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Fish/tetrapod Communities in the Upper Devonian

Maxime Delgehier

Uppsala universitet, Institutionen för geovetenskaper

Examensarbete D/E1/E2/E, Geologi/Hydrologi/Naturgeografi /Paleobiologi, 15/30/45 hpISSN 1650-6553 Nr 301

Examensarbete vid Institutionen för geovetenskaper ISSN 1650-6553 Nr 301

Fish/tetrapod Communities in the Upper Devonian

Maxime Delgehier

Vertebrate communities including tetrapods and fishes are known from a limited number of Late Devonian localities from several areas worldwide.

These localities encompass a wide variety of environments, from true marine conditions of the near shore neritic province, to fluvial or lacustrine conditions. These localities form the foundation for a number of data matrices from which three different sets of cluster analyses were made. The first set practices a strait forward taxonomical framework using present/absent data on species and genus level to test similarity between the various localities. The second set of analyses builds on the first one with the integration of artificial hierarchies to compensate taxonomical biases and instead infer relationship. The third also builds on the previous ones, but integrates morphological data as indicators of relationships between taxa. From this, a critical review was made for each method which comes to the conclusion that the first analysis and the first artificial level of the second analysis provide the distinctions between Frasnian and Frasnian/Famennian locality whereas the second artificial level of the second analysis and the third analysis need to be improved.

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Supervisor: Henning Blom Examensarbete vid Institutionen för geovetenskaper

ISSN 1650-6553 Nr 301

Fish/tetrapod Communities in the Upper Devonian

Maxime Delgehier

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Copyright Maxime Delgehier and the Department of the Earth Sciences Uppsala University Published at Department of Earth Sciences, Geotryckeriet Uppsala University, Uppsala, 2014

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Abstract

Vertebrate communities including tetrapods and fishes are known from a limited number of Late Devonian localities from several areas worldwide. These localities encompass a wide variety of environments, from true marine conditions of the near shore neritic province, to fluvial or lacustrine conditions. These localities form the foundation for a number of data matrices from which three different sets of cluster analyses were made. The first set practices a strait forward taxonomical framework using present/absent data on species and genus level to test similarity between the various localities. The second set of analyses builds on the first one with the integration of artificial hierarchies to compensate taxonomical biases and instead infer relationship. The third also builds on the previous ones, but integrates morphological data as indicators of relationships between taxa. From this, a critical review was made for each method which comes to the conclusion that the first analysis and the first artificial level of the second analysis provide the distinctions between Frasnian and

Frasnian/Famennian locality whereas the second artificial level of the second analysis and the third analysis need to be improved.

Keywords: Palaeobiogeography, vertebrates, Tetrapods, Frasnian, Famennian

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Populärvetenskaplig sammanfattning

Samhällen som innehåller ryggradsdjur, vilken innefattar såväl fiskar som landryggradsdjur (så kallade tetrapoder), är kända från flera lokaler av sen devonsk ålder runt om i världen. Dessa representerar också flera olika miljöer, vilket gör dessa till en bra grund för vår förståelse över hur ekologin påverkade de tidigaste tetrapoderna. Sammansättningen av ryggradsdjur i dessa lokaler ligger till grund för en jämförande analys där tre olika klusteranalyser har använts. Den första analysen använder närvaro/frånvaro-data av traditionellt etablerade arter och släkten för att på så sätt etablera likheten mellan de olika faunorna. Den andra analysen använder samma grundmetod, men dör art och släkten har ersatts av artificiella hierarkier för att på så sätt kompensera för taxonomisk bias och samtidigt ta hänsyn till andra antaganden om släktskapsförhållanden. Den tredje analysen ersätter namn och hierarkier med morfologiska karaktärer för att på så sätt få en annan och taxonomiskt oberoende test av likheterna mellan faunorna hos de olika lokalerna. Dessa tre analyser kunde sedan jämföras, med resultatet och slutsatsen att traditionell taxonomisk jämförelse och till viss del även artificiella hierarkier, ger bättre och mer tolkningsvänliga resultat där studien visar på en tydlig uppdelning i tid mellan lokalerna, medan den tredje metoden bör utvecklas ytterligare.

Vertebrate communities including tetrapods and fishes are known from Late Devonian localities from several areas worldwide. These localities encompass a wide variety of environments. Their

understanding is a key to understand ecologic pressure on these first tetrapods. The vertebrate communities from these localities are the base of data matrices from which three different cluster analyses were made. The first analysis is on forward taxonomical framework using present/absent data on the species and genus to test similarity between the various localities. The second builds on the previous one with the integration of artificial hierarchies to compensate taxonomical biases and instead infer relationship. The third analysis also builds on the previous ones, but integrates morphological data as indicators of relationships between taxa. From this, a critical review was made for each method which comes to the conclusion that the first analysis and the first artificial level of the second analysis provide the distinctions between Frasnian and Frasnian/Famennian localities whereas the second artificial level of the second analysis and the third analysis need to be improved.

Keywords: Palaeobiogeography, vertebrates, Tetrapods, Frasnian, Famennian

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Table of Contents

1.  Introduction ... 1 

2.  Aim ... 2 

3.  Background ... 2 

4.  Methodology ... 12 

5.  Results ... 15 

5.1  First analysis ... 15 

5.1.1  Species level ... 15 

5.1.2  Genus level ... 17 

5.2  Second analysis ... 21 

5.2.1  First artificial level ... 21 

5.2.2  Second artificial level ... 24 

5.3  Third analysis ... 27 

6.  Discussion ... 30 

7.  Conclusion ... 35 

8.  Acknowledgements ... 36 

9.  References ... 36 

Appendix ... 47 

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1. I

localities provide v acanthod diversity vertebrat tetrapods which po Ichthyos only a fe China an Greenlan species d

