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SUPPORTING INFORMATION
Evolution of human–ape relationships remains open for investigation
John R. Grehan and Jeffrey H. Schwartz
Journal of Biogeography
Appendix S2 Explanations of evidence for the removal of characters from the original data
matrix of Lehtonen et al. (2011) as shown in Appendix S1. Removal of characters is
justified on the following criteria: (a) one or more character states are not supported by
documentation, (b) the ingroup character state overlaps the outgroup values, (c) the
ingroup value lies within the outgroup values, (d) the ingroup value lies both above and
below outgroup values. With the overlapping or encompassing range of values there is no
objective way to code the ingroup character states with respect to the outgroup. When
characters included by Lehtonen et al. (2011) comprised two different types of character
states we separated these into separate characters.
(a) Strait & Grine (2004) craniodental qualitative data
Invalid SG7
Nasal cavity entrance. Stepped/overlap = 0; smooth/overlap = 1, smooth/no overlap = 2;
stepped/no overlap = 3
Pongo
Human
Pan
Gorilla
Hylobates
Papio
1
2
0
0
3
3
Character states composite. Separated out provide for the following:
Smooth (1) vs stepped (0)
Pongo
Human
Pan
Gorilla
Hylobates
Papio
1
1
0
0
0
0
and:
Overlap (1) vs no overlap (0):
Pongo
Human
Pan
Gorilla
Hylobates
Papio
1
0
1
1
0
0
We do not include either character as the human condition is autapomorphic so the smooth nasal
floor may not be a shared derived condition. The overlapping premaxilla and maxilla is
characteristic of large bodied hominoids other than hominids. See discussion by Schwartz
(2005).
Invalid SG26
Postorbital constriction. Marked = 0, Moderate = 1, Slight = 2
Pongo
Human
Pan
Gorilla
Hylobates
Papio
1
2
1
0
1
1
Human and gorilla above and below value for outgroup so there is no objective way to code
either with respect to the outgroup.
Invalid CW20
Orientation of zygomatic bone. Frontal = 0, Variable = 1, Frontolateral = 2, Lateral = 3
Pongo
Human
Pan
Gorilla
Hylobates
Papio
2
0
2
2
2
3
Single ingroup character state intermediate to the outgroup range.
0
Invalid CW92
Canine sexual dimorphism. Hyperdimorphic = 0, Strongly dimorphic = 1, Moderately
dimorphic = 2, Monomorphic, small canines = 3, Monomorphic, large canines = 4
Pongo
Human
Pan
Gorilla
Hylobates
Papio
1
3
2
1
4
0
Character states mixed between dimorphism and size, inconsistently identified for each taxon.
(b) Gibbs et al. (2004) characters
Invalid G2
Anterior bellies of digastric in contact in midline
Pongo
Human
Pan
Gorilla
Hylobates
0
1
0
1
0
Orangutan does not have anterior digastric muscle (Schwartz, 2005, p. 236-237)
Invalid G7
Conical filiform [papillae] predominate over cylindrical filiform 0 = no, 1 = yes
Pongo
Human
Pan
Gorilla
Hylobates
0
1
1
1
0
Sources: Sonntag (1921)
Coding cannot be corroborated. Sonntag (1921) makes no reference to humans. Either condition
may predominate in Pan, and the same condition is also found in Gorilla. In Hylobates the
condition appears variable among species, from H. syndactylus where papillae are cylindrical to
H. lar where most are filiform.
Pongo: Sonntag (1921, p.16): “The papillae belong to the filiform and cylindrical types, but the
former predominate”.
Pan: Sonntag (1921, p. 7): “The relative proportions of the filiform and cylindrical types differ
in different tongues. Either type may be in excess, and in some cases be evenly
distributed, but it is unusual to find one form predominating”.
Gorilla: Sonntag (1921, p. 12) “The conical papillae have the same form and arrangements as in
the chimpanzee.”
Hylobates: Sonntag (1921) Hylobates syndactylus (p. 19): “The conical papillae…belong to the
cylindrical type”, Hylobates hoolock (p. 21) “On the anterior two-thirds of the dorsum the
conical papillae are strong and coarse”, Hylobates muelleri (p. 24) “The conical papillae
have the usual arrangement…and belong to the cylindrical and filiform series”, Hylobates
lar (p. 27) “Most of the conical papillae are of the filiform type”.
Invalid G15
Humeroulnar head of flexor digitorum superficialis takes origin from intermuscular
septum 0 = no, 1 = yes
Pongo
Human
Pan
Gorilla
Hylobates
0
1
1
1
0
3
Sources: Beddard (1893), Dwight (1895), MacDowell (1910), Warwick & Williams (1973).
Coding cannot be corroborated because there is no citation for Hylobates, and Beddard (1883)
also indicates an intermuscular septum origin in Pongo.
Pongo: Beddard (1883, p. 211): “Flexor sublimus (perforatus) digitorum arises from the flexor
condule between the flexor ulnaris and flexor carpi radialis, from the septum between
itself and these muscles and also the flexor profundus, and from a part of the radius
behind…”.
