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PlantEvolution • Majoreventsintheevolutionoflandplants – ThePeriodwasatimeofrapidevolutionforthelandplants

– the appearance ofleaves

– and emergenceof seeds PaleozoicFauna Evolutionary Faunas

The Early was a time of adaptive radiation of many faunal groups, following the mass of and nautiloids at end of . Increase in diversity from 150 families -> 400 families

•The Paleozoic fauna (or fauna ): articulate ,stonyand lacy bryozoans, stromatoporoids,cephalopods,crinoids and blastoids, ,graptolites ImportantGroupsofPaleozoic Invertebrates • Porifera – – Corals (Rugosa andTabulata) • – Moss • Brachiopoda – Lampshells (Articulata and Inarticulata) • Arthropoda – Trilobites,Crustaceans,Insects • – Snails,Bivalves,Cephalopods • – CrinoidsandBlastoids

EARLY PALEOZOIC LIFE Unicellular Organisms (Protistans) Foraminifera First appeared in the Cambrian Survive to present

Platysolenites, a Cambrian foram Agglutinated form Calcareous skeleton Nanicella – middle tolate Devonian

Late Frasnian – Eogeinitzina, Eonodosaria Late Famennian - Quasiendothyra EARLY PALEOZOIC LIFE Unicellular Organisms (Protistans)

Radiolaria

First appeared in the Cambrian, more abundant in mid-Paleozoic Survive to present Have a siliceous skeleton

InPaleozoic only Nasselaria. Rockforming roleinthe Devonian – radiolarites.E.g.Ponikev Formation Trilobites

Still abundant and stratigraphically important.Second and last primeinearly Devonian,since middle Devonian onretreat Uralichas

Ordovician

Selenopeltis province Perigondwama

Selenopeltis Ordovician

Asaphus Asaphus province Baltica Aulacopleura konincki, Silur, Phacops DEVONIAN

Reedops . Odontochile EARLY PALEOZOIC LIFE Metazoan Invertebrates Arthropoda Ostracodes

Look like beans Segmented body enclosed

in CaCO 3 and chitin carapace Marine and freshwater Useful in biostratigraphy Early Cambrian to Recent Ostracodes

Since Ordovician diversification of ostracodes • Eoleperditia fabulites Conrad • MiddleOrdovician, RutherfordCo.,Tennessee • Shellsarebivalved,small (1to10mm)andoval • Recrystallization Arthropoda

Subphyla Trilobita Trilobites (extinct) UCMP - Trilobita Subphyla Crustacea Shrimp , lobsters , crabs , barnacles , cladocerans, ostracoids, crayfish , water fleas, and copepods Characteristics Branched antennae Mandibles (chewing mouth parts) Encarta Online - Crustacea - korýši Subphyla Chelicerata - klepítkatci Spiders , scorpions , ticks , mites , sea spiders, and horseshoe crabs Characteristics Lack antennae Chelicerae (pincerlike mouth parts) UCMP - Chelicerata Encarta Online - Chelicerata Subphyla Uniramia Insects , centipedes , and millipedes Characteristics Antennae Mandibles Unbranched appendages Branchiopoda (lupenonožci) jeskupinaprimárně sladkovodních korýšů devon

First Decapoda (shrimps) inthe late Devonian Merostromata EARLY PALEOZOIC LIFE Metazoan Invertebrates Arthropoda Eurypterids Swimming or crawling Some up to 3 m in length Ordovician to Mostly and Devonian

Arachnids

Silurian,Devonian

Gigantocharinus Insects – Apterygota (Colembola Chvostoskoci )

Insect evolution: Six legs good Primitive insect-like creatures called springtails were among the earliest known animals tocolonize the land, early inthe Devonian periodalmost 400 million years ago .

New light shed on the oldest insect MICHAEL S. ENGEL & DAVID A. GRIMALDI Nature 427, 627–630 (2004); doi:10.1038/nature02291 Incontrast,fossils of the earliest known true insects areknown from later oninthe Devonian period.However,reinterpretation of afragmentary insect fossil from the important early Devonian Rhynie cherts of Scotland shows that the enigmatic Rhyniognatha hirsti was notonly a true insect , but relatively derived — that is it hadbeen around long enough tohave accumulated some uniquely insectlike features.Although only the mandibles arepreserved,it is possible that they once belonged toawinged insect.Inany case,the fossil shows that the origin of insects was muchearlier than previously thought. The discovery suggests that insects almost certainly evolved in the Silurian Period , some 438-408 million years ago. Brachiopods

