Lecture 16 Annelids & Nematoda

PhylumAnnelida Keyconcept
Annelidsaresegmentedworms. Thislectureisbasedonpages727729inChapter33andappropriateportionsoftheformand functionchaptersgivenbelow. Annelids are bilaterally symmetrical animals, distributed worldwide. All annelids possess a coelom, and have segmented bodies that show a basic metamerism with comparatively few differencesbetweenthesegment(orsomites).Theyrangeinlengthfromlessthan1mmto3m forthegiantAustralianearthworm.Annelidsliveinthesea,mostfreshwaterhabitats,anddamp soil.Organsystemsarepresentand cephalisation hasresulted inorganssuchastentacles,eye spotsandpalpsbeingwelldeveloped.Thenervoussystemismodified,withcerebralganglia,two closely fused ventral nerve cords with giant fibres running the length of the body, and various gangliawithlateralbranches(Fig33.24,pg728)

Circulation,excretionandmovement
The circulatory system is closed, with muscular blood vessels and aortic arches (hearts), which propelthebloodaroundthesystem.Welldevelopednephridiainmostsegmentshavereacheda stageofdevelopmentwhereremovalofwastefromthebloodaswellasthecoelomoccurs.The gutalsoshowsregionalspecialisationanddigestiveglandsaredeveloped.Incombinationwiththe circulatory system this means that food materials can be supplied to the body much more efficientlythaninflatworms,whosesize(particularlythickness)islimitedbecausealltransferof materialshastotakeplacebydiffusion. The excretory tubules of molluscs and annelids are termed metanephridia, being open to the coelomattheirinnerends.ContrastthiswiththeprotonephridiaofPlatyhelmintheswithclosed innerends.Thedevelopmentofacoelomandofsegmentationalsohasimportanteffectsonthe sizethatan animalcanattainand themovements itiscapable of.Coelomicfluidfunctionsasa hydrostatic skeleton and the important division of the coelom by segmental septa facilitates coordinatedmovementbyperistalsis(Fig50.35,pg1157).

ClassesofAnnelids
The phylum Annelida is divided into three classes: Polychaeta (polychaetes), Oligochaeta (earthworms),andHirudinea(leeches). Polychaeta(manysetae)aremostlymarine,andhaveadistinctheadwitheyesandtentacles. Sexes are separate, and there is a planktonic larva. A variety of lifestyles from predators to deposit and filterfeeders. Each segment of a polychaete has a pair of paddlelike or ridgelike parapodia (almost feet) that function in locomotion. Each parapodium has several chitinous setae.Inmanypolychaetes,therichbloodvesselsintheparapodiafunctionasgills.Manycrawl onorburrowintheseafloor,whileafewdriftandswimintheplankton.Someliveintubesthat thewormsmakebymixingmucuswithsandandbrokenshells.Othersconstructtubesfromtheir ownsecretions. Oligochaeta: Oligochaetes are named for their relatively sparse chaetae, or bristles made of chitin.Thisclassofsegmentedwormsincludestheearthwormsandavarietyofaquaticspecies. Earthworms eat their way through soil, extracting nutrients as the soil passes through the

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alimentarycanal.Undigestedmaterialisegestedascastings.Earthwormstillthesoil,enrichingit withtheircastings.Earthwormsarecrossfertilizinghermaphrodites.Twoearthwormsexchange sperm and then separate. The received sperm are stored while a special organ, the clitellum, secretesamucouscocoon.Asthecocoonslidesalongthebody,itpicksupeggsandstoredsperm and slides off the body into the soil. There is no larva. Some earthworms can also reproduce asexuallybyfragmentationfollowedbyregeneration. Hirudinea: The majority of leeches inhabit fresh water, but land leeches move through moist vegetation,andsomearemarine.Leechesrangeinsizefromabout1to30cm.Thebodyhasa fixednumberofsegments(34:15:30)andhasoralandcaudalsuckers.Setaeareabsent.Theyare hermaphrodite, and have no larva. Movement is by undulatory swimming or "inchworm" looping.Theyaremostlypredatorsofotherinvertebrates,butsomearebloodsuckingparasites thatfeedbyattachingtemporarilytootheranimals,includinghumans.Someparasiticspeciesuse bladelikejawstoslitthehostsskin,whileotherssecreteenzymesthatdigestaholethroughthe skin.Thehostisusuallyunawareoftheattackbecausetheleechsecretesananesthetic.Theleech alsosecreteshirudin,ananticoagulant,intothewound,allowingtheleechtosuckasmuchblood asitcanhold.Untilthe20thcentury,leecheswerefrequentlyusedbyphysiciansforbloodletting. Leechesarestillusedtodrainbloodthataccumulatesintissuesfollowinginjuryorsurgery. Researchers arealsoinvestigatingthe potentialuseofhirudintodissolveunwanted bloodclots fromsurgeryorheartdisease.Arecombinantformofhirudinhasbeendevelopedandisinclinical trials.

Conceptcheck33.3
[Questions3and4].

Vocabulary
At the end of this lecture, you should know the meanings of the following words and, more importantly,youshouldbeabletousetheminwrittenwork: Coelom, metamerism, cerebral ganglia, aortic arches, nephridia, metanephridia, hydrostatic, skeleton, septa, polychaeta, oligochaeta, Hirudinea, parapodia, setae, peristalsis, chaetae, clitellum.

