The ANNALS OF THE SOUTH AFRICAN MUSEUM

are issued in parts at irregular intervals as material

becomes available

Obtainable from the South African Museum, P.O. Box 61, Cape Town

(Cash with order, post free)

Die ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

word uitgegee in dele op ongereelde tye na beskikbaarheid

van stof

Verkrygbaar van die Suid-Afrikaanse Museum, Posbus 61, Kaapstad

(Kontant met bestelling, posvry)

OUT OF PRINT/UIT DRUK

I, 2(1, 3, 5, 7-8), 3(1-2, 5, t-p.i.), 5(2, 5, 7-9);

63 tp) 7(15 3), 8, g(1-2), 10(1-3),

I1(1-2, 7> t.p.i.), 21, 24(2), 31(1-3), 44(4)-

Price of this part/ Prys van hierdie deel

50c

Printed in South Africa by In Suid-Afrika gedruk deur

The Rustica Press Pty., Ltd., Die Rustica-pers Edms., Bpk.,

Court Road, Wynberg, Cape Courtweg, Wynberg, Kaap

THE SKULL OF STRUTHIOCEPHALUS KITCHINGI

LIEUWE DIRK BOONSTRA

South African Museum, Cape Town

(with 11 figures in the text)

CONTENTS

PAGE

Introductionw aa uimesal ec nee 51

Description 5 on 606) .6 ARR

DISCUSSION ETE aoe eee L ne 205

Summative yee) are eee 2

Acknowledgements. . . . 265

IRGUAKANGES 9 5 gs aya) (ot POR

INTRODUCTION

Since Brink’s description in 1958 of the type skull (B.P.I. 284) found on

De Bad, Beaufort West by Kitching, the author with H. Zinn and H. Boonstra

had the good fortune to excavate a second skull (S.A.M. K272) on Perdefontein,

Beaufort West in 1960.

This skull was found zn situ lying upside down with parts of the lower

jaws exposed and the rest intact in a mudstone matrix. Unfortunately the

exposed lower jaws have suffered from weathering which has destroyed the

posterior parts of both rami. The palatal surface of the skull was firmly encased

in a jacket of reinforced plaster and the massive skull could be lifted in one

piece. On preparation the skull proved to be very well preserved and only

very slightly distorted.

On study it became apparent that this specimen shows a number of

structural features much better than the type specimen does. Certain additions

to Brink’s description can thus be made and in the sequel it will also become

clear that I differ in interpretation on a number of points, even if allowance

is made for considerable individual variation.

A correction has also to be made in regard to the type species due to

Brink having misread Broom. The species whaitsi has as its holotype the skull

(S.A.M. 2678) from Vivier, Beaufort West and the second specimen mentioned

by Broom consists of a skull and much of the skeleton (S.A.M. 3012) from

Abrahamskraal, Prince Albert. Broom rightly thought that the second specimen

belonged to the same species as the holotype skull. There are thus two skulls

and one skeleton.

251

Ann. S. Afr. Mus. 48 (14), 1965, 251-265, 11 figs.

252 ANNALS OF THE SOUTH AFRICAN MUSEUM

DESCRIPTION

Build and carriage of the skull

The fronto-nasal boss in my specimen is very like that of the type, but I

do not think that it represents a horn-core. In the collection of the South

African Museum there are over two dozen skulls of Struthiocephalus and Kerato-

cephalus. These all have a fronto-nasal boss developed to a varying degree and

in all of them the sculpturing of the boss is very similar to that of the skull in

general. If the boss were a horn-core one would have expected its surface to

differ from that of the general skull sculpturing. That the boss could have

been used as a battering ram is most probable, but then without a special

horn covering.