Diversification of articulates Ordovician-There was an enormous radiation of brachiopods. Max bar width equals 200 Brachiopod genera abundance through geologic time ARTICULATA

ORTHIDA- Lower CambriantoUpper Permian STROPHOMENA- Lower Ordovician toLower PENTEMERIDA- Middle CambriantoUpper Devonian RHYNCHONELLIDA- Middle Ordovician toRecent SPIRIFERIDA- Middle Ordovician toJurassic TEREBRATULIDA Upper Silurain toRecent Brachiopods Strophomena Stringocephalus Gastropoda

Gastropods they appear inthe Cambrianbut arenotabundant until the Late Paleozoic.

Platyceras BIVALVES

Cardiola

Panenka Tentaculites CLASS: Cephalopoda- This class of the order Molluska is divided up into 7subclasses SUBCLASSES: •NAUTILOIDEA -Thesewere the first cephalopods toappear inthe fossil record. They appeared inthe Late Cambrianand quickly expanded. The only surviving Nautiloidea aremembers of the Nautilus (picture inheading of page). The members of the subclass Nautiloidea have orthoconic or coiled shells. Manyof the straight Nautiloidea secreted deposits intheir older chambers tomake their shells neutrally bouyant. Formoreinformation onthis subclass see this page . •ENDOCERATOIDEA- The mostnoticeable thing about this subclass is the enormous size of its members. The largest Paleozoic fossils,they hadorthoconic shells that could reach up to9m. They hada"short"stratigraphic range,entering the fossil record inthe Ordovician and leaving inthe Silurian. •ORTHOCERATOIDEA - The members of this subclass areknown forthe particular ornamentation ontheir shells. Their orthoconic or cyrtocone (slightly curved)shell have rings,grooves,and/or color. Secondary deposits inthe shell arewell developed. They entered the fossil record inthe Ordovician and left inthe . They arethe probable ancestor toother cephalopoda. •ACTINOCERATOIDEA- They hadlarge straight shells with ablunt end (apex). The siphuncle hasinflated connective rings.They entered the fossil record inthe Ordovician and left inthe Carboniferous . •BACTRITOIDEA- They have shells that areeither straight or slightly curved. Members of this subclass resemble members of the ammonoidea subclass inthat they have abulbous protoconch and asmall marginal siphuncle. They entered the fossil record inthe Devonian and left inthe . •AMMONOIDEA- Their shells aretightly spiraled with complex sutures. Like the bactritoids,they have have a bulbous protoconch and amarginal siphuncle. They entered the fossil record inthe Devonian and left inthe at the famous K/Tboundary. •COLEOIDEA- The common Cephalopoda of today,which include squids,octopuses,and cuttlefish. They have either internal shells or noshell at all. Coleoidea differ from other cephalopods byhaving only one pair of gills and one pair of nephridia (kidneys);all other cephalopods have 2pairs of both gills and nephridia. Coleoidea entered the fossil record during the Devonian and arestill around today.

Nautiloids Lituites littuus ,an odd nautiloid fossil from the Ordovician of China. Reconstruction of the Silurian sea

Ammonites

First occurrence inPragian

Clymenia

The goniatite Goldringia is at center.Behind,the straightshelled cephalopod Michelinoceras is can be seen.At frontleft,the Phacops is moving near aclusterof Paraspirifer brachiopods Coleoidea

Devonian Eoteuthis

Alloteuthis Ordovician reefs FirstTrueReefs The Bryozoa is adiverseone with approximately 4000living known,and almost four times that amount arefound inthe fossil record (Levin,1999). These"moss animals"areoften mistaken forcorals due totheir structural similarities but bryozoans have amuchmorecomplex anatomy. Bryozoans arealso mistaken forplants,hencethe term'moss '. Bryozoans (sometimes referred toas and Ectoprocta)aremicroscopic sea animals that live in colonial structures that aremuchlarger than the individual animal.Because thesestructures areusually composed of secreted calcite ,they commonly form fossils. Bryozoa Moss Animals, Sea Mats,

Stenolaemata --- Cyclostomata Fenestrata Cryptostomata Cystoporata Trepostomata

Gymnolaemata - Ctenostomata Cheilostomata Phylactolaemata Bryozoan Classification Class Stenolaemata

