SUPPLEMENTARY INFORMATION

JOURNAL OF VERTEBRATE PALEONTOLOGY

Insights into the phylogeny and sensory capabilities of terror

FEDERICO J. DEGRANGE,1* CLAUDIA P. TAMBUSSI,1 MATÍAS L.

TAGLIORETTI,2 ALEJANDRO DONDAS,3 and FERNANDO SCAGLIA3

1CICTERRA, CONICET and Universidad Nacional de Córdoba, Av. Vélez Sársfield 1611,

X5016GCA, Córdoba, . [email protected];

2Instituto de Geología de Costas y del Cuaternario, Facultad de Ciencias Exactas y

Naturales, Universidad Nacional de Mar del Plata. CC 722, 7600, Mar del Plata,

Argentina;

3Museo Municipal de Ciencias Naturales Lorenzo Scaglia. Plaza España sin número,

7600, Mar del Plata, Argentina.

*Corresponding author.

APPENDIX S1. Character list used for the phylogenetic analysis of phorusrhacid

intrafamiliar relationships.

Skull and Mandible

(1) Neurocranium, relative size compared with total size of the skull: small (0); big (1).

(2) Upper beak, maxilla and premaxilla: wider than tall (0); taller than wide (1).

Character 1 of Alvarenga et al. (2011); character 3 of Mayr (2002).

(3) Upper beak, distal hook: absent (0); section with triangular shape and poorly developed ventrally (1); section with oval shape and great developed ventrally (2).

(4) Fonticulis occipitalis (Ericson, 1997): present (0); absent (1).

(5) Neurocranium, frontal region: pentagonal (0); triangular (1).

(6) Neurocranium, shape of the foramen magnum: subpentagonal (0); circular (1); oval

(2); triangular (3).

(7) Neurocranium, prominentia cerebellaris: blunt edge (0); sharp edge (1).

(8) Neurocranium, crista nuchalis transversa: absent (0); present, thin and sharp (1); stout

and rounded (2).

(9) Neurocranium, processus parasphenoidalis medialis with accessory tubercules:

present (0), absent (1).

(10) Upper beak, straight or weakly convex above the nares (0); markedly convex above

the nares (1). Character 4 of Agnolín (2009).

(11) Upper beak, nares shape: oval or kidney-shaped (0); triangular (1).

(12) Splachnocranium, zona flexoria craniofacialis: present (0); absent (1).

(13) Splachnocranium, zona flexoria palatina: present (0); absent (1). (14) Splachnocranium, zona flexoria arcus jugalis: present (0); absent (1).

(15) Cranial edge of the fenestra antorbitaria: oblique (0); vertical (1). Compare with

character 4 of Alvarenga et al. (2011) and character 8 of Agnolín (2009)]. Agnolín (2009)

defines both states as oblique against ‘straight’. This last term may be confusing.

(16) Cranial fusion of the lacrimals with the frontals: absent (0); present (1). Compare with character 6 of Alvarenga et al. (2011).

(17) Supraorbital process of lacrimals: short (0); caudally long (1). Compare with character 9 of Alvarenga et al. (2011).

(18) Supraorbital processes of the lacrimals in contact with the orbital edge of the

frontals: present (0); absent (1).

(19) Neurocranium, fossa temporalis, triangular in shape, meeting in the sagittal plane

dorsally: present (0); absent (1). Compare with character 7 of Alvarenga et al. (2011).

(20) Contact between the lacrimal and the jugal bar through the os lacrimale

communicans (present as an independent bone): present (0); absent (1).

(21) Jugal bar, height two times or more than the width: absent (0); two times as tall (1);

more than two times as tall (2). Compare with character 14 of Alvarenga et al. (2011).

(22) Upper beak, nares, prenarial fossa (=fossa premaxillar of Agnolín, 2009): present

(0); absent (1). Compare with character 6 of Agnolín (2009).

(23) Splachnocranium, medial fusion of the maxillopalatine process: absent (0); present

(1).

(24) Palate, fossa palatalis: absent (0); present (1).

(25) Palate, palatine crista lateralis: very extended ventrally (0); poorly extended

ventrally (1). (26) Palate, palatine rostral process: absent or poorly developed (0); present (1).

