Palaeontology — An Introduction by E. W. NIELD Universities Information Unit, Committee of Vice-Chancellors and Principals, London, U.K and V. G>Te TUCKER formerly Yale Sixth Form College, Wrexham. Clwyd. UK & PERGAMON PRESS OXFORD - NEW YORK - BEIJING - FRANKS 5 ‘SAO PAULO : SYDNEY « TOKYO - TORONTO TAbout This Book Tin ic en a a a penne a eee a ane Seon bn Sica Soar ey a eet tect ears nee arn Fee ee ae nian oe os Ce Nene Tee estan or Sepa tpatoaniic ape peri nee eta a Oot we ee osen ee oe Sip ies teen opens uae caes oes, Fees gets beet oo ee ipa ee prpecart eed aerey & wwe aes soto om ee ee eee ese ae emeracondea nee le ty ee ipod yo aeoreshabeertestr open praen ee te oe re se a ela icy epee cade whole range of form present in any group, they may also be scen a additional sources ee ace Tey sell te easly comand and cca capt eee Ny atone wy of pico cn toe ion: secure knowledge of its’ vocabulary. es aek ada wore he 2 renee of aes and ain ao ave rec nogircn picks os trnly bed serra cmc, ok eR te catego be taiotng cf oer stcony, Wels note! Ww give some idea of how science actually works, and of how it regulates itself. So — notwithstanding the great importance of drawing and precise description — we hope Bie aes tn thee 2 caty tae geek fc te vel um een, ie acaty not the end. To id cnrclatin, invxigatecvlution sod 1 Rn ear ie sel icrocsons of mac somal fn vied cng — ie sie fo cert Bee oe to crogntng ol ming) os fonalons by deducing function we ihm Fe ecipnieusprocne which has to ou own ex Soi hoa the idea which Bing che fats together rms, we can analyse inate evolution, and by appreciating that fund: le 10 see, a little more liable correlation and are abl sence. This science, like any other, NContents List of Tables 1. Introduction — Fundamental Questi Whats palscomiologs? 12Whatns tonal? — + 1.3 What features and conditions are favourable for pr- fossl skeletons, it establishes the reasons behind the design of mysterious skeletal features. (Palaeontology is simply the history of nature. and the natural scientific answer to the question “Where do we come from=> With advances in our understanding, many subdivi- sions have arisen within palacontology. The study of fossil plants is known as palaeobotany. and while this would suggest the ttle “‘palacozoology” for the study of fossil animals, the term is not generally used Palaeobiology tends to be used to refer 20 the more interpretative ‘biological’ aspects of the science although these are perhaps more cozrectly combined with the study of fossil communities under the general title of palaeoecology. The studs of small fossils for which a microscope is often (though not zisavs) needed is known as micropalaeontology ‘This science has a special botanical branch. concerned with fossil spores and pollen, called palynology. applying 1.2 What is a fossil? erally, “fossil” means ‘that which is dug up’ —) but the modern meaning of the word has .quired many refinements. For something to be 2 fossil. it ‘must either be the remains of an ancient orz"ism, or the trace of the activity of och an onan. But fossil’ as an adjective may be used to refer to inorganic things: for example. 2 “fossil volcano’ or a “fossil sand dune”. In these case~ reference is made to their former existence before burial and_ preservation. : So there are two types of true fossil — body tossils and trace fossils. Body fossils a: ‘he actual remains of organisms. (Trace fossils are indirect signs of life; saur footprints. worm burrows, trilobite grazing trails and other evidences of life processes, such 2s fossil excremen’ (coprolites). ) 1.3 What features and conditions are favourable for preservation: ‘The soft, fleshy tissues of an organism may bea served in hm way, ‘a dinosaurs shih othe ad oh sta). Dison 2 js shun 2 pretem abBaciing the tty oh Sos which ace pearhy abwas foam a8 fragmen the years A rescanch to ecticks ack be |. which ‘eck eons to wach stem, wihict which comes, wince pode Wo winch Prwess wai > Tine penta oh lem meri » ve Dusiness. Uveryone kame that «st fom ddetorin, trot there is sr Wetthe ates! Femainiong ins cok that i ol prahecetonn n'y Usectiah port boron! © p r J thew bashc wns: ws hangressions, comers petndactiony Ais ener carmnains 1e) 2- in creche Norm oh, saya best =" ‘reas, impressed pom 9 beions schay on it. Whee farms shay p> | nasser, Vicks Varcene plans rman *" mraier, wnaiy a + black Cannas ne, Woy comin, greet” omnes chno™ refers to a spiders wed. ! Greek legend, Arachne 3s * Tyalian maiden who ene! * spinning contest with the &° Homo sapiens Prowospongte fe Srenaseisma djuunbletonensis Anachnophvlluon murchisontddess Minerva and was turned in- to a spider. The scientific name for spiders is ‘Arachnida’. ‘phyl- Jum’ means. leaftike’: (phvllon = eat (Gr)); murchisoni = “of Mur- chison’, after Sir Roderick Impey Murchison (1792-1871), founder of the Silurian and Permian Sys- tems and onetime Director General af the Geological Survey of Great Britain “This system of naming has been with us since it was thar by the Swedish botanist Cor! von Linné 502-78), more commonly known by his Latinized wnacus, in his book Systema Narurae (The see af Nature) in 1758. But it does not end at the nus level. Similar genera are grouped into families, milic; into orders, orders into classes and classes into ‘e greatest division of all. the Phylum. (The names tese groups are slightly different in botany, but the finciple of the classification is the same.) There are over twenty phyla in the Kingdom nimalia, some containing millions of species, seme aly a few. Ard alongside the animal and plant ingdoms (familiar to all of us) there are three oth. ingdonis comprising organisms which cannot be escribed as either animals or plants; these are iscussed in Chapter 10, ‘There has always been some degree of unease about fe fact th some of the phyla have seemed ridicu- ously big, und this has led recently to the creacion of a uperphylum’ wo cnclose all invertebrate animals with \inted appendages. So the new Superphylum Arthro- oda (it used 10 be called simply ‘phylum’) now dmtain- shree-quarters of a million insects (Phylum asecta, formerly a class of the old phylum) as well as ve king crabs, spiders, scorpions, centipedes,,mil- Pedes, true crabs, lobsters, shrimps, water fleas, stracods, copepods and barnacles — together with lany more minor groups and several major extinct roups: ‘such as the trilobites. In contrast, the Phylum. Fiapulida contains a mere eight species. To sim up, we have a basic unit of taxonomy called ne species which has a recognizable existence in the eal world. These species are grouped in progressively ‘igher categories from genera to phyla, but these ligher groups are frequently changed as the state of vur knowledge changes. This means that species may ove from genus to genus: phyla be broken up into ie py ‘or degraded to class level, or elevated 10 & ink of superphylum. Higher classification is an merpretation. Here is the classification of our own Pecies as it stands at present. Kingdom Animalia (Subkingdom Metazoa) all animals all animals consist- ing of more than one cell Intro luction — Fundamental Questions 9 Phylum Chordata Possessing an axial, dorsal nerve cord (Subphylum Vertebrata) cord nerve sur- rounded by a bony spine Class Mar imalia true hair, nursing of young, brain of advanced type mostly tree- dwelling placental ‘mammais, monkeys, apes and ‘man and his im- mediate: ancestors ‘man sensible, judicious. Order Pr mates (Suborder Anthropoidea) Family fomini Genus fomo Species sapiens So th: species is a concept created by biologists: but a biologist deals only with life as it 1s now. A biologist ‘can sce species alive in nature, and can watch them interbreeding. Palacontologists, on the other hand. can not. Moreover, we have to deal with species as they progress through time. We have to think of species in four dimensions, not just in the convention- al thr (The biologist’s species is the biospecies, the true nd full expression of Linnaeus’s concept. But fossil species are describrd by form alone, and are defined solely ou their morphology. They are therefore termed. morphospecies. é.nd since fossil species also have a distribution in the time dimension, from their incep- tion to their eventual extinction, they are als: chronospecies. So palacontoldgists can never be sure that their morphospecies were true biospecies — though it is a reasonable assumption, particularly sin. most living species are clearly recogni/..sle on form. But remem- ber that not only must a palacontologist set limits on the morphological range of a species at any one time; he must also set limits upon the evolutionary variation in form seen through time. Progressive changes in form can therefore become so pronounced that the palacontologist may deem the form to have become sufficiently different for it to receive a new specific name. 1.13 Who creates the classification? When a new species of fossil is discovered, a new name must be chosen. We have seen how it can be descriptive, or refer to the