There is no doubt about the hang-dog carriage of the skull in Struthio-

cephalus. As a matter of fact this applies to all the tapinocephalians in general

—also to those without a naso-frontal boss. The struthiocephalines, with their

long snouts and anteriorly directed upper front teeth, undoubtedly fed on

softer vegetable matter than the moschopines. Their limbs have also been

shown to be more adapted to marshy conditions and Brink’s suggestion that

the struthiocephalines may even have fed duck-like under water is quite

probable. The surface moulding of the bone around the nostrils—especially

of the septomaxilla—suggests the presence of musculature for the closing off

of the nostril.

The structure of the skull

Although the present skull is very well preserved and prepared a number

of sutures cannot be traced with absolute certainty. This is due to a number

of factors such as the rugose nature of the outer surfaces, closure of sutures,

fusion of elements and small displacements. When comparing the figures

given here with those of the type skull, and also with those of other species of

the genus the effects of the pachyostosis should be born in mind. The pachy-

ostosis in the tapinocephalians is to some extent individual with differences

often seen between the relations and extent of the two bones of a pair in the

same skull. Age is also a factor. Differences that have been given as specific

are often due to differences in the tempo of the pachyostosis in adjoining bones.

Thus a strong pachyostotic development in, for instance, the postfrontal,

causes an overlap or overgrowth over the adjoining bones and affects the

relative size and shape of the outer surfaces of these bones. Where, however,

authors show a radical difference in the relations of bones errors of observation

and/or interpretation must be considered probable.

Lateral and dorsal surfaces. (Figs. 1 and 2)

In comparing my figures with those of Brink it is manifest that the type

skull has been subjected to dorso-ventral compression. The effects are especially

obvious in the nature and disposition of the lateral pterygoid flange and the

THE SKULL OF STRUTHIOCEPHALUS KITCHINGI 253

quadrate. It should, moreover, be borne in mind that my figures are ortho-

projections and not perspective drawings. This would in part account for the

fact that in my figures the tabular is visible in dorsal view and in lateral view

forms much of the posttemporal bar. But in my specimen the tabular is really

much more developed and in its forward growth overlaps much of the lateral

Fic. 1. Struthiocephalus kitchingi. S.A.M. K272 x 1/6. Lateral view. Orthoprojection

on to the sagittal plane.

AN—angular. AR —articular. D—dentale. eam —external auditory meatus.

F—frontal. 1P—inter- or postparietal (dermo-supraoccipital). J—jugal. L—lacri-

mal. M—maxilla. N—nasal. P—parietal, PAL—palatine. PM—premaxilla.

PO—postorbital. POF—postfrontal. PRF—prefrontal. PT—pterygoid. Q—

quadrate. QJ—quadratojugal. SA—surangular. SM—septomaxilla. smf—septo-

maxillary foramen. SQ—squamosal. T—tabular. TR —transversum (ecto-

pterygoid).

254 ANNALS OF THE SOUTH AFRICAN MUSEUM

tongue of the parietal which is wedged in between the tabular and the upsweep-

ing process of the squamosal. In my specimen the postorbital is more developed

and this at the expense of the postfrontal. This particularly affects the

appearance of the postorbital bar as seen in dorsal view.

Fic. 2. Struthiocephalus kitchingi. S.A.M. K272 x 1/6. Dorsal view.

Orthoprojection on to the alveolar plane.

Brink figures the squamosal as entering the ventral orbital border, whereas

in my specimen the jugal extends posteriorly, ventral to the postorbital and

forms nearly all of the ventral rim of the orbit with a small contribution by

the postorbital. This is the normal relation, not only in the tapinocephalids

THE SKULL OF STRUTHIOCEPHALUS KITCHINGI 255

but also in all the other Dinocephalia (anteosaurids, titanosuchids, styraco-

cephalids and the Russian brithopids). Brink’s figure indicates an error in

observation.

In the present specimen the outer surface of both septomaxillaries is

well preserved and this shows a distinctive moulding of the posterior border

of the nostril to form a rounded swollen rim set off by the presence of a lateral

groove. I suggest that this moulded structure indicates attachments for a

valvular closure of the nostril when the animal is feeding with the snout

submerged.