& Clack understa is why th Canada) interrela

Figure 1 locality,

Introdu

Vertebrate c s from severa vertebrate as dians, chondr y of vertebrat te faunas is a s, which repr opularly kno stega in Gree ew Upper De nd Latvia), o

nd, USA, Be described and k, 1993; Ahlb and tetrapod e

he elpistosteg (Figure 1) h ationship betw

: Map showin yellow stars r stars repres

uction

communities al areas worl ssemblages c richthyans, a tes in variou also essential

resent a key own as the ‘fi enland by Säv evonian tetra

ne Frasnian- elgium, Latvi d even more berg et al., 19

evolution it i gid localities have been ad

ween localiti

ng the near-tet represent the e sent Famennia

including tet ldwide. Thes constituted by actinopterygi

s palaeoenvi l to understan

event in the ish-tetrapod ve-Söderberg apod localitie

-Famennian l ia and Russia if including 994; Shubin is also impor s (Lode quarr ded to the lis ies and the ev

trapod and tetr elpistostegid lo

an localities, b

trapods and se localities,

y major grou ians and sarc ironmental co and what cou

vertebrate ev transition’ h gh in 1932 w es known (Fi

locality (Aus a). Today the g the yet unde et al., 2004;

rtant to inclu ry in Latvia, st of tetrapod volution of f

rapod localitie ocalities; oran brown stars re

fishes are kn sometimes r ups of verteb copterygians.

onditions (se uld be the sele

volution. Ou as progresse with recent di

gure 1): thre stralia) and s ere are twelv escribed taxa Daeschler et ude the near-t Ellesmere Is d localities u faunas.

es world-wide nge stars repre epresent Frasn

nown from L referred to as rates as agna . These sites ee Backgroun

ection pressu ur understand

d greatly sin iscoveries. T ee Frasnian lo

even Famenn ve tetrapod ge

a that have be t al., 2009). I tetrapods or sland in Can se in order to

e. Green star re esent Frasnian

ian/Famennia

Late Devonia s tetrapod loc

athans, placo show thus a nd). The stud ures on these ding of this e nce the discov There are at p

ocalities (in S nnian localitie enera and fo een found (L In order to

elpistostegid nada and Mig o study the

represents Gog n tetrapod loca an localities.

n calities, oderms, a wide

dy of the e first

vent, very of present

Scotland, es (in

urteen Lebedev

ds. This guasha in

go station alities; red

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Some studies (Young et al., 1992; Talent et al., 2000; Young et al., 2000; Lebedev &

Zakharenko, 2010; Lebedev et al., 2010) were done to understand the global and or provincial relationships between vertebrates during the Devonian as their migrations, their ecology and more particularly on tetrapods (Lelièvre, 2002; Clack, 2006; Blieck et al., 2007). These studies suggest that there are two migration hypotheses for tetrapods: one northern and one southern. The northern one goes through central and southern Asian continental blocks and the southern one along the northern margin of Gondwana (Blieck et al., 2007), neither of which can be confirmed.

2. Aim

The aim of my study is to explore the faunal relationships between different Late Devonian tetrapod yielding localities worldwide. This will be done using several numerical methods using similarity indices, including cluster analysis. Two new methods, based on artificial hierarchies and on morphological data, will be tested.

3. Background

Since the discovery of the first specimen of the well-known tetrapod, Ichthyostega Save- Söderbergh, 1932 in Eastern Greenland (Gauss Halvø and Ymer Ø sites (Figure 2)) more and more tetrapod localities were found world-wide. East Greenland locality have provided a wide diversity of Famennian vertebrates (Blom et al., 2007), including placoderms, chondrichthyans, acanthodians, an actinopterygian, lungfishes, porolepiforms, osteolepiforms and four tetrapod species. The most complete tetrapods are Acanthostega gunnari Jarvik, 1952, Ichthyostega stensioei Säve-Söderbergh, 1932 Ichthyostega watsoni Säve-Söderbergh, 1932 and Ichthyostega eigili Säve-Söderbergh, 1932, while Ymeria denticulata Clack et al., 2012 is recognized based on jaw and tooth morphology only.

All the tetrapods come from Famennian sediments of the Celsius Bjerg Group interpreted as being primarily of fluvial origin (Olsen & Larsen, 1993).

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Tetrapod Russia, A

identifie discover

The Lan fossils w

Figure 2:

ds are also fo Australia, Ch The Strud lo d as Ichthyos ry the origina

Figure 3: M nglier quarry were found in

: Map of East ound in other hina and Sco ocality in Nam

stega-like (C al quarry, wh

Map showing th fossils, near n the same fo

Greenland sh r the localitie otland.

mur Provinc Clément et al hich was aban

he localization Durnal (Fig ormation (Ev

howing the De es around the

e, Belgium ( l., 2004) belo

ndoned for o

n of Strud loc gure 4) were vieux formati

evonian outcro e world, inclu

(Figure 3) pr onging to the one century,

cality in Belgiu added to the ion) with sed

ops (Blom et a uding Belgiu

ovided a part e Evieux form

is now being

um (Gueriau e taxa list of S diments whic

al., 2007).

um, USA, La

rt of a lower j mation. With g studied aga

et al., 2014).

Strud becaus ch suggest a m

atvia,

jaw h this ain.

se these mixed

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alluvio-l environm (bothriol (Clémen 2013) an

Densign based on al., 2004 belongin osteolep interpret

lagoonal env ment (Goema

lepids, phyllo nt & Boisvert

nd tetrapodom

Figure 4: Ma Several tetra athus rowei n a left cleith 4; Daeschler ng to placode iforms (Cres ted as the res

vironment wi aere, 1995; T olepids, groe t, 2006), tetra morph fishes

ap showing th apods have a Daeshler, 20 hrum and a po

et al., 2009) erms, chondr ssler III et al.

sult of a delta

th estuary or Thorez et al., enlandaspids apodomorph s (Clément, 2

he localization lso been foun 000 based on

osterior part . In associati richthyans, ac

., 2010). The aic environm

r tidal delta i , 2006). Thes s), holoptych h fishes in Str

2002; Cléme

n of Durnal loc und in Red H

n a lower jaw of a right low ion with thes canthodians, ese fossils be ment during th

nfluences an se localities p hiid porolepif

rud and acan nt et al., 200

cality in Belgi ill, Pennsylv w, Hynerpeto wer jaw and se tetrapods a , actinopteryg elong to the C he Famennia

nd Strud sedim provide rema forms, rhynch nthodians (De 09).

