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ANNALS OF THE SOUTH AFRICAN MUSEUM

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Volume 69 Band

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ANCHISAURUS CAPENSIS (BROOM)

AND A REVISION OF THE ANCHISAURIDAE

(REPTILIA, SAURISCHIA)

P. M. GALTON & M. A. CLUVER

Cape Town Kaapstad

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ANCHISAURUS CAPENSIS (BROOM) AND A REVISION OF THE

ANCHISAURIDAE (REPTILIA, SAURISCHIA)

P. M. GALTON

University of Bridgeport, Bridgeport, U.S.A.

M. A. CLUVER

South African Museum, Cape Town

(With 13 figures and 3 tables)

[MS accepted 22 October 1975]

ABSTRACT

A complete description of the skeleton of the type specimen (SAM-990) of the pro-

sauropod dinosaur Gyposaurus capensis Broom is given. Comparison with other prosauropods

indicates that this dinosaur represents a valid South African species of Anchisaurus Marsh,

1885 from the Upper Triassic of North America. It is considered that the infra-order Pro-

sauropoda should be divided into three families, viz. Anchisauridae, Plateosauridae and

Melanorosauridae. It is proposed that the family Anchisauridae be restricted to prosauropods

with relatively slender feet, and that broad-footed forms previously assigned to the Anchi-

sauridae be transferred to the Plateosauridae. The Family Anchisauridae is therefore con-

sidered to include the genera Anchisaurus, Efraasia, Thecodontosaurus and several indetermi-

nate species. The genera Ammosaurus, Aristosaurus, Massospondylus (including Gryponyx,

Aetonyx and Dromicosaurus), and Lufengosaurus (including Yunnanosaurus) are included in

the Plateosauridae, while Arctosaurus and Ischisaurus are referred to the suborder Theropoda.

‘Thecodontosaurus’ gibbidens, Spondylosoma absconditum and Teleocrater alphos are placed

within the Thecodontia, and Tanystropheus primus and T. latespinatus in the order Lacertilia.

Thus constituted, the family Plateosauridae becomes the dominant and most widespread

prosauropod family, while the Anchisauridae, in contrast, is known from a geographically

and numerically restricted fossil record.

CONTENTS

PAGE

Introduction , : : : 122

Redescription of An enisaunes ener (Broom) : : z 124

Vertebral column . : : : 5 : : ; 124

Pelvic girdle . : : : : : : : : 126

Hind limb : : ; : . 129

Slender and broad- footed Prosauropods ; ; : a - b3il

Systematic discussion . , : ; ‘ : ? 132

The Family Anchisauridae : : ? : : d 132

Anchisauridae nomina dubia ; F : 141

Prosauropod species incorrectly assigned to Anchisauridae ‘ 143

Prosauropoda nomina dubia Z g : 150

Non-prosauropod species incorrectly assiened to

Anchisauridae : 150

Notes on the Families Anchisauridae and Plateosauridae : 153

Summary . 2 : ‘ : ; ; ; : : 155

Acknowledgements. 3 : : , : : ; 155

References : 3 5 : : : F , : 156

Abbreviations. 3 : 2 P : ‘ ; : 158

IBZ

Ann. S. Afr. Mus. 69 (6), 1976: 121-159, 13 figs, 3 tables.

122 ANNALS OF THE SOUTH AFRICAN MUSEUM

INTRODUCTION

Broom (1906) described a specimen (SAM-990) from the Cave Sandstone,

Stormberg Series (Upper Triassic) of Ladybrand, Orange Free State, South

Africa, and referred it to the prosauropod taxon Hortalotarsus skirtopodus

Seeley, 1894. The holotype of Hortalotarsus skirtopodus was referred to the

genus Thecodontosaurus Riley & Stutchbury, 1836, by Huene (1906) as T.

skirtopodus and, because SAM-990 differed in several aspects from Theco-

dontosaurus, Broom (1911) made it the holotype of Gyposaurus capensis.

Broom (1906) noted several resemblances between SAM-990 and Anchisaurus

Marsh, 1885, from the Upper Triassic of North America, so he referred Gypo-

saurus capensis (SAM-990) to the Family Anchisauridae Marsh, 1885. Galton

(1973, in press) provisionally accepted the validity of Gyposaurus capensis

but noted that the holotype should be carefully compared with Anchisaurus

polyzelus (Hitchcock) to determine whether or not these species are generically

distinct. A comparison of photographs of SAM-990 with a specimen of Anchi-

saurus polyzelus (YPM 1883) showed that Gyposaurus is a junior synonym of

Anchisaurus and that SAM-990 should be redescribed.

Huene in several papers between 1906 and 1932 made important contri-

butions to an understanding of the Family Anchisauridae (as Thecodonto-

sauridae Lydekker, 1890). Charig et al. (1965) considerably enlarged the family

by referring to it genera of Triassic theropods (Family Gryponychidae =

‘Palaeosauridae’) based solely on postcranial material which was_ indis-

tinguishable from that of prosauropods. The generic list of Romer (1966: 370)

includes these changes and the comprehensive list given by Steel (1970) faithfully

but rather uncritically records all the genera and species referred to the family.

A taxonomic revision of the Family Anchisauridae is necessary for several

reasons:

1. Several of the suggested and generally accepted synonymies are probably

incorrect. In assessing these the recognition of slender- and broad-footed

types (Galton 1971, 1973, in press) is useful.

2. The skeletal anatomy of prosauropods is remarkably uniform, so taxa

should be based on specimens which include either most of the specimen or

bones which are diagnostically different from those of other prosauropods.

Because of the limited number of skeletal variations it is necessary to diagnose

taxa on the basis of a combination of characters which, if each was taken in

isolation, would not be diagnostic. Unfortunately several of the taxa listed by

Steel (1970) are based on specimens which are generically and_ specifically

indeterminate.

3. The group has been over-classified and insufficient account has been

taken of the range of individual variation possible within a dinosaurian species;

this can be quite extensive, as shown by the prosauropod Lufengosaurus huenei

Young (see Rozhdestvensky 1966) and the ornithopod Hypsilophodon foxii

Huxley (see Galton 1974).

4. A few of the taxa included within the family are not prosauropods.

ANCHISAURUS CAPENSIS (BROOM) AND A REVISION OF THE ANCHISAURIDAE 123

Fig. 1. Anchisaurus capensis (Broom). Type specimen (SAM-990) showing vertebral column

(top), pubis, ischia and pes (middle) and ilium, femur and fibula (bottom).

124 ANNALS OF THE SOUTH AFRICAN MUSEUM

Summaries of certain aspects of the revision of the family Anchisauridae

have been published earlier (Galton 1971, 1973, in press). The first author

(P. M. G.) is responsible for all sections except the description and illustration

of SAM-990, which is the work of the second author (M. A. C.).

REDESCRIPTION OF ANCHISAURUS CAPENSIS (BROOM)

The specimen (SAM-990) is preserved in a soft sandstone matrix (Fig. 1).

The bone is generally not well preserved and is inclined to crumble during

preparation. Much of the specimen (including most of the vertebral column) is

preserved as bone impression, and details of certain of these portions were

obtained from silicone rubber positives. Chief portions of the preserved skeleton

are 17 vertebrae (including, probably, dorsals, sacrals and caudals), an incom-

plete left pubis, the right illum and pubis and both ischia, and the right femur,

fibula and pes. Several other fragmentary bones, mostly seen as impressions,

are scattered through the block: anteriorly a portion of the right scapula blade

can be made out, while impressions of 13 ribs lie ventral to the dorsal vertebrae.

VERTEBRAL COLUMN (Fig. 2)

Altogether 17 vertebrae are preserved, some very incompletely. Eleven of

these are in articulation and consist of a number of dorsals and possibly two

sacrals. A space separates the last of these from the first of the posterior group,

which have been displaced to the right of the anterior series. The space is

sufficient to accommodate three vertebrae of the size of those on each side of

it, and there is thus the possibility that this gap was originally filled by three

sacral vertebrae. However, as will be shown below, it is more likely that a

parting of the vertebral column between two sacral vertebrae occurred prior

to fossilization of the specimen.

The anterior three vertebrae, imperfectly seen, are not in natural articula-

tion, although still in relatively close association with each other. The centrum

of the second vertebra is opisthocoelous and slightly convex anteriorly, while

the third centrum is concave anteriorly and posteriorly. The fourth vertebra has

a procoelous centrum and is markedly convex posteriorly. This convex rear

meets the apparently convex anterior surface of the fifth vertebra’s centrum,

which also appears to be opisthocoelous. The sixth vertebra is provided with

a biconcave centrum, the neural spine is broad in lateral view, and its postero-

dorsal edge overhangs the postzygapophysis so that a posterior embayment is

formed above the postzygapophysis. The seventh vertebra resembles the sixth

and the succeeding eighth in the shape of the neural spine, and both the seventh

and eighth vertebrae nave biconcave centra, similar to that of the sixth vertebra.

The eighth vertebra shows the neural spine clearly, and probably represents

the condition which existed in the less complete sixth and seventh vertebrae.

The spine, posteriorly situated, is antero-posteriorly lengthened and fairly low.

The neural spine of the ninth vertebra is shorter antero-posteriorly. The

ANCHISAURUS CAPENSIS (BROOM) AND A REVISION OF THE ANCHISAURIDAE 125

Fig. 2. Anchisaurus capensis (Broom). SAM-990. A. Vertebral column in lateral view. Broken

edges shown in dashed outline, hatched areas seen in section only. « 0,5. B. Mould of centra

of dorsal vertebrae.

126 ANNALS OF THE SOUTH AFRICAN MUSEUM

tenth vertebra has the only complete neural spine and there is a shallow notch

above the postzygapophysis. The ninth and tenth vertebrae are possibly pro-

coelous, with a convexity on the posterior articular surface.

