Phylogeny of the Tertiary Giant Anhingas (Pelecaniformes

Phylogeny of the Tertiary Giant Anhingas
(Pelecaniformes: Anhingidae) from South
America
Jorge I. Noriega and Herculano M. F. Alvarenga 1
CICyTTP-CONICET, Matteri y Espafi a, 3105 Diamante, Argentina;
email: [email protected]
1 Museu de Historia Natural de Teubeie, Rua Colombia 99, 12030-520 Teubete,
Brasil; email: [email protected]
and their propulsi on in water consists of swimming very
low on the surface and slow dives to prey on fishe s
(Owre, 1967).
Th e fossil record of the famil y has been reviewed
by Ol son (198 5) , Becker (1986, 1987), Rasmus sen
and Ka y (19 92), Noriega (1994), Alv arenga (1995),
and Campbell (1996). Until recently, fossil darters
were mainly kno wn from North Am erica, Europe, and
Africa. In th e last decade, des cripti ons of three
paleospecies of large-bodied anhin gas (Noriega,
1992 , 1995 ; Al var en ga , 1995 ; Campb ell , 1996)
revealed a great diversity of these birds in the Tertiary
of South A merica. However, knowl edge about the
phylogenetic relationships amon g them remains rather
unclear.
In this pap er we present a hyp othesis about the
phylogeny of th e Miocene da rters Ma cranhinga
paran ens is Nori ega, 1992, Me ganh ing a chilens is
Alvarenga, 1995, and Anhingafraileyi Campb ell, 1996,
based m a inl y o n the mo rpholog y of th e
tarsometatarsus, the pelvis, and the inferences about
their capacities of flight.
Abstract
Recent findings of paleospecies of large­
bodied
darters
(Pelecaniformes:
Anhingidae) have revealed a great div er­
sity of these birds in the Tertiary of South
America. However, knowledge about the
phylogenetic relationships among them has
been rather unclear. An hypothesis of the
phylogen y of the Miocene darters
Macranhinga p aranensis Noriega , 1992,
Meganhinga chilensis Alvarenga, 1995, and
Anhinga fraileyi Campbell, 1996, based
mainly on the morphology of the tarsometa­
tarsus, the pelvis, and the inferences about
their capacities of flight is presented.
Introduction
Anhingas, also known as darters, are pelecaniform
bi rd s belonging to the family Anhingidae of the
subo rder Sulae, which also comprises the families
Sul ida e, Phalacro cor acid ae , and the ext in ct
Plotopteridae. They are fairly large water birds with
very long and slende r necks and pointed bills. Anhingas
alternate flapping flight with soaring glide s in therm als,
Materials and Methods
Fossil materials analyzed herein are housed in the
collections ofthe Museo Argentino de Ciencias Naturales
Bernardino Rivada via (MACN), Museo de La Plata
41
Proceedings of the 5th Symposium of the Society of Avian Paleontology and Evolution
42
(MLP), both in Arge ntina, and M useo Nacion al de
His toria Natur al de Sa ntiago de Ch ile (SG O-PV) .
They include three tarsom etat arsi (MACN 1350 7,
ho!otype, fig . Id; both paratypes, MACN 122 81 and
12293) and three pe lvic girdles (M LP 88-IX-20 -5 ,
fig. 3d; 4 1-XII-13-929; 88- IX-20- 15) of Macran hinga
1°
0 ..'\
pa ranensis; a tar somet atarsu s (SGO-PV 400 I- A ;
fig. lb) and two pelvic gird les (SGO-PV 4001-B and
400 2; fig . 3c) of Meganh inga chilensis. Comparison s
with Anhinga fra ileyi were based on a cast of the
holotyp e (Na tura l History Muse um of Los An ge les
County, LACM 135356; fig . 1e).
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Editors: Zhonghe Zhou and Fucheng Zhang. Beijing: Science Press. 2002
O steolo gical comp ari son s w e re mad e with
s ke le to ns of three s pe c ime n s of th e li vin g
Phalacro corax bra sil ianus (Gme lin, 1789), three of
Phalacrocorax carbo ( Li n na eu s, 175 8) , on e of
Phalacrocorax auritus (Lesson, 1831), six of Anhinga
anhinga (Linnaeus , 1766), two of Anhin g a
nova ehollandiae Penn ant , 1769, one of Anhinga rufa
Daudin , 1802, and one of A nhinga melanog aster
Gould , 1847 (Appendi x 1).