Human: Warwick & Williams (1973, p. 544): “The humero-ulnar head arises from …the
intermuscular septa between it and the preceding muscles…”
Pan: Dwight [1895, p. 36 (as flexor sublimis digitorum)]: “…arises (1) from the
common condyloid muscular mass and septa within it; (2) from the corronoid, and (3)
from the oblique line of the radius”. Beddard (1893, p. 189): “Flexor sublimis
(perforatus) digitorum…arises from the flexor condyle, from the septa between itself and
adjacent muscles, and from the radius.” MacDowell (1910, p. 440): “The flexor
digitorum sublimus (perforatus)…the division for the fith digit arises from the
intermuscular septum between it and the flexor carpi ulnaris…”.
Invalid 16
Flexor carpi radialis orign from intermuscular septum 0 = no, 1 = yes
Pongo
Human
Pan
Gorilla
Hylobates
0
1
1
0
1
Sources: Beddard (1893), Warwick & Williams (1973).
Coding cannot be corroborated in the absence of a citation for Hylobates and Gorilla, and the
same origin cited for Pongo by Beddard (1893).
Pongo: Beddard (1893, p. 210): “…arises from the flexor condyle of the humerus and from the
septa between itself and the pronator radii teres and the flexor sublimis…”
Human: Warwick & Williams (1973, p. 543): “…arises from the medial epicondyle by the
common flexor tendon, from the antebrachial fascia, and from the intermuscular septa
between it and the adjacent muscles.”
Pan: Beddard (1893, p. 189): “It arises from the flexor condyle, from the septa between itself
and the pronator teres, flexor sublimis, and palmaris longus.”
Invalid G17
Flexor carpi radialis fused with flexor digitorum superficialis 0 = no, 1 = yes
Pongo
Human
Pan
Gorilla
Hylobates
1
0
1
1
0
Sources: Sonntag (1923, 1924), Raven (1950)
Coding cannot be corroborated in the absence of a citation for human or Hylobates. It is also not
clear from Raven (1950) as to whether there is fusion between the muscles in the gorilla.
Pongo: Sonntag (1924, p. 368) “It is fused with the flexor sublimis digitorum…”
Pan: Sonntag (1923, p. 348) “…is intimately fused with the pronator radii teres and flexor
sublimus digitorum [flexor digitorum superficialis].”
Gorilla: Raven (1950, p. 44) “Origin: medial epicondyle of the humerus, fused with its
neighbors, and by a flat tendon from the lateral border of the radius…”
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Invalid G18
Flexor carpi radialus insertion into plamar surface of base of MIII
Pongo
Human
Pan
Gorilla
Hylobates
0
1
0
1
?
Outgroup unknown so the ingroup character state coding cannot be specified.
Invalid G20
Flexor carpi ulnaris originates from intermuscular septum 0 = no, 1 = yes
Pongo
Human
Pan
Gorilla
Hylobates
1
1
1
0
0
Sources: Beddard (1893), Warwick & Williams (1973).
Coding cannot be corroborated in the absence of a citation for Hylobates and Gorilla, and the
presence of the same condition for Pongo indicated by Beddard (1893).
Pongo: Beddard (1893, p. 211): “This muscle arises from the flexor condyle between the flexor
ulnaris and flexor carpi radialis, from the septum between itself and these muscles…”
Human: Warrick & Williams (1973, p. 543): “…the ulnar head…and from the intermuscular
septum between it and the flexor digitorum superficialis.”
Pan: Beddard (1893, p. 189): “The muscle arises from the flexor condyle and from a
considerable portion of the ulna, and from the septa between itself and adjacent muscles,
as well as from the fascia covering the arm.”
Invalid G21
Flexor carpi ulnaris gives origin to some fibres of flexor digitorum superficialis 0 = no, 1 =
yes
Pongo
Human
Pan
Gorilla
Hylobates
1
0
1
0
0
Sources: Dwight (1895), Sonntag (1924), Sullivan & Osgood (1927).
Coding cannot be corroborated because no source is given for Hylobates and Gorilla and there is
apparent conflict for Pongo between Sonntag (1924), indicating that it receives rather than gives
origin to fibre of the flexor digitorum superficialis, and Sullivan & Osgood (1927), indicating
otherwise.
Pongo: Sonntag (1924, p. 369): “It is attached by fascia to the olecranon, and it received fibres
from the flexor sublimis digitorum [flexor digitorum superficialis].” Sullivan & Osgood
(1927, p. 222): “On its deep surface the muscle unites with the slip of the superficial
digital flexor for the fifth digit”.
Pan: Dwight (1895, p. 36): “The deep tendon gives origin to some fibers of the flexor sublimis
digitorum).”
Invalid G 26
Flexor pollicis longus gives origin to tendon to digit II 0 = no, 1 = occasional, 2 = often
Pongo
Human
Pan
Gorilla
Hylobates
0
1
2
0
1
Coding cannot be corroborated as is not possible to know whether absence of origin, or frequent
origin, with digit II is the derived state since both are present in the outgroup.
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Recoded G33
Origin of extensor digitorum from radius/ulna 0 = radius and ulna, 1 = ulna only, 2 =
neither forearm bone
Pongo
Human
Pan
Gorilla
Hylobates
1
2
0
0
1
It is not possible to determine how the unique human condition is related to either the ulna or
radius/ulna position of the other apes.
