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Lab handout for External anatomy of the Atelocerata - Salinella

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Insect <strong>External</strong> Anatomy<br />

Insect <strong>External</strong> Anatomy © Houseman – page 1<br />

by Jon G. Houseman<br />

In this lab we continue or observations <strong>of</strong> <strong>the</strong> external <strong>anatomy</strong> we started previously.<br />

This time we look at insects and use <strong>the</strong> grasshopper and cockroach to complete <strong>the</strong><br />

comparative grid started in <strong>the</strong> last lab where we examined panarthopods and o<strong>the</strong>r<br />

insect relatives. We’ll finish up our investigations <strong>of</strong> external <strong>anatomy</strong> with a<br />

comprehensive look at insect mouthparts and some different types <strong>of</strong> insect legs. An<br />

autapomorphy <strong>of</strong> <strong>the</strong> kingdom Animalia is being heterotrophs and a symplesiomorphy<br />

that Insects share with <strong>the</strong>ir sister groups is that <strong>the</strong>y manipulate <strong>the</strong>ir food with <strong>the</strong>ir<br />

appendages. Insect cuticle can be molded into any one <strong>of</strong> a variety <strong>of</strong> structures and<br />

this applies to <strong>the</strong> appendages used <strong>for</strong> feeding. Is it a surprise that insects have shaped<br />

<strong>the</strong>ir feeding appendages into <strong>the</strong> variety <strong>of</strong> structures you will be looking at later?<br />

And, <strong>the</strong> more different ways you can successfully feed on unique foods is certainly a<br />

<strong>for</strong>mula <strong>for</strong> diversification. The same diversification applies to <strong>the</strong> different types <strong>of</strong><br />

insect legs.<br />

Grasshopper: Romalea guttata<br />

Orthoptera, <strong>the</strong> insect order that includes <strong>the</strong> grasshopper, provides excellent<br />

specimens <strong>for</strong> studying many ancestral features <strong>of</strong> insects, such as mouthparts and <strong>the</strong><br />

architecture <strong>of</strong> <strong>the</strong> tagmata that <strong>for</strong>m <strong>the</strong> body. The Lubber grasshopper, Romalea<br />

guttata, is a large species and <strong>the</strong>re<strong>for</strong>e an ideal hopper <strong>for</strong> lab observations. Unlike<br />

most hoppers, however, this species cannot fly because it has small wings; so it<br />

disperses by hopping. It’s found in <strong>the</strong> sou<strong>the</strong>astern United States.<br />

<strong>External</strong> <strong>anatomy</strong><br />

The basic arthropod body plan is metameric, with <strong>the</strong> ancestral segments, metameres,<br />

fused to <strong>for</strong>m larger tagmata. Identify <strong>the</strong> three major insect tagmata in your<br />

specimen: head, thorax, and abdomen (Figure 1). What are <strong>the</strong> functions<br />

associated with each <strong>of</strong> <strong>the</strong> different insect tagmata?<br />

Figure 1 Major anatomical features <strong>of</strong> an insect.<br />

The arthropods, including insects, are covered with an exoskeleton (cuticle) produced<br />

by an underlying epidermis. In <strong>the</strong> most primitive arthropods, ei<strong>the</strong>r two or three


cuticular plates, <strong>the</strong> sclerites, surround each segment. A sclerite on <strong>the</strong> dorsal surface<br />

is called a tergite, <strong>the</strong> plate on <strong>the</strong> ventral surface is called a sternite, and <strong>the</strong> cuticular<br />

plates on <strong>the</strong> sides are called pleurites. In <strong>the</strong> abdomen, <strong>the</strong>re are only two sclerites,<br />

<strong>the</strong> tergite and sternite with and pleural membranes, instead <strong>of</strong> solid pleurites,<br />

connecting <strong>the</strong> two. Membranes between <strong>the</strong> plates allowed adjacent sclerites to move<br />

relative to each o<strong>the</strong>r. For example, <strong>the</strong> pleural membrane allows <strong>the</strong> abdomen to<br />

expand during feeding, or during egg production in <strong>the</strong> female. As metameres became<br />

fused into tagma, <strong>the</strong> articulation between <strong>the</strong> various sclerites <strong>of</strong>ten disappeared, and<br />

what remains <strong>of</strong> <strong>the</strong> junction between <strong>the</strong> two sclerites is a groove or indentation on<br />

<strong>the</strong> surface <strong>of</strong> <strong>the</strong> cuticle and is referred to as a suture. Often, a suture is more than a<br />

line on <strong>the</strong> surface <strong>of</strong> <strong>the</strong> cuticle, and internally <strong>the</strong> cuticle folds to create an internal<br />

ridge. What is <strong>the</strong> advantage <strong>of</strong> <strong>the</strong>se areas <strong>of</strong> infolded cuticle <strong>for</strong> muscle<br />

attachment? Not all sutures reflect ancestral segmentation <strong>of</strong> <strong>the</strong> tagma, and o<strong>the</strong>r<br />

indentations <strong>of</strong> cuticle, apodemes <strong>for</strong> example, also provide internal sites <strong>for</strong> muscle<br />

attachment.<br />

Head<br />

The insect head is a fusion <strong>of</strong> six segments, three <strong>of</strong> which are preoral (pregnathal),<br />

and three are postoral (postgnathal). The appendages on <strong>the</strong> preoral segments include<br />

<strong>the</strong> labrum and antennae; <strong>the</strong>re are no appendages on <strong>the</strong> third head segment. In <strong>the</strong><br />

closely related subphylum <strong>of</strong> mandibulate crustaceans, this third head segment had <strong>the</strong><br />

second pair <strong>of</strong> antennae. In <strong>Atelocerata</strong> <strong>the</strong> second pair <strong>of</strong> antennae have been lost and<br />

this is <strong>the</strong> origin <strong>of</strong> <strong>the</strong> subphylum's taxonomic name. The postoral appendages from<br />

front to back are as follows: mandible, maxillae , and fused labium. The labrum and<br />

labium <strong>for</strong>m <strong>the</strong> top and bottom <strong>of</strong> <strong>the</strong> preoral cavity respectively; <strong>the</strong> mandibles and<br />

maxilla <strong>for</strong>m <strong>the</strong> sides.<br />

Figure 2 Lateral view <strong>of</strong> <strong>the</strong> grasshopper head.<br />

The head capsule (Figure 2 and Figure 3) is almost a solid piece <strong>of</strong> hardened cuticle,<br />

and most <strong>of</strong> <strong>the</strong> sutures and lines visible on its surface are <strong>the</strong> apodemes <strong>of</strong> internal<br />

cuticular ridges that prevent <strong>the</strong> head capsule from collapsing under <strong>the</strong> <strong>for</strong>ce <strong>of</strong> <strong>the</strong><br />

mouthpart musculature. Only <strong>the</strong> posterior sutures near <strong>the</strong> neck reflect <strong>the</strong> original<br />

segmentation <strong>of</strong> <strong>the</strong> head. You will look at <strong>the</strong>se posterior sutures in a moment; <strong>for</strong><br />

now begin with <strong>the</strong> front <strong>of</strong> <strong>the</strong> head. Be<strong>for</strong>e you start your observation <strong>of</strong> <strong>the</strong> external<br />

<strong>anatomy</strong>, be sure that <strong>the</strong> surface <strong>of</strong> <strong>the</strong> specimen is dry; many <strong>of</strong> <strong>the</strong> smaller details<br />

Insect <strong>External</strong> Anatomy © Houseman – page 2


will be hard to see if <strong>the</strong> specimen is wet. If necessary, place <strong>the</strong> specimen on a piece<br />

<strong>of</strong> paper towel and wait <strong>for</strong> it to dry <strong>of</strong>f.<br />

A pair <strong>of</strong> compound eyes is located on <strong>the</strong> sides and near <strong>the</strong> top <strong>of</strong> <strong>the</strong> head (Figure<br />

2). Take a close look under <strong>the</strong> dissecting microscope, and you will see <strong>the</strong> complex<br />

pattern <strong>of</strong> ommatidia, <strong>the</strong> fundamental repeating optic units <strong>of</strong> <strong>the</strong> arthropod eye. The<br />

part <strong>of</strong> <strong>the</strong> ommatidia you see is <strong>the</strong> cornea (lens). In addition to <strong>the</strong> pair <strong>of</strong> compound<br />

eyes, <strong>the</strong>re are three simple eyes called <strong>the</strong> ocelli. A pair <strong>of</strong> <strong>the</strong>se is situated between<br />

<strong>the</strong> base <strong>of</strong> <strong>the</strong> antennae, and <strong>the</strong> compound eyes. A third medial ocellus is located on<br />