Marinebryozoanswithtubularzooidswithcalcifiedwalls.Lophophore is protrudedbyactionofannularmuscles.Includesfivesubgroups: • Trepostomata: Coloniesgenerallyrobust;dendroid,encrusting,ormassive. (Ordovician Triassic) • Cystoporata: Coloniesrobustordelicate.(Ordovician Triassic) • Cryptostomata: Coloniestypicallydelicate;foliateordendroid. (Ordovician Permian) • Fenestrata: Coloniestypicallydelicate;reticulate(netlike)orpinnate. (Ordovician Triassic) • Tubuliporata ,or Cyclostomata (Ordovician Recent) Bryozoa – OrderFenestrata

• Fenestella althaea Hall • Flat colonies of “lacy” bryozoa • EarlyDevonian, AlbanyCo.,New York • Silicareplacement

Porifera FamilyTree Stromatoporoids

Sponge-like, grew in sheet-like calcareous layers. Dominant reef builders, Ordovician Period through the Devonian Period, a period of about 100 million years. Stromatoporoids were marine colonial forms with a calcareous skeleton. They were important contributors to reef building during the Silurian and Devonian. Their relationship to other creatures is uncertain but they show some affinities with Porifera. They consist of calcareous layers which, when weathered, show a charecteristic contour line pattern as seen in the specimen above. Amphipora floatstone in peloidal line mud matrix; note common stylolites" Upper Devonian Leduc Formation Alberta Calcareous sponges

Geologic range: Ordovician to Present

Built reefs in the Silurian (Michigan Basin)

Found in Caribbean Sea, Mediterranean Sea and Pacific Ocean EARLY PALEOZOIC LIFE Metazoan Invertebrates Phylum Porifera (sponges) SPICULES

Spicules –

Composed of

Calcium carbonate EARLY PALEOZOIC LIFE Metazoan Invertebrates Phylum Porifera (sponges) SPICULES

Spicules –

Composed of

Silica EARLY PALEOZOIC LIFE Metazoan Invertebrates Phylum Porifera (sponges) Caryospongia Silurian

Astylospongia Cnidaria – Rugosa (solitary)

• Rugosa areanextinct groupofcoralsthatwere abundantin Middle Ordovician to Late Permian • Solitaryrugosans areoften referredtoas"horn corals" • Rugosa canalsobe colonial • extinctattheendofthe Permian,about245 millionyearsago Cnidaria – Rugosa (solitary)

• Cystiphyllum conifollis • SolitaryRugose coral • MiddleDevonian, Ontario,Canada • Recrystallization

Cnidaria – Rugosa (colonial)

• Arachnophyllum pentagonum Goldfuss • Colonialrugose coral • MiddleSilurian, Kentucky • Silicareplacement PetoskeyStones– recrystallized colonialDevonianrugose corals EARLY PALEOZOIC LIFE Metazoan Invertebrates Tabulata (Tabulate Corals)

Ranged from Ordovician to Permian

Major reef formers, Silurian and Devonian reefs

Always colonial EARLY PALEOZOIC LIFE Metazoan Invertebrates Tabulata (Tabulate Corals)

Favosites EARLY PALEOZOIC LIFE Metazoan Invertebrates Tabulata (Tabulate Corals)

Halysites Phylum Echinodermata

Subphylum Blastozoa ...... Class Eocrinoidea (Cambrian Silurian ,3032genera) ...... Class Parablastoidea (Ordovician,3genera) ...... Class Rhombifera =Cystoidea inpart( Ordovician Devonian ,60genera) ...... Class Diploporita =Cystoidea inpart( Ordovician Devonian ,42genera) ...... Class Blastoidea (Silurian Permian ,95genera)

Subphylum Crinozoa ...... Class Crinoidea sea lilies (Cambrian ? Early Ordovician Recent,1005genera) ...... Class Paracrinoidea (Ordovician Silurian ,1315genera)

Subphylum Echinozoa ...... Class Echinoidea (Sea Urchins)( Ordovician Recent,765genera) ...... Class Holothuroidea (Sea Cucumbers)( Ordovician Recent,200genera) ...... Class Edrioasteroidea (Early Cambrian Carboniferous,35genera) ...... Class Edrioblastoidea (Ordovician ,1genus) ...... Class Helicoplacoidea (Cambrian ,3genera) ...... Class Cyclocystoidea (Ordovician Devonian ,8genera)