ium (Clément vania (Figure

n bassetti Da a possible w are also verte gians, lungfi Catskill Form an.

iments as a d ains of placo hodipterid lu erycke & Cl

t et al., 2009).

e 5) including aeschler et a watcherid (Sh

ebrate remai ishes and mation which

deltaic oderms ungfishes

ément,

g l., 1994 hubin et

ns

h is

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Figure 5:

of bones placoder and a po which ar

Figure 6 Russi

: Map showin

Andreyevka s, scales and rms (Aleksee ossible secon

re interpreted

6: Map showin ia; (B) Locatio

g the location

a in Tula regi teeth belong ev et al., 199 d undetermin d as a marine

ng the localiza on of Andreye represen

n of the Red H

ion, Russia (F ging to sarcop 94). The skele ned tetrapod e environmen

ation of Andre evka locality in nt the position

Hill site, Clinto 2010) Figure 6) pro pterygians, a eton of the te (Lebedev &

nt.

eyevka (A) Lo n the Tula reg n of outcrop. (

on County, Pen

ovide abunda acanthodians

etrapod Tule

& Clack, 1993

ocation of the gion; (C) Andr (Alekseev et a

nnsylvania, U

ant Famennia , actinoptery rpeton curtu 3) were foun

Tula region in reyevka locali al., 1994)

USA. (Cressler

an vertebrate ygians, antiar um Lebedev, nd in limeston

n the Europea ity, the black

r III et al.,

e remains rch

1984 nes

an part of circles

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7). These deposits which be osteolep fragmen

Figure indicate Detail Ketleri Formatio

There are tw e sites belon (Luksevics &

elong to the iforms and te t and a possi

7: Maps show es the extent o

l map of the S i hamlets, the on; light shadi

wo tetrapod lo ng to the Ketl

& Zupins, 20 same fauna a etrapods: Ve ible undeterm

wing the local f Famennian a Skrunda area s

stylized fishe ing, Skervelis Form

ocalities in L leri Formatio 004). Both si and includes entastega cur mined tetrapo

ization of the and later depo showing the lo s mark the fos Formation. T mation. Scale

Latvia dating on constituted

ites are chara placoderms ronica Ahlbe od (Ahlberg

Latvian local osits, the squar ocalities of Pa

ssil localities.

The fossil loca bar 3 km. (Ah

to the Fame d by sandsto acterized by

, acanthodian erg et al., 199 et al., 1994)

ities (a) Map re of dark sha vari and Ketle

Crosses, Zag ality at Ketleri hlberg et al., 1

ennian: Pavar nes interpret rich vertebra ns, porolepif 94 on the bas

.

of Latvia; the ding is the are eri. The house

are Formation lies just below 1994)

ri and Ketler ted as marine ate assembla forms, lungfi sis of mandib

e area of light ea of the next e indicate Pav n; no shading, w the top of th

ri (Figure e

ages ishes,

bular

shading map. (b) ari and Ketleri he Ketleri

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(Figure 8 remains, and meg sandston

Fig Gorno

and desc Shale. T fluvial en rather cl placoder near the tetrapod- (1993) w

In the Oryol 8). This quar , such as plac galichtyds. Th ne and clay in

gure 8: Map sh ostayevka quar

A poorly pre cribed as belo he fish fauna nvironment, ose to the fam rms, lungfish

base of the H -fish assemb which could b

l region (Rus rry provided coderms, aca hese fossils w nterpreted as

howing the loc rry in the Ory

eserved right onging to Me a of placoder

late Frasnian mous Canow h, rhizodonts

Hervey Grou blage was ass be with the p

ssia) near the the tetrapod anthodians, c were found i s a deltaic to

calization of th yol region, a bl

t lower jaw o etaxygnathus rms and lung n or early Fr windra site, w

and tristicho up with evide signed to the previous cons

e town of Liv d Jakubsonia chondrichthy in Famennian

near-shore e

the locality in lack diamond

2004) of a tetrapod

s denticulus gfishes is kno rasnian in age which presen

opteryds. Th ence of a ma se two fauna sideration as

vny, there is t livnensis Le yans, porolep n sedimentar environment.

Russia, an ast d marked the lo

have been fo Campbell &

own from thi e (Campbell nt no tetrapod he vertebrates rine incursio a called ‘Jem s a Frasnian o

the Gornasta ebedev, 2004 iforms, lung ry deposits co

.

terisk shows t ocation of And

ound in Jema Bell, 1977 f is locality an

& Bell, 197 d but show a

s from Canow on dating to F malong-Canow

or Famennian

ayevka quarr 4 and vertebr gfishes, osteo

onstituted by

the location of ndreyevka. (Le

along site (Fi from the Clo nd have sugg 7). This loca

fauna consti windra were Frasnian. The windra’ by Y n age. There

ry ate olepids y

f the ebedev,

igure 9) ghnan ested a ality is

ituted by found e Young e are

(15)

probably discover

Figure 9

panchen heterostr 1950). T from a fl

y more tetrap red (Clack, 1

9: Map of Aus

Near Elgin i ni Ahlberg et racans. The d This vertebrat luvial channe

Figu

pods from Au 997).

stralia showin by tri in Scotland (F

al., 1995 wi dating of this te assemblag el.

ure 10: Map s

ustralia and e

g the localizat iangles (from Figure 10), t ith a fauna co s fauna due t ge was found

showing the lo

especially in

tion of Jemalo Johanson et a the Scat Crai onstituted by to the presen d in sandston

ocation of Elg

Victoria wh

ong and Cano al., 2003 modi ig locality pr y porolepifor ce of heteros nes, these dep

gin in Scotland

here trackway

windra localit ified).