The posterior group of six vertebrae lies slightly to the right of the anterior

row, and appears to be in near-natural association with the right ilium. The

spine of the most anterior vertebra (the twelfth in the column as preserved)

is fairly high and slightly rounded anteriorly; there is only a slight posterior

notch above the postzygapophysis. The spine resembles that of the tenth (and

eleventh?) vertebra of the anterior row fairly closely. The thirteenth vertebra

has a high and narrower spine, posteriorly inclined, and a chevron can be seen

extending back and down from below its centrum. The fourteenth and fifteenth

vertebrae are similar, as far as can be seen. Remnants of chevrons are seen

between the centra of vertebrae fourteen to seventeen.

Identification of vertebral types

Caudal vertebrae are fairly clearly represented by nos. 13 to 17 in the rear

series, with narrower, obliquely inclined spines and a series of chevrons. The

last two vertebrae (10 and 11) of the anterior row have narrower and possibly

higher spines than the preceding ones, and they resemble the first member of

the posterior group. From this it can be argued that the tenth, eleventh and

twelfth vertebrae of the column as a whole are sacrals, separated by the dis-

integration of the pelvic girdle. Vertebra 12 is in fairly natural association

with the right ilium, and 10 and 11 are close to the ilium of the left side. The

left ilium has been displaced sideways and forward relative to the right side,

and this could account for the gap in the series. Moreover, no recognizable

loose vertebral portions can be identified in the surrounding matrix.

In Efraasia (Galton 1973) and Plateosaurus (Huene 1926) there are fifteen

dorsal vertebrae so, if the above interpretation is correct, the specimen as

preserved probably includes dorsal vertebrae 7 to 15, sacral vertebrae | to 3,

and caudal vertebrae | to 5.

Dimensions of the vertebrae, numbered according to their above identi-

fication, are given in Table 1.

PELVIC GIRDLE

The pelvic girdle is represented by the right ilium, both ischia, and an

incomplete left and almost complete right pubis. The right side of the pelvis

(Figs 3, 5) is preserved in almost natural association. The ilium is well pre-

served, but both pubis and ischium are incomplete distally, and do not make any

clear contact with each other. The areas of articulation between ischium, pubis

and ilium are imperfectly preserved and could not be determined.

Tlium

The ilium (Figs 3A, 5) of the right side is uncrushed and complete except

for a portion of the dorsal crest. It is characterized by long anterior and posterior

ANCHISAURUS CAPENSIS (BROOM) AND A REVISION OF THE ANCHISAURIDAE 127

TABLE 1.

Dimensions of Vertebrae (mm). Anchisaurus capensis (Broom) SAM-990

Length Height of

Length of between pre- spine above

centrum Maximum and post- Dost- Length of

Vertebra ventrally height zygapophysis zygapophysis spine

7th dorsal . ; 2 — — == si:

8th dorsal . 5 29 = — aes ae

9th dorsal . ; +30 — = aoe si

10th dorsal . ; 30 — — we ma

11th dorsal . : 31 — = ps pees

12th dorsal . ; 31 40 — 13 30

13th dorsal . ; 33 42 40 16 32

14th dorsal . : 33 40 48 14 30

15th dorsal . : 32 38 44 13 20

Ist sacral . : Sil 43 39 19 18

2nd sacral . ‘ — — oss has ss

3rd sacral . : — — 35 18 Dill

lst caudal . : 18 SI 34 2) 13

2nd—Sth caudals . +18 — = =o eu

processes, and a pre-acetabular process considerably longer than the post-

acetabular process. The anterior process lies slightly external to the more

posterior surface of the bone, and extends as far forwards as the anterior

edge of the pre-acetabular process. The slender finger-like form of the anterior

process (Fig. 10B) is similar to that of Anchisaurus polyzelus (Fig. 10A) and

Ammosaurus (Galton 1971) and in contrast to the small triangle of other pro-

sauropods (Fig. 10C—D).

A prominent pre-acetabular buttress is developed, arising from close

above the tip of the pre-acetabular process and flaring out laterally before

merging with the body of the ilium at the base of the postacetabular process.

The body of the ilium is expanded to a certain extent above the buttress. The

postacetabular process is considerably shorter than the pre-acetabular, and takes

no part in the formation of the buttress. Above the postacetabular process

the ilium is continued posteriorly as a short crest, medial to the base of the

posterior process. The maximum length of the ilium is 130 mm.

Pubis

The proximal part of the pubis (Fig. 3A) is incomplete, and the areas of

articulation with the ilium and, to a lesser extent, the ischium are not fully

preserved. The ventral edge of the bone is deeply notched below the proximal

end, and this appears to be a natural condition. Below this embayment, which

represents an open obturator foramen, the pubis curves medially and forwards

to terminate as a horizontal plate with an average width of 24 mm. The length

of the pubis, as preserved, is 145 mm.

For prosauropods an open obturator foramen is described to date only

128 ANNALS OF THE SOUTH AFRICAN MUSEUM

ant. proc.

pub. ped.

Fig. 3. Anchisaurus capensis (Broom), SAM-990. A. Pelvic girdle in right lateral view.

B. Ischia in ventral view. > 0,5.

ANCHISAURUS CAPENSIS (BROOM) AND A REVISION OF THE ANCHISAURIDAE 129

in SAM-990 (Fig. 10H) and Anchisaurus polyzelus (Fig. 10G, K); in all other

prosauropods it is enclosed ventrally as in Efraasia (Fig. 10J, N). A similar

open obturator foramen is present in most theropods (for Allosaurus and

Ceratosaurus see Gilmore 1920). Romer (1923) noted that as a result of the

more vertical orientation of the archosaurian femur there is a trend amongst

archosaurs to reduce that part of the pelvis equivalent to the central portion

of the pubo-ischiadic plate of primitive reptiles. The loss of the ventral border

of the obturator foramen in Anchisaurus (Fig. 10G, H, K) and most therapods

probably represents the loss of that portion of the m. pubo-ischio-femoralis

externus 2 which originated ventral to the acetabulum in most prosauropods

and in all sauropods (see Romer 1923: fig. 2, Camarasaurus).

Ischium

Both ischia (Fig. 3), with a preserved length of 136 mm, are present,

but are incomplete posteriorly. In each the widened proximal portion is curved

outwards and carries two embayments, one above a ventral hook-shaped

keel and the other, less clearly defined, lying more dorsally. The proximal

portions of the ischia are separated by an ovoid space, but the shafts are closely

appressed and form a dorsally open trough.

HIND LIMB

Femur

The femur (Fig. 4A), which is seen in dorsal (anterior) view, is broad

and fairly powerful but, with a preserved length of 194 mm, it is incomplete

proximally and distally. The proximal end, as shown by what is still preserved,

was inclined fairly sharply inwards. Below the proximal end the femur is

strongly built and raised to a smooth crest, which runs from proximo-laterally

to disto-medially where it merges into the flat distal end. No condyles are

preserved, and only an indication of the base of the fourth trochanter can

be made out, high in the upper half of the bone.

Fibula

The right fibula (Fig. 4B) lies in its natural position between the femur

and the pes and is seen in lateral (external) view. The proximal half is stoutly

built and leads to a crest formed in the middle of the bone, directed sideways

and forwards. The distal part of the bone is slender and tapers off to the articu-

lation with the pes. Both proximal and distal ends are incomplete and the

preserved length of the bone is 176 mm.

Pes

The pes (Figs 4C, 6, 11K) is seen in ventral view, the bones being either

eroded or indicated by bone impressions. The calcaneum is incomplete laterally,

and probably extended out slightly farther than shown. The astragalus is

transversely elongated, with a rounded anterior surface curving back to the

130 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 4. Anchisaurus capensis (Broom), SAM-990. A. Right femur in dorsal (anterior) view.

B. Right fibula in lateral view. C. Right pes in ventral view. 0,5.

narrow lateral corner of the bone. Both calcaneum and astragalus are pre-

served as impressions, both approximately 15 mm long.

Two small distal tarsal elements are preserved, probably nos 3 and 4.

No. 3 is no more than a bony nodule, while 4 is more robust with posterior

and medial surfaces at right angles to each other and a convex anterior surface

facing metatarsals IV and V. The considerable space between the astragalus

and the proximal ends of metatarsals I and II suggests that the two distal

tarsals are laterally displaced.

All five metatarsals are preserved, in several cases as somewhat imperfect

impressions. There is the usual overlapping of the proximal ends with each

metatarsal slightly underlying its medial fellow. Metatarsal V is short and

narrow but provided with a strong and wide base. Metatarsal IV is about

twice as long and of more or less equal width over its entire length. Meta-

ANCHISAURUS CAPENSIS (BROOM) AND A REVISION OF THE ANCHISAURIDAE 131

tarsal III is the longest and most robust of the series, while II is slightly shorter

than IV. The distal ends of metatarsals III and IV are squarely truncated but

metatarsal II terminates in an oblique surface, so that the medial inclination

of the row of phalanges seems to be a natural one. This could be true, too,

of the short metatarsal I, although this element is incomplete distally.

A small fragment of bone in front of the fifth metatarsal probably repre-

sents a vestigial phalanx. The form and degree of preservation of the phalanges

are apparent from Figures 4C and 6, and from Table 2.

TABLE 2.

Anchisaurus capensis (Broom), SAM—990. Dimensions of pes (in mm).

Metatarsal Phalanx length

length (proximal to distal)

Metatarsal I 44 Digit I 30 34

Il 76 II 30 22 31

Ul 86 Il 30 17 14 30

IV 75 IV 24 16 12 12 21

Vv 48 V =

SLENDER- AND BROAD-FOOTED PROSAUROPODS

When the feet of anchisaurids and plateosaurids are drawn so that digit II

of the manus (Fig. 7) or digit III of the pes (Fig. 8) are reduced to unit length

then two groups are distinguishable, those with slender feet (Figs 7A, C—D, I,

8D-G, 81) and those with broad feet (Figs 7B, E-H, J-Q, 8A—C, H, J-S).

