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.
Supervisor: Henning Blom Examensarbete vid Institutionen för geovetenskaper
ISSN 1650-6553 Nr 301
Fish/tetrapod Communities in the Upper Devonian
Maxime Delgehier
Copyright Maxime Delgehier and the Department of the Earth Sciences Uppsala University Published at Department of Earth Sciences, Geotryckeriet Uppsala University, Uppsala, 2014
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
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
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
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
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).
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
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
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
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
(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
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
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
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
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
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
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
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Laccognathus, Glyptolepis and Tiktaalik) show characters in plesiomorphic states.
5. Results
5.1 First analysis
5.1.1 Species levelIn 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.
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
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
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.
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
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
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.
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
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
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.
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
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.
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
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.