Informative only for Pan and Gorilla. Recoded for radius/ulna origin as shared derived state
Pongo
Human
Pan
Gorilla
Hylobates
0
0
1
1
0
Invalid G34
Extensor digitorum originates from antebrachial fascia 0=no, 1=yes
Pongo
Human
Pan
Gorilla
Hylobates
0
1
1
0
0
Sources: Beddard (1893), Sonntag (1923), Warwick & Williams (1973)
Coding cannot be corroborated because no source is given for Hylobates and Gorilla.
Pongo: Beddard (1893, p. 188): “Extensor communis digitorum [extensor digitorum]…arises
from the extensor condyle, and lies between the extensor carpi radialis brevior and the
extensor minimi digiti, from the septa between which muscles and itself it also arises…”
Human: Warwick & Williams (1973, p. 548): “…arises from the lateral epicondyle of the
humerus by the common extensor tendon, from the intermuscular septa between it and
adjacent muscles, and from the antebrachial fascia.”
Pan: Beddard (1893, p. 188): “Extensor communis digitorum…arises from the extensor condyle
of the humerus, from the septa between itself and the adjacent muscles, and from the
fascia covering the deep extensors…”Sonntag (1923, p. 351): “The extensor communis
digitorum arises from the external epicondyle, the fascia over it, and the intermuscular
septa on either side”.
Invalid G40
Lateral head of triceps brachii originates from lateral intermuscular septum 0=no, 1 = yes
Pongo
Human
Pan
Gorilla
Hylobates
0
1
1
0
0
Sources: Beddard (1893), Warwick & Williams (1973)
Coding cannot be corroborated with respect to Gorilla and Hylobates and there is no clear
exclusion of any intermuscular origin for Pongo by Beddard (1893) and a lack of difference
noted by Anderton (1988) for Pongo and Pan.
Pongo:Beddard (1893, p. 208): “The outer head arises from the humerus commencing about half
an inch below the insertion of the teres minor: the origin of the inner head commences a
little below that of the outer head.”
Anderton (1988, p. 332-333) noted for the orangutan that the lateral head has “a notably
restricted origin from the posterior aspect of the humerus, just superior to the surgical
neck and between the insertions of the teres minor superiorly, and the extended lateral
origin of the brachialis inferiorly.” He also notes (p. 333) that the chimpanzee described
by Sonntag (1923) “exhibited the same characters of the lateral head as seen in my orangutan…”
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Human: Warwick & Williams (1973, p. 541) “The lateral head arises by a flattened tendon from
a narrow ridge on the posterior surface of the shaft of the humerus, from the lateral
border of the humerus and the lateral intermuscular septum.”
Pan: Beddard (1893, p. 187) “Both humeral heads arise not only from the humerus, but also
from the septa between themselves and adjacent muscles.”
Invalid 42
Extensor digitorum inserts into interphalangeal joints 0 = no, 1= yes
Pongo
Human
Pan
Gorilla
Hylobates
1
0
1
0
0
Sources: Sonntag (1923), Sullivan & Osgood (1927), Warwick & Williams (1973)
Coding cannot be corroborated in the absence of sources for Hylobates and Gorilla, and
apparently the same condition is described for humans by Warwick & Williams (1973)
Pongo: Sullivan & Osgood (1927, p. 218): “The tendons of the long extensor[extensor digitorum
longus=extensor digitorum communis]…central part of each tendon attaches to the base
of the intermediate phalanx…”
Human: Warwick & Williams (1973, p. 549): “The extensor digitorum extends the fingers at the
metacarpophalangeal and interphalangeal joints…”
Pan: Sonntag (1923, p 351): “The extensor communis digitorum [extensor digitorum]…tendons
inserted into the bases of the ungulal phalanges”
Invalid 46
Abductor pollicis longus origination from intermuscular septum 0 = no, 1 = yes
Pongo
Human
Pan
Gorilla
Hylobates
1
0
1
0
0
Sources: Beddard (1893), MacDowell (1910).
Coding cannot be corroborated because of absence of information on Gorilla and ambiguous
information for Hylobates. Absence of an origin from intermuscular septum in humans is not
indicated by Warwick & Williams (1973).
Pongo: Beddard (1893, p. 211): “Flexor profundus (perforatus) digitorum. – This muscle
appears to represent both the muscle so called and the flexor pollicis longus of human
anatomy, since it arises both from the radius and ulna; it also arises from the interosseous
ligament.”
Human: Warwick & Williams (1973, p. 550): “It arises from the lateral part of the posterior
surface of the shaft of the ulna and below the anconeus, from the interosseous
membrane…”
Pan: Beddard (1893, p. 189): “Flexor profundus (perforans) digitorum…arises chiefly from the
ulna and from the interosseous membrane…”
MacDowell (1910, p. 443): “The Flexor digitorum profundus arises in the
chimpanzee…(b) from the radial half of the interosseous membrane.”
Hylobates: Beddard (1893, p. 445): “In a gibbon, Hepburn found an origin of the profundus
from the internal condyle.”