™ ‘ = Wy -

2. Sy,

Fig. 5. Anchisaurus capensis (Broom), SAM-990. Stereophotograph of right ilium.

132 ANNALS OF THE SOUTH AFRICAN MUSEUM

The difference is clearest for the manus (digits II to IV) and, where both fore-

and hind feet are known, the manus and pes are both slender (Figs 7C, E, I,

8D) or broad (Figs 7B; 8H); 7E, 8K; 7G, 88; 75, 8N; 7K, 80; 7Q78@)s ihe

difference between slender and broad feet is not growth related since there

are small prosauropods with broad feet (Figs 7B, 8A—C, H). These appear to

be juveniles of species that grew much larger (Figs 7B, H, K, 8H, O-P—all

Ammosaurus major, see Galton 1971) and all the larger prosauropods are

broad-footed. The prosauropod families Plateosauridae (Figs 7-8) and Melano-

rosauridae (see Bonaparte 1972a: figs 62, 70; Raath 1972: figs 9f, 10a—b) are

all broad-footed, whereas the Anchisauridae as currently classified include

both slender and broad-footed forms (Figs 7-8). ‘Gyposaurus’ capensis (Figs 3C,

5, 7G, 10K) and Thecodontosaurus antiquus (Figs 8F, 11G) are both slender-

footed species but broad-footed species have been incorrectly referred to both

genera.

SYSTEMATIC DISCUSSION

Order SAURISCHIA

Suborder SAUROPODOMORPHA

Infra-order PROSAUROPODA

Family Anchisauridae Marsh, 1885

Diagnosis

Smaller forms, skull lightly built, shallow posterior half of lower jaw with

articulation in line with tooth row, manus and pes slender.

Genus Anchisaurus Marsh, 1885

(includes Megadactylus Hitchcock, Amphisaurus Marsh, Gyposaurus Broom)

Diagnosis

Centra of dorsal vertebrae low, broad bases to neural spines of anterior

caudal vertebrae, ilium with long anterior process, pubis with open obturator

foramen and a relatively narrow distal part that is not apron-like.

The characters of the skull, neck and manus listed below (p. 133) for

Anchisaurus polyzelus may also be diagnostic of the genus, but these regions

are not known in A. capensis.

Anchisaurus polyzelus (Hitchcock, 1865)

Megadactylus polyzelus Hitchcock, 1865: 40, pl. 9 (fig. 6). Cope, 1870: 122A-G, pl. 13 (preocc.).

Amphisaurus polyzelus Marsh, 1882: 84 (preocc.).

Anchisaurus polyzelus Marsh, 1885: 169; 1892: pl. 16 (fig. 3) pl. 17 (fig. 6); 1896: 147, pl. 3

(figs 4-5). Lull, 1915: 119, figs 14-17; 1953: 99, figs 12-14a. Galton, 1971: 782, fig. 7C;

IWS sity AVES VEG Ibe Oy 12s Ay

Anchisaurus colurus Marsh, 1891: 267; 1892: 543, pl. 15, 16 (figs 1, 2); 1893: 169, pl. 6; 1896:

148, pl. 2 (figs 1-3), pl. 3 (figs 1, 2), pl. 4. Huene, 1906: 6, figs 1-6, pls 1-3; 1914b: 69,

figs 1-11; Lull, 1912: 414, figs 2-3; 1915: 130, figs 18-21, pls 4, 10.

Thecodontosaurus polyzelus Huene, 1906: 19, figs 10, 10a; 19145: 75, figs 23-24; 1932: 116.

Yaleosaurus colurus Huene, 1932: 119, pl. 14 (fig. 1), pl. 54 (fig. 3). Lull, 1953: 107, figs 15-18,

pl. 4.

ANCHISAURUS CAPENSIS (BROOM) AND A REVISION OF THE ANCHISAURIDAE 133

Types

Incomplete skeleton (AM 41/109) from Longmeadow Sandstone, upper

part of Newark Series (Upper Triassic) of Springfield, Massachusetts, U.S.A.

(A. polyzelus). Almost complete skeleton (YPM 1883) from Portland beds,

upper part of Newark Series near Manchester, Conn., U.S.A. (A. colurus).

Diagnosis

+ 9 maxillary teeth, 16 dentary teeth, basipterygoid processes very small,

cervical vertebrae elongate, metacarpal I broad, digits If and III of manus

subequal in length, ungual I of pes smaller than ungual II.

Discussion

Megadactylus Hitchcock being preoccupied, Marsh (1882) replaced it with

Amphisaurus (also preoccupied) and then Anchisaurus (Family Anchisauridae

also proposed). Huene (1906) referred the material of Megadactylus polyzelus

Hitchcock to Thecodontosaurus as T. polyzelus (Hitchcock) and used the Family

Thecodontosauridae (originally proposed by Lydekker, 1890: 246) to replace

Anchisauridae. Marsh (1891) made YPM 1883 the holotype of a new species

of Anchisaurus, A. colurus, but he did not indicate how it differed from A. poly-

zelus (AM 41/109). Huene (1906) suggested that Anchisaurus polyzelus resembled

Thecodontosaurus and differed from Anchisaurus colurus in several features,

discussed here together with others noted later by Huene (1907-08, 1932) (for

full discussion see Galton, in press):

1. Shortness of cervical vertebrae. Huene (1932) noted that AM 41/109

resembled Thecodontosaurus (Fig. 11A) in the shortness of the anterior

cervical vertebrae, which are elongate in YPM 1883 (Figs 9H, 11B). How-

ever, this comparison was based on misidentification of part of the neural

arch of a dorsal vertebra (Galton in press, fig. 3a—c) as a cervical vertebra

(Huene 19144: fig. 23a).

2. Shortness of dorsal vertebrae. Huene (1906) originally noted that AM

41/109 differed from Thecodontosaurus and resembled YPM 1883 in having

very elongate dorsal vertebrae. Later Huene (1914) figured an extremely

short centrum of AM 41/109 as that of a dorsal vertebra. Although not

stated, this implied that the dorsal vertebrae of AM 41/109 are extremely

short as in Thecodontosaurus. The isolated centrum figured by Huene

(19146) could not be located but judging from the proportions it was

probably part of an anterior caudal vertebra. The proportions of an

isolated neural arch (Galton in press, fig. 3a—c) and of a centrum (Galton

in press, fig. 5c) show that the dorsal vertebrae of AM 41/109 were probably

elongate (i.e. the centra were low) as in YPM 1883 (Fig. 9K).

3. Slenderness of neural spines of anterior caudal vertebrae. This comparison

by Huene (1906) was based on misleading figures given by Cope (1870)

and Marsh (1893, 1895, 1896). Cope (1870: pl. 8 (fig. 7): see Lull 1953:

134 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 6. Anchisaurus capensis (Broom), SAM-990. Stereophotograph of right pes in

ventral view.

fig. 12a) did not indicate that the neural spines of the anterior caudal

vertebrae of AM 41/109 were damaged and incomplete (Fig. 9P); originally

the neural spines were broader and not as narrow as in the vertebrae

referred to Thecodontosaurus (Fig. 9S) by Huene (1907-08: pl. 77 (fig. 4);

19146: fig. 40).

Most of the supposed differences between AM 41/109 and YPM 1883 were

either the result of misinterpretation (1-3 above, different position of fourth

trochanter of femur) or the result of differences in preservation (form of radius,

metacarpals, tibia, fibula: see Galton in press). YPM 1883 does differ from

AM 41/109 in having a proportionally long centrum to the last dorsal vertebra

(but this might be sacral vertebra 3) and a proportionally smaller manus with

a less trenchant first ungual phalanx (Fig. 7A, C). However, these differences

probably represent individual variations within a species because individuals

of the ornithopod dinosaur Hypsilophodon foxii show a much wider range of

morphological variation (see Galton 1974). YPM 1883 should be referred to

Anchisaurus polyzelus because, on the basis of available material, it cannot be

distinguished from AM 41/109 by any characters of taxonomic significance

and, in addition, AM 41/109 does not show any unique resemblance to Theco-

dontosaurus. Consequently, Anchisaurus colurus Marsh is a junior synonym of

Anchisaurus polyzelus (Hitchcock) and, as A. colurus is the type species of the

135

ANCHISAURUS CAPENSIS (BROOM) AND A REVISION OF THE ANCHISAURIDAE

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136 ANNALS OF THE SOUTH AFRICAN MUSEUM

genus Yaleosaurus (Huene 1932: 122), Yaleosaurus is a junior synonym of

Anchisaurus Marsh, 1885.

Anchisaurus capensis (Broom, 1911)

Hortalotarsus skirtopodus (non Seeley, 1894) Broom, 1906: 201, pl. 3.

Gyposaurus capensis Broom, 1911: 293.

Type

Partial skeleton (SAM-990) from the Cave Sandstone, Stormberg Series

(Upper Triassic) of Ladybrand, Orange Free State, South Africa.

Diagnosis

Ungual | largest on pes.

Discussion

Because of its nature, SAM-990 can be distinguished from Anchisaurus

polyzelus (YPM 1883) oniy by the relative size of ungual | of the pes—large

in SAM-990 and small in YPM 1883. Anchisaurus capensis (Broom) is dis-

tinguishable from all prosauropods other than A. polyzelus (Hitchcock) by

the following combination of characters:

1. Centra of posterior dorsal vertebrae (about the tenth) are proportionally

low so that the ratio of central length to height CG) is 2,1 (Figs 2; 916):

2. Anterior process of ilium is elongate (Figs 3, 5, 10B).

. Subacetabular part of pubis is emarginated ventrally so that the obturator

foramen is open (Figs 3, 10H).