Subphylum Asterozoa (= Stelleroidea) ...... Class Asteroidea starfish (Early Ordovician Recent,430genera) ...... Class Ophiuroidea Brittle Stars (Ordovician Recent,325genera)

Subphylum Homalozoa ...... Class Stylophora (Cambrian Devonian ,32genera) ...... Class Homoiostelea (Cambrian Devonian ,1213genera) ...... Class Homostelea (Cambrian ,3genera) ...... Class Ctenocystoidea (Cambrian ,2genera)

EARLY PALEOZOIC LIFE Metazoan Invertebrates Echinodermata Crinoidea Crinoids Middle Cambrian to Recent

Scyphocrinites EARLY PALEOZOIC LIFE Metazoan Invertebrates Echinodermata Blastoidea Blastoids Early Cambrian to Permian

Echinodermata Asterozoa

• Devonaster eucharie (Hall • Middle Devonian, Ulster Co., New York • External Mold in shale

Graptolites

Graptolites range from the middle Cambrian to the Carboniferous. Dendroidea are found across this entire span while Graptoloida are found from the Ordovician until the earlyDevonian. Graptolites are most commonly found in deep water, dysoxic facies (black shales), but do extend into shallow facies. Because they did not biomineralize an easily preservable skeleton they are nearly always carbonized. The process of carbonization combined with the highly compressible nature of shales made most graptolite fossils extremely flat and therefore difficult to study. Graptoloidea EARLY PALEOZOIC LIFE Metazoan Invertebrates Graptolithina (Graptolites) Marine Planktonic or epiplanktonic Sessile benthonic Chitin skeleton Cambrian to Mississippian?

Good index fossil for Ord-Sil EARLY PALEOZOIC LIFE Metazoan Invertebrates Graptolithina (Graptolites) • Silurian graptolite shales Climacograptus Didymograptus from Victoria,Australia (Lower Ordovician) Graptolites range from the middle Cambriantothe Carboniferous. Dendroidea arefound across this entire span while Graptoloida arefound from the Ordovician until the early Silurian. Graptolites aremost commonly found indeep water,dysoxic facies(black shales),but doextend into shallow facies. Because they did notbiomineralize an easily preservable skeletonthey arenearly always carbonized. The dendroid graptolite Rhabdinopora ,lower Ordovician.Did it float suspended from a"bubble",like aPortuguese Manowar,or asepiplankton,attached toseaweed?Acolony like this could become at least afoot long.

A. Portions of dendroid graptolite colonies preserved as carbonized films on the surface of black shale of Early Paleozoic age Dendroid graptolites were probably benthic organisms (approximately life size). B. Fragments of graptoloid graptolite colonies (stipes ) preserved as carbonized films on the surface of black shale of Early Paleozoic age. Graptoloids were probably planktic organisms and may have attached their colonies to other floating objects as illustrated in morphology figure above (approximately life size). KINGDOM: ANIMALIA PHYLUM: CHORDATA

SUB-PHYLA: UROCHORDATA (sea squirts) HEMICHORDATA (pterobranchs, graptolites *) CAMB. CEPHALOCHORDATA (lancets) CAMB.

CRANIATA () CAMB.

CLASSES: CONODONTA * CAMB.

AGNATHA (jawless fish) CAMB. ACANTHODI (spiny sharks)* SIL. PLACODERMI (armored fish)* SIL. CHONDRICHTHYES (cartilaginous sharks) DEV OSTEICHTHYES (bony fish) SIL.

AMPHIBIA (amphibians) DEV. REPTILIA (reptiles) CARB. AVES (birds) JURASSIC MAMMALIA (mammals) TRIASSIC

SubphylumVertebrata

• SubphylumVertebratahasseveraldivisionsyouneedtobe familiarwith : – Superclass Agnatha – JawlessFish;Lampry Eel; Ostracoderm (fossil) – Superclass Gnathostomata • ClassPlacodermi – FirstJawedFish(Fossils) • ClassChondricthyes – Cartilagenous Fish;Sharks;Rays • ClassOsteicthyes – SubclassActinopterygii – RayfinnedFish;Goldfish;SeaHorse – SubclassSarcopterygii – LobefinnedFish;Coelocanth

GeologicRangesof MajorFishGroups EARLY PALEOZOIC LIFE Vertebrates Fish EARLY PALEOZOIC LIFE Vertebrates Conodonts Resemble teeth Proterozoic to Triassic Calcium phosphate CONODONTA