ovided bone ms, lungfish stracans is Fr posits were in

d (Ahlberg, 19

ys have been

ty, fossil site i

es of Elginerp hes, placoderm

rasnian (Gro nterpreted as

998).

n

indicated

peton ms and ss, s coming

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1977 wa sandston shallow in verteb acanthod heterostr species w

Figure 1

have pro with a di were fou (Jia et al

Velna-Ala (F as found in O ne, siltstone, waters of a n brate remains dians, actinis racans has da was found bu

1: A, B Local

The Frasnian ovided only o iverse fauna und, is still pr

l., 2010). The

Figure 11) in Ogre Formati

dolomitic m narrow gulf o s and the fau stians, porole ated the Ogre ut the locality

lization of Lan

n sediments one poorly pr

of vertebrate roblematic to e depositiona

n Latvia is th on. This form marl, clay and

of the Baltic una provides epiforms, lun e Formation y is not preci

ngsede site in near Sabile of Ningxia H reserved tetr es. The preci o date (Bliec al environme

he locality wh mation is com d gypsum, wh

palaeobasin some specim ngfishes and

to Frasnian isely known

Europe and in e (Luksevics e Hui autonom rapod lower j ise age of the ck et al., 200

ent of this lo

here Obruch mposed of a hich are inter by Sorokin mens of heter tristichopter in age. In we

and could b

n Latvia; C St et al., 2011).

mous region in jaw named S e Zhongning 7) and needs cality is inter

evichthys gra siliciclastic s rpreted to be

(1978). This rostracans, pl yds. The pre estern Russia e in the Novg

tar indicate loc

n northweste Sinostega pan

Formation, s further bios rpreted as no

racilis Vorob sequence of e formed in th s is a relative

lacoderms, esence of

a the same te vgorod distric

cation of Lang

ern China (Fi ni Zhu et al., where these stratigraphic on-marine.

byeva

he ely poor

etrapod ct.

gsede site

igure 12) , 2002

fossils research

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Figure 12 of the

and relev found. E (Canada

actinopte rhombol problem

2: (A) Locatio Zhongning Fo

In order to u vant to inclu Elpistostegid a). All of thes Lode quarry erygians, act lepis Gross, 1 matic (Figure

on of Ningxia ormation in fo

understand th de to the loc localities are se sites are F y (Figure 13) tinistians, lun 1941 from th 13). The sed

Hui Autonom our selected D

he interrelatio alities where e composed b

rasnian.

provide a fa ngfishes, oste he top part of diments of the

mous Region; ( Devonian secti

2010).

onships betw e elpistostegi by Lode (La

auna compos eolepiforms f Lode Form e quarry wer

(B) Devonian ions mainly ba

ween these tet id (also calle atvia), Ellesm

ed by agnath (Upeniece, 2 mation. Also t

re interpreted

outcrops in N ased on Pan et

trapod locali d near-tetrap mere (Canada

hans, placode 2001) and Pa the precise da d as deltaic e

Ningxia; (C) C t al., 1987 (Jia

ities, it is imp pods) have b a) and Migua

erms, acanth anderichthys

ating still environment.

Correlation a et al.,

portant een asha

odians, s

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Figure Forma

2006 wa porolepi Formatio interpret

Figure

e 13: Map sho ation (there is The Ellesme as found here tforms (Dow on which is c ted as the dep

e 14: Location

owing the loca s uncertainty a ere Island is i e in one local wns et al., 20 characterized posits of mea

n and stratigrap T

alization of Lo about the posit in the Canad lity with othe

11), lungfish d by alternati andering stre

phic position Territory, Can

ode quarry in L tion of the Giv dian Arctic (F er fish remai hes and osteo

ing resistant eam systems

of the site (NV nada. (Daeschl

Latvia and the vetian-Frasnia Figure 14). T ins such as an olepiforms. T sandstones a (Embry & K

V2K17) on so ler et al., 2006

e stratigraphic an boundary) (

Tiktaalik rose ntiarch placo The come fro and recessive Klovan, 1976

outhern Ellesm 6)

c column of th (Forey et al.,

eae Daeschle oderms, holo om the Fram e siltstones, a 6; Embry, 19

mere Island, N he Lode

2000).

er et al., optychiid

and 91).

Nunavut

(19)

Devonia fauna wa acanthod 2011) an interpret

Figure 1 Group i

For the p Strud as China. O Appendi

4. Me

faunal re done clu

Miguasha (F an faunas in t

as found in th dians, actinop nd Elpistosteg ted as estuari

15: a. Localiza includes the F

present analy the Belgium Other names

ix.

ethodol

The methodo elationships a uster analysis

Figure 15), in the world du

he Escumiac pterygians, a ge watsoniW ine (Cloutier

ation of Migu Fleurant and E Escum yses, Gauss H m locality, Ke

are used unc

logy

ological core and similarit s on several d

n Québec Pro e to an excep c Formation a actinistians, p Westoll 1938 r et al., 2011)

uasha in easter Escumiac Form miac Formation Halvø and Ym etleri and Pav changed. The

e of this stud ty between th data: the first

ovince (Cana ptional conse and includes porolepiform 8. The deposi

).

rn Québec, b.

mations), c. Ge n (see in b) (C mer ø localit vari as Pavar e complete li

dy is to use se he different t

t based on ta

ada), represe ervation (Jan s anaspids, os ms, lungfishe

itional enviro

Geological m eneral view of Cloutier et al.,

ties are referr ri, Velna-Ala st of taxa pre

everal sets of tetrapod loca axonomical fr

nts one of th nvier & Arsen

steostracans, s, osteolepifo onment of th

ap of the Mig f the east side

2011).

red to as the a as Ogre and

esent in each

f analyses in alities worldw framework, th

he best know nault, 2009).