Occiput. (Fig. 3)

The occiput is low and broad with the squamosals bulging laterally to

form prominent ‘cheeks’. A strong rounded thickened upper and lateral border

formed by the interparietal and the tabulars prominently demarcates a deep

bipartite area of origin for the nuchal muscles. The interparietal and the

tabulars have large posterior faces, whereas the supraoccipital is very low. The

large face of the paroccipital is directed much more ventrally than posteriorly.

The ridge on the squamosal bounding the external auditory meatus medially

is very prominent. The posttemporal fossa is all but closed by the downgrowth

Fic. 3. Struthiocephalus kitchingi. S.A.MM. K272 x 1/6. Occipital

view. Orthoprojection at right angles to the sagittal and alveolar

planes.

BO—basioccipital. EO—exoccipital. fq—quadrate foramen.

jf—jugular foramen. mp—mastoid process of the paroccipital.

pin—pineal (parietal) foramen. POC—paroccipital. ptf—post-

temporal fenestra. ptpocf—pterygo-paroccipital fenestra.

SO —supraoccipital. ST —stapes. stf—stapedial foramen.

Other lettering as for Fig. 1.

256 ANNALS OF THE SOUTH AFRICAN MUSEUM

of the tabular. The stapes is seen to lie diagonally with the distal end lying

low down in the recess on the quadrate. Little is seen of the posterior face of

the quadrate in this view because of its horizontal disposition. The occipital

condyle is directed much ventrally with the exoccipitals forming most of the

posterior face and the basioccipital facing mainly ventrally.

Ventral surface. (Fig. 4)

Comparing my figure of the ventral surface to that given by Brink a

number of differences are apparent, apart from the fact that mine is a projection

and Brink’s a perspective drawing. The differences are in main due to the fact

that the present specimen being uneroded is in a much better state of preserva-

tion. The regions mainly affected are the transverse processes of the pterygoids,

quadrate and stapes, the occipital condyle and the relations of the supra-

occipital, interparietal, tabular and squamosal.

In the present specimen the lateral flanges of the pterygoids form well

demarcated deep transverse ridges extending far ventrally in their lateral

parts, lying far below the level of the quadrate rami.

In its lateral part the quadrate ramus forms a deep vertical flange of

bone which meets the quadrate along a large synchrondrotic face. More

medially the pterygoid is deeply vaulted and sends a process posteriorly which

abuts against the paroccipital at a level higher than the quadrate process of

the paroccipital.

The quadrates are well preserved and carry cotyli shaped as shown in the

figure. The stapedial recess is well preserved on both sides.

Both stapes are well preserved and are seen to lie diagonally with the

distal ends lying well anterior to the plane of the fenestrae ovales. The occipital

condyle is a prominent large knob roughly circular in outline with its articular

face directed only slightly posterior off the ventral. The exoccipitals form

more of the articular face than is shown in Brink’s figure.

The paroccipital has a large ventral face. Medio-anteriorly it has a

process meeting the prodtic above the level of the fenestra ovalis. Latero-

anteriorly there is a long quadrate process applies to the quadrate and ter-

minating posterior to the stapedial recess. At a higher level the quadrate

process of the paroccipital has a thin flange of bone whose anterior edge meets

a process of the quadrate ramus of the pterygoid. Posterolaterally the par-

occipital develops an everted thickened edge underlying the squamosal to

form a mastoid process. The supraoccipital is wide but low and the posttemporal

fossa all but obliterated by the overgrowth of the large paroccipital. The

tabular has a large ventral face which, however, does not extend anteriorly

between the paroccipital and squamosal as shown by Brink in his figure of the

type.

The squamosal carries a prominent ridge, roughly comma-shaped in

outline, and this forms the internal limit of the external auditory meatus

developed as an antero-posteriorly directed groove.