(27) Palate, basipterygoid articulation: present (0); absent (1). Compare with character 23 of Mayr and Clarke (2003).

(28) Pterygoid, with articulation process for the basipterygoid process: absent (0); present

(1). Compare with character 11 of Alvarenga et al. (2011).

(29) Quadrate, condyle number: two, caudal condyle untied with the lateral condyle (0); three, caudal condyle separated from the lateral condyle (1). Compare with characters 52 and 54 of Livezey (1998).

(30) Quadrate, pneumatic foramen in the medial surface of the orbital process: one (0); two (1); absent (2).

(31) Quadrate, caudal pneumatic foramen on the corpus: present (0); absent (1).

(32) Symphysis: ventrally curved (0); straight (1); dorsally curved (2). Compare with character 17 of Alvarenga et al. (2011).

(33) Rami mandibulae, mandibular fenestra: simple (0); double (1); triple (2).

(34) Fossa articularis quadratica, processus lateralis mandibulae: very developed (0); poorly developed (1).

(35) Processus retroarticularis: absent (0); present, caudolaterally directed (1); present, laterally directed (2).

Vertebral Column

(36) Third cervical vertebra, an osseous bridge linking the processus transversus to processus articularis (postzygapophysis) making a dorsal fenestra: absent (0); present (1).

Character 52 of Mayr and Clarke (2003); character 21 of Alvarenga et al. (2011). (37) Bifurcate neural spines in cervical vertebrae: absent (0); present (1).

Thoracic Girdle

(38) Sternum, spina externa: absent (0); present (1).

(39) Sternum, craniolateral process: present but poorly developed (0); present and very developed, describing a stout spine (1).

(40) Sternum, caudal margin with: one pair of incisions (0); without incisions (1).

Character 13 of Mayr (2002).

(41) Coracoid, procoracoidal process: developed (0); absent or poorly developed (1).

Modified from character 28 of Alvarenga et al. (2011). According to Alvarenga and

Höfling (2003) and Alvarenga et al. (2011), the procoracoidal process is absent in all phorusrhacids.

(42) Coracoid, acrocoracoidal process: present (0); absent (1). Character 29 of Alvarenga et al. (2011).

(43) Coracoid, an osseous bridge linking the acrocoracoidal and the procoracoidal process: present (0); absent (1). Character 6 of Mayr (2002); character 30 of Alvarenga et al. (2011).

(44) Coracoid, coracoid fused to clavicle: absent (0); present (1). Character 31 of

Alvarenga et al.[(2011).

(45) Scapula, acromion cranially projected: absent (0); present (1). Character 33 of

Alvarenga et al. (2011).

(46) Scapula, extremitas cranialis scapulae, group of foramina ventrally located to facies articularis humeralis: absent (0); present (1).

Forelimb

(47) Humerus, fossa m. brachialis: shallow (0); deep (1).

(48) Humerus, processus flexorius: present (0); absent (1). Compare with character 129

of Livezey (1998); character 39 of Alvarenga et al. (2011).

(49) Humerus, distal end strongly oblique in relation to longitudinal axis of the shaft:

absent (0); present (1). Character 17 of Mayr (2002).

(50) Humerus, incisura intercondylaris: present (0); absent or poorly developed (1).

(51) Humerus, fossa olecrani: absent or poorly excavated (0); present and deep (1).

(52) Ulna, greatly abbreviated, measuring only about three-fourths (or less) of the length

of the humerus: absent (0); present (1). Modified from character 18 of Mayr (2002).

(53) Carpometacarpus, Os metacarpale minus, more extended distally than the os

metacarpale major: present (0); absent (1). Modified from character 67 of Agnolín

(2009).

(54) Carpometacarpus, medial process located at the base of the os metacarpale minus:

present and distally projected (0); present and ventrally projected (1); absent (2).

Pelvic Girdle

(55) Preacetabular region: elongated and not compressed laterally (0); elongated and compressed laterally (1); compressed laterally but not elongated (2). Modified from character 24 of Mayr (2002) and character 43 of Alvarenga et al. (2011). Previous authors

(Mayr, 2002; Alvarenga and Höfling, 2003; Agnolín, 2009; Alvarenga et al., 2011) considered that all phorusrhacids have a long and compressed pelvis. However, the psilopterine phorusrhacid does not possess a fully compressed pelvis.