EARLY PALEOZOIC LIFE Vertebrates Conodonts Pre-1964 examples

Ordovician-29-39 Silurian-25-27 Devonian-16,17,21-24,28 Mississippian-4,8-15,18-20 Pennsylvanian-1-3,5-7 EARLY PALEOZOIC LIFE

Vertebrates Cambrian to Recent Fish Agnatha (Agnathids) Jawless fish

EARLY PALEOZOIC LIFE Vertebrates Fish Agnatha (Agnathids) Jawless fish

Astrapis Astraspis DevonianJawlessFish

~20cm

AgnathaAgnatha

EARLY PALEOZOIC LIFE Vertebrates Fish Placodermi (Placoderms) Plate-skinned fishes Late Silurian to Permian

Bothryolepis

DEVONIAN

CAMBRIAN ORDOVICIAN SILURIAN DEVONIAN CARBONIFEROUS PERMIAN EARLY PALEOZOIC LIFE Vertebrates Fish Placodermi (Placoderms)

Dunkleosteus ACANTHODI EARLY PALEOZOIC LIFE Vertebrates Fish Acanthodii (Acanthodians) Early jawed fish Late Silurian to Permian DevonianSeafloor

acanthodian rayfinnedfish (Parexus ) (Cheirolepis )

placoderm (Bothriolepis ) ostracoderm (Hemicyclaspis )

Acanthodii Chondrichtyes

Ctenacanthus sp, a Late Devonian and Carboniferous shark

The very earliest signs of sharks areminute fossil scales and teeth which arefound inrocks from the late Silurian to early Devonian period{around 400million years ago).It becomes moreand more difficult,however,toidentify shark scales inolder rocks because they closely resemble those from jawless fishes called the lodonts,which lived at the same time.Only microscopic differences separate shark and the lodont scales,and the two kinds seem tobecome moreand morealike the further one goes back.

Cladoselache (top left, Middle Devonian), Ischyodus (top right, Upper Jurassic) and Hybodus (bottom, Lower Jurassic) © •Bony fish (Osteichtyes) •There aretwo groups of bonyfish 1.Ray-finned fish (Actinopterygii) began their evolution inDevonian lakes and streams (freshwater)and then spread tothe sea.They arethe dominant fishes of the modern world . 2.Lobe-finned fish Lobefinned fish have muscular fins with articulating bones.There aretwo groups of lobefinned fish. a.The lungfish Lungfish live today infreshwater. b.The crossopterygians This is an important group of lobefinned fish because it gave rise to the amphibians during the Devonian. RayFinnedandLobeFinnedFish

• Arrangementoffinbones for

(a)arayfinnedfish

(b)alobefinnedfish – musclesextendinto thefinallowing greaterflexibility Actinopterygii

Chondrostei Dipnoi Neoceratudos forsteri

Protopterus ssp

Lepidosiren paradoxa Crossopterygii

Eusthenopteron Latimeria chalumnae

Ichthyostega CrawlingoutoftheWaterRhynochodipterus

Eustheonopteron

A mass extinction occurswhenalarge fractionofalllivingspeciesbecomes rapidlyextinct. Thefossilrecordshowsthatatleastfive majormasshaveoccurredinthe past500millionyears. ImpactsofasteroidsonEartharesuspected asaprimarycauseofmassextinction. ThreeofthefivemajormassextinctionsoccurredduringthePaleozoicera:

– Attheendofthe Ordovician period, – duringthelate Devonian period,and – attheendofthe Permian period.

The graph shows when the five extinctions occurred. As you can see, the Permian extinction was the most severe. EARLY PALEOZOIC LIFE Mass Extinctions Ordovician Mass Extinction Second most devastating in earth history Caused by glaciation and associated lowering of sea level Thisextinctionoccurred Since more water atendoftheperiod, was in ice form, the about440450mya. sea level lowered all over the world, Itisthoughttobecaused causing a reduction byaglobalcooling, of space for life on whichcausedthe continental shelves. continentGondwana to glaciate.Geologists The most havefoundglacial affected animal depositsintheSaharan group was the desert,whichprovided marine invertebrates , theevidenceforthis in which more than theory. 100 families were wiped out.

