture. The insect leg is composed of six segments and the tarsus is further divided into up to five segments, and pretarsus with its claws. The divisions of the tarsus gives the distal end of the appendage an even more multisegmented appearance. Don't confuse these with the true leg segments. Later in the course you’ll be counting tarsal and prestarsal segments and you’ll need to know the difference! The tubular exoskeleton means that only linear or planar articulations occur. Try to flex each of the segments. Do they all move in the same direction relative to each other? Identify the major leg segments include: coxa, trochanter, femur, tibia and tarsus and the last pretarsal segment which has a claw. A fleshy pad is often associated with the claw at the tip of the organism. This pad when located between the claws is referred to as an arolium while when located under and at the base of the claw it is called a pulvilli. Figure 5 Parts of the typical insect leg. © BIODI- DAC Tarsomere Claw Trochanter Coxa Tarsus Tibia Femur The wings of an insect are often membranous and strengthened by a series of ridges or veins. This arrangement of veins and cross veins is an important taxonomic tool for identification of insects. Identify the following wing veins costa, cubitus, subcosta, radial, medial, cubital and anal. Are all of these veins visible on the grasshopper wing? Abdomen The apparent or visible segmentation of the insect abdomen varies in different insect groups and but primitively this tagma was composed of 11 segments. A typical abdominal segment consists of a sclerotized tergite and sternite and membranous pleura. Spiracular openings are PAGE: 6 EXTERNAL ANATOMY © JON G. HOUSEMAN Entomology BIO 3323 located on each of the segments and the terminal segments bear appendages involved in copulation and egg laying . Identify the cerci and ovipositor. VARIATIONS ON THE BASIC PLAN The insect head Insect heads may have one of three different orientations based on the position of the mouthparts relative to the remainder of the head capsule. When the mouthparts are directed downward the head is considered hypognathous compared to prognathous, when the mouthparts are directed forward. In both these last two cases the oral appendages are located anteriorly but when positioned in the posterior region the head is opisthognathous. Figure 6 Three different types of insect heads From left to right, hypognathous, prognathous and opisthognathous © BIODIDAC There are some distinct advantages to these different orientations. Many larval insects have prognathous heads because they live in the same medium on which they feed. Forward directed mouthparts are an obvious advantage. Take a look at the mealworm larva as our example of a prognathous head. Others such as the cicada and a variety of plant sap feeding insects must have the mouthparts penetrate the vascular plant tissue. The achieve this requires considerable strength and the opisthognathous head allows the insect to apply the pressure required to penetrate the plant. Identify the different mouth part types on the specimens. Insect Mouthparts One reason for insect success is their ability to exploit a wide variety of food sources and their mouthparts have been extensively modified to accomplish this. These modifications included changing the shape of the of the various mouthparts and even the complete loss of some parts are whole mouthparts. Identify the components in the different PAGE: 7 - EXTERNAL ANATOMY © JON G. HOUSEMAN BIO 3323 Entomology mouthpart types by using following examples (Text fig . 1-5) and compare the various types to the descriptions provided in your text book. Grasshopper Chewing mouthparts are the least specialised and the grasshopper is an example of this type. Take a second look at the mouthparts from your specimen and the prepared slides. Be sure to identify the components so that you can keep track of them in the remaining specimens. House fly Sponging mouthparts. are found in the adults of some of the higher Diptera such as the housefly, Musca domestica. In these animals the labium is enlarged in a fleshy labellum with numerous small channels which by their capillary action draw up the fluids during feeding . These are called canaliculi. The labium and labrum form the food channel up which the liquefied food moves. The salivary canal is separate from the food canal and allows the salivary secretions to be pumped out while the ingested food passes into the organisms. Mandibles have disappeared in the flies and only the palps are the only maxillary features that remain. Figure 7 Sponging mouthparts of the Housefly. © BIODIDAC Compound eye Antenna Maxillary palp Labrum Labium