PhylumNematoda Keyconcept
Nematodes(roundworms)arenonsegmentedpseudocoelomatescoveredbytoughcuticles. Roundwormsarefoundinmostaquatichabitats,wetsoil,moisttissuesofplants,andthebody fluidsandtissuesofanimals.Theyrangeinsizefromlessthan1mmtomorethanameter.The cylindricalbodiesofroundwormsarecoveredwithatoughexoskeleton,thecuticle.Astheworm grows,itperiodicallyshedsitsoldcuticleandsecretesanew,largerone. They have an alimentary tract and use the fluid in their pseudocoelom to transport nutrients since they lack a circulatory system. They have a characteristic thrashing motion due to contractionoflongitudinalmuscles,lackingcircularmuscles. Nematodesusuallyreproducesexually.Thesexesareseparateinmostspecies,andfertilizationis internal. Females may lay 100,000 or more fertilized eggs per day. The zygotes of most nematodesareresistantcellsthatcansurviveharshconditions.

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 Abundant, freeliving nematodes live in moist soil and in decomposing organic matter on the bottom of lakes and oceans. There are 25,000 described species, and perhaps ten times that number actually exists. If nothing but nematodes remained, it has been said, they would still preservetheoutlineoftheplanetandmanyofitsfeatures. They play a major role in decomposition and nutrient recycling. The soil nematode, Caenorhabditiselegans,hasbecomeamodelorganismindevelopmentalbiology. Thenematodesincludemanyspeciesthatareimportantagriculturalpeststhatattackplantroots. Other species parasitize animals. More than 50 nematode species, including various pinworms and hookworms, parasitize humans. Trichinella spiralis causes trichinosis when the nematode wormsencystinavarietyofhumanorgans,includingskeletalmuscle.Theyareacquiredbyeating undercooked meat that has juvenile worms encysted in the muscle tissue. Parasitic nematodes areabletohijacksomeofthecellularfunctionsoftheirhosts.Plantparasiticnematodesproduce molecules that induce the development of root cells that provide nutrients to the parasites. Trichenella in human muscle cells controls the expression of muscle cell genes that code for proteinsthatmakethecellelasticenoughtohousethenematode.Themusclecellalsoreleases signalstoattractbloodvessels,supplyingthenematodewithnutrients.

Conceptcheck33.4
[Question1].

Vocabulary
At the end of this lecture, you should know the meanings of the following words and, more importantly,youshouldbeabletousetheminwrittenwork: Cuticle,pseudocoelom.

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Lecture 17 Arthropoda
PhylumArthropoda Keyconcept
Arthropodsaresegmentedcoelomatesthathaveanexoskeletonandjointedappendages. Thislectureisbasedonpages730732inChapter33.Coverageofthisgroupisinadequateinthe textbook,soadditionalinformationissupplied. Thearthropods(PhylumArthropoda="jointedfoot")areanexceedinglysuccessfulgroup.More than63%ofalldescribedspeciesoforganismsarearthropodsandtheyconstituteabout80%of theknownspeciesofanimals(Figure1).Ofthese,about75%areinsects.Inadditiontoinsects, the phylum Arthropoda includes spiders, mites, ticks, scorpions, centipedes, millipedes, crabs, crayfish, shrimps and barnacles, plus many other extant (living) forms. It also contains many extinct animals such as the trilobites. Arthropods can be found in almost every environment: marine,freshwaterandair.
Figure1Biodiversitywithindifferenttaxa(CopiedfromWheeler,1990)

Fromwhattypeofanimaldidthearthropodevolve?
They arose either from primitive annelids (the subject of the last lecture) or from ancestors common to both. The relationship between the arthropods and the annelids can be seen from severalsharedfeatures.Mostnotably,bothgroupsaremetameric.Thatis,theybothshowserial repetitionoflikesegments.Primitively,eacharthropodsegmentbearsapairofappendages,just as each segment of a polychaete (Phylum Annelida) bears a pair of parapodia. In most modern members of the Arthropoda the underlying metameric (serial) repetition of like segments is maskedbythefusionandmodificationofdifferentregionsofthebodyforspecialisedfunctions. Thisspecialisationofgroupsofsegmentsisknownastagmatisation.Morewillbesaidaboutthis

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laterwithreferencetospecificexamples.Atthisstageitissufficienttopointoutthattwomajor arthropod classes (Insecta and Crustacea) have three distinct tagmata (segments): head, thorax and abdomen. In some Crustacea the head and thorax are fused to form a cephalothorax (= headthorax).

Whyhavethearthropodsbeensosuccessful?
As just mentioned, the arthropods evolved from an annelidlike ancestor. Annelids are soft bodied,andtheouterbodywallisgenerallyflexible.Annelidsmovebythrowingtheirbodiesinto twistsandbends,withtheirmusclesworkingagainstaninternal"hydrostaticskeleton".Thebody wallisthin,andcanserveasasurfaceforgasexchangesolongasitiskeptmoist.Tohelpachieve this,andtoprotectagainstabrasion,theyoftensecretealayerofmucusovertheoutsideoftheir body.Occasionallytheannelidsalsosecreteaprotectivelayerofcuticle,buteventhenthecuticle remains permeable to both water and gases. For this reason annelids are restricted to moist environments. Theprimaryfactorinthesuccessofthearthropods,then,wastheabilitytosecreteanouterlayer of cuticle that was no longer permeable to water and gases. This tough outer layer is called an exoskeleton(="externalskeleton").Theexoskeletonwasmadewaterproofandgasproofbythe secretionofanouterwaxylayerofepicuticle(Figure2).Thecuticleunderneaththiswaxylayeris called the procuticle. The procuticle was also hardened by "tanning" via a process called sclerotisation. This involves the formation of crosslinkages between the protein chains in the cuticle.Thecombinationoftheouterwaxyepicuticlemeantthattheancestralarthropodswere nowencasedintoughwaterproofcasesthatprotectedthemfromtheenvironmentandenabled someofthemtobecometrulyterrestrial.
Figure2(CopiedfromPechenik,1991)

Butiftheywereencasedinatough,rigid,impermeableexoskeleton,howdidtheymoveandhow didtheygrow?