<strong>the</strong> anterior surface <strong>of</strong> <strong>the</strong> head, below <strong>the</strong> groove that runs between <strong>the</strong> antennae and<br />

<strong>the</strong> compound eyes. Cross sections through an insect head are available that show <strong>the</strong><br />

compound and simple eyes. Be sure to look at <strong>the</strong>se specimens and compare <strong>the</strong>m to<br />

<strong>the</strong> figures in you lecture notes. The number and position <strong>of</strong> <strong>the</strong> ocelli varies in<br />

Figure 3 Anterior view <strong>of</strong> <strong>the</strong> grasshopper head<br />

different insects, but this arrangement <strong>of</strong> three is <strong>the</strong> ancestral pattern <strong>for</strong> <strong>the</strong> ocelli.<br />

Can ocelli <strong>for</strong>m an image; what is <strong>the</strong>ir function?<br />

A single pair <strong>of</strong> antennae is characteristic <strong>of</strong> <strong>the</strong> <strong>Atelocerata</strong>, and each antenna is<br />

composed <strong>of</strong> three basic parts: <strong>the</strong> scape, which is most proximal, <strong>the</strong> pedicel, and <strong>the</strong><br />

flagellum, which is most distal. The flagellum is a single segment, but <strong>the</strong>re is a series<br />

<strong>of</strong> annulations that give it a segmented appearance; <strong>the</strong> annulations help it bend. How<br />

does <strong>the</strong> arrangement <strong>of</strong> <strong>the</strong> internal musculature differ between annulations and<br />

a segment in <strong>the</strong> antennae? Examine <strong>the</strong> antennae closely using <strong>the</strong> dissecting<br />

microscope to see <strong>the</strong> sensory setal hairs that cover its surface. Although some setal<br />

hairs are mechanoreceptive, <strong>the</strong>y primarily have a chemosensory role. During flight,<br />

<strong>the</strong> antennae bend and move, providing important sensory in<strong>for</strong>mation on wind speed,<br />

direction, and resistance. This twisting and bending <strong>of</strong> <strong>the</strong> flagellum generates <strong>for</strong>ces<br />

in <strong>the</strong> cuticle <strong>of</strong> <strong>the</strong> pedicel, and <strong>the</strong>se pressures are detected by a special<br />

mechanoreceptive structure known as Johnston’s organ. All flying insects have a<br />

Johnston’s organ that provides in<strong>for</strong>mation that enables <strong>the</strong> insect to adjust its flight to<br />

compensate <strong>for</strong> wind currents.<br />

The o<strong>the</strong>r appendages on <strong>the</strong> head are mouthparts. The fleshy lip is <strong>the</strong> labrum, which<br />

is attached to <strong>the</strong> head by <strong>the</strong> clypeus. Look at <strong>the</strong> side <strong>of</strong> <strong>the</strong> head to see <strong>the</strong> powerful<br />

and hardened mandibles (Figure 2).Behind <strong>the</strong> mandibles, <strong>the</strong>re are <strong>the</strong> maxillae with<br />

<strong>the</strong>ir sensory maxillary palps. The last set <strong>of</strong> appendages on <strong>the</strong> head is <strong>the</strong> fused<br />

Insect <strong>External</strong> Anatomy © Houseman – page 3


appendages <strong>of</strong> <strong>the</strong> labium, which also have sensory labial palps. You will take a<br />

closer look at prepared slides <strong>of</strong> <strong>the</strong> hopper mouthparts later in <strong>the</strong> lab when we<br />

compare <strong>the</strong> ancestral pattern with mouthparts that are specialized <strong>for</strong> different types<br />

<strong>of</strong> food.<br />

The head capsule is divided into a number <strong>of</strong> regions, or plates, by sutures and lines.<br />

The top <strong>of</strong> <strong>the</strong> head is <strong>the</strong> vertex (Figure 3). From <strong>the</strong> vertex a set <strong>of</strong> lines appears as<br />

an inverted Y; this is <strong>the</strong> epicranial suture and when <strong>the</strong> hopper molts, <strong>the</strong>se lines<br />

break open, and <strong>the</strong> insect emerges from <strong>the</strong> old cuticle. The cuticle on <strong>the</strong> sides <strong>of</strong> <strong>the</strong><br />

head and behind <strong>the</strong> epicranial suture is <strong>the</strong> gena, and <strong>the</strong> part <strong>of</strong> <strong>the</strong> cuticle between<br />

<strong>the</strong> branched base <strong>of</strong> <strong>the</strong> epicranial sutures is <strong>the</strong> frons. Behind <strong>the</strong> gena <strong>the</strong>re are two<br />

more saddle-like plates that cover <strong>the</strong> dorsal and lateral surfaces <strong>of</strong> <strong>the</strong> head: <strong>the</strong><br />

occiput and postocciput, separated from each o<strong>the</strong>r by <strong>the</strong> postoccipital suture (Figure<br />

3). The postoccipital suture is <strong>the</strong> only suture on <strong>the</strong> head that identifies <strong>the</strong> original<br />

segments <strong>of</strong> <strong>the</strong> head, <strong>the</strong> fusion <strong>of</strong> <strong>the</strong> fifth and sixth segments. The posterior opening<br />

to <strong>the</strong> head is <strong>the</strong> <strong>for</strong>amen.<br />

Two connected sutures run horizontally across <strong>the</strong> head, above to <strong>the</strong> attachment <strong>of</strong><br />

<strong>the</strong> mouthparts. Inside <strong>the</strong> head and beneath <strong>the</strong> two sutures is a ridge <strong>of</strong> cuticle that<br />

encircles <strong>the</strong> head. The mouthparts articulate with <strong>the</strong> head in this region, and this<br />

ridge <strong>of</strong> cuticle rein<strong>for</strong>ces <strong>the</strong> head against <strong>the</strong> <strong>for</strong>ces generated by <strong>the</strong> musculature <strong>of</strong><br />

<strong>the</strong> mouthparts. The small pits at <strong>the</strong> lateral edges <strong>of</strong> <strong>the</strong> suture are <strong>the</strong> apodemes that<br />

<strong>for</strong>m <strong>the</strong> endoskeletal tentorium <strong>of</strong> <strong>the</strong> head. The o<strong>the</strong>r apodemes <strong>of</strong> <strong>the</strong> tentorium are<br />

located at <strong>the</strong> base <strong>of</strong> <strong>the</strong> postoccipital suture. The tentorium creates internal “flying<br />

buttresses” from <strong>the</strong> back and <strong>the</strong> front to <strong>the</strong> top <strong>of</strong> <strong>the</strong> head, ensuring that <strong>the</strong><br />

cranium doesn’t collapse under <strong>the</strong> <strong>for</strong>ces created by <strong>the</strong> mouthparts. Cleared<br />

specimens <strong>of</strong> <strong>the</strong> hopper head are available. Please handle <strong>the</strong>m carefully and look<br />

inside <strong>the</strong> head through <strong>the</strong> <strong>for</strong>amen to see components <strong>of</strong> <strong>the</strong> tentorium.<br />

Figure 4 Major sclerites and sutures <strong>of</strong> <strong>the</strong> prothoracic segment <strong>of</strong> an insect<br />

Thorax<br />

Insects have two pairs <strong>of</strong> wings, and in <strong>the</strong> grasshoppers <strong>the</strong> two wings have a<br />

different appearance. The <strong>for</strong>ewings, attached to <strong>the</strong> mesothorax, are lea<strong>the</strong>ry and<br />

Insect <strong>External</strong> Anatomy © Houseman – page 4


protect <strong>the</strong> hindwings underneath. This is <strong>of</strong>ten <strong>the</strong> case among <strong>the</strong> insects, and beetles<br />

perfect this strategy by having hardened <strong>for</strong>ewings called elytra that are not used <strong>for</strong><br />

flying, but as protective wing covers. You can see <strong>the</strong> difference between <strong>the</strong> wings by<br />

unfolding <strong>the</strong> hindwing and comparing it to <strong>the</strong> <strong>for</strong>ewing. The wings are supported by<br />

a series <strong>of</strong> veins, but don’t let <strong>the</strong> term vein confuse you; in insects <strong>the</strong> word has<br />

nothing to do with <strong>the</strong> circulatory system <strong>of</strong> an insect. The veins are ridges <strong>of</strong> cuticle<br />

that streng<strong>the</strong>n <strong>the</strong> wing. In <strong>the</strong> Lubber grasshopper, both sets <strong>of</strong> wings are small and<br />

not large enough <strong>for</strong> flight.<br />

The thorax is composed <strong>of</strong> three segments: <strong>the</strong> prothorax, mesothorax, and<br />

metathorax. The thoracic segments can be divided into three major sclerites that have<br />