(56) Pelvis postacetabularly compressed laterally: absent (0); present (1).

(57) Crista iliaca dorsalis, preacetabular edge: straight (0); curved, forming a bow (1).

(58) Tuberculum preacetabulare: very developed (0); absent or poorly developed (1).

(59) Crista trochanterica: low (0); tall, very developed (1). Compare with character 44 of

Alvarenga et al. (2011).

(60) Supratrochanteric crest, lateral projection: more projected than the antitrochanter (0);

less projected than the antitrochanter (1).

(61) Iliac platform (sensu Rovereto, 1914): absent (0); present (1).

(62) Iliac shield (sensu Rovereto, 1914): absent (0); present (1).

(63) Ala ischii, caudal margin: ‘V’-shaped (0); bowed (1).

(64) Process caudally located to the ilioischiadic foramen: very developed (0); poorly

developed (1); absent (2).

(65) Pubis incomplete: absent (0); present (1). Character 46 of Alvarenga et al. (2011).

Hind Limbs

(66) Femur, trochanter majus: prominent proximally (0); absent or not prominent (1).

Character 49 of Alvarenga et al. (2011).

(67) Femur, crista trochanteris, relative height to the caput femoris in cranial view: equal

(0); more proximal (1); more distal (2).

(68) Femur, linea intermuscularis cranialis: well marked (0); absent or tenuous (1).

(69) Femur, fossa poplitea: shallow (0); deep (1). Character 50 of Alvarenga et al. (2011). (70) Femur, osseous bar uniting the lateral and medial condyli, limiting caudodistally the popliteal fossa: present (0); absent (1).

(71) Tibiotarsus, pons supratendineus with distal lip (sensu Degrange and Tambussi,

2011): present (0); absent (1).

(72) Tibiotarsus, tubercle laterodistally located to the pons supratendineus: present and very developed (0); absent or poorly developed (1).

(73) Tarsometatarsus proportions: less than 70% the length of tibiotarsus (0); between

70–80% (1); between 80–90% (2); more than 90% (3).

(74) Tarsometatarsus long and slender: the ratio of total length/width of middle of diaphysis is smaller than 12 (0); the ratio is greater than 12 (1). Character 55 of

Alvarenga et al. (2011).

(75) Tarsometatarsus, shaft width: constant along the whole shaft (0); distally narrowed

(1).

(76) Tarsometatarsus, medial and lateral cotyla: located at the same level (0); lateral cotyla more distal (1).

(77) Tarsometatarsus, furrow in the base of the eminentia intercotylaris: present (0); absent (1).

(78) Tarsometatarsus, triangular hypotarsus: absent (0); present and symmetric (1); present and asymmetric (with one of its flanges more projected laterally) (2).

(79) Tarsometatarsus, hypotarsus, proximal edges caudally projected (Alvarenga and

Höfling, 2003): present (0); absent (1).

(80) Tarsometatarsus, foramina vascularia proximalia, position: same level (0); different level (1). (81) Tarsometatarsus (dorsal view), trochlea metatarsi II: deflected medially (0); almost parallel to the trochlea III (1); articular surface extended medially (2). Character 59 of

Alvarenga et al. (2011).

(82) Tarsometatarsus, trochlea metatarsi II, with posteromedial projection: very developed (0); absent or poorly developed (1).

(83) Tarsometatarsus, trochlea metatarsi III, widened distally: present (0); absent (1).

(84) Tarsometatarsus, trochlea metatarsi IV, posterocaudal projection caudally directed: very developed (0); absent or poorly developed (1).

APPENDIX S2. Supplementary phylogenetic analysis: protocol and results.

Within Neornithes

In order to establish the phylogenetic position of phorusrhacids within Neornithes, we used the data matrix of Mayr and Clarke (2003) This proposal uses a reasonable number of characters, which are mainly osteological and easily understood which facilitates scoring. Livezey and Zusi’s (2006, 2007) work was criticized by Mayr (2008) because it includes more than 2,950 very complex characters, many of which are difficult to understand (and therefore hard to score), repetitive (in some cases), and in others not applicable to the fossil record (more than 500 characters are for soft tissue features).