The principles of ph ylo g enetic s ystemat ics
follo wed in this study are those described by Henn ig
(1 968), Wiley ( 198 1), and E ld redg e and Crac ra ft
(1980). Hypotheses of cha racter-state pol ar ity are
proposed based on out- group comparison (Watrou s and
Wheeler, 1981; Maddison et al., 1984).
Cladistic Analysis
Th e ph ylo gen etic ana lysis was performed usin g
Henni g86 pro gram ve rs io n 1.5 ( F a rris, 198 8 ),
employing the implicit enumeration option (ie), which
ensures that minim al length trees are found .
A total of 2 1 equa lly weighted ch aracters were
selected: seventeen fro m the tarsometatarsu s, three
from the pelvic girdle, and one related to the capacity
of flight (Appendix 2). Three characters have multiple­
step transformations and they are treated as additive .
The matri x of character states and taxa is given in
Appendi x 3. Dat a that are missing because of the
fragmentary state of pre servation of fossil mater ials
.are indicated by question marks. The matrix includes
four terminal taxa plu s two outgroups: the form er are
the three Miocene paleospecies of giant dart ers plus
Anhinga anhinga.
The genu s A nh inga is assumed to be monophyletic ,
this fact being c onfirmed by the strong ske leta l
resemblance among all its modem spe cies compared.
FIGURE 1. Tarsometatarsi of (a) Phalacrocorax
carbo (MHNT 989), (b) Meganhinga chilensis
(SGO-PV-4001-A), (c) Anhinga anhinga (MHNT
1039), (d) Ma cranhinga paranensi s (MACN
13507), and (e) Anhinga fraileyi (Cast of LACM
135356) in anterior and posterior views; (a), (b),
and (c) modified from Alvarenga (1995); (e) modi­
fied from Campbell (1996).
43
Based on this ass um ption, the same hyp otheses of
character state pol arit y were established for all the
living form s of Anhinga, and Anhinga anhinga was
sele cted as rep resent ative of them because it is resident
in Americ a . The liv ing species Ph al a cro corax
brasilianus and Phalacrocorax auritus ( Phalacro ­
coracidae) were chosen as the outgroups to polarize
the characters be cause the Phalacro cora cidae is
traditionally c o ns ide r ed the sister- group of the
Anhingidae (O lson , 1985 ; Becker, 19 86 ; Si ege l­
Causey, 1988).
Results
The cladi stic analysis results in onl y one tree , with
a total length of 31, Consistency Index (C.L) of 77,
and Retention Inde x (R.L) of 68 (Fig . 4).
Three syn apomorphies shared by all members of
the famil y Anhingidae support their monoph yly (Node
A: 11,8 1, 131; fig. 4) and separate them from the spec ies
of Phalacrocoracidae, whi ch composes the out group
taxa. The third calcaneal ridge of hypo tarsus (Character
1) evol ves from an incipient (Node A: 11 ; figs. Ib, 2c)
to a well ped iceled tubercle (Node B: 12; figs . 1c, d,
2a, b) within darters. The other two derived characters
of the tar som etatarsus of the Anhin gidae are the
presence of a cons picuous internal mar gin of the
anterior metatarsal groove (8 1) , and the cotyla lateralis
with a less pronounced slope anteriad than cormorants
(13 1) .
M eg anh inga chilensis appears as the most basal
taxon among the selected species of darters , close to
the cormorants. Meganhinga chilensis is characterized
by one autapomorphy (21 1) , which is its flightlessness.
Alvarenga (199 5) inferred that Meganh inga chilensis
was flightl ess based on the morphology of its wing
bones and the different proportions that it had between
the fore - and hind limbs than modern s pec ies of
Anhinga .