4. The pes is slender (Figs 4C, 6C, 11K).

5. Broad bases to neural spines of anterior caudal vertebrae (Figs 2, 9Q).

oS)

The form of these elements in the species of Anchisauridae is summarized

in Table 3 to facilitate comparisons. The systematic position of other species

incorrectly referred to ‘Gyposaurus’ are discussed below (pp. 141, 143, 147).

TABLE 3.

Comparison of species attributed to ‘Gyposaurus’ with other anchisaurids.

Dorsal Neural Proximal

Centra: spine of Anterior pubis|

length| anterior process of obturator

height caudal ilium foramen Pes

‘“Gyposaurus’ capensis - : Pal wide long shallow/open slender

‘Gyposaurus’ erectus 1,4 wide short deep/closed broad

“Gyposaurus’ sinensis 1,1 — short deep/closed broad

Anchisaurus polyzelus : Dl wide long shallow/open slender

Thecodontosaurus antiquus ed narrow short = slender

Efraasia diagnostica 25) narrow short deep/closed slender

ANCHISAURUS CAPENSIS (BROOM) AND A REVISION OF THE ANCHISAURIDAE 137

Genus Efraasia Galton, 1973

Diagnosis

Basipterygoid processes of medium length, cervical vertebrae elongate,

centra of dorsal vertebrae low, narrow bases to neural spines of anterior caudal

vertebrae, slender metacarpal I, digit I! of the manus robust and appreciably

longer than digit III, ilium with short triangular anterior process, pubis with

closed obturator foramen with an apron-like distal part, ungual | largest in pes.

Efraasia diagnostica (Huene)

Thecodontosaurus diagnosticus Fraas, 1913: 1098 (nomen nudum).

Palaeosaurus (?) diagnosticus Huene, 1932: 52, 73, figs 1, 2, 7-8, pls 4-6.

Palaeosauriscus diagnosticus Charig, 1967: 712.

Efraasia diagnostica Galton, 1973: 247, figs I[A-E, 2-15, 16A, 17C-D.

Syntypes

An almost complete skeleton (SMNS 12667) (Berckhemer 1938) together

with additional material (SMNS 12668) from the Stubensandstein (Upper

Triassic) of Pfaffenhofen, Wiirttemberg, West Germany (see Galton 1973).

Diagnosis

As for genus.

Discussion

As shown by the manus (Fig. 71) and the pes (Figs 8D, 11M), Efraasia

diagnostica undoubtedly represents a slender-footed prosauropod. Apart from

that of Anchisaurus polyzelus (YPM 1883, Fig. 12B; Huene 1906: pl. 1), this

is the only reasonably complete and well-preserved skeleton (Fig. 12A; Berck-

hemer 1938) of a slender-footed prosauropod described to date. Efraasia

diagnostica (Figs 71, 8D, 91, M, 10D, J, N, 11M) resembles Anchisaurus capensis

in several features (Table 3) but differs in three important respects, viz. the

ilium has a short triangular anterior process (Fig. 10D), the subacetabular

part of the pubis is deep with a complete obturator foramen (Fig. 10J, N), and

the bases of the neural spines of the anterior caudal vertebrae are narrow

(Fig. OR).

Genus Thecodontosaurus Riley & Stutchbury, 1836

Diagnosis

At least 21 dentary teeth (in holotype, Fig. 9B); from referred specimens

without teeth diagnosis tentatively expanded as follows: elongate basiptery-

goid processes, cervical vertebrae proportionally short compared with other

anchisaurids, high centra to dorsal vertebrae, narrow base to neural spines of

anterior and caudal vertebrae, high placed deltopectoral crest on proximal

third of humerus, manus with slender metacarpal I and digits II and III subequal

in length, short triangular anterior process to ilium.

2 SCRE

le!

: =e :

Z SS

= .

See = = =

em = OES pa a

ANCHISAURUS CAPENSIS (BROOM) AND A REVISION OF THE ANCHISAURIDAE 139

Thecodontosaurus antiquus Morris

Thecodontosaurus Riley & Stutchbury, 1836: 398; 1840: 352, pl. 29 (figs 1-2).

Thecodontosaurus antiquus Morris, 1843: 211.

Type

Incomplete dentary with teeth from the Magnesian Conglomerate (Upper

Triassic) near Bristol, England.

Diagnosis

As for genus.

Discussion

Riley & Stutchbury (1836, 1840) did not give a specific name for Theco-

dontosaurus and this oversight was rectified by the proposal of T. antiquus

Morris, 1843.

Thecodontosaurus was the first genus of prosauropod to be described

so it is unfortunate that there is no articulated association between teeth of

the type and the postcranial material referred to the genus by Seeley (1895a)

and Huene (1907-8, 19145). Indeed, the only articulated bones referred to

Thecodontosaurus are a few short sequences of vertebrae (Huene 1907-8:

figs 214, 218-220) and a fore limb with scapula, cervical vertebra and dorsal

ribs (Fig. 11A—G). The description and skeletal reconstruction of Thecodonto-

saurus antiquus given by Huene (1932: 116, pl. 54 (fig. 1)) are based on many

specimens. However, the postcranial remains from Bristol indicate the presence

of a slender-footed prosauropod (Fig. 11A—G) and it is reasonable to refer

this material to Thecodontosaurus antiquus. Species of Thecodontosaurus from

other parts of the world (see next section and pp. 145, 147, 152, 153) are incor-

rectly referred to this genus.

Fig. 8. Comparison of the pes in various prosauropods, either right in dorsal view or left in

ventral view (A, C, E, G, I,), all drawn to digit III unit length, scale lines represent 5 cm. These

genera are divided by Romer (1966) between the families Anchisauridae (B, C, E-I, L, N, Q)

and Plateosauridae (M, O, P, R, S). A. Ammosaurus major, YPM 209. B. Aristosaurus erectus,

from Van Hoepen (1920a). C. Thecodontosaurus browni, from Huene (1932). D. Efraasia

diagnostica, SMNS 12668. E. Anchisaurus polyzelus, YPM 1883. F. Thecodontosaurus antiquus,

metatarsal II], from Huene (1907-08). G. Anchisaurus capensis, SAM-990, H. Lufengosaurus

huenei, figured as Gyposaurus sinensis by Young (1941). I. Hortalotarsus skirtopodus, figured

as Thecodontosaurus skirtopodus by Huene (1906). J. Ammosaurus major, YPM 208. K. Ammo-

saurus cf. major, from Galton (1971). L. Massospondylus harriesi, figured as M. browni by

Yan Hoepen (1920b). M. Plateosaurus gracilis, from Berckhemer (1938). N. Massospondylus

harriesi, from Broom (1911). O. Lufengosaurus huenei, figured as Yunnanosaurus magnus by

Young (1947). P. Lufengosaurus huenei, figured as Yunnanosaurus robustus by Young (1951).

Q. Gryponyx africanus, from Broom (1911). R. Plateosaurus robustus, from Huene (1932).

S. Plateosaurus sp., figured as Pachysaurus wetzelianus by Huene (1932).

140 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 9. Comparison of skulls and vertebrae of prosauropods, scale = 2,5 cm, A, B, D-F

x 0,5. A. Skull of Anchisaurus polyzelus, YPM 1883, lateral view. B. Thecodontosaurus anti-

quus, left partial dentary in lateral view, from Riley & Stutchbury (1840). C. Skull of Plateo-

saurus in lateral view, from Romer (1966). D. Massospondylus harriesi, left partial lower jaw

in lateral view, from Haughton (1924). E-G. Braincases in ventral view: E. Anchisaurus

polyzelus, YPM 1883. F. Thecodontosaurus antiquus, YPM 2192. G. Efraasia diagnostica

basisphenoid, SMNS 12667. H—J. Third cervical vertebra in lateral view: H. Anchisaurus

polyzelus, YPM 1883. I. Efraasia diagnostica, SMNS 12667. J. Lufengosaurus huenei, figured

as Gyposaurus sinensis by Young (1941). K—O. dorsal vertebrae (tenth to twelfth) in lateral

view: K. Anchisaurus polyzelus, YPM 1883. L. Anchisaurus capensis, SAM-990. M. Efraasia

diagnostica, SMNS 12667. S. Thecodontosaurus antiquus, from Huene (1907-08).

ANCHISAURUS CAPENSIS (BROOM) AND A REVISION OF THE ANCHISAURIDAE 14]

Anchisauridae nomina dubia

Hortalotarsus skirtopodus Seeley, 1894

Hortalotarsus skirtopodus Seeley, 1894: 411, figs 1-3.

Thecodontosaurus skirtopodus Huene, 1906: 44, figs 72-78, pls 13, 14. Haughton, 1924: 370.

Huene, 1932: 117. Haughton & Brink, 1954: 35.

Gyposaurus skirtopodus Charig, 1967: 712.

Type

Incomplete hind limb in the Albany Museum, from the Cave Sandstone,

Stormberg Series, of Barkly East Division, Cape Province, South Africa.

Discussion

The assignment of Hortalotarsus skirtopodus to Thecodontosaurus by

Huene (1906) was based in part on the characters of isolated bones found at

localities different from that of the type specimen. The pes of the type (Fig. 81)

is obviously that of a slender-footed prosauropod, and Huene (1906) could

not distinguish it from the pes of Thecodontosaurus. However, this pes (Fig. 81)

is also indistinguishable from those of Anchisaurus capensis (Fig. 11K), Anchi-

saurus polyzelus (Fig. 11L) and Efraasia diagnostica (Fig. 11M). This specimen

is generically and specifically indeterminate so Hortalotarsus skirtopodus

Seeley is a nomen dubium.

Thecodontosaurus browni (Seeley, 18956)

Massospondylus browni Seeley, 18956: 118, figs 13-14.

Thecodontosaurus browni Huene, 1906: 141, pl. 12 (figs 7-8); 1932: 118. Broom, 1911: 293.

Haughton, 1924: 370.