THE SKULL OF STRUTHIOCEPHALUS KITCHINGI 257

fm Ptf

Fic. 4. Struthiocephalus kitchingi. S.A.M. K272 x 1/6. Ventral

view. Orthoprojection on to the alveolar plane.

fm—foramen magnum. fo—fenestra ovalis. ic—internal

carotid foramina. iptv—interpterygoid vacuity. PBS—para-

basisphenoid. PROT —prodtic. qp—quadrate process of the

paroccipital. rt—replacing teeth. V—vomer. Other lettering

as in previous figures.

The position of the stapes. (Fig. 5)

In order to describe the relations of the stapes to the bones with which it

makes contact I have sketched the various bones disarticulated but lying in

their relative positions.

258

ANNALS OF THE SOUTH AFRICAN MUSEUM

Fic. 5. Struthiocephalus kitchingi. S.:A.M. K272 x 1/3. The stapes and

contiguous bones as disarticulated. Ventral view. Orthoprojection on

to the alveolar plane.

con—condyles of the quadrate. cpt—contact of pterygoid (quadrate

ramus) with the quadrate, paroccipital and para-basisphenoid. cpot—

contact of paroccipital with the proétic. cq—contact of quadrate with

the pterygoid and the stapes. csq—contact of the squamosal with

the quadrate and the quadratojugal. np—notochordal pit in the

basioccipital.

Other lettering as in previous figures.

1— wedge of the pterygoid applied to the medial edge of the quadrate

(4)-

2—area on the posterior process of the quadrate ramus of the pterygoid

overlapping the distal end of the stapes.

3 —surface of the quadrate overlapping the stapes dorsal to the stapedial

recess proper.

4—area on the medial edge of the quadrate making contact with the

posterior process of the quadrate ramus of the pterygoid.

5—surface of the quadrate underlapped by the paroccipital.

6—area on the dorsal process of the stapes in contact with the

paroccipital (7).

7—area on the paroccipital in contact with the dorsal process of the

stapes (6).

8—area on the paroccipital underlapped by the stapes.

g—area on the paroccipital underlapped by the proximal end of the

stapes.

10—area of the proétic underlapped by the stapes.

11—hollow in the distal end of the stapes which receives a process of

the quadrate.

THE SKULL OF STRUTHIOCEPHALUS KITCHINGI 259

The footplate of the stapes lies fitted into the fenestra ovalis on both sides

so that its proximal end cannot be seen.

Antero-dorsally the proximal end of the stapes is applied to a face on

the prodtic and further posteriorly underlies a face on the paroccipital. Both

these contacts are not very intimate—definitely not synostotic—but rather

syndesmotic or synchondrotic. Ventrally the footplate is in contact with the

edges of the fenestra ovalis formed by the paroccipital, basioccipital and the

para-basisphenoid.

The distal end of the stapes makes contact with the quadrate and

paroccipital.

Postero-laterally the distal end of the stapes has a fairly short truncated

process—the tympanic process—whose tip is connected directly or by the

intercalation of a cartilaginous extra-stapes to the tympanum.

Antero-medially of this tip the distal end of the stapes fits into an elongated

groove-like stapedial recess in the quadrate, lying in a plane nearly parallel

to the quadratic cotyli. The fit is not tight and movement in this diagonal plane

is possible even with the presence of cartilage or connective tissue.

The anterior part of the distal surface of the stapes has an oval hollowed-out

articulatory face which makes contact with a process of the quadrate lying

anterior to the elongated stapedial recess and posterior to the surface of contact

with the end of the quadrate process of the pterygoid.

On the postero-distal surface of the stapes, posterior and medial to the

tympanic process, and at a higher level than the main shaft of the stapes lies

the dorsal process of the stapes, which makes contact with the quadrate process

of the paroccipital. This contact was apparently synchrondrotic.