, placoderms forms (Clouti his site was

guasha area (M e of the synclin

Greenland l d Ningxia Hu h locality is in

n order to exp wide. For thi

he second on n Late . This s,

ier et al.,

Miguasha ne of the

ocality, ui as n the

plore the s I have n

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artificial hierarchies of the previous taxa and the third on characters from taxa. Cluster analyses were selected, because they are simple to use and provide results which are rather strait forward to analysis.

Cluster analyses using the Jaccard similarity index have been used for each database to explore the faunal relationships between localities. Other indices were explored but they don’t provide conclusive results. This is why this index was preferred because it emphasizes datasets which have only few common data (Hammer et al. 2001). PAST version 1.56b (Hammer et al. 2001) was used to perform the cluster analyses.

To do these analyses, the first step was to identify relevant Late Devonian tetrapod localities.

In that case, there are ten localities (see Background part), but in order to expand the list of localities and include a broader definition of tetrapods, near-tetrapod localities were added. Thus three localities were added: Ellesmere, Lode and Miguasha. Canowindra (Australia) was also added because of the proximity of this locality with Jemalong, these sites are sometimes considered as having the same fauna (Young, 1993). In that respect, each analysis was done with these two localities separated and grouped called JeCa. The whole of localities is considered as the ingroup. Finally the reference locality without tetrapods or elpistostegids is the Gogo station (Western Australia) where an

exceptional preservation (Long & Trinajstic, 2010) provided a great number of species of placoderms, actinopterygians, lungfishes and sarcopterygians.

The basis for all analyses is to use presence or absence of taxa to compare the localities. The first step is the listing of taxa for each and one of the localities. Some localities as Miguasha or Gogo provide lot of Frasnian fossils which give a good picture of the vertebrate fauna during the Late Devonian, but others such as Ellesmere record only some vertebrates or contain still undescribed material. These taxa are then placed in the matrix and subdivided into the different vertebrate groups:

Anaspida, Osteostraci, Heterostraci, Galeaspida, Placodermi, Chondrichthyes, Acanthodii, Actinopterygii, Actinistia, Porolepiformes, Dipnoi, Osteolepiforms, Elpistostegalia and Tetrapoda (taxa list in Appendix part). Finally the matrix is coded: absent taxa as 0 and present taxa as 1. Then the matrix is run with the software PAST version 1.56b (Hammer et al. 2001) in order to get

dendrograms which will be described and interpreted to understand the faunal relationships.

The second analysis is based on artificial hierarchies of taxa, which is a new method based on assumed interrelationships from available phylogenies rather than traditional taxonomical hierarchies.

There are two levels of hierarchies: the first level could be assimilated to a classical family level, the second level which group some of first level taxa could be assimilated to a sub-order or super-family level. Also each localities of the previous analysis are used for this analysis. Taxa for which there are not phylogenetic analyses or which were not included in phylogenetic analyses, the current family level is used. A taxon not described or an indeterminate taxon cannot be grouped with other taxa. In that respect these taxa get a level for them. In the case of the indeterminate taxon Acanthodii indet. A, the artificial level 1 is Acanthodii 7 and the artificial level 2 is Acanthodii C (see the list of taxa). The

(21)

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(22)

is Ellesmere which have the greater number of characters in plesiomorphic state. This is explained by the few number of taxa present and whose which are present (Asterolepis spp., Holoptychius,

Laccognathus, Glyptolepis and Tiktaalik) show characters in plesiomorphic states.

5. Results

5.1 First analysis

5.1.1 Species level

In this first cluster analysis (Figure 17), in which Jemalong and Canowindra are treated as separate localities, presence/absence data for 284 species resulted in very low similarity indices between the various localities. This is most likely due to the low number of common taxa. The y axis represents the similarity index and the x axis represents the number of localities.

The most distinct cluster, with rather high levels of similarity, includes Pavari, Ogre, Oryol and Andreyevka. These localities from this cluster come from Latvia and Western Russia. So this cluster can be called the Baltica cluster. Ogre is the only locality of the cluster which is Frasnian.

This cluster is also grouping with another distinct cluster, including Greenland, Jemalong, RedHill and Belgium. Jemalong is present in this cluster because some taxa as Bothriolepis and Phyllolepis species from this locality are only present in Famennian localities of the Northern Hemisphere whereas Jemalong is Late Frasnian to Early Famennian in age. Except this locality, Greenland, RedHill and Belgium belong to Laurussia this is why this cluster can be considered as a Laurussia cluster.

There is one additional cluster showing some similarity between Ellesmere and Lode which are two near-tetrapod localities. Only one porolepiform is common to these localities. This cluster shows no similarity with any of the other localities, including the Miguasha locality.

Other localities (Miguasha, China, ScatCraig and Canowindra) show such low similarity index, considered null. This could be explained by the endemism and the exceptional conservation in Miguasha and by a low number of taxa from China, ScatCraig and Canowindra localities.

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Figure 17: Dendrogram showing faunal relationship between near-tetrapod and tetrapod localities during the Late Devonian for the species level when Canowindra and Jemalong localities are considered as separated.

In the cluster analysis (Figure 18), in which Jemalong and Canowindra are treated as the same localities called JeCa, presence/absence data for 284 species also resulted in very low similarity indices between the various localities. This is again most likely due to the low number of common taxa. The y axis represents the similarity index and the x axis represents the number of localities.

The most distinct cluster, with rather high levels of similarity, includes Pavari, Ogre, Oryol and Andreyevka. This cluster can be identified as the Baltica cluster because these localities coming from Latvia and Western Russia. All localities are Famennian except Ogre

This cluster is also grouping with another distinct cluster, including RedHill, Belgium, Greenland and JeCa. JeCa are Late Frasnian to Early Famennian in age and present some taxa as Bothriolepis and Phyllolepis species. These taxa are only present in Famennian localities of the Northern Hemisphere whereas JeCa is Late Frasnian to Early Famennian in age. This cluster can be

0 1,6 3,2 4,8 6,4 8 9,6 11,2 12,8 14,4 16

0 0,1 0,2 0,3 0,4 0,5 0,6 0,7 0,8 0,9 1

Similarity Canowindra Ellesmere Lode Andreyvka Pavari Oryol Ogre Greenland Jemalong RedHill Belgium Miguasha Gogo China ScatCraig

(24)

considered as a Laurussia cluster because the three other localities come from the Northern Hemisphere.