Arthropodmovement
Thesecrettoarthropodmovementishintedatbytheirname(rememberArthropoda="jointed foot").Thecuticleisnotuniformlythickorhardovertheanimal'sbody.Inmanyplacesthecuticle is thin and flexible, forming joints (Figure 3). Within these joints pairs of muscles (flexors and extensors)operateantagonistically(inopposition)toeachother,muchastheydoinvertebrate joints.Butinarthropods,ofcourse,theskeletontowhichthemusclesattachisontheoutsideof

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the animal. The jointed flexible exoskeleton has enabled arthropods to adopt many different lifestyles,includingflight.

Figure50.34Comparisonofantagonisticmusclesina humanarmandinaninsectleg.

Figure3Arthropodmovement (CopiedfromPechenik,1991)

Arthropodgrowth
Unlikemolluscs,arthropodscannotgraduallyaddtotheprotectivecoveringatagrowingedgeas the soft internal tissue grows. Once the cuticle hardens the arthropod is literally encased in its armour,exceptwherethearmourispiercedbysensoryhairsandglandopenings. In addition, the front and hind parts of the arthropods digestive system are also lined with cuticle.Togrow,thearthropodhastoshedthecuticle,includingthatliningthegut.Itmustthen grow larger, and then harden a new layer of cuticle to cover the bigger body. Before the old cuticleisshed,itispartiallydigestedbyenzymessecretedbytheepidermis.Theoldcuticleisthen split by the uptake of water and air, and increased blood pressure, which cause the animal to swell. Duringthisprocessanewlayerofcuticleissecreted,butthisdoesnothardenuntilthearthropod hassheditsoldcuticleandswelledtoitsnewsize.Theprocessofsheddingtheoldexoskeletonis called ecdysis, from the Greek work for "an escape". A consequence of this process is that arthropodsgrowinaseriesofdiscretesteps(Figure4).Ininsects,theperiodbetweenmoultsare calledinstars.

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Figure 4 The moult cycle of a crab (CopiedfromBarnes,1987)

Above we have considered some of the basic features of the arthropods including their origin, how they move, and how they grow.Wewillnowconsiderdetails aboutafewselectedarthropods.

Thetrilobites(subphylumTrilobita)
The trilobites are the most primitive of the known arthropod groups, and are now all extinct. About 4,000 species have been described from the fossil record. They were especially common 500 million years ago to 250 million years ago. They were marine arthropods that showed a considerable degree of ecological diversity. Some forms burrowed, some walked, and others swam.Mosttrilobitespeciesrangedfrom3to10cminlength,althoughsomeplanktonicforms wereonly0.5mmlong,andotherformsgrewtonearly1mlong.Thetrilobitebodywasflattened dorsoventrally and divided into three sections (tagmata). Sections I and III were covered by a continuousunjointedsheetofexoskeleton(acarapace)onthedorsalsurface(Figure5).Fromthe ventralsurfacethemetamericnatureoftheseanimalsisclear.Posteriortothemoutheachbody segmentboreapairoftwobranched(biramous)appendages.Thesegmentsoftheouterbranch borelongfilaments,whichmayhavebeen,gillfilamentsorusedforswimming.Theinnerbranch ofeachlimbwasdevoidofthesefilaments,andtheywereprobablyusedforwalking.Thisserial repetition of segments with identical appendages down the body was the primitive arthropod condition. From this condition various degrees of specialisation have occurred. Often this specialisation has involved the reduction or loss of one or both branches of the primitive biramouslimbs.

Figure5 (Copiedfrom Pechenik,1991)

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Kingcrabs(SubphylumChelicerata,ClassMerostomata)(pgs732733)
The king crab, sometimes called horseshoe crab, is not a crab but is more closely related to spiders. The fossil record of these animals extends back to the Cambrian period, but only five speciesarecurrentlyliving.Kingcrabsburrowthroughthesurfacelayersofmuddysubstrateand eatsmalleranimalsthattheycomeacross. Seen from above it appears to have three segments (tagmata): a large rounded brown or dark olivegreendome(prosoma=headandthorax)hingedtoahardandroughlytriangularabdomen (opisthosoma),andendinginalongmovabletailspike(telson).Ontheventralsurfaceareaseries ofpairsofjointedlimbs.Behindthefirstshortpair(chelicerae)arefourpairsoflongerlimbsthat areallalike.Eachoftheseendsinlittlepincers(chelae)exceptforthesecondpair(orsecondand thirdpairs,accordingtospecies)intheadultmale.Nextintheseriesisonemorepairoflegswith nopincers.Insteadtherearespinesthatarisefromthesecondtolastjointandhelptheanimalin gettingagriponthesand.Finallyontheprosoma(headandthorax)isapairofsmallstructuresof uncertainfunctionknownaschilaria. Theseriesoflimbscontinuesontheabdomen(opisthosoma),firstwithasortofcoverorgenital operculumwithpairedgenitalopeningsonitsunderside,andthenfivepairsofgillflaps.Eachof theseismadeupofupto200thinleafletsorgills. Kingcrabsarescavengersandfeedonmolluscs,worms,andotherorganismsincludingbottom dwellingalgae.TheyaresometimesaseriouspredatoronclambedsontheAmericancoast.The mouth is located just behind the chelicerae. Food is picked up by the chelate (pincerlike) appendages and passed to the base of the legs which have spiny protuberances (gnathobases) usedtochewthefood.