Stapes. (Fig. 6)

In ventral view the stapes presents a main portion consisting of a proximal

end forming the footplate, a greatly expanded distal end and a shaft with a

waist-like constriction. From the posterior edge of this main portion the rest

of the bone lies at a higher level. About halfway along the shaft a large oval

stapedial foramen pierces this flange. Postero-distally of the foramen there is a

process which meets the paroccipital—this is the dorsal process.

The distal end bears a process directed postero-distally—this is the

tympanic process. Anterior to the tip of the tympanic process the distal end

has an elongated face which fits into the stapedial recess in the quadrate.

Anterior to this the distal end presents a concave oval articulating face,

which articulates with a convex process on the quadrate lying immediately

posterior to the contact face on the quadrate, which receives the quadrate

process of the pterygoid.

In dorsal view the stapes has the surface of its main portion and the

surface of the dorsal process lying in the same plane. On the upper face of the

dorsal process there rises a conical protuberance with its tip directed medially.

This I have labelled the medial dorsal process. The function of this process

260 ANNALS OF THE SOUTH AFRICAN MUSEUM

Br ieee Ss ae nes

. eo U ie t Bao Og

Euan eR ROR anit 7 ip =P Opec ier

‘

A B Cc D

Fic. 6. Right stapes of Struthiocephalus kitchingi. S.A.M. K272 x 1/3.

A~—anterior. B—posterior. C—dorsal. D—ventral. FPOC—facet on

the dorsal process which is applied to the quadrate process of the

paroccipital. FQ facet fitting into the elongated stapedial recess in

the quadrate. FQP—concave facet applied to a convex process on the

quadrate lying medial to the stapedial process proper. MDP—medial

dorsal process. PDP—proximal dorsal process. STF—stapedial

foramen. ‘TY—tympanic process.

would appear to be to receive a tendon probably attached to the inner edge .

of the quadrate.

In anterior and posterior view this median dorsal process is seen to be

both strong and prominent.

Quadrate. (Fig. 7)

The quadrate is a large bone—robust in its cotylar region and in the

parts making contact with the squamosal, quadratojugal and the quadrate

ramus of the pterygoid. But its inner portion overlying the paroccipital consists

of a thin sheet of bone with a free edge.

Its stout pterygoid process bears a large roughly oval area for the reception

of the quadrate ramus of the pterygoid, which at a higher level has an additional

process meeting the quadrate in a groove on the lower part of the inner edge

of the quadrate. The articulatory area on the pterygoid process is a hollowed

out area with, in its middle part, a longitudinal ridge. This is matched on the

quadrate ramus of the pterygoid by an articulating face bearing a median

groove flanked by two longitudinal ridges.

The nature of this joint is such that it admits of movement between the

quadrate and the pterygoid in a parasagittal plane. The roughness of both

articulating faces indicates the presence of synchondrotic cartilage.

Just above the quadrate-pterygoid articulation there is a transverse

everted flange of bone. The upper edge of this flange carries a transverse

groove which thus lies between two lips. This groove is the stapedial recess.

The outer end of this everted flange develops a thin free standing process to

which the tympanum was probably attached.

261

THE SKULL OF STRUTHIOCEPHALUS KITCHINGI

*(rodord ssasaq Terpodejs) sodeys ay} Jo uotdada4 ay} Joy ssodar payeSuoja 9y3—Y_LS ‘sedvis oy} jo puo [eistp

OY} UO JOR} DAVOUOD 9Y} YIM UOLV[NoIL Joy sso901d XaAUOI— JG “sse004 Terpade}js ay} Jo diy 101193s0d ay} Sururs0y 9Spo]—T. L$

‘Tesnfoyerpenb oy} Jo pus yesyUsA 9y} BUTATII—I ssov0I—[OY “Uoues0j ayerpenb ay} jo so9ps0q JQuUI 9Y} Suro; yoI0U—NO

‘oyerpenb oy} jo ssav01d prosAiajd— gd ‘aApuoo [eustoyx9—9Q ‘afApuoo [eusajyuI—Dy “proSA193d oy} Jo snures ayerpenb oy}