Type

Limb bones (BMNH R3302) from the Red Beds, Stormberg Series of Telle

River, Herschel, Cape Province, South Africa.

Discussion

Seeley (18954) noted that the proportions of the phalanges of the pes are

very similar to those of Hortalotarsus (Fig. 8H) so this is probably another

generically and specifically indeterminate specimen of a_ slender-footed

prosauropod.

Thecodontosaurus minor Haughton, 1918

Thecodontosaurus minor Haughton, 1918: 468; 1924: 376, fig. 21.

Thecodontosaurus browni: Huene, 1932: 118.

Type

Left tibia, a cervical vertebra and a portion of a left ilium (SAM-3451)

from the Red Beds, Stormberg Series, from road-cutting at Naude’s Nek,

Pitsing, Maclear District, Cape Province, South Africa.

Discussion

The tibia is slender so this may be a slender-footed prosauropod but, on

142 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 10. Comparison of ilium and pubis of prosauropods. Scale = 2,5 cm. A—F. Lateral view

of left ilium: A. Anchisaurus polyzelus, YPM 1883. B. Anchisaurus capensis, SAM-990. C.

Lufengosaurus huenei, figured as Gyposaurus sinensis by Young (1941). D. Efraasia diagnostica,

SMNS 12667. E. Aristosaurus erectus, fromVan Hoepen (1920a). F. Thecodontosaurus antiquus,

from Huene (1907-08). G—J. lateral view of left pubis: G. as A. H as B. Las C. J as D. K-N.

Ventral view of right pubis (L, M) or dorsal view of left pubis (K, N): K as A, G. Las C, I.

M as E. N as D, J.

the basis of available material, this specimen is a generically and specifically

indeterminate prosauropod and Thecodontosaurus minor is a nomen dubium.

Thecodontosaurus macgilivrayi (Seeley)

Agrosaurus macgilivrayi Seeley, 1891: 161, figs 1-6.

Thecodontosaurus macgilivrayi Huene, 1906: 147, figs 86-90; 1932: 52.

Type

Tibiae, radius, an ungual and tooth (BMNH 49984) from York Peninsula,

Queensland, Australia.

Discussion

Huene (1932: 52) subsequently referred Agrosaurus macgilivrayi to the

Theropoda (Coelurosauria) as do Romer (1956, 1966), Steel (1970) and White

(1973). However, the latero-distal surface of the tibia is notched (Fig. 11H—J)

(Huene 1906: fig. 86a, d-e) to receive the central ascending process of the

astragalus in typical prosauropod fashion. This material undoubtedly represents

ANCHISAURUS CAPENSIS (BROOM) AND A REVISION OF THE ANCHISAURIDAE 143

a prosauropod but is generically and specifically indeterminate, so Agrosaurus

macgilivrayi is a nomen dubium; it may represent a slender-footed prosauropod.

PROSAUROPOD SPECIES INCORRECTLY ASSIGNED TO ANCHISAURIDAE

Family Plateosauridae Marsh, 1895

Diagnosis

Larger forms, skull massively built, deep posterior half to lower jaw with

articulation offset ventral to line of tooth row, manus and pes broad.

Ammosaurus major Marsh, 1889

Anchisaurus major Marsh, 1889: 331, fig. 1.

Ammosaurus major Marsh, 1891: 267; 1892: 545, pl. 16 (fig. 4), pl. 17 (fig. 3); 1896: 150,

pl. 3 (figs 3, 6). Huene, 1906: 15, pls 5-9; 1907-08: 303-04, figs 297-298; 1914a: 13;

19145: 74, figs 20-22; 1932: 26. Lull, 1915: 148, figs 24-25; 1953: 123, figs 19-20. Galton,

1971: 786, figs 9, 11A.

Anchisaurus solus Marsh, 1892: 545; 1896: 149. Huene, 1914b: 72, figs 12-19. Lull, 1915:

144, figs 22-23; 1953: 120.

Anchisaurus (?) solus Huene, 1906: 14, pl. 4.

Ammosaurus solus Huene, 1932: 27, pl. 49 (fig. 1).

Type

Pelvis and hind limbs (YPM 208) from the Portland Beds, upper part of

Newark Series near Manchester, Connecticut, U.S.A. (Ammosaurus major).

Almost complete skeleton (YPM 209) from the same locality and horizon

(Anchisaurus solus).

Discussion

Ammosaurus has long been regarded as a primitive theropod dinosaur

but it is considered as an anchisaurid by Steel (1970) and by Galton (1971),

who provides a detailed discussion of the taxonomic position of this genus

(Galton in press). The pes of the holotype (Fig. 8J) and of the referred specimens

(Fig. 8A, K) plus a referred manus (Fig. 7E) are of the broad type. Ammo-

saurus is a broad-footed prosauropod characterized by the following combina-

tion of characters: centra of dorsal vertebrae low, slender sacral rib 3, elongate

anterior process to ilium, subacetabular part of the ischium emarginated

ventrally (Galton, in press).

Aristosaurus erectus van Hoepen, 1920a

Aristosaurus erectus yan Hoepen, 1920a: 82, figs 1-6, pls 9-10. Haughton 1924: 379, Haughton

& Brink, 1954: 33.

Gyposaurus capensis Huene, 1932: 123, pl. 54 (fig. 2).

Gyposaurus erectus Charig, 1967: 712. Steel, 1970: 49.

Type

An almost complete skeleton as slab and counterpart (TM 130) from the

Cave Sandstone, Stormberg Series, near Roosendal, Senekal District, Orange

Free State, South Africa.

144 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 11. Thecodontosaurus, Agrosaurus and comparisons of the anchisaurid pes. Scale = 5 cm,

A-J + x 0,45. A-G. Thecodontosaurus antiquus, YPM 2195, from Durdham Down, Bristol,

England. Matrix indicated by stipple, broken bone by diagonal shading: A. Right side of

anterior cervical vertebra. B. Right scapula in lateral view. C. Left humerus in medial view.

D. As C, in anterior view. E. Left ulna in proximal, lateral and distal views. F. Left radius

in lateral view. G. Left manus in lateral or dorsal view. H-J. Agrosaurus macgillivrayi, distal

end of left tibia in lateral view (H), anterior view (1), and distal view (J), all from Huene (1906).

K-—M. Anchisaurid pes, drawn to digit III unit length: K. Anchisaurus capensis, SAM-990,

compare with Fig. 5. L. Anchisaurus polyzelus, YPM 1883. M. Efraasia diagnostica,

SMNS 12668.

ANCHISAURUS CAPENSIS (BROOM) AND A REVISION OF THE ANCHISAURIDAE 145

Discussion

Romer (1956, 1966), Charig (1967), Steel (1970) and White (1973) follow

Huene (1932, 1956) in regarding Aristosaurus as a junior synonym for Gypo-

saurus but this is unlikely because, in contrast to the situation in Gyposaurus

capensis (Table 3):

1. In photographs of the skeleton of Aristosaurus erectus in Van Hoepen

(1920a: pls 9-10) and in the reconstruction (Fig. 12D) given by Huene

(1932) the dorsal vertebrae are proportionally higher with a central

length to height ratio of about 1,4 for dorsal 10.

2. The anterior process of the ilium is short (Fig. 10E).

3. The obturator foramen of the pubis is closed ventrally (Fig 10M).

4. The hind feet of Aristosaurus erectus appear to be of the broad type.

There is no reason why Aristosaurus erectus should be referred to the

genus Anchisaurus. Aristosaurus erectus appears to be a valid taxon of broad-

footed prosauropod, but further preparation and illustration of the holotype

is needed.

Genus Massospondylus Owen, 1854

(includes Leptospondylus Owen, Pachyspondylus Owen, Aetonyx Broom,

Gryponyx Broom, Dromicosaurus Van Hoepen)

Massospondylus carinatus Owen, 1854

Massospondylus carinatus Owen, 1854: 97. Seeley, 1895b: 102, figs 1-12. Huene, 1906: 36,

figs 43-70, pls 13-16. Broom, 1911: 241. Haughton, 1924: 383. Huene, 1932: 124.

Leptospondylus capensis Owen, 1854: 97.

Pachyspondylus orpenii Owen, 1854: 97.

Type

Isolated bones from the Red Beds, Stormberg Series of Beaucherf, Harri-

smith, Orange Free State, South Africa. The holotype in the Museum of the

Royal College of Surgeons in London was destroyed during World War II

but casts of this material are in the National Museum of Southern Rhodesia,

Bulawayo (J. Attridge, pers. comm.).

Massospondylus harriesi Broom, 1911

Massospondylus harriesi Broom, 1911: 299, pls 15-17. Haughton, 1924: 384, figs 21-29.

Huene, 1932: 125.

Massospondylus browni (non Seeley, 1895h): Van Hoepen 19204: 118, pls 17-22.

Aetonyx palustris Broom, 1911: 304, figs 20-23. Haughton, 1924: 404, fig. 30. Huene, 1932: 91.

Gryponyx africanus Broom, 1911: 294, figs 1-9. Haughton, 1924: 417, figs 36-38. Huene,

1932: 88, pl. 7 (figs 1-4).

Gryponyx taylori Haughton, 1924: 420, fig. 39. Huene, 1932: 90.

Dromicosaurus gracilis Van Hoepen, 1920b: 103, figs 8-21, pls 13-16. Haughton, 1924: 405.

Thecodontosaurus dubius Haughton, 1924: 377.

146 ANNALS OF THE SOUTH AFRICAN MUSEUM

Types

Massospondylus harriesi: Bones of the fore limb (SAM-3394) from the top

of the Red Beds, Stormberg Series of Foutanie, Fouriesburg, Orange Free

State, South Africa.

Aetonyx palustris: Partial skeleton (SAM-2768, 2769, 2770) from the top

of the Red Beds (but listed as from Cave Sandstone by C. E. Gow in Anderson

& Anderson 1970), Stormberg Series from Foutanie, Fouriesburg, Orange

Free State, South Africa.