place where it was found, or bbe samed after someone (this can be anyone — even the author's pet dog — but it cannot be the author hisself). The description of the new species, together with abundant diagrams and photographs, is released in aPalacomtolony — An Introduction cil scientific journal such ax Palaeontology. which is published by the Palaeontological Association. ‘The description. in the form of a scientific paper, is examined by the editors of the journal and by a number of referces who are chosen for their special knowledge of the particular group to which the new species belongs. Once the manuscript has been corrected and the revised version accepted, it will (in due course) appear in print. This is how the basic work of palaeontology is cartied out, Without it. and without adequate classification, no science would be possible. In order that future sev wists may re-examine the material, all the fossils illustrated in the paper are deposited in a museum. In most cases this selection ‘will include one special specimen chosen by the author 0 represent the characteristics of the new species to the fullest degree. Ihis is the ‘type specimen’. Its location and its inuseum number will appear in the paper when it is published. Because taxonomy is s0 importint, no taxonomic work is recognized unless these procedures are carried out. There is a complex set of rules governing the establishment of classification and ensuring that it remains stindard. It is known as the International Code of Zoological Nomenclature (there is a separate fone for botanists) laid down by an international ‘commission, 1.14 What is ‘numerical taxonomy’? When biologists first began classifying organisms, they found that they were able to perform this task by referring particularly to certain features which seemed. 10 offer the closest approximation to the true deriva- tion of the animal or plant, Linnacus, for example, used floral auributes to classify flowering plants, rather than, say, the shape of their leaves. Ammonite specialists use the complex pattern of suturing which marks the junction between the outer shell and its internal walls, ‘These things are thought by specialists, Wo be less sensitive to shorter changes, and therefore more expressive of an W's origins, Recently, there has been a move away: from this Mtvle of approach, and the new methods are converts in palacontology also, Crities of the methods see the choice of special (or ‘characters as subjective and! unscientific. Rather than making “before-the-fact’ decisions about which tea Nes ‘are important, they prefer 1 amass ever jbervable measurement and chaructristie about the Specimens under review, and then to use a computer to make an objective assessment of hovw similar one set ‘of characters is (9 another, without “Weighting” any particular feature beforehand. "The Oxfond. Linglish Dictionary detines “objectivity* as “treating a subject 30 88 to exhibit the actual fiets, not coloured. by the feelings oF opinions of the write, >, numerical taxonomy aims to do * Although it is a new field, the concery. be realized practically until compu. developed that cw. id deal with the mmcrce Jaborious calculation. Using in apy. ro0 # computer can analyse similarity over inc ce" of attributes, plotting i results in « displays these resemblances graphically: There are many types of array 0 choose like the dendrogram shown, sing 2“; approach. Others may come in the form ¢ Points, whose closeness of cluster represen. similarity So, ina typical array, the most closely ‘Spaced poi may represent one species. Each species clr n occur in one ‘of two groups of ‘clusters, and thee Broupings may represent genera method does not tell the taxonomist where to drs line between species, or between species and sexe, This is left to subjective judgement, because s mately all taxonomy must bh. a Matter for fall! human decision. Numerical methods merely resin: % similarity o° 2 0 100 So oc Ficuny 14 This, agra is know 3.4 ‘derdrogeam’, and ste wniely os! 2 numerical txonomy TE ee i at afferent paresis tween aay of species sormpt hay termina the Sorrespondence etwcen the = Characters om ach specimen th 5 set et epee th 8 ivanching pte hn Hand 9 bike, The si 8 Mee where to uw sec toundaces. Ar * Ch TES fe De EA + Eland G sash ines) Soest alin srs go 4 a CES) hat the final al 2° mactsty se e The the an) [DAEG 60 i he emus?the degree to which such subjective assessments affect imp ofthe major advantss ofthe metho is ha, by removing the necessity for years of personal acquaintance with a particular group, the taxonomy can be worked out much more quickly. The work is also ‘repeatable’ — i.e. another scientist can easily the results and come £0 the same conclusions with the same data. But it must be said that the new f science of taxomenics usually finds itself confirming the work of classical taxonomy. ts advantages are chiefly those of speed 1.15 Welcome to palacontology We hope that the brief ousline set out in this chapter has encouraged yor" 10 find out more about the science Introduction — Fundamental Questions 4) of palacontology. The next chapter deals with the ways in which life originated on earth, and describes some of the oldest Known’ fossils, from rocks of the Precambrian, From Chapter 3 on, all the major fossil groups ofthe Phanerozoic (Cambrian to the present) are describes and Chapter 12 attempts to bring together the major themes which have emerged within a discussion of the principle of evolution, ‘We hope that you will make use of the many annotated diagrams to make your own scaled drawings ‘of fossil specimens. The easiest way to learn isto see things tor yourself, and to record them. Its also vital to understand the reasons behind she appearances. AS a matter of necessity we ask the question ‘what?": but only so that we may then ask “why?The Origin of Life and the Earliest Fossils JE When and how ilght tite awe Beaman? ‘hive Cart yeas favimedd ABOU HOOD HTTION VERE AHO, ut fe probably spent about 100 villian lifeless yeare efore the fet primitive bacterial organinme devel ape pan Ah Dini Uwe tvmtene peel Caleost ywiee 8 Tonge we the perloat frown the Cambrian (0 (he ‘ovawent) the erent was Likely (9 have been extremely: wwobile aad wubjeet fo aveh qarnie liiluenees ae meteor Homibartivent, ‘Phere is abe evitenee that, jrametine cturiing (Nie early: pase, the entire ation: ‘phere wane tripped away ann showwty: rentonedl as hare: aves emanates! (hom the planet's Unterior “Tivis atmaghere WOMKE Nave been Fel i carbon dHiawiste, ear bon twelve, ANNA, Aysleagen ant nitaen, There noxious wokeanie Anes: -suovrmnest aeranmn Ue Sore EK ann HOS WHAT, HADI Fo poroteet the auefice oF une earahy fhom the lethal ‘ultraviolet raatiation oF the sun Howe coal fe have: ‘avinen natty neler auch coneitiona? Strangely, (Ne: ‘ion of Highning discharges (ant other sources of energy) apo the prinvoatial atmmoaphere woukt have ‘reenable (0 create COMPRAR MoleCteN CONIA OF Jong carbon ator etains, Anat these ate the conte pours upon whieh the ehemiatey oF We depends, Tw a famous experamenty fo}. Mitten of the University oF Ohioauo placenta MNTuFE OF RaNER SANITAE 16 (hat envisaged Far the early atimogphere (te a coset ‘oni Fagettien ait some water (PIR. 21. 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WARING aL ORAHAN G0 evvates Bckeat UNE NWRARIMRANE AREER WHR, ‘td the tHE FR aK ANGER tS CONN toon m= DNAS But expiants ice he Hate HERS vst cory aties Nave ge a RANE AY TOMKUAAE RON ‘Rat actos ais coh al Nave Beet Ae AN NE de Naghtoat ashi en Ue NARS ite WRT Was ANAS (NEA © Sone geRN GReFEIINE NOL WAtURAL prRRANE Ad Dave crewtoa the COMANCHE HALEHIAME Hehe A the activity of Diwchemnical processes Within the ty calllike capsules WHICH MAY Nave Bee the ecutoor OF atl WS ork cath JNO oan wags Lo wn about Abin ea Oe thy catient rms of tte hadl tevehapet, thet + mma of gro th ana sri wohl have beet bo Ke the organic tokeules around then as hw! * Ns coulkt Nave continied Fora ite tine, But wane wer She supply woul Nave ruin O48, the reer * Ss alias (whieh ingest other ike Roeas) hy Nol * Dav the resources because timate. thes all els ~ Wn plats, WRK CaN MAKE orgaMKC MAHER HO ; ‘ateriats using the energy: of the sun. For 9 develo farther thom ite earliest canton, totonyuthenis, FWNIEN Ie Ne plate’ biochemical RR Wau have to evulve, Its event anypeatance |) Senet the most momentous events ot only hoe ie LoMis Bue FOr the earth. itsell, ." like — execpt, perhaps, the growth of bacteria Petei dish of nutrient agar. Each bacterial cell divides once every few min, Growth iy slow at first; after the first period, on hecomes two. two become four, and ss doubling every time In no time at all there millions of cells, yet the individual bacteria arc... dividing any faster. Try it on your calcuis., robe ill 4 multiplying one by two, then three, then four...) see how long it takes you to reach a million This is the way populations giow sen ther a, limits to growth. Of course, the rate slows event because of competition for