An additional cluster, showing some faunal similarity between Ellesmere and Lode, is present.

These near-tetrapod localities show only one porolepiform species common taxa. This cluster shows no similarity with any of the other localities, including the Miguasha locality.

Other localities (Miguasha, China and ScatCraig) show such low similarity index, considered null. This could be explained by endemism and the exceptional conservation in Miguasha and by a low number of taxa from China and ScatCraig localities.

Figure 18: Dendrogram showing faunal relationship between near-tetrapod and tetrapod localities during the Late Devonian for the species level when Canowindra and Jemalong localities are considered as grouped.

5.1.2 Genus level

In this first cluster analysis (Figure 19), in which Jemalong and Canowindra are treated as separate localities, presence/absence data for 175 genera resulted high similarity indices between the

0 1,6 3,2 4,8 6,4 8 9,6 11,2 12,8 14,4

0 0,1 0,2 0,3 0,4 0,5 0,6 0,7 0,8 0,9 1

Similarity Ellesmere Lode Greenland RedHill Belgium JeCa Pavari Oryol Ogre Andreyvka Miguasha Gogo China ScatCraig

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various localities. This is most likely due to the increase number of common taxa which is explained by the genus level: a higher level gives by default a higher similarity index. The y axis represents the similarity index and the x axis represents the number of localities.

The most distinct cluster, with rather high levels of similarity, includes Pavari, Ogre, Oryol, Andreyevka and ScatCraig. Except ScatCraig, these localities from this cluster come from Latvia and Western Russia. So this cluster can be called the Baltica cluster. There are two Frasnian localities which are Ogre and ScatCraig. This latter occupy a basal position in the cluster. This position could be explained by the number of taxa from this site and only few taxa (Holoptychius, Concochus) are common with other localities of the Baltica cluster.

This cluster is also grouping with another distinct cluster, including Greenland, Belgium Jemalong, Canowindra and RedHill. Jemalong and Canowindra are the closest localities with nearly the totality of taxa in common certainly due to the geographical proximity (see Background part).

Jemalong and Canowindra are present in this cluster because some taxa as Bothriolepis species from these locality are only present in Famennian localities of the Northern Hemisphere whereas there two localities are Late Frasnian to Early Famennian in age. Except this locality, Greenland, RedHill and Belgium belong to Laurussia this is why this cluster can be considered as a Laurussia cluster. These three Laurussian localities show the presence of Groenlandaspis, Phyllolepis, Soederberghia and Eusthernodon as common taxa. Greenland and Belgium are close and the common taxa Ichthyostega which is the only tetrapod which is not considered restricted to a single locality.

There is on additional cluster showing some similarity between Ellesmere, Lode and China.

Common taxa are represented by Asterolepis, Laccognahus and Glyptolepis. The China locality falls to the base of this cluster because of endemic taxa as antiarchs (Jingxilepis, Ninjxialepis and Sinolepis) and other taxa. This cluster shows no similarity with any of the other localities, including the

Miguasha locality. The last localities Miguasha show such low similarity index, considered null.

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Figure 19: Dendrogram showing faunal relationship between near-tetrapod and tetrapod localities during the Late Devonian for the genus level when Canowindra and Jemalong localities are considered as separated.

In this second cluster analysis (Figure 20), in which Jemalong and Canowindra are treated as grouped localities, presence/absence data for 175 genera resulted higher similarity indices between the various localities. This is most likely due to the increase number of common taxa which is explained by the genus level: a higher level gives by default a higher similarity index. The y axis represents the similarity index and the x axis represents the number of localities.

The most distinct cluster, with rather high levels of similarity, includes Pavari, Ogre, Oryol and Andreyevka. All these localities come from this cluster come from Latvia and Western Russia. So this cluster can be called the Baltica cluster. Ogre is the only locality which is Frasnian. This cluster show common taxa as Devononchus, Holoptychius, Glyptolepis, Cryptolepis, and Glyptopomus.

This cluster is also grouping with another distinct cluster, including Greenland, Belgium JeCa and RedHill. JeCa are present in this cluster because some taxa as Bothriolepis and Phyllolepis species

0 1,6 3,2 4,8 6,4 8 9,6 11,2 12,8 14,4 16

0 0,1 0,2 0,3 0,4 0,5 0,6 0,7 0,8 0,9 1

Similarity Miguasha China Ellesmere Lode ScatCraig Ogre Andreyvka Pavari Oryol Greenland Belgium Canowindra Jemalong RedHill Gogo

(27)

from these locality are only present in Famennian localities of the Northern Hemisphere whereas there two localities are Late Frasnian to Early Famennian in age. Except this locality, Greenland, RedHill and Belgium belong to Laurussia this is why this cluster can be considered as a Laurussia cluster.

These three Laurussian localities show the presence of Groenlandaspis, Phyllolepis, Soederberghia and Eusthernodon as common taxa. Greenland and Belgium are close and the common taxa

Ichthyostega which is the only tetrapod which is not considered restricted to a single locality. Then, there is the ScatCraig locality which is not grouped with another locality.

There is on additional cluster showing some similarity between Ellesmere, Lode and China.

Common taxa are represented by Asterolepis, Laccognahus and Glyptolepis. The China locality falls to the base of this cluster because of endemic taxa as antiarchs (Jingxilepis, Ninjxialepis and Sinolepis) and other taxa. This cluster shows no similarity with any of the other localities, including the

Miguasha locality. The last locality, Miguasha, is in the basal position.

0 1,6 3,2 4,8 6,4 8 9,6 11,2 12,8 14,4

0 0,1 0,2 0,3 0,4 0,5 0,6 0,7 0,8 0,9 1

Similarity Miguasha China Ellesmere Lode ScatCraig Greenland Belgium JeCa RedHill Ogre Andreyvka Pavari Oryol Gogo

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Figure 20: Dendrogram showing faunal relationship between near-tetrapod and tetrapod localities during the Late Devonian for the genus level when Canowindra and Jemalong localities are considered as grouped.