<strong>the</strong>ir own special names: <strong>the</strong> dorsal notum (tergite), lateral pleurites, and <strong>the</strong> sternum<br />

(sternite). Each <strong>of</strong> <strong>the</strong>se thoracic segments bears a pair <strong>of</strong> appendages and wings<br />

located on <strong>the</strong> mesothoracic and metathoracic segments, which toge<strong>the</strong>r <strong>for</strong>m <strong>the</strong><br />

pterothorax.<br />

Figure 5 Major sclerites and sutures <strong>of</strong> <strong>the</strong> insect metathorax.<br />

The wingless prothorax is <strong>for</strong>med from <strong>the</strong> fusion <strong>of</strong> <strong>the</strong> four sclerites : a notum,<br />

sternum and two lateral pleurites . The dorsal notum, <strong>the</strong> pronotum, is enlarged and<br />

saddle-like, and extends down <strong>the</strong> sides <strong>of</strong> <strong>the</strong> segment. Ancestrally, each <strong>of</strong> <strong>the</strong><br />

thoracic and abdominal segments had spiracles. Locate <strong>the</strong> spiracle on <strong>the</strong> prothorax.<br />

Articulation <strong>of</strong> <strong>the</strong> prothoracic leg with <strong>the</strong> thorax follows <strong>the</strong> same pattern as <strong>the</strong><br />

pterothorax (Figure 4). On each segment <strong>of</strong> <strong>the</strong> pterothorax, lateral pleurites are fused<br />

with <strong>the</strong> ventral sternites, <strong>for</strong>ming a three-sided thoracic box. The notum is divided<br />

into <strong>the</strong> anterior alinotum and behind it, <strong>the</strong> postnotum (Figure 5). The postnotum is<br />

continuous with <strong>the</strong> posterior edges <strong>of</strong> left and right pleurites, and it <strong>for</strong>ms <strong>the</strong> upper<br />

surface <strong>of</strong> <strong>the</strong> rigid thoracic box.<br />

Insect <strong>External</strong> Anatomy © Houseman – page 5


The alinotum is connected to <strong>the</strong> membranous part <strong>of</strong> <strong>the</strong> pleura that lies below, and<br />

this allows <strong>the</strong> alinotum to move up and down relative to <strong>the</strong> box underneath. Look<br />

closely at <strong>the</strong> pleural plates <strong>of</strong> <strong>the</strong> mesothorax and metathorax. A central pleural<br />

suture divides <strong>the</strong> pleurite in half and <strong>the</strong> pleural process is an extension <strong>of</strong> cuticle at<br />

<strong>the</strong> dorsal end <strong>of</strong> <strong>the</strong> pleural suture. It provides <strong>the</strong> central point <strong>of</strong> articulation <strong>for</strong> <strong>the</strong><br />

wings, which pivot on <strong>the</strong> upper edges <strong>of</strong> <strong>the</strong> pleural process when pushed up and<br />

down by <strong>the</strong> flight muscles. As <strong>the</strong> notum rises, <strong>the</strong> wings are lowered; as it falls, <strong>the</strong><br />

Figure 6 Anatomy <strong>of</strong> a typical insect leg.<br />

wings are raised. Similar to <strong>the</strong> structure <strong>of</strong> <strong>the</strong> head, <strong>the</strong> pleural suture has a ridge <strong>of</strong><br />

cuticle folded inside, and this cuticle braces <strong>the</strong> thoracic box against <strong>the</strong> muscular<br />

<strong>for</strong>ces <strong>of</strong> <strong>the</strong> flight muscles. Like <strong>the</strong> pleurites, <strong>the</strong> sternite is also divided into anterior<br />

and posterior portions separated by a suture that functions as an internal rein<strong>for</strong>cing<br />

apodeme. Take a close look at <strong>the</strong> suture and locate <strong>the</strong> external apodemes, apophyseal<br />

pits, which, along with <strong>the</strong> plural suture, maintain <strong>the</strong> rigidity <strong>of</strong> <strong>the</strong> thoracic box when<br />

<strong>the</strong> flight muscles contract.<br />

The three pairs <strong>of</strong> legs are attached on <strong>the</strong> ventral side <strong>of</strong> <strong>the</strong> pleurites and articulate<br />

with <strong>the</strong> ventral end <strong>of</strong> <strong>the</strong> pleural suture and with <strong>the</strong> pleurite. Each leg has six tubular<br />

segments, which means that each joint can only move in one direction (Figure 6). As<br />

you flex each <strong>of</strong> <strong>the</strong> segments, do <strong>the</strong>y all move in <strong>the</strong> same direction relative to<br />

each o<strong>the</strong>r? The segment closest to <strong>the</strong> body is <strong>the</strong> coxa, and this is followed by <strong>the</strong><br />

trochanter, <strong>the</strong> large femur, and <strong>the</strong> b. In <strong>the</strong> hopper, <strong>the</strong> trochanter is fused with <strong>the</strong><br />

femur; so be sure to find <strong>the</strong> suture between <strong>the</strong> two. The fifth segment is <strong>the</strong> tarsus,<br />

and in insects it is divided by a series <strong>of</strong> up to five articulations called tarsomeres<br />

with <strong>the</strong> pretarsal segment and claws at <strong>the</strong> tip <strong>of</strong> <strong>the</strong> last tarsomere. The prothoracic<br />

and mesothoracic legs are very similar to each o<strong>the</strong>r; <strong>the</strong> metathoracic leg is modified<br />

<strong>for</strong> jumping and inside are <strong>the</strong> muscles that elongate <strong>the</strong> leg <strong>for</strong> <strong>the</strong> jump.<br />

Abdomen<br />

Ancestrally, <strong>the</strong> insect abdomen had eleven segments, but it’s <strong>of</strong>ten difficult to see all<br />

<strong>the</strong>se because some have become fused toge<strong>the</strong>r, or reduced in size, or modified<br />

because <strong>of</strong> <strong>the</strong> genitalia associated with <strong>the</strong> posterior segments. The first seven<br />

abdominal segments have <strong>the</strong> usual sclerotized tergite and sternite, and membranous<br />

pleura and spiracular opening. The terminal abdominal segments bear appendages<br />

involved in copulation and egg laying.<br />

Identify <strong>the</strong> tergite, sternite, pleural membrane, and spiracle on any <strong>of</strong> <strong>the</strong> first seven<br />

pregenital segments <strong>of</strong> your specimen (Figure 7). With a few exceptions, <strong>the</strong>se<br />

Insect <strong>External</strong> Anatomy © Houseman – page 6


segments will look identical. The exceptions include <strong>the</strong> tympanum on <strong>the</strong> first<br />

abdominal tergite, and <strong>the</strong> extension <strong>of</strong> <strong>the</strong> metathoracic sternite into a groove on <strong>the</strong><br />

first abdominal sternite. The whole first abdominal segment is fused to <strong>the</strong> thorax, and<br />

Figure 7 <strong>External</strong> <strong>anatomy</strong> <strong>of</strong> <strong>the</strong> grasshopper abdomen<br />

this modification <strong>of</strong> <strong>the</strong> sternite helps to lock <strong>the</strong> abdomen to <strong>the</strong> thorax. The air sac<br />

behind <strong>the</strong> tympanal membrane is connected to <strong>the</strong> rest <strong>of</strong> <strong>the</strong> tracheal system and it<br />

opens to <strong>the</strong> outside through <strong>the</strong> spiracle beside <strong>the</strong> tympanum. Why is it important<br />

to have an air sac behind <strong>the</strong> tympanum? The remaining segments are <strong>the</strong> genital<br />

segments; <strong>the</strong>se retain ancestral appendages that have been modified into <strong>the</strong> genitalia<br />

<strong>of</strong> <strong>the</strong> male and female. Look at both male and female specimens.<br />

In <strong>the</strong> female, <strong>the</strong> genital plate is <strong>the</strong> eighth sternite, <strong>the</strong> last one visible on <strong>the</strong> ventral<br />

surface <strong>of</strong> <strong>the</strong> body. Although <strong>the</strong> eighth tergite is unmodified, tergites nine and ten<br />

are partially fused. The eleventh tergite, an enlarged triangular plate, is <strong>the</strong> epiproct,<br />

which has <strong>the</strong> anus underneath it. Identify <strong>the</strong> short sensory circus, a modified<br />

eleventh appendage, located below <strong>the</strong> eleventh tergite. In <strong>the</strong> general plan <strong>of</strong> <strong>the</strong><br />

female genitalia, <strong>the</strong> paired appendages <strong>of</strong> segments eight and nine <strong>for</strong>m <strong>the</strong><br />

ovipositor. The valves surround a central egg canal inside, and <strong>the</strong> gonopore opens at<br />