Gryponyx africanus: Pelvis and hind limb, right and left manus, vertebrae

(SAM-3357-9) from the top of the Red Beds (but listed as from the Cave

Sandstone by C. E. Gow in Anderson & Anderson 1970), Stormberg Series of

Foutanie, Fouriesburg, Orange Free State, South Africa.

Gryponyx taylori: Pelvic girdle and sacral vertebrae (SAM-3453) from the

top of the Red Beds (but listed as from Cave Sandstone by C. E. Gow in Ander-

son & Anderson 1970), Stormberg Series of Fouriesburg, Orange Free State,

South Africa.

Dromicosaurus gracilis: Partial skeleton (TM 123) from Red Beds of

Naaupoort Nek, Bethlehem, Orange Free State, South Africa.

Thecodontosaurus dubius: Larger portion of a skeleton (SAM-3712) from

the Cave Sandstone, Stormberg Series of Ladybrand, Orange Free State,

South Africa.

Discussion

On the basis of the phalanges Seeley (1895b) stated that Massospondylus

carinatus had a broad hind foot and Huene (1906) separated this genus from

Plateosaurus mainly because of its Thecodontosaurus-like tibia. Massospondylus

was the first genus of broad-footed prosauropod to be described from South

Africa.

The manus (Fig. 7F—-G, J) and the pes (Fig. 8L, N) of Massospondylus

harriesi (SAM-3394) are obviously of the broad type. The material (Figs 7G,

8L, 12E) described by Van Hoepen (19204) as Massospondylus browni should

be referred to this species (Haughton 1924). The manus of Aetonyx palustris

Broom (Fig. 7O) is of the broad type and the pes is similar, as indicated by

the measurements given by Huene (1932: 92). J. Attridge (pers. comm.) regards

Aetonyx palustris as a junior synonym for Massospondylus harriesi. The manus

(Fig. 7Q) and pes (Fig. 8Q) of Gryponyx africanus are of the broad type and

Gryponyx africanus is probably a junior synonym for Massospondylus harriesi.

Gryponyx taylori Haughton is a nomen dubium because the material is

generically and specifically indeterminate; it probably represents another

specimen of Massospondylus harriesi.

The manus and pes of Dromicosaurus gracilis Van Hoepen are not pre-

served but were probably of the broad type because Dromicosaurus was regarded

as being closely allied to Aetonyx and Massospondylus by Van Hoepen (19205),

Haughton (1924) and Huene (1932). J. Attridge (pers. comm.) regards Dromico-

ANCHISAURUS CAPENSIS (BROOM) AND A REVISION OF THE ANCHISAURIDAE 147

saurus gracilis as a junior synonym for Massospondylus harriesi. The type

specimen of Thecodontosaurus dubius Haughton has never been figured, but

Huene (1932: 92) referred it to Aetonyx palustris so it 1s presumably a broad-

footed form; J. Attridge (pers. comm.) refers this specimen to Massospondylus

harriesi.

Stratigraphically and geographically, Massospondylus harriesi is the most

ubiquitous prosauropod in southern Africa.

Plateosaurus gracilis (Huene, 1907)

Thecodontosaurus (?) hermannianus Huene, 1907-08: 216, fig. 236, pl. 144 (fig. 1).

Plateosaurus gracilis Huene, 1932: 303.

Type

Right maxilla with teeth from the Stubensandstein (Upper Triassic) of

Heslach, in Stuttgart, West Germany.

Lufengosaurus huenei Young, 1941a

Lufengosaurus hueni Young, 1941a: 1, figs 1-25, pls 1-6; 1947: 41; 1951: 50, fig. 11, pl. 12.

Rozhdestvensky, 1965.

Gyposaurus sinensis Young, 1941b: 205, pls 1-9; 1948: 91, pls 1-5; 1951: 49.

Yunnanosaurus huangi Young, 1942: 64, figs 1-17; 1951: 56.

Lufengosaurus magnus Young, 1947: 2, figs 1-14.

Yunnanosaurus robustus Young, 1951: 58, figs 12-14, pls 7-10.

Types

Several incomplete skeletons from the lower Lufeng Series (Upper Triassic,

Rhaetic) of Lufeng, Yunnan, China.

Discussion

Rozhdestvensky (1966) restudied 70 specimens from the Lufeng Series

and decided that Lufengosaurus huenei, L. magnus, Yunnanosaurus huangi,

Y. robustus and Gyposaurus sinensis of Young were all differentiated only on

size-related characters and are conspecific (as Lufengosaurus huenei Young,

1941a). Rozhdestvensky (1966) noted that, judging from the original diagnosis

of Young (19414), Gyposaurus sinensis is hardly distinguishable from G. capensis,

but that without visual comparisons or more detailed descriptions it is impos-

sible to decide the relationship between these two species. Galton (1973, in

press) noted that ‘Gyposaurus’ sinensis was incorrectly referred to the genus

Gyposaurus, and may represent a new genus or be based on juveniles of either

Lufengosaurus or Yunnanosaurus. ‘Gyposaurus’ sinensis should not be referred

to the genus Gyposaurus (or Anchisaurus) because of the following anatomical

features (Table 3):

1. Centra of posterior dorsal vertebrae are proportionally high with a

central length to height ratio of 1,1 (Fig. 90).

2. Anterior process of ilium is short and triangular (Fig. 10C).

3. Subacetabular part of pubis is broken but originally this region was

148 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 12. Skeletal reconstructions of prosauropods. Scale = 20 cm. Tails of A and B diagram-

matically folded over. A. Efraasia diagnostica, SMNS 12667, 12668, from Galton (1973).

B. Anchisaurus polyzelus, YPM 1883, AM 41/109, from Galton (1973). C. Lufengosaurus huenei,

based on figures of individual bones given by Young (19415) as Gyposaurus sinensis.

ANCHISAURUS CAPENSIS (BROOM) AND A REVISION OF THE ANCHISAURIDAE 149

. eh

‘eg ae Se PC Ree |

LE aati

~~ et:

WES ace >

Fig. 12. (cont.)

D. Aristosaurus erectus, modified from Gyposaurus capensis of Huene (1932). E. Massospondylus

harriesi, modified from Thecodontosaurus browni of Huene (1932), based on specimen described

as Massospondylus browni by Van Hoepen (1920a).

deep with a complete obturator foramen (Young 1941b: 222; 1948: 96)

(Figs 10I, L).

4. Pes (Fig. 8H) and manus (Fig. 7B) are broad.

Young (19414) did not cite the papers of Broom (1906, 1911) so the assign-

ment as Gyposaurus sinensis was probably based on the skeletal reconstruction

of Aristosaurus erectus given by Huene (1932) as Gyposaurus capensis. However,

the skeleton of ‘Gyposaurus’ sinensis (Fig. 12C) differs greatly in several aspects

from that of Aristosaurus erectus (Fig. 12D) and this is especially true for the

form of the neck vertebrae (Fig. 9J) and fore limb. It should be noted that the

skeletal reconstruction of ‘Gyposaurus’ sinensis given by Young (19415, pl. 9)

bears practically no resemblance to a reconstruction (Fig. 12C) based on

figures of the bones of the same specimen given by Young (19410). The correct-

ness of the contention of Rozhdestvensky (1966) that Gyposaurus sinensis

150 ANNALS OF THE SOUTH AFRICAN MUSEUM

Young, 1941b is a junior synonym of Lufengosaurus huenei Young, 1941a is

shown by the proportionally short neck and proportionally massive manus

of the two types.

Prosauropoda nomina dubia

Gryponyx transvaalensis Broom, 1912

Gryponyx transvaalensis Broom, 1912: 82, figs 3-4. Van Hoepen, 1920b: 102. Haughton,

1924: 420. Huene, 1932: 91, pl. 7, fig. 5.

Type

Ungual | of the manus and a metatarsal (in the Transvaal Museum) from

the Bushveld Sandstone (Cave Sandstone), Stormberg Series of Wiepe 1258,

northern Transvaal, South Africa.

Discussion

This material is probably prosauropod but is generically and specifically

indeterminate.

NON-PROSAUROPOD SPECIES INCORRECTLY ASSIGNED TO ANCHISAURIDAE

Order SAURISCHIA

Suborder THEROPODA

Arctosaurus osborni Adams, 1875

Arctosaurus osborni Adams, 1875: 177. Lydekker, 1889: 352.

Type

Isolated cervical vertebra (NMI 62 1971) from Heiberg Formation (Upper

Triassic) of north-west extremity of Cameron Island, Bathurst Group, Arctic

Archipelago, Canada.

Discussion

Arctosaurus was described as reptilian by Adams (1875) but subsequently

Lydekker (1889) referred it to the family Anchisauridae. Regarded as a turtle

by Huene (1906) and White (1973) but referred to the prosauropod family

Melanorosauridae (as Plateosauravidae) by Huene (1956) and to the Anchi-

sauridae (as Thecodontosauridae) by Romer (1966). The region of the dia-

pophysis is slightly damaged (Fig. 13B, F) but from the adjacent curves of the

neural arch (Fig. 13E) it is obvious that the diapophysis was very small and,

as a result, this vertebra is from the anterior part of the series and is probably

either the third or fourth cervical vertebra. It is proportionally very much

shorter than the equivalent vertebrae of Anchisaurus (Figs 9H, 12B), Efraasia

(Figs 9I, 12A) and Plateosaurus (Huene 1926). The only prosauropods with

cervical vertebrae proportionally as short are Thecodontosaurus (Fig 11A) and

Lufengosaurus (Figs 9J, 12C) but in both cases the vertebrae are proportionally

much lower, the neural spines are not so well developed and there is no pleuro-

coel (exaggerated in Arctosaurus because of crushing). Arctosaurus osborni is

ANCHISAURUS CAPENSIS (BROOM) AND A REVISION OF THE ANCHISAURIDAE 15]

Fig. 13. Arctosaurus osborni Adams, holotype NMI G2 1971, anterior cervical vertebra in:

A. Anterior view. B. Left lateral view. C. Posterior view. D. Ventral view. E. Dorsal view.

F. Stereo photograph of left side, compare with B. Scale = 2,5 cm. Broken bone indicated

by diagonal shading.

not a prosauropod and, on the basis of the general form-of the vertebra and

the presumed presence of a pleurocoel, this specimen is tentatively regarded as

Theropoda incertae sedis as listed by Steel (1970).

