A Phylogenetic Analysis of the Early Late Ordovician Orthograptinae (Graptolithina)

A Phylogenetic Analysis A Phylogenetic Analysis of the Early Late of the Early Late

Ordovician Ordovician Orthograptinae Orthograptinae (Graptolithina)(Graptolithina)

Sue KlostermanSue Klosterman

Charles Charles MitchellMitchell

Dan GoldmanDan Goldman

MacroevolutionMacroevolution

Large scale project on Large scale project on evolutionary trendsevolutionary trends

Phylogenetic Phylogenetic reconstruction of the reconstruction of the Orthograptinae is an Orthograptinae is an essential componentessential component

Interpretation of Interpretation of trends are dependent trends are dependent on phylogenyon phylogeny

DDO RadiationDDO Radiation Rapid expansion in Rapid expansion in

the Early Late the Early Late Ordovician, about Ordovician, about 458 million years ago458 million years ago

Difficulty in Difficulty in determining determining evolutionary evolutionary relationshipsrelationships

Character AnalysisCharacter Analysis

Commonly Commonly preserved preserved on black on black shales shales

3-D specimens 3-D specimens extracted extracted from from limestone limestone matrixmatrix

Character ExamplesCharacter Examples 56 parsimony-56 parsimony-

informative informative discrete discrete characterscharacters

Thecal Shape

Apertural Orientation

10 continuous 10 continuous characters – characters – measurement measurement datadata

2TRD2TRD

Colony Width

PAUP

Tree AnalysisTree Analysis

MacCladeMacClade

Branch & Bound Branch & Bound SearchSearch

RC Index RC Index Character WeightingCharacter Weighting

16 Taxa16 Taxa

Outgroup SelectionOutgroup Selection

HustedograptusHustedograptus as primitive outgroup as primitive outgroup

Consensus Tree - Consensus Tree - Discrete characters onlyDiscrete characters only

9 MPR – all variations within O. quadrimucronatus group

Generally compatible with current systematics

H. u

plan

dicu

s

H. t

eret

iusc

ulus

O. c

alca

ratu

s inc

isus

O. a

picu

latu

s

O. p

agea

nus

O. s

pini

geru

s

O. q

uadr

imuc

rona

tus

Prim

itive

Rec

togr

aptu

s

R. a

mpl

exic

aulis

G. p

ygm

aeus

G. t

ypic

alis

A.p

raet

ypic

alis

A. m

axw

elli

A. p

erex

cava

tus

O. r

uede

man

ni

O. e

ucha

ris

Individual CladesIndividual Clades

OrthograptusOrthograptus

OrthograptusOrthograptus is a is a paraphyletic taxon paraphyletic taxon from which from which RectograptusRectograptus, , AmplexograptusAmplexograptus and and GeniculograptusGeniculograptus have have been derivedbeen derived


Orthograptus Orthograptus calcaratuscalcaratus group is group is the primitive sister the primitive sister group to the rest of group to the rest of the derived the derived orthograptinesorthograptines

Derived orthograptines


A – R – G subclades

Orthograptus Orthograptus quadrimucronatusquadrimucronatus group is group is the sister group to the the sister group to the AmplexograptusAmplexograptus, , RectograptusRectograptus and and Geniculograptus Geniculograptus subcladessubclades


RectograptusRectograptus is is the primitive the primitive sister group to sister group to the the AmplexograptuAmplexograptuss and and GeniculograptuGeniculograptuss sub-clades sub-clades

Thecal DiversificationThecal Diversification

Thecal Thecal diversification diversification from primitive, from primitive, straight tubes to straight tubes to more complex more complex geniculate thecae geniculate thecae for this subclade.for this subclade.

Cladogram SupportCladogram Support

CI = .84CI = .84

RI = .89RI = .89

RC = .74RC = .74

Consistency Consistency Indicies – low Indicies – low levels of levels of homoplasyhomoplasy


0

2

4

6

8

10

12

14

16

18

1 11 21 31 41 51 61 71 81 91 101 111 121 131 141

Cladogram Length

Fre

qu

en

cy

PTP Test – No PTP Test – No permutated permutated cladograms are shorter cladograms are shorter in length than original in length than original cladogramcladogram

Original Original

Permutated Permutated


H. u

plan

dicu

s

H. t

eret

iusc

ulus

O. c

alca

ratu

s inc

isus

O. a

picu

latu

s

O. p

agea

nus

O. s

pini

geru

s

O. q

uadr

imuc

rona

tus

Prim

itve

Rec

togr

aptu

s

R. a

mpl

exic

aulis

G. p

ygm

aeus

G. t

ypic

alis

A.p

raet

ypic

alis

A. m

axw

elli

A. p

erex

cava

tus

O. r

uede

man

ni

O. e

ucha

ris

Bremer Support Indicies

6

2

1

6

1

11

4

100

0

Consensus Tree - Consensus Tree - Measurement Data Measurement Data

includedincluded

H. u

plan

dicu

s

H. t

eret

iusc

ulus

O. c

alca

ratu

s inc

isus

O. a

picu

latu

s

O. p

agea

nus

O. s

pini

geru

s

O. q

uadr

imuc

rona

tus

Prim

itve

Rec

togr

aptu

s

R. a

mpl

exic

aulis

G. p

ygm

aeus

G. t

ypic

alis

A.p

raet

ypic

alis

A. m

axw

elli

A. p

erex

cava

tus

O. r

uede

man

ni

O. e

ucha

ris

Consensus Tree - Consensus Tree - Measurement Data Measurement Data

includedincluded

H. u

plan

dicu

s

H. t

eret

iusc

ulus

O. c

alca

ratu

s inc

isus

O. a

picu

latu

s

O. p

agea

nus

O. s

pini

geru

s

O. q

uadr

imuc

rona

tus

Prim

itve

Rec

togr

aptu

s

R. a

mpl

exic

aulis

G. p

ygm

aeus

G. t

ypic

alis

A.p

raet

ypic

alis

A. m

axw

elli

A. p

erex

cava

tus

O. r

uede

man

ni

O. e

ucha

ris

O. ruedemanni becomes sister taxon to A - R - G subclade

Relationship of O. quadrimucronatus group is better defined

Future WorkFuture Work

Start to incorporate Start to incorporate the orthograptinae the orthograptinae cladogram with cladogram with DDO fauna and DDO fauna and other graptolite other graptolite groupsgroups

Lasiograptids Dicranograptids

Orthograptids

AcknowledgmentsAcknowledgments

NSF Grant Ear - 0106844NSF Grant Ear - 0106844

Documents

A Phylogenetic Analysis of the Early Late Ordovician Orthograptinae (Graptolithina)