<strong>the</strong> base <strong>of</strong> <strong>the</strong> egg canal. The first paired valves <strong>of</strong> <strong>the</strong> ovipositor are on segment<br />

eight, and <strong>the</strong> second and third valves are on segment ten. In <strong>the</strong> hopper, valves one<br />

and three are large and heavily sclerotized; <strong>the</strong> second valve is reduced in size. The<br />

two valves in each pair interlock and are used to dig into <strong>the</strong> soil be<strong>for</strong>e <strong>the</strong> eggs are<br />

laid.<br />

In <strong>the</strong> male, <strong>the</strong> ancestral appendages at <strong>the</strong> tip <strong>of</strong> <strong>the</strong> abdomen are involved in<br />

transferring sperm, usually inside a spermatophore, to <strong>the</strong> female. It’s not easy to<br />

define a generalized plan <strong>for</strong> <strong>the</strong> male, and a variety <strong>of</strong> different morphologies and<br />

terms are used <strong>for</strong> <strong>the</strong> male genitalia in insects. Instead <strong>of</strong> being constructed from <strong>the</strong><br />

appendages <strong>of</strong> <strong>the</strong> eighth and nine segments, male genitalia are <strong>for</strong>med from <strong>the</strong> ninth.<br />

In your specimen, both <strong>the</strong> eighth and ninth sternites are visible, and <strong>the</strong> ninth <strong>for</strong>ms<br />

both <strong>the</strong> genital plate and <strong>the</strong> ancestral appendage that surrounds <strong>the</strong> male aedeagus<br />

that passes sperm to <strong>the</strong> female. On <strong>the</strong> dorsal surface, <strong>the</strong> tergites resemble those <strong>of</strong><br />

<strong>the</strong> female, although <strong>the</strong> ninth and tenth are not fused.<br />

Insect <strong>External</strong> Anatomy © Houseman – page 7


Cockroach: Periplaneta americana<br />

Cockroaches, in <strong>the</strong> insect order Blattodea, represent some <strong>of</strong> <strong>the</strong> most ancient and<br />

successful insects, and <strong>the</strong> best example is <strong>the</strong> roach Periplaneta americana that you<br />

will study in this lab. This cosmopolitan species can be found wherever <strong>the</strong>re are<br />

humans. Despite <strong>the</strong> word americana in <strong>the</strong>ir name, <strong>the</strong>y originate in Africa and are<br />

believed to have first appeared in North America in <strong>the</strong> early 1600s, soon after <strong>the</strong><br />

continent became colonized. Their universal distribution is <strong>the</strong> result <strong>of</strong> global<br />

shipping, from <strong>the</strong> age <strong>of</strong> sailing vessels to <strong>the</strong> cargo planes <strong>of</strong> today. Their success<br />

can be attributed to <strong>the</strong>ir generalized body plan, which also makes this an excellent<br />

species <strong>for</strong> examining <strong>the</strong> anatomical features <strong>of</strong> an insect.<br />

<strong>External</strong> <strong>anatomy</strong><br />

Use <strong>the</strong> preserved roaches <strong>for</strong> your observations <strong>of</strong> <strong>the</strong> external <strong>anatomy</strong>.<br />

Figure 8 Dorsal view <strong>of</strong> <strong>the</strong> main external features <strong>of</strong> a cockroach<br />

Head<br />

The head capsule <strong>of</strong> <strong>the</strong> roach is very small, and you will need to use <strong>the</strong> dissecting<br />

scope to see <strong>the</strong> various structures (Figure 8 and Figure 9). Two large, paired,<br />

compound eyes are visible on <strong>the</strong> sides <strong>of</strong> <strong>the</strong> head and like you did <strong>for</strong> <strong>the</strong><br />

grasshopper look <strong>for</strong> <strong>the</strong> individual ommatidia that are <strong>the</strong> fundamental units <strong>of</strong> <strong>the</strong><br />

compound eye. Most insects have an epicranial suture that starts at <strong>the</strong> top <strong>of</strong> <strong>the</strong> head<br />

and branches below <strong>the</strong> eyes; it splits open when <strong>the</strong> insect molts, but un<strong>for</strong>tunately, in<br />

<strong>the</strong> roach <strong>the</strong> suture is almost impossible to see as are many <strong>of</strong> <strong>the</strong> cranial sutures you<br />

saw in <strong>the</strong> grasshopper. In addition to <strong>the</strong> compound eyes, <strong>the</strong>re are ocelli, known as<br />

simple eyes, but <strong>the</strong>se will be hard to see in your specimen.<br />

A single pair <strong>of</strong> antennae is characteristic <strong>of</strong> <strong>the</strong> <strong>Atelocerata</strong>, and each antenna is<br />

composed <strong>of</strong> three basic parts: <strong>the</strong> scape, which is most proximal, <strong>the</strong> pedicel, and <strong>the</strong><br />

annulated flagellum, which is most distal.<br />

Insect <strong>External</strong> Anatomy © Houseman – page 8


The mandibulate mouthparts <strong>of</strong> <strong>the</strong> roach resemble those <strong>of</strong> <strong>the</strong> generalized<br />

“grasshopper-like” or primitive mouthparts you will examine later in <strong>the</strong> lab when we<br />

compare <strong>the</strong> mouthparts <strong>of</strong> different insects. Because <strong>the</strong> head capsule <strong>of</strong> a roach is<br />

very small, it is almost impossible to remove <strong>the</strong> individual mouthparts. Try and<br />

differentiate between <strong>the</strong> labrum, and <strong>the</strong> labium with its labial palps. Without<br />

removing mouthparts <strong>the</strong> labium and maxilla are best seen as a set <strong>of</strong> palps. If you pull<br />

<strong>the</strong> set <strong>of</strong> labial and maxillary palps back you should also be able to see <strong>the</strong> outer<br />

surface <strong>of</strong> <strong>the</strong> mandible.<br />

Thorax<br />

The two pairs <strong>of</strong> wings that are found in insects have a different appearance in<br />

cockroaches (Figure 8). The <strong>for</strong>ewings, attached to <strong>the</strong> mesothorax, are lea<strong>the</strong>ry<br />

tegmen and protect <strong>the</strong> more delicate hindwings on <strong>the</strong> metathorax. You can see <strong>the</strong><br />

difference between <strong>the</strong> wings by unfolding <strong>the</strong> hindwing and comparing it to <strong>the</strong><br />

<strong>for</strong>ewing.<br />

Figure 9 Ventral view <strong>of</strong> <strong>the</strong> main external features <strong>of</strong> a cockroach.<br />

The wings are supported by a series <strong>of</strong> veins, but don’t let <strong>the</strong> term vein confuse you;<br />

in insects <strong>the</strong> word has nothing to do with <strong>the</strong> circulatory system. The veins are ridges<br />

<strong>of</strong> cuticle that streng<strong>the</strong>n <strong>the</strong> wing, and, starting at <strong>the</strong> anterior edge, include <strong>the</strong> costa<br />

on <strong>the</strong> leading edge and <strong>the</strong> subcosta behind it. The radius, medial, and cubital veins<br />

are behind <strong>the</strong> subcosta, and are part <strong>of</strong> <strong>the</strong> remigium. Behind <strong>the</strong> remigium is <strong>the</strong><br />

vannus region. This region in roaches is very large and includes <strong>the</strong> highly branched<br />

vannal and anal veins. Most <strong>of</strong> <strong>the</strong> folding <strong>of</strong> <strong>the</strong> wing involves this vannus region.<br />

Identify <strong>the</strong> three thoracic segments (Figure 8): prothorax, mesothorax, and<br />

metathorax and how <strong>the</strong>y are divided into <strong>the</strong> dorsal notum (tergite), lateral<br />

pleurites, and <strong>the</strong> sternum (sternite) and <strong>the</strong> two pairs <strong>of</strong> wings attached to <strong>the</strong><br />

pterothorax. In <strong>the</strong> roaches <strong>the</strong> pronotum is enlarged into a pronotal shield that<br />

protects <strong>the</strong> underlying body; this is an autapomorphy <strong>for</strong> <strong>the</strong> family. Spiracles are<br />