Ma cranhinga paranensis and A nhin ga fraileyi are
sister taxa and, together with A nhinga anhinga , they
form a clade that is the sister group of Meganhinga
chilensis . This group is defined by three
synapomorphies (Node B: 12,2 1,14 1; fig. 4): a) the
first calcaneal rid ge of hypo tarsus less projected
posteriorly than in Meganhing a chilensis and in the
outgoups (2' ; figs. 2a, b). Regardless of the fact that
Proceedings of the 5th Symposium of the Society of Avian Paleontology and Evolution
44
12
21
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Editors: Zhonghe Zhou and Fucheng Zhang. Beijing: Science Press. 2002
45
E
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FIGURE 3. Pelves of (a) Phalacro corax brasilianus (MHNT 232), (b) Anh inga anhinga (MHNT 1039), (c)
Meganhinga chilensis (SGO-PV-4001-B and 4002), and (d) Macranhinga paran ensis (MLP-88-IX-20-5) in
dorsal views; (a), (b), and (c) modified from Alvarenga (1995).
the hypo tarsus is missing in Anhinga fraileyi, the
polarity of this character state was inferred by
analyzing the morphology of its well preserved
hypotarsal ridge (crista medianoplantaris) . Crista
medianoplantaris in Anhinga fraileyi is developed as
a low ridge from level of foramina vascularia
proximalia medialis distad (much more prominent
ridge in Macranhinga paranensisi and more distally
extended than in Meganhinga chilensis (see Campbell,
1996 for detailed comparisons). Both characters show
that the crista medianoplantaris of the hypotarsus in
Anhinga fraileyi meets the shaft smoothly, at a higher
angle (more open) than in Meganhinga chilensis and
FIGURE 2. Tarsometatarsi of (a) Macranhinga
paranensis (MACN 13507), (b) Anhinga anhinga
(MHNT 1039), (c) Meganhinga chilensis (SGO­
PV -400 I-A) , and (d) Phalacro corax carbo
(MHNT 989) in cranial and medial views; (b), (c),
and (d) modified from Alvarenga (1995).
cormorants, as it would be expected for the presence
of the first calcaneal ridge with lesser posterior
projection; b) the proximal end of shaft wider in
anterior view, narrowing less distal to cotylae than in .
Meganhinga chil ensis and the outgoups (14 1; figs. le­
e). See Campbell (1996) for detailed description of
this character in Anhinga fraileyi; and c) the more
derived state of character 1 cited above.
Anhinga anhin ga, as a representative of the living
genus, is defined by three synapomorphies (17 1, 19',
20 1) : a) the impression of the M. extensor hallus longus
origin well developed on anterior aspect, being only
insinuated on medial view (17 1; fig. 2b); b) the cranial
portions of the posterior iliac crests more divergent
caudally on dorsal aspect. Consequently, the post­
acetabular region of the pelvis is wider (19 1; fig. 3b);
whereas these crests are almost parallel caudally in
the primitive character state and, consequently, the
post-acetabular region of the pelvis is narrower (Figs.
3a, c, d) ; and c) the post-trochanteric process situated
at the level of the acetabulum on dorsal aspect and
46
Proceedings of the 5th Symposium of the Society of Avian Paleontology and Evolution
more laterally (20 1; fig. 3b). The plesiomorphic state of
thi s character consists in the presence of the post­
trochanteric process at a more caudal and medial position
to the acetabulum on dorsal view (Figs. 3a, c, d).
The monophyly of the clade composed of
Macranhinga paranensis and Anhinga fraileyi is
supported by four derived ch aracters (Node C: 51,
7 1,9',12'; fig. 4): a) the angle between the hypo tarsal
ridge (crista medianoplantaris) and the shaft is high
(more open), meeting both smoothly (5'; figs. l e, 2a) ;
this condition determines that the fossa
parahypotarsalis medialis is larger and more extended
downshaft than that of Anhinga anhinga and
Meganhinga chilensis ; see Campbell (1996) for
detailed description of this character in Anhinga
frail eyi ; b) the groove where M . extensor hallus
longus passes from the anterior to the posterior
surface is situated on medial border of the shaft
distally, i.e. at least on midshaft or more distal part
of shaft (7 1; figs. Id, e); c) the broad midshaft region,
widening more as it approaches trochlea metatarsi II
(9 1; fig s. l d, e); and d) the broad and short trochlea
III (12 1; figs . I d, e) are the remaining syn apomorphies
that define this clade.
Anhinga fraileyi is characterized in this cladistic
analysis by presenting the autopomorphy of a high and
narrow eminentia intercondylaris (4 1) . Other characters
interpreted as specific, e.g. the orientation of trochlea
III , are listed in the original diagnosis (Campbell,
1996).