Figure6Atick,Dermacentorvaria (CopiedfromBarnes,1987)

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Mitesandticks(SubphylumChelicerata,ClassArachnida,OrderAcari)(pgs732733)
Themitesandticks(OrderAcarina)arethemostimportantofthearachnids(spiderlikeanimals) from the standpoint of human economics. Many species are parasitic on humans, our domesticatedanimalsandourcrops.Othersdestroyfoodandotherproducts.Mitesareoneof themostubiquitous(widespread)groupsintheAnimalKingdom.Mitescanbefoundinalmostall habitats including Polar Regions, deserts and hot springs. They are also found in the nostrils of seals,amongthe gillsofcrayfish,inthehearingorgansofmoths,andcommonlyinhumanhair follicles. Terrestrialspeciesareextremelyabundant,particularlyinmoss,fallenleaves,humus,soil,rotten wood and detritus. Their success seems to be related to their small size and ability to exploit microhabitats. The number of individuals is enormous. Despite their abundance the taxonomy andbiologyofmitesarestillnotwellknown. Themajorityofadultmitesare1mmorlessinlengthalthoughticks,thelargestmembersofthe order,canreach3cminlength.Astrikingfeatureofmitesandticksistheapparentlackofbody divisions. Abdominal segmentation has disappeared and the abdomen has fused with the prosoma. Except in a few primitive forms segmentation, when present, is secondary. So the positions of the appendages, the eyes and the genital orifice are the only features that differentiate the original body regions. Coinciding with this fusion, the entire body has become coveredwithasinglescleratisedshield(carapace)inmanyforms.

Conceptcheck33.4
Youshouldbeabletoanswerbothquestion2,plusunderstandthecontentoftable33.5.

Vocabulary
At the end of this lecture, you should know the meanings of the following words and, more importantly,youshouldbeabletousetheminwrittenwork: Metameric, tagmatisation, cephalothorax, exoskeleton, epicuticle, procuticle, sclerotisation, jointedlimbs,ecdysis,instar,carapace,biramouslimbs,tagmata,prosoma,opisthosoma,telson, chelicerae,chelate,gnathobases,

References
Barnes,R.D.1987."InvertebrateZoology"5thEdition. SaundersCollege.Philadelphia. Pechenik,J.A.1991."BiologyofInvertebrates".2ndEdition. Wm.C.BrownPublishers.Dubuque. Wheeler, Q.D. 1990. Insect diversity and cladistic constraints. Ann. Entomol. Soc. Am. 83:1031 1047.

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Lecture 18 Crustacea
Keyconcept
Crustaceaarearthropodsthathavetwopairsofantennaeandbiramouslimbs. Thislectureisbasedonpages737738inChapter33.Coverageofthisgroupisinadequateinthe textbook,soadditionalinformationissupplied. ThesubphylumCrustaceacontainsmorethan42,000knownspeciesincludingshrimps,crayfish, crabsandbarnacles.Unlikethemites,ticks,spidersandinsects,Crustaceaareprimarilyaquatic. Mostaremarine,butsomeliveinfreshwaterandafewareterrestrial. WhatarethecharacteristicfeaturesofCrustacea?
Figure 1. Archetypal plan of a malacostracan (left) and parts of a biramous crustacean appendage (third maxillipedofacrayfish)(right).CopiedfromHickmanetal.(2009),Figs13.17and13.18.

Afeaturethatdistinguishesthecrustaceansfromallotherarthropodsispossessionoftwopairs ofantennae.Thefirstpairisbelievedtobehomologoustotheantennaeofinsects.Thesecond antennaearefoundonlyincrustaceans,andareprobablyhomologoustothechelicerae(fangs)of thearachnids.Otherheadappendagesthatareconsistentacrossthecrustaceansareonepairof mandiblesandtwopairsofmaxillae.Theseareusedinfeeding.Thetrunkofcrustaceansvariesa lot from one class to another, and shows varying degrees of regional specialisation. In many Crustaceathethorax iscoveredbya carapace (a partoftheexoskeleton thatextends ventrally fromthebackofthehead).Insomecrustaceans(e.g.ClassBranchiopodaandClassOstracoda) thecarapaceextendsdownwardsoverthesidesofthebodytoencloseitliketheshellofabivalve mollusc. Thelimbsofcrustaceansarebiramous,(i.e.twobranched,asseenearlierinthetrilobites).The branchontheoutsideofthelimbiscalledanexopodite,whilethebranchontheinsideiscalled an endopodite. The trunk appendages are extremely variable across the Crustacea, and have beenspecialisedforahugerangeoffunctionse.g.feeding,swimming,eggbroodingandsperm transfer.

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 Whatistheexoskeletonofcrustaceanslike? Theexoskeletonofcrustaceansisquitedifferenttothatoftheinsectsandmostotherarthropods. Incrustaceansboththeepicuticleandprocuticlecontaincalciumcarbonate(referFigure2inthe ArthropodDiversitylecture),andtheepicuticleisnotwaxyandwaterprooflikethatoftheinsects orarachnids.Thecuticleofcrustaceansisoftenbrightlypigmented. Howdocrustaceansbreathe? Being largely aquatic, most larger crustaceans breathe via gills. Their gas exchange system is closed (cf. the open system of insects), and the oxygen that diffuses through the gills is either transported in simple solution in the haemolymph, or bound to blood pigments such as haemoglobinorhaemocyanin.Wewillnowconsiderafewspecificexamplesofcrustaceans.

ClassMalacostraca
The class Malacostraca is the largest class within the subphylum Crustacea, containing about threequartersoftheknownspecies.ThethoraxoftheMalacostracahaveeightsegments,each bearingapairoflimbs(maxillipedsorlegs).Theabdomenhassixsegments,andthefirstfiveof these bear a pair of biramous pleopods. The sixth (termina) segment has a pair of flattened uropods. We will consider two orders within the class malacostraca: the Decapoda and the Euphausiacea.