Food canal Labium Hypopharynx Salivary canal Labium Labellum Stable fly Stable flies evolved from this same group of insects and had to modify the basic plan of the sponging mouthpart into piercing structure that allows them to obtain their blood meal. To do this the sponging labellum becomes hardened and only its tip, with the teeth like structures, remains flexible. Stable fly, Stomoxys calcitrans, mouthparts are available on a prepared slide. How does labial PAGE: 8 EXTERNAL ANATOMY © JON G. HOUSEMAN Entomology BIO 3323 modification allow for penetration through the skin in order to obtain the blood meal? Compare the stable fly to the tsetse fly that you’ll be looking at later in the lab. Compound eye Antenna Figure 8 Piercing mouthparts of the stable fly. © Maxillary palp BIODIDAC Labrum Labrum Food canal Salvary canal Maxillary palp Labium Hypopharynx Hypopharynx Labium Labellum Labellum Compound eye Antenna Figure 9 Mouthparts of the mosquito. © BIODI- Food canal Labrum DAC Mandible Hypopharynx Maxilla Salivary canal Labium Maxillary palp Mandible Maxilla Labrum Labium Hypopharynx Mosquito Piercing-sucking mouthparts are seen in insects such as mosquitoes, which represent the Diptera, and Hemiptera, such as Anasa tristis. In one case these mouthparts pierce skin and the other plant seeds. Look under the scope at the needle like maxilla and mandibles of these two different species. Butterfly Siphoning mouthparts are characteristic of Lepidoptera or butterflies. In this case the major part of the mouthpart is the galea and mandibles and labium are lost. Figure 10 Siphoning mouthparts of a butterfly. © BIODIDAC PAGE: 9 - EXTERNAL ANATOMY © JON G. HOUSEMAN BIO 3323 Entomology Antenna Compound eye Labrum Labial palp Maxillae Maxillae Nectar channel Horse fly Horse flies, Tabanus sp., and deer flies, Crysops sp., are more primitive dipterans than are the house fly and stable fly that you’ve already looked at. In these flies all of the ancestral mouthparts are still present and the horseflies use them in a very different way. The mandibles and maxilla resemble scissor blades and when the fly feeds it opens and closes these two sets of scissors cutting the skin surface. Blood pools inside the ragged wound and the fleshy labellum, with its sponging tip, sucks up the pooling blood. You’ll begin to understand why a horse fly bite is so painful! Figure 11 Mouthparts of horseflies or deerflies. Compound eye Antenna Maxillary palp Labrum Food canal Mandible Hypopharynx Maxilla Labium Hypopharynx Maxilla Labium Labella Mandible Labrum Tsetse fly The tsetse fly is a very important medical insect in Africa and it transmits African sleeping sickness. Tsetse’s also transmit a similar version of the disease to domestic cattle and its because of that large parts of the African plains are unavailable for agriculture (Depending on which side of the fence you sit, conservation vs. development, this is either a good or bad thing). Compare the mouthparts of the Tsetse PAGE: 10 EXTERNAL ANATOMY © JON G. HOUSEMAN Entomology BIO 3323 fly Glossina to those of the stable fly. How similar are they? Both of these flies are advanced Diptera and they have only the labellum and labrum and the hypopharynx inside to penetrate the host’s skin and ingest the bloodmeal. “Bugs” - Hemiptera The Hemiptera are known as the true bugs and the whole order specializes in piercing into their food source and sucking out the liquids. Originally that involved plants and the mouthparts worked their way through the plant tissues and tapped into the vessels that moved fluid through the plant. With the mouthparts anchored into these plant tubes the bug simply sucked up it’s meal. There are also blood feeding bugs and it’s assumed that over time some of these plant feeders probably pierced through the cuticle of other insects to feed and along with that became predacious. From there it was piercing vertebrates and drinking blood. Figure 12 Mouthparts of an hemipteran. Compound eye Labrum Maxilla Mandible Maxilla Food canal Salivary canal Labium Labium Mandibles Antenna Proboscis We have the plant bug Anasa tristis for you to take a look at. The maxilla form interlocking stylets with the mandibles on either side. The mandible dig in and then anchor in place and then the maxillary stylets are pushed forward looking for a vessel to pierce. If there isn’t one they then anchor in place the mandible move forward to dig a little deeper. The process repeats itself until a meal source has been located. Both the mandibles and maxilla are wrapped inside the labium to form the proboscis Fleas Adult fleas are unusual in that they have lost their wings and become ectoparasitic on vertebrate hosts. The