To accomplish this analysis, the phorusrhacids Mesembriornis milneedwardsi,

Llallawavis scagliai, marshi, and lemoinei were chosen because they are represented by the most complete specimens. Also, the anseriform Brontornis burmeisteri was included in the analysis. This taxon was previously considered a member of by Alvarenga and Höfling (2003) and Alvarenga et al. (2011).

The matrix constitutes 148 characters and 51 taxa, including the four phorusrhacids and Brontornis burmeisteri. Of the 148 characters, 138 are binary, 10 are multistate, and 43 are polymorphic for determined taxa. Missing characters were coded as

‘?’ Polymorphic characters are enclosed with brackets (i.e., [01]). For cladogram construction, the principle of parsimony was followed (Hennig, 1968).

The analysis was performed using TNT (Tree analysis using New Technology), v.

1.1 (Goloboff et al., 2008). For the analysis, no implied weighting was used because its use is not exempt from criticism (Farris 1969, 1989; Goloboff, 1993; Kluge, 1997).

Characters 55, 71, and 91 were treated as ordered and, following Mayr and Clarke

(2003), the taxon Gaviidae (loons) was excluded because it produced polytomies in the consensus tree of Mayr and Clarke (2003).

The analysis was performed based on the random generation of 10 Wagner trees with 100 addition sequences, using the TBR algorithm and saving 100 trees per replicate, collapsing the trees after the search.

Fourteen trees of length 746 steps were obtained. In all of them, the

Phorusrhacidae constitutes a monophyletic group that is the sister taxon of Cariamidae, without a relationship with the rest of ‘.’ The sister group of Phorusrhacidae +

Cariamidae is Opisthocomidae and the sister group of this more inclusive clade is

Cuculidae + Musophagidae (Cuculiformes). Also, in all trees obtained, Brontornis was excluded from Phorusrhacidae and included among (waterfowls), constituting the sister group of Anhimidae (screamers). The consensus tree (Fig. S1) is 806 steps in length and has a consistency index

(CI) of 0.20596 and a retention index (RI) of 0.45399. In this tree, and contrary to Mayr

and Clarke (2003:fig. 3), the great majority of the relationships among Neoaves are not

resolved (only two of the 14 individual trees are similar to the relationships among

Neoaves found in the consensus tree of Mayr and Clarke [2003]).

According to the analysis, Cariamidae + Phorusrhacidae (node 68)—defined here

as —is characterized by the following unambiguous synapomorphies: (11)

processus maxillopalatini fused along its midline (i.e., palate directly desmognathous:

this character is discussed below); (12) the descending process of the lacrimal bone does

not contact the jugal bar; (13) supraorbital process of the lacrimal bone caudally

projected; (21) vomer forming a medial lamina that is narrow and tall; (83) proximal end

of the ulna dorsoventrally compressed and cranioventrally flexed; (103) hypotarsus

without well-developed cristae/sulci; and (110) hallux strongly reduced.

Phorusrhacidae (node 85) is characterized by the following synapomorphies: (2)

premaxila ended in a hook; (23) basipterygoid articulation present; (24) basipterygoid process with a large and ovoid facet for articulation with the pterygoids; (31) pila otica without small pneumatic foramina; (53) osseous bridge that links the costal process with the medial part of the vertebral body present, at least in the 7th and 8th cervical vertebrae;

(73) caudal margin of the sternum without notch or fenestra; (74) uncinate process

absent; (81) humerus scapulotricipital sulcus poorly developed; (84) ulnar radial depression well marked; (90) pelvis very elongated and strongly mediolaterally compressed; (93) preacetabular tuberculum developed; (101) presence of a tubercle laterodistally located to the supratendinal bridge of the tibiotarsus; and (102) distal

margin of the tibiotarsus medial condyle markedly excavated.

Supplementary Discussion

Phorusrhacidae constitutes a monophyletic group and sister taxon of the

Cariamidae, a position advocated by previous authors (Brodkorb, 1967; Cracraft, 1968,

1971; Mourer-Chauviré, 1983; Alvarenga and Höfling, 2003; Agnolín, 2009; Alvarenga

et al., 2011). Also, we corroborate the relationship of Brontornis to Anseriformes in this

cladistic analysis as proposed by Agnolín (2007) (contra Alvarenga et al., 2011).