OrderDecapoda(Deca=ten,poda=feet)
ANewZealandexampleofadecapodisthefreshwatercrayfishorkoura(=Paranephrops).Koura occurinawidevarietyofhabitatsincludinglakes,ponds,streamsandswamps.Kouraarebottom dwellers,andtheirthoraciclimbsareusedforwalking.Thefirstpairoflegsbarelargepincersor chelipedswhichareusedforbothoffenceanddefence.Ifafightgetstoomuchforthemtheyare able to travel very rapidly backward for short distances by bending their abdomen and tail fan violentlytowardsthethorax.Thepleopodsontheabdomenoffemalesareusedtocarrytheeggs, whicharefirmlyattachedwithaform ofglue.Oncehatchedtheyoungremainattachedtothe femaleuntiltheirthirdmoult.

OrderEuphausiacea(Krill)
Krillarepelagic(livingintheopenocean)shrimplikecrustaceansapproximately5cminlength. Theappendageshavedifferentstructuresandfunctionsdownthelengthoftheanimal.Krillare filter feeders. The first six appendages on the thorax function as a filtering apparatus. Each of the leglike endopodites has a long fringe of setae (hairs) on one side, and together with the other limbs of that side they form one half of a funnelshaped net or basket as they swim. All but the first pairofthoraciclimbshaveagillattheir base. Much of the propulsion for swimmingcomesfromthepleopodson theabdomen. Figure2(CopiedfromBarnes,1987)

KrillliveingreatswarmsinthesouthernoceanaroundAntarcticaandarethechieffoodofmany whalesandpenguins.Bluewhalescanconsumeatonneofkrillatonefeedingandmayfeedupto fourtimesaday.Theswarmsarepatchybutmayreachdensitiesof63,000individualspercubic metre.

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Krillisreceivingincreasingattentionasapotentialhumanfoodsource.Currentlyaround400,000 tonnesareharvestedeachyear.Mostofthisiscrushedandpressedtoobtainjuicesfromwhich theproteinisextractedandusedtoenrichotherfoods.Amajorconcernaboutharvestingkrillis theunknowneffectthismayhaveontheAntarcticecosystem.PracticallyeverythinginAntarctica dinesonkrill.Theyarethebigmiddlelinkofwhatisessentiallyathreepartfoodchainalgae, thekrill,theneverythingelse,fromwhalestopenguins.

ClassMaxillopodasubclassCirripedia
ThesecondclassofCrustaceawewillconsideraretheCirripediaorbarnacles.Theyaretheonly groupofcrustaceansthatspendmostoftheirlifeattachedtoasubstrateratherthanfreeliving. They are exclusively marine and attach to a wide range of objects such as rocks, shells, coral, floatingtimer,orevenwhales,turtlesandfish.Theyaresodifferentinappearanceandlifestyle fromtheothercrustaceansthatuntil1830theywerebelievedtobemolluscs. There are two types of barnacles, stalked and stalkless. The stalked type are believed to have evolved first, and given rise to the stalkless barnacles. The stalk of stalked barnacles is called a peduncle. Most stalked barnacles live attached by their peduncle to floating objects such as driftwood. The rest of the barnacle's body is called the capitulum. The stalkless barnacles are sometimescalledacornbarnacles,andconsistalmostentirelyofcapitulum. T.H. Huxley once described barnacles as animals that 'stood on their heads and kicked food in theirmouths'.Thisisessentiallytrue.Inacornbarnaclesthebarnacle'sheadiscementedfirmlyto thesubstratebysecretionsfromthefirstpairofantennae(=antennules).Thebarnacle'sbodyis enclosedinfivecalciumcarbonateplates.Barnacleshavesixpairsofbiramouslimbscalledcirri (hencetheirnamecirripedia)thatarefringedwithstiffhairs.Barnaclesfeedbykickingtheircirri outandretractingthem,therebycreatingwatercurrentfromwhichsmallorganismsarefiltered.

Conceptcheck
WhatarethedistinguishingcharacteristicsofCrustacea?

Vocabulary
Youshouldknowandunderstandthefollowingannelidfeaturesandconcepts: Antennae, carapace, branchiopoda, ostracoda, biramous, exopodite, endopodite, Crustacean, exoskeleton, haemolymph, haemoglobin, haemocyanin, malacostraca, maxillipeds, pleopods, uropods,chelipeds,decapoda,euphausiacea,pelagic,cirripedia,setae,peduncle,capitulum,cirri.

References
Barnes,R.D.1987.InvertebrateZoology5thEdition.SaundersCollege.Philadelphia. Hickman, C.P., Roberts, L.S., Keen, S.L., Larson, A, Eisenhour, D.J. 2009. Animal Diversity, 5th edition.McGrawHill.Boston,USA.

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Lecture 19 Insecta
Keyconcept
Insectshaveasimpleyethighlyadaptablebodyplanthathasenabledthemtobecomethemost abundantanddiversegroupofanimalsonearth. Thislectureisbasedonpages734737inChapter33andappropriateportionsoftheformand functionchaptersgivenbelow. TheClassInsecta(orHexapoda=`sixfooted')containsmorethan750,000describedspecies.This notonlymakesthemthelargestclasswithintheArthropoda,butalsomorenumerousthanallthe otheranimalgroupscombined.Insectsoccurinnearlyeveryhabitatexceptthedeepsea.Many speciesareaquatic,eitherasadultsorlarvae,andafewspecies(theoceanstriders)liveonthe surfaceoftheopenoceans.Mostspecies,however,areterrestrial.