mandible is completely missing and most of the work in feeding is done by the maxilla instead. The lacinia of the maxilla do the piercing and cutting while what were the galia form fleshy lobes. The maxillary lacinia combine with the PAGE: 11 - EXTERNAL ANATOMY © JON G. HOUSEMAN BIO 3323 Entomology epipharynx (there is some debate as to whether this is the hypopharynx or a new cuticular extension of the buccal cavity) and surrounded by the labial palps to form the food canal. Figure 13 Mouthparts of a flea. Antenna Ocellus Maxillary palps Maxilla Labial palps Epipharynx Maxillary lacinia Honey bee In the honey bee one ofthe biggest differences from that of other insects is the labium and maxilla are enlarged. The central glossa of the labium is modified into a curved tongue. The bee extends the “tongue” and it becomes covered in nectar. After that it’s pulled back into the surrounding tube that is created by the two maxilla and the labial palps. From there the ingested nectar is sucked up and into the digestive system. The mandibles are still present and used for eating pollen and for work that is done on the hive which is essentially built using the mandibles. When the mandibles are used the other mouthparts are folded back and out of the way. Figure 14 Mouthparts of a bee. Compound Eye Antenna Labrum Mandible Maxillary palp Maxilla Labial palp Labium Dragon fly nymph Dragon fly nymphs are deadly predator in freshwater lakes and streams the whole labium has been modified into a viscous prehensile structure with opposable labial palps at the tip that grap PAGE: 12 EXTERNAL ANATOMY © JON G. HOUSEMAN Entomology BIO 3323 onto the prey. The overlapping visual field of the large compound eyes is located precisely at the point of full extension of the labium. When a potential meal wanders into this field of view, the mouthparts shoot forward and immediately retract carrying the trapped prey to the mouth. Figure 15 Mouthparts of an Dragon fly nymph. Labrum Compound eye Prementum (Labium) Prementum (Labium) Maxilla Hypopharynx Labrum Postmentum (Labium) Prementum (Labium) Labial palp Legs Just as the mouthparts can be extensively modified so can insect legs. The basic insect is used for walking and this type is the primitive or ancestral form of the appendages. In many insects all three legs or only certain pairs are modified in different ways so that they can perform specialised functions such as swimming , climbing , jumping , digging and gathering specialised foods. The femur and tibia in running legs (cursorial) are long and thin. The increased length of the appendage means that the lever like motion of arthropod movement causes the distal end to move over a greater distance for the same amount of muscle movement. Jumping legs (saltatorial) have large femurs which contain the enlarged muscle PAGE: 13 - EXTERNAL ANATOMY © JON G. HOUSEMAN BIO 3323 Entomology required to extend the tibia. In jumping legs the femur and tibia are close to each at rest and the tarsal claws are often well developed so that the insect can get a good grip on the substrate when jumping . Raptorial legs are specially modified to grasp and retain prey. The classic example of this type of leg is the front leg of the preying mantid. The inner surfaces of the tibia and femur in raptorial legs are often equipped with sharp spines to help crush and immobilise the captured prey. Swimming (Natatorial) legs are modified to increase the available surface that pulls against the water during the swimming stroke. This can be achieved in one of two ways. Either the segments of the leg are flattened or are fringed with flexible hairs Digging legs are short and hardened and often flattened into shovel like shapes. They may also have heavy toothed projections that also assist in moving the soil. Male water beetles are faced with a unique problem and their front legs have been modified with suction cup like suckers so that they can remain attached to the female during copulation. WINGS The major taxa within the class Insecta are characterised by the presence or absence of wings and their structure and venation when present. The wingless insects are the Apterygota and the winged forms the Pterygota. The most primitive wings are found, as the name implies, in the Paleoptera with the rest of the winged insects being in the Division Neoptera. Paleopteran insects include the Odonata, the dragonflies and damselflies, and possibly the Ephemeroptera, the mayflies or shad flies as they are called locally. Entomologists working on Insect Phylogeny sometimes separate the two but we'll consider both as Paleoptera. Insects in the Odonata and Ephemeroptera are unable to fold their wings back and