space or food. But pers those incredible 10 million years at the beginnins the Phanerozoic were the “log phase’ (Fig. 2.3) of development, where instead of individuals, it vas species which were being created. $319908 40°0N Te Bison 24. A. sigmoid cane AMustating the “Tog. phase) — 2 fetid where Togaehmaic 060 fn numbers takes place. Svs? 2 tft may” help. to explain the Sanden development of is a he Tse of the Cambrian But there are other factors to consider. For # 8 time, the sudden appearance of fossils has ' attributed 0 the first development of hard pars other words, we are witnessing a sudden © preservation potential rather than in life itell ‘The arpument is partly valid. Certainly, skeletons were not found in the Eiacara Fauna, °°" any other from that time, Hard exteriors my" ‘ident’ in the first instance, perhaps with Seip sr ep procs te sm Ba Bey geen hard shell, competition would ensure that similar the advantages? Protection, certainly: but a skeleton Re peace peer Mee toes etiam teste Ac ma Thee aid oe oboe cdesnaer would strongly favour ‘skeletonized” groups. with soft-bodied equivalents. This is probably not true, Firstly, no such ‘eq’: valents’ have turned up; ‘hard parts’ from the kinds of animals which use them. The Origin f Life and the Karliest Fossils 45 Without their hard parts, the biology of skeletonized animals would not make sense. You cannot have a soft-bodied arthropod. Such animals need their hard parts to be the way they are, and without them their life would be impossible To sum up, then, the sudden appearance of fossils at the opening of the Phanerozoic is partly an illusion, because of the increased preservation potential. But the origin of hard skeletons also amounted to such an innovation in animal structure that it led to the explosive radiation of new and rapidly developing groups into a world where there were no constraints of space or available food. Revolutionary progress in design and unlimited opportunities for expansion made the Lower Cambrian the single most important Period in the history of life3. Trilobit Phoyfame Triioties (L_ Camie 0 Prem. ugerpeytam Actenprute (ate saeco fas eee pt wr fy Meter (IS73. W977) ats items at me ~ ‘This is the scientific term for the moulting of the euriicleton. Kecause this suir of armour does not shows that this is the result of two deep longitS" sa expand, it must be sloughed off. and a new, larger one furrows which divide the dorsal surfsce into 28° ed | M ees pleara 5. pIUoO- i p the ‘sof’ ste, when the animal is extcemely vulner- ‘oie it might be 25 well 0 punt 0X @ Cael from sof sell 1 eventual exes i called an (aii Diffecent groups of arthropods have different sys- aad see tems of ecdysis. Some moult only until the adult form the rear or tail-end, posterior, oreephan Mentihah, song |_]dene Perewnenae fof 7” fan, Pere vere omni way cont NCI veo jonni oni | (CVV BE Nason Jt caunatonvvestnn eve men Mees ames» mn, ng al es een aeavants a Re every her that the OES Reaturen Common (is many’ Rawer invertes oreR, where the segments are wor fase et ar p Figure 2.1% RARE segmentation of he body, Thin ina late WN ne anne eeakrene Retains nant out thar PREMPR seen to beat vantage in Worms, PUTIN WN ARERR EHCIN, amt URere are apecte: raat’, ‘back’. “SOAR segmentation WE BEM lH UHihOites, VEVOTINES HL ARE RIRECIOR FO athe Ue enraltinent wala ancl the SY peobahty valved from 9 warmtike ancestor he ‘hckedt Ww) affontig even greater xeewrity, Mas dnote on i Ne he Precumt ". w atiest iy Uke eRtRNT State, NUt RY RO means: Mt i Wer ASME llshiekt oF pytiun, an AN thee Tach segment af the Horan Deane @ aie of Waal ‘ws Ue ouiginal segmentation tas ts Wea ant a pai of feat rahe WAR A NE eck ation Nas teen auyynresiet eg aint aE OF FeatNeey apyantages id Sampteety Ah the cepahon than in the Tet enteral ah at are nen saa git Beanahen ‘itd == Wadd the exoskeleton nas become fused (Fig 86) They ake desorbed aiare Milt Bek “NEM Plat, Berween thee two shih Hen the There Wee Abe NHN. DOMAIN the eeNaNON Ai \ I 18 Palaeontology — An fntroductwn pygidium, and from the anterior of the headshield there projected a pair of sensory antennae Deuails of dorsal morphology commonly seen in trilobites are shown and explained in Figs. 3.1-3.5, Soine of these, however. demand further description. 3.3.1 Facial sutures “The facial suture is important in classifi it was also important o the living trilobite, since it was the line along which the cephalon opt open during ‘ecaysis. Typically, this suture runs along the front of the glabelle, around the eyes and then either towards the anterior margin, the posterior margin or the sgenal angle (Fig. 3.3). During moulting, the fixed cheek remained at- taches! th. glabella, while the free cheek fell away: ‘The compound eyes we shell Heft attached to the 3.5.2 Ewes compound ev of the trite example of efficient vision in the anna ‘Their similarity to the eyes of insets ane is obvious, but tis doubifu whether the crn ancestral form of these. More like!,, compensa developed separately in these different group zach eye is divided into many lenses, each ot» covers a small portion of the visual field. In mot arthropods, each lens ligs at the top of a \,. cylindrical structure called an ommatidium, refractive system focuses light onfo a semory sry called a thabdom. The trilobites probably h. 5 the eaticg Glabetar furrow straight or curved groove extwoding across ‘lalla from the dorsal furrow Gabetar lobe area of Glabolla bounded by Short glabelar furrows Doras furrow ~ Glabellar segment — 62 of glabella bounded by short glabeliar furrons 9 furrow — transverse groove in front of hindmost slobelor segment -/ Decipital node—tubercle at centre of occipital segment Free cheeke— ace separated trom the cranigium by the Tacal suture Fixed cheek that portion of the cranigium flanking the label. The plane of symmetcy * Givdes one xed cheek rom ite ‘mirror image symmetry Figure 3.2. Cephalic morphology. Diayrammatic dorsal views Eye—curved plan, facing outwards C-anidium—centr cephalic area comprising fixed cheeks sod glabella (chdeal similar interna The vertical proba which ‘Man so. lan aiffere are fur eves u ‘Gena angle —sbrupt change in direction of Cephale margin posterolteraly10 the sor Akin represent an pound eve ns cNot Which In moxtery of a long HA, Whos bly hid a t ‘cular re narroneaevate Copal tbat ‘Oplathoparian facil suture cus similar syste ‘ternal str The trilobite probably saw the world in a series of Vertical strips of light, and the function of its eves was Probably the detection of movement — a function which compound eyes are well suited to fulfil Many trilobites had no eyes, while others had eyes ipttte they even dwarfed the glabella, ‘These uiferences obviously tell us about mode of life, and ars further discussed belois. Rare trilobites bore their S¥es “upon stalks, and some, though apparently Possessed of eyes, were probably blind nevertheless We believe this e eye structure, when Famined in dewsil, shows variable numbers of lenses om one specimen to another nore or but as yer litle is known of their eyes’ because t Trlobites 49 ear tparian Fecal mitre cuts ‘margin at pena orgle Caphalie tings oad shalt ahong ‘Stalk prolongation bearing eve at apex This is thought to indicate less disordered pattern. functional degeneracy speci exceptional conditions. They wi sd of two elements, lying above it, a gill preset biramous, that is, compo eine walking leg and the othe though this is not certain, and it may have had other functions too, ant bore two such biramous axis, There appears (© ‘of limbs as is forelimbs may be segn each side of tl ‘no functional dil seen, for example, in erabs whoa locton tlle for ratve ‘movement of horace ments ~ Facet sharply dowobent ar. \ ‘och en during neaiment (ahade) tera! portion of thorate segment) “Articlatog hattsng the sched, ‘Shier arte of etl reg wanes ‘roject unger samantha ~ Axial node — tubercle placed centraity on wi orn furrow hig eit ng on Piura furrow exten omerion rom inne ron a 1 Enroll specimen of Acatte downingioe Ail ngs under tension, exposing their arculainghalesngs Pleura of micshoraci region ‘clonly articulated slong foes Frau 3.4. Thoracic morphology. Diagrammatic dors view modified into pincers. The only variation in trilobite limbs was the gradual decrease im size towards the posterior, mirroring the taper of the body itself. Even limbs beneath cephalon and pygidiuin were of the same standard design. Since trilobite limbs were so uniform, since they appear to have shown little evolutionary development and are, in any case, so rarely preserved, they are not useful either in classification or in determining mode of lf. Most of their morphology has been worked out from exceptionally well-preserved fossils in the Burgess Shale (M. Cambrian, British Columbia) and Utica Shale (L. Ordovician, New York) with a few rarities from other localities sround the world. The technique of using X-rays to look at fossils: within blocks of rock has also shown up many interest features, not only of the legs but also of the internal struts of the exoskeleton to which muscles were attached. The two branches of the limb join at an element called the coxa. In some trilobites the coxac of a few anterior thoracic segments bear_sharp_ projections ‘which could have had some function in working over the sediment prior to ingestion (Fig. 3.6). Before proceeding any further with the text, you should familiarize yourself with ell the structures shown in the diagrams; and with their scientific names. When you feel confident of your powers of recognition, you should draw your own scaled, annotated diagraris from specimens or rep copying the diagrams in this book will be of help to you in remembering these forms correct terms. Ultimately, you should aim to be able to draw label any specimen you are given, after the masse shown in Fig. 3.7. Remember that accurate ob: tion and naming are vital to the identifica’ species and to the correct understanding evolution of fossil structures from one form to It is the fundamental job of the pelaeontolos' which everything else in the science ultmate'y ce pends. 