5.2 Second analysis

5.2.1 First artificial level

In this first cluster analysis (Figure 21), in which Jemalong and Canowindra are treated as separate localities, presence/absence data for 126 artificial grouping (first level) resulted higher similarity indices between the various localities. The y axis represents the similarity index and the x axis represents the number of localities.

The most distinct cluster, with rather high levels of similarity, includes Pavari, Greenland, Oryol and Andreyevka. All the localities are Famennian. This first cluster shows several common taxa as Acanthodii 1, Chondrichthyes 5, Porolepiformes 1, Dipnoi 2 and 3, Onychodontida 2,

Osteolepididae 1 and Tristichopterydae 1. Two sister-groups are presents: an Oryol and Pavari grouping and an Andreyevka and Greenland grouping.

This cluster is also grouping with another distinct cluster, including Canowindra, Jemalong, ScatCraig, RedHill and Belgium. There are ScatCraig and a grouping Canowindra and Jemalong.

These two localities are the closest localities with nearly the totality of taxa in common. The rest of the cluster is composed by RedHill and Belgium localities are still together. This cluster is due to some common taxa as Phlyctaenii 1, Phyllolepida, Dipnoi 3, 11, Osteolepididae 1 Megalichthydae 1 Tristichopterydae 1.

There is on additional cluster showing some similarity between Ellesmere, Lode, China, Ogre and Miguasha. The Miguasha locality falls at the base of this cluster certainly because of the presence of endemic taxa from this locality. Then there two grouping with Antiarcha 1 as common taxa:

Ellesmere and Lode group and Ogre and China.

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Figure 21: Dendrogram showing faunal relationship between near-tetrapod and tetrapod localities during the Late Devonian for the first artificial level when Canowindra and Jemalong localities are considered as separated.

In this second cluster analysis (Figure 22), in which Jemalong and Canowindra are treated as grouped localities, presence/absence data for 126 artificial grouping (first level) resulted higher similarity indices between the various localities. Of course, a higher level gives by default a higher similarity index. The y axis represents the similarity index and the x axis represents the number of localities.

The most distinct cluster, with rather high levels of similarity, includes Pavari, Oryol and Andreyevka. All the localities are Famennian and come from Western Russia and Latvia. This is why this cluster can be called a Baltica cluster. This cluster shows several common taxa as Antiarcha 2 Acanthodii 1, Porolepiformes 2 and Osteolipiformes 1. Andreyevka fall at the base of this group but this group shows always the strong faunal connection between its members.

0 1,6 3,2 4,8 6,4 8 9,6 11,2 12,8 14,4 16

0 0,1 0,2 0,3 0,4 0,5 0,6 0,7 0,8 0,9 1

Similarity Miguasha Ellesmere Lode Ogre China Greenland Andreyvka Pavari Oryol RedHill Belgium ScatCraig Canowindra Jemalong Gogo

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This cluster is also grouping with another distinct cluster, including JeCa, Greenland,

ScatCraig, RedHill and Belgium. The ScatCraig is in a basal position due to Heterostraci taxa. The rest of the cluster is defined by taxa as Antiarcha 2, Phyllolepida, Dipnoi 1, 3 and Tristichopterydae 1.

Then, there are two grouping: the Greenland and Jeca localities are together as RedHill and Belgium.

There is on additional cluster showing some similarity between Ellesmere, Lode, China, Ogre and Miguasha. The Miguasha locality falls at the base of this cluster certainly because of the presence of endemic taxa from this locality. Then there two grouping with Antiarcha 1 as common taxa:

Ellesmere and Lode group and Ogre and China.

Figure 22: Dendrogram showing faunal relationship between near-tetrapod and tetrapod localities during the Late Devonian for the first artificial level when Canowindra and Jemalong localities are considered as grouped.

0 1,6 3,2 4,8 6,4 8 9,6 11,2 12,8 14,4

0 0,1 0,2 0,3 0,4 0,5 0,6 0,7 0,8 0,9 1

Similarity Miguasha Ellesmere Lode Ogre China Greenland JeCa RedHill Belgium ScatCraig Andreyvka Pavari Oryol Gogo

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5.2.2 Second artificial level

In this first cluster analysis (Figure 23), in which Jemalong and Canowindra are treated as separate localities, presence/absence data for 90 artificial grouping (second level) resulted higher similarity indices between the various localities. Of course, a higher level gives by default a higher similarity index. With this level of grouping, Gogo fall in the ingroup. The y axis represents the similarity index and the x axis represents the number of localities.

The most distinct cluster, with rather high levels of similarity, includes Canowindra, Jemalong and ScatCraig. This locality falls at the base of the cluster. The rest of the cluster is composed by the Canowindra and Jemalong localities which are the closest localities with nearly the totality of taxa in common.

This cluster is also grouping with another distinct cluster, including Greenland, Andreyevka, RedHill, Belgium and Ogre. This Ogre locality fall at the base because of the presence of taxa Heterostraci C, Antiarcha C, Dipnoi D and Tetrapoda K. The rest of the cluster is composed by two grouping: the Greenland and Andreyevka localities are together as the RedHill and Belgium sites. This last part of the cluster is due to common taxa: Acanthodii B, Chondrichthyes D, Actinopterygii A and Dipnoi A. Then, there is the Pavari locality.

A cluster which shows similarity between Gogo and Oryol is also present. There are two additional clusters showing some similarity between Ellesmere, Lode, China and Miguasha. This cluster is due to some taxa as Antiarcha A and B, Acanthodii B, Actinopterygii D, Porolepiformes A, Tristichopterydae B. Then, there are two grouping: Ellesmere and Lode group and Ogre and China.

Moreover these localities should be considered as the outgroup for this analysis.