Invalid 53
Teres minor shares origin from intermuscular septum with teres major 0 = no, 1 = yes
Pongo
Human
Pan
Gorilla
Hylobates
7
0
1
1
0
0
Sources: Beddard (1893), Warwick & Williams (1973).
Coding cannot be corroborated because in the absence of sources for Hylobates and
Gorilla, and same condition for Pongo indicated by Sullivan & Osgood (1927).
Pongo: Beddard (1893, p. 208): “…entirely fleshy origin partly from the axillary border of
the scapula and partly from the septum between itself and the infra-spinatus.”
Sullivan & Osgood (1927, p. 210): “…just below and parallel to the infraspinuatus. It has
its scapular attachment to the upper two thirds of the sillary margin dorsal to the
attachment of the long head of the triceps and extending from the m. teres major to the
neck of the bone”.
Human: Warwick & Williams (1973, p. 539): “arises from the upper two thirds of a flattened
strip on the lateral part of the dorsal surface of the scapula …and from two aponeurotic
laminae, one separating it from the infraspinatus, and the other from teres major.”
Pan: Beddard (1893 p. 186): “…and also from the septa between itself and the following
Muscles, viz., deltoid, infraspinatus, triceps, and teres major.”
Invalid 54
Latissimus dorsi may originate from inferior scapular angle 0=no, 1 = yes
Pongo
Human
Pan
Gorilla
Hylobates
0
1
1
0
0
Sources: Hepburn (1892), Sonntag (1923, 1924), Sullivan & Osgood (1927), Miller (1952).
Coding cannot be corroborated in the absence of information on Hylobates and Gorrilla, and the
absence of reference to an inferior scapular angle origin in the sources cited. Hepburn (1892)
specifically indicates the absence of any other origin other than those observed.
General: Hepburn (1892, p. 151): “In the Gorilla and Orang it arose from the spinous process
and supra-spinous ligaments of the three lower dorsal vertebrate. (In the Chimpanzee and
Gibbon this part of the origin was mutilated)…In no case was there any additional origin
from the inferior angle of the scapula over which the muscle ran to its insertion…”
Pan: Sonntag (1923, p. 338): “arises from the lower five dorsal spines…none from
the inferior angle of the scapula”.
Miller (1952, p. 193) for the chimpanzee: “arises by means of the lumbar aponeurosis
from the spinous processes of the last 5 thoracic, the lumbar, and the sacral vertebrae. It
also takes origin from the middle third of the crest of the ilimum and by digitations from
the last 5 ribs”.
Pongo: Sonntag (1924, p. 360): “arises from the lower five dorsal spines and supraspinous
ligaments…no slips are derived from the inferior angle of the scapula, and all Apes agree
with one another and differ from man in this respect.”
Sullivan & Osgood (1927, p. 207): “The proximal attachment is through an aponeurosis,
the lumbodorsal fascia, to the spines of the tenth, eleventh, and twelfth thoracic vertebrae,
and the dorsal third of the iliac crest…We found no scapular or costal attachments…”
Invalid G55
Extent of costal origin of latissimus dorsi 0 = three or four ribs, 1 = three, four, or five ribs,
2 = five ribs, 3 = 6 ribs.
Pongo
Human
Pan
Gorilla
Hylobates
6 ribs
3-4 ribs
3-5 ribs
6 ribs
5 ribs
8
3
0
1
3
2
Coding cannot be corroborated. Ingroup range of values is above and below outgroup so it is
impossible to know the derived state.
Invalid G72
Tensor fascia latae normally fused proximally with gluteus maximus 0 = yes, 1 = no
Pongo
Human
Pan
Gorilla
Hylobates
0
1
1
0
0
Sources: Hepburn (1892), Swindler & Wood (1973), Sigmon (1974).
Coding cannot be corroborated because the tensor fascia is absent in orangutan so its
origin prior to loss is unknown. Swindler & Wood (1973) also indicate that fusion is
infrequent in Homo and the anthropoids. There is also no clear indication of proximal
fusion in Hylobates by Sigmon (1974).
Great apes: Hepburn (1892, p. 324): “In the Orang there was no trace of the tensor fasciae
femoris muscle, but in the Gorilla and Chimpanzee it was present as a feeble muscular
slip inserted into the fascia of the thigh close to the insertion of the gluteus maximus.”
Pan/Homo/Papio: Swindler & Wood (1973, p. 246): “On the lateral side of the thigh, note the
tensor fascia latae, and in Papio, the gluteus maximus…their anterior position should be
noted in Papio, especially that of the gluteus maximus. These two muscles are fused in
their upper portions, a condition only infrequently encountered in Homo and the
anthropoids.”
Hylobates/Pan/Gorilla: Sigmon (1974, p. 165): “Tensor fasciae latae present in the gibbon,
chimpanzee, and gorilla. It is not found in the orangutan…in the former three
pongids…originates in the gluteal fascia.”
Invalid G74
Gluteus maximus fused with biceps femoralis 0 = no fusion, 1 = at origin, 2 = more distally
Pongo
Human
Pan
Gorilla
Hylobates
2
0
1
1
1
Coding cannot be corroborated because if fusion at the origin is assumed to be the primitive
state, it is not possible to know if the attachment was previously distal or at the origin for humans
where no fusion is the current state.