3.4 The nature of the trilo'ite cuticle “The exoskeleton of arthropods is built uP fei’ rather a a motor cris mais up ol. halara pressed stch Ln most arthropods this cus ° Way habe sade of chitin o of chitin strengthened by == orth mineral = “The trilobite cuticle was rather different, HOW'S" aes being bul up of tiny elite crystals arrane! 28 Te earth surface and bound together 2005 sae saL ‘This organic material reais & Y= ot (escribed, It may have been chitin, and er sci seg ‘eee cre ibe cca "minerals Sg ok sin bas ever actualy bret (UNS.g of the y anothe ist, upon ately de of panels y of panels cuticle is «i by some ho ad at right an organic as yet un many test ped chitin’ nd Pleural taro, Meh lutating Tilo “ie Wonly catied by mont Borsa fro groove ep ‘he asa trom the he < a = Wevron tuted t9alacent peur F ] —taterptaurat groove non functions mpyotum Scause of fhe tegen \ ey 5S ) Byala 10 cephalon) Front 3.8. Pygitial morphology. Diagrammatic dorsal views (904 0 * Life cycle and development Although none has ever bew firmly identified, itis sumed that the trilobite began as an egg, from ‘larva eventually hatched, The first larval stage was capable of preservation (and of which. srefore, there is evidence) is known as the protaspis. ‘is tage encompasses all growth up to the develop: ‘of a transverse ridge and furrow which divides ‘Cephalon from the rest of the body. Protaspd Danete 028-1 mm in size stank the second phase called the meraspis, the ‘evr portion of the body developed recognizable reeset one by one. ‘The meraspis ste is “icfore divisible into degrees, beginning with the Upution of the ridge an: furrow (degree 0) ancl ing with the number of the lastformed thoracic ment, As each new segment forme’. the larva had ) U | inatee Macropyoous Apyoktirn of greater area ale) to undergo eedysis. Once the final segment was made, the adult form was complete, and this: maturity is called the holaspis, which means ‘com: plete shield’: Growth continued, however, proceeding by further ecdyses until the death of t The process of development during called ontogeny ‘Many’ changes took pice durin From smallest protaspis (encompassing therefore ontogenetic growth) we 6 400.fold increas Although a SO-fold increase is more common, During larval de clopmient, the gradual formation of the eyes took place, as did theie migration from the cephli margin to their final sites, In some species, the facial Suture began as a proparian type, andl migrated during Ontogeny through a gonatoparian state fo an eventval hoparian conklition, this ontogeny largest holaspis Tm22 Patacontology — An Introduction {and entarged thorecle ppendoe Ia Gill-branch. 7z os Leobranch __ V me Tro ee oreo icncite ae Reeucpescon ne Plewron a —_ > ane hee se Ccophaton, Articul 1 Axiat Vertical Section showing of sot parts (shaded) Larval stages are useful in taxonomy, because f with apparently very different holaspids may be very much alike in their early stages. ‘This can often be taken as indicating a common d&scent 3.6 Classification ‘The question of what makes a satisfactory classifi tion was touched on in Chapter 1, and there have been many attempts to create one for the trilobites, For es imple, one early system used certain heavily weighted Characters (notably, the type of facial suture) to create three Orders. It was assumed that the suture types (Proparian, opisthoparian, gonatoparian) were natural labels which reflected three «i rinet phylog lines. ‘The discovery of larval stages whoxe ontogeny involved the migration of sutures trom one condition {0 another (mentioned above) severely weakene ease for retaining this old scheme As is so often the case in palagonte.ogy, we have to FY to classify together those forms which, quite ‘simply, look the same, ‘This is a practical thing to do, 1 Ventral vi ofthe Onoviciantiobite Triartrus trght) Cephalic woendag on sc worn | | cm Sd it “ lomi Gillbraneh outer rome) Telopod) atin thoracic segments Hee 46, Trilobite limbs and internal anatomy (not 49 sake ‘Anus . me = 10 parts back) an racontructad anatomy € it makes identification easy and memorable Whether these groups based on appearance (phenetic roups) actually reflect natural phylogenetic or enetically related categories is, and will remains 4 matter for debate 7 Mode The classification used he bby the authors of Volume O (Trilobites) ofthe Treurie The mar dn Incertebrate, Paleonivlogy. edited. by the seri lifferent m founder, R. C. Moore, The Treatise is published by een temp the University of Kansas, and while its pronounce: that featu nents lo not quite poses the authority of Holy Wri Pt it the nearest thing that palacontoogits have to 8 UC Bile, Is masive effort aims to cover all lds of the sua subject and has harnessed the services of specialist vretations wshor all ver the work! providing what is best 1300 based on that devised dleseribed ay a tate-oFtie-art consensus of expert? OM ba entioned | The ‘Treatise classification recognises fifteen it ge ent types of suture, uses the features of the label gone and also pays attention to the pyxi:/ium. But, happily i a the Orders are easily recognizable, and Figs. 3.83.12 a set out their basie features, These are also set out for 78h ease of reference in Table 3.2. is aan refore decid‘Darmanites 008 (0. PHACOPIDA) Losow (8) Tritobites 23 Inflated glabeta and tim cheeks ort of ater me” rseor manga Sorte rhe) ye yr Paiperal lobe ara to owt main Preparan suture nut ras hin como Fined cheek arrow and party {reef wbercies moult roglon Gana angle extend into ‘Occpitat fing bear's cena se sean acho Pontrioreaphatie margin xa ‘teaetieghtsng one Somtuiate oade-snaped) ponenorty aida sigity eater nar ‘an eeohaion “2 soppy ble Hex Bygia emer exryirg heat ‘win 14 vole agments | nd races of tubercles Patium caries axa ine ponenoriy Le: DORSAL view jorable. Drown trom relia henetic etic oF Ficsme 3.7. Dolmanies mops Pidiode of Mis For example, those forms with small eyes combined devised with a wide axial region and smooth carapace, whose Treavise | The many forms seen in trilobites probably reflect thoracic segments appear to have fitted very closely, series’ Hfferent modes of life, and palaeortologists have long may have been burrowers, The wide axis possibly shed by [been tempted to make speculative suggestions as to reflects the presence of strong muscles, and the nounce- (What feature: correspond to which tife style. With smoothness of the exterior aided passage through ly Writ, HPerewsing knowledge of not only general skeletal sediment. There would have'been some need for the ive to a BlTUcture ut also of the microstructure of its surface exclusion of grit from the gaps between the segments, is of the APA surface features, it is now at last becoming hence their close fit 5 peciaist. POSSible 10 support of dis, rove many of these inter- Prychocephalus vigilans (Fig. 3.13 si) is a trilobite is best. PFetations which have lain around inconclusively for which is often found in life position, its eephalon long, peeping up above the bedding plane, and the rest is Let us look first of all at the eyes, It has been body buried vertically in the sediment. This was ntioned that the eyes can show various degrees of probably a posture adopted while waiting for is prey {expert 1 differ velo 1 Forms with very large eyes, such as Cyclone, Jabella Me%clopment or suppression. Trilobites which were m apnily, Blind may have been nocturnal in habit, or may have could ot have been Benthic, sine this fe sa 3.4-5.13_ fived in cavities such as occur in reefs. Alternatively, would have buried their enormous ocular suustt jn the sediment, Such a degree of all-round vision 8 difficult feature wo which to ascribe any definite could only have been of use to a. free swine: tt a ce ingle cause. It is byat weigher oiher features They may have swum upside down te cdi, byt wenger Watt other features Tht ng a sharp look-out for prey or predstort: stout for EPY might