Insect <strong>External</strong> Anatomy © Houseman – page 9


located on <strong>the</strong> last two segments <strong>of</strong> <strong>the</strong> thorax but appear between <strong>the</strong> pro- and<br />

mesothorax and <strong>the</strong> meso- and metathorax.<br />

Each leg has six tubular segments (Figure 9). Closest to <strong>the</strong> body is <strong>the</strong> large coxa, and<br />

this segment is followed by <strong>the</strong> trochanter, <strong>the</strong> large femur, and <strong>the</strong> tibia. The sixth<br />

segment is <strong>the</strong> tarsus, and in insects it is divided by a series <strong>of</strong> up to five articulations<br />

called tarsomeres. Like <strong>the</strong> antennal articulations observed on <strong>the</strong> flagellum, <strong>the</strong><br />

tarsomeres are not segments. The claws at <strong>the</strong> tip are <strong>the</strong> nearly invisible pretarsus.<br />

Don’t let <strong>the</strong> name <strong>of</strong> <strong>the</strong> pretarsus confuse you; it is located after <strong>the</strong> tarsus, not<br />

be<strong>for</strong>e it as <strong>the</strong> name implies. A fleshy pad is <strong>of</strong>ten associated with <strong>the</strong> claw at <strong>the</strong> tip<br />

<strong>of</strong> <strong>the</strong> organism. This pad, when located between <strong>the</strong> claws, is referred to as an<br />

arolium; when located under and at <strong>the</strong> base <strong>of</strong> <strong>the</strong> claw, it is called a pulvillus.<br />

Abdomen<br />

Identify <strong>the</strong> typical abdominal segment components including: sclerotized tergite,<br />

sternite, membranous pleura, and spiracular openings. On <strong>the</strong> terminal segments<br />

identify <strong>the</strong> appendages involved in copulation and egg laying. Most insects have a<br />

complex ovipositor <strong>for</strong>med from <strong>the</strong> ancestral appendages <strong>of</strong> <strong>the</strong> eighth and ninth<br />

segments. But this is missing in roaches, an autapomorphy <strong>for</strong> <strong>the</strong> order, because <strong>of</strong><br />

<strong>the</strong>ir use <strong>of</strong> an oo<strong>the</strong>ca. In <strong>the</strong> female <strong>the</strong> eighth and ninth segments are reduced and<br />

hidden by <strong>the</strong> seventh, which is enlarged into two genital plates on ei<strong>the</strong>r side <strong>of</strong> <strong>the</strong><br />

genital opening where <strong>the</strong> oo<strong>the</strong>ca <strong>for</strong>ms. Unlike in <strong>the</strong> male specimen you will<br />

observe next, only <strong>the</strong> cerci are present. In <strong>the</strong> male <strong>the</strong> eight and ninth segments <strong>of</strong><br />

<strong>the</strong> abdomen resemble <strong>the</strong> ancestral <strong>for</strong>m. The tergites <strong>of</strong> <strong>the</strong> tenth and eleventh<br />

segments have a central notch and extend towards <strong>the</strong> back <strong>of</strong> <strong>the</strong> roach. The cuticular<br />

plates on ei<strong>the</strong>r side <strong>of</strong> <strong>the</strong> anal opening are <strong>for</strong>med from <strong>the</strong> eleventh sternite; <strong>the</strong><br />

tenth sternite is reduced in size and not visible. Males have a pair <strong>of</strong> cerci and stylets<br />

at <strong>the</strong> tip <strong>of</strong> <strong>the</strong> abdomen.<br />

The Insect Head<br />

Insect heads may have one <strong>of</strong> three different orientations based on <strong>the</strong> position <strong>of</strong> <strong>the</strong><br />

mouthparts relative to <strong>the</strong> remainder <strong>of</strong> <strong>the</strong> head capsule. When <strong>the</strong> mouthparts are<br />

directed downward <strong>the</strong> head is considered hypognathous compared to prognathous,<br />

when <strong>the</strong> mouthparts are directed <strong>for</strong>ward. In both <strong>the</strong>se last two cases <strong>the</strong> oral<br />

appendages are located anteriorly but when positioned in <strong>the</strong> posterior region <strong>the</strong> head<br />

is opisthognathous.<br />

There are some distinct advantages to <strong>the</strong>se different orientations.Many larval insects<br />

have prognathous heads because <strong>the</strong>y live in <strong>the</strong> same medium on which <strong>the</strong>y feed.<br />

Forward directed mouthparts are an obvious advantage. Take a look at <strong>the</strong> mealworm<br />

larva as our example <strong>of</strong> a prognathous head. O<strong>the</strong>rs such as <strong>the</strong> cicada and a variety <strong>of</strong><br />

plant sap feeding insects must have <strong>the</strong> mouthparts penetrate <strong>the</strong> vascular plant tissue.<br />

To achieve this requires considerable strength and <strong>the</strong> opisthognathous head allows <strong>the</strong><br />

Insect <strong>External</strong> Anatomy © Houseman – page 10


insect to apply <strong>the</strong> pressure required to penetrate <strong>the</strong> plant. Identify <strong>the</strong> different mouth<br />

part types on <strong>the</strong> specimens when you examine <strong>the</strong>ir mouthparts.<br />

Figure 10 Three different types <strong>of</strong> insect heads From left to right,<br />

hypognathous, prognathous and opisthognathous<br />

Insect Mouthparts<br />

One <strong>of</strong> <strong>the</strong> autapomorphies defining <strong>the</strong> Kingdom Animalia is heterotrophy, and<br />

animals must consume high-energy chemical bonds in <strong>the</strong> <strong>for</strong>m <strong>of</strong> food and <strong>the</strong>n break<br />

<strong>the</strong>se bonds to fuel <strong>the</strong>ir life’s processes. It should come as no surprise that animal<br />

diversity is <strong>of</strong>ten reflected in <strong>the</strong> varied diets <strong>of</strong> animals. Specialists feeding on one<br />

type <strong>of</strong> food, phytoplankton <strong>for</strong> example, have a very different morphology than<br />

burrowers living in <strong>the</strong> sediments <strong>of</strong> <strong>the</strong> bottom <strong>of</strong> <strong>the</strong> ocean. It’s <strong>the</strong> same with<br />

insects, and one <strong>of</strong> <strong>the</strong> reasons <strong>for</strong> <strong>the</strong>ir success is <strong>the</strong>ir ability to exploit a wide variety<br />

<strong>of</strong> food sources, from plant sap or blood to fea<strong>the</strong>rs, wood, or leaves. In order to<br />

consume <strong>the</strong>se different types <strong>of</strong> food, insect mouthparts have become modified in<br />

various ways to create specialized tools <strong>for</strong> processing different types <strong>of</strong> food <strong>for</strong><br />

consumption.<br />

An insect’s mouth lies at <strong>the</strong> base <strong>of</strong> a preoral cavity <strong>for</strong>med by <strong>the</strong> feeding<br />

appendages. This arrangement <strong>of</strong> appendages is ideal <strong>for</strong> <strong>the</strong> manipulation and<br />

modification <strong>of</strong> food be<strong>for</strong>e ingestion. The preoral cavity is like an open box: <strong>the</strong><br />

paired mandibles and maxillae <strong>for</strong>m one pair <strong>of</strong> opposing sides, and <strong>the</strong> labrum and<br />

labium <strong>for</strong>m <strong>the</strong> o<strong>the</strong>r pair <strong>of</strong> sides. The front <strong>of</strong> <strong>the</strong> preoral cavity is <strong>the</strong> opening to<br />

<strong>the</strong> box; at <strong>the</strong> bottom is <strong>the</strong> mouth and <strong>the</strong> opening <strong>of</strong> <strong>the</strong> salivary glands. The preoral<br />

cavity is divided in two by <strong>the</strong> tongue-like hypopharynx; this creates a dorsal cibarium<br />

that leads to <strong>the</strong> mouth and, below <strong>the</strong> hypopharynx, a ventral salivarium with <strong>the</strong><br />

opening <strong>of</strong> <strong>the</strong> salivary glands. What appendages on an insect’s head are not<br />

involved in feeding?<br />

One <strong>of</strong> <strong>the</strong> earliest innovations in insect mouthparts was a change from a mandible<br />

with one joint (condyle) connecting it with <strong>the</strong> head capsule to a double-jointed<br />

mandible articulating with <strong>the</strong> head capsule. The symplesiomorphy <strong>of</strong> <strong>the</strong> dicondylic<br />