Next,interms of severity,was probably the Ashgillian (latest Ordovician)event when perhaps 50% of marine species went extinct. The Ordovician extinction occurred at the end of the Ordovician period,about 440450 million years ago.This extinction,cited asthe second mostdevastating extinction tomarine communities inearth history,caused the disappearance of one third of all brachiopod and bryozoan families,aswell asnumerous groups of conodonts , trilobites ,and graptolites . Muchof the reefbuilding faunawas also decimated.Intotal,morethan one hundred families of marine invertebrates perished inthis extinction. Echinoderms, trilobites, nautiloids and manyother groups suffered significant losses,. • Thiseventistheorizedtobecaused byanepisodeof global cooling similartothatoftheOrdovician extinction. • Thistime,the glacial deposits have beenfoundinnorthernBrazil. • However, meteorite impacts have alsobeenthoughtapossiblecauseof thismassextinction,althoughthe evidenceremainsinconclusive. • Thisextinctionhadlittleeffecton landanimals,mostlyaffecting(again) the ,inparticular,the reef-builders . InFamennian strata only about 15percentof Frasnian brachiopod genera arefound. Ammonoids trilobites, and conodonts experienced asimilar decline,and manytypes of gastropods and trilobites disappeared aswell.The reef community became forever changed.After the Famennian, tabulate corals, stromatoporoids ,and rugose corals arerare.Thesehadachieved their greatest faunal diversity during Middle Devonian time and hadbeen important reef contributors for120million years.The time of extinctions was also when acritarchs (the only phytoplankton with an extensive Devonian fossil record), became rare. Placoderms almost disappear at this time.Until then,during the Devonian, they hadbeen the dominantpelagic carnivores.Onthe land,life appears tohave been little affected,although the marine record is that world climate may have cooled significantly at this time .The evidenceforthis (inNew YorkState)is that glass sponges,which today are restricted tocool waters,began tothrive where formerly successful tropical marine faunahad become extinct. Evidencesupporting the Devonian mass extinction suggests that warm water marine species were the mostseverely affected inthis extinction event.This evidencehaslead many paleontologists toattribute the Devonian extinction toan episode of global cooling,similar to the event which is thought tohave causethe late Ordovician mass extinction. DevonianTimeline

Acritarchs from the lower Paleozoic Upper Ordovician

Upper Ordovician Upper Ordovician-SEM photo PLANTANCESTRY

• Plants represent a monophyletic group – Evolved from a common ancestor – Who was this common ancestor? • Multiple lines of evidence indicate that plants evolved from a group of green algae termed Charophytes – What is this evidence? Figure 29.1 What Is a Plant? MAJORPLANTGROUPS

• Fourmajorperiodsofplantevolution – Newstructuresevolved,adaptiveradiationsfollowed • Originofplantsfrom aquaticancestors • Diversificationof vascularplants • Originofseeds • Emergenceof floweringplants D. Evolution of land plants

1. Latest Ordovician

2. Early and Middle Silurian

3. Late Silurian E. Devonian plantscape

i. Vascular tissue (e.g. rhyniophytes)

ii. Advantages E. Devonian tree 4. Early Devonian (Archaeopteris, 30m) Adaptive radiation Plant evolution 5. Late Devonian

i. Seeds

a. Advantages

ii. Adaptive radiation

Plant evolution PLANTEVOLUTION

• Plantancestrycanbeviewedintermsofa nestedsetofmonophyleticgroups flowering green zygophytes, charophytes bryophytes lycophytes horsetails ferns cycadsginkgos conifers gnetophytes plants algae related groups

seed plants euphyllophytes vascular plants embryophytes (land plants) (closely related groups) BRYOPHYTES – first in Ordovician

• Thegametophyteisthedominantgenerationinthe lifecyclesofbryophytes

BRYOPHYTE ) ) n n SPOROPHYTE (2 SPOROPHYTE GAMETOPHYTE ( GAMETOPHYTE

GR ALGA BRYOPHYTE FERN GYMNOSPERM ANGIOSPERM VASCULARPLANTS

Adaptations of vascular plants • Differentiatedbodies – Subterraneanrootsystems(water&minerals) – Aerialstemsandleaves(photosynthesis) • Vasculartissue – Xylem(water&minerals) – Phloem(organicnutrients) • Lignin – Cellwallcomponentprovidingmechanicalsupport Psilopsida &Rhyniopsida

• Earliest land plants • Does not possess leaves or true roots • Stems photosynthesize • Simple, dicotomous branching • Apical reproductive structures, spores in sporangia • Rhynia is earliest fossil • Reconstruction of Early Devonian landscape. – showingsomeoftheearliestlandplants

