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ANNALS OF THE SOUTH AFRICAN MUSEUM

ANNALE VAN DIE SUID-AFRIKAANSE MUSEUM

Volume 72 ~ Band

January 1977 Januarie

Part 6 Deel

NEW PROCOLOPHONIDS

FROM THE TRIASSIC CYNOGNATHUS ZONE

OF SOUTH AFRICA

C. E. GOW

Cape Town Kaapstad

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NEW PROCOLOPHONIDS FROM THE TRIASSIC CYNOGNATHUS

ZONE OF SOUTH AFRICA

C. E. Gow

Bernard Price Institute for Palaeontological Research, Johannesburg

(With 9 figures)

[MS. accepted 11 August 1976]

ABSTRACT

Previously only scraps of procolophonids have been known from the Cynognathus zone.

One is redescribed here and four new species are described: Thelegnathus oppressus, T. perfor-

atus, T. contritus and T. spinigenis. Taxonomy is based on tooth morphology, the species

being adapted for specialized herbivorous diets. One of the new species exhibits Captorhinus-

like tooth replacement. The suggestion is made that procolophonids were upland animals.

CONTENTS

PAGE

Introduction’ 23) sane een OS)

Systematics and description. . . 110

DISCUSSION eae cat nee enemy,

SUMMAKVAl eae Ue teen nel 23)

Acknowledgements: = 2 3.3. . . 4123

IRCLETENCES a uecaar iene ere 124

INTRODUCTION

Broom (1905) described a new genus and species of procolophonid from

the Cynognathus zone which had been collected by Alfred Brown; this he named

Thelegnathus browni. The specimen was reconsidered and illustrated thirty years

later (Broom 1936). Since that time the specimen has been further prepared and

will be illustrated and further described in detail below.

Recently Kitching has collected ten more procolophonids from three locali-

ties in the Cynognathus zone; these are all skulls or partial skulls, some with a

little associated postcranial skeleton. Detailed examination of particularly the

teeth of these animals leads to the conclusion that there is a group of species

present all broadly related to Thelegnathus. Owing to the small number of

specimens this overall relationship will be assumed to be at the generic level for

present purposes. The species exhibit various specializations for an herbivorous

diet.

In this paper maxillary teeth are referred to as molars. Tooth replacement

terminology follows Edmund (1960).

109

Ann. S. Afr. Mus. 72 (6), 1977: 109-124, 9 figs.

110 ANNALS OF THE SOUTH AFRICAN MUSEUM

SYSTEMATICS AND DESCRIPTION

Class REPTILIA

Subclass ANAPSIDA

Order COTYLOSAURIA

Family Procolophonidae

Thelegnathus

Diagnosis

Procolophonids in which the maxillary teeth exhibit progressive mesio-

distal broadening posteriad.

Thelegnathus browni Broom, 1905

Rigo

Holotype

SAM-5869, a left maxilla containing six teeth.

Locality

Unknown, but from a Cynognathus zone bone breccia.

Revised diagnosis

The curve of the occlusal plane and the nature of tooth wear are specifically

distinctive.

Etymology

Named for the collector, Alfred Brown.

Description

The six molar teeth progressively increase in size posteriad. The teeth have

undergone some polishing post mortem and before incorporation in the breccia.

Fig. 1. Thelegnathus browni SAM-5896. The type maxilla in

labial and occlusal views.

NEW PROCOLOPHONIDS FROM THE TRIASSIC CYNOGNATHUS ZONE 111

In occlusal view the tip of M1 is seen to lie in line with the labial cusps of the

following teeth. Posterior teeth are more heavily worn, particularly on the lingual

cusp, with M6 having actually been broken some time well before death. The

occlusal plane, i.e. the best fit curve linking the occlusal surfaces of the teeth, is

slightly convex ventrally.

Thelegnathus oppressus sp. nov.