over the abdomen because the axillary sclerites that allow this in other insects remain fused to the wing veins. The result is that at rest a dragonfly's wings stick out from the side and damselflies and ephemoropteran wings are held above the abdomen. Some of the Neoptera also hold their wings in a similar manner but the difference is that either the forewing , or most often the hind wing , has additional folds not seen in the Paleoptera or the wings surround or partially cover the abdomen. Although it's not unique to them the large numbers of cross veins are also a good way to identify the Paleoptera. Dragonflies and damselflies are available for you to examine. PAGE: 14 EXTERNAL ANATOMY © JON G. HOUSEMAN Entomology BIO 3323 Figure 16 Ephemeroptera are often included in the Paleoptera and they hold their unfolded wings high above the body - a characteristic of the Paleoptera Within the Neoptera there are a variety of different ways that wings are modified and, as you'll see when we start identifying insects, not only the folding , but the similarities and/or differences between the forewing and hindwing and their venation are important taxonomic tools. Stoneflies, Plecoptera, have lots of cross veins in their wings but look closely at how they are positioned over top of the abdomen. Take a closer look at the hind wing and you'll see out it unfolds like a fan. This difference in shape is one of the ways that insect forewings may have a different appearance than the hindwing . Figure 17 Stoneflies have lots of cross-veins in their wings this and their mandibualte appearance shouldn’t confuse you. The hindwings fold and these are neopterans. © BIODIDAC The Orthoptera, which includes crickets and grasshoppers among others, have leathery forewings compared to the membranous hindwings folded underneath. With your first look at the grasshoppers you might disagree but look closely that hind wing is folded on itself a number of times. Unfold it to see the difference in texture between the two wings, another way wing pairs can differ from each other. The Hemiptera, or true bugs, also have forewings that differ from the hind wings. In this case the forewing as thicker at it's base and membranous at the overlapping tips. The half hardened wing is the origin of their name the Hemiptera. Look for these characteristics on the giant water bug . PAGE: 15 - EXTERNAL ANATOMY © JON G. HOUSEMAN BIO 3323 Entomology Figure 18 The true bugs, Hemiptera have a characteristic forewing that is leathery at its base and membranous at the overlapping tips. This dorsal view makes it easy to identify Hemiptera. © BIODIDAC The Homoptera include plant sap feeding insects such as aphids, leaf hoppers and cicadas, to name a few. Their wings differ in shape but both have a membranous texture. The forewings are held over the abdomen covering both the dorsal and lateral sides. In the keys this is referred to as roof-like and shouldn't be confused with the way the damselflies and mayflies hold their wings flat against each other and completely above the abdomen. A cicada is available for you to look at. Figure 19 Homopteran wings are both membranous but are helf like o roof covering the dorsal and lateral sides of the abdomen © BIODIDAC The greatest difference between the two pairs of wings occurs in the beetles, Coleopetra. Here the front wing is hardened into elytra protecting the delicate hind wings underneath. The degree of hardness varies from leathery to hard. The elytra are not involved in flight and are lifted out of the way as the insect flies using its hindwings. Soldier beetles and dogbane beetles are available. If during your observations of the hind wing the forewing breaks off discard the specimen. Not all insects have four wings, the Diptera or flies are characterised by only two, a single pair, of visible wings. The second set is actually still there, but reduced to halteres. Find them on the blowflies that have been provided. Only the Diptera have two pairs of wings but there are others that can fool you. Many insect groups lock the two pairs of wings together PAGE: 16 EXTERNAL ANATOMY © JON G. HOUSEMAN Entomology BIO 3323 so that they function as a single wing . A good example of this is seen in the wasps (Hymenoptera). A series of hooks on the leading edge of the hind wing hold it the trailing edge of the forewing . Look for the coupling mechanism on the wasps that have been provided. Lepidoptera also couple their wings together using frenula, a long bristle that arises at the base of the hindwing and fits into a set of hook like scales on the forewing . Geometrid butterflies are available for you to look at. PAGE: 17 - EXTERNAL ANATOMY © JON G. HOUSEMAN