SUTATO0a1 ayerpenb oy} Jo ssooo1d prosd193d oy} uo yoorJ—].qaq ‘Tesourenbs oy} YIM yowzUOD IOy sovjins—QOsSD ‘[esnfojespenb

24) YIM JowIUCD Jo sovJIns— [OD ‘prosdsajd ay} Jo snurest oyerpenb ay} jo ssooo1d sJoraysod ay} Suratsoa1 9}e1penb ay} Jo aSpa

[erp oy} UO so"jINsS—T[dD ‘sliepnqrpueu-nides ‘ur ay} jo yred Sutsnoy sovjAns (101sayUe) [esIOp UI 9ACOIS—Hy *(TeUOA

Aqjeorsojoydiour) ro11eyue—( ‘Terpeur Jo 1guUT—r) *(s0L19} Ue ATTeotS0;oydsour) ;es1op—g *(aor1za3sod Ayjeorso;oydasout) peryusA —y

E/1 xX Loy WeV'S “Wumysny snjoydasoryynys jo ayexrpenb yySry °L ‘ory

2) ) 20 230 2} )

262 ANNALS OF THE SOUTH AFRICAN MUSEUM

In between the quadrate-pterygoid joint and the outer lip the everted

flange has a rounded process to which is articulated the hollow face on the

distal end of the stapes.

On its posterior face the quadrate is greatly thickened where it is applied to

the squamosal and quadratojugal. Medial to this thickened area lies a wide

groove with a smooth surface. This housed part of the body of the m. capiti-

mandibularis.

A moschopid and jonkeriid quadrate. (Figs. 8 and 9)

I am including comparable figures of a quadrate of a moschopid and a

jonkeriid. Essentially these are of the same type as that of Struthzocephalus,

except that in the jonkeriid quadrate a very definite recess is developed on the

posterior face well above the stapedial recess. I can offer no suggestion as to

what was received in this recess.

STR

STL

PTP

Ic oc IC

Fic. 8. Left quadrate of Moschops sp. S.A.M. 11701 X 1/3.

The Tympanum

A depressor mandibuli muscle originating from the prominent ridge on

the squamosal just medial of the groove housing the external auditory meatus

and inserted on the process of the articular would allow ample room for a

tympanum with a diameter of about 30 mm.

The tympanum having one point of attachment on the free tip of the

lip of the ledge bounding the stapedial recess could lie in a parasagittal plane

and receive the tympanic process of the stapes meeting it at right angles.

The external auditory meatus extending from the tympanum along the

groove in the posterior face of the squamosal could have its opening just above

the level of the upper limit of the squamosal ridge.

The nature of the joint of the stapes with the quadrate as described above

would allow of movement of sufficient amplitude for the conduction of sound

waves.

THE SKULL OF STRUTHIOCEPHALUS KITCHINGI 263

Fic. 9. Right quadrate of Jonkeria haughtoni. S.A.M. 4343 X 1/3.

Dentition. (Fig. 10)

Brink maintains that in the type specimen the marginal teeth are disposed

in a double row both functional at the same time. This view is manifestly

incorrect. The fact is that the replacing teeth arise lingually of those in use

and each new tooth is thus a younger member of the same tooth family of

264 ANNALS OF THE SOUTH AFRICAN MUSEUM

Cc D

Fic. 10. First left incisor of Struthiocephalus kitchingi.

S.A.M. K272 x 4.

A— posterior. B—anterior. C—labial view.

D—lingual view. H—heel. R—root. T—talon.

the tooth it replaces. The position is thus as in the Titanosuchia. The structure

of the teeth is of the talon-and-heel type thoughout the series, with decrease in

size in posterior direction.

Fic. 11. Skull of Moschosaurus longiceps. Type.

S.A.M. 3015 xX 1/6.

A—dorsal. B—lateral.

THE SKULL OF STRUTHIOCEPHALUS KITCHINGI 265

Discussion

In a recent paper I have suggested that all the described species

of Struthiocephalus could well be considered conspecific and I arranged them in

a growth series.