Insect <strong>External</strong> Anatomy © Houseman – page 11


mandible is used to separate <strong>the</strong> most primitive hexapods from <strong>the</strong> insects. Are <strong>the</strong>re<br />

any o<strong>the</strong>r differences between insects and o<strong>the</strong>r more primitive hexapods? This<br />

chewing, mandibulate condition is primitive, and <strong>the</strong> grasshopper is <strong>the</strong> classic<br />

example <strong>of</strong> this morphology. Fluid-feeding insects have haustellate mouthparts. One<br />

<strong>of</strong> <strong>the</strong> reasons insects are so successful is that <strong>the</strong>y are able to modify <strong>the</strong>ir body<br />

cuticle into an amazing array <strong>of</strong> shapes and <strong>for</strong>ms; think <strong>of</strong> a dragonfly, a house fly,<br />

and a beetle, and you can see just how dramatic <strong>the</strong>se changes can be. The same<br />

applies to <strong>the</strong>ir mouthparts. For example, a large grinding mandible can be modified<br />

into a delicate syringe-like structure that penetrates <strong>the</strong> circulatory vessels <strong>of</strong> plants<br />

and animals to suck up fluid plant sap or blood. Not only are <strong>the</strong> shapes <strong>of</strong> <strong>the</strong> original<br />

mouthparts changed, <strong>the</strong>y <strong>of</strong>ten disappear completely. In this lab, you will get a<br />

chance to see some <strong>of</strong> <strong>the</strong>se modifications.<br />

Mandibulate: Chewing and biting<br />

The ancestral mouthparts were mandibulate, and teeth on <strong>the</strong> mandibles ripped <strong>the</strong><br />

food into pieces that are ground up by <strong>the</strong> molar surface <strong>of</strong> <strong>the</strong> mandible be<strong>for</strong>e being<br />

swallowed. As was mentioned earlier, <strong>the</strong> chewing and biting mandible is <strong>the</strong> most<br />

primitive type <strong>of</strong> insect mouthpart, and <strong>the</strong> grasshopper is a good example <strong>for</strong> studying<br />

this type <strong>of</strong> mouthpart.<br />

Grasshopper - mandibulate<br />

The mandibulate mouthparts <strong>of</strong> <strong>the</strong> grasshopper resemble those <strong>of</strong> <strong>the</strong> generalized or<br />

Figure 11 Mouthparts <strong>of</strong> <strong>the</strong> grasshopper.<br />

primitive insect (Figure 11). Prepared slides are available <strong>of</strong> <strong>the</strong> hopper mouthparts<br />

and as you examine <strong>the</strong>m, locate <strong>the</strong>m on <strong>the</strong> head <strong>of</strong> <strong>the</strong> preserved specimen.<br />

The labrum is suspended from <strong>the</strong> clypeus, and toge<strong>the</strong>r <strong>the</strong> two pieces <strong>for</strong>m a platelike<br />

appendage on <strong>the</strong> ro<strong>of</strong> <strong>of</strong> <strong>the</strong> preoral cavity; <strong>the</strong> underside <strong>of</strong> <strong>the</strong> labrum is<br />

referred to as <strong>the</strong> epipharynx. Although not all entomologists agree, it is generally<br />

accepted that <strong>the</strong> labrum is <strong>for</strong>med from <strong>the</strong> fused appendages <strong>of</strong> <strong>the</strong> first head<br />

segment. The mandibles, highly sclerotized and hardened, have a cutting (incisor)<br />

and a grinding (molar) surface; find <strong>the</strong>se on your specimen. The mandible articulates<br />

Insect <strong>External</strong> Anatomy © Houseman – page 12


at <strong>the</strong> front and back <strong>of</strong> <strong>the</strong> gena and is dicondylic with cuticular extensions known as<br />

condyles that fit into grooves or slots. The presence <strong>of</strong> two joints means that <strong>the</strong><br />

mandible swings in and out when <strong>the</strong> animal chews. Try to locate <strong>the</strong> articulations and<br />

<strong>the</strong> condyle.<br />

The maxillae are also paired, and <strong>the</strong> basal segments include <strong>the</strong> cardo that articulates<br />

with <strong>the</strong> gena and <strong>the</strong> stipes, with its tooth-like lacinia, a galea, and a leg-like<br />

maxillary palp. The teeth on <strong>the</strong> lacinia help <strong>the</strong> mandible to tear up food, while<br />

sensory hairs and bristles on <strong>the</strong> galea and palp are used to taste and feel <strong>the</strong> food. The<br />

labium is <strong>the</strong> fused appendages <strong>of</strong> <strong>the</strong> sixth head segment, and it <strong>for</strong>ms <strong>the</strong> floor to <strong>the</strong><br />

preoral cavity. It attaches to <strong>the</strong> head by <strong>the</strong> gula, which articulates across <strong>the</strong> back <strong>of</strong><br />

<strong>the</strong> head. The gula has a fleshy inner ligula and outer sensory palps. The<br />

hypopharynx is not an appendage but lies in <strong>the</strong> preoral cavity as a tongue-like<br />

structure that has an oral opening above and a salivary opening below: <strong>the</strong> cibarium<br />

and salivarium, respectively.<br />

Haustellate: Fluid feeders<br />

Mouthpart morphologies in fluid-feeding insects are classified based on where <strong>the</strong><br />

ingested fluid is found and how it is ingested. Piercing mouthparts are modified into<br />

long needle-like structures, called stylets, that pierce into a plant or animal and <strong>for</strong>m a<br />

drinking tube to suck up its internal fluids. Insects can also feed on fluids by ei<strong>the</strong>r<br />

sponging <strong>the</strong>m <strong>of</strong>f a surface or by using a drinking tube to feed on hidden or hard-toreach<br />

liquid foods.<br />

Piercing mouthparts are an autapomorphy that defines <strong>the</strong> Hemiptera who feed on a<br />

range <strong>of</strong> fluids from plant sap to vertebrate blood. The mosquito, horsefly, house fly,<br />

stable fly, and tsetse fly are all in <strong>the</strong> Order Diptera, which is divided into two groups:<br />

<strong>the</strong> lower and higher flies. One <strong>of</strong> <strong>the</strong> differences between <strong>the</strong> two dipteran groups is<br />

found in <strong>the</strong> morphology <strong>of</strong> <strong>the</strong>ir mouthparts. Lower flies have modified versions <strong>of</strong><br />

all five mouthparts; higher flies have lost <strong>the</strong> mandible, and <strong>the</strong> palp is <strong>the</strong> only part <strong>of</strong><br />

<strong>the</strong> maxilla that remains. Both piercing and sponging mouthparts are found in <strong>the</strong><br />

Diptera. Adult Lepidoptera are all siphon feeders, using <strong>the</strong>ir reduced mouthparts to<br />

feed on <strong>the</strong> nectar <strong>of</strong> flowers. The fleshy mouthparts <strong>of</strong> a house fly, which extend as<br />

<strong>the</strong> fly crawls across a table are an example <strong>of</strong> <strong>the</strong> sponging type, and <strong>the</strong> long<br />

proboscis <strong>of</strong> a butterfly sipping nectar from within a flower is an example <strong>of</strong> <strong>the</strong><br />

siphoning type <strong>of</strong> haustellate mouthpart.<br />

Insect <strong>External</strong> Anatomy © Houseman – page 13


Mosquito - piercing<br />

When a female mosquito feeds, <strong>the</strong> labrum, maxillae, and mandibles (Figure 12) all<br />

penetrate <strong>the</strong> skin, and <strong>the</strong> labium folds out <strong>of</strong> <strong>the</strong> way as <strong>the</strong> mouthparts cut deeper in<br />

search <strong>of</strong> a capillary. Most <strong>of</strong> <strong>the</strong> cutting work is done by <strong>the</strong> maxilla and mandible.<br />

Once a capillary is located, <strong>the</strong> maxilla anchors <strong>the</strong> mouthparts, <strong>the</strong> mandibles cut<br />

through <strong>the</strong> capillary wall, and <strong>the</strong> tip <strong>of</strong> labrum is inserted into <strong>the</strong> blood vessel.<br />