Ischisaurus cattoi Reig, 1963

Ischisaurus cattoi Reig, 1963: 10, figs 4B, 5. Colbert, 1970: 27. Bonaparte, 1972b: 673, fig. 22.

Type

Two incomplete skeletons from the Ischigualasto Formation (lower Upper

Triassic) of Argentina.

Discussion

Ischisaurus is listed as an anchisaurid by Steel (1970) and as Saurischia

incertae by Bonaparte (1972b: 674), who notes that ‘the suggested affinities

SZ ANNALS OF THE SOUTH AFRICAN MUSEUM

with the Coelurosauria appears as one of the possible relationships’. Romer

(1966) and Colbert (1970) list Zschisaurus as a coelurosaurian theropod.

Order THECODONTIA

‘Thecodontosaurus’ gibbidens Cope, 1878

Thecodontosaurus gibbidens Cope, 1878: 177. Huene, 1921: 571, figs 14-15.

Type

Isolated teeth from the Upper Triassic of Pennsylvania, U.S.A.

Discussion

The isolated teeth are almost circular rather than oval in cross-section as

in prosauropods (Thecodontosaurus antiquus Riley & Stutchbury, 1840; Anchi-

saurus, YPM 1883; Plateosaurus. AMNH 6810), so these teeth are provisionally

referred to the Ornithischia (Galton, in press).

Spondylosoma absconditum Huene, 1935

Spondylosoma absconditum Huene, 1935: 247, pl. 30, figs 1-13. Charig, 1967: 712. Colbert,

1970: 19. Bonaparte, 19726: 674.

Type

Scapula, humerus, femur, tibia (all incomplete) and eight vertebrae from

the Santa Maria Formation (Upper Triassic) of Brazil.

Discussion

Huene (1935, 1942) regarded Spondylosoma as a saurischian but did not

make a more specific assignment for this genus. Romer (1956, 1966) referred

Spondylosoma to the Anchisauridae (as Thecodontosauridae) as did Colbert

(1970) and Charig (1967), who noted at the same time the possibility of its

being a prestosuchid pseudosuchian. Bonaparte (19725: 674) notes that ‘there

are doubts regarding its assignment to the Saurischia, or even to Prosauropoda.

Unfortunately there are not sufficient diagnostic pieces to define better its

taxonomic position’ and Spondylosoma is listed as Saurischia incertae.

Teleocrater alphos (Haughton, 1932)

Thecodontosaurus (?) alphos Haughton, 1932: 662, fig. 19.

Teleocrater alphos Charig, 1967: 712.

Type

Two cervical vertebrae (SAM-10654) from the Manda Formation (Upper

Triassic, Anisian) of Tanzania.

Discussion

Charig refers this material to the pseudosuchian Te/leocrater.

ANCHISAURUS CAPENSIS (BROOM) AND A REVISION OF THE ANCHISAURIDAE 153

Subclass LEPIDOSAURIA

Order LACERTILIA

Huene (1932, not 1931 as cited by Steel 1970) considered that the follow-

ing Middle Triassic (Muschelkalk) species were based on specimens which

should be referred to the eosuchian Tanystropheus, a lacertilian according to

Wild (1974); the non-prosauropod nature of these species is also discussed by

Colbert (1970):

Tanystropheus primus (Huene, 1907-08)

Thecodontosaurus primus Huene, 1907-08: pl. 42 (figs 8-9).

Tanystropheus primus Huene, 1932: 6.

Tanystropheus latespinatus (Huene, 1907-08)

Thecodontosaurus (?) latespinatus Huene, 1907-08: figs 237-245.

Tanystropheus latespinatus Huene, 1932: 6.

NOTES ON THE FAMILIES ANCHISAURIDAE AND PLATEOSAURIDAE

The infra-order Prosauropoda is currently divided into three families:

Anchisauridae (= Thecodontosauridae), Plateosauridae and Melanorosauridae

(see Romer 1956; Colbert 1964; Charig et al. 1965; Bonaparte 1972a). Post-

cranially the separation is clearest between melanorosaurids and non-

melanorosaurids (Romer 1956: 617; Bonaparte 1972a: 160). Galton (1971,

1973) suggests that the range of morphological variation is insufficient to

warrant the retention of two families of non-melanorosaurid prosauropods.

However, the skulls of Anchisaurus (Fig. 9A) and Plateosaurus (Fig. 9C) are

very different and, because of this, Galton (in press) now considers that they

should not be included in the same family. Fortunately the genera concerned

are the basis for the first two valid prosauropod family names to be proposed:

Anchisauridae Marsh, 1885, and Plateosauridae Marsh, 1895.

In only one case (Anchisaurus polyzelus, YPM 1883) is a well-preserved

skull found in natural association with a skeleton of a slender-footed pro-

sauropod. Consequently the referral of Efraasia and Thecodontosaurus to the

Family Anchisauridae is tentative. Contrary to the impression given by Huene

(1932: fig. 7; 1956: fig. 10), the skull of Efraasia is very incomplete but, as

noted by Galton (1973), Efraasia is an ideal ancestor for the more recent

Anchisaurus. The holotype of Thecodontosaurus antiquus 1s an incomplete

dentary but, judging from what is preserved (Fig. 9B), the complete lower

jaw was probably more like that of Anchisaurus (Fig. 9A) than that of Plateo-

saurus (Fig. 9C). Although considered unlikely, the discovery of additional

material may show that the restriction of the family Anchisauridae to slender-

footed forms is artificial. However, the criterion is practical and with it most

taxa and specimens of non-melanorosaurid prosauropods are readily referable

to either the Family Anchisauridae or the Family Plateosauridae.

Plateosaurus engelhardti Meyer, 1837 from the Keuper (Upper Triassic)

154 ANNALS OF THE SOUTH AFRICAN MUSEUM

of Germany is the earliest taxon of plateosaurid to be described, but since

then a multitude of taxa have been erected for material from the Triassic of

Germany (Steel (1970) lists 9 species as Plateosaurus, 8 species as Gresslyo-

saurus). However, this material is probably extremely overclassified and all

the taxa of European plateosaurids listed by Steel (1970: 53-56) should be

provisionally regarded as junior synonyms of Plateosaurus engelhardti Meyer,

1837. All the European plateosaurid material can probably be referred to (at

the most) three species of Plateosaurus, but a restudy of all the holotypes is

needed to determine the other valid species.

Lufengosaurus (see Young 194la, b, 1942, 1947, 1951) and Plateosaurus

(see Huene 1907-8, 1926, 1932) possess the features listed above as charac-

teristic of plateosaurids. The skulls of Ammosaurus (see Galton, in press) and

Aristosaurus (see Van Hoepen 1920a) are not well enough preserved to tell

anything about the form of the skull. However, the holotype of Massospondylus

harriesi includes a lower jaw (Fig. 9D), the posterior part of which is deep

with the articulation offset ventral to the line of the tooth row. J. Attridge is

studying two skulls of Massospondylus harriesi (SAM- K388 and K1314) and

has found that both skulls show the features listed above (pers. comm.).

Massospondylus is the most ubiquitous prosauropod in southern Africa

and its previous classification as an anchisaurid made the prosauropod fauna

of Africa unique, because in other areas with abundant, well-preserved pro-

sauropod skeletons, plateosaurids are the most common form. However,

Cox (1973: 213) notes that ‘it is clear that land connections between all the

continents existed for much, at least, of the Triassic’. As regards prosauropods

the presence of Anchisaurus in North America and South Africa and the presence

of Plateosaurus in Germany and South America (Casamiquela 1964; Bonaparte

19725) indicated that this was the case for the continents on either side of the

Atlantic. With the transfer of broad-footed forms, previously listed under the

Anchisauridae, to the Plateosauridae, this family becomes the dominant and

cosmopolitan prosauropod family of the world. In marked contrast, the Anchi-

sauridae have an extremely restricted fossil record (total of about 10 articulated

specimens for North America, Europe and South Africa) with no remains

discovered to date from Asia (Young 1951; Rozhdestvensky 1966) or South

America (Bonaparte, pers. comm.). Haughton (1924) noted that the Stormberg

Series of South Africa was deposited under conditions of progressively increasing

aridity, and it is interesting that skeletal remains of melanorosaurids occur in

the lowermost levels (Passage Beds, Charig et a/. 1965; basal Red Beds,

Haughton 1924), most plateosaurid skeletons occur higher in the Red Beds,

and those of anchisaurids are found in the overlying Cave Sandstone (Charig

et al. 1965; Haughton 1924: Haughton & Brink 1956). Charig et al. (1965)

report the presence of small tridactyl footprints in the Passage Beds and in the

lower Red Beds and note (p. 204) that these ‘. . . may indicate the movement

of thecodontosaurids from one upland region to another via a lowland area’.

The world-wide rarity of anchisaurid skeletal remains is presumably because

ANCHISAURUS CAPENSIS (BROOM) AND A REVISION OF THE ANCHISAURIDAE 155

these species occupied the drier upland areas, which are rarely represented in

the fossil record.