Invalid G95
Incidence of peroneus tertius 0 = low (0-5% of specimens), 1 = moderate (30-50%), 2 = high
(~95%)
Pongo
Human
Pan
Gorilla
Hylobates
0-5%
95%
0-5%
30-50%
30-50%
0
2
0
1
1
Coding cannot be corroborated because it is not objectively possible to designate the human
character state where the incidence is higher than the outgroup values compared with both Pongo
and Pan where the incidence is below the outgroup values.
Invalid G107
Origin of transverse head of adductor hallucis with metatarsophalangeal joints
9
Pongo
Human
Pan
Gorilla
Hylobates
rd
th
th
th
2-3
3-5
2-4
2-4
2-4th
0
2
1
1
1
Uninformative as both Pongo and human conditions may be derived from 2-4th
metatarsophalangeal joints so there is no objective way to know which is closer to Pan and
Gorilla.
Invalid G108
First dorsal interosseious originates from MI and MII. No=0, yes=1
Pongo
Human
Pan
Gorilla
Hylobates
0
1
1
0
0
Sources: Champneys (1872), Brooks, (1887), Dwight (1895), Sonntag (1924), Boyer (1935),
Raven (1950), Miller (1952), Warwick & Williams (1973).
Coding cannot be corroborated. The literature cited indicates that the origin in the chimpanzee is
from MI and MII, but there is lack of clarity as to whether this excludes the cuneiform. If it does,
the chimpanzee and human share this feature in common. However, the gibbon origin is limited
to MII only, so it is not possible to know whether the MI-MII configuration of humans and
chimpanzees, or the MI-MII plus cuneiform of orangutans and gorillas represents the derived
state for large bodied hominoids. There is also uncertainty as to whether the gorilla configuration
is also unique in lacking any contact with MI. Hepburn (1892, p. 348) distinguished the
orangutan from African apes in having the first interosseous muscle “in which the inner head of
the muscle arose from the internal cuneiform bone (fig. 6), but his illustration of the Gorilla (fig.
7) also shows this connection.
Pan: Miller (1952, p. 225): “Each one arises by two heads from the sides of the two adjacent
metatarsal bones…”
Dwight (1895, p. 41): “Dorsals. First muscle from the first and second metatarsal
bones…”
Pan, Pongo, Gibon: Champneys (1872, p. 206): “The first dorsal was much the largest, and had
a broad origin from the base of met. I. as well from the side of met. I.” Brooks (1887, p.
92): “In the Chimpanzee it arises from the extreme base of the metatarsal of the hallux
(PI. III. fig. 3, abd2); in the Orang it takes origin from the internal cuneiform bone (which
in this ape is placed at a different angle to the remaining cuneiforms, thus resembling the
trapezium in the hand. In the hand of the Gibbon (PI. III. fig. 5, abd2) it arises from the
trapezium and base of the thumb metacarpal; in the foot of the same animal it is a singleheaded muscle, arising entirely from the tibial side of the index metatarsal bone.”
Pongo: Sonntag (1924, p. 380): “…the inner head of the first dorsal
interosseous muscle, which arises, according to Hepburn (35), from the internal
cuneiform bone. In my specimen the arrangements are similar to those described by
Hepburn.”
Boyer (1935, p. 237): “It arises by two heads, one from the first cuneiform bone and the
base of the phalanz of the first toes, the other from the medial half of the dorsal surface of
the proximal portion of the second metatarsal bone and a narrow strip of the dorsal part of
the medial surface along the length of the metatarsal bone.”
Gorilla: Raven (1950, p. 64): ”…originates from the dorsodistal aspect of the first cuneiform
and most of the dorsal, medial, and plantar aspects of the second metatarsal.”
Human: Warwick & Williams (1973, p. 583): “each arising by two heads from the adjacent
10
sides of the two metatarsal bones.” Fig. 5.111A illustrates origin at MI and MII only.
Invalid 109
Flexor digitorum brevis originates from plantar aponeurosis 0 = no, 1 = yes
Pongo
Human
Pan
Gorilla
Hylobates
0
1
1
0
0
Sources: Chapman (1878), Warwick & Williams (1973).
Coding cannot be corroborated because of absence of information on Hylobates, Pongo and Pan.
Vereecke et al. (2005) state that the plantar aponeurosis in gibbons and bonobos is not as
extensive and strong as in humans and a longitudinal foot arch is lacking. In gibbons and
bonobos, the plantar aponeurosis originates from the calcaneal tuberosity and from the
intermuscular septum between the hallucal and digital flexors. The origin of the flexor digitorum
brevis is identified as the medio-plantar side of the tuber calcanei, but the positional relationship
to the plantar aponeurosis is not described. Anderton (1988, p. 340) notes that “the origin of the
superficial head of the flexor digitorum brevis from the plantar aponeurosis is limited in
orangutans due to the lateral…origin of the abductor hallucus, which overlies the former
structure; otherwise, the disposition of the muscle is rather consistent among hominoids.
Gorilla: Chapman (1878, p. 390): “The part supplying the second and third toes…only in part
from the calcaneum.” makes no reference to the plantar aponeurosis.