However, our results suggest that it was more closely related to screamers (Anhimidae)

than to ducks (Anatidae).

There is consensus among different authors (Patterson and Kraglievich, 1960;

Mayr, 2002; Alvarenga and Höfling, 2003; Agnolín, 2009; Alvarenga et al., 2011)

regarding some of the possible synapomorphies of phorusrhacids. These are: maxilla

dorsoventrally deep and strongly compressed mediolaterally, endowed with a hook on its

distal end; basipterygoid process well developed; pterygoid with a well developed

articular facet on its medial portion that contacts the basipterygoid process;

acrocoracoidal process reduced; flexor process very developed; uncinate processes

absent; and pelvis strongly compressed mediolaterally (among others). Several authors

have particularly noted the presence of a desmognathous palate as a synapomorphy

(Patterson and Kraglievich, 1960; Alvarenga and Höfling, 2003; Agnolín, 2009;

Alvarenga et al., 2011), establishing that the Cariamidae () have a

schizognathous palate (Zusi and Livezey, 2006; Agnolín, 2009). However, Mayr and Clarke (2003) proposed that Cariamidae have a desmognathous palate, because the

maxillopalatine processes are united and fused along its midline (character 11 of Mayr

and Clarke, 2003). Revision of cranial material of Cariamidae establishes that seriemas do not have maxillopalatine medial fusion, although these bones do contact each other.

Following Huxley’s (1867) definition, seriemas do have a desmognathous palate because the maxillopalatine processes are very well developed and unite (though not necessarily through fusion) along its midline. This kind of palate clearly stands out from the schizognathous palate defined by the same author and typically found in Charadriiformes,

Columbiformes, and some ‘Gruiformes,’ such us Psophia, Grus, Otididae, and Eurypyga.

Agnolín (2009) misinterprets the presence of a vomer in Cariama as typical of a schizognathous palate. We verified for the first time the presence of the vomer in

Llallawavis; it could be present also be in Patagornis marshi NHMUK A516 (Degrange,

2012).

Van Remsen’s proposal to the South American Classification Committee

(Proposal 290, approved in October 2007) suggested that extant seriemas are in

Cariamiformes. Degrange and Tambussi (2011) proposed the use of Cariamiformes

Verheyen, 1957 to include Cariamidae and Phorusrhacidae, removing them from the order ‘Gruiformes’. This is in agreement with the idea that ‘Gruiformes’ is a polyphyletic group (Mayr and Clarke, 2003; Cracraft et al., 2004; Fain and Houde, 2004; Fain et al.,

2007). Cariamiformes might also include the fossil taxa Paleopsilopterus,

Elaphrocnemus, Salmilidae, Bathornithidae, Ameghinornithidae, and Idiornithidae, all of which are related to Cariamidae (Mourer-Chauviré, 1981, 1983; Mayr, 2002; Agnolín, 2009; Alvarenga et al., 2011) although a new and more inclusive phylogenetic analysis is

necessary.

In this context, a synapomorphy of Cariamiformes is the presence of a desmognathous palate. Another derived feature of this clade is the presence of the ossification of the lacrimojugal ligamentum, named the ‘os lacrimale communicans’ by

Burmeister (1854), who highlighted that this was a unique and exclusive feature of the seriemas. Degrange (2012) pointed out that the os lacrimale communicans is also present

as an independent bone in Psilopterinae and Mesembrionithinae and fused to the

descending process of the lacrimal in the other terror birds.

The definition of the synapomorphy in relation to the palate of the Phorusrhacidae

is here emended in the following way: phorusrhacids have a craniocaudally extended

medial fusion of the maxilloplatine processes (i.e., directly desmognathous palate).

SUPPLEMENTARY FIGURE

FIGURE S1. Consensus tree obtained from analysis of the character matrix of Mayr and

Clarke (2003), with the addition of four of Phorusrhacidae and the anseriform

Brontornis. Characters 55, 71, and 91 were treated as ordered and the taxon Gaviidae was excluded. Tree length = 806 steps, Consistency Index = 0.20596, Retention Index =

0.45399.

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