Figure 12 Mouthparts <strong>of</strong> <strong>the</strong> mosquito.<br />

Saliva from <strong>the</strong> hypopharynx is added to <strong>the</strong> blood just be<strong>for</strong>e it is sucked up <strong>the</strong> food<br />

canal into <strong>the</strong> cibarium and <strong>the</strong>n into <strong>the</strong> mouth. Only female mosquitoes are blood<br />

feeders, and <strong>the</strong>ir saliva contains an anticoagulant that is mixed with <strong>the</strong> blood so it<br />

doesn’t clot as it passes through <strong>the</strong> food canal. Like male mosquitoes, females also<br />

feed on nectar, and <strong>the</strong> sugar is used to fuel flight in both. The protein <strong>of</strong> <strong>the</strong> blood<br />

meal consumed by <strong>the</strong> female is used to produce eggs. Technically it’s not a piercing<br />

mouthpart when <strong>the</strong> mosquito feeds on nectar: Why and what type <strong>of</strong> a<br />

mouthpart is it?<br />

Although whole mount slides <strong>of</strong> mosquitoes can be used <strong>for</strong> observation <strong>of</strong> <strong>the</strong><br />

mouthparts, it will be difficult to see <strong>the</strong> individual stylets that are usually inside <strong>the</strong><br />

labial sheath.<br />

Insect <strong>External</strong> Anatomy © Houseman – page 14


Horse fly – Cutting and sponging<br />

Horse flies, Tabanus spp., and deer flies, Crysops spp., are also lower Diptera. All <strong>the</strong><br />

ancestral mouthparts are present, but highly modified and divided in two main<br />

functional components (Figure 13). The cutting blades are <strong>for</strong>med from <strong>the</strong> mandibles<br />

Figure 13 Cutting and sponging mouthparts <strong>of</strong> <strong>the</strong> horsefly or deer fly<br />

and <strong>the</strong> lacinea <strong>of</strong> <strong>the</strong> maxilla. They open and close much like a pair <strong>of</strong> scissors as <strong>the</strong>y<br />

cut through <strong>the</strong> skin. It is <strong>the</strong> combined action <strong>of</strong> <strong>the</strong> paired mandibles and maxillae<br />

that makes <strong>the</strong> bite <strong>of</strong> <strong>the</strong>se two flies so painful. The hypopharynx is elongated and has<br />

a central salivary canal that releases saliva into <strong>the</strong> blood that pools in a wound. The<br />

labellum is made up <strong>of</strong> <strong>the</strong> fleshy lobes <strong>of</strong> <strong>the</strong> labium, and each <strong>of</strong> <strong>the</strong>se lobes is a<br />

modified labial palps. Take a close look at <strong>the</strong> surface <strong>of</strong> <strong>the</strong> labellum and you will see<br />

it is covered in small channels referred to as pseudotrachea. When <strong>the</strong> labellum<br />

contacts <strong>the</strong> blood in <strong>the</strong> wound, <strong>the</strong> pseudotrachea fill by capillary action. Partial<br />

rings <strong>of</strong> cuticle rein<strong>for</strong>ce <strong>the</strong> pseudotrachea; this prevents <strong>the</strong> channels from collapsing<br />

when blood is sucked into <strong>the</strong> food canal that is <strong>for</strong>med from <strong>the</strong> inner surface <strong>of</strong> <strong>the</strong><br />

labrum. Are maxillary palps present?<br />

Insect <strong>External</strong> Anatomy © Houseman – page 15


House fly - sponging<br />

Figure 14 Mouth parts <strong>of</strong> a house fly.<br />

Although <strong>the</strong> sponging labium <strong>of</strong> <strong>the</strong> horse fly and house fly are similar, <strong>the</strong> house fly<br />

is a higher fly and has no mandibles, and <strong>the</strong> maxilla are reduced to maxillary palps<br />

and an internal cuticular rod that supports <strong>the</strong> attachment <strong>of</strong> <strong>the</strong> modified labium to <strong>the</strong><br />

head. Like that <strong>of</strong> <strong>the</strong> horse fly, <strong>the</strong> labium <strong>of</strong> <strong>the</strong> house fly, Musca domestica, is a<br />

large fleshy structure (Figure 14). When sensory hairs on <strong>the</strong> fly’s feet detect food, <strong>the</strong><br />

labium is lowered <strong>for</strong> feeding. The large labellum is <strong>for</strong>med from modified labial<br />

palps, and pseudotrachea use capillary action to absorb liquid foods. The mechanism<br />

is similar to how <strong>the</strong> labellum is used to sponge up blood in <strong>the</strong> horse fly. It’s a little<br />

different with solid foods. In this case, <strong>the</strong> fly salivates on <strong>the</strong> food, uses small teeth on<br />

<strong>the</strong> surface <strong>of</strong> <strong>the</strong> labellum to rasp and scrape at <strong>the</strong> food, combines <strong>the</strong> food with<br />

salivary secretions, and <strong>the</strong>n sucks <strong>the</strong> nutrients up and into <strong>the</strong> mouth. How do <strong>the</strong><br />

salivary secretions reach <strong>the</strong> tip <strong>of</strong> <strong>the</strong> labellum?<br />

Insect <strong>External</strong> Anatomy © Houseman – page 16


Stable fly and tsetse fly - piercing<br />

The stable fly, Stomoxys sp., and <strong>the</strong> tsetse fly, Glossina sp., are higher flies. Similarly<br />

to o<strong>the</strong>r members <strong>of</strong> <strong>the</strong> group, <strong>the</strong> labium is <strong>the</strong> most prominent mouthpart; <strong>the</strong><br />

mandible is missing and <strong>the</strong> palps are all that remain <strong>of</strong> <strong>the</strong> maxillae (Figure 16). The<br />

Figure 16 With <strong>the</strong> exception <strong>of</strong> <strong>the</strong> labellar teeth <strong>the</strong> piercing mouthparts <strong>of</strong><br />

<strong>the</strong> stable fly and <strong>the</strong> tsetse fly are similar.<br />

tsetse and <strong>the</strong> stable fly are both blood feeders; to be able to pierce through skin, <strong>the</strong><br />

fleshy, large labium typical <strong>of</strong> <strong>the</strong> higher flies has been modified into a strong, sturdy<br />

shaft or rostrum. The hollow cavity <strong>of</strong> <strong>the</strong> rostrum <strong>for</strong>ms <strong>the</strong> food canal, and prestomal<br />

teeth at <strong>the</strong> tip <strong>of</strong> <strong>the</strong> rostrum are used to pierce through <strong>the</strong> skin. The teeth, which are<br />

part <strong>of</strong> <strong>the</strong> modified labellum, move in and out <strong>of</strong> <strong>the</strong> opening to <strong>the</strong> buccal cavity as<br />

<strong>the</strong>y rasp and dig away at <strong>the</strong> skin until it is pierced; in <strong>the</strong> process <strong>the</strong> mouthparts are<br />

pushed deeper into <strong>the</strong> wound (Figure 15). The hypopharynx extends to <strong>the</strong> tip <strong>of</strong> <strong>the</strong><br />

Figure 15 <strong>Lab</strong>ellar teeth in <strong>the</strong> Stable fly (A) and <strong>the</strong> tsetse fly (B).<br />

rostrum, and <strong>the</strong> hollow core <strong>for</strong>ms <strong>the</strong> salivary canal, which delivers an<br />

anticoagulant-rich salivary secretion to <strong>the</strong> blood be<strong>for</strong>e it enters <strong>the</strong> food canal.<br />

Blood-feeding insects are <strong>of</strong>ten carriers, or vectors, <strong>of</strong> disease: Are ei<strong>the</strong>r <strong>of</strong> <strong>the</strong>se<br />

insects vectors, and if <strong>the</strong>y are, what disease(s) do <strong>the</strong>y carry?<br />

Insect <strong>External</strong> Anatomy © Houseman – page 17


"Bugs": Hemiptera<br />

Hemipterans are specialists at piercing, and an autapomorphy <strong>of</strong> <strong>the</strong> order is <strong>the</strong>ir<br />

unique structural “beak” with its interlocking maxillary stylets inside (Figure 17). Like<br />

that <strong>of</strong> <strong>the</strong> mosquito, <strong>the</strong> labium <strong>for</strong>ms a sheath surrounding <strong>the</strong> mouthparts that are<br />

modified into stylets. These are used to feed on a variety <strong>of</strong> foods ranging from seeds<br />

to plant fluids and blood. As a hemipteran feeds, <strong>the</strong> labium folds back out <strong>of</strong> <strong>the</strong> way,<br />