Figs 2-4, 8A, C

Holotype

BPI 155, a small, nearly complete but distorted skull with well-preserved

dentition.

real Sn

at uate

B 1 POT

5 mm pe ete

jae

(e ay (Gud

ee ae

-T>

D 1 cm

Fig. 2. Thelegnathus oppressus BPI 115. A. Labial view of left dentition. B. Labial view of

right dentary teeth. C. Occlusal view of same. D. Dorsal outline of the skull.

2 ANNALS OF THE SOUTH AFRICAN MUSEUM

Paratypes

BPI 4586, a beautifully preserved small skull with teeth occluded, and

BPI 4584, the left half of a skull, somewhat larger and with good molar teeth.

Both from the same locality.

Locality

The farm Hugoskop, district Rouxville, from the Cynognathus zone.

Diagnosis

Molar teeth bulbous at the base with the crowns pinched up to present a

small occlusal area.

Etymology

From the Latin apprimo = crush, refers to the powerful crushing teeth.

Description

BPI 155 is a small skull found high in the Cynognathus zone (J. W. Kitching,

pers. com.). Preservation is patchy, apart from the dentition. The roughened

quadratojugal projects very slightly obliquely backwards. A subhorizontal break

has exposed the right dentary teeth (Fig. 2B—C) and facilitated their preparation.

The left dentition (Fig. 2A) has only been developed in external aspect. The

dental formula of Iz M$ is what would be expected in Procolophon of this

size (Gow, in press), however, the teeth are much more massive and differently

shaped. They are broad and bulbous at the base, and the crowns, although still

transversely widened, are pinched to present a much smaller occlusal surface.

The pattern of wear on the dentary teeth (C) produces a short, narrow, flat,

transverse terminal facet set off from a deep facet on the anterior surface of the

tooth (facets demarcated by a solid line). In molars three and four, wear is so

heavy that a dark discoloration caused by the pulp cavity is visible at the centre

(black areas).

Especially interesting is the groove which deeply undercuts the molar row

on the labial surface, forming the curious sulci at the base of each tooth first

reported by Ivachnenko (1974) in Contritosaurus.

The paratype BPI 4686 is shown in outline in Figure 3. A notable feature

is that the teeth are set well in from the jaw margins. No suture detail is visible.

Lateral views of the teeth of this specimen (A) and the type (B) are shown in

Figure 4, from which it can be seen that they are substantially the same.

Thelegnathus perforatus sp. nov.

Figs 4-5, 8B, 9

Holotype

BPI 4585, a small, badly crushed skull with well-preserved teeth.

Locality

The farm Hugoskop, district Rouxville, from the Cynognathus zone.

NEW PROCOLOPHONIDS FROM THE TRIASSIC CYNOGNATHUS ZONE 1133

Diagnosis

Dentary incisiform teeth large and robust. Occlusal plane forms distinct

angle with the tooth-bearing surface of the dentary. Multiple tooth rows present

in maxillae and dentaries.

Etymology

From the Latin perforo = pierce, refers to the piercing molars.

Description

This is a badly crushed skull with well-preserved dentition. The right maxilla

is covered by the right lower jaw, which is displaced and closely applied against

the exterior surface of the maxilla, which has thus not been prepared at this stage.

The full left maxillary dentition is preserved (Fig. 5A, D), and contains

elements of three Zahnreihen. The labial row comprises one large incisiform tooth

followed by five molars of which the last, and youngest, is the only unworn tooth.

These teeth are massive pointed cones; wear is due to tip-to-tip pounding, and

there is also a slip-wear facet on the lingual surface of the three central molars

caused by rubbing against the corresponding lower teeth—hence we know that

the lower teeth occluded within the uppers.

alernany

Fig. 3. Thelegnathus oppressus BPI 4586.

Paratype: outline drawings of the skull.