have been burrowers. The absence of eyes24 Palacontolog, — An Introduction Tams: $2, Trobe Clasifiation ce serumarsut ora teen: pane ve tact Thi woupa arab aor canon any. The ata ERS acticin san anh Otomo Bueryursue Seva, uae Mane Large a rests. BI me ong gn pie conron, =e cvmane, Tomo © apes paropna run sracnien ome ‘ements Sree estar dey Ince. 4 eg Preparian sutures, 19 tora gman ad ‘neiedersioped sovalimnt mechenan ‘ecoot, aman, Ace eral ropa 0 of ky appar pei Ovohon Pra pinescomma 9-10 tore sopra Very ginineie group ot arg ies Onehaporen pata ae he uon nm i w u at ovina, hay ga, ty pra The wide cc xy are, in fact hharpinids hav time. The t ts of cuticle ich look, on as pillars se Ifway down th ich closes off ‘hat was the fe races left by U face suggests t I hole or depi tv ahe hole wi jce these traces sion when mals were aligns ssibly, sinall se ‘long the brim Amal if it was ¢‘Olenoides (0. CORYNEXOCHIDA) o label anteriorly ‘Small eve mpands Eight thoracic fe onte with unvatoned) inal pyaiia! yaidium large-slmost tsppyaous NOTE: Other species in this order may cary occipital and thoracic axial spines Ans aspie (0. PHACOPIDA) Cephalon semicircuar “Sward anterior margin Genat angle rounded — ‘Small, semicircular ysidism (mieropy ous) — Furrowed slabel — Properan suture Twelve thoracic segments fe £8. Olenndes and Amana The wide ccjihalic fringes seen in the trinucleids M harpinids have been a subject for speculation for a hs time. The trinucleid brim is made up of two 'S of cuticle joined by roughly cylindrical tubes Bich look, on the surface, like pits, but which really 1 Pillats separating the two cuticular sheets vay down these cvlinders there is a delicate wall Bich closes off the tube pu for a mi What was the function of this brim? Evidence from {races left by trinucleids resting upon the seuiment mis SUesests that rhe animal used its legs to dig @ H(PE OF depression, and that it then positioned ee en hole with the eephal: augled downwards « these traces appear to point consistently in one Fan Mhe® found together, it is assumed that the nals were aligned towar.’. some prevailing current pibly, small sensory hairs (setae), situated in the plone the brim, acted as ‘weather vanes" to tell the Mal it way correctly aligned. The implication oscopic central from this is that trinucleids were filter feeders. The deep anterior arch of the cephalic fringe may have allowed currents (0 pass under the creature and towards its Jong, filamentous gill branches, ‘The brim of the harpids is quite difterent, It is flat, nd. its irregularly spaced pits perforase the brim ‘completely. This was probably a resting structure, and may even have acted as a “snow-shoe’ to prevent sinking in soft sediment. Alternanvely ‘or addition: ally). it could have been a device to render its owner unpalatable 10 predators. Spines on trilobites are common, especially. in certain orders, and it his been suggested that they act! as baffles to increase water resistance and so help to jicvent sinking. In most cases this seems rather unlikelys the spines would probably have had only a marginal effect upon such relauively large creatures, Perhaps the answer here also lies in unpalatability Spines on the ventral surface probibly acted as supports, but dorsal spines may have to be envisaged in the enrolled position before their protective fun = tion can be fully. appreciated26 Palaeontology Calymone, Pari Am Introduction Occipital ving Inciatinet 13 thoreele Plauron to ta aveta, before ent margin Curved border forming the dou ‘om the ventral margin CGonatoparian suture Eyes small, but prosninent = Cephalon semicircular (Ceanidium large; rion ‘alto moderately broad ‘Lichas (0. LICHIDAD Ti thorscc segments _— Axil segments dstne, ‘ors! furrow deep Longitudinal glabllar furrows eren the oabelirfobes Opisthoparien suture ‘Gonal angle extended into curvad spine ‘snrolled(0, PHACOPIOA) Wenlock (Sil) lating halt-segmants visible segment short xed, with strongly indented pleural 8 viiby fused segments, Pleura fete overlapping due to enrollment 18 composite glaballr body by itolating Pleural termination pinose and unretixed Pysidium pl dn ‘Peronoptis (0. AGNOSTIDA) ‘Tepering late piece bie _Note diminutive sae and hort of anterior ee le ) 2 theracie segments 3 constricted tapering axial ‘of pyaidium tmm es ‘ Fucus 39. Calymene, 3.8 Geolowical history and stratigraphic value ‘The Cambrian trilobite faunas underwent a crisis at the end of that period, so few of the uld stocks made it imto the Ordovician. There was a brief period of experimentation, when short-lived groups attempted to fill the ecologic! gaps left behind by these ‘extinctions, but it was in the late “Fremadoc that the ‘major and ‘typical’ Ordovician groups became estab lished. ‘After this there seems to have be development in the overall design of trilobites, ‘only minor variations on the basic themes stated in the arly Ordovician, Those-which survived that period ‘vere Tong-ranged, and many went on through the cen little further ‘with 5 Lichas ad Peronopss yn, But it was anothet Silurian and into the Devoniat he end of that period ‘major crisis, from the middle to hich dealt the trilobites their mortal blow. A for managed to survive through the Carboniferous, the last seem to have died out in the later Permit Rares of development were high enough it Be Cambro-Ordovician for trilobites to be used 3s 20 Tae Tor correlation, Theit major di-nivantage U8 fossils For crence is not worldwide. iflerent POPU: rae being separated into distinct faunal ‘ovine tions Being “Pyent or make more troublesome business of long-distance correlation Fn being dominantly shel(-dwellins Marcia’ of the relatively shallow-water ‘shelly 15> catble the dovetailing of these sediment with the ==‘Senopeltis (0. ODONTOPLEURIDA) Tem Onvaiccarelie (0, PTYCHCPARIIDA) Cephalon age & semiircu. ssi barge : ‘tobe istnet a Senor expanses //@ Shor ibe furrows ond tine 8 tnoraie segments swt sha sre! Pypidal border Fear 30. (Onnia (0, PTYCHOPARIIOA) spudeshapied, with pitted margin, Cheeks large, convex, lccking eves Focal suture merginal Foreshortened thorax of 6 segments Triangular pyeisium Axis narrow but distin Tem another periods A few ous, Dut ermine yin the as zone Encrinurvt (0. PHACOPIDA) Cephaton triangu Liveigens very restricteg (suture — runs around Base of eye stalk) ge is that Broed axa re, deep, popula: dorsal furrow rovinces’ some i ‘most nepygous PYoidiat pleura strongly 1 ve contrast to thorecte pleura Traces of ax acteristic Large, int Trilobites 27 _ Narrow ocul de {bate gata apni 9 thoracic segments Pleural spines much ‘Small tuberculaie pyoidiumy (strongly micropygous) Large, crescent eves brood free cheeks Suture opithoparian Genal spine ve y short Selmopets and Oxgwvarsa _>- Outer cephalic margin tree of pts ©, 56 roms of pits, deen. its wi 10 margin Giabelia inflated, lacking furrows ‘ending well short of anterior margin Pleural terminations sharp. Pleurat furrows somewhat fant Genal spine over twice body lenath and flexed Pyaiaial segmentation evident. td labels, expanding beyond ‘Much of eranidium tuberculate Eyes perched upon shor’ 0! — Traces of facial suture around short geral 39106 3rd thorecie eg" ie troracic segments narrowing only aightly €0 posterior ain, separated by deen Interpleural furrows ~< Bite agimants convex: marginal tubercles present ~ragering ail region of pawn Nas rove segments than fused leurs 8 visible pygiaia! segments, c2"ving pleural encing to yordium projets posteriorly in spine = Hike structure Ficene 3.11, of Onma and Enernurs28 Palecontology — An Tanradiactum Eyes connected to globe bby oculer dae Enlarged thoraeie segment (enseroploura) ‘Axial spine C1elson") Narrow posterior thoracic textonsion (opisthothorax) Pygidium consists oF sing Bumastue (O. PTYCHOPARIIDA) Opisthoparisn Generaty inated, bulbous Smoo' raptolites occur, Grapto- fe vigorous. sedimentary facies in which the urvive the 0 environments of shel The very provinciality of trilobsic tsunay may be put to use, however (Fig, 3.13 i). It had long been noticed that the trilobites in western Ireland and Scotlind showed more resemblance to those of North Americit, nd Newfoundland than to those found in England, Wales, Massachusetts. New Bruyswick and south-east Ireland The reason for this can be found an plate Tn the Lower Palaeozoic, the present sy North America and dN. Ireland. wd by a huge tonic ‘movements, Adlantig Ocean had not forme Greenlans! were joined 1 Scotland Hut these regions were widely sepa Otenetus Paedeurvae) (0. REDLIGHI:DA) Vat 3.12, Olen Pad Gutela strongly segmented Strong gens! and pleural spines 14 thoracie segments batore pisthothorax coe Tetbie to Vem Glabetta poorly detines Eve narrow, crescent 10 thoracic segments, dorsal furrow oy weak but axis Broad 3 oa and urns ‘ocean from those places we now call England, Wale Ea Massachusetts and New Brunswick. ‘This. lor and ot vi, Holm, vanished ocean has heen called Tapetus 7, Html (On either side of lupetus, separate trilobite fate tert eveloped which were