Discussions
Campbell (1996) considered Meganhinga to be a
junior synonym of Anhinga, arguing that the
characters used by Alvarenga (1992) to erect the
genus are typical of a larger version of a foot­
propelled diving bird . In addition, Campbell (1996)
rejected the incapacity of flight of Meganhinga as a
valid generic character based on the fact that the
Phalacrocorax brasilianus
Phalacrocorax auritus
Meganhinga chilensis
A
Anhinga anhinga
B
Macranhinga paranensis
C
Anhinga fraileyi
FIGURE 4. Cladogram with the non-ambiguous synapomorphies supporting each node; Character state indi­
cated as super-index. Node A: 11,8 1, ]3 1; Node B: 12,2 1,14 1; Node C: 5' ,7 1,91, 121•
Editors: Zhonghe Zhou and Fucheng Zhang. Beijing: Science Press. 2002
variation between fore- and hindlimb lengths
observed in this taxon may be common in anhingas .
He added that the latter may be a situation analogous
to that seen in cormorants, where only one species of
Phalacrocorax is flightless , whereas the remaining
species are volant. However, the cladogram obtained
in this analysis shows that the first divergence splits
Meganhinga chilensis from the remaining species as
a separate taxon. Then, the recognition of a separate
genus is largely justified if these ch aracters , i.e. the
morphology of the first and third calcaneal ridges of
hypotarsus of Meganhinga and its flightlessness , are
analyzed in a phylogenetic context. Moreover, the
same argument of flightlessnes s may be used
conversely to validate the genus Meganhinga if the
alternative systematic hypothesis, supporting that the
nonvolant cormorant of the Galapagos Islands
belongs to a genus (Nannopterum) distinct from
Phalacrocorax, is considered (see Witherby et al.,
1940; Ono, 1980; Siegel-Causey, 1988).
It must be noted that three of the four
synapomorphies defining the clade composed of
Macranhinga paranens is and Anhinga fraileyi (Node
C: 5 1, 7 1,12 1) are included in the original diagnosis of
the genus Macranhinga (Norieg a, 1992). These
characters, together with those described of the femur,
tibiotarsus , pelvic girdle , humerus , and
carpometacarpus (Noriega, this volume), justify the
validity of the genus Macranhinga that was recently
considered to be a junior synonym of Anhinga
(Campbell, 1996).
Finally, the cladistic analysis supports the
taxonomic revision of Anhinga fra ileyi. We consider
Anhinga fraileyi to be a valid species, but it probably
should be included in the genus Ma cranhinga. A set
of characters, not only of the tarsometatarsus, but from
the humerus (i.e . the morphology of the margo
caudalis, fossa pneumotricipitalis, impressio M.
coracobrachialis cranial is, attachment ofM. pectoralis,
processus flexorius , and the olecranal fossa) and
tibiotarsus (i.e. the massive shaft and the proximal
morphology of groove for peroneus profundus) of the
paleospecies described by Campbell (1996), are shared
with Macranhinga paranensis (Noriega, this volume).
However, until new and better preserved fossil
materials of Anhinga fraileyi are found, it is advisable
to maintain its present status.
47
Conclusions
1) Meganhinga chilensis is the most basal
paleospecies within the Miocene radiation of giant
darters.
2) Macranhinga paranensis and Anhinga fraileyi
are sister taxa, conforming a well defined clade.
3) Macranhinga and Meganhinga are valid genera,
distinct from the extant genus Anhinga.
4) Anhinga fraileyi is a valid species, but it must
be revised to determine its pos sible inclusion in the
genus Macranhinga.
Acknowledgments
We express our gratitude to the organizing com­
mittee of the 5th International Meeting of SAPE
for providing us the opportunity to participate in
the symposium, as well as for their special atten­
tion to our case. We are grateful to K. E. Campbell
for providing a cast of the holotype of Anhinga
fraileyi. We thank Dr. Jose Bonaparte (Museo
Argentino de C ien cias Naturales Bernardino
Rivadavia) and curatorial personnel of the Museo
de La Plata for granting acces s to fossil specimens;
and CONICET for partial financial support of this
research study.
Literature Cited
Alvarenga, H.M.F., 1995. A large and probably flightless
Anhinga from the Miocene of Chile. In Peters,
D.S., editor, Acta palaeomithologica, third Sym­
posium SAPE; 5 Internationale Senckenberg­
Konferenz 22-26 Juni 1992. Courier
Forchungsintitut Senckenberg, 181: 149-161
Becker, J.J., 1986. Reidentification of "Phalacrocorax"
s ubvo la ns Brodkorb as the earliest record of
Anhingidae. Auk, 103: 804-808
Becker, J.J., 1987. Additional material of Anhinga grandis
Martin y Mengel (Aves: Anhingidae) from the late
Miocene of Florida. Proc. Biol. Soc. Washington,
100(2): 358-363
Campbell, K.E ., 1996. A new species of giant Anhinga
(Aves: Pe1ecanifonnes: Anhingidae) from the up­
per Miocene (Huayquerian) of Amazonian Peru.