114 ANNALS OF THE SOUTH AFRICAN MUSEUM

In the left dentary (Fig. 5B, E-F), there are two mature incisors followed

by a tooth space, followed by three worn and two unworn molars comprising a

Zahnreihe. The three anterior (therefore oldest) incisors of the next Zahnreihe

are preserved in position. One or two tooth buds have probably been lost behind

these.

The right dentary dentition (Fig. 5C, G) differs in having the first incisor

broken off and in having only four molars in the oldest Zahnreihe; the second

Zahnreihe is also more advanced.

The teeth of this specimen are shown in Figure 4C. The relatively enormous

size of the incisors compared with those of 7. oppressus is clear. In T. oppressus

the slope of the occlusal plane is parallel to the tooth-bearing surface of the

dentary, whereas in 7. perforatus the occlusal plane forms a pronounced angle

with the surface of the dentary.

t 1 cm

Fig. 4. Comparisons of the teeth of several new species in labial view. A. Thelegnathus oppressus

BPI 4586. B. Thelegnathus oppressus BPI 155. C. Thelegnathus perforatus BPI 4585.

D. Thelegnathus contritus BPI 3512.

NEW PROCOLOPHONIDS FROM THE TRIASSIC CYNOGNATHUS ZONE lS)

Thelegnathus contritus sp. nov.

Figs 4, 6, 8D

Holotype

BPI 3512, a skull with some associated post-cranial bones, lacking the snout

and with the right side of the skull weathered away.

Locality

The farm Winnaarsbaken, district Burghersdorp, from the Cynognathus zone.

Fig. 5. Thelegnathus perforatus BPI 4585. Details of the dentition of the type. A. Left maxillary

dentition in occlusal view. D. The same in labial view. B. Left dentary dentition in occlusal

view. E. The same in labial view. F. The same in lingual view. C. Right dentary dentition

in occlusal view. G. The same in labial view. Zahnreihen are indicated by dashed lines. Teeth

of younger Zahnreihen replace those to which they are joined by dotted lines. Occluding teeth

are linked by dashed arcs.

116 ANNALS OF THE SOUTH AFRICAN MUSEUM

Diagnosis

One tooth of the molar series almost twice as broad in lateral aspect as any

other. (The position of this tooth in the row will vary with the age of the indivi-

dual. Tooth replacement is demonstrated in another paper (Gow, in press).)

Etymology

From the Latin contero = pound.

Description

All the preserved teeth are probably molars, though there could be some

doubt about the anterior dentary teeth. Posterior molars are all broadened mesio-

distally, but the second of these thickened teeth in both jaws is almost twice as

1cm

Fig. 6. Thelegnathus contritus BPI 3512. A. The skull in dorsal outline. B. The skull in lateral

view. C. Labial outline of the dentition. D. Sketches of crown shape in mesiodistal and

occlusal aspects.

NEW PROCOLOPHONIDS FROM THE TRIASSIC CYNOGNATHUS ZONE 7

thick as any other. In this feature this species differs substantially from those

described above. The occlusal plane is curved and the molars are all heavily worn

to the form sketched in Figure 6D. The quadratojugal of this specimen is small

and smooth, but the taxonomic significance of this is unknown. (In Procolophon

the morphology of the quadratojugal varies with the age of the individual.)

Thelegnathus spinigenis sp. nov.

Fig. 7

Holotype

BPI 4299, a reasonably complete and undistorted skull.

Paratypes

BPI 3894, 4300, 4587 and 4588, all skulls.

Locality

The farm Lemoenfontein, district Rouxville, from the Cynognathus zone.

Diagnosis

Quadratojugal produced postero-laterally in the form of a large blunt spine.

(No species diagnostic dental information available.)

Etymology

From the Latin spina = spine, and gena = cheek.