Human: Warwick & Williams (1973, p. 580): “It arises by a narrow tendon from the medial
process of the calcanean tuberosity, from the central part of the plantar aponeurosis…”
Invalid G120
Origin of radial recurrent artery from brachial and radial arteries 0 = radial artery, 1 =
variable, 2 = brachial artery
Pongo
Human
Pan
Gorilla
Hylobates
brachial
radial
variable
radial
radial
2
0
1
0
0
Coding cannot be corroborated. Not possible to relate Pongo character state to that of Pan where
the condition is variable.
Invalid G137
Muscular branches of profunda femoris for hamstring 0 = no, 1 = yes
Pongo
Human
Pan
Gorilla
Hylobates
0
1
1
0
0
Sources: Sonntag (1923), Warrick & Williams (1973).
Coding cannot be corroborated in the absence of information for Gorilla and Hylobates.
Pan: Sonntag (1923, p. 390): “The profunda femoris gives off the lateral circumflex and a
branch
passing back under the rectus femoris to the gluteus medius. It then passes through the
middle head of the adductor magnus, supplies the adductor longus and vasti, and ends in
the biceps”. These citations support supply of profunda femoris to the hamstring muscles
(at least the biceps in the chimpanzee), but give no indication of the situation in the other
apes.
Human: Warrick & Williams (1973, p. 677): “Numerous muscular branches arise from the
arteria profunda femoris;…give branches to the hamstrings…”
11
Invalid G139
Number of digits supplied by median nerve 0 = normally 2.5, 1 = normally 3.5
Pongo
Human
Pan
Gorilla
Hylobates
0
1
1
0
0
Sources: Chapman (1878), Hepburn (1892), Sonntag (1924), Raven (1950), Warwick &
Williams (1973).
Cannot be corroborated in absence of information on Hylobates and citations do not show the
orangutan to be different from humans or African apes.
Pongo: Sonntag (1924, p. 431): “The median nerve…At the wrist divides into two branches. The
first, or lateral branch, supplies the thenar muscles and gives cutaneous branches to both
sides of the thumb. The second, or medial branch, divides to supply the adjacent sides of
the first, second, and third fingers, and it sends a branch to the first two lumbrical
muscles.”
Pongo/Gorilla: Hepburn (1892, p. 184): “…median nerve…in the Gorilla and Orang the
cutaneous and muscular supply were exactly the same as in Man. In the Chimpanzee, the
cutaneous supply resembled that found in Man; but in regard to the supply of muscles, it
had the 3rd lumbrical muscle supplied by the median nerve, while all the other muscles
were supplied just as in Man. The greatest variety, however, existed in the Gibbon. In it
the median nerve merely supplied the two and a half radial digits with digital cutaneous
branches; and of muscles it supplied the two outer lumbricales, the abductor pollicis
(brevis), the opponens pollicis, and both heads of the flexor brevis pollicis…”
Human: Warwick & Williams (1973, p. 1043): “In most cases the median nerve [of humans]
supplies…the lateral three and one-half digits (thumb, index, middle and lateral side of
the ring finger);...”
Gorilla: Chapman (1878, p. 392): “The median nerve, Plate VI, fig 1, f, supplies, in the Gorilla,
the thumb, index, middle, and the inner side of the ring finger;…”
Raven (1950, p. 45-46): “M. opponens pollicis…Mm. lumbricales…Invervation: first and
second by median nerve; third and fourth by the deep branch of the ulnar nerve...” But
Plate 44 p. 137 shows enervation of same digits as for humans.
Invalid G141
Gangliform enlargement at junction of radial and posterior interosseous nerves 0 = no, 1 =
yes
Pongo
Human
Pan
Gorilla
Hylobates
0
1
1
0
0
Sources: Champneys (1872) Warwick & Williams (1973)
Coding cannot be corroborated because sources do not refer to a junction between radial and
posterior interosseous nerves, and while they support the presence of a pseudoganglion in
humans and chimpanzee, they do not refer to its presence or absence in any other primate.
Pan/human: Champneys (1872, p. 14): “Gangliform enlargements over the back of the carpus,
at the end of the posterior Interosseous nerve, and on the branch of the Circumflex going
to the Teres minor were present in CHIMP. as in man.”
Human: Warwick & Williams (1973, p. 1045): “…posterior interosseous…reaches the
12
dorsum of the carpus, where it presents a flattened and somewhat expanded termination
(‘pseudoganglion’) from which filaments are distributed to the ligaments and
articulations of the carpus.”
Invalid G144
Number of lumbricals innervated by ulnar nerve 0 = normally one, 1 = normally two, 2 =
normally three.
Pongo
Human
Pan
Gorilla
Hylobates
Two
two
one
three
two
Coding cannot be corroborated because there is no objective way to determine polarity of Pan
and Gorilla above and below the value given for the outgroup (Hylobates).