Nat. Hist. Mus . Los Angeles Co., Contrib. Sci. , 460:
1-9
48
Proceed ings of the 5th Symposium of the Society of Av ian Paleontology and Evolut ion
Eld red ge , N . and 1. Crac raft, 1980 . Phylogeneti c Pat­
terns and the Evo lutionary Process. 349 pages.
New York: Co lumbia Univers ity Pre ss
Farris, 1.S., 1988. Henn ig 86 Program and document a­
tio n fo r ve rsion 1.5
Henn ig , W., 19 6 8 . Elementos de un a Siste mati ca
Filogenetica . 353 pages. Bue nos A ires : Eudeba
Mad dison, w.r, M J . Donohu e and D .R. Maddison, 1984.
O utgroup analysis and pa rsimony . Sy st . Zo o!.,
33( 1): 83- 103
Noriega, J.I., 1992. Un nuevo ge nero de Anhingidae (Aves:
Pe leca niformes) de la Fo rmaci6 n Itu z ai n g6
(Mioceno superior) de Argentina. No tas del Museo
de La Plata, 109: 2 17223
Noriega, J. I., 1994 . Las Aves del "M esopotamiense" de
la provincia de Entre Ri os, Argentina. Facultad de
Cie ncias Natu rales y Museo, Univers idad Nacional
de La Plata, Argentina , Ph.D. dissertat ion , No . 61 1.
162 page s
N oriega , J .1., 19 9 5. The avifauna fro m the " Meso­
potam ian " (Ituzai ng6 Form ation; Upp er Mi ocene)
o f E n tre Ri os P rovin ce , Arge n tina. Co urier
Forchu ngsinst itut Senckenberg, 181 : 141-1 48
O lson, S.L., 1985. The fos sil record of bird s. In Fa nner,
D.S., J.R. Ki ng and K.C. Parkes, editors, Avian
Bio!., 8: 79-256. New York: Academ ic Pre ss
O no, K., 1980. Compa rative osteo logy of three spec ies
o f J ap an e se Cormora n ts of th e ge n us
Phalacrocorax (Aves, Pelecanifonnes). Bull. Na­
tional Sci . Mus., Tokyo, 6(4): 129- 151
Owre,O.T., 1967. Adaptations for locomotion and feed­
ing in the Anhinga and the Do uble -Crested Cor­
morant. Ornithol. Mo nogr., 6: 1-138
Rasmussen , D.T. and R.F. Kay, 1992 . A Miocene Anhinga
fro m Colomb ia, and comments on the zoogeo­
graphic relation sh ips of South America 's Tertiary
av ifau na . In Campbell, K .E ., edi tor, Pap ers in
Avian Paleontology honoring Pierce B rodkorb.
Na t. Hist. Mu s. Los Ang eles Co., Sci. Series 36 :
225-230
S iegel-Causey, D ., 19 8 8. Phylogeny of th e
Phalacrocoracidae. Condo r, 90: 885-905
Watrous, L., and Q .D. Wh eeler, 1981. The out-group com ­
pari son meth od of chara cter analys is. Sys t. Zoo i.,
30(1) : 1-11
Wiley, E.O ., 198 1. Phyl ogen etics: The Theo ry and Prac­
tice ofPhy logenetic Systemat ics. 439 pages. New
York : J. Wiley and So ns
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1940 . Handbook Br itish Birds, 4 : 1-14
Appendix 1: List of Specimens Used in
the Osteological Comparisons
Phala crocorax brasilianus (M use u d e Hist6 r ia
Natura l d e T aub at e, M HNT 26, 232; Ce n t ro d e
In v e s t ig ac ion e s Cie n tificas y T ra nsfe re ncia d e
Tec no logia a la P roducci6n de Di am ant e, C ICyTTP
152), Pha facro corax ca rbo (MHNT 989, 113 4 , 1190),
Pha lacrocorax au ritu s (M HNT 63 5) , Anhinga anhinga
(M HNT 2 5, 882, 92 4, 10 39 ; Smithso n ian Institution ,
Natural Museum ofNatu ra l Hi story, Di vi sion of B ird s ,
NMNH 182 59; Mu seo Naciona l de Hi st oria Natu ra l
d e P ara gu a y, M NH NP u nc at al o gu ed s peci men),
A n h ing a n o va eh o l la nd ia e (M HNT 1210 , 1195 ) ,
A nhinga rufa ( N MN H 4 30 6 80 ) , a n d An hi ng a
melan og aster (NMNH 488772 ).