Whyhaveinsectsbeensoenormouslysuccessful?
Perhaps the singlemost important factor contributing to the success of this group was the evolutionofflight.Flyinganimalscanescapemanypredators,findfoodandmates,anddisperse tonewhabitatsfasterthanorganismsthatmustcrawlontheground.Nootherinvertebrates,and relativelyfewvertebratespecies,haveevolvedthiscapability.Theevolutionofflightopenedupa vast range of lifestyles that no previous organisms had been able to exploit. The possession of wings provided, and continues to provide, extraordinary dispersal capabilities. Flight made possiblethedevelopmentofcompletelynewwaysoffindingmates,seekingfood,andescaping frompredators.Manyinsectshaveoneortwopairsofwingsthatemergefromthedorsalsideof thethorax.Wingsareextensionsofthecuticleandarenottrueappendages. Severalhypotheseshavebeenproposedfortheevolutionofwings.Inonehypothesis,wingsfirst evolvedasextensionsofthecuticlethathelpedtheinsectabsorbheatandwerelatermodified forflight.Asecondhypothesisarguesthatwingsallowedanimalstoglidefromvegetationtothe ground. Alternatively, wings may have served as gills in aquatic insects. Still another hypothesis proposesthatinsectwingsfunctionedforswimmingbeforetheyfunctionedforflight. Insectshavealsoshowngreatadaptabilityintermsoftheirbodyplan.Thishasenabledthemto live in widely different climatic conditions and temperature ranges: from 50oC for some polar speciestoover60oCinsomedesertdwellers.Manyinsectsare parasitesonorinthe bodiesof otherorganisms.Othersliveinseeminglyimpossibleenvironments.ThelarvaofanEphydridfly (Psilopapetrolei),forexample,livesinpoolsofcrudepetroleuminCalifornia! Whatisthegeneralbodyplanoftheinsects? Theinsectbodyisdividedintothreetagmata:head,thoraxandabdomen. i) Thehead Theheadbearsonepairofantennaeandonepairofcompoundeyes.Thecompoundeyeismade upofmanyindividualunitscalledommatidia(ommatoisGreekfor`eye';iumisGreekfor`little'). Thelightfromeachommatidiumisintegratedbytheinsect'scentralnervoussystemtoforman image.Inadditiontothecompoundeyesthereareoftenthreesingleuniteyescalledocelli.Ocelli are sensitive to varying light intensity and, sometimes, specific wavelengths (colours), but generallydonotformanimage. Three pairs of appendages contribute to the mouthparts: one pair of mandibles, one pair of maxillae,andthelabium.Althoughthelabiumappearssingle,itisactuallytheproductoffusion

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between a pair of second maxillae. In front of the mandibles is a labrum. This extension of the front of the headfunctionsasanupperlip.Arisingbehindthemaxillae nearthebaseofthelabrumisafurtherstructurecalledthe hypopharynx that aids in swallowing. This primitive arrangement of the mouthparts suits insects that chew. Many insects show modifications of this arrangement, usuallytosuitasuckingorpiercingfeedinghabit.Youwill get to see some of these modifications in the Arthropod laboratory.

Text Figure50.16 Compoundeye.

TextFigure50.9Aninsectear.

ii) Thethorax Themiddleregionoftheinsectbody(thethorax)comprisesthreesegments,eachbearingapair oflegs.Thelegsaregenerallyadaptedforwalkingorrunning.However,sometimesoneormore pairsaremodifiedforspecialfunctionssuchasgraspingprey(e.g.preyingmantis),jumping(e.g. fleas),swimming(e.g.waterboatmen)ordigging(e.g.molecrickets). The thorax also usually bears two pairs of wings. The wings are outfolding of the thoracic integumentandarecomposedoftwothinsheetsofcuticle.Thewingshaveveinsthatopeninto thebodyandcontaincirculatingblood(haemolymph). In beetles, the posterior wings function in flight, while the anterior wings act as covers that protecttheflightwingswhenthebeetleisonthegroundorburrowing. iii) Theabdomen The third insect tagma is the abdomen, which has 9 to 11 segments. Larval insects may have a varietyofabdominalappendagesonthese,servingarangeoffunctions,butinadultinsectsthe only appendages here are a terminal pair of sensory cerci. The abdomen bears the external genitaliathatarediverseinstructure.Insectsaredioecious(maleandfemalesexorgansoccurin differentindividuals:cf.theannelidsinwhichmanyindividualsarehermaphrodite). Theexoskeletonofeachsegmentinboththethoraxandtheabdomenismadeupoffour`plates'. Thedorsalplateiscalledatergum,theventralplateasternum,andeachofthelateralplatesa pleuron.Eachthoracicandmostabdominalsegmentsalsohaveapairofspiracles(oneoneach pleuron).Thesearetheopeningsintotheinsect'srespiratorysystem.