Description

The five skulls from one locality all clearly belong to one species. Drawings

are taken from the type except for the teeth, which are those of 4300. In all cases

bone is soft and the nodular red rock matrix extremely hard, thus, while the

teeth are seen to broaden posteriad and to be heavily worn, no detail of the

occlusal surfaces is available. There is no suture detail that is anything but

typically procolophonid. What all the skulls have, however, is the enormous

quadratojugal which extends the line of the cheek backwards in the form of a

large, blunt spine which in life may have borne a sharp keratinous spine protect-

ing the neck and shoulder region.

Remarks

This form must be regarded as a distinct palaeontologic species. There is

the possibility that it might be the adult form of 7. oppressus; however, 4300, the

smallest of the five, is not much bigger than T. oppressus so the inclination is to

believe that 7. spinigenis is a true biologic species.

DISCUSSION

The multi-rowed dentition of Thegnathus perforatus

In the left maxilla (Fig. 5A, D), initiation of anlagen at each tooth position

clearly followed rapidly on the formation of their predecessors, as lingual to the

marginal row of teeth the next row is nearly completely formed. This can be

118 ANNALS OF THE SOUTH AFRICAN MUSEUM

stated another way, by saying that the Zahnreihe slope (dashed lines connecting

teeth, Fig. 5) is roughly parallel to the jaw. Of the oldest Zahnreihe the incisor

has already been shed, while the interesting point about the second Zahnreihe is

that the first two molars are already partially worn. A third Zahnreihe is repre-

sented by an incisor and first molar. In the maxillary dentition, wear on molars

of the second Zahnreihe is interesting. The only way this wear could have

I 5cm j

I 1cm j

Fig. 7. Thelegnathus spinigenis BPI 4299.

Skull in dorsal and lateral views.

BPI 4300. Labial outline of teeth.

NEW PROCOLOPHONIDS FROM THE TRIASSIC CYNOGNATHUS ZONE 119

A B

Fig. 8. A. Thelegnathus oppressus dorsal view of paratype BPI 586; x1. B. Thelegnathus

perforatus dorsal view of type BPI 4585; «1. C. Thelegnathus oppressus BPI 586. Stereo-

photographs of left dentition; x2. D. Thelegnathus contritus BPI 3513. Stereophotographs of

left dentition; 2.

120 ANNALS OF THE SOUTH AFRICAN MUSEUM

Fig. 9. Thelegnathus perforatus BPI 4585. Stereophotographs of the left lower jaw and teeth.

NEW PROCOLOPHONIDS FROM THE TRIASSIC CYNOGNATHUS ZONE 121

occurred would have been by contact with the first two lower molars before the

latter reached their present position, hence the wear on a tooth may be due in

part to contact with another long since replaced.

With respect to the dentary dentitions and due probably to the marked

heterodonty as between molars and incisors, it is not possible to incorporate both

types of teeth within the same Zahnreihe and it seems probable that replacement

operated independently (as appears to be the case in Procolophon (Gow, in press)).

Replacing teeth in the dentary are interpreted as having formed beside the teeth

to which they are linked by dotted lines (Fig. 5), but replacing those to which

they are linked by solid lines. This means that the functional teeth moved antero-

labiad before being finally shed.

The dentition of T. perforatus as described above is strikingly similar to that

of Captorhinus aguti (Bolt & De Mar 1975). One inference from this is that

multiple rowed dentitions in cotylosaurs are a Permian specialization retained

in T. perforatus. The alternative possibility is that the same dental arrangement

has been evolved independently at different times in these two animals, i.e. that

there is a genetically inherent predisposition to the attainment of this condition

in cotylosaurs.

The teeth of most known procolophonids are labio-lingually broadened and

distinctly bilobate (this is described elsewhere (Gow, in press)). Exceptions to

this are some of the primitive Russian forms and Owenetta. It is becoming clear

from much data on tooth replacement in reptiles by various workers, including

the present writer, that rate of replacement is often variable with the age of the

individual (typically, it seems, slowing with age). This affects Zahnreihen spacing

and the number of teeth in a replacement sequence. These remarks are necessarily

condensed as this is primarily a taxonomic paper; they lead to the following

tentative statement.