Appendix 2: List of Characters Used
in the Cladistic Analysis
1. T hird calcanea l ridge of hypotarsu s: a low and small
prominence (0); an inci p ient raised and flatt ened tu­
bercle (1); a we ll pediceled tubercle (2) .
2. First calcaneal ridge of hyp otarsu s: well projected
posteriorly (0); less projected posteriorly (1) .
3 . Small fossa above trochlea III, on anterior view:
present (0); absent ( 1).
4. Eminentia intercondylaris: low and broad (0); high
and narrow ( 1).
5. Angle between hypo tarsal ridge and shaft: low (closed),
meeting both abruptl y (0); high (open), smoo thly (1).
6. Lateral hypota rsal crest: hardly pron ou nced and rathe r
straight (0); pronou nced and co nvex (1); p ronoun ced
and less co nvex (2) .
7 . Positi on at medi al border of shaft of groove for pass­
ing M . extenso r hallu s longus from anterior to poste­
rior surface: proxim al to midshaft (0) ; mid- or more
distal shaft ( 1).
8. Intern al ma rgi n of anterior me tatarsal groove : low,
less marked (0); high , co nspicuous (1).
9. M ids ha ft region : narrow (0) ; broad (1) .
10. Media l expa ns ion of medial side of shaft at the fossa
metatarsal I: not pro nounc ed (0); pro nounced (1).
II. Distal extension of trochl ea II: sho rter than trochl ea
III (0); equal or longer than trochl ea III ( I) .
Editors : Zhonghe Zhou and Fucheng Zhang, Beijing: Science Press. 2002
•
12. Trochl ea III: nan-ow and ex ten ding more anteriad
(lo nger), wit h thin troch lear rims and central groove
(0); broad and extendi ng less anteriad (shorter), with
w ide trochle ar rims and central groo ve (1) .
13. Cotyla lateral is: slopi ng steepl y anteria d (0); less pro­
nou nced sloping ( 1).
14. Narrowing of proximal end of shaft distal to coty lae
in anterior view : we ll marked (0) ; less pronounced
(1).
15. Spatial relati on betwe en pro ximal lateral ridge and
dista l end of crista me dioanop lantaris: not meeting
(0); mee ting at low ang le (1); meeting at hig h ang le
(2) .
16. Proximal mo rphology of shaft, in medial view : curv ­
ing strongly craniad to meet lip of cotyla medialis
(0); curv ing sligh tly (1).
49
17. Impression of the M. extensor hallus longus origin:
we ll deve loped on medial aspect (0); well developed
on anterior aspect , on ly insinuated on media l aspect
(1).
18. Hyp apo physis of the synsacral thoracic and lum bar
vertebrae: present (0); absent (1) .
19. Cranial portions of the posterior iliac crests on dorsal
aspect: almost parallel caudally. Consequen tly, the
pos t- ace tabular region of the pelvis is nan-ower (0);
divergent caudally. Consequently, the post-acetabul ar
reg ion of the pelvis wider (1).
20 . Position of the post- trochanteric process on dorsal
aspect: caudal and medial to the ace tabulum (0) ; at
the level of the acetabulum and more lateral ( I).
21. Capacity of flight: pres ent (0); absent (1).
Appendix 3: Matrix of Character States and Taxa
•
~
Taxa
Characters
Phala crocorax bra silia n u s
Ph a l acrocorax aur i t u s
An h inga anhing a
Mega nh i nga ch i l e ns is
Macranh i n g a pa r anens is
Anhinga fr a i l e yi
10
00000000 0 0
00 0 0 0 00000
21 100 20 0 0 1
1 01 0 01 0100
2 1 10 1 1 1 1 1 1
? 1 1 1 12 1 1 1 0
20
0 00 0 0 0 0 000
000 0 0 0 0000
1 0 01 21 1011
? 01 0 2 0 0000
0 1 11 1 10 100
1111 100 ? ??
0
0
0
1
0
0