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Howdoinsectsbreathe?
Each spiracle opens into a complexly branching network of tracheae, which further branch into tracheolesthatarelessthan1mindiameter(Figure1;seealsotextFigure42.24).Thesebranch againuntilallthecellsintheinsect'sbodyareclosetoatracheole.Thissystemiscalledanopen respiratorysystembecauseoxygeniscarrieddirectlytotheinsect'stissuesbythesetubes,and theblood(haemolymph)isnotinvolvedinthetransferofgases.
Figure1

(CopiedfromPechenik,1991)

Whataretheinsect'sdigestiveandexcretorysystemslike?
The basic layout of the insect digestive tract is presented in Text figures 41.18 and 44.13. The mouth opens into a pharynx that may form a muscular pump in blood or plantjuice feeding insects. This is followed by the oesophagus, which is often enlarged posteriorly to form a crop thatservesasastoragechamberforthefoodbeforeitispassedontothemidgut.Togettothe midgutthefoodmustpassthroughthegizzardorproventriculusthatgrindsthefoodintosmall fragments. The midgut is the region in which most digestion and absorption occurs. It frequently has a numberofgastriccaeca,whichsecreteenzymesthataiddigestion.Betweenthemidgutandthe hindgutisaregionwheretheMalpighiantubulesconnectwiththegut.TheMalpighiantubules, whichnumberupto250pairs,arethemajorexcretoryorgansininsects.Theyareresponsiblefor the active transport of metabolic waste products from the haemolymph into the gut, and subsequentlyoutofthebody. The hindgut carries the undigested food and the metabolic waste products to the rectum and anus.Thehindgutalsoselectivelyabsorbswater,salt,andaminoacidspreviouslyremovedfrom thehaemolymphbytheMalpighiantubules.

Howimportantareinsectstopeople?
Insectsimpingeonusinmanyways.Abouttwothirdsofallfloweringplants,includingmanyof agricultural importance, depend on insects for pollination. Many insects, like mosquitoes, lice, fleas, bedbugs and many flies bite us and suck on our blood. These, and other insects, are also significant vectors of diseases like malaria, elephantiasis, bubonic plague, typhoid fever, yellow fever, sleeping sickness and dysentery. Many other insects munch their way through our crops both before and after they are harvested, and vast sums of money are expended annually on

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insect pest control. On the brighter side, some species such as the silkworm and the honeybee havegreatlybenefitedhumans.

Conceptcheck
You should be able to explain why insects have been able to become so numerous, diverse and widely distributed.

Vocabulary
At the end of this lecture, you should know the meanings of the following words and, more importantly,youshouldbeabletousetheminwrittenwork: Ommatidia, ocelli, mandibles, maxillae, labium, labrum, hypopharynx, thorax, abdomen, haemolymph,tergum,sternum,pleuron,spiracles,tracheae,tracheoles,openrespiratorysystem, pharynx,oesophagus,crop,midgut,gizzard,proventriculus,gastriccaeca,malpighiantubules.

Reference
Pechenik,J.A.1991.BiologyoftheInvertebrates.2ndedition.Wm.C.BrownPublishers,Dubuque.

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Lecture 20 Echinodermata
Keyconcept
33.8Echinodermsandchordatesaredeuterostomes. This lecture is based on pages 706707 in Chapter 32 and pages 738740 in Chapter 33 and appropriateportionsoftheformandfunctionchaptersgivenbelow. Deuterostomedevelopment ThephylumEchinodermatadeuterostomes.Itisimportantthatyouunderstandthedifferencesin developmentbetweenprotostomesanddeuterostomes(pgs706707,Figure32.9). Deuterostome development is a type of embryonic development shared by echinoderms, hemichordates and chordates and defines the second major evolutionary line of the animal kingdom. The other major line is protostome development, which is shared by flat worms, annelids, arthropods and molluscs. In deuterstome development the mouth arises as a new openingandtheanusformsfromorneartheblastopore(intheprotostomesthemouthformsat the blastopore), cleavage is radial (not spiral as in protostomes) and the fate of blastomeres is fixedmuchlaterindevelopment.

Echinodermcharacteristics
The name of the phylum means spiny skin (from the Greek echin, spiny, and derma, skin) and many of these animals have spiny projections on their outer surface. It is an exclusively marine phylum and includes the common starfish (Class Stelleroidea). Sea stars and most other echinoderms are slowmoving or sessile marine animals. A thin skin covers an endoskeleton of hardcalcareousplatesorossicles.Mostechinodermsarepricklyfromskeletalbumpsandspines. Unique to echinoderms is the water vascular system(seefigure22.3thewatervascularsystem ofaseastar,Hickmanetal.(2004)),anetworkof hydraulic canals branching into extensions called tube feet that function in locomotion, feeding, sensory perception and gas exchange (Figure 33.39). Sexual reproduction of echinoderms usually involves separate male and female individuals that release their gametes into the water.
Figure33.40Anatomyofaseastar,anechinoderm

Theinternalandexternalpartsofmostechinodermsradiatefromthecentre,oftenasfivespokes. However, the pentamerous radial symmetry of the adults is a secondary characteristic, not an ancestral one as in the Cnidaria, because the larvae are bilateral. Some adults e.g. the Holothuroidea,showatendencytowardsbilateralsymmetry.Furthermore,thesymmetryofadult echinodermsisnotperfectlyradial.Forexample,theopening(madreporite)ofaseastarswater vascularsystemisnotcentralbutshiftedtooneside. There is an endoskeleton of small calcareous ossicles in the body wall and the whole outer surface,includingthespines,whicharecoveredbytheepidermis.

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Themouthisusuallyinthecentreofonesideoftheradialbody.Inmostechinodermsthissurface the oral surface is towards the substrate. The anus is usually on the aboral surface the oppositesideofthebodytothemouth. Internal transport is mainly by the coelomic fluid. Exchange of gases and waste between the coelomicfluidandseawateroccursacrossthetubefeetandothersurfacestructures. Thesexesareseparateandtheembryonicdevelopmentresultsinaciliatedlarva.

Echinodermdiversity
Living echinoderms are divided into six classes Figure 33.4133.45: Asteroidea (sea stars), Ophiuroidea (brittle stars), Echinoidea (sea urchins and sand dollars), Crinoidea (sea lilies and featherstars),Holothuroidea(seacucumbers),andConcentricycloidea(seadaisies).