at their most divergent in UM Lanvin. As the ocean began o close, in the same Way on that the Pacific Ocean is closing today, the faunas its. opposing closer together. Tht resemblance increased accordingly 4 2 migration of larval torms bees harrowing sea. This narrossing. process continu Timtil the two faunas were identical, and eventually to the complete closufe of the Seaway the Caledonian Orogeny shores came ne possible across th ‘Se 24 es yas 33Trlobites « 4 Newtoundiand 2S Neate the dichotomy, but trough the Ordovician the owen = ch the Paciie ces” narrowing today ~ by the consumption of ocean oor beneath adocent ALi procen continoes.onetramding 8nd migration became possible and He two Yaunal Prowinces nt Wenty The tetobite Prychacephaluswiilans Vo ts prmwnad ite orantaion, ‘head shied hor ronta ae prota 208 the ea i wh there ‘of the boxy a buried, Ordowieion ot Eg, lowe, Redrwn fr Ager 1963 TPiarot h: Trmacdocian (Ora) 24 meme te ile Sage 1 rote4. Graptolites Fini Hemera Scbpyium Protsordata Clas (rapt ones { ventas a terete a the dak sty sages the Lower lee, yor sind 3 reas ord cae of Sang pope. Thr none coo ras Sine cer chee ner opens Sem ciketeg pace Mowe apc os wi ok Ihe oe of mere churn Had, ‘eh tir Kee merpbepeal dent. They bok ‘Sseetc yet they can be id sate ad (Serie the ec Orson ad Saran Sos Tie meng he mead tk be se Gecclimws w ploy it bong member of he Piyien Hemeberda tae any reed 2 sete mais — sd cd ‘Altiage emacs eg ic sagas Bsc lounges whch Bip snag hao sail! cin Mat oe pom see ty spams pe, Se say ‘Sees eine of mepbeogy tobe dcr. Th ‘lf he pape of sempre ot ‘Sed comet de srs eh tek Pee etme plat proc a te ‘lien ll exept seal cd crt sed evs io eco he Grepieica (L. Ont -L- Dev Riper oes ook place ike tis. Prom the sul eettoacca pe on (Fg 4.1208 fossa large aototec ad he other = Fp stains then grew abead and bene pace a tds, and sce. hen fe vcothes ould tin 40 fecefoating (pom fe ming) a domunate he planton throughout the Ors Sdarian. They ranged fely over very la. ocean aad occur widely i the eepater ‘those periods The alo lived in shallow = tree et prod thare boouse of the « Jad he atuen burtonng sa browsing sch wogeber destroyed most of the sk “The enmense practical us of graptaites = strange fascinate ofan intricate frm, on by a superbly sacienfol group of creates at whose toe Mentity we can a 42 morphology The skcewn of 3 proline is trove stabdosome- Ths may ate bed esichite ons cle pes. ch spe 3 blow tbe slong whi many tml os ine gn ethan wach eth seed cle Sg the tips ef etic Sat cs the tor ined spam po Sina coed ‘The wth ofa dense eraptlie een 1.9 Seam oe st leeph of emt. snd ecending tthe se ech deagher theca. a tr reat led ‘Solon. Ths beiced to have brs some Kd of ‘Rulncetar! acre cod, and a ta fee WC Paces capes in the Hembra “What ten na the fonction othe ashes and ‘bebe? Irv have been ta he pe attiese Cotane a cmale td andthe er mle. Sack refald divison cf arocure wal seal pe st sin oor own secs wd led ea Gimorphinn oe Cn 6. Tl “Snce mos deni as wet (he same BES) sree the, gromen stached tthe 4 bee Se Secome (ecfowing Thee are charac 332 Palotlagy — Am fia io I erciesee, Got ae ty fos ihn one ee four meen cutout erie ate ate ey poly hums _crample the erry tranches of hes may crete ea tetera ticnwetrcuts musrpesnce of cxmlenty. All he seer on ry event eed ey caibty anion thabssone ee of mil haps, A the Ite te thee pomted downwards Pesactuetral deals cused Below. wee fe. i i acs th man rom dnd raps dm pee eee ty tind The benching of silo, abou thre wat roy tre slr fares dodo tan nets, aac siracture ping down, the Cm 1 ee nc dnd pater oftranchng_ Tis never posi seinen The ean sicla wo sar vo that of den Bi it rrr tatencd upon dling. ois Kos onavrge alee | m0 akan, Si Peaeeete faces icesimersml cnet wos cca apc mga cons the Ts» cial, Gap: MRSS th csepiments remains uated in pi: Frojctng spine mas developed, brown he igi Upon the adc ofthe sel there ree an ntl bud fromrafach the fis thea developed his mainuined or ahs opcted, Reguar- cont vith thesculrzoid visable (8:4 Vand a rcercl the numice af branches with lsubssoent tyre Uy the common canal. The sar and iesenal way, his_ shapes ofthese ary enormously 7 SOUP wTOUP ‘kerr rough neal took pl tte (Fig. 4.2, and these esa ase rene Sirs et more suctesful derivations, the apto- sraigrepicllywsetl specs, The wxonony of he ropintes rests pay on ths. party upon the omer on teen ae i Me apes rebtive to the sco (FE. 4.2 i. 422 Gnpoits [AM nonscandent thabosomes have stipes wi Gepideldsa wade in mentees. betwen econ onesie oly. They maycommony cms fracture. TA the plane mde of ie was adopted by more snd nore forms, certain tends 1 the evouton of em twimoloyfbewoptstie see Varian tea metry e Deon er Site = { ey 1 Temioey etd tpn me aon 0h ‘ance orzo wc 5» or our (aederamons my or te ose ro rows ofthese hiner Moura (avs oil 9 1 Mp arong upon unsere sytcce TN + thay be ile pon ation ok ple 1.) rent essed ck ‘Sat But members te Base wside, eer tes fod isn commen ex 5 daerminne pratt Ps Sut whih soup fears aie csnocacheber,s 9a he festhesyiesinatichenth nce mientocrr cnr nner ied ide oss 0 ict mcd toe gently sr Be athe was vane ee Ad om the sige na bo rhe scat bs tee afer miso ploialdscinon ofthe many dnd raed ames fe gwen io Figs. 45-111 Mor the gropiloids in Tale 3.2 or 49.7 nto te ome Gen tts ed fee of he ere tats ene ‘icity i toes toe Oe Tony na anc tea Seep ce and mt eam one mat ne ug ie ae oo ee eV ceca see al cdi boo nc ts ther sfuncs meeah ee They tr meal rea sa Soh tn le el et Tact el meget een te OH Ses Mme ae ce cso POSS SS mee’ Se ee Sethian ken on can oe sea poe tesa eee4 Patmeray’— Ae tettion e eran Lerrocname a ae i A iss § Fs irction of tuner oo 9 Det commen Paluncute forms (hose beating > pele) Poli emerges trough i "The smaller valve isu a the brachial va so my pies Bear om iter ue > ov led racine elo. i ter books yo maya ce the peice ‘alle the “venta” andthe brachial valve “Sor! Should be undertoo that hee words donot 2 espns for secreing em sound the edge of the male te amc ht i eri ee en may case he ll 1 $9 os oe re posteie nd ofthe shel te ta ered em am wy Smart wih HY tema Tanne evo ame Chacha) othe propre hry organ whose oni 8 leone, age the wate nd 8 tng ue nvge a eli a0 ‘rachtumeted 8 sees of cia and + thelr ihe mtlriin within the rv. Tee 175 ete, but th corresponding ‘ont and (eae ad posterior are very convent sshown in Figs. 5.1 and 52 ig 5.3) does oot ins tore eras alls sc ‘ae From teas kc See laments cht i fs suspended. THe f ibe saves and sce below real sures16 Palo An nt sticky mucus This cntaps fl pts which the Acorcand) decisions about 9st major unit sytem hs Be feroung together f pei ant fcr ih snes ad ier ei ss ae rer ht ie ‘etoence to few hs" charcens, Newt sind po ecu in om the wate anf ead er Sain fee peat cpeentnes of ch Tale 52 PS SF ara ae dv io fe onder of ot peal nent Te a th O. Lin fe 0 here ie 3.9 OP nga and he or saesaped th 0 Png ae oral sadestupe. tho gen ineec saves of sabegeal sao Betweencae. 2. ono [trypsin were ost abun ‘isi acopods checoasscmee vewtmas view ME ecsoce. Tics etoee ne as many of our owt Tfarcerizd by soph hii fanestrardiaryadapatiy. The dope a crested Me Te Sharan specs Taratio YOU 2 suophomenite regard ote bs nes uurte piel Sveum BO a girs ich Mes) umes Renin 18 ree vith tee moepogs TF een ihe bachor| ihe Figs 83-816, You Sous oie deta oil nf ato th oven though he Bes Nor esl vee Suse ainda me1 me, na. RFERION 5.0, SHRIFERIOINA) ad 7 pea es jLoctiny-ArborturyDonet_(0.TERESRATULIDA) ay ep aes a Seer peer — Dresrene ee Seen" Cea rot ogee omg wi pr Ft spnstponty ‘ a Sata Fee caenen a eo A Las cent eusenoy te SOLES persia a onan vin ial te min er ym FB nan orem te a enn es cee .... sty erst ‘. Bh estan, Tire ns 2, THF pay an el } sae org iw oe errs nes fn cnc aaa Geers er Zoe a i cea Mapp ve he bates wer2 Palaonsngy — Ar traction ‘rminae in hry baat serene) Peretti ee = ...