Invalid G147
Origin of subscapular nerves 0 = C5, C6, 1 = C5-7, 2 = C5-8, 3 = C5-8 and T1
Pongo
Human
Pan
Gorilla
Hylobates
C5-7
C5
C5-8/T1
C5-8/T1
C5-8
1
0
3
3
2
Spilt mixed states G155
Muscular branches of medial plantar nerve
Pongo
Human
Pan
Gorilla
Hylobates
Two/hallucis one
Two/hallucis one
two
2
0
2
0
1
separate into number lumbricals and presence or absence of adductor hallucis
(a) number of lumbricals
Pongo
Human
Pan
Gorilla
Hylobates
Two/
one
Two
one
two
0
1
0
1
0
(b) adductor hallucis present
Pongo
Human
Pan
Gorilla
Hylobates
hallucis
no
hallucis
no
no
1
0
1
0
0
Invalid G162
Axillary organ present or absent 0 = absent, 1 = present
Pongo
Human
Pan
Gorilla
Hylobates
0
1
1
1
0
Incorrect. Axillary organ also present in orangutan (see references in Grehan, 2006).
Invalid G167
Scrotum normally postpenial 0 = no, 1 = yes
Pongo
Human
Pan
Gorilla
Hylobates
0
1
1
0
0
Sources: De Beaux (1917), Miller (1933), Wislocki (1936), Harrison-Matthews (1946), Hill &
Harrison-Matthews (1949), Hill (1958), Hill & Kanagasuntheram (1959).
13
Cannot be corroborated as sources indicate a similar position in all great apes and possibly
some gibbons.
Great apes/gibbons: Wislocki (1936, p. 338): postpenial in all simians except a majority of
gibbons where it is parapenial or partially prepenial (also De Beaux, 1917; HarrisonMatthews, 1946; Miller, 1933; Hill, 1958).
Pongo: Wislocki (1936, p. 332): “In this specimen the postpenial scrotum is darker…The
scrotum consists of two pigmented halves situated postpenially…”
Humans: Warwick & Williams (1973, p. 1345) make no reference to scrotal position with
respect to the penis.
Pan: Wislocki (1936, p. 328): “…the scrotum is postpenial…”
Gorilla: Wislocki (1936, p. 334): “The genitalia of a mountain gorilla…testes which are
parapenial to postpenial in position…(p. 335)…an adult…the small scrotum is placed
immediately behind the base of the penis…”
Hill & Harrison-Matthews (1949). No reference to position.
Hylobates: No statements about position by Hill & Kanagasuntheram (1959), although
illustrations in both articles show a postpenial position.
Invalid G169
Relative testes size [weight] (ratio of observed/predicted body testis size) 0 ≤ 0.4, 1 ≥ 0.4
Pongo
Human
Pan
Gorilla
Hylobates
0
1
1
0
0
Source: Schultz (1938).
Coding cannot be corroborated. Schultz (1938) does not document the ratio of observed body
testis size against predicted, but only gives testis weight as a percentage of body width: Pan
(0.27), human (0.08), Hylobates (0.08), Pongo (0.05), and Gorilla (0.02). Harcourt et al. (1981)
shows that the ratio of testis to body weight of Pan (0.27) is higher than humans (0.06), Gorilla
(0.02) and Pongo (0.06), and these values overlap those of various monkeys. Dahl et al. (1993, p.
234) suggest the two orangutans measured by Schultz (1938) were immature.
Invalid G171
Development of transverse rugae of vagina 0 = little developed, 1 = well developed
Pongo
Human
Pan
Gorilla
Hylobates
0
1
1
1
0
Sources: Deniker (1885), Gerhardt (1906), Sonntag (1923), Wislocki (1932), Dempsey (1940),
Atkinson & Elftman (1950).
Coding cannot be corroborated. Wislocki (1932) indicates that rugae are virtually absent in Pan
as well as Pongo, and while Sontage refers to transverse folds there is no specific comparison
with the human condition. Descriptions are variable for gorilla and gibbon, with the illustration
and description by Gerhardt (1906) only referring to fine transfuse folds between the longitudinal
folds in the lower half of the vagina.
Apes: Wislocki (1932, p. 184): “…these folds are conceded to be less pronounced at any period
of life than in the human…In the chimpanzee, orang-utan and gibbon they are practically
absent…In the gorilla…there are moderate numbers of low transverse rugae…more
numerous in the upper part of the vagina, the middle portion of the vagina being nearly
smooth, while the lower parts exhibit low longitudinal rugae.”
Pan: Sonntag (1923, p. 401): “…the mucosa has transverse folds.”
14
Gorilla: Atkinson & Elftman (1950, p. 205): “Rugation of the gorilla vagina is not marked.”
Denniker (1885, p. 243 for foetus): “Plus loin commence le vagin (id., v), dont les parois
sont absolument lisses (translated as “Further the vagina (id., v) starts, whose walls are
absolutely smooth”).
Gerhardt (1906, p. 636): “Diese runzeln, die an der vorderen wand zu einer deutlichen,
4.5 cm langen, 1.6 cm breiten säule von queren falten zusammenfliessen, sind sehr fein
und dicht gestellt.Ihre ausbildung ist sehr viel geringer als beim menschen. Die oberen
partien der vaginalwand sind glatt. Alle falten der scheide vereinigen sich an deren
unterem ende zur bildung eines deutlichen hymens, der als ca. ½ cm breite
halbmonförmige falte, besonders an der hinteren vagainalwand, deutlich ins lumen der
scheide vorspringt. Die form des hymens ist also die beim menschlichen weibe häufigste.