ClassAsteroidea
Thecommonstarfishoftherockyshoreareinthisgroup.Theytypicallyhave5arms,whichare not sharply set off from the central disc. Some, such as sun stars, have many arms. The arms radiatefromacentraldisk.Theundersurfacesofthearmsbeartubefeet,eachofwhichcanact likeasuctiondisk.Byacomplexsetofhydraulicandmuscularactions,thesuctioncanbecreated or released (see Figure 33.40). The tube feet emerge from an ambulacral (treelined) groove down each arm. The sea star adheres firmly to rocks or creeps along slowly as the tube feet extend,grip,contract,release,extend,andgripagain.Seastarsalsousetheirtubefeettograsp prey,suchasclamsandoysters.Thearmsoftheseastarembracetheclosedbivalve,hangingon tightly by the tube feet. The sea star then turns its stomach inside out, everting it through its mouth and into the narrow opening between the halves of the bivalves shell. The digestive systemoftheseastarsecretesjuicesthatbegindigestingthesoftbodyoftheMolluscwithinits ownshell. Seastarsandsomeotherechinodermshaveconsiderablepowersofregeneration.Seastarscan regrowlostarms,andmembersofonegenuscanevenregrowanentirebodyfromasinglearm. The nervous system is comparatively simple in asteroids, with no central neural control of locomotion.

ClassOphiuroidea
Thesearebrittlestarsandbasketstarsandarethelargestgroupofechinoderms.Theyareclosely relatedtotheAsteroidea,butdifferinhavingacentraldiskthatissharplysetofffromtheirlong slenderarms.Thesearmsareverymobileandbrittlestarsmovebyflexingtheirarmsandpushing withthemratherthanbywalkingwiththeirtubefeetastheasteroidsdo,insteadtheymoveby serpentinelashingoftheirarms.Thearmsareeasilybrokenhencethecommonname. Compare the aboralsurface of an ophiuroid with that of an asteroid (see Hickman et al. (2004) figure22.12,Theaboralsurfaceofanophiuroid). Somespeciesaresuspensionfeeders;othersarepredatorsorscavengers.

ClassEchinoidea
Thisclasscontainstheseaurchinsandsanddollars.Seaurchinshavenoarms,buttheydohave five rows of tube feet that function in slow movement. They also have muscles that pivot their longspines,whichaidsinlocomotion.Theyhaveasphericalbody,inwhichtheskeletalossicles arefusedtogethertoformflattenedplates,whichformarigidinternalshell.Basicallythebody

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formislikethatofanasteroidwhichhashadthetipsofthearmsraisedandjoinedsothatthe oralsurfaceformstheoutersurfaceandthenthegapsbetweenthearmsarefilledin. (SeeHickmanetal.(2004)figure22.18,Theinternalstructureofaseaurchin) Themouthofaseaurchinisringedbycomplex,jawlikestructuresadaptedforeatingseaweeds and other food. Sea urchins are roughly spherical, whereas sand dollars are flattened and disk shaped.

ClassHolothuroidea
Holothuroidsareseacucumbers.Theiroral/aboralaxisisgreatlylengthenedandtheylieontheir side.Theirshapeiscucumberlikeandthebodywallisleathery,lackingspines,andtheossicles arereducedtomicroscopicsize.Inspecieswhichliveonhardsubstratesthereareusuallythree ventralambulacralgrooveswithwelldevelopedtubefeetwhichareusedformovementandthe tube feet of the other two ambulacral grooves are reduced. Some of the tube feet around the moutharedevelopedasfeedingtentacles.Theirabilitytoejecttheirvisceraisunusual. (SeeHickmanetal.(2004)figure22.22,Theanatomyofaseacucumber).

ClassCrinoidea
Crinoids are sea lilies and feather stars. The crinoids are believed to be the oldest living echinoderms and are thought to show the ancestral mode of existence and function. Fossilized sealiliessome500millionyearsoldareextremelysimilartopresentdaymembersoftheclass. Thecrinoidsdifferfromallotherlivingechinodermsinhavingtheiroralsurfacedirectedupwards and,inthecaseofthesealilies,beingsessileanimalspermanentlyattachedtothesubstratebya stalk.Featherstarshavethestalkduringtheirdevelopment,butitisnotpresentintheadultsand theyaremotile,swimmingwiththeirarms,butspendinglongperiodsattachedtothesubstrate bycirri.Crinoidsaresuspensionfeeders,usingtheirtubefeettocaptureparticlesfromthewater and then transporting the particles entrapped in mucous via the ambulacral grooves to the mouth.Feeding,ratherthanlocomotion,isthoughttohavebeentheoriginalfunctionofthetube feet.

ClassConcentricycloidea
Sea daisies were discovered in 1986, and only two species are known. Both live on submerged woodoffthecoastofNewZealandandtheBahamas.Theirarmlessbodyistypicallydiskshaped, hasafivefoldsymmetry,andmeasureslessthanacentimeterindiameter.Theedgeofthebody isringedwithsmallspines.Seadaisiesabsorbnutrientsthroughthemembranesurroundingtheir body. The relationship of sea daisies to other echinoderms remains unclear; some taxonomists considerseadaisiestobehighlyderivedseastars.

Conceptcheck33.5
[Questions1and2].

Vocabulary
Deuterostome, water vascular system, tube feet, madreporite, pentamerous radial symmetry, calcareous ossicles, oral and aboral surfaces, stelleroidea, ambulacral groove, ophiuroidea, holothuroidea,crinoidea,concentricycloidea.

Reference
Hickman,C.P.,Roberts,L.S.,Larson,A,IAnson,H.2004.IntegratedPrinciplesofZoology.McGraw Hill.Boston,USA.

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