Protolepidodendron \

Dawsonites /

- Bucheria

SEEDLESS VASCULAR PLANTS

• Dominated forest landscapes of Devonian and Carboniferous period Three living divisions – Lycophyta – Horsetails (Shenophyta) – Ferns (Pterophyta) LYCOPHYTES

• Division Lycophata • EvolvedinDevonianperiod – PrevalentinCarboniferousperiod – Woodytreelineage • BecameextinctnearendofCarboniferousperiod – Herbaceouslineage • Representedtodayby~1,000species

LYCOPHYTES

Plavuň vidlačka Sphenophyta • Divisionhorsetail • Ancientlineageofseedless vascularplants – Datesbackto Devonian Prevalentduring Carboniferous • Modernsurvivorsinclude ~15speciesinthegenus Equisetum – MostcommoninNorthern hemisphere – Generallyfoundindamp locations,streambanks FERNS

• Division Pterophyta • Ancient ancestry – Origins in Devonian period – Prevalent in Carboniferous period • Currently most prevalent seedless vascular plant – >12,000 species exist today – Most diverse in tropics SEEDPLANTS Key adaptations of seed plants • Reductionofthegametophyte – Minutegametophytesretainedwithinand protectedbythesporophyte • Adventoftheseed – Seedsreplacedsporesasameansofdispersing offspring • Evolutionofpollen

– Eliminated the liquid H 2O fertilization requirement PlantEvolution • Majoreventsintheevolutionoflandplants – TheDevonianPeriodwasatimeofrapidevolutionforthelandplants

– the appearance ofleaves

– and emergenceof seeds EvolutionofSeeds

• TheevolutionoftheseedduringtheLateDevonian – liberatedlandplantsfromtheirdependenceonmoistconditions – andallowedthemtospreadoverallpartsoftheland

• Intheseedmethodofreproduction – thesporesarenotreleasedtotheenvironment – butareretainedonthesporebearingplant – wheretheygrowintothemaleandfemaleforms Progymnosperms • Seedlessvascularplants– butlikelyprogenitorsof seedplants • Unlikeotherseedlessvascularplants, progymnosperms hadsecondaryvasculartissue (bothxylemandphloem)anditsstructureisvery likethatofmodernconifers • Somehadaeustele andwereheterosporous Reconstructionofprogymnosperm Archaeopteris EvolutionofConiferSeedPlants

• Theappearanceofheterospory – wasfollowedseveralmillionyearslater – bytheemergenceofprogymnosperms • MiddleandLateDevonianplants • withfernlikereproductivehabit • andagymnospermanatomy – whichgaveriseintheLateDevonian – tosuchothergymnospermgroupsas • the seed ferns • andconifertype seed plants Gymnosperms

• Paraphyletic groupingofunrelatedwoodyseedplantsthat arejustnotAngiosperms • Flowerless,seedbearingplants • Possesscones,pollencarriedbywind • Seedsdispersedbygravityorsomeanimals • Importantgroups: – Pteridosperms “seedferns” – Cordaitales straplikeleaveswithparallelvenation – Cycads – Ginkgos – Conifers (pines&firs) nextlab EvolutionofSeeds

• TheevolutionoftheseedduringtheLateDevonian – liberatedlandplantsfromtheirdependenceonmoistconditions – andallowedthemtospreadoverallpartsoftheland

• Intheseedmethodofreproduction – thesporesarenotreleasedtotheenvironment – butareretainedonthesporebearingplant – wheretheygrowintothemaleandfemaleforms Pteridosperms

• Seedferns • Havesimilarphenotypiccharactersasthe trueferns,butwithseeds&conesinsteadof spores Fossils • OldestseedsfromDevonian(365mya) • Elkinsia • Archaeosperma

Gymnospermphylogenyisamess.

MYA --Epoch 50Tertiary 100 150Cretaceous Angiosperms 200Jurassic 250Triassic Cycads Ginkgos 286Permian Gnetophytes 360Carboniferous Conifers Seed Ferns Ferns Progymnosperms 400Devonian Lycophytes Zosterophyllophytes Trimerophytes 425 Rhyniophyta Cooksonia 450 500Silurian 600CambrianOrdovician

SeedlessVascularPlant

• Generalizedlife historyofa seedlessvascular plant • Themature sporophyteplant producesspores – whichupon germination growintosmall gametophyte plants SEEDLESSVASCULARPLANTS Phylogeny