SUMMARY

The holotype (SAM-990) of the prosauropod dinosaur Anchisaurus

capensis (Broom, 1911) is characterized by the following combination of

characters: low centra to posterior dorsal vertebrae, broad bases to neural

spines of anterior caudal vertebrae, ilium with long anterior process, pubis

with open obturator foramen, and with slender pes with first ungual phalanx

the largest. SAM-990 differs from the North American Anchisaurus polyzelus

(Hitchcock) only in the large size of the first ungual of the pes, and SAM-990

is referred to Anchisaurus Marsh, 1885 as Anchisaurus capensis (Broom). Taxa

which have been incorrectly referred to the genus Gyposaurus (= Anchisaurus)

are Gyposaurus erectus (Van Hoepen) (= Aristosaurus erectus Van Hoepen),

Gyposaurus sinensis Young (= Lufengosaurus huenei Young) and Gyposaurus

skirtopodus (Seeley) (= Hortalotarsus skirtopodus Seeley, nomen dubium). The

Family Anchisauridae is restricted to those species with slender feet, viz. Anchi-

saurus polyzelus, A. capensis, Efraasia diagnostica, and Thecodontosaurus

antiquus. Taxa with broad feet previously classified as anchisaurids (Aristo-

saurus, Ammosaurus, Gyposaurus sinensis (as Lufengosaurus huenei), Masso-

spondylus (including Aetonyx, Dromicosaurus, Gryponyx africanus, G. taylori,

Thecodontosaurus dubius), Yunnanosaurus (= Lufengosaurus) are transferred to

the family Plateosauridae, the dominant and cosmopolitan family of

prosauropods.

ACKNOWLEDGEMENTS

P. M. Galton is grateful to the following for the loan or use of specimens

from their respective institutions, listed in alphabetical order: Drs W. Coombs,

Jr and A. E. Wood, Amherst College Museum, Amherst, Massachusetts;

Drs E. H. Colbert and E. S. Gaffney, American Museum of Natural History,

New York; Dr A. J. Charig, British Museum (Natural History), London;

C. E. O’Riordan, National Museum of Ireland, Dublin; Dr K. D. Adam,

Staatlichen Museum fiir Naturkunde in Stuttgart; Drs J. H. Ostrom and E. L.

Simons, Peabody Museum, Yale University, New Haven, Connecticut. Galton

thanks Mr J. Attridge, Birkbeck College, University of London; Dr D. Baird,

Princeton University, New Jersey and Dr J. Bonaparte, National University

of Tucuman, Argentina for helpful discussions and personal communications

as cited above, and Drs J. H. Ostrom and E. L. Simons of Yale University

for the continued use of the research facilities of the Division of Vertebrate

Paleontology of the Peabody Museum of Natural History. The photographs

were taken by Mr N. J. Eden, Department of Palaeontology, South African

Museum (Figs 1, 5, 6) and Mr Alan Coleman, Yale Peabody Museum (Fig. 13F).

156 ANNALS OF THE SOUTH AFRICAN MUSEUM

Support for this research was provided to the senior author by the Connecticut

Research Commission (Grant no. RSA 680) while at Yale University and by

a Faculty Research Grant from the University of Bridgeport.

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Mag. nat. Hist. 2: 468-469.

HAuGHTON, S. H. 1924. The fauna and stratigraphy of the Stormberg series.— Ann. S. Afr.

Mus. 12: 323-497.

HauGutTon, S. H. 1932. On a collection of Karroo vertebrates from Tanganyika Territory.

QO. Jl. geol. Soc. Lond. 88: 634-671.

HAUGHTON, S. H. & Brink, A. S. 1954. A bibliographic list of Reptilia from the Karroo Beds

of Africa.—Palaeont. afr. 2: 1-187.

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Abh. (N.F.) 8: 99-156.

HueENE, F. von. 1907-1908. Die Dinosaurier der Europaischen Triasformation mit Beruck-

sichtigung der aussereuropdischen Vorkomnisse.— Geol. paldont. Abh. (Suppl.) 1: 1-419.

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silium Cat. (Animalia) 4: 1-21.

HUuENE, F. von. 1914b. Nachtrage zu meinen fruheren Beschreibungen triassischer Saurischia.

—Geol. paldont. Abh. (N.F.) 13: 69-82.

HUuENE, F. von. 1921. Reptilian and stegocephalian remains from the Triassic of Pennsylvania

in the Cope Collection.— Bull. Am. Mus. nat. Hist. 44: 561-574.

HUuENE, F. von. 1926. Vollstandige Osteologie seines Plateosauriden aus dem Schwabischen

Keuper.— Geol. paliont. Abh. (N.F.) 15: 134-180.

HUENE, F. von. 1932. Die fossile Reptil-Ordnung Saurischia, ihre Entwicklung und Geschichte.

— Monogrn Geol. Paldont. (1) 4: 1-361.

HUuENE, F. von. 1935-1942. Die Fossilen Reptilien des Sudamerikanischen Gondwanalandes.

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HUuENE, F. von. 1956. Palaontologie und Phylogenie der niederen Tetrapoden. Jena: Fischer.

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geol. nat. Hist. Sury. 81: 1-331.

LYDEKKER, R. 1890. Catalogue of fossil reptilia and amphibia. Part 4. London: British Museum

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Marsh, O. C. 1885. Names of extinct reptiles.— Am. J. Sci. 37: 331-336.

Marsh, O. C. 1889. Notice of new American Dinosauria.— Am. J. Sci. 37: 331-336.

Marsh, O. C. 1891. Notice of new vertebrate fossils.— Am. J. Sci. 42: 265-269.

Marsu, O. C. 1892. Notes on Triassic Dinosauria.— Am. J. Sci. 43: 543-546.

Marsh, O. C. 1893. Restoration of Anchisaurus.— Am. J. Sci. 45: 169-170.

Marsh, O. C. 1895. On the affinities and classification of the dinosaurian reptiles.— Am. J. Sci.

50: 483-498.

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(Mesotriasico Superior) de las Provincias de San Juan y la Rioja (Republica Argentina).

—Ameghiniania 3: 3-20.

158 ANNALS OF THE SOUTH AFRICAN MUSEUM

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Cape Colony.— Ann. Mag. nat. Hist. 14: 411-419.

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ABBREVIATIONS

acetabulum acet

anterior process of ilium ant proc

astragalus as

centrum Cc

caudal vertebra ca

calcaneum cal

chevron ch

dorsal vertebra dor

neural spine ns

pleurocoel - pl

postzygapophysis poz

prezygapophysis prz

ANCHISAURUS CAPENSIS (BROOM) AND A REVISION OF THE ANCHISAURIDAE 159

pubic peduncle pub ped

sacral vertebra sac

Amherst College Museum, Amherst, Massachusetts AM

American Museum of Natural History, New York AMNH

British Museum (Natural History), London BMNH

National Museum of Ireland, Dublin NMI

South African Museum, Cape Town SAM

Staatlichen Museum fiir Naturkunde in Stuttgart SMNS

Yale Peabody Museum, New Haven, Connecticut YPM

6. SYSTEMATIC papers must conform with the International code of zoological nomenclature

(particularly Articles 22 and 51).

Names of new taxa, combinations, synonyms, etc., when used for the first time, must be

followed by the appropriate Latin (not English) abbreviation, e.g. gen. n., sp. n., comb. n.,

syn. n., etc.

An author’s name when cited must follow the name of the taxon without intervening

punctuation and not be abbreviated; if the year is added, a comma must separate author’s

name and year. The author’s name (and date, if cited) must be placed in parentheses if a

species or subspecies is transferred from its original genus. The name of a subsequent user of

a scientific name must be separated from the scientific name by a colon.

Synonymy arrangement should be according to chronology of names, i.e. all published

scientific names by which the species previously has been designated are listed in chronological

order, with all references to that name following in chronological order, e.g.:

Family Nuculanidae

Nuculana (Lembulus) bicuspidata (Gould, 1845)

Figs 14-15A

Nucula (Leda) bicuspidata Gould, 1845: 37.

Leda plicifera A. Adams, 1856: 50.

Laeda bicuspidata Hanley, 1859: 118, pl. 228 (fig. 73). Sowerby, 1871: pl. 2 (figs 8a—b).

Nucula largillierti Philippi, 1861: 87

Leda bicuspidata: Nicklés, 1950: 163, fig. 301; 1955: 110. Barnard, 1964: 234, figs 8-9.

Note punctuation in the above example:

comma separates author’s name and year

semicolon separates more than one reference by the same author

full stop separates references by different authors

figures of plates are enclosed in parentheses to distinguish them from text-figures

dash, not comma, separates consecutive numbers

Synonymy arrangement according to chronology of bibliographic references, whereby

the year is placed in front of each entry, and the synonym repeated in full for each entry, is

not acceptable.

In describing new species, one specimen must be designated as the holotype; other speci-

mens mentioned in the original description are to be designated paratypes; additional material

not regarded as paratypes should be listed separately. The complete data (registration number,

depository, description of specimen, locality, collector, date) of the holotype and paratypes

must be recorded, e.g.:

Holotype

SAM-—A13535 in the South African Museum, Cape Town. Adult female from mid-tide region, King’s Beach,

Port Elizabeth (33.51S, 25.39E), collected by A. Smith, 15 January 1973.

Note standard form of writing South African Museum registration numbers and of date.

7. SPECIAL HOUSE RULES

Capital initial letters

(a) The Figures, Maps and Tables of the paper when referred to in the text

*...1in C. namacolus (Fig. 10)...”

(b) The prefixes of prefixed surnames in all languages, when used in the text, if not preceded

by initials or full names

e.g. Du Toit but A. L. du Toit

Yon Huene _— but _-F. von Huene

e.g. Therocephalia, but therocephalian

Punctuation should be loose, omitting all not strictly necessary

Reference to the author should be expressed in the third person

Roman numerals should be converted to arabic, except when forming part of the title of a

book or article, such as

“Revision of the Crustacea. Part VIII. The Amphipoda.’

Specific name must not stand alone, but be preceded by the generic name or its abbreviation

to initial capital letter, provided the same generic name is used consecutively.