Unterhalb des hymens, in der wand des vestibulum vaginae, liegen tief nischen, zwischen
denen falten oder runzeln vorspringen, doch stehen sie viel weiter voneinander entfernt
als in der vagina”. Translated as “At its upper end before and behind portio vaginalis
uteri, the vagina forms a shorter front and a longer rear fornix vaginae. The vagina itself
is a channel 4.75cm in length, its circumference, measured at the lower part, is 3.8cm. In
the lower half, particularly on the frontal wall, there are recognizable and distinct
longitudinal wrinkles that are slightly diagonal. Within these wrinkles, on the frontal wall
are distinct 4.5cm long and 1.6cm wide columns of transverse folds that blend together,
are fine and closely spaced. Their development is far more constrained than that in
humans. The upper parts of the vaginal wall are smooth. All folds of the vagina merge
posteriorly to form a distinct hymen, which is an approximately 0.5cm half-moon shaped
fold, particularly at the hindmost vaginal wall, clearly protrude in the lumen of the
vagina. The shape of the hymen is also common in the female human. Beneath the
hymen, in the vestibulum vaginae wall, lay deep recesses between the folds or where
wrinkling occurs, but are spaced further apart as in the vagina.”
Human: Wislocki (1932, p. 184): “These are most conspicuous in the young and virginal
vaginas, having a tendency to disappear with age and after childbirth.”
Hylobates: Dempsey (1940, p. 234): “…mucosal folds and ridges. These ridges vary in size and
texture, depending upon the phase of the reproductive cycle. They are large and course in
animals whose ovaries contain large follicles…During the luteal phase of the cycle, on
the contrary, the folds are smaller and quite smooth.
Deniker (1885, p. 250 for foetus): “Le vagin est lisse, excepté dans sa partie tout à fait
postérieure voisine du col de l'utérus (id., v'), où l'on aperçoit une série de rides
transversales sur la paroi postérieure aussi bien que sur la paroi antérieure, mais pas sur
les parois latérales. Ce sont là évidemment les homologues des plis transversaux du vagin
de la femme (colonnes rugueuses). (Translated as “The vagina is smooth, except in its
part completely posterior in the neighborhood of the collar of the uterus (id., v'), where
one also sees a series of transverse wrinkles on the posterior wall of the former wall, but
not on the side walls. They are there obviously the counterparts of the transverse folds of
the vagina of the woman (rough columns)”.
(c) Character changes to G&S
15
Coding modified GS4
Chest hair density
Pongo
Human
Pan
Gorilla
Hylobates
Papio
1
0
1
0
2
1
Recoding by Lehtonen et al. (2011) results in only one character state uniquely shared within
large bodied hominoids (human, gorilla) so we have retained our original coding which provides
the same information:
Pongo
Human
Pan
Gorilla
Hylobates
Papio
0
1
0
1
0
0
Coding modified GS14
Type I aorta proporation
Pongo
Human
Pan
Gorilla
Hylobates
Papio
2
0
0
0
2
1
Coding by Lehtonen et al. (2011) results in only state 0 being shared within the ingroup.
Recoded to original which provides the same information.
Pongo
Human
Pan
Gorilla
Hylobates
Papio
0
1
1
1
0
0
Coding modified GS27
Ethmolacrimal contact
Pongo
Human
Pan
Gorilla
Hylobates
Papio
2
2
1
0
2
2
Coding by Lehtonen et al. (2011) results in only two informative character states linking Pan and
Gorilla. Recoded for outgroup state = 0, Pan and Gorilla as 1, which provides the same
information:
Pongo
Human
Pan
Gorilla
Hylobates
Papio
0
0
1
1
0
0
Coding modified GS28
Average number of sacral vertebrae
Pongo
Human
Pan
Gorilla
Hylobates
Papio
2
2
2
2
1
0
Coding by Lehtonen et al. (2011) renders this character (average number of sacral vertebrae)
uninformative. Since chimpanzees and gorillas share a higher average number (six) of sacral
vertebrae than other primates we have retained our original coding to reflect this fact with
respect to an average of five for humans and orangutans, five for Hylobates, and three for Papio.
Coding modified GS49
Ethmoid-sphenoid contact
Pongo
Human
Pan
Gorilla
Hylobates
Papio
2
2
2
1
0
0
Coding by Lehtonen et al. (2011) clusters Pan with Pongo and human, but this obscures the fact
that orangutan (77%) is more similar to Homo (97%) than Pan (77%) relative to the outgroup (025%). We have recoded this as follows:
16
Pongo
3
Human
3
Pan
2
Gorilla
1
Hylobates
0
Papio
0
Coding modified GS61
Mammary gland spacing
Pongo
Human
Pan
Gorilla
Hylobates
Papio
2
1
1
0
0
0
Coding by Lehtonen et al. (2011) clusters human with pan, but this obscures the fact that human
spacing (71%) is more derived with respect to Pongo (90%) than Pan (52%) or Gorilla (46%).
We have therefore recoded as a multistate character as follows:
Pongo
Human
Pan
Gorilla
Hylobates
Papio
3
2
1
0
0
0
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