Figure 17 Mouthparts <strong>of</strong> a hemipteran define <strong>the</strong> insect order because <strong>of</strong> <strong>the</strong><br />

zipper mechanism that locks <strong>the</strong> maxillary stylets toge<strong>the</strong>r.<br />

and salivary secretions are important in preparing <strong>the</strong> food <strong>for</strong> ingestion. In seed<br />

feeders, salivary secretions liquefy <strong>the</strong> seed; in aphids, pectinases dissolve <strong>the</strong> glue<br />

between plant cells so <strong>the</strong> mouthparts can reach <strong>the</strong> vascular tissues in <strong>the</strong> plant. In<br />

blood feeders, salivary secretions include anticoagulents. Unlike <strong>the</strong> o<strong>the</strong>r insects you<br />

have examined, <strong>the</strong> Hemiptera don’t have a hypopharynx to deliver <strong>the</strong> salivary<br />

secretions to <strong>the</strong> tip <strong>of</strong> <strong>the</strong> mouthparts. Instead, <strong>the</strong> salivary canal is part <strong>of</strong> <strong>the</strong>ir<br />

unique interlocking maxillary. The paired maxillary stylets, linked to each o<strong>the</strong>r in a<br />

ziplock-like seal, create <strong>the</strong> food canal and <strong>the</strong> salivary canal. Mandibles are <strong>the</strong> main<br />

cutting structure and are found on ei<strong>the</strong>r side <strong>the</strong> maxillary stylets. <strong>Lab</strong>ial palps have<br />

disappeared along with <strong>the</strong> maxillary palps.<br />

Hemipteran mouthparts can be examined from prepared slides, <strong>for</strong> example, <strong>of</strong> <strong>the</strong><br />

squash bug Anasa tristis or whole mounts <strong>of</strong> <strong>the</strong> bloodsucking bug Rhodnius prolixus.<br />

The mouthparts on <strong>the</strong> slides may still be surrounded by <strong>the</strong> labium.<br />

Insect <strong>External</strong> Anatomy © Houseman – page 18


Honey bee - Combination<br />

The haustellate mouthparts <strong>of</strong> <strong>the</strong> honey bee are used to drink <strong>the</strong> nectar <strong>of</strong> flowers, a<br />

reward <strong>for</strong> <strong>the</strong> bee’s services as a pollinator. Nectar is a near-perfect food: an almost<br />

pure solution <strong>of</strong> sucrose and its monosaccharide components <strong>of</strong> glucose and fructose.<br />

Plants produce nectar <strong>for</strong> no o<strong>the</strong>r reason than to attract pollinators, and <strong>the</strong> honey<br />

bee’s mouthparts have been modified to ingest this sweet solution. However, honey<br />

bee mouthparts have a second function that isn’t related to feeding. Their mouthparts<br />

work <strong>the</strong> waxes that build <strong>the</strong> hive’s honeycomb, which stores pollen, honey, and <strong>the</strong><br />

incubation cells <strong>of</strong> larval bees. Strictly speaking, honey bees have haustellate<br />

mouthparts, but this second function <strong>of</strong> manipulating wax classifies <strong>the</strong>m as having<br />

combination mouthparts. Horse flies could also be placed in <strong>the</strong> same category:<br />

why?<br />

The food canal in a honey bee is different from <strong>the</strong> simple food tube or “straw” in<br />

o<strong>the</strong>r insects (Figure 18). The difference is related to one <strong>of</strong> <strong>the</strong> properties <strong>of</strong> <strong>the</strong> insect<br />

cuticle. Insects are able to survive in a desiccating terrestrial environment because <strong>of</strong><br />

<strong>the</strong>ir waxy, waterpro<strong>of</strong> epicuticle. Although it hasn’t been mentioned previously, this<br />

is a problem <strong>for</strong> insects with haustellate mouthparts made <strong>of</strong> cuticle: <strong>the</strong> waxy, waterrepelling<br />

epicuticle also lines <strong>the</strong> food canal! In most cases, <strong>the</strong> pumping action <strong>of</strong> <strong>the</strong><br />

pharynx is sufficient to overcome <strong>the</strong> problem. Honey bees, however, have ano<strong>the</strong>r<br />

way <strong>of</strong> solving <strong>the</strong> problem.<br />

Figure 18 The mouthparts <strong>of</strong> a honey bee lap up nectar and <strong>the</strong>n swing out <strong>of</strong><br />

<strong>the</strong> way <strong>for</strong> mandible to manipulate wax.<br />

The donut-shaped food canal in a honey bee is made <strong>of</strong> a central, cuticular tongue,<br />

surrounded by a sheath <strong>for</strong>med by <strong>the</strong> o<strong>the</strong>r mouthparts; it is not like <strong>the</strong> simple tube<br />

<strong>of</strong> <strong>the</strong> o<strong>the</strong>r insects you have examined in <strong>the</strong> lab (Figure 9). The central tongue is<br />

<strong>for</strong>med from <strong>the</strong> fleshy glossa <strong>of</strong> <strong>the</strong> labium and is covered in setal hairs that trap<br />

liquid between <strong>the</strong> hairs. The tongue functions as cuticular sponge, whereby fluid<br />

beads up between <strong>the</strong> water repellent hairs like a droplet <strong>of</strong> water beads on a freshly<br />

waxed surface. When <strong>the</strong> honey bee feeds, its tongue is extended into <strong>the</strong> sugar<br />

solution and it becomes coated in nectar. When <strong>the</strong> tongue-like glossa is retracted, it<br />

Insect <strong>External</strong> Anatomy © Houseman – page 19


ecomes surrounded by a sheath that is <strong>for</strong>med from <strong>the</strong> galea <strong>of</strong> <strong>the</strong> maxilla and<br />

modified labial palps. On <strong>the</strong> ventral surface <strong>of</strong> <strong>the</strong> glossa a groove extends up and<br />

into <strong>the</strong> cibarium and <strong>the</strong> mouth, and ei<strong>the</strong>r capillary action or suction pulls <strong>the</strong> nectar<br />

up <strong>the</strong> donut-shaped food canal and into <strong>the</strong> mouth.<br />

As already mentioned, insects are covered in a waxy cuticle, and <strong>the</strong> abdominal<br />

sclerites <strong>of</strong> <strong>the</strong> worker bees produce extra wax, beeswax. The worker bee scrapes <strong>the</strong><br />

excess wax from <strong>the</strong> abdomen and chews it with <strong>the</strong> mandibles, molding it into <strong>the</strong><br />

hexagonal honeycomb <strong>of</strong> <strong>the</strong> hive and <strong>the</strong> caps that cover <strong>the</strong> cells. When mandibles<br />

are used, <strong>the</strong> o<strong>the</strong>r mouthparts are folded back and out <strong>of</strong> <strong>the</strong> way. What type <strong>of</strong><br />

mouthparts do larval honey bees have?<br />

Butterflies and moths – Drinking tube<br />

Adult Lepidoptera, butterflies and moths, feed on <strong>the</strong> nectar <strong>of</strong> plants, and most <strong>of</strong> <strong>the</strong><br />

ancestral mouthparts are ei<strong>the</strong>r lost or highly modified. The coiled proboscis is <strong>for</strong>med<br />

from <strong>the</strong> galea <strong>of</strong> <strong>the</strong> maxilla; <strong>the</strong> maxillary palps have been lost (Figure 19). The<br />

galea is hollow and <strong>the</strong> space inside is an extension <strong>of</strong> <strong>the</strong> hemocoel. The proboscis is<br />

extended by hydrostatic pressure and recoiled by a combination <strong>of</strong> internal<br />

musculature and cuticular elasticity. The food canal is <strong>for</strong>med between galea, which is<br />

held toge<strong>the</strong>r by hooks <strong>for</strong>med from cuticular hairs. Mandibles are missing, and <strong>the</strong><br />

hypopharynx is part <strong>of</strong> <strong>the</strong> cibarial pump. <strong>Lab</strong>ial palps are all that remains <strong>of</strong> <strong>the</strong><br />

labium.<br />

Figure 19 The mouthparts <strong>of</strong> Lepidoptera are perfect drinking straws <strong>for</strong><br />

nectar.<br />

Insect <strong>External</strong> Anatomy © Houseman – page 20


DragonFly nymph<br />

Dragon fly nymphs are deadly predators in freshwater lakes and streams. The whole<br />

labium has been modified into a viscous prehensile structure with opposable labial<br />

palps at <strong>the</strong> tip that grasp onto <strong>the</strong> prey. The overlapping visual field <strong>of</strong> <strong>the</strong> large<br />

compound eyes is located precisely at <strong>the</strong> point <strong>of</strong> full extension <strong>of</strong> <strong>the</strong> labium. When<br />

a potential meal wanders into this field <strong>of</strong> view, <strong>the</strong> mouthparts shoot <strong>for</strong>ward and<br />

immediately retract carrying <strong>the</strong> trapped prey to <strong>the</strong> mouth. We only have a few <strong>of</strong><br />

<strong>the</strong>se slides so <strong>the</strong>re will be one placed on demonstration in <strong>the</strong> lab; be sure to take a<br />

look.<br />

Figure 20 Mouthparts <strong>of</strong> a dragon fly nymph<br />

Insect <strong>External</strong> Anatomy © Houseman – page 21

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