Captorhinus and procolophonids presumably share common ancestry from

romeriid captorhinomorphs. Captorhinus has attained a multiple tooth-rowed

state by lengthening of Zahnreihen, and the same condition is seen in Thelegnathus

perforatus; what then is the odontological history of the more typical broadened

bilobate procolophonid tooth? At this stage one can suggest no more than a

possibility, and this is done here in the hope of generating further interest, that

possibility being that the bilobate molar may represent a fusion of two tooth

buds from adjacent Zahnreihen.

One is not, in fact, faced with a choice between direct relationship between

T. perforatus and C. aguti or the development much later in time of a Captorhinus-

like condition in Thelegnathus. Rather, these apparently dramatic dental patterns

are achieved through the plasticity inherent in the reptilian dental replacement

cycle.

General

The sudden appearance in the Cynognathus zone of five species of procolo-

phonid is most unexpected. That one of these should in addition possess a pattern

[22 ANNALS OF THE SOUTH AFRICAN MUSEUM

of tooth replacement apparently more primitive than that in any other known

procolophonid regardless of age, is remarkable.

Several points arise, but perhaps the overriding consideration is the inference

that some aspect of procolophonid biology has mitigated against their preserva-

tion in the fossil record except under certain peculiar circumstances.

Hotton & Kitching (1963) suggested that what was then known as the

Procolophon zone, and indeed the overlying Cynognathus zone as well, may have

been laid down discontinuously on an erosion-surface of considerable relief.

Unfortunately this important concept still remains to be tested by a regional

geological study. What is certain is that procolophonids from both these zones

occur in hard red rocks typical of terrestrial redbeds.

Colbert (1946) discusses the geological setting of Hypsognathus in some

detail. Hypsognathus was found in sandstone of the Newark Series. Taking a

closer look at this series we find that it was laid down in a block-faulted area of

high relief. Deposition of the Newark was very rapid, hence the lower section is

nearly all sandstone, but redbeds towards the top (over part of the area of

deposition) are cited by Pettijohn (1957) as a classic terrestrial redbed sequence.

Newark plant fossils correlate floristically with those of the southern African

Molteno formation (J. T. Brown, pers. com.).

Procolophonids are also known from the Triassic fissure fillings of Britain

(Robinson 1967); these are as yet undescribed. These fillings contain an upland

fauna.

These few observations suggest that the reason procolophonids occur only

sporadically in the fossil record is that they were upland animals and would not

have occurred in large depositional basins.

All the broad-toothed procolophonids were almost certainly plant-eaters.

If it can be assumed that the large, well-known anomodonts were by and large

lowland forms, then the procolophonids as upland herbivores would have had

little competition from other known reptiles between the disappearance of the

edaphosaurs and the arrival of the gomphodonts and bauriamorphs, with the

exception of some of the rhynchosaurs and possibly small anomodonts like

Myosaurus (this might also account for the extreme rarity of the latter). Thus it

is possible that for much of the Permian and Triassic the procolophonids were

not offered any serious competition as the only upland herbivorous reptiles.

However, their depredations on particularly the reproductive parts of the vegeta-

tion very likely resulted in an interaction producing evolutionary responses on

both sides.

This last observation seems particularly to hold good for the late Triassic

as regards both plant (J. T. Brown, pers. com.) and animal. Thelegnathus browni

clearly had powerful crushing teeth (even to the extent of the most recently

erupted molar in the type having broken in use). Such an animal would make

a very efficient seed-eater. T. oppressus was an even better crusher of hard seeds

with its large, bulbous, sharp-pointed molars. In T. perforatus the incisors are

large and powerful, such as might be used for grabbing and wrenching off the

NEW PROCOLOPHONIDS FROM THE TRIASSIC CYNOGNATHUS ZONE 123

seed of a particularly tough plant. The first maxillary tooth is a piercing tooth

which does not meet in tip-to-tip occlusion with any other. Other molars,

although the tips do wear, seem to be primarily piercing teeth. On this basis it

is suggested that this animal utilized only the soft outer coating of certain seeds.