& vena view \ eng rocks he diversity fndes wat never eesablished, deep harming habit one ino which sb {iu the plum sams to ane been in delne ight ower ly entered ‘tom the en of the Pc Brschopods together wih nb “The reson wal che for ht i the immense used pide nest Pla 2 _sneraentun othe abed ells (Ch Sh whch where papas are rly fund Src presmed hae diplacel tem eclogialy. But Sunn gralte ames ate wll {he cnnot Bethe whale wath Beuse the Hales correspon asemBlapes of hs! ‘Gee their sncess ai ther explnton ofthe tame cue the depres feline | fries og et nee te aS. Me hn eb pnt th Laon (Stead cena estan nth then mathe Se Dean stp fr Ml ye geo onbeoclige Comets Rowe, ch ootMol Phytam Motte 4.1 Iatovetion Thee of atures vey 0 fair wus hs underpin vty, Sich eran these with ‘natu, Wh snp ‘arent ‘ieee fram ach ete tha ne ‘Wet comin Bins ia gs teh stapes wl + host ott nd a al wn vm, whch tthe xo ment Ashpoda M00 living 35,40 ol pies Smet age age of has deep sea ‘maunain tps. Vere are Beers and ee 10 man lengthy att ‘ant Mid white curd at 4m asia. vs Aho en her mars see omaha ogo wal ‘Vet th superbe vary of frm, sie art wade 44 like hae teen deel! vp ‘Musgvint which hough seal here amt thse oman festgniably const tvs Th is eval finan the Phylum tain 62 General motlanean anatomy 1 hs etn we hal ot be eng wh he ‘amy any Aon ain athe esha ‘he run pan lt aro anatomy ih he dn tases saan ot enn yr Yee ~ ive hen eg ay Wl ht ee ‘he Fegan de Sse cy aan os lib 6. luscs ‘teins called tanya (in ths ‘cpa, which ay srs water cy Sin oor the einen women of the nh Soles — 4 cst vps in the cept ad pen ct speed hx Rm be ike ms whch be the Spine repeed na Teaco ara ma pan. have Pree To re fhe amen Oe Toca ster oud cee sesame ot eared bythe eeu came ea emule tae en eed 8 Srey ace ney ve tly as ee fonder Cored + FS mide aes a saline cry ce ae ne he {her ound fr a hr oe mae "aa event Howeer, mole rally sm al skeet S204 eray of atts ‘32, Srmeniion i the repent pte 8 loathe nth the ah eo on rgthsovendage, ec am evotion ly em {oat nt an bese ey ra ai Tas hwah we Rave ready encoumered Unie wnbie tran wetted he men are ey tre ut tha they YEno this he Mat wher who py Motes 1 gastaoroos Onder Arcs (LCamb-Res ass ont Cac “ype Onde ‘Shr sod utecyte whch a rn ah et Sse enc ae feed cae a Times Feshs mane erm ine ow ee di TEC Mpa cages uns er cote, Th ee of ea cen acetates ht et ome of5 carson et owt 0 5 pr he opt Tecrhnme sath teen eet ehh el oes tera ne fas smh sti 1g es Pe OO A he eet orate, fom | cement seat. oan no = fet es ee ay entation, ra ee ergot recy. tw opcneng Saeco a So Srey spar Ne re prc er pt ts ear at ch my oe =e eee ance the wit pas ay hdr im emergencies. Viowever. om - canaries balla a containg Srp of te sa ret ging of 3 ball of perm Jeers ars me nner aypomame Ss scoped io fei the marie } Aisle desing with ving cpap nin mate cory Se presen ane oars etry ace unable to apply 2 ma Oster anaiomicl int Cine in te ie ot cephatopds ace borne sn 1x0 #2 Stent, or they ace exeral he the Mths meitone. As re, external shells ab er omecoma T ie ps eee rr 0) french at FEPNS treat inde Most were proce by on the sell maya rnc with ridges (hee) and a ries themselves nay aso ave Fe 6 wy sh _- as eee a patoreda Order elommida ae ‘J suveiss Noutden Amour he Subelass Coeds isopods, only the. blemies Bae mporant in the fs record ie Bice het sells were covered wih ing att gall consider them in thet the com se gts (Fig. 6.10 i) fret Fa gon fund ithe posirioe portion. + aped object alle the guard sic depresin called the alvetas. Ina ral he end of a delicate chambcred phragmo- ths a very similar to the phragmacone of 3 eld maui but it has a ene ih ir extend out beyond the lve, and wa Thethind and ast por tthe formofa ad. Ie dlcay sch hati teres preserved It called the poe We regarded as an-complcte tsincem. and Hm hr chambe. can tel by the way the .oard appears to Ba Feith no the back ofthe phratmncone that nis mist have extended ight ove the bl in ‘emits, Many diferent shapes of gard are seen ot neaty al of shem have the same, streamlined pe shape, This ells ht ke modern cute fis the direction of motion (a as ring escape rnpcnres) was backwards, guand-fist. The fo To ofthe guard ic dscurced i section 6.17 below Telemnites ca be very easy ond especial whe the rch which contin them ad the fw an be wathed out y the weather or by wave action 416 Ammon Re Anni as Cephalopod Subelss Naioen The hee sutlases with which we are concerned fe cach large grou, comprising many nde, ANd Het the Belemnte compe one order (alee; the ammonites makeup the Orde AM. i ofthe Ammen They ate ¥0 pera ‘tio pablo ste ee 4 Aronies dominaed the Mean certo sings rapidly and becoming ple Aber amon ede. the Sonia underwent sam et les setae nomi the Pero Carbone, hich spe eset they What distinguishes smmontes fom other members oissublss~ the pte or ane ket ammonvid. the anvmonts fended hc ele bans shell td ventions Cr Brotoconeh the orga of the Sp ‘But ammonites in parser had eens ctr: ane suru nes. Lack he cum ofthe pet forthe sepa necks wo pt loans. whe side of he Shel an eee me geo shel 6.17 Made of ite The exvtonmcntal see with which aquatic animals must del come in to eaten. Hyde “dynam elects orerte when Mus (or ‘hoop th fd) Hyde het han. ae doe 10 the sow wth 9S ae all anor wih these leet prope fo ben oe sam ns fe i estes of rum ede oe ime ee fBcaomenon of Rett re hare ers a eh sen ns ae aw ee of ped fo haa LEN pose seat de et a plkbends servoing, Be eee panned ae he ee oe ere see een a Seem Foe change he sb “eyo ofthe Sp a ser Bos. ek yal bodept oe mpl pret ch met a St ‘Seino etal omen compen we he borecy o Nes stoned by the shin rt aka he ters te ere, tng ote twop ote emanate, ‘Sts led te ih ea St op Do yey hor the ban ecw he ge i le et 8 ‘et meme pat permis re hws far he sgl ws tc Sy Keni ome tie seh treo 4 frome ort nage ora Sree ye Mr he cd fem et om he pr th ‘mayors b tem hrc brn tes et te eh Net the ps sot ha ch me ‘ieee ewer a he ea ns wer a th he amt se tol ma eee pt eee i Se pene ato Cres ever ring 00 spec ee comenutiged. N hay mes i oral as techn we to sore ken te ne cnsee res ia kes ee pone Hany fh a ms er cide = ok ms mace he ne Tee van» faba eer fee base a rhs canes Toblerone spenahepiy Mar Ghareaar or oy, Foil chanced poms Orga or re fees bet fore av han cos Seeariss ping nes Cet on tr ord vrices BE Rtas et te ie oonion f eden resin Det br id hy cla Geese) Tk dt inten sey fim eons trea AN Bg laters cnter at as secs sree Sad Wag te One wend eddie adel ro booynty 0 ton oe coe sey Soe st gts ted ir They ted nt ely the psn chr therm ele we have ten te 3) ds peda othe vel deo fhe dmbers Thor ec couche Kn the sch to pp, whe hy ao owl vlog soe the ing a SE abe tht te ttenoe guard wt tech be shen whe sles Sth Bees, her come oho ib pu thei en ie of the parce When savy sts upon a body. the a imitation exried by the art, net abe sre at en ete ‘meg the emir oh yravity, This 6 mh Aromatic middle Consider the be ie maa 6 he Man epreen the effec of sing camera dernis Tal haters rms fing th ene ‘orc dei os ech se ihe wih wo prs hc of aa Tin wre here's seman 7 so ie Hm hermes byecy sd «fet poe ew a venieal powar arrangement (Fit 6.11 stem tome ms Ph pom mt sl si ‘ed rts ae cn rs aeteres yeh Seger : fae eee cic F oan smsare nev ge mg er don fer ex a gt nthe se eh ge chy oi me en ee ond | , Simemele meager, wth! tral Fett ton, wo oper tt 1 ok llereces enh a sexual Teriay han come To nen” oat he cay 6:18 Serast Svorphiom eae ste, etre the The Ammonode:cvolved to the Deena, de we cant bl Sea ete, oct om sy a ae yes afnce Sido lv et dacnnd sb Ts ce men secs. Bat | 5. Sie rere! a bak ee aimee Sirorphines has reveled that PORDING 8 She eeoeee shapes yer because the propcrion of male Heh tn 8 dct hoster from era pl form The ‘emnmonsty 1 one mma rleo chub i i Armmonoids di become pent member Bia th recone Xs Sed pe tae! tie Fees Sire ee pores. 1 ar in te ToS yrvwing armen stare wpe (Fig. 69.9 Fre tigraphic valve the na ra eee ced a fats ba wera eo cremaa In semy-momte 56 MMONDIOEA O-ANAREESTION) Pain fy decay of scavengers, you will notice that they Jus Sonstant Opening force is applied by a struct rata nah mena ‘hugh the gatopeds to ‘the cephalopods, wo Rave snp tal oy ee me Erion My tc Sah oO on te ee io cheea Sateen iar. Eutbgesopmen of predatory ie snes inthe Saab mathe hy (neem ee eter decoed nervous system fr improved and) the concentration S09 ed mouth a econ Hea! (Hen Inc vason he ter hand, he ite ens tetas ht had the opposite effect. filter, feeder Se xs pe, an ie corinai Aah nade rr. The et baa te hea an the al eto tie ome ke the salon Par hve oe mm eloped “ees” all Mane Theses selon = 3 the pulmonate sor soe ue hen they became adased fF Pacem developed 2 visual *ste st hig 6 he rie neem: ai es ples eid The math een from the he moa at cecing A serene an ana pore win es Tee eye ieee ont oil wre ces i fo meme, en, a ae om ine5 —_— c 8 the ene et them Ww exp the ft ‘omen fonts ite Sochin (e 6 Lacan a et rsacmetemmnity#9625 Seal nd bg cy Ara, eo sues Ain ten tty dei ional ution and tus provide the animal with 5 6222 fat Rete nine fons ft Eh he 2 shin of ho ead yf ORY threads made of ead beg ya Rand near the hse a the Mats 99 eet Sepa amos byssate forms th anata rsh commis plane sel heh In much sy ig 620 he mt potions much reduced, andthe Te pts upwards rom the sb, 5 ame even Se reckng of idk ii pcine dere, mang Mo nent by predators sic he seg SEAR The ts emerges fom» fio tinbemerovenal otter mths yaa notch ma bee SP mal ng ct many Dae ne {Shier apes In turner and fee pay quik. Forms whose du or ae fete ne on set Seu characteristic tm adultos Ho 3° < Dagan fovon oun maton, ron Pens es it Ps scape dng <8 my Re xl S08 eevee the mane sry tet TS The eng salon In uthoa ten 81 vate gtr Wate rin sound the sl — ad mom cee at TN Aha conse tng aes 0 ah serena ac ote eae ‘te hae veh develope shel a IRs oe ler tend eee sn ice bose Ca ng joss nts es ered een FE an teciva ch) a ty vite of their Shae, ‘Pe st So neevan teehee ‘tr, ae oe Fen es (Snr Pa ed ter Haue Et) #8 Gee supports wrt ie ie ono woe ict 2 ee sige saga rs he {he fed oie Some see fe (Keser Aorta wey set ae oe. " “Sor tse ee a ee Eee tay sets ca sorte: se aca ae