I would now go further and suggest that Moschosaurus could very well be

the youngest form of such a series. (Fig. 11)

SUMMARY

A detailed description of the skull of Struthiocephalus kitchingi is given,

based on a second skull from Beaufort West. This specimen shows a number of

features much better than the type specimen and leads to some differences in

interpretation.

ACKNOWLEDGEMENTS

The skull here described was obtained on a collecting trip by the South

African Museum which was in part financed by a grant from C.S.I.R. to

whom we tender our thanks.

The Trustees of the South African Museum are grateful to the Council

for Scientific and Industrial Research for a grant to publish this paper.

REFERENCES

Boonstra, L. D. 1963. Diversity within the South African Dinocephalia. S. Afr. F. Sct. 59:

196-207.

Brink, A. S. 1959. Struthiocephalus kitchingi sp. nov. Palaeont. Afr. 5: 39-56.

INSTRUCTIONS TO AUTHORS

MANUSCRIPTS

In duplicate (one set of illustrations), type-written, double spaced with good margins,

including TaBLE oF ConTENTs and Summary. Position of text-figures and tables must be

indicated.

ILLUSTRATIONS

So proportioned that when reduced they will occupy not more than 4? in. X 7 in. (7% in.

including the caption). A scale (metric) must appear with all photographs.

REFERENCES

Authors’ names and dates of publication given in text; full references at end of paper in

alphabetical order of authors’ names (Harvard system). References at end of paper must be

given in this order:

Name of author, in capitals, followed by initials; names of joint authors connected by &,

not ‘and’. Year of publication; several papers by the same author in one year designated by

suffixes a, b, etc. Full title of paper; initial capital letters only for first word and for proper

names (except in German). Title of journal, abbreviated according to World list of scientific

periodicals and underlined (italics). Series number, if any, in parenthesis, e.g. (3), (n.s.), (B.).

Volume number in arabic numerals (without prefix ‘vol.’), with wavy underlining (bold type).

Part number, only if separate parts of one volume are independently numbered. Page numbers,

first and last, preceded by a colon (without prefix ‘p’). Thus:

SmirH, A. B. 1956. New Plonia species from South Africa. Ann. Mag. nat. Hist. (12) 9: 937-945.

When reference is made to a separate book, give in this order: Author’s name; his initials;

date of publication; title, underlined; edition, if any; volume number, if any, in arabic numerals,

with wavy underlining; place of publication; name of publisher. Thus:

Brown, X. Y. 1953. Marine faunas. 2nd ed. 2. London: Green.

When reference is made to a paper forming a distinct part of another book, give: Name of

author of paper, his initials; date of publication; title of paper; ‘In’, underlined; name of

author of book; his initials; title of book, underlined; edition, if any; volume number, if any,

in arabic numerals, with wavy underlining; pagination of paper; place of publication; name

of publisher. Thus:

SmirH, C. D. 1954. South African Plonias. Jn Brown, X. Y. Marine faunas. and ed. 3: 63-95.

London: Green.

SYNONYMY

Arranged according to chronology of names. Published scientific names by which a species

has been previously designated (subsequent to 1758) are listed in chronological order, with

abbreviated bibliographic references to descriptions or citations following in chronological

order after each name. Full references must be given at the end of the paper. Articles and

recommendations of the International code of zoological nomenclature adopted by the XV International

congress of zoology, London, July 1958, are to be observed (particularly articles 22 and 51).

Examples: Plonia capensis Smith, 1954: 86, pl. 27, fig. 3. Green, 1955: 23, fig. 2.

When transferred to another genus:

Euplonia capensis (Smith) Brown, 1955: 259.

When misidentified as another species:

Plonia natalensis (non West), Jones, 1956: 18.

When another species has been called by the same name:

[non] Plonia capensis: Jones, 1957: 27 (= natalensis West).