Molar tip wear would then simply be due to contact with the hard seed.

T. contritus has well-worn pounding teeth rather like those of Procolophon: this

species may have been a foliage feeder. Nothing can be said concerning the diet

of 7. spinigenis until a specimen turns up which is more amenable to

preparation.

SUMMARY

The description of the procolophonid Thelegnathus browni from the

Cynognathus zone is expanded and four new species referable to this genus are

described.

Taxonomic separation is based entirely on details of tooth morphology, the

various species being specialized for different herbivorous diets. T. browni and

T. oppressus are thought to have been crushers of hard seeds. 7. perforatus may

have lived on the softer outer coatings of seeds. 7. contritus was a foliage

eater.

One of the new species, 7. perforatus, had multi-rowed marginal teeth. The

Zahnreihen are parallel to the jaw and follow in rapid succession, which suggests

the possibility that the specialized broadened bicuspid teeth of typical procolo-

phonids may represent a fusion of two anlagen in adjacent Zahnreihen.

Preliminary evidence is presented in the discussion which suggests that

procolophonids were upland animals. Aside from fissure fillings they are typically

associated with discontinuous redbed facies. This may account for the patchy

appearance of these animals in the fossil record. It may also account for their

success as they would then be removed from competition with most known

anomodonts.

ACKNOWLEDGEMENTS

I thank Dr Michael Cluver of the South African Museum, Cape Town for

the loan of the genotype. Dr James Kitching of the Bernard Price Institute for

Palaeontological Research collected all the new material with his usual skill and

enthusiasm. Dr John Brown, also of this Institute, contributed lively discussion

on plant-animal interactions.

124 ANNALS OF THE SOUTH AFRICAN MUSEUM

REFERENCES

Bott, J. R. & DE Mar, R. 1975. An explanatory model of the evolution of multiple rows of

teeth in Captorhinus aguti.—J. Paleont. 49: 814-832.

Broom, R. 1905. Preliminary notice of some new fossil reptiles collected by Mr. Alfred Brown

at Aliwal North, S. Africa.— Rec. Albany Mus. 1: 269-275.

Broom, R. 1936. The South African Procolophonia.— Ann. Transy. Mus. 18: 387-391.

CocsertT, E. H. 1946. Hypsognathus, a Triassic reptile from New Jersey.— Bull. Am. Mus. nat.

Hist. 86: 227-274.

EDMUND, A. G. 1960. Tooth replacement phenomena in the lower vertebrates. — Life Sci. Contr.

R. Ont. Mus. 52: 1-190. :

Gow, C. E. In press. Tooth function and succession in Procolophon trigoniceps.— Palaeontology.

Hotton, M. & KITCHING, J. W. 1963. Speculations on upper Beaufort deposition.—S. Afr.

J. Sci. 59: 254-258.

IVACHNENKO, M. F. 1974. New data on the early Triassic procolophonids.—Paleont. Zh. (3):

68-74.

PETTIJOHN, F. J. 1957. Sedimentary Rocks. New York: Harper.

RoBInson, P. L. 1967. Triassic vertebrates from lowland and upland. — Sci. Cult. 33: 169-173.

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. nov., sp. nov., comb.

nov., syn. nov., 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°51’S 25°39’E), collected by A. Smith, 15 January 1973.

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

7. SPECIAL HOUSE RULES

Capital initial letters

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

e.g. ‘... the Figure depicting C. namacolus...’; ‘. . . in 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; Von 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.’ eae

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. ;

Name of new genus or species is not to be included in the title: it should be included in the

abstract, counter to Recommendation 23 of the Code, to meet the requirements of

Biological Abstracts.