are fee aay rs A. Gest Nmtiry ma eg phe rn er Poce tee ema mucin i ae Btn Mv wo ise ogre on a tm Ser Oh wie se vss, fe Roeser hun Thee ecto fms Ssenly het as oping the |. Coahi, Ad « ee et Oy ange oeahy of he ran He fms ary Meo eae were tr Fe tan mn ame Sth yelp ave Mews ay tear, fg of he cote of te The fe Wien Hes pen w bse 6815 Comme Gorn 7 sconces fae Sh ead poate. ers 3 woe» Tepe wih ty Ze Echinoderms Snes iterate Restore crewmen inte sei Ser tee aa memes toa se nna See ererancc ore fi et ra Sete Ate eer | « ecnmoz0A — c1A88 EOmODEA eS Sa! ‘its oT ste ta IN set loti Serer um, fei cet Pada om sd fu oa tle te oo cae Be a the een a eso ox test, 08 1 called — Fi. speaking oo SNE gr the ONS aa sae conti jee the ee cpr eee chord tenis szungsy vown'd that of aed aborat 1 a cd ih macs amd sowed 30 he msesso Pomme — Me HN ay . to eh ro a rea ta pe it ome» Bet em mah se ner te te es, WA yi Sir ee At of apis Tec nme he we ru imadrepoie Se te ose roe ise pv ring hes ing of cal le cch v8 cc pre Ths frei ‘ets her crs, nd Seal be hc ot mayb die in ea ames evening eer he ap i nde eh, eee emacs ae fier ake, the tet rie save, minute pcr sometimes ‘pein glands Fin 7.3 Bahamian ira are compose fo 088 Abul nc ear pore pais nse her ose S85 exh pir mie ens to a snl tbe foot iow. Bath abs nitrate 10 the sis ad nar apn on he oral sure, whee ty mt th peistome, This sa ge ole Mae the oh us the complen in apart nana rl’ lantern oc = : loam cet the een the peste at sal shee repo ils prota rm he es Fei the te ft tn as tine eo he gut extends upwards tout he ‘ty cv towards the ans (FB. 72 Soe the anus the madeponite piste T eed rns. cae she stone caal a ccumroal carl Ts inks ph aay chibe radial vests extend, cash one ps8 UP HM fan amulorumm an terminating 1 3 Sn ibe whieh protrudes throu she sine he ela pae aS hese rail vessels, paved tubes branch Hach Teas the hase of a ube fot muscular scr aml. TO 1S rt sore pig cade The *plseed do ove he tet ihe cas fro "ogee pi CONC oe np ct weno ened. The herrea a mich 10 cing to ie agen : Graco the ample fr ta 2 ‘heanimal i rape sng, can any hundred sch tig gn SOL ‘ 73 Inept ecicite Jn ieguar chins the symm ad es on x ‘ater than eur outa (Fy Soret Mater en ig. 75 i, But thee ae many other it 10) binealy vhs stoped, ® 75 We ene fo the be empl we and, insead of scing ol apeareh, the tno ang iin he pc se a dae een tone i. 75) rer paar ee rlcinga mich ig em eal oy rej cnte he ot Sac a 73 “ne aber se pena uae , Denied eee Sg NMHC ra, eater) Humps of hee mrrones | Sromyagae aoauenees, wat fe Sa fen xk pointe em LS) Adenine “Cana, Rt, 2 Temas cas Fe anne ees i at ati sy an w te | Recon open amy eng oe Ri eran sc Sag Mn Pie root | a RS cr oak TE Se rs ermal etek haps he mon nny meets’ | Sa Si loner gre le lat od or come Oar tee Bnet Srecture resembling opder® web 8 SHB SLY eden. He shown in Fig. 82.mas. nucnenas eins #0" oscar ca" De saseme stents eet ws ae aerate ars foarvens we | oyaaanvnnran tami ao " sewer amen ican 1 Bi rere acute E s con wi en actin tesa scene ote miosis ia ar Coat ee _— ts deen oferta soto on hn soca (not ype) Caemtay aren in Wie we 8 we smpimanm =" LonarTupinal section me nent cot tacnion pasos SS Te arse ar Serine ta grows vo ural Meroe cries ce 4 Coens present when they ae posily me tees ran Secemmrarisniter acinar he eee serena Me cme: Sage props eT : Se yp maybe ae up 8mm (Te spectacular de hee oe leractnian polyp ovechangs its calYs ‘ne snl ewig td Eat oak os ° ecm csey 1 the subsrae 204 @ Could Sram yy ert srvconie [er antnow _ ie mm on tg ot ihe min here Sot ent — Tay Tee a to be horn-shaped. and IM IMTS Gas teem tee Sg These = rai coral Dio, for wastance (FE A ques MCT re ly Flees 2 tere ton n meron tt mgt 8 Tying in valley between comme ese manu m Tenge, Sc cnc 6 ite etm Oi srcnians owe tir sucess poy ©ste, Thy aot, enone. of och srt phic we, bt ty can Pad ani fe ‘ery mal als AMtoorh semsupnis are probly more Sect vd opens tan ca thy ae ‘ected here heme yd wf ond on ‘etka wre cin Pca Te elo fies of th perplexing. exinctsrovp Sach any ten ced ee ook ‘Noein rene, Ma sh 4,8 Prion Stomnaprie, rt we noe, [lb scaned, andthe fon ferme pooper Cow. of the Phykom Pores ihe ‘Pree We tal ave Wo wits Smogon ‘ain fom (SES, and dome scenes mee, cena epee tat the ay oa opchnaeg Seema Wika dee ive snd ofen sommes sn teres he pascy of cord growth and he dna ‘Sioa fas to whch they fll ae sng ito of sch ecg zonation nfs we eeeel te ura cxhi'g wd chalcgig pre feo puaccony, in sn area wher hs ae Bake imprest note aid dpe of ‘Bhimenogy, An tnderanding of oral clogs peru the merci ofthe cron tdihe gat reser pti of rfl od ap fede oimry to nance much research ints soni eid 1. stnomaTopoROS Phylum Stomatoproies (Camb -Cret.) layers ar Koon as latilaminae snd rocord growth Petia ke the rings ofa tree. But each ofthe ‘conic of very fine horizontal and veri prem In some, the horizomal‘lements are like tbub. sold calcareous Noor, separated by pillars to [ar Balle. The whole siructire resembles 3 mx opie muli-storey ca-park. In others, the hoz fomponcnts are alo bars, so that the coenoste Iooks more tke 2 mase of safflding On the llaminr surface, small hammocks ele mamelons may be seen, end raditing from th Ssimmits one sometimes observes caliting #9 (lod astorkinse, The function of thee. and nde the hole ature ofthe soft pars of somone 18 mater for conjure, ‘Mesowkcsomatopooide were not generally ft Trlr, but Palscazsic representatives were oP ther dominant component. Dificulesn Neale ham. however, have rendered their pastas Saphic we ext to impasse Sstitaton Y Seen Gers een tae Tee cen age aaSevens T= con i= Yas = Sel ono nenon fmt Bw we wm PO Do aden, ae noe Brckshang wes Wo acme toe a ta tae Ae nd te 9. Vertebrates eng Ths fur is called a notochord, and by lorie bony or cartilaginous, as well as ty fain 0 2 to envelop and protect the nerve cod, uted into hs jointed spinal or vertebral columa tenin the vertebrates (Subphylum. Vertebrata — {ike91), Only they, of the Pivlum Chordata, havea fal cord, and an exceedingly complex one iis Fay the internal skeletons of vertebrates may be gious rather than bony, and thus are ikely btepreered. Also, dsariculation ofthe bons after fais very common, making complete finds very fe. Nevertbles, the excitement of findins the snes vererate relic (a shark's tooth, some fi sks ora replian vertebra) is easily matched by te fone fascination of an evolutionary story which imsly, leads directly to ov:selves Todo justice 1o the Subphylum Vertebrata would ‘Site a chapicr as Tong 2° the rest of this book: ‘ae drs not make her units in nat, comparable Prdazes So, to iry and make sense ofthe sory, We Pn sity Sperion “omer 91. A Cafe of eV shal concen at on the ve ite og mach teed to afl fom ie fla indent een 92 The vertebrate pla ree vn fe oa See tee rs iin aes ten See aS oe ee aii ee Shall sce the congestalanf ake newest rc alterations in. soft-part anatomy, phsslogy smd sent ee ae oa soho em ‘we must rar w ving representatives ofthese ance pee7 Geog and lic vide aoetpl lene son sg sare ey verb at He a aoe soit. they prea pal es ko devo Imre wily ere nth aie dime Devan pn wich hn oe | 94 The origin of jaws and lateral fi skeletons, forward propulsion 8 Vide y the ona fe To Aer. To assume pedatry oaks on oped sae Toeidenaly. stan heya hae and 8 Tring. naturally bao Bi we oe wt Veet i eres “A ca the amcor glace wt mero it heb mis "ost when arch forme part he 1 Abn wraton he hey sme Cen ae er) The second vanished when the atees Supporting it rotted to Become the uppet a Tower fawbones (Fig. 9.34), Th hie sit disappeared when "earch moved close othe jaw in ode 5 The origin of the jo hy il arch main felch ny jaws and al fing, th fh as stacted on what tare the andar verebate len. Bat ts Bucy was established sah ‘ay. Stange, ‘experimental aes oud near 2 placoderma) ae av found, which difer ate {une “Exams sone te ate Sehing mand had masive hens the conn ie 9.5 The oriin of hone and anes yw ibe (aut fe) impression that the changes Shon cured wok pe presi an {Saroag to astme tha the Ges vere wee Caringnous and mee radu inproved by the trent af tone The nn ff neers Tan hick Race ser hin be elton patito ate, had Bony ed let {ioc team an evcuonary pint of sew, «re frumi.fortan te cargoes star Soe (Palen beny fishes (Ones) say 2Furies: ay from the U Instat the wld pase These forms (bic nce mamma from Aardvaks te Zebra, taking in clephans, rodents, -carmores, tors whsis, monkeys and sels) hint 2 very pro: Toned gestation made possible bythe placenta This i frmed from one ofthe old egg membranes. the Slisois (Fig. 95 ti), which hes become sacha ‘he wall ofthe wer od permit the blood systems of mother and cid to cxchange oxygen, carbon die {nd food without actully txng. (Mixing of eure frond be disstrus, The mothe’ imme sn Soekl rec ie had lca ages fat vemplant pants will ssc thei done organs nou special drugs 10" suppres tht” satura munity. Moreover, she rcther's bod group sed that of her child may not be the sme ke shrews and eds, and thes fet

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