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Figure 23: Dendrogram showing faunal relationship between near-tetrapod and tetrapod localities during the Late Devonian for the second artificial level when Canowindra and Jemalong localities are considered as

separated.

In this second cluster analysis (Figure 24), in which Jemalong and Canowindra are treated as grouped localities, presence/absence data for 90 artificial grouping (second level) resulted higher similarity indices between the various localities. With this level of grouping, Gogo fall in the ingroup.

The y axis represents the similarity index and the x axis represents the number of localities.

The most distinct cluster, with rather high levels of similarity, includes Greenland,

Andreyevka, JeCa, RedHill and Belgium. This cluster is supported by Antiarcha A and B, Phlyctaenii, Phyllolepida Acanthodii A B, Chondrichthyes F, Actinopterygii A, Dipnoi A, B and P and

Tristichopterydae A. It is possible to distinguish two grouping: the RedHill and Belgium locality are together whereas the other grouping is composed by Andreyevka, Greenland and JeCa. This last one falls at the base of the grouping.

0 1,6 3,2 4,8 6,4 8 9,6 11,2 12,8 14,4 16

0 0,1 0,2 0,3 0,4 0,5 0,6 0,7 0,8 0,9 1

Similarity China Ellesmere Miguasha Lode Oryol Gogo Ogre Greenland Andreyvka RedHill Belgium ScatCraig Canowindra Jemalong Pavari

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This cluster is also grouping with another distinct cluster, including Greenland, Andreyevka, RedHill, Belgium and Ogre. This Ogre locality fall at the base because of the presence of taxa Heterostraci C, Antiarcha C, Dipnoi D and Tetrapoda K. The rest of the cluster is composed by two grouping: the Greenland and Andreyevka localities are together as the RedHill and Belgium sites. This last part of the cluster is due to common taxa: Acanthodii B, Chondrichthyes D, Actinopterygii A and Dipnoi A. Then, there are the Pavari, ScatCraig and Ogre localities which are not grouped with other localities.

The next cluster has as common taxa Arthrodira B, Ptyctodontida, Dipnoi E and G and Osteolepididae A. So, this cluster shows similarity between Gogo and Oryol. There is on additional cluster showing some similarity between Lode and Miguasha. This cluster shows some taxa in common as Actinopterygii D, Actinistia and Tristichopterydae B which could explain the position of these localities in the basal position.

Then, there is a cluster which shows similarity between the Ellesmere and China localities.

Moreover these localities should be considered as the outgroup for this analysis.

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Figure 24: Dendrogram showing faunal relationship between near-tetrapod and tetrapod localities during the Late Devonian for the second artificial level when Canowindra and Jemalong localities are considered as

grouped.

5.3 Third analysis

Ellesmere becomes the outgroup locality in this analysis because of the great number of plesiomorphic characters (Figure 25 and 26).

In this first cluster analysis (Figure 25), in which Jemalong and Canowindra are treated as separate localities, characters from phylogenies are used. With this analysis, Ellesmere becomes the outgroup locality due to the number of characters coded as 0. The y axis represents the similarity index and the x axis represents the number of localities.

The most distinct cluster, with rather high levels of similarity, includes Belgium, RedHill, Andreyevka, Greenland and Oryol supported by characters 1, 2, 5 and 6 from Antiarcha, 30, 33, 35, 36, 42 and 73 from Dipnoi, 8, 18, 22, 46, 47, 60 and 61 from Tristichopterydae and 50, 74, 80, 81, 92

0 1,6 3,2 4,8 6,4 8 9,6 11,2 12,8 14,4

0 0,1 0,2 0,3 0,4 0,5 0,6 0,7 0,8 0,9 1

Similarity Miguasha Lode Oryol Gogo Ogre Greenland Andreyvka JeCa RedHill Belgium ScatCraig Pavari China Ellesmere

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and 96 from Elpistostegalia/Tetrapoda.. At the base of this cluster, the Oryol locality is present. The rest of the cluster is supported by some common characters as 44, 70and 72 from Actinopterygii, 70 from Dipnoi, 6, 9, 11, 34, 50, 51 and 52 from Tristichopterydae and 36, 50, 68, 73, 111 and 112 from Elpistostegalia/Tetrapoda. Then, there are Greenland and Andreyevka which are not grouped with other localities. The grouping composed by Belgium and RedHill is supported by the character 2 from Phlyctaenii which is not present in the other group. This locality falls at the base of the cluster. The rest of the cluster is composed by the Canowindra and Jemalong localities which are the closest localities with nearly the totality of taxa in common.

This cluster is also grouping with another distinct cluster, including Lode, Miguasha, China, Canowindra and Ogre. A cluster with Lode and Miguasha is present and supported by the characters 9, 23, 31, 58, 70, 72 and 76 from Actinopterygii .The characters 1, 2, 5 and 6 from Antiarcha, 6, 9, 11, 22, 46, 47 50, 51 ,52, 60 and 61 from Tristichopterydae support the rest of this cluster is composed by China, Canowindra and Ogre. The China locality fall at the base of this group whereas the grouping Canowindra-Ogre is supported by characters 2 from Antiarcha, 30, 33, 35, 36, 42, 73 from Dipnoi and 8, 18, 34 from Tristichopterydae.

A cluster which shows similarity between Gogo and Oryol is also present. There are two additional clusters showing some similarity between Ellesmere, Lode, China and Miguasha. This cluster is due to some taxa as Antiarcha A and B, Acanthodii B, Actinopterygii D, Porolepiformes A, Tristichopterydae B.

The ScatCraig locality is in the basal position of the cluster composed by two other localities:

Jemalong and Pavari. This grouping is supported by characters 1 to 3 and 4 to 5 from Antiarcha, 30, 33, 35, 36 and 73 from Dipnoi, 50, 74, 77, 81 and 112 from Elpistostegalia/Tetrapoda.

Finally, the Gogo locality is in the basal position because of all characters are absent except characters coming from Arthrodira, Actinopterygii and Dipnoi and three characters from

Osteolepididae.

References

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